HOPS

HOPS
R.A. Neve
OBE, PhD. Dip Agric Sci, FRASE, FI. Brew, MI Biol

Formerly
Director, Hop Research Department, Wye College

SPRINGER-SCIENCE+BUSINESS MEDIA, B.V.

First edition 1991
@ 1991 R. A. Neve
Originally published by Chapman aud Hall1991
Softcover reprint ofthe hardcover Ist edition 1991
Typeset in 10/ 12pt Times by Witwell Ltd, Southport

Bury St. Edmunds, Suffolk

ISBN 978-94-010-5375-4

AII rights reserved. No part of this publication may be reproduced or transmitted, in

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made to the publisher.

The publisher makes no representation, express or implied, with regard to the accuracy
of the information contained in this book and cannot accept any legal responsibility or
liability for any errors or omissions that may be made.

British Library Cataloguing in Publication Data

Neve, R. A. (Ray A.) 1922-
Hops.
1. Hops. Production
1. Title
633.82
ISBN 978-94-010-5375-4 ISBN 978-94-011-3106-3 (eBook)
DOI 10.1007/978-94-011-3106-3
Library of Congress Cataloging-in-Publication Data

Available
Contents

Introduction IX
Units and Abbreviations xi

1 Botany 1
1.1 The genus 1
1.1.1 Humulus lupulus 1
1.1.2 Humulus japonicus 9
1.1.3 Humulus yunnanensis 10
1.2 Sex ratios 10
1.3 Cytology 11
1.4 Polyploidy and sex determination 12
1.5 Centre of origin 16
1.6 Photoperiodic response 16
1. 7 Dormancy 18
1.8 Photosynthesis and carbohydrate reserves 20

2 The cultivated hop 25

2.1 History 25
2.2 Use 29
2.2.1 Hop resins 33
2.2.2 Essential oils 38
2.2.3 Tannins 43
2.3 Hop processing 45
2.4 In vitro production 47

3 Production methods 49
3.1 Supporting structures 49
3.2 Cultural operations 53
3.2.1 Non-cultivation 54
3.2.2 Training 56
3.2.3 Stripping 57
VI Contents
3.3 Soils 59
3.4 Manuring 59
3.4.1 Nitrogen 60
3.4.2 Phosphate and potash 61
3.4.3 Magnesium 62
3.4.4 Trace elements 62
3.5 Irrigation 62
3.6 Spraying 64
3.7 Low trellis 66
3.8 Seeded and seedless hops 68
3.9 Growth substances 70
3.10 Propagation 72
3.11 Factors affecting quality 75

4 Harvesting 79
4.1 Picking 79
4.1.1 Hand picking 79
4.1.2 Machine picking 84
4.2 Drying 89

5 Pests and diseases: historical review 101

6 Pests 115
6.1 Damson-hop aphid: Phorodon humuli 115
6.1.1 Description 115
6.1.2 Chemical control 120
6.1.3 Biological control 121
6.2 Spider mite: Tetranychus urticae 126
6.2.1 Description 126
6.2.2 Chemical control 129
6.2.3 Biological control 130
6.3 Nematodes 131
6.3.1 Dagger nematode: Xiphinema diversicaudatum 131
6.3.2 Hop-root eelworm: Heterodera humuli 132
6.4 Other pests 133
6.4.1 Clay-coloured weevils: Otiorrhynchus singularis 133
6.4.2 Rosy rustic moth: Hydroecia micacea 134
6.4.3 Flea beetle: Psylliodes attenuata 135
6.4.4 Earwigs: Forficula auricularia 135
6.4.5 Wireworm: Agriotes spp. 136
6.4.6 Slugs: Agriolimax reticulatus and Arion hortensis 136
Contents Vll

7 Fungal diseases 137

7.1 Downy mildew: Pseudoperonospora humuli 137
7.1.1 Overwintering 141
7.1.2 Secondary infection 143
7.1.3 Hygiene 144
7.1.4 Chemical control 145
7.1.5 Epidemiology 146
7.1.6 Resistant cultivars 149
7.2 Powdery mildew: Sphaerotheca humuli 150
7.2.1 Overwintering 153
7.2.2 Secondary infection 154
7.2.3 Hygiene 155
7.2.4 Chemical control 156
7.2.5 Epidemiology 156
7.2.6 Resistant cultivars 157
7.3 Verticillium wilt: Verticillium albo-atrum 160
7.3.1 Description 160
7.3.2 Hygiene 165
7.3.3 Cultural methods 165
7.3.4 Chemical control 167
7.3.5 Biological control 167
7.3.6 Resistant cultivars 168
7.3.7 Verticillium dahliae 169
7.4 Other fungal diseases 170
7.4.1 Fusarium canker: Fusarium sambucinum 170
7.4.2 Black root rot: Phytophthora citricola 171
7.4.3 Grey mould: Botrytis cinerea 171
7.4.4 Alternaria alternata 172
7.4.5 Black mould: Cladosporium 172
7.4.6 Armillaria root rot: Armillaria mellea 172

8 Virus diseases 175

8.1 Hop mosaic virus (HMV) 176
8.1.1 Control 180
8.2 Hop latent virus (HLV) 180
8.3 American hop latent virus (AHLV) 180
8.4 Arabis mosaic virus (AMV) 182
8.4.1 Nettlehead 183
8.4.2 Split leaf blotch 184
8.4.3 Bare bine or spidery hop 184
8.4.4 Hop chlorotic disease 185
8.4.5 Control 187
Vlll Contents
8.5 Prunus necrotic ringspot virus (NRSV) 189
8.5.1 Control 191
8.6 Other viruses 191
8.7 Viroids 192
8.7.1 Hop stunt viroid (HSVd) 192
8.7.2 Hop latent viroid (HLVd) 192

9 Varieties and breeding 195

9.1 Historical introduction 195
9.2 Aroma hops 198
9.3 Bitter hops 202
9.4 Varietal acreages 207
9.5 Future developments 210
9.5.1 Pesticides 210
9.5.2 Dwarf hops 210
9.6 Inheritance of important characters 211
9.6.1 Resin contents 211
9.6.2 Yield 213
9.6.3 Resistance to verticillium wilt 213
9.6.4 Resistance to downy mildew 214
9.6.5 Resistance to powdery mildew 214
9.6.6 Reaction to viruses 215
9.7 Polyploidy 215
9.8 Mutations 216
9.9 Wild hops and gene pools 217
9.10 Characteristics of principal cultivars 218

10 The hop trade 225

References 235
Index 261
Introduction

It is 25 years since Dr Burgess wrote his invaluable book on hops and in the
intervening period there have been very many advances in hop research and
hop production techniques. When invited to produce a replacement for that
book, therefore, the problem was not finding enough new material but
deciding on what to include.
People interested in reading about the hop are likely to fall into very diverse
categories. Hop growers will be looking for practical advice on production
methods while research workers with specialist knowledge in one field may
want detailed information about research in other disciplines. In addition,
there are many people for whom hops are of much more general interest and
for them a source of basic information about the crop will be required.
The aim has not been to produce a detailed growers' handbook, since
techniques vary considerably from district to district and I believe that it is
better to obtain advice from neighbouring growers or from specialist advisers
than from any book. What I have attempted is to outline the basic principles
upon which production methods should be based. At the same time, I have
tried to include material that will be of general interest both to those who work
with hops and to those to whom they might otherwise remain a complete
mystery. In doing this my own personal interests have inevitably played an
important part. Since it is difficult for most people to obtain access to
historical accounts I have included extracts from those that have fascinated
me. In particular, there are several extracts from Reynolde Scot's Perfite
Platforme of a Hoppe Garden. Reference has, been made throughout, to the
second (1576) edition of this book as it includes some material that is not to be
found in the first edition of 1574.
One book cannot possibly include all the information that readers may
require and in the course of writing it I have become aware of a great deal of
literature that was new to me. The attempt to record some of the history of hop
growing, the achievements of early workers which might otherwise be over-
looked and details of current work has resulted in a very extensive bibliogra-
phy. By some this may be considered excessive but it is hoped that many will
find it a useful source of information.
x Introduction

It was decided not to invite specialists to write individual chapters since this
might result in a more disjointed text than one written by a single author. As
director of the Hop Research Department at Wye for many years I have been
associated with a dedicated team of workers dealing with most aspects of hop
research. That certainly does not mean that I am an expert in them all and, in
order to avoid gross errors and omissions, most of the chapters have been
referred to colleagues for their comments. While most gratefully acknowledg-
ing the help of Dr Tony Adams, Dr Colin Campbell, Dr Peter Darby, Dr Greg
Lewis, Mr Mike Shea, Dr Philip Talboys, Dr Mike Thresh and Mr Arthur
Winfield, I would emphasise that they cannot be held responsible for any
errors that may remain.
An exception to the general scheme was made when it was suggested that
there should be a section on hop chemistry. This was something which I was
not competent to write and Dr Roger Stevens very generously provided a
section that is incorporated in Chapter 2. To him, I am particularly grateful.
My thanks also go to Mr Bob Barrar for preparing new drawings for Figures
1.1, 1.2 and 1.3 as well as for permission to re-use Figure 3.10.
The Hop Research Department at Wye and East Malling Research Station
have recently been grouped together as part of the Institute of Horticultural
Research and illustrations from those sources are acknowledged as from IHR,
Wye and IHR, East Malling.
The book has, throughout, had the support of the members of the Scientific
Commission of the International Hop Growers Convention and I am most
grateful for the encouragement that they have given me.
Hop research is notable for the very close co-operation that exists, not only
between workers in the various hop growing countries but also between
research workers, hop growers and brewers. It is this friendly and stimulating
atmosphere that has made working on the crop such a rewarding personal
experience.
Finally I would thank my wife, Lesley, for encouraging me to undertake the
task of writing the book and for enduring the consequences.

R.A. Neve
Units and abbreviations

Since the book is likely to be referred to by practical farmers it was decided to

use the more everyday units in preference to the SI units which are likely to be
confusing for some people. Generally, both metric and imperial units have
been quoted, priority being given to metric except when imperial was used in
the publication being quoted. The only unit used that may be unfamiliar to
some readers is the zentner (zr) which is equivalent to 50 kg and is the standard
unit for commercial hop dealings in Europe.
Where initials are used to identify countries these are given according to
their own convention. Thus the English form is used for the UK and the USA
but DDR is used for the area formerly known as East Germany and BRD,
similarly, for West Germany.
CHAPTER 1

Botany

1.1 THE GENUS

The genus Humulus consists of three species, H. iupuius, H. japonicus and H.
yunnanensis. The cultivated hop, H. lupulus, is indigenous throughout much of
the Northern hemisphere between the latitudes of approximately 35° and 70° N.
H. japonicus is widespread throughout much of China and Japan but has not
been recorded elsewhere. Remarkably little is known about H. yunnanensis of
which there are very few herbarium specimens and no plants in cultivation.
Humulus and Cannabis are the only two genera in the family Cannabinaceae
and there are many similarities between hemp (Cannabis sativa) and the
cultivated hop. The nettle family is also rather less closely related being in the
same order, the Urticales. It is possible to produce viable grafts between hops and
hemp and it is reported that pollination of hops by hemp, annual nettle ( Urtica
urens) or perennial nettle (Urtica dioica) stimulates cone development but only
abortive embryos are produced (Schnarf, 1929. Quoted by Davis, 1956).

1.1.1 Humulus lupulus

This is a dioecious perennial climbing plant without tendrils, the bines twining
round any available support in a clockwise direction with the aid of hooked
hairs located on the angles of the stem. Darwin (1882), while ill in bed, watched
a hop plant growing on his window-sill and noted that the tip of the bine
completed a revolution in two hours. This was later confirmed by Shea
(unpublished) using time-lapse photography. Darwin concluded that the
twining was brought about by a wave of growth travelling around the shoot in
the extending region. Bell (1958) investigated the twining of the hop on
supports with diameters ranging from 0.25 in (0.635 cm) to 3 in (7.62 cm) and
found that the steepness with which the stem twined diminished with increas-
ing diameter of the support, suggesting that the radius of curvature of its helix
remained constant.
The leaves are normally borne in pairs at each node although a tricussate
arrangement is sometimes found. The leaves have a toothed margin and the
shape varies from cordate to 7-lobed although 3-5 lobes are the most common
2 Botany

in mature leaves (Fig. 1.1). Wild American hops generally have more lobes and
are more deeply divided than European or Japanese hops and most cultivars.
The flowers of the male plants are produced in loose panicles, and have a
perianth of five yellowish-green sepals and five anthers on short filaments. The
anthers, which have a furrow in which a few resin glands are located, dehisce to
produce large quantities of pollen which is wind-borne (Figure 1.2).
Female flowers occur in inflorescences that consist of a condensed central
axis; on each node there is a pair of bracts. Each bract subtends a pair of
bracteoles and each bracteole has a small flower enclosed in a fold at the base.
The flower consists of an ovary, enclosed in the perianth, with a pair of
papillated stigmas. As these inflorescences mature, the central axis elongates
and the bracts and bracteoles enlarge to produce the strobiles (commonly
referred to as 'cones) which form the commercial product of the plant (Figure
1.3). Flowers that are pollinated develop to form an achene enclosed in the
papery perianth and their development stimulates the bracteoles in which they
lie to extend much further beyond the bracts than do those bracteoles without
seeds. Pollination also results in elongation and thickening of the central strig
(Figure 1.4), and the nodes bearing seeded flowers are frequently pigmented.
These changes make it easy to distinguish between seeded and seedless cones.
The commercial value of hops lies in the lupulin glands which contain resins
(of which the a-acid is the most important fraction) which give beer its
bitterness and essential oils which contribute hop flavour. Whereas male
flowers develop only a small number of resin glands, these are abundant on the
cones of the female plants (Figure 1.5). They are most numerous on the
bracteoles with relatively few on the bracts. They are dense on the pericarp and
in seeded hops up to a third of the a-acid in the cone is produced there. In
seedless cones the pericarp remains vestigial and contributes only a small
proportion of the whole. There are great variations between different geno-
types in the number and size of the glands in the cones and these differences are
commercially very important since it is the glands that contain everything that
is of value to the brewer. Neve (1968) investigated the distribution of a-acid on
the different parts of seeded and seedless cones of the three cultivars Fuggle,
Northern Brewer and Bullion with the results shown in Table 1.1.
Although resin glands were found on the strigs, examination indicated that
these had not been formed there but had become detached from other parts
during the dissection of the cone and were adhering to the fine hairs which cover
the strigs. Resin glands are also produced on the underside of the leaves but are
not sufficiently abundant to make the leaves of any value as brewing material.
Thomas (1980a) has published electron microscope photographs showing
the stages of development of the glands on cones (Figure 1.6). The separation
of the sexes between plants means that self-pollination is not normally possible
and consequently hop plants are genetically very heterozygous. Therefore,
when plants are raised from seed the progeny are extremely variable and the
The genus 3

Figure 1.1 Variations in hop leaves (after R. F. Farrar).

4 Botany

Figure 1.2 (a) Young shoot; (b) male flowers; (c) male and female flowers of
monoecious plant; (d) anthers with resin glands in the furrows (after R. F. Farrar).

majority are of little value commercially. For this reason, hop gardens are
invariably planted with material that has been propagated vegetatively from
one of the established cultivars.
The above-ground parts of the plant die back to ground level each winter but
the below ground 'rootstock' is perennial and can survive for very many years.
Although referred to as a rootstock the upper part is, in fact, stem tissue (Figure
1.7). In a mature plant the root system can extend downwards for 1.5 m or more
and laterally for 2-3 m although the extent of lateral spread is less on deeper
soils (Beard, 1943a). There are two types of roots: horizontal roots that are
tough, wiry and branching which produce much fibrous growth within the top
20-30 cm of the soil and vertical roots which arise from the crown or from
horizontal roots and are fleshy, irregularly swollen and easily broken. The roots
do not produce buds and regeneration each spring is from buds on the branched
stem tissue which lies below ground level and forms the upper part of the
rootstock. These buds produce a mass of shoots in the spring - far more than
are needed to grow a crop - so that the grower has to remove most of them.
Some buds also develop into runners which can spread sideways, just below
ground level, for 1 m or more although most will emerge above ground closer to
the rootstock.
The genus 5

Figure 1.3 (a) 'Pin'; young flowering shoots developing in the leafaxils; (b) 'Burr';
young female inflorescence with papillated stigmas. Mature cones of (c) Fuggle; (d)
Wye Target; (e) Yeoman; (f) 'Strig'; central axis of cone; (g) bract; (h) bracteole with
enclosed seed; (i) bracteole with seed; (j) removed (after R. F. Farrar).

There are considerable differences between the plants to be found in various

parts of the world and botanists have on several occasions suggested that the
variation is sufficient to justify classifying them into different sUb-species or
even into separate species. The wild hops of North America have been named
H. americanus or H. neomexicanus while the type to be found in Japan (and
possibly China) with cordate leaves is named H. cordifolius. All these forms
6 Botany

Figure 1.4 Strigs of unpollinated (left) and pollinated (right) cones.

are totally interfertile and most workers do not now separate them. Davis
(1957) examined a range of specimens and separated them into three mor-
phological complexes which were represented by English, German-Czech and
American (Late Cluster) cultivars. Wild American hops had characters in
common with Late Clusters and Japanese wild hops but the variation within
the wild Americans was such that further investigation would be necessary
before the various forms could be separated.
In the most recent review of the genus, however, Small (1978) has moved in
the other direction and proposed dividing the species into five varieties, two of
which are new. Under his proposed nomenclature the European hop is H.
lupulus var. lupulus, that of Japan remains as H. lupulus var. cordifolius and
the form native to western North America as H. lupulus var. neomexicanus.
He considers these to be distinct from the plants native to the American
Midwest and the eastern part of the United States for which he proposes the
names H. lupulus var. pubescens (var. nov.) and H. lupulus var. lupuloides
The genus 7

Figure I.S Cones of traditional cultivar (left) and high a-acid cultivar Yeoman (right)
showing resin glands.

(var. nov.) respectively. Davis (1956) , while recognizing differences between

the hops of the West, Midwest and East, considered that these differences were
too slight to warrant separate names. Moreover, Hampton (personal commu-
nication) reports that he was unable to recognize a distinct type when
collecting wild hops in the Midwest.

TABLE 1.1 Alpha-acid in each part of the cone as a percentage of the total.
Seeded Seedless

Fuggle N. Brewer Bullion Fuggle N. Brewer Bullion

Bract 14.3 14.4 15.0 23.7 21.9 21.8

Bracteole 55.0 66.5 50.2 66.7 74.1 71.2
Seed 28.7 16.6 32.2 5.7 1.8 4.5
Strig 2.0 2.6 3.0 3.9 2.2 2.6
 (a) ...._ .

(c)

(e)
Figure 1.6 Scanning electron microscope photographs showing development of resin
glands on the following dates: (a) 30.7; (b) 6.8; (c) 10.8; (d) 29.8; (e) 15.8; (f) 7.9;
(Thomas, I 980a).
The genus 9

New shoot

Old stem

New root

Figure 1.7 Perennial storage organ of young plant consisting of new shoot, old stem of
original trimmed sett and new storage roots (Williams, 1960).

1.1.2 Humulus japonicus

This is typically an annual species but there are suggestions that it may
sometimes survive for more than one season. It is also dioecious and the cones,
although similar in structure to those of H. /upu/us, look quite different, ripen
sequentially and have few, if any, lupulin glands. They are therefore of no
value as brewing material. The leaves are normally 7-1obed and the plants are
covered with very strong hooked hairs which can make them quite painful to
work with. It is a strong climber and is sometimes grown horticulturally to
provide a leafy screen.

1.1.3 Humulus yunnanensis

Attention has been drawn to the existence of this species by Small (1978) but
little is known about it apart from a few herbarium specimens. It occurs,
apparently, at rather high altitudes in southern China, particularly in Yunnan
Province at a latitude of approximately 25°N. Small states that it is not known
whether it is an annual or perennial plant but information from China (Jinyu,
10 Botany
personal communication) is that it is a perennial. Since it grows at lower
latitudes than other members of the genus it could have potential as breeding
material to extend the areas in which hops could be grown commercially.

1.2 SEX RA nos

The fact that female hop seedlings normally occur more frequently than males
was first recorded by Winkler (1908) who suggested that it might be due to the
production of parthenogenetic seed. Tournois (1914) demonstrated with very
careful experiments that no seed was set when pollen was completely excluded,
confirming similar reports by Salmon and Amos (1908) and Winge (1914). In a
later paper Winge (1923) was the first to suggest that the excess of females was
the result of pollen competition.
Dark (1951) reviewed the published results on sex ratios and pointed out
that many of the reports gave no indication of germination percentage of seed
but that where it was quoted, it was very low. He concluded that this 'must
profoundly modify any genetical interpretation of the sex ratios counted in
mature families '.
Sex ratios were the subject of PhD theses by Neve (1961) and Smithson
(1963) who showed that when germination percentages were low there was a
greater preponderance of females and that females tended to germinate earlier
then males. If seed was subjected to adequate cold treatment, good
germination was obtained; it was then found that pollen competition was the
main reason for the excess of females. Even when this was eliminated,
however, by diluting the pollen or by placing single pollen grains on the styles,
females were still in excess. In most families the ratio was about 2: 1 but when
the female parent was the Saazer variety it was as high as 4: 1. Limited studies
indicated that half the male seedlings of the Saazer crosses developed into
pagoda dwarfs which survived for such a short time that their sex could only be
determined cytologically. This would account for the difference between
Saazer and other crosses but, in the absence of evidence to account for the
general 2: 1 ratio, Smithson could only postulate an incompatability system
operating on 50% of the male gametes.

1.3 CYTOLOGY
H. lupulus normally has a diploid chromosome number of 20 in both male and
female plants but H. japonicus has 16 chromosomes in the female and 17 in the
male. Nothing is known of the cytology of H. yunnanensis.
Occasional triploid plants have been identified as having occurred naturally
in H. lupulus. The first report was by Tournois (1914) who examined plants
that were basically male but had occasional female flowers at the ends of the
laterals and reported that these were triploids. This is the most common form
but at least two female cultivars developed from breeding programmes using
Cytology 11
diploid parents were later found to be triploids. These were AGG8 bred at Wye
and Huller Bitterer from Hull in Germany.
Both H. lupulus and H. japonicus have recognizable sex chromosomes and
these have been the subject of various investigations. Winge (1923), Derenne
(1954) and Jacobsen (1957) working with plants of European origin reported
the occurrence of a heteromorphic pair of sex chromosomes in the meiotic
divisions of male plants. At the same time the Japanese workers Sinoto (1929a,
b) and Ono (1937) were claiming that in the wild Japanese hop they found a
chain of four chromosomes that represented a sex chromosome complex while
in a later paper Ono (1955) reported on a wide range of male hops in which he
found five different sex chromosome types as follows:
1. 'Winge type': a heteromorphic pair with a size ratio of 10:5.2, and nine
bivalents;
2. 'New Winge type': a heteromorphic pair with a size ratio of 10:8 and nine
, bivalents;
3. 'Sinoto type': a quadrivalent with size ratios of 10.0:12.0:14.2:7.2 and eight
bivalents;
4. 'New Sinoto type': a quadrivalent with size ratios of 10.0:12.7:10.9:3.4 and
eight bivalents;
5. 'Homotype': ten bivalents.
Neve (1958a) studied meiosis in six male plants in which a quadrivalent
association occurred together with a heteromorphic pair which he identified as
the sex chromosomes. He suggested that this raised doubts about the
interpretation by Sinoto and Ono of their quadrivalents as a sex chromosome
complex. Ono and Suzuki (1962) examined wild American plants sent to them
by Neve and confirmed Neve's interpretation in some plants but in others
reported that the sex chromosomes formed part of the quadrivalent.
Neve (1961) found a heteromorphic pair of sex chromosomes corresponding to
Ono's 'Winge type' in all the European males that he examined while in males
from the USA or Canada there was either a heteromorphic pair corresponding to
the New Winge type or an equal sized pair as in the Homotype. Any chains offour
chromosomes that he found were in divisions where there was also a sex
chromosome bivalent of the New Winge type, identifiable by the unequal lengths
of the X and Y chromosomes (Figure 1.8). If a chain of four occurred in plants
with homotype sex chromosomes it would be difficult to be certain whether the
sex chromosomes were part of the quadrivalent or not. This may account for the
different interpretations put forward. Neve did not examine any Japanese wild
hops in which Ono and Suzuki (1962) have described tetrapartite and hexapartite
sex chromosome complexes.
In all the plants he examined, Neve (1961) found that the X chromosomes
could be clearly identified in mitotic divisions by means of the unstained region
in the short arm when using pre-treatment with 8-hydroxyquinoline as
12 Botany

(a)

Figure 1.8 New Winge type males with and without chain of four autosomes. XY
bivalent shaded; (a) male CG24; (b) male 321.

described by Jacobsen (1957) while the Y chromosome in Winge and New

Winge types was recognizable as the only one with a short arm ending
diffusively (Figure 1.9). The Y chromosome of homotype males was not
distinguishable in this way.

1.4 POLYPLOIDY AND SEX DETERMINA nON

The use of polyploidy in hop breeding was first reported by Ono (1948) and
was subsequently developed by Dark (1953) at Wye. The commercial reasons
for this are described in Section 9.7 but the availability of a wide range of
polyploid plants provided Neve (1961) with an opportunity to investigate the
cytological and genetic basis for sex determination.
Although hops are dioecious, some diploid female plants occasionally
develop some male flowers which are infertile. Weston (1960) reported the
production of male flowers on the female cultivar Fuggle following treatment
with a(2-chlorophenylthio )propionic acid. He did not state whether the male
flowers produced any pollen and it is likely that they did not do so. From
Australia, three male plants have been reported to occasionally produce female
cones with both sexes being fertile (Versluys, personal communication). When
grown at Wye College, England, two of these have never produced cones while
the third has produced a few cones in some seasons.
x

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 - - WE owe to the kindness of Mr. DARWIN the
communication oC the following, with the specimen to
which it refers : -
II I inclose a spr.cimen of the male Hop with appa-

rently female flowers at the tips of the branches. on

FIG. 37. -MOXCP.CIOUS I/OI'.

the chance of its having some interest for the naturalist.

I observed it this morning, and though accustomed
to walk Hop grounds for years [ have never seen the
two sexes on the same Hop plant before. Perhaps,
however. it is but the growth of the flower into a male
catkin.
"There are other male plants in the same ground,
but I have not seen any other instance of this peCUliarity.
The whole Hop hill groa.vs in the same way. If we
obtain seed, might not it be "possible to select a strain of
Hops which are uniformly monrecious on the same plant?
(Certainly. ]
II The Hop ground is in Boughton Monchelsea,
facing south. very warm, and of strong rich soil.
L. Lewis, East Farleiglz."

Figure 1.10 An early account of a monoecious male plant similar to those now known
to be triploids with XXY sex chromosomes.
Polyploidy and sex determination 15
Occasionally, in the progeny of diploid parents, plants are found that are
predominantly male with a terminal cone on some of the laterals but differing
from the Australian plants in that they shed little pollen. Such plants were
described on two separate occasions over 100 years ago (Masters 1852, Lewis
1874, Figure 1.10) and examination of recent examples has shown them to be
triploids.
Amongst the progeny of polyploid parents, however, there are many plants
with both male and female flowers (Ono, 1959) and the male flowers frequently
produce viable pollen. Neve (1961) examined the chromosomes in such plants
and established that there was a clear correlation between the sex of the plants
and the ratio of X: Y : A (autosome sets). In plants that had complete
chromosome sets the relationship was as follows:
Ratio X : Y: A Most frequent sexual expression
1:2:3 male
1:1:2 male
2: I :3 male with occasional female flowers
3: I :4 approximately equal male and female flowers
2 : 0 : -2 female
3:0:3 female
4:0:4 female
This relationship could be interpreted as a balance between X and Y, X and
A or Y and A chromosomes. Examination of aneuploid plants (having
chromosomes additional to or missing from the sets of 10) indicated that the Y
chromosome had little effect upon the sex of the plant and that most of the
male determining genes were located on the autosomes. It was the ratio of X:A
that was the main determinant of sex. Although the Y chromosome apparently
had no influence on the sexual form of the plants, it played an essential role
since male plants which lacked any Y chromosome failed to produce viable
pollen, development ceasing at the pollen tetrad stage. This is very similar to
the situation found in Drosophila. Although pairing between the sex
chromosomes is very restricted, the normal pairing and crossing over amongst
the autosomes allows for genetic segregation of the male-determining genes
and this would account for the wide range of sexual expression between plants
having similar chromosome complements.
The fact that male hops are of no value for brewing creates problems for
hop breeders in selecting males for use as parents in their breeding pro-
grammes. Males can be tested for such characters as disease resistance but
brewing quality cannot be directly assessed. There would be great advantages
if plants that produced hop cones could also be used as sources of pollen.
Since some diploid plants produce both male and female organs attempts
have been made to induce sex changes by chemical treatment but without any
success.
16 Botany
Neve (1965) suggested that tetraploid plants that were both male- and
female-fertile could be selected on their cone characters and the best used as
male parents. Some work was carried out along these lines but the two stages
of having first to breed good tetraploids and then to use them as parents in the
commercial programme were too time-consuming to meet the urgent needs of
the industry.
Another approach was based on the Australian diploid that was occasio-
nally monoecious. This is cytologically a male plant with XY chromosomes
that is presumably deficient in male determining genes in the autosomes. By
using this as a parent it might be possible to develop a diploid breeding line in
which many of the 'male' plants produced sufficient cones for their brewing
quality to be assessed. Since these males would also have XY chromosomes
they could produce viable pollen yet their female progeny would not be liable
to develop any male flowers. This programme was also commenced but the
frequency of the monoecious character was so low that no useful results could
have been expected for a very long time (Neve, unpublished).
The female flowers of tetraploid monoecious plants are no longer receptive by
the time the male flowers shed pollen so self-fertilization will not normally occur,
but Haunold (1972) achieved it by having clonal plants of the same genotype at
different stages of development. Extensions of this technique, or of the diploid
monoecious material could be used to develop more homozygous lines than are
at present available and these could provide useful breeding material.

1.5 CENTRE OF ORIGIN

The occurrence of three species in China suggests very strongly that this area is
the centre of origin of the genus. Migration eastwards to America and
westwards to Europe would have resulted in distinct populations in these two
areas and this conforms with the observations that the Y chromosomes in
European hops are different from those in America. This conclusion is
supported by Davis's observation of similarity in morphological characters
between wild hops from America and Japan.

1.6 PHOTOPERIODIC RESPONSE

The very first reports of flowering in plants being controlled by daylength were
in two classic papers by Tournois (1912, 1914) which described this phenome-
non in Humulus japonicus and the closely related hemp Cannabis sativa. He
demonstrated that flowering was induced when plants were raised in the winter
or when plants growing in the summer were covered to reduce the period of
exposure to light. Such plants would now be described as short-day plants.
Jackson (1955), who experimented with H. lupulus in Kenya, reported that
under natural conditions at that latitude the plants slowly died after making
only a few inches of growth but that they grew successfully if the natural
Photoperiodic response 17
daylength was extended to 15 hours by using artificial light. He accordingly
described the hop as a long-day plant.
Detailed determination of the photoperiodic responses of the cultivated hop
was reported by Thomas and Schwabe (1969) who demonstrated that it is a
true short-day plant responding to both length of day and light breaks in the
dark period. However, ifthe daylength is too short the plant becomes dormant
and fails to flower. A further controlling factor is that bines of less than a
critical size, as measured by the number of nodes, are unable to initiate
flowering, even in short days. Their results indicated that the Fuggle variety
was ripe to flower when 23 nodes were visible while the corresponding node
numbers for CC31 (a very early flowering variety) and New York hop (very
late flowering) were 12 and 20 respectively. Under experimental conditions (in
a glasshouse) the critical daylength for flowering was approximately 16.5 hours
for Fuggle and 15.5 hours for New York hop. At lower temperatures the
critical daylengths were longer.
In a later publication Thomas (1982a) introduced the term 'minimum'
daylength as that below which the plant ceased vegetative growth and formed a
dormant terminal bud while he retained 'critical' daylength to describe that
daylength beyond which flowering would not be induced. Any daylength
between these two would induce flowering but, if it was close to the minimum,
initiation would be rapid with few flowers being produced. Increasing day-
length resulted in slower initiation but more abundant flowering so maximum
cone production was achieved with daylengths slightly shorter than the critical.
Thomas and Schwabe (1970) described how the diameter of the apical buds
of plants in the field showed a marked decrease in diameter at about the time
that they became ripe to flower. That this was not, however, a morphological
indication of ripeness to flower was demonstrated by keeping other plants
under long day conditions well past the ripe to flower stage and in these the
reduction in diameter did not occur until they were transferred to short days.
They concluded that the sudden change in diameter of the meristems indicated
the commencement of flower initiation.
Bhat et al. (1978) reported that, in Kashmir, early trained hops had two
flushes of flowers; one before the longest day and one after. This would be
accounted for by the fact that at such latitudes the daylength when such plants
became ripe to flower was short enough to initiate flowering but this was then
suppressed by the lengthening days and was resumed as the days again became
short enough. Under Kashmiri conditions, with the hops grown on high
wirework, it would not be feasible to harvest the two crops as the bines would
have to be taken down to do so. In China hops are grown on wirework only 2 m
high (Chapter 3) and it has been reported that in the southern regions two crops
are harvested in the year.
The importance of growing cultivars with daylength requirements suited to
the locality was demonstrated in an international variety trial conducted by the
18 Botany
Scientific Commission of the International Hop Growers Convention (Neve,
1983). In this trial three cultivars with a common female parent that had been
bred and selected in England, the Federal Republic of Germany or Yugoslavia,
at latitudes of 51°,48° and 46° N respectively, were all grown in those three
countries and also in France at 47°.
All three cultivars flowered earliest at the lower latitudes, the difference
between Yugoslavia and England being 10-14 days. These differences were
reflected in the yield figures. The English and Yugoslav cultivars both showed
a steady reduction in yield as the sites became more remote from their place of
origin. The results for the German variety were more erratic but the yield was
lowest when grown in England.
Hop varieties differ not only in the critical daylength for flowering but also
in their minimum daylength requirement for vegetative growth. Neve (1961)
described how some seedling plants, on reaching the four-leaf stage, turned
dark green, ceased to elongate and developed a swollen hypocotyl. They
remained in this condition for a variable length of time but eventually
elongation recommenced as the daylength increased. It was also noted (but not
published) that plants overwintered in the glasshouse sometimes assumed a
similar appearance as a result of commencing growth earlier than they would
have done outside and that the cultivar Saaz was especially liable to do so.
Thomas (1968) attributed stunted and slow growth of OT48 (Bramling Cross)
in the field to warm conditions in early spring inducing the plants to emerge at a
time when the daylength was less t)1an the minimum required by this cultivar.
The flowering dates of male plants are much more variable from season to
season than are those of females and Thomas (personal communication) has
suggested that males might be daylength neutral. An experiment by Brooks
(1963), however, suggests that their basic response to daylength is similar to
females'. Early-, medium- and late-maturing males were trained on different
dates and, whereas the period from training to flowering of the early clones
was little affected by training date, the late clones did not flower until late in
the season whenever they were trained. This can be explained on the basis that,
whereas the early clones had a long critical daylength and could initiate flowers
as soon as they were ripe to flower, the late clones had a short critical
daylength so flowering could only occur late in the season when the daylength
had shortened.

1.7 DORMANCY
At the end of the growing season hop plants and their seeds both enter a
dormant phase which has to be broken before growth can recommence for the
next season.
Williams and Weston (1958) reviewed the literature on conditions affecting
the germination of hop seeds. Raum (1929), Smith (1939) and Holubinsky
Dormancy 19
(1941) had shown that some six weeks exposure of moist seed to low
temperatures (ranging from -12°to 5°C) were required for good germination
to be achieved. Bressman (1931) had increased germination by scarifying the
seed but Keller (1953) failed to do so. Paine (1951) obtained rapid germination
by chipping the hard testa. Keller and Paine failed to improve germination
by chemical treatments. Williams and Weston's own experiments confirmed
these findings although they showed that chemical treatment with concen-
trated sulphuric acid for 9 minutes gave a moderate increase in germination,
apparently having a similar effect to scarification, but that it caused damage to
the root tips. Other chemical treatments (gibberellic acid, malic acid,
potassium nitrate, hydrogen sulphide, bis-cyclohexanoneoxalyldihydrazone,
dimethylglyoxime and ethylenediamine tetracetic acid) were ineffective, while
thiorea had only a very slight effect. The lack of activity may have been
because the chemicals were unable to penetrate the seed coat.
Haunold and Zimmermann (1974) described their techniques for collecting
pollen, making crosses and germinating the resulting seeds which included a
period of 6-8 weeks under moist conditions in a refrigerator at 2-3° C prior to
sowing.
Williams et af. (1961) reported that the dormancy of established plants
involves two major stages. The first is the 'onset of dormancy' which is
complete when the plant will not make new growth even when given good
growing conditions. The second stage, the 'breaking of dormancy', involves the
gradual removal of growth inhibition and is considered complete when the
resting buds will break into new growth when climatic conditions permit.
The onset of dormancy is initiated by changes in the daylength and involves
the death of the shoots and the finer root system, the transfer and accumula-
tion of food reserves in the storage roots and the development of relatively
large resting buds on the perennating shoot system below soil level.
Experimental plants grown in a warm glasshouse during the winter under
artificially extended daylength continued vegetative elongation and developed
large fibrous root systems containing little storage material. Plants under the
same conditions but exposed to the naturally shortening days of autumn
ceased all vegetative growth by early September. The leaves remained viable
until late October or early November when the shoot system died completely to
near soil level. During this period much storage material, mainly starch,
accumulated in the roots which increased in thickness. The fibrous roots died
and large dormant buds developed.
By exposing other groups of plants to different temperatures for varying
periods before moving them to a growth room at 20°C, it was found that, even
in a mild winter, exposure to outdoor temperatures for the whole of November
and December was sufficient to break dormancy completely. Plants that had
been exposed outdoors only until 2nd December remained dormant. One of
the three plants moved to the growth room on 14th December and all three
20 Botany
Ib lacre

4000 .......... Main bine and leaves

_._._ Main bine and leaves +
laterals and leaves
- - Main bine and leaves +
laterals and leaves + cones
3000

2000
,.,.

1000 ./
800
600 /
400
..'
200
Ob=r=~==~--~--~~--~
Feb Mar Apr May June July Aug

Figure 1.11 Dry matter accumulation of above ground portion of the hop plant Ibl acre
(Thomas, 1967).

moved on 20th December made some growth but this stopped when the shoots
were between 20-40 cm long.
Other plants were placed in a refrigerator at 3°C at the end of October and
samples removed at weekly intervals, potted up and placed in the growth
room. For the first five weeks these plants made no growth but those moved on
the sixth and subsequent weeks all grew strongly.
Hops grown in Kenya where there is no exposure to low temperatures have
been found to break dormancy very erratically. Under artificially extended
daylength some plants grew vigorously, others made only very limited growth,
just like Williams' experimental plants that had insufficient cold treatment,
while some made no growth at all (Neve, unpublished). Owino (personal
communication) considered that exposure to drought conditions helped to
break dormancy.
Thomas (1965) reported that dormancy could also be broken by treatment
with gibberellic acid (growth occurring within 14 days of treatment) but that
the shoots were often thin and spindly. Attempts by Owino to overcome the
lack of low temperatures in Kenya by gibberellic acid treatment have been
unsuccessful.
Photosynthesis and carbohydrate reserves 21

300
Mature
plants

200
OJ
.....
.!:
OJ
Qj
3: 100 / / Young plants

O~--~--~--~--=---~--~-·=------~"--------~--~---~·~/~/~--~~--~--
Feb Mar Apr May June July Aug Sept Oct Nov Dec Jan

Figure 1.12 Dry weight of rootstock throughout the year (Thomas, 1967).

1.8 PHOTOSYNTHESIS AND CARBOHYDRATE RESERVES

Williams coupled his work on dormancy with studies on the changes in
carbohydrate balance. He and his colleagues published their work on young
plants (Williams and Weston, 1959; Williams, 1960; Williams et al., 1961;
Williams, 1962) while his subsequent work on mature plants was published
after his death by Thomas (1967). The results for young and old plants of the
cultivar Fuggle were similar in that the carbohydrate in the rootstocks fell from
a high level at the end of the winter to a very low level in May. It then
commenced to rise in June and July with a more rapid rise from August to
mid-October. During the winter months much of the starch component was
converted to soluble sugars.
These changes were first associated with the rapid growth in the spring when
the plants produce long shoots with little leaf area and are dependent on the
reserves in the rootstock. Once the leaves expand, carbohydrates are produced
in excess of that required for growth and accumulate in the rootstock, most
rapidly during the shortening days of August and September when extension
growth has ceased. Although this pattern was the same for both young and old
plants, the percentage of starch in old rootstocks during the winter period was
considerably lower than young plants, presumably because of the greater
accumulation of fibrous material in the mature plants.
Changes throughout the year in the dry weight of the above-ground portion
of mature plants are illustrated in Figure 1.11, of the rootstock in Figure 1.12,
while Figure 1.13 shows the total starch and soluble sugars contents of mature
plants and the total starch of young plants (Thomas, 1967).
 22 Botany

50

-....
40

........ ..--.

01
30 ,!

.··
..... ,
.r:. I
i
01 i
'ij 20 i
~ , .i
:' / ........ ,._ ...:_..J

10 !
';_""''/
,...-__ .-'"/
I- T otal starch } Mature plants
--._.. Total soluble sugars
I,' •••••• Total starch-young plants
...... ~...........
....... _::=~::':.~-::':'..:::..f.~.----'
O~--~==~~~--~--~~--____--__~--
Feb Mar Apr May June July Aug Sept Oct Nov Dec Jan

Figure 1.13 Total starch and soluble sugars in the rootstock throughout the year
(Thomas, 1967).

From Figure 1.11 it can be seen that total dry matter has accumulated in
three phases. Up to the end of June production consisted largely of the main
bines and leaves with a total dry weight of about 125 kg/ha. During July
growth consisted almost entirely of laterals and their leaves which contributed
a further 125 kg to the total. Cones commenced to develop in the third week of
July and by the end of August their dry matter amounted to some 250 kg so
that they comprised approximately 50% of the total above-ground growth.
Peat and Thomas (1974) measured the carbon dioxide exchange of develop-
ing cones and found that, although older cones showed a clear response to light
intensity indicating photosynthetic activity, this was negligible in young cones.
Stomata in the young leaves of hops are covered by a continuous cuticular
membrane which eventually ruptures to form the stomatal pore (Royle and
Thomas, 1971b). Such a membrane would permit only very low levels of
gaseous diffusion and it is probable that a similar development in the stomata
of hop cones accounts for the sudden increase in photosynthetic activity as the
bracts and bracteoles commence expanding rapidly. Although older cones
were photosynthetically active, this rarely exceeded respiratory loss so made
little contribution to their growth.
Kenny and Rohrbach (personal communication) found that leaves more
than 75% expanded had maximum photosynthetic potential but that those
shaded by the canopy achieved very low rates. They also found that different
genotypes varied in their photosynthetic rates.
Photosynthesis and carbohydrate reserves 23
The carbohydrate reserves of the plant normally accumulate in the rootstock
but a feature of hop bines is that any portion of them placed in the dark
becomes a storage organ. If the base of a bine is earthed up, or laid along the
ground and covered with soil it will accumulate food reserves, thickening up as
it does so, while the buds also enlarge. Instead of dying at the end of the
season, this swollen tissue becomes perennial and it can be planted the
following season. This is the basis of some methods of propagation (Section
3.l0). Even sections of the bine at a considerable height above ground can be
enclosed in black paper and will behave in this fashion although sections of
bine above and below will not be affected.
CHAPTER 2

The cultivated hop

2.1 HISTORY
An early record of the use of hops in brewing is to be found in the Finnish saga
The Kalevala (Lonnrot, 1963) in which the Beer Lay begins as follows:
The origin of beer is barley, of the superior drink the hop plant,
though that is not produced without water or a good hot fire.
The hop, son of Remunen, was stuck in the ground when little,
was plowed into the ground like a serpent, was thrown away like a stinging nettle
to the side of a Kaleva spring, to the edge of an Osmo field.
Then the young seedling came up, a slender green shoot came up;
it went up into a little tree, climbed to the crown.
The father of good fortune sowed barley at the end of the newly cultivated Osmo field;
the barley grew beautifully, rose up finely
at the end of the newly cultivated Osmo field, on the clearing of a Kaleva descendant.
A little time passed. Now the hop vine cried out from the tree,
the barley spoke from the end of the field, the water from the Kaleva spring:
'When will we be joined together, when to one another?
Life alone is dreary; it is nicer with two or three.'
An Osmo descendant, a brewer of beer, a maiden, maker of table beers,
took some grains of barley, six grains of barley,
seven hop pods, eight dippers of water;
then she put a pot on the fire, brought the liquor to the boil. .. '.
This saga, although it is reputed to go back some 3000 years, was not written
down until the 19th century and as it was handed down orally, could have been
subjected to considerable alteration or addition. It is, therefore, impossible to
judge how old this reference to hops really is.
The earliest written evidence of hop cultivation appears to be that concern-
ing the hop garden of a Wendish prisoner near Geisenfeld in the Hallertau
district of Germany, in 736 AD (Linke and Rebl, 1950). The Wends were Slavs
and it is thought that the Slav word for hops, 'hmelj', may have a Finnish
origin hence the reference in the Kalevala may genuinely indicate that their use
originated there. There is further documentary evidence from the 9th-12th
centuries for hop cultivation in Bohemia, Slovenia and Bavaria so there seems
to be little doubt that this area was the centre from which the practice spread to
the rest of Europe and eventually to the rest of the world. Fric (1985) states
26 The cultivated hop
that 'In a list of goods exported from Bohemia as compiled in the year 1101 we
can find also hops. They used to be shipped to Hamburg where they were
appraised on the famous Forum humuli by specially trained experts'. Fric also
states that hop growing declined in Czech countries during the Thirty Years'
War and that hop cuttings were used for planting expanding areas in
Brandenburg, Silesia, Bavaria, Styria, Baden, Russia and elsewhere. With
major concentrations of production in Bavaria and Bohemian Czechoslovakia
today it is still one of the most important hop growing regions.
A detailed account of hop growing in Eastern Europe, including its
historical background, is given by Strausz (1969). In the Ukraine, Russia, there
is an old tradition of hop usage for brewing which may even predate that of
Bohemia. Ivan the Terrible passed laws that favoured vodka production over,
beer - in contrast to the current policy there - and brewing did not develop
commercially until the 19th century when both brewing and hop production
were, to a large extent, in German hands. Although originally local hop
varieties had been grown, these were unpopular with the German brewers and
so better quality but lower yielding foreign cultivars were grown almost
exclusively.
In Poland there are very early records of hop growing which expanded
considerably in the 18th century. In Yugoslavia, too, there are definite
references to hops being cultivated from about 1160 but again it did not
expand into a significant commercial operation until after 1870.
From central Europe hop growing spread westwards to Flanders during the
14th century where their cultivation was associated with a local specialization
in hopped beer as opposed to unhopped ale.
Hops were not cultivated on a commercial scale in England until about 1524
when Flemish planters were brought over to introduce their cultural tech-
niques. This was at a time when the enclosure of common lands was making it
more feasible to grow such a crop (Burgess, 1964), but there is some evidence
that they were grown on a small scale earlier than that. Baker (1976) believes
that hops were grown before the Norman conquest (1066) and refers to Anglo
Saxon deeds from Himbleton 'Hymel-tun' in Worcestershire which indicates a
hop yard. He also quotes Harrison (1577) as evidence that the 16th century
plantings were a revival of hop growing that had lapsed. 'Hops in the past were
plentiful in this land: afterwards also their maintenance did cease: and now
being revived, where are anie better to be found'.
A so-far unexplained discovery is that of the Graveney boat in England.
Graveney is on the north Kent coast and drainage work there in 1970
uncovered the remains of a boat that has been carbon-dated to about 950 AD.
The abundant hop remains associated with this boat indicate that these must
have been the cargo that it was carrying. There is little evidence that hops were
in general use in England at this time although Wilson (1975) refers to
documents in Canterbury Cathedral suggesting that they were in use in Kent
History 27
in the 11th century while Parker (1934) quotes another 11th century document
from Westminster Abbey that refers to hopis de brassio. Baker (1976) quotes
from the Anglo Saxon Herbarium of Apuleius that from the hop 'hymele' was
produced a 'wort' which was 'that laudable that men mix it with their usual
drinks'. It is possible, therefore, that the hop cargo in the Graveney boat was
destined for this country but an alternative explanation could be that the boat
was heading for a continental destination and was blown off-course.
Several authors have questioned whether the hop is indigenous to Britain
but the discovery of hop fruits in an archaeological dig at Shippea Hill in
Cambridgeshire, dated to some 3000 BC (Clarke and Godwin, 1962) and
pollen residues from other sites show conclusively that it is a native species.
From Europe the cultivated hop was introduced into North America by the
Massachusetts Company in 1629 and was grown for domestic use in New
Netherlands and Virginia in the mid-17th century. It did not become an
important field crop until the early 19th century, the first commercial hop
yard being established in New York in about 1808 (Burgess, 1964). Produc-
tion was centred in New England until problems with disease led to the
industry moving to the west coast where the drier conditions were more
favourable.
Settlers were also responsible for the introduction of hops into newly
developed countries such as South Africa, Australia and New Zealand. A
detailed account of their introduction to Australia is given by Pearce (1976).
The early governors of New South Wales were most anxious that a brewing
industry should be developed because they felt that beer would be a better and
more wholesome drink than the rum which the workforce of convicts seemed
to need to enable them to complete even the simplest of tasks. The first attempt
to transport hop plants from England, organized by Sir Joseph Banks, was
made in 1799 but they failed to survive the voyage because they were 'so
constantly exposed, from the smallness of the ship, to the washing of the sea,
and other circumstances that tended so much to their hurt'. It appears that the
first plants to be grown in Australia were raised from seed in New South Wales
in 1803 and 1804 while the first positive record of vegetative material arriving
from England is that brought by William Shoobridge, 'an experienced hop
cultivator' who left Maidstone in Kent to settle in Tasmania in 1822. There
may have been earlier introductions as a result of urgent requests for material,
but there are no records of any having arrived.
The variety of hops brought by Shoo bridge is not mentioned but there are
accounts of subsequent introductions of Canterbury Goldings and Early White
Grapes and of 'new kinds of hops'. An intriguing question about such
introductions is whether male plants would have been included with them,
either deliberately or accidentally. There are plenty of male plants in Australia
but these could all have been derived from seedling material and not
introduced as imported plants or cuttings.
28 The cultivated hop
Although the first attempts at hop growing were made in New South Wales
these were never very successful, possibly because of the daylength at latitude
34° S; it was in Tasmania that production was first really successful, to be
followed rather later by Victoria.
The growing popularity of beer worldwide led to the further extension of
hop cultivation. In Japan hops were introduced to Hokkaido in the 1880s and
regular cultivation was begun at Nagano during the 1914-18 war when imports
were stopped (Ono, 1959). Hops are said to have been first introduced into
China from Japan in 1921 (Zheng and Pen, 1987) although Simmonds (1877)
refers to a shipment of 1331b from Chefoo in 1874. However, Neve
(unpublished) was once sent a sample of Chinese 'hops' that consisted of seed
of an unidentified plant so it is possible there was a similar confusion over
nomenclature in Simmonds' reference also. Chinese hop production was on a
very small scale until 1949 when the industry spread into Xinjiang, Gansu,
Neimeng, Beijing, Shanghai, Jiangxi and Zhejiang. The enterprise was
abandoned in the last three areas because low yields resulted from heavy
damage by moulds and downy mildew. Since these areas lie between latitudes
27° and 31° it is likely that short daylength was also a major factor.
Problems with availability of foreign currency to buy hops from the
traditional producing countries have led to attempts being made, with varying
success, to extend production to other areas. In India, a large number of hop
cuttings were taken to Kashmir in the 1880s but although the first reports of
the experiment were encouraging (Simmonds, 1877) for some reason the crop
did not become established there at that time. More recent efforts have been
much more successful and India is now largely self-sufficient.
The areas where hops can be successfully grown are limited by the
photoperiodic requirements of the plants as described in Chapter 1. The
lowest latitudes at which they are produced commercially at present are
approximately 34°S in Cape Province, South Africa and 34°N in Kashmir,
India. However, even here there are problems because the daylength is too
short for optimum yields and in South Africa electric lights are being used
commercially to extend the daylength artificially while in Kashmir training
has to be delayed to ensure that growth takes place during the period of
longer daylength.
In Mexico there are reports of success in breeding varieties that are adapted
to latitudes as low as 25°N. These have been described as daylength neutral but
since good yields depend upon the inhibition of flowering while the plants
make sufficient vegetative growth to bear the crop it is more likely that these
plants merely have a shorter critical daylength.
There is interest in hop production at even lower latitudes and it is known
that trials are under way in Colombia, Kenya, Zimbabwe and Burma. Earlier
attempts at such latitudes have failed and it is now known that this is largely
due to the photoperiodic requirements of the plant. These are now being met
Use 29
by using artificial light to extend the natural daylength. There is, however, a
further problem as most of these areas lack a sufficiently cold dormant season
for normal vernalization to occur and it is difficult to simulate this artificially.

2.2 USE
Although hops are today used almost exclusively for brewing, they were first
taken into cultivation for their herbal and medicinal properties. Gerarde
(1636), in his Herball says that:
The buds or first sprouts which come forth in the Spring are used to be eaten in sallads;
yet are they, as Pliny saith, more toothsome than nourishing, for they yeeld but very
small nourishment.
The floures make bread light, and the lumpe to be sooner and easilier leavened, if the
meale be tempered with liquor wherein they have been boiled.
The manifold vertues of Hops do manifest argue the wholesomenesse of beere above
ale; for the hops make it a physicall drinke to keepe the body in health, than an ordinary
drinke for the quenching of our thirst.

The original distinction between unhopped ale and hopped beer should be
noted; a distinction that has disappeared since no unhopped ales are now
produced.
Culpeper (1653) referred to many virtues:
It is under the dominion of Mars. This, in physical operations, is to open obstructions
of the liver and spleen, to cleanse the blood, to loosen the belly, to cleanse the reins from
gravel, and provoke urine. . .. In cleansing the blood they help to cure the French
diseases, and all manner of scabs, itch and other breakings-out of the body; as also all
tetters, ringworms and spreading sores, the morphew and all discolouring of the skin.
The decoction of the flowers and hops, do help to expel poison that anyone hath drank.
Half a dram of the seed in powder taken in drink, kills worms in the body, brings down
women's courses, and expels urine. A syrup made of the juice and sugar, cures the
yellow jaundice, eases the head-ache that comes of heat, and tempers the heat of the
liver and stomach, and is profitably given in long and hot agues that rise in choler and
blood . . . By all these testimonies beer appears to be better than ale.

In Germany, von M1thren (1701) wrote as follows (quoted by Gross, 1900):

The principal use of hops is for making beer, in which it acts as a saline or aromatic; if,
however, too much is used, the beer is too bitter and affects the head. Young hop shoots
taken with the food purify the blood, heal the itch, and relieve the liver and spleen.
Distilled hop extract cleanses the blood from all impurities, tumours and flatulence, and
cures skin diseases and other complaints if taken in regular morning doses of 4 to 5
lothe [the loth = ~oz.].

A somewhat more recent American writer (Mrs Longshore-Potts, 1887) says

that, following childbirth, 'If 'after pains' become an annoyance, the appli-
cation of hot flannel, or steamed hops over the abdomen will be found
beneficial, or a draught of warm hop tea.'
30 The cultivated hop
Other parts of the plant have been used in various ways. The bines contain
fibres similar to those of its close relative hemp and these have been used to
make cloth, sacking or paper while an extract of the leaves or bines was used to
'dye woollens a fine cinnamon brown'. The bines and leaves have also been
used for 'tanning light skins instead of oak bark' (Lance, 1838). The possibility
of using hops as a rather uninteresting, but very permanent, dye will not
surprise anyone who has had their clothes disfigured by contact with hop sap!
Gerarde's uses for hops have slightly more relevance to the present day than
do Culpeper's since hop shoots are still eaten and hop cones are used in some
countries for bread making. In England it is suggested that the young green
shoots can be eaten rather like asparagus but it is not a common dish. In
Belgium, however, the blanched underground shoots are regarded as a delicac~
and there has been a recent report of a technique for growing hop plants
specifically to produce such shoots over a much longer period of the year
(Maton, 1986).
The recipes that are to be found for using hops in bread making do not
indicate the role that they play but it is probable that they help to keep the
yeast cultures free from contamination. Davis (1956) suggests that their use in
bread making may have preceded their use in brewing.
Howard (1964) states that 'They were apparently not introduced into
England until about AD 1400, and the flavour they gave was sufficiently
unpopular to delay complete acceptance until as late as the nineteenth century.
Originally they were used by the brewer not because their flavour was liked but
because they protected his product against bacterial spoilage'. Reynolde Scot
(1576), however, held a different view about their flavour for he says:
For if your Ale may endure fortnight, your beere through the benefite of the Hoppe shall
continue a Moneth, and what grace it yieldeth to the taste, all men maye judge that have
sense in their mouths, and if the controversie be betwixt Beere and Ale, which of them
two shall have ye place of preheminence: it sufficeth for the glorie and commendation of
the Beere, that here in our owne countrie, Ale giveth place unto it, and that most part of
our Countrymen doe abhore and abandon Ale, as a lothsome drincke . . . .
There had long been a liking for beers containing bittering or flavouring
substances as indicated by de Candolle (1884):
In the Middle Ages . . . the Kelts, the Germans, other people of the north and even of
the south who had the vine, made beer either of barley or of other fermented grain
adding in certain cases different vegetable substances - the bark of oak or of the
tamarisk, for instance, or the fruits of Myrica gale.
The substitution of hops for the various mixtures of herbs (sometimes called
gruit) would have provided the preservative action which the other substances
lacked although it is said that Myrica gale is still used on some farms in
Scandinavia. It contains resins similar to those of the hop, but with methyl
groups instead of isoprenyl groups, and having some preservative value
(Malterud and Faegri, 1982).
Use 31
One minor, present-day herbal use for hops is in hop pillows, in which they
are included for their reputed soporific effect.
Whatever their original purpose, modern brewing techniques rely so heavily
upon hygiene and pasteurization to produce a sterile product that the
preservative function of the hops is now of little importance and they are used
for the bitter taste and hop flavour that they impart. These are derived from
the resins and essential oils that are produced in the lupulin glands of the hop
cone. The chemistry of these compounds has been recently summarized
(Hough et 01., 1982).
The traditional method of assessing the quality of hop samples is to examine
them for visual defects, to rub them in the hands when the degree of stickiness
gives a rough estimate of the amount of resin, and to smell them to evaluate
their aroma. There are many brewers and members of the hop trade who are
very skilled in this but the value of such organoleptic assessment is probably
limited. The essential oil components that predominate in a sample rubbed in
the hand will be the most volatile ones and it is these that are most rapidly lost
when hops are added to boiling wort, and even the fraction that is not lost is
likely to undergo chemical change.
Accumulated experience doubtless makes it possible to examine a sample of
an established cultivar and determine the intensity of aroma and whether there
are any abnormalities. Such a hop will only have become an established variety
if it has been found in practice to produce good beers. For the reasons given
above, however, it must be doubted whether it is reliable to predict the
suitability of new varieties for brewing simply by smelling them. Trials have
shown, in fact, that experimental hops with quite unpleasant aromas have
made very satisfactory beers. In spite of the limitations of the technique, simply
smelling the cones has the advantage for the hop breeder in that it is a very
rapid way of assessing the large numbers of seedling varieties from which to
select. It is unlikely that brewers would be interested in using a variety that had
an unusual aroma even if it did prove satisfactory in brewing trials, so it is
logical to discard them at an early stage on that basis.
Gas chromatographic analysis provides a less subjective method of examin-
ing the composition of the essential oils and it will give similar results for
samples of the same cultivar even when these have been grown under widely
different conditions. Figure 2.1 shows gas chromatograms of three different
cultivars and it is apparent that there are major differences between them.
There is some variation between samples of the same variety depending upon
the stage of ripeness at which the hops were picked, particularly in the height of
the myrcene peak as this is synthesized most rapidly as the crop nears maturity.
Nevertheless, by comparing the ratios of the other major peaks it is generally
possible to identify the cultivar from which the oil sample was obtained
(Green, 1986).
Traditionally hops are boiled with wort for 1~-2 hours during which
32 The cultivated hop

Figure 2.1 Gas chromatographs of the essential oils of: a) Rersbruck; b) Pride of
Ringwood; c) Saaz (lRR, Wye).
Use 33
time some of the resins go into solution to provide the bittering principles of
beer but the bulk of the essential oil constituents are evaporated. To overcome
this loss of essential oil flavour some brewers reserve a portion of choice
'aroma' hops and add them late in the boil, 15-20 minutes before copper
casting. British brewers may add 'dry' hops to unconditioned beer, either in
cask or tank, to introduce more essential oil components into the beer. The
flavours introduced by 'late' hops or by 'dry' hops can be distinguished and
either can be reproduced by using appropriate fractions of the essential oil.
Hops added at the start of the boil, rich in resin but with a poorer 'aroma', are
sometime referred to as 'alpha' or 'bitter' hops, but there is no clear distinction
between these and 'aroma' hops.

2.2.1 Hop resins

The hop resins are peculiar to the hop and have not been found in any other
plant species. The lupulin glands consist largely of a mixture of soft and hard
resins. From the brewing point of view it is the soft resins, soluble in hexane,
that are important. These consist of the a-acids, ,B-acids and the so-called
uncharacterized soft resins. It is the a-acids that are the precursors of the bitter
principles of beer, the iso-a-acids. The a-acids are a mixture of analogues,
humulone (1a), cohumulone (1b) and adhumulone (1e) which differ only in the
nature of the acyl side chain R. The ,B-acids are a similar mixture of analogues,
lupulone (2a), colupulone (2b) and adlupulone (2e), as are the deoxyhumul-
ones (3), the probable biological precursors of the a- and ,B-acids. Both the a-
and ,B-acids can exhibit tautomerism and exist as a mixture of readily
interconvertible structures. Those given are thought to represent the principal
tautomers.
Neither the a-acids nor the ,B-acids are very soluble in water. At 25°C the
solubility of humulone is 6 mg/ I and lupulone 1.5 mg/1. They are more soluble
at the boiling point but any extra that is dissolved will precipitate out on
cooling. However, during wort boiling, the a-acids are isomerized into the iso-
a-acids which are much more soluble. The ,B-acids, being much less soluble, are
largely unchanged during wort boiling. The iso-a-acids now have two chiral
centres and so exist as a mixture of cis-(4) and trans-(5) isomers. The bittering
principles of beer thus consist of at least six compounds: cis- and trans-
isohumulone, cis- and trans-isocohumulone and cis- and trans-isoadhumulone.
The overall level in beer is 20-50 mgjl. Cis- and trans-isohumulone have
similar bitterness and it is thought that the other analogues have similar
intensity. Further hydrolysis of the iso-a-acids gives trans-humulinic acids (6),
which lack bitterness, but this transformation occurs to a negligible extent in
the brewers' copper. The proportion of cohumulone in the a-acids can vary
from about 20%-50%. At one time it was thought that cohumulone was better
utilized (converted into isocohumulone) than the other a-acids and that a high
34 The cultivated hop

OH

3 2
Deoxyhumulones a-Acids !'I-Acids
R = CH2CHMe2 (a) (a) Humulone Lupulone
R = CHMe2 (b) (b) Cohumulone Colupulone
R = CHMeCH 2CH 3 (c) (c) Adhumulone Adlupulone

~ A...-~\JuCOR
o

""'= ~ COR
HO I OH HO- -(--i'OH [OJ
I
CO CO

). ~
5 4
trans Iso-a-acids cis

\ H 0
J
~"'~COR
HO.+-!lOH
I
I
H
6
Trans-humulinic acid 7
Hulupone

Figure 2.2 Hop resins and their transformation products.

proportion of it was, therefore, desirable. This was subsequently questioned

and it was later claimed that cohumulone produced less acceptable beers than
the other analogues (Rigby, 1972) but this is not everywhere accepted (Laws et
al., 1976).
Use 35
The ability of hops to give a bitter taste to the beer is their principal function
and many hops are bought solely for this purpose. Their a-acid content is then
of prime importance and this can vary from as low as 3% in traditional
cultivars to as much as 14% in the newer sorts now in production while even
higher levels are to be expected from breeding programmes now under way. It
is now common practice for the contract price for such 'bitter' hops to be based
on the weight of a-acid they contain rather than the weight of the hops
themselves. Accurate analysis of the a-acid content is, therefore, commercially
important. Since different methods of analysis frequently give different results
it is important that the method used should be specified.
Three properties of the a-acids are commonly used for their estimation: (a)
lead salt formation; (b) ultraviolet light absorption; and (c) optical rotation.
The J3-acids do not form insoluble lead salts, do not rotate polarized light and
absorb ultraviolet light at different wavelengths from the a-acids. When a 4%
solution of lead acetate in methanol is added to a methanolic solution
containing a-acids, a yellow salt is precipitated. Originally this was weighed
but now it is more usual to follow the electrical conductance of the solution
during the titration as this changes dramatically when excess reagent is present.
Since uncharacterized resins and oxidation products may react with the lead
acetate, the result of the analysis is usually termed the lead conductance value
although with fresh hops the result is very close to the percentage a-acids. This
is the most common method for the estimation of the a-acids in Europe and is
recommended by the European Brewery Convention (EBC) as the basis for
commercial transactions.
In the USA however, the measurement of ultraviolet light absorption is
the method of choice. Most hop resins absorb ultraviolet light at a character-
istic wavelength which also depends on whether the solution is acidic or
basic. In the method recommended by the American Society of Brewing
Chemists (ASBC), measurements are made in alkaline solution at 325 nm
(Amax for the a-acids), 355 nm (Amax for the J3-acids) and 275 nm (Amin for both
a- and J3-acids - background reading) and regression equations applied to
calculate the percentage of both a- and J3-acids. This is the only simple
method to give a value for the percentage of J3-acids. The regression
equations assume that the hop resins are a mixture of pure a- and J3-acids
with a constant background. This assumption may be fairly true for fresh
hops but does not apply when they are old. From the results of this analysis,
Likens et al. (1970) proposed that the ratio of optical density at 275 nm to
that at 325 nm should be used as a hop storage index (HSI). The ratio
increased from 0.24 in fresh hops to 2.5 in completely oxidized lupulin. Over
several seasons the regression equation
%(a+ J3) lost = 110 10g(HSIj 0.25)
was arrived at and used to compare the storage characteristics of many
commercial cultivars.
36 The cultivated hop
Ultraviolet light absorption is the only easy method for estimating the iso-a-
acids in beer. In the internationally agreed method, degassed, acidified beer
(10.0 ml) is extracted with iso-octane (2,2,4-trimethylpentane) (20.0 ml) and,
after centrifugation, the absorbance of the iso-octane layer is read at 275 nm in
a I cm cell against a blank of pure iso-octane. The result is expressed in
Bitterness Units (BU) (= 50 x absorbance). In beers brewed with fresh hops the
bitterness units are almost equiValent to mg/ I iso-a-acids. This analysis would
be interfered with by unisomerized a- and J3-acids but these are not normally
present in beer.
The optical rotation method depends upon the fact that a-acids rotate the
plane of polarized light and can thus be measured by polarimetry. The
difficulty is to prepare a solution of hop resins that is sufficiently transparent to
enable a reading to be taken. The chlorophyll and degradation products must
be removed without removing any a-acid but suitable grades of charcoal are
usually satisfactory. A more serious disadvantage is that on storage, or in the
preparation of hop extracts, the a-acids may racemize and not be detected by
this method although racemic a-acids are still capable of bittering beer.
More detailed analyses are now obtained by high pressure liquid chromato-
graphy (HPLC). Individual analogues can be separated, though with varying
degrees of precision. It is more difficult, for example, to resolve humulone from
adhumulone than to resolve these isomers from cohumulone. In the method
recommended by the Institute of Brewing (1988) four peaks are resolved:
cohumulone, humulone + adhumulone, colupulone and lupulone + adlupulone.
Hops undergo oxidation and degradation during storage, some cultivars
more rapidly than others. Since they are only harvested at one season of the
year, part of each crop needs to be stored for at least 18 months before use
unless supplies are imported from the other hemisphere. To minimize deterio-
ration many brewers either keep their hops in cold store or have them
processed into pellets or extract.
As a result of oxidation the amount of soft resins decrease while the hard
resins (insoluble in hexane but soluble in ether) increase. It is possible that even
more bitter products that are insoluble in ether but soluble in water are formed
but these would be more difficult to detect. Compounds present in the hard
resin of fresh hops include xanthohumol (8), iso-xanthohumol (9) and the
flavone (10) but these have no known brewing value.
Oxidative cleavage of the acyl side-chains of the a- and J3-acids is manifest in
the 'cheesy' aroma, in old hops, of the isobutyric, isovaleric and 2-methylbuta-
nic acids. No oxidation products of the a-acids have been characterized as
bittering agents in beer but oxidation of the J3-acids gives hulupones that are
bitter. When hops are boiled in the copper using traditional brewing methods,
the loss of bitterness of the a-fraction during storage is partly compensated by
the gain from the J3-fraction. The utilization of the J3-acids is probably most
significant when hops are boiled with the wort more than once, as in stout
Use 37

HO =--- OH trH-o OH
: ,. I C"
CH

CH 3 0 II
o
8
Xanthohumol
Iso-xanthohumol

10

Figure 2.3 Constituents of the hard resins of hops.

brewing. In the first boil the molten J3-acids spread over the surface of the hop
cones as a thin layer and are readily oxidized during the air rest between boils.
Procedures have been described for utilizing the J3-acids by these routes with
improved yields in commercial trials (Moir, 1988) but they are not yet widely used.
Whereas newer cultivars have much higher a-acid contents than older sorts,
there has not been a corresponding increase in the J3-acids. Whether this is
simply because there has been no selection for the J3-acids or whether there is
competition between a and 13 for a common precursor as suggested by Likens
et af. (1978) is not clear.
Because of the high proportion of J3-acids: a-acids in the old cultivars the
changes in storage did not greatly reduce the bittering power of the hops when
used in the copper, but with the modern, bitter varieties the proportion of 13-
acids: a-acids is much lower and so the losses of bittering power are greater.
Storage stability has, consequently, become a much more important criterion
for hop breeders when making their selections. It is even more important if the
hops are to be used for extraction when, at present, only the a-acid is required.
Cultivars that store well can be processed economically over a longer period
after harvest thereby enabling the capital cost of the processing equipment to
be better utilized.
It was reported by Warmke and Davidson (1944) that hop scions grafted
onto Cannabis stocks produced cannabinoid resins and this led to interest in
the technique as a means of producing such material while avoiding legal
restrictions (Drake, 1970). An observation that soil applications of carbamate
38 The cultivated hop
insecticides appeared to reduce a-acid levels in treated hop plants also
suggested that rootstocks might have an influence on resin production.
However, trials by Crombie and Crombie (1975) involving Humulus/ Cannabis
grafts failed to show any change in the type of resins produced as a result of
such unions, while Neve (1973) made grafts between hop cultivars with high
and Iowa-acid contents and these too showed no influence of the stock on the
resin production of the scions:

2.2.2 Essential oils

The other important hop product is the essential oil which is also produced in
the lupulin glands and normally represents about 0.5-1.5% of the weight of the
dried cones. Some cultivars contain more than others while seedless hops have
a considerably higher content than seeded cones of the same cultivar and
contents as high as 3-4% have been reported in some situations.
Whereas the hop resins give beer its bitterness, the essential oil gives it aroma
and flavour. Although the chemistry and value of the resins are well
understood, there is still much to be learned about the essential oil which is a
complex mixture of 200 or more components. These components can be
separated by gas chromatography and many of them have been identified but
this has by no means provided a complete understanding of their use in the
brewery.
By definition the essential oil is volatile in steam and is estimated by
steam distillation, the oil being retained in a trap while the condensed
aqueous phase returns to the boiler; a process known as cohobation. In the
Institute of Brewing method, steam distillation is carried out for three hours.
Commercial hop oil, prepared by this process, has been available for many
years and used by some brewers to impart a hop flavour to finished beers, but
most brewers can distinguish between beers that have been late hopped, dry
hopped or those treated with such hop oil. Some changes occur during
cohobation and hop oil preparations obtained by steam distillation under
high vacuum at low temperatures, or by liquid carbon dioxide extraction,
give flavours more akin to those obtained by dry hopping. Liquid CO2
extracts can be fractionated to provide preparations that give mainly late hop
flavour (floral and citrus) or dry hop flavour (resinous, spicy and citrus) to
beers.
It appears that hop flavours are due to synergistic mixtures of compounds
rather than single impact compounds. The essential oils can be divided into
three classes of compounds: (a) hydrocarbons; (b) oxygenated derivatives; and
(c) compounds containing sulphur. In most cultivars the hydrocarbons
predominate but they are the most volatile and few survive wort boiling, even
with late addition, but they will be slightly more soluble in the dilute alcoholic
solution of beer and traces will dissolve during dry hopping.
Use 39
The main hydrocarbons in the essential oil are the monoterpene (C IO)
myrcene (12) and the sesquiterpenes (CIS) caryophyllene (24) and humulene
(23). Other sesquiterpenes, such as farnesene (22) and p-selinene (26) are found
in some cultivars but not in others. The ratios of humulene:caryophyllene and
humulene:selinene appear to be characteristic of cultivars and are useful for
their identification (Green, 1986). The essential oil of the 'aroma' hop,
Hersbrucker, contains several sesquiterpenes not found in other cultivars
(Tressl et ai., 1983).
Current biogenetic theory suggests that the terpene hydrocarbons are
formed from oxygenated intermediates and in the ripening hop oxygenated
compounds are found before the hydrocarbons. Of the hydrocarbons, the
cyclic sesquiterpenes are found before farnesene and myrcene. As the hop
ripens the synthesis of myrcene appears to be the dominant pathway. Myrcene
and other monoterpenoids are thought to be formed from geranyl pyrophos-
phate (11). Many of these intermediates are now thought to contribute to the
hoppy flavour of beers. Figure 2.4 shows the relationships of some of these
compounds, the flavour they produce and their concentrations in beers
(Sharpe, 1988). Another compound in hop oil giving a floral flavour to beer is
undecan-2-one (15 ppb). This compound is not terpenoid in nature and
presumably is formed as a by-product of fatty acid biosynthesis. Smaller
amounts of related methyl ketones such as tridecan-2-one also occur in hop oil.
The sesquiterpenes are formed in a similar manner to the monoterpenes
from farnesyl pyrophosphates (20) but cyclization of the farnesyl cation can
lead to many cyclic structures such as the monocyclic humulene (23), the
bicyclic caryophyllene (24) and the tricyclic aromadendrene (25) found in
Hersbrucker hops (Figure 2.5).
Oxygenated compounds can also be formed from hydrocarbons during
storage (Figure 2.6). Oxygen can add to carbon-carbon double bonds to form
cyclic epoxides. The epoxides of caryophyllene (24) and humulene (23) have
been detected in hop oil and beer. Humulene can form three monoepoxides
(28-30) and each of these, and diepoxides, have been found. Reductive ring
scission of the epoxides gives rise to alcohols such as humulol (31) and
humulenols I (32) and II (33) found in oil and beer. It is possible that epoxides
and ketones in wort could be reduced to alcohols during fermentation. Similar
transformations to those given in Figure 2.5 occur with other sesquiterpenes
but, although many of the oxygenated products have been detected in hop oil
and beer, it is not known, with the possible exception of humulenol (see
below), what flavours they contribute to beer.
By fractionation of a liquid carbon dioxide extract of hop oil, two fractions
were obtained, one rich in ketones such as undecan-2-one and the other rich in
alcohols, especially linalol and humulenol. Both imparted late hop character-
istics to beer. The ketone-rich fraction was the more floral but also contributed
some spicy / mouthfeel and astringent characteristics. The alcohol-rich fraction
40 The cultivated hop

( )

13 11 12
Linalol Geranyl pyrophosphate Myrcene
(Floral: 40 ppb) (Spicy, resinous:
200ppb)

1
2H'OH ~(--- __~~ \CH'O~CH'
16 14
Nerol 15
Geraniol
Geranyl acetate
(Geranium, floral:
(Fruity: 15 ppb)
5ppb)

9: 2
I
~
+
17 18 19
a-Terpineol Limonene Terpinolene
(Woody, resinous: (Citrus, fruity: (Woody, resinous:
5ppb) 80-100ppb) 20ppb)

Figure 2.4 Monoterpenoids in hop oil and beer (after Sharpe, 1988).

was less floral but exhibited more of the spicy / mouthfeel and astringent
characteristics. Both fractions produced a similar degree of grapefruit char-
acter in the beers and contributed to the body (Murray et al., 1987).
The essential oil of hops contains only trace amounts of sulphur compounds
but since these have potent aromas they may influence the overall flavour of
the essential oil and beer. Hops in the field may be treated with elemental
sulphur to control mildew and it used to be a common practice to burn sulphur
 ~
(1)
'"

I..
" ~
~ , r
I ~H h ~
I opp
II opp
~ 21
R 20
~ c;::y 22
Nerolidol Farnesyl pyrophosphate
fJ-Farnesene

/ ~
H

lJ 23
1;ts 24
,~ 25
~ H

Humulene Caryophyllene Aromadendrene 26

fJ-Selinene

Figure 2.5 Sesquiterpenoids in hops.

.j:>..

-
42 The cultivated hop

[OJ

24 27
Caryophyllene Caryophyllene epoxide

/
23
Humulene

V 28

31 33 32
Humulol Humulenol II
HumulenolI
Figure 2.6 Transformation of sesquiterpenes.

in the oast to bleach the hops. Both of these treatments may alter the spectrum
of sulphur compounds in the oil. For example, the sesquiterpenes caryo-
Use 43
phyllene (24) and humulene (23) react with elemental sulphur under mild
conditions to form episulphides (as 27-30 but with sulphur replacing oxygen).
The essential oil contains polysulphides such as CH 3SSSCH3, CH 3SSSSCH 3,
and CH 3SSCH2SCH3, which are formed from S-methylcysteine oxide,
CH 3SOCH2CH(NH2)COOH, during steam distillation. This intermediate is
largely destroyed when sulphur is burnt on the kiln but slowly regenerates
during storage. Thioesters are also present in hops and hop oil (<1000 ppm) and
some of these, such as S-methyl 2-methylbutanethiolate, are introduced into
beer by dry hopping.
Sharpe (1988) describes how hop flavour is added to some commercial beers
by the addition of CO2 extracts post-fermentation and identifies the key
components of the essential oil and their flavour characteristics. Thus, much is
known about the composition of hop oil and its effect upon beer flavour, but
this is still not enough to enable the plant breeder to make selections on this
basis.

2.2.3 Tannins
The tannins are a group of water soluble polyphenolic compounds which react
with proteins in the wort or beer to form insoluble precipitates. Those
precipitated during wort boiling or cooling are known as the hot break and
cold break respectively and are removed by filtration. Some protein and
polyphenol materials still remain, however, and these will continue to react to
produce a haze which can make the beer cloudy and unacceptable. Much of
this haze can be induced to form by conditioning the beer at low temperatures
and filtering it off but, even after this, haze formation can continue in the
finished beer, the shelf life of which is largely determined by how soon the
cloudiness develops.
Normally about 70-80% of the polyphenols in the wort come from the malt
and only 20-30% from the hops. Opinions differ among brewers as to whether
the hop contribution is important and this uncertainty is reflected in the fact
that some of those who use hop extracts require them to contain the water
soluble fraction, that contains the tannins, while others do not.
The most reactive of the polyphenols are the proanthocyanidins. These are
made up of catechin units (Figure 2.7). Procyanidin B-3 (35), for example,
contains two such units while procyanidin C-3 (37) contains three. On
treatment with acid they yield catechin (34) and the pigment cyanidin (36).
Other procyanidins contain additional hydroxyl groups and give delphinidin
etc. on treatment with acid. Recent barley breeding work by Carlsberg has
developed cultivars that are free of these. Brewing with malt from these barleys
produces beers with a considerably extended shelf life and if all the malt is of
this type the proanthocyanidin content of the hops used may become
important. Determinations made by Carlsberg of the amounts present in
44 The cultivated hop

,O
-...:OH

°
H0CX:XO
.& OH

I OH
OH HO~vl'

~H
HOW AOH
.d
HO I I
OOH

HO OH
34
Catechin HO OH
(817-2821 mg/kg)
35
Procyanidin 8-3
(428-1472 mg/kg)

H 0 O :0: X•
O~
I
OH

OH

~ I OH I
OOH
HO
HO ~
ro o.
I 0 ~OH

H0O::X- . " OH
::::,..
HO, OH O
I OH
36
Cyanidin
HO OH
37
Procyanidin C3
(287-875 mg/kg)

Figure 2.7 Flavenoids in hops: Alsace Record, Hallertauer, Tettnang, Saaz (after
Jerumanis, 1985).

various hop cultivars have shown considerable differences (Table 2.1). It is not
only the proanthocyanidin content that is important but also the amount of
a-acid. This is because the weight of hops added would be calculated from the
quantity of a-acid required and so it is the ratio of proanthocyanidin:a-acid
that matters.
The proanthocyanidin-free barleys so far in production are probably not
commercially acceptable for other reasons but these objections are likely to be
overcome. Once such improved barleys are available it can be expected that
some brewers, at least, will be looking for hop cultivars that have low P:a
ratios and this is another factor that hop breeders need to anticipate.
Hop processing 45
TABLE 2.1 Determination of proanthocyanidin content of a number of hop cultivars

Cultivar Proanthocyanidin a-acid Ratio

% % P:a
Yeoman 1.2 5.6 0.2
Target 3.9 12.6 0.3
Northem Brewer 3.1 7.1 0.4
Fuggle 2.9 5.1 0.6
Saaz 5.1 3.5 1.5

Alternatively, brewers could use CO2 extracts which are very low in proan-
thocyanidins.
Jerumanis (1985) describes a method for quantitative analysis of flavanoids
in hops using HPLC and in limited trials found that (+)-catechin and
procyanidin B3 were the most representative followed by procyanidin C2• The
values he found for these three major components in Saaz hops totalled about
5000 mg/ kg (0.5%) which is only one-tenth of the total value recorded for that
cultivar by the Carlsberg workers.

2.3 HOP PROCESSING

Hops pressed into the conventional bales or 'pockets' are bulky to store and
they gradually lose a-acid and aroma through oxidation which can be slowed,
but not eliminated, by cold storage. Various methods of processing hops which
help to overcome these disadvantages have been developed.
The first of these is the use of organic solvents such as hexane, methanol,
methylene chloride or trichloroethylene to extract the brewing materials from
the hops. The solvents are then evaporated and during this process the more
volatile essential oils are lost. The finished extracts are green viscous liquids
containing as much as 50% a-acid.
These extracts are packed in cans which are far less bulky than an equivalent
quantity of baled hops and they are very stable in storage. Although they are
useful as a source of a-acid they have lost a great deal of the essential oils and
some breweries are reluctant to use them in case they contain residues of the
solvent used. The cost of the extraction process is compensated by the
elimination of storage losses and by a small increase in the utilization of a-acid
in the copper.
A process that retains most of the character of the hop cones is to grind them
to a powder which is then compressed into pellets. Packing the pellets in
vacuum or inert gas ensures they have a long shelf life with limited losses in
brewing value. Again they are far less bulky than whole hops and are much
46 The cultivated hop
easier to handle. They are particularly well suited to whirlpool separators, the
use of which has been stimulated by the use of high a-acid hops. With
traditional cultivars sufficient hops are added to the copper for them to form a
filter bed when the wort is run off from the hop back. With high a-acid hops
much smaller quantities are required and the bed is too thin to provide an
effective filter. Whirlpools perform better in this situation and work more
effectively with pellets than 'with whole hops. Systems have also been
developed that enable pellets to be loaded automatically into the copper.
By suitable blending during processing, pellets can be produced with a
standard a-acid content, again making them far easier to use than baled hops
which require an a-acid analysis of each batch so that the bitterness levels of
the beer can be controlled. The a-acid content can be adjusted by blending
hops with different alpha contents, discarding some of the cone material that is
free of lupulin or by sieving out lupulin from one batch of hops to add to
another batch to produce enriched pellets.
In recent years a better method of extracting hops has been developed by
using carbon dioxide as the solvent. One method uses liquid CO2 at 5-15° C at
about 50 atmospheres pressure. Above the critical temperature of 31.1 ° C the
gas will not liquefy at any pressure and the second, supercritical method uses
CO 2 gas at a temperature of 45-50° C and pressures as high as 400 atmos-
pheres. Supercritical CO2 is a more powerful solvent but the liquid process is
more selective and produces an extract containing a- and f3-acids together with
the essential oils but without the hard resins, polyphenols, fats, waxes and
chlorophyll which are found in supercritical extracts (Laws et al., 1977). With
either process, there are no objectionable solvent residue problems.
A further advantage of CO2 extraction is that it is possible to collect separate
fractions that are rich in either the essential oils or the a-acid. Other methods
of collecting the oils waste the a-acid. The latest development has been to
separate further the essential oils into fractions that can be added to beer after
fermentation to give results similar to either late hop addition or dry hopping
(Haley and Peppard, 1983).
Because liquid CO2 extraction produces the purest source of a-acid, it makes
an excellent starting point for the preparation of pre-isomerized extracts. As
already mentioned, the utilization of hops or hop products added to the copper
is low but this can be overcome by pre-isomerizing the a-acids by treatment
with dilute alkali and adding this material to the beer after fermentation. In
this way almost 100% conversion to isohumulone can be achieved and the
losses on yeast are avoided. Small losses do occur in the process but an overall
utilization of about 85% can be achieved. A review of the different types of
processed hops and their characteristics is given by Moir (1988).
The most recent development has been a report from the Brewing Research
Foundation (1988) that extrusion cooking of hops prior to addition to the
copper converts substantial quantities of a-acids into iso-a-acids and that
In vitro production 47
overall utilization of hops for the bittering of beer is greatly enhanced. The
process has been patented.
With any of these methods of processing hops the costs are the same for a
given quantity of hops regardless of their a-acid content. High a-acid cultivars,
however, yield a more valuable product and it is only economic, in general, to
process these. More than 60% of the world hop crop is now processed in one
way or another and this has played a large part in the increased demand for the
new high a-acid selections.
These comments are true for the bulk of the crop that is bought simply for its
bittering value. Some of the crop, however, is more highly valued for the fine
quality of its aroma and such hops command a premium on the market at the
present time. For brewers who demand such cultivars and also want the
benefits of processed hops the costs of processing may be justified but this
represents only a small proportion of the hops that are processed.

2.4 IN VITRO PRODUCTION

Various attempts are at present being made to produce plant products by
means of cell or tissue culture techniques and some work has been done using
these methods for a-acid production from hop cell cultures. Heale et af. (1988)
have reviewed this work which has not, so far, given any positive results. It is
suggested by Robins et af. (1985) that this failure may be due to peroxidase
activity degrading any a-acid that is produced in hop suspension cell cultures.
Support for this suggestion came from the rapid disappearance of exogenous
a':acid that was added to the suspension. The possibility of selecting cell lines
that would not degrade the a-acid has been considered.
The possibility of brewers obtaining their hop material from cell cultures in
rather the same way that they maintain yeast cultures probably has some
attraction. Such a source should be more reliable as it would not be subject to
the vagaries of the weather or market fluctuations but it would first have to be
shown that it was a suitable replacement for whole hops. As a source of a-acid
for pre-isomerization it would presumably be adequate but it might not
contribute all the constituents of the hop, such as the essential oils and tannins,
that are utilized when hops are added to the copper.
There would also be a major difference in scale between maintaining a yeast
culture and producing the quantity of a-acid that is required. In vitro
techniques are only likely to be economically, as opposed to technically, viable
when the product has a very high commercial value and it is doubtful whether
a-acid would fall into this class. The developments already achieved in hop
research, especially in the production of new varieties, have led to a big
reduction in the price of a-acid grown on the farm and further advances can be
expected so it will be increasingly difficult for cell culture to prove competitive.
CHAPTER 3

Production methods

3.1 SUPPORTING STRUCTURES

The hop is a climbing plant and in order to bring it into cultivation it is

necessary to provide a support for it. Originally this was done by providing
each plant with a pole or poles up which it could climb. These were pulled from
the ground at harvest time and stored until the beginning of the following
season. In the earliest description of hop growing in England written by
Reynolde Scot (1576) it is said:
.. .if your hylles be distant three yardes a sunder, provide for every hyll foure Poales, if
you will make your hylles nearer togither, three Poales shall suffice ... Your Poales
maye not be above, xv, or xvi, foote long at the most, except your ground be very riche
or that you have added there unto great labour in raysing up your hylles, or else except
your hylles stande too near togither: if anye of these chance to be, or if all these three
thynges meete in one Garden, the best waye of reformation, is to set the fewer Poales to
a hyll, or to let them remain the longer. Otherwise the hoppes will growe from one Poale
to another, and so overshadowe your Garden, the fault thereof being especially to be
imputed to the nearnesse of the hylles. Therefore chiefly you must measure your Poales
by the goodnesse of youre grounde.

Since the end ofthe 19th century it has become standard practice to provide
a permanent structure of poles and wire to which each year string or thin wire
is attached to provide support for the hop bines. The height of such wirework
structures varies from country to country. In continental Europe it is usually
7-8 m, in England rarely more than 5 m, while in the USA an intermediate
height is normal.
Originally all poles were, of course, wooden but there has, during the past 30
years, been some change towards steel or concrete poles. Concrete has been
quite widely adopted in Eastern Europe because of shortage of suitable fir
poles (Gjurov and Christov, 1963; Kiss, 1965) but it is much more expensive
than wood (Gerber, 1966) and has been rarely used elsewhere.
When low-tensile wire is used with short spans between poles the wires are
strained quite tight so that there is little sag. Systems that use high-tensile wire
with wide spans achieve greater strength by leaving the wires lightly strained so
that they sag in a catenary. Attaching the string or wire supports directly to
50 Production methods

I
\ I

1'1'1'1 ,,",'///
'- ,t

Figure 3.1 Wide span wirework system showing horizontal stringing wire suspended
from the sagged load-bearing wire (after Kamm, 1969).

the catenaries would mean that the bines were of varying length and this would
complicate the harvesting process. A second horizontal wire is therefore
suspended from the catenary by supports of different lengths to hold it level
(Figure 3.1).
In the 1960s there was a series of papers presented at the meetings of the
Technical Commission of the European Hop Growers Convention describing
the new developments in wirework systems and these included calculations of
the various stresses which needed to be considered in designing the systems
developed in the following countries: Gjurov and Christov (1963) for Bulgaria,
Bailey (1963) for England, Kamm (1965 and 1969) for BRD, Kiss (1965) for
Hungary, Gerber (1966) for France, and Thomas (1967) for Belgium.
In England coir string is used for the hops to climb and this is attached at the
bottom to a ground peg at each hill and, at the top, to hooks clamped onto the
top wires. Stringing is carried out from the ground using a continuous length
of string and a pole long enough to reach the top wire (Figure 3.2). In
continental Europe cut lengths of thin, soft iron wire are commonly used and
these are attached to the tpp wire by means of a device called a cuckoo (Figure
3.3) and dibbed into the soil beside the hop hill. In the USA and Australia, cut
lengths of string are tied to the top wire by workers standing on a tractor-
drawn platform and also dibbed into the ground.
In continental Europe the rows of plants were traditionally close together
(1.6 m) with 1.4 m between plants in the row and one wire provided for each
plant. These narrow rows meant that only very narrow tractors and imple-
ments could be used so more recently there has been a change to a distance of
Supporting structures 51

Figure 3.2 English hop stringing (IHR, Wye).

2.8-3.2 m between rows. With a spacing of 1.5 m between hills within the row
each plant is provided with two wires or, if the spacing is halved, then only one
is used.
In England also there has recently been a change to wider spacing. The
traditional layouts were either the umbrella system with 2 m between and
within the rows and four strings to each plant, or the Worcester system with
wider rows (up to 3 m), approximately 1 m within the row and only two strings
per plant. In the USA the spacing has remained fairly constant at about 2.25 m
between and within the rows and two strings per plant.
These variations in spacing result in big differences in the number of strings
(and hence the number of bines) per hectare. The results of experiments in
52 Production methods

Figure 3.3 'Cuckoo' used for attaching training wires to the top wire in Germany.

England at Wye and Rosemaund have been summarized by Thomas (1979).

These trials included wirework of 14 ft, 16 ft and 18 ft (4.3, 4.9 and 5.5 m)
heights, and it was found that Bullion performed best at 18 ft but Fuggle and
Northern Brewer were best at 16 ft. All cultivars produced the highest yield per
plant with the widest spacings but the yield per hectare was greatest with the
closest spacings. In Czechoslovakia, Stranc et al. (1979) obtained the highest
yield per plant at a spacing of 3 m x 1 m and 3 bines up a single string while the
highest yield per hectare was obtained from a 3 m x 0.75 m spacing with 2 bines
on each of two strings per plant.
To calculate which spacing would give the best economic return is quite
complicated since it is necessary to determine the costs that are related to the
number of plants and strings (such as planting, stringing, training and
harvesting costs) and those related to the area of the garden (ground rental,
wirework, manuring and spray materials).
Examination of the growth on the different heights showed that the bending
over, or breaking, of the bines at the top wire stimulated lateral development.
Cultural operations 53
Since this happened earlier on the lower wirework the amount of head
development was greatest at 14 ft and least at 18 ft. The growth on the 18 ft
system was most suitable for machine picking as there were fewer tangled
laterals but the reduced lateral development gave lower yields except for the
very vigorous cultivar Bullion.
It is worth noting that Reynolde Scot had anticipated this finding in 1576
with the following comment:
The hoppe never stocketh kindlye, untill it reache higher than the Poale, and returneth
from it a yarde or two, for whylest it tendeth climbing upwarde, the branches which
growe out of the principall stalke (wherein consisteth the abundance of encrease) growe
little or nothing.

The various heights and spacings in the Wye experiments resulted in the
strings having different angles of slope and yields were highest on the flattest
slopes. In a more detailed study on the effect of slope Thomas (1969) compared
growth on strings inclined at 45°, 65° and 85° from the horizontal. The flatter
the slope, the slower the rate of growth and the shorter the internodes on the
main bine. The highest yields were obtained from the 65° slope as it had the
greatest number of laterals which were also the longest and had the most cones.
A similar study in Germany (Rossbauer and Christl, 1979) indicated that a
slope of 72-8° was best for the cultivars Hallertauer, Hersbrucker, Huller and
Brewer's Gold but that Northern Brewer was best when grown on vertical
strings.
A recent development in the USA has been a system of growing hops on
wirework only 3 m high and without any supporting wires running across the
rows and in England there are plans for a similar system using dwarf cultivars
(see section 3.7).
In China a totally different method of supporting the hops is employed. The
wirework structure is only 2 m high and the top wires are only about 30 cm
apart. The plants are grown in rows 3 m apart with 1 m spacing within the row.
Two bines from each plant are trained up a vertical wire and on reaching the
top they are trained horizontally in opposite directions in such a way that they
and their lateral shoots form a complete canopy. All side shoots are stripped
from the vertical growth, so that the cones are formed entirely upon the
horizontal area with the majority of them on the upper surface but with some
below. Under this system mechanization is practically impossible but this is not
a problem in a country where plenty of hand labour is available.

3.2 CULTURAL OPERA nONS

When establishing a new hop garden it is important that the site is well drained
and that the ground has been deep-ploughed and well cultivated to ensure that
it is clear of perennial weeds. Since the hop plant is perennial and remains in
54 Production methods
the ground for many years, soil cultivations in established gardens are
concerned principally with controlling weeds, incorporating manures or
fertilizers, and controlling the growth of the rootstock.
If left untended, hop hills will spread extensively by means of underground
runners with the result that many shoots emerge in the spring too far from the
strings to be trained satisfactorily. The traditional method of dealing with this
problem was to plough one or two furrows away from the hop row on each
side during the winter and then to dig around the hill by hand in the spring to
expose the rootstocks so that they could be trimmed with a knife, cutting back
the old shoots to below ground level and removing runners and side shoots,
leaving the hills neat and compact. A series of cultivations with tined or disc
harrows during the spring and summer then moved the soil back over the
rootstocks and kept weeds under control. Such cultivations could be quite
deep at the beginning of the season but the depth would be reduced later so as
to avoid damage to the fibrous feeding roots that developed as the season
progressed.
It was also a common practice, half way through the season, to 'earth-up' the
hop hills by using a pair of mould boards to sweep the top soil from the centre
of the alleys over the hop rows. This helped to control weed growth in the rows
while the covered bases of the bines produced additional roots and thickened
up so that they could be cut off in the winter to provide 'strap cuttings' for
further plantings. Every effort is made today to reduce the amount of hand
labour required and various machines have been developed to cut the hills
mechanically, the commonest comprising two overlapping disc coulters. This
operation is not suited to English conditions because of the permanent soil
pegs that are used for holding the strings.

3.2.1 Non-cultivation
The majority of hops in England are now grown under a non-cultivation
system which has also been adopted to some extent in Australia and New
Zealand. Under this system weeds are controlled by herbicides and the hills are
left undisturbed so that a strong root system develops near the surface (Figure
3.4). This greatly reduces the labour requirement and has been found to have
the added advantage that after rain tractors can travel much sooner on
uncultivated than on cultivated ground. Under non-cultivation, cracks develop
in most soils and these provide drainage channels down which surface water
soaks away far quicker than on cultivated ground where such channels are
constantly being destroyed. Some silty soils form a cap after rain which
prevents such drainage, reSUlting in run-off and soil erosion. It has then been
found useful to run a sub-soiler down the middle of the alleys in the winter to
provide an alternative drainage channel.
Figure 3.4 Root system of a) cultivated; and b) non-cultivated hop hills {lRR, Wye).
56 Production methods
Trials have shown that yields are at least as good, and frequently better, under
non-cultivation and as soil cultivations have been shown to be a very important
cause of verticillium wilt disease spreading within an infected garden (Keyworth,
1939). There has been considerable incentive for growers quickly to adopt non-
cultivation in the areas affected by this problem. Although most rapidly adopted
in the wilt-infected areas it is now widely used in all districts.
The frequent soil cultivations of the traditional systems destroyed soil
structure and heavy dressings of bulky organic manures were required to
counteract this. There were fears that the inability to incorporate such manures
under non-cultivation would lead to a breakdown of soil structure but this
does not appear to happen. Since the soil is not disturbed the hop root systems
extend throughout the soil profile and help to provide structure while there are
none of the mechanical operations which are the main cause of damage.
Non-cultivation does, however, have the disadvantages that powdery mil-
dew disease is more difficult to control and that the spread of runners from the
hills can be a problem. In order to control runners a technique was introduced
of planting hop setts in sleeves which, being open at the bottom, allowed the
roots to emerge but checked the runners which develop from the upper part of
the rootstocks (Thomas and Farrar, 1976, 1977). This method was first
observed in Czechoslovakia where rigid pipes were being used but in trials at
Wye these were found to be too restricting and polythene tubing was found to
be much better.
The standard treatment with herbicides in the non-cultivation system
consists of an application of paraquat in the autumn, to burn off any weeds
that may have developed during the latter part of the season, followed by
simazine in the spring to prevent the establishment of new seedlings. This
controls most weeds but alternative herbicides may be required to deal with
such problems as docks (Rumex spp.), perennial nettles (Urtica dioica),
creeping thistle (Cirsium arvense), cleavers (Galium aparine) or couch grass
(Agropyron repens). Difficulties have also started to arise with simazine-
resistant strains of weeds such as groundsel (Senecio vulgaris) that were
originally susceptible.

3.2.2 Training
In general two or three bines are trained up each of the supporting strings or
wires as this produces the optimum density of growth for high yield and good
quality. The bines are trained when they are about 0.5 m long and it is usually
found to be best not to select the most vigorous of all. Under the English
system in which there are four strings to each hill, there may not be the
required 8-12 shoots ofthe right type at the first training and a second round is
then necessary. Established plants eventually produce many more shoots than
are needed and once training is completed it is necessary to remove the surplus.
Cultural operations 57
This can be done by hand by the trainers but more commonly the unwanted
growth is burned off by chemical defoliants. Leaving the excess shoots to form
a dense growth around the base of the plants would create a major disease
problem since the conditions would be ideal for the mildew diseases to develop
there.
Training is one of the most labour-demanding operations and there have
been several attempts to simplify or eliminate it. Various training aids that can
be placed around the hill to direct the shoots towards the strings have been
tried but not found to be economical. The idea of planting the setts in
containers was initiated in Czechoslovakia with the intention of simplifying
training by ensuring that all the shoots emerged close to the supporting wires
and it was found to be effective. In England a series of trials were carried out at
Wye and Rosemaund to see whether the bines could be left to self-train. The
conclusion at Rosemaund was that hand training consistently produced higher
yields than 'assisted self training' (Anon, 1984). The Wye report was more
encouraging though it pointed out that much depended upon the habit of
growth of the cultivar concerned. The upright-growing variety Early Bird when
given assisted self-training, or even complete self-training, gave yields not far
short of hand-trained controls. On the other hand few bines of Wye Challenger
climbed the strings when left to self-train, although assisted self-training was
quite successful (Thomas and Farrar, 1975). Assisted self-training consisted of
initially putting a handful of bines between the strings and later furnishing any
empty strings.
After training, some cultivars will climb the supports less successfully than
others and, especially under windy conditions, many bines may fall away from
the strings or wires, necessitating more hand labour to retrain them. If they can
still be reached from the ground the work is not too difficult but if the trouble
occurs later it is necessary to use a forked stick to reach them or else the
trainers may be driven through the garden on a trailer that is high enough for
them to reach.

3.2.3 Stripping
In addition to removing all the excess shoots it is normal practice to strip the
leaves and laterals from the lowest part of the bines. If left unrestricted these
provide an excellent site for downy and powdery mildews or red spider mites to
become established and then spread upwards into the main canopy. Stripping
is usually started when the bines are about 2 m high and is carried on later in
the season as required. The height to which it is done depends upon the
circumstances. On young plants it is kept to the minimum so that some leaves
will be left on the basal part of the bines that is left attached to the plant after
harvest. This allows some additional photosynthetic activity to continue which
helps to build up food reserves in the rootstock. Past experience of disease
58 Production methods

Figure 3.5 Basal growth removed by chemical defoliation (IRR, Wye).

problems in each garden can modify the amount of stripping that the grower
considers necessary. The introduction of bine-pulling machines at harvest has
required that bines be stripped to the height at which they are cut. Because of
these various factors, bines may be stripped to anything between 1-2 m high.
Stripping was originally a hand operation but today it is much more
commonly done by spraying with a chemical defoliant (Figure 3.5). This was
first done in the USA using dinoseb as the defoliant whereas the material used
in England for many years was tar oil, sometimes mixed with sodium
monochloracetate, but this was then largely superseded by dinoseb until the
use of this was banned. After the bines have reached the top wire it is usually
safe to use diquat or paraquat but these materials can cause damage if applied
to young bines. In Germany much use has been made of concentrated solutions
of nitrogenous materials such as ammonium sulphate or urea, generally mixed
with defoliants such as bromfenoxim or diquat. In England it is usually
necessary to spray three or four times in the season to cope with the renewed
flushes of growth.
In Australia and New Zealand, where there is no need for early-season
application of fungicides or pesticides, sheep are often put into the hop gardens
to graze and defoliate the bottoms of the bines.
Soils 59
3.3 SOILS

Although hop growing is concentrated in relatively small areas of each

producing country, soil type has probably been only one of the factors leading
to this localization. Hop plants will thrive on a wide range of soils although
different cultivars do appear to be better suited to some areas than others.
Even light, sandy soils can produce good crops provided the water supply is
augmented by irrigation and manuring is adequate. Hops will grow well on
heavy clay soils but these create real difficulties for the cultivator when he is
faced with the work of digging round the hills and dressing them in a wet
winter or cultivating in a dry summer. However, for hop production to be
economically viable the grower needs to keep costs down and yields high and
this can only be achieved on land that is easy to manage and suited to the
cultivars that he wishes to grow.
The main requirements for good yields are probably a sufficient depth of soil
for this deep-rooted plant and an adequate supply of moisture without any
danger of waterlogging. It has been suggested that a high water table during
the winter can kill off the lower roots and that the plants suffer during the
following summer, especially if there is a shortage of water then. The
importance of a well-developed root system has been noted in the breeding
garden at Wye in dry summers when two-year old seedlings have been checked
much more than three-year olds. In wet summers there has been little
difference between the two age groups.
Although hop production is restricted to areas where the soils are suitable
there are many other areas with similar soils where hops are not grown. One of
the reasons for such localization used to be the availability of sufficient labour
for harvesting by hand and in England this restricted production to places with
good access to the major cities of London and Birmingham. Although labour
is no longer drawn from those sources, production has not spread from the
traditional areas, largely because the necessary expertise and equipment are
not readily available elsewhere.

3.4 MANURING
Hops do not thrive under acid soil conditions and liming should be carried out
when necessary to prevent the pH from falling below 6.5.
Although hops require higher levels of fertility than many other crops,
recent fertilizer trials have shown that there is little if any benefit from the very
high levels of fertilizers that have frequently been advocated. Brown (1980)
records that Burgess's recommendation for the Guinness farm in 1934 was for
336 kg N, 230 kg P20S and 224 kg K20 per ha, in addition to 37 tonnes of
farmyard manure or 2.5 tonnes of shoddy. In the years 1934-7 the rates of
artificial fertilizer applied were considerably in excess of the recommendation
60 Production methods
averaging 432, 1132 and 342 kgjha respectively although very little farmyard
manure was used.
There can be considerable advantages on some soils of heavy dressings of
bulky organic manures that improve the moisture-holding capacity of the soils
and maintain soil structure and this may have led to the assumption that high
levels of artificial fertilizers are also needed. Under non-cultivation systems
however, trials have shown only small benefits, if any, from applications of
dung or straw mulches. Heavy straw mulches can be disadvantageous since
they keep down soil temperatures in the spring, delaying the growth of the
plants and increasing the risk of verticillium wilt.

3.4.1 Nitrogen
In spite of numerous fertilizer trials, there are still considerable differences of
opinion as to the levels of manuring that are required, especially for nitrogen.
In England, Burgess (1950) reported that maximum yields were obtained from
300 kgjha N but at present the official recommendation is considerably lower,
ranging from 150 kgjha if there have been frequent applications of farmyard
manure in previous seasons, to 225 kgjha when no FYM has been used for
several years (Ministry of Agriculture, 1985). However, several trials have
shown little response to applications of more than 135 kgj ha even in the
absence of FYM.
In Germany the recommended rates have been based upon the quantities of
nutrients removed by the crop each year and the figure quoted for nitrogen
with a crop yield of 37 zrjha is 120 kg N. Since the plants only utilize some
65% of the nitrogen applied it is calculated that such a crop requires 185 kgj ha
while for a crop of 45 zrjha this needs to be increased to 224 kgjha
(Kohlmann and Kastner, 1975). More recently however, growers have been
encouraged to adopt the Nmin system whereby nitrogen applications are based
on soil analyses which are interpreted with reference to the cultivar and its
expected yield to calculate the minimum application of nitrogen that is
required (Rossbauer and Zwack, 1989). The use of this method can lead to
reductions of 50-80 kg N for some cultivars.
The previous standard rates are considerably higher than those suggested for
the Yakima Valley in the USA where the recommendations are based upon the
soil test values for nitrogen in the soil. Even for the lowest nitrogen soils the
suggested rate is only 140 lbj acre (160 kgjha) while for the highest nitrogen
soils no further application is advised (Roberts et al., 1985). Since verticillium
wilt disease is more severe when heavy applications of nitrogen are used it is
recommended in England that 135 kgj ha is the maximum that should be
applied when this disease is present but some growers prefer to apply even
lower rates.
A further reason for keeping nitrogen manuring at as Iowa level as possible
Manuring 61
is the increasing concern about the quantities of nitrates that are consumed in
food and drink. Provided that brewers take steps to reduce the nitrogen
content of the brewing water, hops are the major source. Although more work
needs to be done to establish all the factors determining the nitrogen level in
hop cones it does appear that it increases with higher rates of nitrogen
fertilization.
The manurial recommendations in different localities may differ because
nitrogen is sometimes not a factor limiting yield. As the vigour of hop plants
increases, yields will initially increase also, but if the growth is too strong the
increased shading effect will have an adverse effect. It is to be expected
therefore that plants that are widely spaced would respond to higher nitrogen
levels than plants that are more closely planted. Since English hop gardens
usually have a much higher density of bines than other countries the nitrogen
requirement may also be lower. For the same reason it has been found that the
less vigorous cultivars may require heavier applications while the supply needs
to be restricted on the most vigorous sorts in order to avoid excessive growth.

3.4.2 Phosphate and potash

After several years of manuring hops at much higher rates than those used for
most arable crops, soil analyses can show very high residual values for
phosphate and potash. The application rates recommended in England for
these elements therefore vary according to the soil analysis. For phosphate the
maximum advised is 300 kgjha P20S but for most established gardens it would
be only 50-100 kgjha. For potash the maximum is 450 kgjha K20 but in most
cases only 75-150 kgjha. With the highest residual values no additional
phosphate or potash is considered necessary.
In Germany the levels recommended are again based upon the quantities of
each element removed by the crop and the recommendations for a crop of
45 zrjha, are 225 kg P20S, and 270 kg K20.
The recommended rates for soils in Yakima with the lowest soil test values
are very similar to the German figures but lower rates are suggested for the
more fertile soils and again no additional treatment is advised for the soils with
the highest soil test figures.
Although a deficiency of phosphate will result in a progressive reduction in
yield there are no visual symptoms to identify the cause. Phosphate is
particularly important for the stimulation of root development and it is
therefore necessary to ensure an adequate supply for newly planted hops.
Deficiency of potash also results in poor growth and reduction in yield
which is associated with a bronzing of the interveinal areas which later become
necrotic. Excessive levels of potassium should be avoided as these lead to
magnesium deficiency.
62 Production methods
3.4.3 Magnesium
At one time the magnesium requirements were supplied by the heavy dressings
of farmyard manure or crude potash fertilizers such as kainit, but without
them it is frequently necessary to include magnesium in the fertilizer pro-
gramme. Deficiency is recognizable as chlorotic yellowing of the interveinal
areas followed by necrosis and some cultivars are more liable to show these
symptoms than others (Marocke et al., 1979). Soil analysis should indicate the
need for additional magnesium before the symptoms occur and applications of
30-100 kg/ ha Mg are recommended.

3.4.4 Trace elements

Various trace element deficiencies occur in some areas. Perhaps the common-
est is zinc deficiency which is quite common in Europe and the USA. The
problem is known in Czechoslovakia as kaderavost and in Germany as
Krauselkrankheit (= curling disease) and was for many years thought to be due
to a virus infection. The leaves become light yellow in colour with upward
cupping, are sharply toothed and have an elongated middle lobe. In severe
cases the teeth wither and the leaves are brittle to touch. The condition can be
improved by spraying with fungicides that contain zinc but as these go out of
use it is necessary to spray with zinc sulphate.
In the USA zinc deficiency occurs most frequently in the heavily irrigated
desert areas such as the Yakima valley but has been much reduced since
growers included zinc in their fertilizer programmes (Roberts et al., 1985).
The other trace element deficiency that has been quite widely reported is
boron. It occurs in Germany and has also been reported from New Zealand
where the symptoms were described as delayed development of new shoots
accompanied by crinkling, often causing malformation of the leaves. The
internodes were short, laterals developed at an early age and the growing point
was killed (Askew and Monk, 1951). In England, Cripps (1956) further
described a rigidity of the young tips of the bines which had small leaves, large
stipules and short internodes. These symptoms were described as 'fluffy tip'.

3.5 IRRIGA nON

Hops require a good supply of water during the growing season to produce
high yields but in western Europe this is normally provided by natural rainfall.
In the drier regions of the world, however, irrigation is essential for hop
production, not only because of the low rainfall but also because under hot,
dry conditions the hop plants have a higher water requirement. It is reported
(Evans, 1985) that their requirement in Yugoslavia and the Willamette Valley
of Oregon is between 450-500 mm (18-20 in) whereas in the Yakima Valley in
Irrigation 63
Washington State they require nearly 760 mm (30 in). The Yakima Valley and
the Xinjiang Region of China, which lies at the western end of the Gobi desert,
are two of the most arid areas where hops are grown. In both these areas the
water for irrigation is provided by melting snow from the nearby mountain
ranges and it is distributed in the hop gardens by furrow irrigation rather than
overhead sprinklers. The combination of a controlled water supply and hot
sunny weather provides some of the best conditions for hop production.
Between these two extremes there are areas where hops can be grown
without irrigation but will not give economic yields. Pearce (1976) describes
how Australian growers, who initially used the methods with which they had
been familiar in Europe, suffered heavy losses from dry summer conditions
until a visit to Tasmania in 1840 by the Polish Count Strzelecki, who had
travelled widely in North America, did much to encourage the use of irrigation.
This was initially done with furrow irrigation but later was replaced largely by
overhead sprinkler systems.
In the Backa area of Yugoslavia conditions are dry enough for irrigation to
be a standard requirement and it is provided by overhead sprinkler systems.
Kisgeci (1980) reported on physiological changes to hops in irrigation trials
carried out there. Measurements of osmotic pressure showed only a low
correlation with soil moisture in the top 60 cm layer. It was concluded that this
was because the hops were deep rooted and were able to obtain water from
greater depths but there was a good response to irrigation because most of the
soil nutrients are in the top layer which irrigation made available. Irrigation is
also used in some parts of Czechoslovakia and other eastern European
countries.
In England some growers have used overhead systems of irrigation in dry
seasons but various trials have indicated that this only occasionally increased
yield significantly. Because of the expense of moving and maintaining the
equipment the system has largely lost favour although it is still used occasio-
nally on young hops during spells of dry weather. The development of trickle
irrigation systems, where plastic lines remain down permanently and less water
is needed, presented a new possibility and trials were commenced at Wye in
1973 (Thomas and Farrar, 1975). The first trial on Fuggle hops gave increased
yields in two years out of four but when the plants were grubbed it was found
that the hills from the irrigated plots were smaller with a peat-like mat of fine
fibrous roots close to the crown. A subsequent trial with the cultivar Wye
Saxon gave no significant increases in yield.
Evans (1985) suggested that trickle irrigation is probably the most efficient
method of irrigating hops but there were problems if the emitters become
blocked while pipes that were laid on the surface interfered with cultural
operations. Nor were underground systems to be recommended because of
maintenance problems. Wample and Farrar (1980) found that trickle irrigation
with fertilizer injection provided several crop management advantages but
64 Production methods
required changes in cultural practices. In the second year of their trial they
obtained the highest yield from trickle plots which received the least water,
fertilizer or labour.
Following reports that yields of fruit trees could be increased by the use of
intermittent overhead sprinkling with water in order to reduce moisture stress,
a similar treatment was tested at Wye (Thomas, 1981). In some treatments the
sprinkler treatment was combined with trickle irrigation. Measurements of leaf
water potential and stomatal conductance showed that, after sprinkling, the
water stress fell and the stomata were more open than in control plants.
Increased photosynthesis and increased growth and development were
expected to result. Non-significant yield increases were recorded from both
sprinkling and trickle irrigation except in 1982 when yields were lower in the
sprinkled plots than in the controls. This was attributed to either the excessive
('housey1 growth or to the downy mildew which was evidently encouraged by
the humid conditions (Thomas and Farrar, 1983).

3.6 SPRAYING

The very serious damage that can be inflicted on hops by various pests and
diseases makes it essential, in nearly all countries, to carry out regular and
efficient spraying operations. The crop does not provide an easy target for
spraying, at least during the later stages of growth. Not only is it grown at
heights of up to 7 m, but as the growth becomes more dense it is increasingly
difficult to achieve penetration of the spray to all areas. Although some
chemicals are now available with systemic action it is necessary in most cases to
achieve good cover of all parts of the plant to ensure good control of the
various pests and diseases.
Most sprayers in use today rely on fans to carry the spray droplets to the
necessary height (Figure 3.6) but some growers in continental Europe still rely
on purely hydraulic systems with the nozzles placed at varying heights on a
vertical boom in order to reach the crop at all levels.
Without air assistance the droplets must be relatively large in order that they
should travel far enough to reach the target but this is not necessary when they
are transported in an air stream. One of the problems with hop spraying is that
the use of large volumes of water means the sprayer has to return to the filling
point frequently, thus wasting a considerable amount of time. By reducing
droplet size it is possible greatly to reduce the total volume of spray that is
required so some air-assisted sprayers will work with very small droplets at
very low volumes.
In England the majority of growers use high volume, air-assisted sprayers
and apply from 600 1/ ha (50 gall acre) at the start of the season to a maximum
of 2200 l/ha (200 gal/acre) when the plants are fully grown. In Germany
Spraying 65

Figure 3.6 Fan-assisted sprayer (IHR, Wye).

volumes as high as 3300 I/ ha are recommended for this type of sprayer and as
much as 5000 1/ ha for hydraulic sprayers.
A few growers in England have ch~nged to low volume application with
rates of only 280 l/ ha (25 gal/ acre) and are satisfied that they are achieving
good control. In some cases an electrostatic charge is induced on the droplets
as they leave the sprayer so that they are attracted to the leaf surface but there
is little evidence so far to show how effective this modification may be. With
any sprayer there are difficulties, in a crop like the hop, in achieving a uniform
distribution of spray between the leaves close to the spray output and those
further away so there are advantages in providing outlets at different heights.
66 Production methods

Figure 3.7 American mobile harvester on 3m high wirework (Hopunion, USA).

When weather conditions are bad there is frequently only a limited period in
which spraying can be carried out and speeding up spraying operations not
only reduces costs but is also necessary to ensure that the whole crop can be
sprayed at critical times. Whereas a high volume sprayer may only cover a half
hectare at each filling, a low volume sprayer can cover up to eight times as
much. Transporting heavy loads of water through gardens when the ground is
wet can cause a lot of soil compaction and low volume spraying is better in this
respect also.
Whichever system of spraying is used, it is essential that the sprayer is well
maintained and correctly adjusted with the appropriate nozzles and pressure
and with the tractor driven at the correct speed. In many cases where the
grower complains that the sprays are not working effectively, the problem lies
in the way in which the spraying has been carried out.

3.7 LOW TRELLIS

A system has been developed in the USA of growing the normal cultivars on
wirework that was at first only 2 m high but it was found necessary to increase
this height to 3 m. Mobile harvesters have been constructed which straddle the
row (Figure 3.7) and these appear to be far more successful than the mobile
harvesters that have been used on normal, high wirework (Figure 3.8).
Low trellis 67

Figure 3.8 American mobile harvester for traditional high wirework (IHR, Wye).

Since the crop is harvested in situ, the bines remain attached to the wirework
and it has been found that these can be used to support the next season's
growth, thus eliminating the need for restringing each year. The harvester
should pick the hops from the laterals as cleanly as possible so there is no need
to complicate the machine by adding a lateral picker. The laterals are then left
attached to the main bines and when the next year's shoots start to climb they
frequently grow out along these laterals instead of going upwards. It is
therefore necessary to remove the old laterals during the dormant season.
Very little training is required and hops on low trellis offer a much better
target for spraying to control pests and diseases, reducing spraying costs and
producing cleaner crops.
68 Production methods

Figure 3.9 Prototype harvester picking dwarf hops on 2m high wirework (lRR, Wye).

It was found in England that the standard cultivars were not suitable for
growing under low trellis and a breeding programme is in progress to develop
dwarf types that should be more successful. It was intended that these would be
grown on 2 m high wirework (Figure 3.9) but it now appears that it may also
be necessary to increase this to about 2.5 m in order to achieve economic
yields. Whereas most hop yards in the USA are quite level, many English hop
gardens are on sloping land and the extra height may create difficulty with
stability of a harvester that has to be high enough to straddle the row.
Low trellis offers considerable cost savings on such operations as stringing,
training, spraying and harvesting so efforts are likely to continue in developing
a commercially acceptable system.

3.8 SEEDED AND SEEDLESS HOPS

Since the male and female flowers of hops are produced on separate plants, the
females will only be pollinated and produce seed if there are male plants
sufficiently close for the wind-borne pollen to reach them.
Many brewers insist that the presence of seed is undesirable, usually with no
explanation, on the grounds that oxidation of the seed fat can give rise to
Seeded and seedless hops 69
rancid flavours. There is little experimental evidence to support the objection
to seed although Zattler and Krauss (1970) found that lager brewed with
seedless hops was preferred after storage for six weeks. Several other in-
vestigations have failed to detect such differences (BIRF Ann. Report, 1984;
Harrison, 1971; Pfenninger et al., 1978; Neame et al., 1980) but brewery
demand remains such that, in most countries, hops are grown seedless by
ensuring that all male plants in the hop growing areas are destroyed. In
Germany, for example, there are regulations that require all wild hops to be cut
down or, where possible, grubbed by 15th June at the latest (Kohlmann and
Kastner, 1975).
In Europe the seed content of dried hops has to be below 2% for them to be
classed as seedless while in the United States the limit is 3% but this is
equivalent to 4.2% when determined by the same method as in Europe.
Salmon and Amos (1908) published a report indicating that the yield of hops
was considerably higher when grown seeded and they encouraged English
growers to plant males by offering to supply them free of charge. Until then
there had been no definite policy regarding the use of male plants but from that
time on English hops were invariably grown seeded. Brewers in Britain did not
share the dislike of seeded hops that German brewmasters had advocated as
they spread their influence around the world. Since the English hops were
bought mainly by British brewers there was every advantage to be gained by
the grower from planting male hops in the gardens.
The development of high a-acid varieties in England offered an opportunity
for the export of such hops and in 1963 an experiment at Wye again indicated
that yields of seedless hops were reduced (Neve, 1964a). There was a further
stimulus to overseas trading as a result of Britain's entry into the European
Economic Community and this led to more accurate experiments being started
in 1971 (Thomas and Neve, 1976). These showed that a reduction in yield of up
to 30% resulted from growing hops seedless but that there was, with the high a-
acid varieties in particular, an increase in a-acid content that largely com-
pensated for this since the price of such hops is based upon their a-acid analysis.
With the traditional cultivars Fuggle and Golding there was no increase in a-
acid (nor would such an increase enhance the price of these aroma hops) and it
was therefore uneconomic to consider growing them seedless.
In the USA too it proved uneconomic to grow Fuggle seedless so males had
to be planted in this variety whereas it is standard practice to grow other hops
seedless. This stimulated the breeding of cultivars with similar qualities to
Fuggle that could be grown economically without seeds.
One breeding technique which could do this was the production of triploid
cultivars which is described in section 9.7. Triploids are very infertile and
although, if fully pollinated, their seed content may exceed the European limit
of 2% for seedless hops, this limit can be achieved without very stringent
elimination of male plants from the garden and its surrounds.
70 Production methods
The ordinary diploid hop cultivars vary considerably in the amount of seed
that they set when pollinated and it has been found the English high a-acid
cultivar Wye Target has a low seed content so by removing males from gardens
of this hop the European limit of 2% seed can be readily achieved, even when
neighbouring farms still have male plants in their hop gardens.
An alternative to breeding infertile female cultivars is to use male plants
whose pollen will stimulate cone development but will not produce seed. Davis
(1959) found that the annual species Humulusjaponicus when used in this way
stimulated the development of bracts, bracteoles and internodal length though
to a lesser extent than ordinary male hops and he said that its commercial
practicality had not been tested. This method is currently being tried in
England (Gunn, personal communication).
Another method, initiated by Haunold (1975), was to use triploid male
plants of H. lupulus. Because of irregularities in the meiotic divisions leading
to pollen formation in such males, many of the gametes which they give rise to
are not viable and so only empty seed cases develop. In field trials (Haunold
and Nickerson, 1979) the overall seed content of hops pollinated by triploids
was 4% compared with 2% in an adjacent yard which contained no males and
the yield was 30% higher. In the females close to the triploid males, however,
the seed content was much higher, averaging 10.3%. Although the reduction in
seed weight may be sufficient to qualify for the seedless limit of the USA it
would be less likely to meet the more stringent European standard.

3.9 GROWTH SUBSTANCES

Seeley and Wain (1955) attempted to stimulate cone development by applying
synthetic growth substances or pollen extracts to early or fully developed burr.
They reported that 2-naphthoxyacetic acid and a(2-naphthoxy)propionic acid
caused a marked initial stimulation of cone growth but the effect did not
persist. After 20 days the control cones were as large as those that had been
treated while at harvest both the cone weight and a-acid content of the treated
hops were lower. There was no effect from pollen extracts.
Weston (1960) found that a(2-chlorophenylthio)propionic acid applied to
female hop plants induced the formation of some male flowers. The earliest
applications (2 and 4 July) had most effect with fewer male flowers from the 6
July treatment. He did not report that any viable pollen was produced.
Nash and Mullaney (1960) reported in Australia that applications of
gibberellic acid at a concentration of 12.5 ppm to hops when the stigmas were
lis in (3 mm) long advanced the maturity of the developing cones by 10 days
and increased the yield by 40%. However, the a-acid content was reduced from
10.16% to 1.8% so the treatment was not commercially beneficial. Roberts and
Stevens (1962) repeated the treatment in England on the cultivar Bullion and
found that, although the treated hops were initially more advanced, by the time
Growth substances 71
of harvest the differences had disappeared and the yields were lower than the
untreated controls and the a-acid contents were the same. They suggested that
their results might have been different from those in Australia because of the
different environmental conditions. They did not suggest another explanation,
that the Australian hops were harvested at an earlier stage of maturity before
the untreated hops had time to catch up with the treated ones.
Zimmermann et al. (1964) treated Fuggle hops in America with gibberellin
A3 at various stages of growth. They agreed with the British workers that
application at the burr stage did not influence maturity date but that the
treated hops were liable to shatter badly when harvested. However, a
concentration of 5 ppm applied at the 1.5 m (5 ft) stage resulted in a large set
of small cones and a 25% increase in yield. Hartley and Neve (1966) tried the
same treatment on Fuggles in England and found that the treated plants also
produced a greater number of smaller cones but that there was no increase in
yield. They suggested that the different result might be related to the fact that
in England the plants were spaced 2 m x 2 m (6.5 ft x 6.5 ft) with four strings
to each plant compared with the American system of 2.4 m x 2.4 m (8 ft x 8 ft)
and three strings per plant. With the denser plant in England there could be a
limit to the ability of the plants to support the development of the increased
number of cones.
Further trials in England (Thompson and Thomas, 1971) demonstrated no
increase in yield of similar treatments to Fuggle, Eastwell Golding or Bullion
but the yield of Northern Brewer was sometimes significantly increased and
never, apparently, reduced. There was an additional benefit that the number of
leafy cones ('cock' hops), which are an undesirable feature of this cultivar, was
frequently reduced. The treatment was therefore recommended for Northern
Brewer and later for Wye N orthdown which is a seedling of Northern Brewer.
Thomas and Goldwin (1976) tested mixtures of growth substances that were
effective on apples on hops. The most effective mixture consisting of GA3
(5 ppm), NN'-diphenylurea (3 ppm) and 2-naphthoxyacetic acid (0.4 ppm)
applied at cone formation gave small but significant yield increases. Responses
differed with cultivars, Northern Brewer showing yield increases of 9-12%
while Wye Challenger only gave 4-5% increases. A subsequent report (Thomas
and Farrar, 1979) indicated little benefit from the treatment in a range of trials.
There have been a number of trials with growth retardants, mainly with the
object of reducing the excessive growth of some cultivars. The retardants (2-
chloroethyl)trimethyl ammonium chloride (CCC) and N-dimethylamino succi-
namic acid (= daminozide, B9 or Alar) were tested for several years at Wye.
Both compounds caused shortening of the internodes but the effect lasted for
only about 10 days (Hartley, 1966). Repeated applications led to a general
shortening of the internodes but yields were much reduced (Hartley, 1967).
Thomas (1968) reported that B9 could induce flowering in plants growing
under continuous light. In natural dayIength it reduced yield but increased
72 Production methods
a-acid content. The final conclusion (Thomas, 1973) was that this material was
of doubtful value. Only on Fuggle had increased yields of a-acid been
recorded; on other cultivars the increased a-acid content had not compensated
for the reduced yield.
Similar transient shortening of the internodes has been recorded with
paclobutrazol (PP333 or Cultar) applied as a foliar spray but yield was
reduced. When applied as a soil drench there was no effect until the bines
reached the top wire when the intermodes and laterals were shortened. Both
sprays reduced a-acid content and yield. The effects of the soil treatment
persisted into the following season, initial growth being slower although by
harvest differences had disappeared (Thomas, 1982b, 1983). One trial indicated
that early application of paclobutrazol delayed shoot development but that
growth was vigorous when the effect of the treatment wore off. It was thought
that this could be a useful management tool to hold back the development of
some hops on a farm so that everything did not require training at the same
time (Thomas, personal communication).

3.10 PROPAGATION
Hop gardens are invariably planted with clonal material of one of the standard
cultivars and this has to be produced by vegetative propagation. This can be
done very easily by taking cuttings from established plants in commercial
gardens. In England this was traditionally done by earthing up the bines during
the latter part of the growing season, inducing the covered bases to thicken and
develop perennial buds. These bine bases could be cut off from the hills in the
winter to provide 'strap cuts' which would be planted directly into a hop
garden, usually putting 2 or 3 at each hill position. Alternatively they could be
grown on in a nursery for one year to produce 'bedded setts' which would
establish quicker than strap cuts (Figure 3.10).
In the USA the corresponding technique was to remove underground
runners from the stock during the winter and plant several of these at each hill
position.
The spread of verticillium wilt in England resulted in a demand for large
quantities of planting material as resistant varieties became available and this
led to the development of more rapid methods of propagation.
Instead of earthing up the base of the bine more cuttings could be obtained
by lowering the bine from the top wire, laying a length of it along the ground
and covering this with soil, while the tip was retrained upwards along another
string, usually that of the next hill in the row. In this way the whole of the
layered portion thickened up and could be cut up into single-node sections to
provide several layer cuts (Figure 3.10). Plants in a commercial garden could
be layered in this way, provided that cultivations were to be carried out in one
direction only, but an alternative was to establish permanent layer beds on full
Propagation 73

Cutting

A Basal cutting
with several nodes

Set
1 foot

Figure 3.10 a) Bedded sett; b) layered bine. Dashed lines indicate suitable positions for
dividing into cuttings (drawn by R. F. Farrar).

height wirework and to use these solely for propagation (Keyworth et al.,
1948).
An alternative to layering, which is termed air layering, has been suggested
by Vasek and Tassell (1984). The technique consists of placing black polythene
sleeves (20 cm diameter, 60 cm long) round the plants in the spring and, once
the bines reached the top wire, filling the sleeves with pulverized bark to a
height of 50 cm. By treating up to ten bines per hill in this way they were able
to produce 40-50 one-node cuttings which were superior to those obtained
from layering below ground.
Permanent layer beds had the disadvantage that there was a delay of a whole
season during which the plants were becoming established. A more rapid
system was to plant strap cuts or bedded setts in nursery rows, under 2 m high
wirework, and to layer the bines that they produced in the first season. A
technique was devised using two top wires, one of which could be lowered
when the bines had made sufficient growth, so that a length of each bine lay on
the ground where it could quickly be covered with soil (Keyworth and Wilson,
1948).
A more rapid multiplication would be achieved if it were possible to root
soft-wood cuttings quickly during the growing season; this was also attempted
at East MaIling by Beard and Wilson (1946 and 1947) who concluded that the
methods they used were not of wide application.
74 Production methods

Figure 3.11 Cuttings prepared for mist-propagation (right) two-node cutting; (left) half-
node cutting (IHR, Wye).

Soft-wood propagation became a very practical proposition following the

work of Williams and Sykes (1959) in which they adapted the mist-propa-
gation system for use with hops. In their paper they suggested that existing
methods were adequate for commercial propagation and that the more rapid
multiplication made possible by the new technique would be useful mainly for
research projects.
Cuttings rooted in a mist bench (Figure 3.11) will rapidly make sufficient
growth to provide approximately ten further cuttings within 4-6 weeks. Since
cuttings are only in the mist unit for 10- 14 days, two or three batches can be
put through in each 4- 6 week period. Quite soon the factor limiting the rate of
multiplication is the capacity of the mist bench rather than the number of soft
wood cuttings available.
The advantages of this form of rapid multiplication were soon appreciated.
New cultivars can be bulked up for large-scale planting much more rapidly
than was previously possible and this is extremely important at times when
growers have an urgent need to change varieties. Since only small nuclear
stocks are required to commence the process each season it is possible to
ensure that these are true to type and, even more importantly, that they are free
Factors affecting quality 75
from infection with the various virus diseases that can severely affect produc-
tivity (Chapter 8).
When cuttings were taken from plants in a commercial hop garden there was
no check on their virus status and this resulted in considerable spread of such
diseases. With the adoption of mist propagation of virus-tested stocks there
was a rapid improvement in the situation. In England for example, a survey
showed that all the available stocks of the established cultivars were 100%
infected with prunus necrotic ringspot virus but this changed very rapidly as a
result of large-scale planting of new varieties that were available free from this
virus. Over the two seasons 1973 and 1974 some 25% of the total English hop
area was replanted with healthy stocks and the process has continued at a
slower rate since then. It was calculated that in 1976 the value of the English
crop was increased by £ 1 million as a direct result of the virus-free status of the
newly planted material (Thresh, 1979).
Another possible method of rapid propagation could be by tissue culture
which has been adopted on a commercial scale in South Africa but elsewhere it
offers little advantage for a crop that can be multiplied so rapidly and
successfully by mist propagation.

3.11 FACTORS AFFECTING QUALITY

The quality of hops is assessed by a combination of chemical analysis and

evaluation of appearance and aroma. The appearance of a sample can be
adversely affected by damage from pests or diseases, by wind bruising, by the
presence of extraneous matter or by faulty drying methods. These topics are
dealt with in the relevant chapters.
Chemical analysis is concerned principally with the a-acid content of the
sample and there have been several investigations into the factors that might be
responsible for the considerable variations in a-acid that occur in samples of
the same cultivar between farms and between years.
It has been difficult to relate quality to different cultural practices other than
effective control of pests and diseases. Burgess (1935) showed that omitting any
of the major nutrients from experimental plots for 10 years resulted in
considerable reductions in yield but that quality was not affected. Keller and
Magee (1952) found that applications of nitrogen in excess of normal practice
resulted in significant lowering of the total soft resin and B-fraction but not of
the a-acid contents. There was no significant correlation between yield and
resin content. Thompson and Neve (1972) reviewed the evidence on manuring
and concluded that only small effects of manuring on a-acid content had been
demonstrated. Where there had been a response, the most consistent effect had
been a decrease in a-acid with increased nitrogen applications whereas high
potash usually had the reverse effect.
76 Production methods
Thompson (1957) reported largely inconsistent effects of height of wirework
or plant spacing on a-acid content of four cultivars in 1954 and 1955 while a
subsequent height and spacing trial also indicated negligible effects
(Thompson, 1967). In both these trials there were significant differences in
yields of cones between treatments and these led to corresponding differences
in the yields of a-acid per hectare.
One cultural treatment that does have a considerable effect upon hop quality
is the elimination of male plants in order to produce seedless cones as described
above (section 3.8).
The effects of replacing hand picking by picking machines have been
investigated and at Wye Thompson and Neve (1972) reported a-acid contents
from 7-11 % lower in machine-picked samples. The lower values were partly
due to dilution of the cone material by leaf and stem extraneous matter but was
mainly due to loss of lupulin from whole and apparently undamaged cones.
More recent experience with the cultivar Yeoman has shown that the belts of
picking machines working on seeded hops rapidly become coated with thick
layers of lupulin whereas this does not happen with the less open cones of
seedless hops. In Yugoslavia, Spevak and Kisgeci (1982) reported losses as high
as 20% in machine-picked hops compared with those picked by hand.
The major factors affecting quality are undoubtedly environmental con-
ditions. Burgess (1964) quotes Meneret and Svinareff (1955, 1956) who
concluded that a-acid content depended upon the amount of sunshine during
the development and ripening of the cones but that it might also be influenced
by the amount of rain during maturation. Zattler (1960a) had shown that
shaded hops had a lower resin content than unshaded ones and that those
produced in a glasshouse had a higher content than those grown outside at a
lower temperature. Zattler and Jehl (1962) concluded that a hot, dry summer
with much sunshine during cone formation gave lower resin contents than a
moist summer with lower temperatures and a normal amount of sunshine.
Burgess himself examined data for the English crop and found strong
indications that total soft resin content increased rapidly with higher August
temperatures with no falling off at the highest temperatures. There appeared to
be a similar increase in a-acid but with a lessening response at the higher
temperatures. July temperatures did not appear materially to influence either
total soft resin or a-acid levels.
Smith (1970) examined data relating to several cultivars grown in various
European countries and concluded that there was convincing proof that mean
air temperature over the last 40-60 days before maturity exercised a strong
influence on the a-acid content. He also suggested that very high temperatures
(over 26°C) had an adverse effect. Lyashenko (1985) found that temperatures
as high as 18°C reduced a-acid content.
Farrar (1971) carried out a survey of several gardens of Bullion hops in
England during 1969 and 1970 but was unable to identify any factor that was
Factors affecting quality 77
of practical value in helping a grower to improve or maintain the a-acid level
of the hops. He concluded that any small effects of changes in management
would be masked by climatic and geographical considerations that would be
beyond the control of the grower. However, there was little correlation
between the a-acid contents of each site in 1969 and 1970 which is difficult to
explain in terms of climate or geography since they were all located within a
relatively small area.
Thomas (1980b) submitted the analyses of the whole of the English crop for
the years 1971-1978, which were available from the laboratory of the Hops
Marketing Board, to multiple regression analysis in an effort to establish which
weather factors were related to a-acid content. He found a high correlation
with the mean temperature from 24 May-21 June whereas temperature or
sunshine during the cone-ripening period of August appeared to be of
secondary importance. Lewis and Thomas (1982) investigated the variations in
a-acid content within the same cultivar and found they were due to differences
in the ratio of resin gland numbers per unit weight of cone material and/ or
differences in the a-acid content of the glands themselves.
The correlations found from the 1971-8 seasons were used to predict the a-
acid levels of the 1979-83 seasons. Whereas good predictions were made in 1979,
1980 and 1982, there were wide divergences in 1981 and 1983 (Thomas and
Darby, 1984). This led to the proposition that the final a-acid content could be
the result of a series of interacting weather factors which they listed as follows:
1. factors affecting cone weight;
(a) during flower initiation;
(b) during pollination (seeded crops only);
2. factors affecting resin gland numbers per cone;
3. factors affecting resin gland size;
4. factors affecting the percentage of resin in the glands.
The final a-acid content would result from a combination of these and possibly
other factors. A mathematical model was constructed which assumed, amongst
other things, that these factors combine in a multiplicative rather than additive
manner. This model was used and indicated some highly significant associ-
ations which supported the contentions of Thomas that May / June weather
and of other workers that the cone-ripening period were of crucial importance.
It is unfortunate that this hypothesis has not been tested in subsequent seasons.
Versluys (1982) examined Australian data and found low levels of correl-
ation from applying Thomas's hypothesis. He found that minimum and
maximum temperatures during January and February fluctuated less in years
when a-acid contents were high and deduced that there were high and low
temperature thresholds which, if exceeded, depressed a-acid synthesis. He
calculated the best fit of these two limits with a-acid content were when he
chose a critical period from 18th January-7th February and threshold
78 Production methods
temperatures of 10.40 C and 23.1 0 C. That critical period coincided with the
flowering stage of the main Australian cultivar. He introduced, as an
additional factor, the fluctuations of minimum and maximum temperatures
during the critical period and developed a formula to fit the data for the years
1973-80 which he then used successfully to predict the a-acid content of the
1981 crop.
He applied his method to English data for the two cultivars Northern Brewer
and Bullion with similar results except that there was no negative effect of
temperatures above a threshold maximum. When he examined the limited
German data he decided that the temperature effects were the same as in
Australia.
Diseases can also have very significant effects upon the a-acid content of
infected plants. Royle (1969) reported that increasing severity of infection with
downy mildew significantly reduced a-acid content. Virus infections, especially
with prunus necrotic ringspot virus, have been shown to have a very important
effect upon a-acid content. This is described in more detail in section 8.5.
CHAPTER 4

Harvesting

4.1 PICKING
Possibly the most economically important development in the methods of hop
production was the transition from picking hops by hand to the use of
machines. Mechanical harvesting commenced in the USA in the early part of
the 20th century but did not become normal practice until labour shortages
during World War II accelerated its adoption. The Productivity Team Report
(1951) states that by 1951 more than 85% of the USA hops were picked by
machine.
The American lead in this was followed by England when an American
machine was imported in 1922 but proved unsuccessful. In 1935 the first
English machine was installed and by 1952 some 10% of the crop was harvested
by machine. This figure had increased to 65% by 1958 and 90% by 1963.
In the Federal Republic of Germany, the first picking machine was one
imported from England in 1955 but thereafter locally produced machines were
installed very rapidly and by 1963 they were being used on 85% of the crop.
There were similar developments in other countries. In Czechoslovakia too the
first machines were imported but much effort was then put into developing
their own equipment.

4.1.1 Hand picking

Over the years there have been many changes in the way hops have been grown
and these have sometimes necessitated corresponding changes in harvesting
methods.
Because most hop cones are produced out of reach from the ground, the
bines need to be lowered in order that the hops can be picked. When the bines
were trained up poles, the poles themselves had to be lowered. However since
they were firmly embedded in the ground it was necessary first to raise them
clear. The method was described by Reynolde Scot (1576)
And bicause, when the hylles are made great, and raysed high; you can neyther easily
pull up any, nor possiblye pull up all your Poales except you breake them ... I thought
good to shewe you an Instrument wherewith you shall pull them up without disease to
80 Harvesting
your selfe, destruction to your Poales, or expence of your money, the charge beinge
onelye fourtene or fiftene pound of iron wherewith the Smith will make you a paire of
Tongues ... or Pynsers, of the fashion here set downe.
The pole was gripped by tongs (Figure 4.l) and prised out of the ground
using a block of wood as a fulcrum. First, however, the bine had to be cut at
the base to separate it from the rootstock so that the pole around which the
bine was tightly twined was free to be lifted.
With the introduction of permanent wirework systems, in which the bines
were trained up either string or thin wire, it was possible to lower the bines in
order to pick them without having to sever them from the rootstock (Figure
4.2) and they could be left attached until after harvest was completed. Hall
(1902) suggested that leaving the bines uncut might benefit the plants in the
following season by allowing additional nutrient reserves to be translocated to
the rootstock. In 1897 and 1898 one plot had been grown on poles and the crop
in 1898 and 1899 was below that of the rest of the garden where the hops were
under wirework. In 1899 the system was changed and the previously poled plot
was left uncut at harvest. There was a partial recovery of yield in 1900 and this
was complete by 1901.
Once the bines had been lowered to the ground the cones could be picked off
by hand. This required the services of large numbers of people and the
availability of an adequate supply of such temporary labour was one of the
factors limiting the districts in which hops could be grown. Most of this labour
for southern England came from London and for the west Midlands from
Birmingham. Farmers had to provide shelter for the pickers during their stay
on the farm though this was usually of a very poor standard. Nevertheless, in
the days before paid holidays, hop picking provided the only opportunity that
most of the pickers ever had to visit the countryside and the work was popular
for that reason.
In England the members of the same family or a group of friends might pick
as a team. The hops were picked into bins or baskets. Some leaves would
inevitably collect amongst the hops, and pickers were expected to remove these
in order to obtain as clean a sample of hops as possible. At the end of each day
the hops were measured out by volume on which payment was based. In the
USA, on the other hand, pickers were paid by the weight of green hops
(Productivity Team Report). This difference in procedure is possibly
accounted for by climatic differences; in England the weight of the hops would
be greatly affected by heavy dew or rain but these would not have had so much
influence under the drier conditions in the USA.
In the days when many hop pickers in England were illiterate the quantities
of hops they had picked were recorded by means of wooden 'tallies', the two
parts of which were laid together and marked with a file. Alternatively, the
farmer would issue 'tokens' which were usually similar to coins that were
locally made of a design unique to each farm (Burgess, 1964).
Picking 81
22 vi'peifite platforii1e
aake benoe tI)e b~Ok£t1 19oale, ani) to tpe tbe
b~ok£n poal£ to tbe fame) wbtcb Inare bplJollle
tbe faloe b~ollctl ~oale,at1n p~rfrtne tbe ~oppe.
JlftlJe t90ale be onlp b~olletl at tbe netber enDe,
pon tllape (houe tbe fame ~~oaIe againe, into tbe
~tllJ anti fo leaue it.

Of pulling vp Poalcs,~

tf\,li ~ ~ bicaufe, bJben toe bpUe~ ate maOe

~ J gtent, anD rarrell bfgb; you can neptber
~. eafilp pull bp anl', no~ polTiblpe pull bp all
pont lfjoales e,:cept POll b~fake tbeln. etc. erpeci~
nllp if tbe metbrt o~ tbe gronnO be o~te, o~ dfe tbe
l~oale~ olile o~ ftllall, '] tl)ongl)t ~roo to ftJewe
ron nn lnlltument wbererottb )'on llJall pull
tbem bp witbont ilifeafe to pout felfe,tJeftructfon
to l'rmc t0oaleS, 02 rtorner of rOUt monep, tbe
rF=========~=======~ cbarge

~ M""WO""'.'f:: ~o~~~! teneo~

fiftene
., pounD.
..______ =======--=_of]~
tOli,Un;n;UJ'~'J ~y(: ~lilH(l) ~inL lHd6f fOU a patre
of '([:ongne~ , o~ (tatber ron tnape caU tbetn) a
pnp~t of ~pnrers, oftbe fal1)ton bere ret norone,
tIle wl)icb tllnrenlfo be Inane witt) wrooe if POU
ti)mllc gruD.
Figure 4.1 'Pynsers' for pulling up hop poles (Reynolde Scot, 1576).
82
83
84 Harvesting

Hand picking has been very largely abandoned in favour of machine picking
in most countries, partly because of the cost but also because it has become
increasingly difficult to recruit sufficient casual labour to do the work. One
exception is China where there is not only sufficient labour available but the
wirework system would not be suitable for mechanized bine collection.

4.1.2 Machine picking

Although there have been experiments with mobile mechanical harvesters that
could pick the hops in situ as with hand picking, only static machines have so
far proved of any importance and these require the bines to be cut in the field
and transported to the picker. This again raised the question of the effects of,
cutting the bine on yield in the following season and led to fresh investigations.
Thompson (1959) reviewed six years of trials which showed that overall any
loss of crop was negligible. He pointed out, however, that all the hops were
harvested when fully ripe and suggested that the time of harvest might have an
important influence. This comment is supported by observations in com-
mercial practice where it is frequently necessary to start picking a cultivar
before it is fully ripe in order that it is not too over-ripe by the end. Experience
of many growers is that the first-picked gardens do suffer in the following
season and this is particularly noticeable with the earlier maturing cultivars.
Williams (1962) carried out an experiment on hop plants in their first
growing season to test the effects of stripping the leaves to a height of 1 m in
late May and up to 1.5 m in early June, and also of cutting the bines at a height
of 1.5 m at the end of the first week of September. Both stripping and cutting
reduced yields in the following season and these were more than halved by the
combination of both treatments. The reduction in yield was closely correlated
with the starch reserves in the rootstocks at the end of the first season.
The Productivity Team Report (1951) described the types of picking
machine then in use and there have been only relatively minor modifications
since then. It was during the 1950s that attempts were made to develop the
Harvestex machine which employed a totally different mechanism. Brown
(1980) describes it as designed to operate by sucking the hop cones into circular
orifices of special design; round the circumference of each rotated a knife
which cut off the hops. The hops fell onto a belt while the leaves were carried
away in the airstream. Although originally thought to be a promising idea
several years' work failed to produce an effective machine.
Although such alternative mechanisms have been tried, all the machines in
commercial use are equipped with a system that removes the cones from the
laterals by means of wire loops (or 'fingers) which are usually mounted on
rotating drums. The bines are transported past these, either hanging upside
down between two banks of fingers or drawn horizontally, base first, over a
line of picking drums.
Picking 85

Figure 4.4 Layout of horizontal hop picking machine (Productivity Team Report,
1951),

Figure 4.5 Layout of lateral picker (Productivity Team Report, 1951).

The earliest American machines had horizontal pickers (Figure 4.4) but in
the 1950s vertical pickers were developed and these then became dominant
(Riel, 1970). In Europe vertical pickers have been most widely used, the only
exception being the Allaeys machine which had a horizontal unit.
In practice some laterals, with cones still attached, are stripped from the
bines in the picking unit and these have to pass through a 'lateral picker'
(Figure 4.5). The picked material then has to undergo a cleaning process to
separate the hops from pieces of leaf, stem or other extraneous material. This is
done by a combination of coarse mesh conveyors and spaced rollers through
which the hops will fall while large pieces of leaf and bine will be carried to
waste. This is followed by fan cleaners which suck leaf material onto a wire
mesh, and then by inclined belts, travelling upwards, down which the cones
will roll while flatter waste products are carried upwards and away (Figure
4.6). In many cases there is also a visual inspection on the final conveyor belt.
The bines are carried through the picking mechanism on a continuous bine
track which is fitted either with clamps into which the bine bases are fixed or
with hooks around which they are twisted. After they emerge from the picking
unit the clamps are triggered to release the picked bines or a blade cuts the
bines close to the hooks. The spent bines may then be chopped up and blown
onto the waste heap or carried away unchopped.
86 Harvesting

Unpicked lateral
" square wire mesh
conveyor
M

Jigging tray:
uq:::~~~~~,~300 st~okes per min

I
1-4-----i-
Vibrating screen
of fine wire
mesh
R

,,
\
I\
I , I
\ I
, I
I
I \I
"tI
I "'\
~ \\
'----.,......
1 ,..
Hop Leaf and Petal
flow stem

Figure 4.6 Layout of fan cleaner and inclined belts (Productivity Team Report, 1951).

Although the basic design of the picking machines has not changed there
have been several modifications to ensure more efficient working. The most
significant has been the installation of magazines which hold a reserve of
clamps ready loaded with bines. This helps to smoothe out any irregularity in
the supply of bines to the machine from the field.
The magazines can only guard against minor interruptions in supply and
much attention has therefore been paid to the methods of collecting the bines
from the field. They have first to be cut to separate them from the rootstock
and this is usually done at a height of about 1-1.5 m since there are few, if any,
hop cones below this point and the length of bine to be dealt with by the
machine should be kept as short as possible. The bines then have to be
detached from the top wire and the least sophisticated way of doing this is
simply to pull them, from the ground, hard enough to break the supporting
Picking 87

string. The bines are then thrown onto a trailer which carry them to the picking
machine. This is hard work, not only pulling the bines down but also picking
them up from where they have fallen and throwing them up onto the trailer.
The work has been greatly simplified by using portable crows nests which
could be fitted into a socket on each trailer while it was being loaded and then
removed and used on the next trailer. A worker climbs into the crows nest and
cuts the strings close to the top wire and the bines fall into the trailer. The butts
of the bines would mostly be laid between upright posts at the end of the trailer
so that they were readily available for loading onto the picking machine
without tangling. In the USA the crows nest was more commonly replaced by
elevated tractor-mounted frames (Shea, 1980) (Figure 4.7).
Whereas in Europe it was normal to load the bines onto tractor-drawn
trailers (Figure 4.8), in the USA trucks were frequently used, especially for
long hauls which involved travel on public highways (Productivity Team
Report).
More recently in the USA the operation has been mechanized by means of a
reciprocating or rotating blade positioned to cut the bines just below the top
wire by means of an hydraulically operated arm that is mounted on the front of
a tractor which pushes the collecting trailer or truck along the row.
Alternatively in Europe, mechanized bine pullers have been introduced that
cut the base of the bine and grip the severed end. As the machine moves
further along the row it pulls the bines from the top wire and they fall onto a
trailer which is towed behind the bine puller. In one system a single unit is
attached to the tractor which serves a number of trailers while another system
uses a considerably cheaper device, one of which is mounted on each of the
trailers.
Labour costs for bine collection and transport to the machine are a major
item for consideration and it is for this reason that so much attention has been
paid to mechanization of the process. A better solution would be a machine
that would harvest the hops in the field, thereby eliminating the need for bine
collection entirely. Shea (1980) described mobile harvesters that had been
developed in the USA. These picked the hops but did not clean them so they
had to be transported to the static machine for this operation to be carried out.
He reported that the picking machines were little used and attributed this to
their capital cost and difficulties with maintenance.
Mobile harvesters are, however, the only suitable method for picking hops
grown on the low trellis system. This wirework is low enough for the machines
to straddle the rows and pick the hops by means of a bank of picking fingers on
each side of the row. Because the bines are not inverted, as they are when
loaded onto a static machine, the picking fingers work in an upward direction.
Such a machine is in production in the USA to work on the 3 m high low trellis
developed there while a prototype machine is being used in England for the
dwarf hops growing 2 m high.
88 Harvesting

Figure 4.7 Tractor-mounted crows-nest in USA (Hopunion, USA).

Far greater economies would be achieved if the hops could be not only
picked but also cleaned in the field and a machine has been developed in the
Federal Republic of Germany which will do this. To ensure that the cleaning
operation is successful this machine has a self-adjusting hydraulic system to
keep it level and this contributes to the considerable size and weight of the
Drying 89

Figure 4.8 Hop bines transported by tractor-drawn trailer in England (IHR, Wye).

machine (12 tonnes). Trials with the two machines that were built showed that
harvesting costs were reduced by 27% (from 3000 to 2200 DMjha) but this
advantage was offset by the high cost of the machine. The limited potential
market for it meant that there was no scope to automate the manufacture and
it would have to be constructed entirely by hand (Richtsfeld and Gme1ch,
1982). So far it does not appear to have attracted much support in Germany
where most hop farms are too small for its use to be justified and it is
understood that the machines are now working in Czechoslovakia where the
large collective farms are more suitable for it.
Another mobile machine which both picked and cleaned the hops was
developed in England using a novel picking mechanism in which the bines were
propelled round the inside of a large drum past the picking fingers. It was not
so heavy as the German machine and it was claimed that its more compact
form and design allowed it to turn within the hop garden, thereby eliminating
the need for wide headlands (Catchpole, 1982). A prototype and one pro-
duction model were produced but the results were not entirely satisfactory and
production has now ceased.

4 .2 DRYING
Hop cones at harvest have a moisture content of nearly 80% and this needs to
be reduced to no more than 12% or the hops will not keep in good condition in
90 Harvesting
storage. Apart from the risk of the hops going mouldy, Maton (1982) has
recorded considerable losses of a-acid in hops dried to final moisture contents
of more than 11% while Henderson (1973) has reported evidence of spoilage
with moisture contents above 10.5%. To allow for variations within a batch of
dried hops it is safer to aim for a maximum of 10%. In continental Europe the
drying and packing of the hops is not completed on the farm where they are
only dried to a maximum of 14% and loose packed for movement to a
merchant's warehouse. At the warehouse several lots of the same variety and
similar quality are blended, re-dried, sulphured to improve their appearance
and baled. In other countries the whole procedure of drying and packing is
carried out on the farm.
Makovec et 01. (1978) described how in Czechoslovakia hops were, until the
19th century, dried with cold air in lofts and other such spaces. Houses built
with such drying lofts are still to be seen in Europe, notably in the town of
Spalt. These authors state that hot-air drying did not become the normal
method until the end of the 19th and beginning of the 20th centuries. This is
surprising since, in England, Scot (1576) described how to build and use an
oast for drying with hot air which was based upon 'such an Oste as they drie
their Hoppes upon at Poppering' and the use of hot air appears to have been
standard practice since then in England. As an appendix to his thesis Baker
(1976) includes An Account of Hopps by a Kentish Gentleman which was
probably written between 1707 and 1712. This account states that:

The best way of drying hopps is with a charcoal fire on an oast or kiln covered with a
hair cloth, of the same form and fashion which is used to dry malt on, which every
carpenter or bricklayer, in countries where hopps grow or malt is made, knows how to
build.

Makovec describes how the first kilns were very simple and consisted of a
single layer on which the hops were spread and states that this type survived in
the UK and Belgium whereas in Czechoslovakia this was replaced, as early as
1880 in some places, by several layers fitted with tilting, louvred floors so that
the hops could be dropped from one layer to the next as drying proceeded.
Such multitier kilns (Figure 4.9) became standard in central and eastern
Europe but have only more recently been introduced in England.
The air was at first heated by burning charcoal but replaced later by coal.
Since the hot air from the fire passed directly through the hops care had to be
taken that it was free from contaminants. The fire had to be carefully stoked to
produce the minimum of smoke while in England it was also necessary to use
coal that came from selected pits where the arsenic content was extremely low.
This followed the appointment of a Royal Commission in 1901 to inquire into
cases of arsenical poisoning in Lancashire although the investigation found
that the quantities of arsenic found in hops had in all cases been minute (Select
Committee, 1908).
Drying 91

Fan

8asculating
tier
Movable
tier
0
U"l
Pressure C')
<Xl
chamber
Warm air
duct
Chimney

Heat
exchanger
Burner

Figure 4.9 Layout of three-tiered kiln (Zeisig, 1970).

Contamination can be avoided by the use of heat exchangers which transfer

the heat from the exhaust air to fresh air. Although Burgess (1964) states that a
patent was granted as early as 1635 for a kiln in which the products of
combustion did not come in contact with the hops, heat exchangers were first
introduced on coal-fired kilns in America. Today most kilns are heated by oil
and it is especially important to ensure that the hops are not contaminated by
unburnt oil. In many countries heat exchangers avoid this risk but where they
are not used, great care has to be taken with the setting of the oil burner to
ensure efficient atomization of the fuel and sufficient air for complete
combustion of the oil.
In the older systems the air flow through the hops depended upon natural
draught and the traditional cowls of the English hop kilns turned with the wind
to maximize the air flow. Burgess (1964) quotes air flows measured in such
kilns as ranging from practically nil to a maximum of 19 ft/ min (0.1 m/ s). At a
later stage electric fans were installed and these made it possible to dry deeper
loads of hops and gave more control over the air flow through them.
There have in recent years been several investigations into ways of im-
proving the efficiency of hop drying. When oil was cheap a major consider-
ation was to reduce the labour requirement either by mechanization of the
operation or by reducing drying times so as to minimize overtime payments to
92 Harvesting

~_-+ Recirculation
duct

Pressure chamber
: : : : : : : : : ::-_-_-..:-: ::::-_-_ -":-:QI Dry hops

Figure 4.10 Layout of three-tier continuous drier (Zeisig, 1970).

the staff. Although shorter drying times could, in theory, allow growers to
increase their drying capacity without additional capital cost, in most cases this
would not happen because the time saved would be insufficient to allow for an
additional drying cycle before picking started again the next day.
Following sharp rises in oil costs however the emphasis swung towards
saving fuel, rather than labour. The most heat-efficient system would be one in
which, after passing through the hops, the exhaust air would always be close to
saturation. Single-tier batch drying is inefficient in the final stages because, as
drying proceeds, the quantity of moisture removed becomes less and the
exhaust air has only a relatively low moisture content. With multi-tier systems,
fresh green hops are loaded onto the top tier each time that dry hops are
removed from the bottom tier and this ensures that the exhaust air is at all
times more nearly saturated so that less heat is wasted. The capital costs of
such systems are, however, higher than those of single-tier systems.
A modification of the multi-tier system is the continuous drier in which the
hops pass through the plant on three tiers of moving belts, falling from the
ends of the upper and middle belts onto the one beneath (Figure 4.10). Driers
of this type were developed in Germany, Czechoslovakia and Yugoslavia. The
capital costs of such driers are again high and there can be rather more
difficulty in controlling the system so that the hops emerge with the desired
moisture content but they have the advantage that very little labour is
required.
Burgess worked for many years on the principles of drying and the results of
his and other work are summarized in his book (Burgess, 1964). Because the
time taken for the hops in a load to dry depends upon their position in the load
he introduced the concept of minimum time of drying as that required to dry
Drying 93
TABLE 4.1 Minimum times of drying at different temperatures. Mean values for a
range of air speeds. (Summarized from Burgess, 1937).

Maximum Minimum
temperature time
(OF) (min)

104 1072
122 532
140 329
149 265
158 237
176 123
194 82
212 56

an infinitely shallow layer of hops - which in practice meant a layer one cone
deep. The additional time required to dry loads of greater depth was found to
be proportional to the depth of the load and he called this the 'extra time'.
The relationship that he found between the temperature of the drying air and
the minimum time of drying is shown in Table 4.1. Air speed had little effect
upon the minimum time but the extra time was inversely proportional to the
air speed within the bed of hops (Burgess, 1937).
The thermal efficiency of drying can be greatly modified by adjustments to
the air speed and temperature. As the heated air passes through the hop bed
and evaporates some of the moisture in the cones, its temperature will fall and
its moisture content will increase. Its drying capacity will be greatest as it
passes through the hops at the bottom of the bed and least when it reaches the
top layer. If the input temperature is high and the air speed is low, so much
moisture may be extracted from the lower layers of hops that the air becomes
saturated before reaching the top and may then deposit moisture onto the
cooler hops at the top of the load causing them to be discoloured - commonly
referred to as 'stewed' or 'reeked'. For this reason it was traditional to
commence drying at a lower temperature and raise it once the risk of such
condensation was over.
Care must be taken that the temperature is not raised to the point where
damage is done to the hops. Burgess (1931) carried out trials in which the air
temperatures ranged from 40-100°C and found that the a-acid content of the
finished hops fell steadily as the air temperature increased. At 100° C the a-acid
level was little more than half that of the hops dried at 40° C. The colour of the
hops also deteriorated as the temperature was raised. In most cases the adverse
effect of the high temperatures on a-acid content continued in the subsequent
storage period.
94 Harvesting
The hops being dried do not, however, reach the same temperature as the air
with which they are in contact because they are cooled by the evaporation of
moisture from them. This cooling effect is greatest at the commencement of
drying when the hops contain the most moisture and the temperature
differential decreases as drying proceeds. It should therefore be possible to use
hotter air at the beginning of the drying cycle without heating the hops to the
point where damage occurs but this would also require higher airspeeds to
avoid reeking.
Zeisig (1970) showed that increasing the air speed through a 30 cm (12 in)
deep bed of hops at 60° C could only reduce drying time by about 20% and that
it had practically no effect once the moisture content of the hops had fallen
below 45%. He therefore carried out experiments using temperatures ranging
from 60-120°C (140-248° F) and air speeds from 0.28-1.27 m/s
(55-10 1 ft/ min) and found the drying time at the lowest temperature was 11.5
times as long as at the highest temperature and air speed. He realized that there
would be damage to the hops under some of these treatments and added a
curve to indicate the boundary of what he judged to be acceptable conditions.
A further problem was that as the air speeds were increased above 0.3 m/ s
(60 ft/min) the hops were lifted by the air stream and because of uneven
loading they would be blown around causing damage and loss of resin glands.
He therefore suggested a design for a three-tier belt drier in which the air
speeds would be high enough to lift and hold the hops to the underside of the
belt above them, the top belt being provided with a mesh screen above it to
restrain the hops. This required avoiding air speeds from 0.3-0.5 m/ s
(60-100 ft/min) since within this range the hops would not be stable on either
the belt below or above them. Moreover, he advocated that for a bed of hops
27 cm (11 in) deep the distance between the top of that bed and the belt above
it should be not more than 5-7 cm (2-2.75 in). By introducing a second air
stream at a higher temperature to the green hops on the first section of the top
belt he was able to recommend a two-stage regime with air for the green hops
at around 100°C (212°F) at a speed of 1.45 mls (285 ft/min) with the second
stage operating at about 75° C (167° F) and 0.9 m/ s (177 ft/ min). In this work
Zeisig was assessing the efficiency of the kiln in terms of drying time and thus
the quantity of hops that could be processed in a given time and did not include
any estimates of fuel consumption.
Maton (1982) dried hops at temperatures ranging from 50-75°C at two air
speeds - 0.2 and 0.1 m/s (44 and 22 ft/min.) and found that at the lower air
speed a-acid losses increased greatly as the temperature increased but at the
higher speed there was little loss of a-acid at any temperature.
Shea (1971) carried out high temperature drying runs which started with air
at 96°C and 0.6-0.8 mls (120-60 ft/min) which was reduced in two stages to
71°C at 0.4-0.5 m/s (80-100 ft/min). The following season he carried out
another series of experiments at temperatures between 60-100° C which were
Drying 95
held constant throughout drying, again using high air speeds (Shea, 1972). In
neither of these series was there any significant loss of a-acid although the
appearance of the samples was seriously affected.
In order to operate at these high air speeds it was necessary to restrain the
hops by means of a thin wire mesh laid on top of the bed otherwise the hops
would have entrained in the air stream. Bailey (1958) found that dried bracts
were moved by air speeds between 0.4 and 0.5 m/s (80-110 ft/min) while dry
whole hops moved at about 0.76 m/ s (150 ft/ min). In practice it is not possible
to operate with air speeds as high as this because uneven loading of the kiln
results in spots with lower resistance through which the air speed is greater
than average and these can become 'blow holes'. The problem is greater with
deep loads and towards the end of the drying cycle. Thompson et aZ. (1985)
have reported that hops can lift with air speeds as low as 0.15 m/ s (30 ft/ min)
but this figure is surprisingly low and the limit of 0.3 m/ s (60 ft/ min) suggested
by Henderson and Miller (1972) agrees with that recorded by Zeisig and is
more realistic.
The air speeds that are quoted in these experiments are the averages but the
speeds through different parts of the bed will differ as the loading can never be
completely uniform and the resistance to air flow will vary as a result. With
very shallow loads the resistance is so low that there can be only very small
differences across the bed. With deep loading there is much more resistance
with correspondingly greater variability.
The danger of blow holes developing as the hops dry and become less dense
is not, however, only due to uneven loading but may also result from the
shrinkage that takes place in the bed. This is very noticeable in the depth of the
bed but it also occurs in the horizontal plane although this is not so obvious.
One consequence of the lateral shrinkage is that a small gap develops between
the hops and the kiln wall thus the air speed increases in this area and a
significant amount of heat is wasted. This can be overcome by blanking off a
15-20 cm strip all round the edge of the floor. Cracks also tend to develop
within the bed itself and, as the air accelerates through these, the surrounding
hops begin to lift, the resistance to air flow decreases still further and blow
holes can develop.
Various workers have used computer modelling to calculate the most
efficient system for hop drying. Doe (1984) concluded that with a bed of 1.0 m
(3.3 ft) depth the optimum air speed would be 0.194 m/s (38 ft/min) at 75°C.
Kranzler et aZ. (1984) used a computer simulation to compare various drying
programmes and concluded that the most efficient would be MAT (Modified
Airflow and Temperature) which commenced with a temperature of 82°C and
airflow of 0.3 m/ s (60 ft/ min), these conditions being reduced to 70°C and
0.1 m/ s (20 ft/ min) as drying proceeded. They comment that this method
would require a more sophisticated control system as well as fan and furnace
modifications. The next most efficient system was one in which some of the
96 Harvesting

exhaust air was recirculated once the low-humidity air had broken through the
drying bed. They suggested that this would require additional ductwork and
fans but Ellis and Winch (1983) have reported that it will work successfully
with a cheap type of ducting leading from the kiln wall above the hop bed
down to the main fan. The difference in pressure between these two points is
sufficient to ensure recirculation and no additional fans are necessary.
The drier advocated by Zeisig introduced additional hot air to the green
hops so that they could be dried at a higher temperature than those with a
lower moisture content. A similar system is used on a few multi-tiered kilns. In
most cases, however, it is the driest hops that are exposed to the hottest air.
There could therefore be advantages in developing a system in which the hops
and the air move in the same direction instead of the usual counter-currenj
arrangement. This was one of the features of the prototype auto-continuous
drier, designed to achieve short drying times at high temperatures with the
minimum of labour, developed at Wye (Shea and Sykes, 1981). In spite of
some promise the project suffered from many technical problems and was
eventually abandoned.
Another development in recent years has been the conditioning of hops after
drying. When hops are unloaded from the drying kilns there are considerable
variations in their moisture content. Not only do the hops from the bottom of the
load have a lower average moisture content than those from the top but there are
also variations within the cones themselves. Whereas the bracts and bracteoles
(the 'petals') dry quickly, the central strigs are sheltered within the cones and dry
more slowly. Watson (1954) recorded moisture contents of 15% in the strigs of
Fuggle hops at the end of drying when the bracts contained only 4.5%.
Bracts with such low moisture contents are very brittle and shatter badly if
the hops are baled in this condition. If the dried hops are left in heaps on the
cooling floor there will be a redistribution of moisture between and within the
cones so that they reach a more uniform moisture content and will suffer less
damage when pressed up. There will, in addition, be some transfer of moisture
between the hops and the surrounding atmosphere, depending upon the
relative humidity of the air and the extent to which the hops are exposed to it.
Skilled workers could manipulate the conditions on the cooling floor to
some extent by covering or uncovering the heaps of hops or by opening and
closing windows at times of the day to suit the ambient conditions. This was
not only rather unreliable but the time required to achieve uniformity through
the load could run into days, requiring more cooling space than was generally
available. More rapid results can be achieved by blowing air at the correct RH
through the hops and this process is referred to as conditioning and was first
adopted commercially in Czechoslovakia (Vent and Makovec, 1970).
Henderson and Miller (1972) suggested using steam to bring the RH of
conditioning air to 72-4%, which would bring the moisture content of the hops
to 10-11 % while Henderson (1973) published results of experiments to
establish the equilibrium relationship between the relative humidity (RH) of
Drying 97
14

12

10

...c:
2 8
c:
o
u
Q)

...5 6
en
'0
~

I 1
o 10 20 30 40 50 60 70 80
Relative humidity ("10)
Figure 4.11 Equilibrium conditions for hops when drying, etc. (Henderson, 1973.
Courtesy of J. Agric. Eng. Res.)

the atmosphere and the moisture content of hops exposed to it (Figure 4.11).
His results suggested that a moisture content of 10.5% was the maximum that
would avoid spoilage and this would be in equilibrium with air at 69% RH.
When freshly dried hops are exposed to such air the bracts will mostly
absorb moisture while the strigs will generally lose it. Moreover, as with the
drying process the bracts will respond more quickly than the strigs and will
reach the equilibrium moisture content while the strigs are still too moist. If the
hops are baled in that condition there would be a serious risk of spoilage. If
conditioning was continued long enough the strigs would also reach the desired
moisture content but this would take too long. In practice it is generally
necessary to dry hops that are to be conditioned to a lower mean moisture
content than those which will be left to equilibrate on a cooling floor with
restricted exposure to the atmosphere. Conditioning has the advantages that
the hops can be ready for baling in a matter of one or two hours after drying,
without requiring cooling floor space, and the final moisture content can be
close to the intended limit so that there is the maximum weight of hops for sale.
The large-scale hop enterprises on the state farms of Eastern Europe were
especially suited to the introduction of mechanized harvesting and drying and
Vent and Macovec (1970) described the development of the picking machines
and multi-tiered drying and conditioning equipment that was developed in
Czechoslovakia.
98 Harvesting

Figure 4.12 American hop kiln with automatic loading (IHR, Wye).

Although the principles of hop drying have been well researched and the
fuel-saving benefits of multi-tier driers are well understood, they have not been
adopted everywhere, probably because of the capital cost while single-tier
operations can be operated very efficiently. This is particularly true in the USA
where the very large hop farms are concerned to achieve a very rapid
throughput with the minimum of labour.
Shea (1980) has described how some of the American ranches operate with
large, single-tier units comprising five or six separate 'floors' each measuring
13.4 m x 13.4 m(32 ft x 32 ft) (Figure 4.12). These are loaded automatically and
the even way in which the hops are spread makes it possible to dry deep loads
with high air speeds without risk of the blowing through less dense spots in the
load. It also results in more even drying so that the process does not have to be
extended to deal with dense patches that dry more slowly.
Loads are 0.76-1.0 m (30-40 in) deep and the air temperature is 60-65°C
(140-150°F). The hops are also unloaded mechanically by winding the hessian
floor cloth onto rollers so that the dried hops fall either onto a conveyor or
directly into a deep silo for cooling. Cooling times are not more than 24 hand
are sometimes only 2 h. The hops are frequently cooled on the kilns with
ambient air slightly heated by passage through the fan furnace unit. Since a
large volume of air is drawn through the furnace unit, the lightly constructed
walls of which retain little heat, the conditioning air soon resumes a relative
humidity high enough to condition the hops.
Drying 99
In Belgium, Maton (1982) developed a fully automatic three-tiered kiln
using an air speed of 19 m/ s (63 ft/min) at a temperature of 65°C which gives a
drying time of approximately 6 h. When the hops on the bottom tier are dry
the fan and burner are switched off, and switching on a timer then automati-
cally carries out the reloading sequence. The hops on the bottom tier are
dropped onto a mesh belt which then conveys them through the kiln wall to
another conveyor outside. The other two tiers are then dropped to the layer
below. A container outside the kiln is loaded by blowing hops from the storage
bin into it through a telescopic tube fitted with a distributor at the end. This
container, which also has a louvred floor, is then wheeled into the kiln and the
floor actuated to drop the hops in an even load onto the top floor of the kiln.
At the end of this programme, which takes about 13 min, the fan and burner
are switched on again.
Davey (1982) described the kilns that were in operation in Scottsdale in
Tasmania. This is a four-tiered kiln with another loading floor above which
can be ready filled with hops so that when required its solid, louvred floor can
be operated to drop the hops onto the top floor of the kiln.
The objective with this operation is to achieve good fuel economy by trying
to maintain the exhaust air completely saturated and to do this its RH is
monitored by a direct-reading recorder. In practice the air cannot be main-
tained at 100% RH constantly and the floors are changed when the RH has
dropped to 70% but the change from 100% RH to this level only takes 5 min or
less. The best depth of load for fuel efficiency and throughput has been found
to be 30 cm (12 in). The airspeed used is 0.5 m/s (100 ft/min) and the air
temperature is maintained at noc (162°F).
This kiln is probably unique in having a pelleting plant in line with it. The
hops are dried to approximately 6% moisture content and then moved to
conveyors on which they can be conditioned by blowing moist air through
them to bring them up to about 8.5% moisture content, thereby helping to
equalize the moisture contents of the strigs and petals to aid hammer-milling
the hops prior to pelleting.
In Czechoslovakia a great deal of work has been devoted to developing
drying equipment and Fric et al. (1985) have described the TPD-3-K model
which has a capacity of 1500 kg/h. It has a regulator which can control drying
periods of 4-9 h and conditioning times of 1-2 h. Drying is carried out at 50-
65°C (122-149° F) and the hops are dried to a moisture content of7% which is
brought back to II % by conditioning.
Hops are sometimes exposed to the fumes of burning sulphur or to S02 to
improve their appearance and possibly to improve their storage stability. It is
particularly effective in removing some of the brown discoloration of hops that
have been damaged by wind, pests or diseases.
In England and the USA the burning of sulphur in the plenum chamber of
hop kilns during drying was a standard practice. It was abandoned first in the
100 Harvesting

USA but was continued in England until quite recently even though it made
working in the kilns very unpleasant and it caused a lot of corrosion of metal
fittings and equipment. Growers were reluctant to refrain from using it in case
their hop samples appeared inferior to those of other growers who did use it.
Eventually the brewing industry produced evidence that sulphured hops were
inferior from their point of view and the practice was abandoned. In the
merchants' warehouses in Europe some sulphuring is still carried out when the
hops from the different farms are bulked, blended and redried.
In countries where the processing of hops is finished on the farm the dried
hops are pressed into 'pockets' or bales. Pockets are the traditional pack used
in England and to fill these it was necessary to fill the pocket with loose hops
and press these several times before the pocket was full and could be sewn up.
Today, many farms have installed bale presses with which a rectangular shaped
bale can be filled with only two charges. This speeds up the operation and the
bales take up less space in storage.
In Europe, where the hops are to be reprocessed by the merchant, they are
only lightly pressed into large sacks on the farm so that they can be unpacked
at the warehouse without too much breakage of the cones.
CHAPTER 5

Pests and diseases: historical

revIew

Hops are subject to attack by several pests and diseases and the most serious of
these, if not controlled, will reduce the crop to a level at which it is not worth
harvesting. The few cones that might be produced would probably be
themselves so discoloured that they would be unmarketable.
Because there was in England, for very many years, a tax on hops, there are
nearly complete records since 1711 of the quantities harvested while since 1807
the acreage has also been recorded. From the latter date therefore it is possible
to calculate the average yield for the country (except for 1862-5) (Table 5.1,
Figure 5.1). These figures demonstrate tremendous fluctuations in yield from
season to season. In 1825, for example, the average yield for the country was
only 2.69 zr per hectare while in the following season it was up to 27.72 zr. In
1854 there was another bad year with an average yield of 4.09 zr, again
followed by a very good year with 32.19 zr.
The unpredictability of the crop led during the 19th century, to the amount
of tax collected each year becoming a popular subject for gambling. A book
written by Lance in 1838 introduced the subject by saying: 'The gentlemen who
anticipate the amount of duty while the crop is growing, and make wagers on
what that amount will be, may find some assistance from the following table
... ' There followed two pages which include a table showing the amount of
duty that would be raised from a range of average yields with comments which
conclude as follows:
The speculations on the duty, and the spirit of gambling which it inculcates, and which
is so prevalent amongst the growers, dealers, and others, in the neighbourhood of hops,
is by some considered to have an injurious effect, and appears to be the most dangerous
attendant on hop growing, in nowise according with the sober meetings and habits of
British farmers, who are in general industrious thoughtful husbandmen.
The main pest and disease problems that Lance described were flea beetles,
aphids and mould (powdery mildew). From his and other accounts it is clear
that the aphids were the most important cause of the damage which led to the
great fluctuations in yield, though mould could also have serious effects. It is
therefore remarkable that the only reference by Reynolde Scot in 1576 to pests
or diseases is what appears to be a description of the flea beetle:
102 Pests and diseases: historical review

TABLE 5.1 English hop yields from 1807 to 1986. From 1946 the Hops Marketing
Board included headlands in the area recorded as being under hops. This reduced the
calculated yield by up to 10%.

Year zrjha Year zrjha Year zrjha

1807 13.6 1847 21.3 1887 18.1

1808 33.9 1848 21.8 1888 12.1
1809 8.5 1849 9.8 1889 21.6
1810 9.8 1850 27.9 1890 13.1
1811 21.1 1851 15.6 1891 19.6
1812 4.0 1852 24.9 1892 18.3
1813 17.3 1853 14.3 1893 18.1
1814 17.8 1854 4.0 1894 26.9
1815 15.1 1855 32.4 1895 23.6
1816 5.3 1856 22.8 1896 21.1
1817 7.3 1857 20.8 1897 20.3
1818 21.1 1858 24.9 1898 18.1
1819 24.6 1859 33.6 1899 32.1
1820 14.8 1860 5.5 1900 17.1
1821 17.6 1861 11.3 1901 31.9
1822 24.1 1862 no record 1902 16.3
1823 3.3 1863 no record 1903 22.1
1824 17.8 1864 no record 1904 14.8
1825 2.5 1865 no record 1905 35.7
1826 27.6 1866 13.3 1906 13.1
1827 14.6 1867 9.5 1907 20.8
1828 18.3 1868 18.3 1908 30.4
1829 3.5 1869 11.0 1909 16.6
1830 9.8 1870 29.1 1910 23.1
1831 19.1 1871 20.8 1911 24.9
1832 15.3 1872 25.9 1912 26.9
1833 16.6 1873 21.3 1913 17.8
1834 19.1 1874 11.5 1914 34.6
1835 22.6 1875 25.4 1915 18.3
1836 18.8 1876 15.8 1916 24.6
1837 16.3 1877 16.3 1917 33.4
1838 16.1 1878 24.6 1918 23.9
1839 20.3 1879 6.0 1919 28.1
1840 4.0 1880 16.6 1920 30.1
1841 16.6 1881 17.6 1921 23.9
1842 20.1 1882 4.5 1922 29.6
1843 16.1 1883 20.6 1923 23.3
1844 16.3 1884 15.3 1924 45.4
1845 17.1 1885 17.8 1925 33.9
1846 24.4 1886 27.9 1926 32.6
Pests and diseases: historical review 103
TABLE 5.1 Continued
1927 27.9 1947 32.9 1967 28.2
1928 25.6 1948 30.1 1968 27.9
1929 37.7 1949 28.4 1969 31.5
1930 32.6 1950 41.7 1970 34.7
1931 21.8 1951 35.9 1971 32.7
1932 28.6 1952 31.9 1972 26.0
1933 32.1 1953 31.1 1973 30.9
1934 36.1 1954 29.6 1974 31.2
1935 34.1 1955 31.6 1975 25.8
1936 34.6 1956 23.1 1976 26.9
1937 32.6 1957 32.9 1977 24.4
1938 34.9 1958 35.9 1978 32.1
1939 38.4 1959 27.4 1979 36.2
1940 36.4 1960 31.1 1980 34.1
1941 36.4 1961 25.9 1981 32.2
1942 35.7 1962 32.9 1982 35.1
1943 37.4 1963 33.1 1983 30.3
1944 32.6 1964 30.4 1984 31.0
1945 35.4 1965 31.4 1985 27.4
1946 29.4* 1966 28.2 1986 24.0

* Change to 'board acres'

The Hoppe that lykes not his entertaynment, namely his seate, his grounde, his keeper,
his dung, or the manner of his setting, commeth up greene and small in stalke, thicke
and rough in leaves, very like unto a Nettle, which will be commonly devoured, or much
bitten with a little black flie, who also will do harme unto good Hoppes where the
Garden standeth bleake, or the Hoppe springeth rath, but be not discomforted
herewith, for the heate of the Summer will reforme this matter, and the later springs will
be little annoyed wyth this flie, who (though she leave the leafe as full of holes as a
nettle) yet she seldome proceedeth to the utter destruction of the Hoppe.

With such a detailed description of the flea beetle it is most unlikely that he
would not also have commented upon aphids or mould if they had been
prevalent at that time so it is interesting to enquire when these were first
reported. An invaluable source of references for this is Parker (1934) who
provides an exhaustive historical review.
The earliest account of either of these problems appears to be in a book by
Worlidge (1669) who 'said of hops: 'They are the most of any plant that grows
subjected to the various mutations of the Ayr, from the time of their first
springing, till they are ready to be gathered; over-much Drought, or wet,
spoyles them, mill-dews also sometimes totally destroys them.' Although this is
a clear reference to powdery mildew there is still no reference to anything
resembling aphids.
104 Pests and diseases: historical review

zr/ha

Years

Figure 5.1 English hop yields from 1807 to 1986. (See Table 5.1.)

Leonard Meager (1697) referred to the attacks of insects: 'If you perceive the
Flies, or any other Insects that bite and affect them, sprinkle the Vines with
Water wherein Wormwood has been boiled, and it will preserve them.' This
appears to be the first reference to control measures but is not very helpful in
identifying the insects involved which are probably, again, flea beetles.
The earliest recognizable reference to aphids in England comes from Ellis
(1750) who described the problems affecting the hop as follows:
Lice, Fly, Worm, Blight and the Mould or Dwindle. Lice are bred by Mill or
Honeydews or Fogs. At a greater age its called the Collier Fly because it turns black and
will then feed on the Sap of the Vines and Hops ... Put Stone Lime on each Hill and
the first rain will cause a fume that will do Service ... Or you may make Use of the
Dutch Squirt that casts its Water twenty Feet high and thereby washes off the Breed of
Lice, Lady Bird and Slug or Snail that are bred and nourished by the Honey-dews.
A few years later, Mills (1763) described a disastrous attack of mould (or
fen) in 1723 when the most flourishing and promising hops were all infected.
He quoted the authors of the Journal Oeconomique on this disease:
... no other remedy from nature against this mischance, except rain sufficient to wash
the plant, and clear it entirely from this fatal dew: but as rain seldom comes quite
seasonably to the relief of the plant thus affected, artificial means have been sought for
insuring it against this accident. Some have surrounded their hop-grounds with hogs
dung; others have employed persons to go through the ground with vessels full of beech
Pests and diseases: historical review 105
ashes, and to throw them upon the hops while the mildew was falling; and both sides
... have even proceeded so far, as that each affirm their's to be the only remedy. Those
who use hog dung say, that the ashes may probably hinder the action of the dew upon
the plant; but that they must, at the same time, stop up its pores, and deprive the soil of
its humidity ... The partisans of the ashes say, that they cannot comprehend how hog
dung laid around the hop-ground in the spring, should preserve such virtue as to
destroy the bad quality of this mildew in the summer.

He also quoted an account of the 1725 season by Mr Austen of Canterbury,

... 'who was a very great planter and an accurate observer'. According to this
account the hops were infested with flies towards the end of April and, since
this would be too early for migrant aphids, it probably refers to flea beetle. But
then about the 20 May: ... 'the flies appeared upon the leaves of the
forwardest vines ... About the middle of June the flies increased, yet not so as
to endanger the crop; but in distant plantations they were exceedingly
multiplied' and this is most probably a reference to the hop aphid.
Mills referred elsewhere to honeydew which is a common feature, from this
time onwards, in reports on aphids and its seems unlikely that earlier writers
would not also have noted it if aphids had been a pest of hops in their time. It
does seem likely, therefore, that they did not become a problem until sometime
in the 17th or early 18th centuries.
While there is no definite date for the first appearance of hop aphids in
Europe, their arrival in North America appears to have been accurately
recorded. Parker (1913) stated that they first appeared in New York in 1863
and that they reached the Pacific coast in 1890. It is very unlikely that live
aphids would have been carried to America on fresh hop material and it is
much more likely that they were introduced as eggs on shoots of Prunus which
is their winter host.
Honeydew was the subject of considerable disagreement in the 19th century,
some maintaining that it was excreted by aphids, others asserting that this was
not true. Lance, for example, said that:
The strange notion is entertained, that this matter is the excrement of aphides when in
reality it is a disease of the vegetable occasioned by their attacks ... Bonnet has asserted
that the juice emitted by this vine fretter, plasters over the upper side of the leaf, and this
idea has been taken up by all writers on entymology since his time, without examining,
or reflecting on the subject.

He then proceeded to argue that 'juice' from the aphids could not be
distributed in the way that the honeydew was but, not surprisingly, he quoted
no experimental evidence to support his view. He did however make more
worthwhile comments upon the activities of the ladybird as a predator of the
aphids (describing it as 'the principal antidote) and there are other dramatic
accounts at that time of how the appearance of ladybird larvae (or 'niggers' as
they were commonly called) saved the day when the crop was apparently
doomed by heavy infestations of the aphids.
106 Pests and diseases: historical review
In the early 19th century efforts were made to control aphids but these were
not very effective. Lance again: 'When once the leaves of hops are infected by
aphides, it is almost useless to attempt a cure.' But a little later he said, 'Means
have been tried, and sometimes with good effect, to destroy them in the
progress of their growth, by fires and fumigating with sulphur and tobacco,
which, when burnt immediately under them, is sure to occasion their destruc-
tion.'
It was not until spraying with a solution of soft soap was started about 1865,
to which quassia was added a few years later, that any worthwhile control was
achieved and such spraying did not become general until about 1883 (Burgess,
1964). The subsequent introduction of nicotine as an insecticide and the
widespread adoption of sulphur to control powdery mildew resulted in a
marked improvement in yields. The last really bad crop was in 1882 with a
yield of 4.5 zrjha. Since then the yield has never fallen below 13 zr whereas this
had happened 19 times in the preceding 72 years.
Cousins (1895) described the use of soft soap and quassia but advocated as a
better alternative the use of 'Paranaph' Soap which was a mixture of paraffin,
naphthalene and soft soap which had been patented. The details of the mixture
were not given and growers were warned that they needed a demonstration of
how to make the mixture as it could cause serious scorching if prepared
incorrectly.
Some early writers showed a remarkable appreciation of the way that
spraying to destroy aphids could upset the ecological balance and they
questioned whether the destruction of natural predators did not outweigh the
direct benefit achieved by spraying. Any attempt to produce hops today without
the use of insecticides results in even greater losses than those of the pres praying
era, presumably because the widespread use of non-selective insecticides over
the past 100 years has seriously depleted the predator popUlations.
Blattny and Osvald (1950) described the importance of predators as follows:
Nature, which is superior to us, looks after both the increase and decrease of aphids.
She enables the species to survive and she also created the natural enemies. Those
predators do not permit the aphids to have control over hop plants and so over the hop
growers. These natural enemies are helpmates in the struggle with aphids. The most
important of them are the seven-spotted and the two-spotted ladybirds. We can say that
without ladybirds and the other natural enemies we should have to spray against aphids
every year.
It was not until the latter part of the 19th century that the life history ofthe
aphid, overwintering on Prunus and migrating to hops in the summer, was
understood. Ormerod (1890) says:
It does not seem now to be open to doubt that a great part of the yearly attack of Hop
Aphis, or 'Fly', comes on the wing from Sloe, Damson, or plants of the Plum tribe. This
was long ago stated by German entomologists, also laid down by at least some of our
Hop-growers ... In 1887 Prof Riley set the matter of migration from Plum to Hop
Pests and diseases: historical review 107
beyond doubt by his observations of which a part was read before our own British
Association.

It is understandable that the available records do not indicate when the long-
established pest and disease problems first attacked the crop. In many cases the
knowledge available at the time was insufficient to understand the cause of the
symptoms observed. Even today the cause of some abnormalities is not known
while, in the case of prunus necrotic ringspot virus, it is only recent research
that has indicated the existence of a symptomless disease that seriously affects
production.
In the 1920s, however, a new problem arose in Europe which was
immediately identified and reported. This was downy mildew which had first
been recorded in Japan in 1905 and later on wild hops in the USA in 1909. It
was first observed and identified in Europe by Salmon and Wormald (1923)
who found it in October 1920 at Wye on young plants raised from Italian seed.
It could not be found in 1921, which was a dry year, in spite of a careful search,
but 1922 was a wet year and it reappeared in the same garden on a number of
plants in September. In 1924 it was reported as being endemic on wild hops in
the south of England (Salmon and Ware, 1924) and by the autumn there was a
heavy epidemic on cultivated and wild hops in several southern counties
(Salmon and Ware, 1925). Because it occurred on wild hops at localities far
from any known source of infection, the authors thought this indicated that the
disease was indigenous and not introduced from abroad.
In 1924 there was a severe outbreak at Wiirttemberg in Germany (Lang,
1925) and it was also present in Belgium (Marchal and Verplancke, 1926). In
1925 it was severe in Belgium (Lindemans, 1925), was widespread in France
(Rials, 1926), made its first appearance in Bavaria (Korff, 1925) and caused
some browning of cones in Czechoslovakia (Blattny, 1925). It was also
reported in Yugoslavia from both the Vojvodina (Vrbovsky, 1928) and Savinja
Valley (Terzan, 1927a) regions. In Germany the disease is referred to as
'Falscher Mehltau' (False Mildew) to distinguish it from the old powdery
mildew which is now called 'Echter Mehltau' (True Mildew).
By 1926 it had been discovered in Italy near Perugia (Curzi, 1926) in Poland
(Siemaszko, 1927) and on wild hops in north-west Russia (Naoum off, 1928)
while in Bavaria it devastated 60% of the crop (Sgd, 1927).
In 1927 it was reported on wild hops in Sweden (Juel, 1928) while in Russia
it was found chiefly on wild hops in several widely separated localities,
including the Caucasus ' ... in virgin forest, scarcely penetrated by man'. In
1928 a systematic search revealed its presence in several districts of West
Prussia (Jaczewski, 1929).
This apparent spread throughout Europe led Salmon and Ware (1926) to
change their mind about its origin and to conclude that it probably had been
introduced from abroad and was not indigenous. Although this is now
108 Pests and diseases: historical review
generally accepted, the rate of spread over such wide areas is quite remarkable,
especially for an organism whose spores are only short-lived (section 7.1.2). It
is interesting to note too that many of the records report it first on wild hops
far from areas where the crop was cultivated.
The original account of the disease from Japan (Miyabe and Takahashi,
1905-6), as quoted in Salmon and Wormald (1923), states that it was first
noted on cultivated hops at the Hokkaido Experimental Station in the early
summer and that by mid-June it had spread to an alarming extent through the
field. In view of this rapid spread it is surprising that they inferred that the
disease had been present there for many years without attracting attention. It
was also found on wild hops in Japan and Miyabe concluded that the mildew
was, without doubt, indigenous to the country and had found the cultivated
hops introduced from America and Europe to be a more congenial host
(Salmon and Wormald, 1923).
The first American record on wild hops in Wisconsin in 1909 was by Davis
(1910) who found it again in 1920 (letter quoted by Salmon and Wormald, 1923)
and did not doubt that it was indigenous there. It was not recorded on cultivated
hops anywhere in North America until 1928 when Salmon and Ware (1929)
described how it was prevalent on cultivated hops at Sardis, British Columbia in
1928 and surmised that it was present in 1927 and possibly earlier. Since the
disease was unknown in hop gardens in the USA they assumed that it had been
introduced on setts imported from Europe. In the same year it was reported for
the first time on commercial hops in the USA on a Bavarian hop farm in Otsego
County, New York where ' ... it was evidently of recent introduction' (Dunn,
1929). In 1929 it appeared in the Willamette Valley, Oregon (Barrs, 1930) and in
the same year Heald (1930) reported it in western Washington while it had
spread to California by 1934. Although found on ditchside hops in eastern
Washington in 1937 it did not affect commercial hops in the Yakima Valley
until ' ... the calamitous year of 1947' (Romanko, 1964a).
This evidence suggests that the disease was so uncommon in America before
1920 that it is very unlikely to have been introduced in Europe from there
(although Japan remains as a possible source). On the other hand, it was quite
possibly introduced into American commercial production from Europe in a
more virulent form than the indigenous disease reported by Davis.
A curious fact is that Salmon observed the disease in Wye in 1920 but did
not report this until 1923. In the meantime the disease had been included in the
Destructive Insects and Pests Order of 1921 on the grounds that it had been
introduced from Japan to the USA and was there spreading but that it had not
yet reached Britain. It had, in fact, reached Britain and there appears to be no
evidence of it having been introduced into America from Japan nor of it
spreading in the USA at that time.
Several of the sources quoted above suggest that the pathogen was present in
Europe prior to the first reports. Rials thought that it had probably existed in
Pests and diseases: historical review 109
the south of France for some years. Because of its widespread distribution and
the fact that there had been no hop plants introduced, laczewski believed that
it had a ubiquitous origin of long-standing on wild hops in Russia and that the
various epidemics were brought about by unascertained ecological conditions.
Curzi noted that the first report at Wye was on plants raised from Italian seed
and suggested that the disease had existed previously in Europe. Moreover
Baudys (1927) thought that it had been present for some time in Cze-
choslovakia without attracting attention and that it was definitely present on
wild hops near Weisskirchen (Moravia) in 1924.
Marchal and Verplancke considered that it could have been introduced to
Belgium on plants imported from England but nowhere else was movement of
hop plants suggested as a possible source of infection while Lang states that
there had been none to account for the first outbreak in Germany.
The most remarkable report of all is that the first serious occurrence of the
disease was as early as 1894 in Vojvodina at the manor of Graf Kotec in Old
Futog when it reduced the yield of the hops by a half (Terzan, 1927a, b, 1928).
If this report could be confirmed it would certainly support the suggestions
that the pathogen was indigenous and it would then be necessary to explain
what could have led to the sudden epidemic throughout Europe. Romanko
(1964a) suggested as several possibilities: the mutation of existing mild forms,
hybridization of mild forms brought together by planting imported rootstocks,
a shifting of the general world climate or changes in production practices.
Hybridization would only lead to new forms if the resulting oospores were able
to germinate and infect hop plants. As described in Chapter 7, oospore
germination appears to be very infrequent. Could it be that hybridization had
occurred throughout Europe before 1920 but that it was not until then that
there was a year, or years, that were particularly favourable for oospore
germination?
Once established, the disease caused dramatic reductions in yield in
Germany in some seasons, Zattler (1928) reporting that in 1926 the average
yield was only 2.15 zr/ha but that with an energetic spraying programme in
1927 it increased to 8.2 zr/ha. Magie (1942) reported that American growers in
New York State who did not spray lost at least one-third of their crop in the
average year. The damage caused by downy mildew was largely responsible for
a shift in hop cultivation away from the Eastern States to the drier conditions
of Washington and Idaho in the west (Skotland, 1961) but powdery mildew
was also a contributing factor. Powdery mildew was first reported in New
York State in 1909 and then gained steadily in importance (Blodgett, 1915).
Whereas both diseases caused serious losses in the eastern states, downy
mildew is much less of a problem in the west while powdery mildew does not
occur at all.
In England, Beard and Derbyshire (1957) recorded a marked decline in
yields of a garden of Golding hops over a seven year period during which most
110 Pests and diseases: historical review
of the rootstocks became infected. Coley-Smith and Beard (1962) working in
the same garden showed that the yield of infected hills was 27.7% lower than
healthy plants. It is surprising, therefore, that the English yield figures do not
indicate any general decline following the appearance of the disease in the
1920s. This may be because the other varieties being grown were not as
susceptible as the Goldings. Fuggles, in particular, were noted as being highly
resistant, not only in Englan·d but also in Europe where they were called
Goldings, a confusing nomenclature that persists until now in Yugoslavia.
Although downy mildew has spread through all the hop growing areas of the
northern hemisphere (with the possible exception of some of the Chinese
regions), and also into South America, it has been excluded from South Africa
and Australasia by strict quarantine measures. Salmon and Ware (1931) give a
fascinating account of how these quarantine precautions were initiated in
Australia:
Hearing that a consignment of hop roots from this country was being sent (with the
sanction of the Ministry of Agriculture and Fisheries) to Australia, we wrote to the
official mycologist and to Dr E.1. Butler, Director of the Imperial Bureau of Mycology
in this country, pointing out the great danger of this importation. Thanks to the prompt
help given by Dr Butler, and to the action taken locally by the mycologist and
agricultural authorities, a Proclamation was issued prohibiting the importation of hop
plants into the Continent of Australia. Although the hops were landed, the Proclama-
tion was just in time; the cases were never opened, and were eventually destroyed. It is
very possible that the introduction of Downy Mildew into Australia has thus been
prevented. It is satisfactory to know that New Zealand has recently legislated to the
same effect.

Because Professor Salmon believed that he was responsible for the introduc-
tion of downy mildew to Europe on material imported from the USA, he was
only too conscious of the potential danger to the Australian hop growing
industry of that consignment of planting material.
Whereas in most countries the response to the outbreaks of downy mildew
was to concentrate upon chemical control, in Germany there was also a more
positive approach with the establishment at Hull of a research institute having
the primary objective of breeding varieties that were resistant to the disease
(section 7.l.6).
Although the history of verticillium wilt also seems to be well documented
there are increasing doubts about how this disease developed in England. It
was first recorded in England near Tonbridge in 1924 (Harris, 1927) but it
seems that this was merely the first time it had been identified and that it was
already widespread in the hop growing areas but not sufficiently serious to
attract attention. In 1938 it was noted that there were two types of attack
which were either mild ('fluctuating') or severe ('progressive') in character
(Keyworth, 1939). This was later shown to be due to mild and virulent strains
of the fungus but more recent work has shown that there is now a range of
Pests and diseases: historical review 111
strains of varying pathogenicity and it is thought possible that the original
mild strain has changed to a more pathogenic form on more than one
occasion.
No chemical control measures are available for the control of this disease
and growers have to exercise careful hygiene to prevent its spread or, if that
fails, plant cultivars that are resistant to it. Had it not been for the successful
breeding of such cultivars, the virulent strains of the fungus would have
devastated the English hop industry. Even so, there have been considerable
losses in production though these have been due less to the direct damage to
the hop gardens, which have usually been grubbed before too many plants
were affected, than to losses caused by replanting with new cultivars and the
delay before these were in full production. Many gardens in south-east
England have had to be replanted more than once as the first of the resistant
cultivars were superseded by newer selections that were either more acceptable
to brewers, or able to withstand the attacks of the more virulent strains of the
pathogen which developed.
In 1952 verticillium wilt was first recorded in the Hallertau district of
Germany where it spread rapidly through the whole of the area and
subsequently into the Jura and Hersbruck districts (Kohlmann and Kastner,
1975). The strains found there are not as virulent as the worst of the English
types and the impact has not been so serious as in England. Even so there have
had to be great changes in the types of hop that could be grown in the affected
areas.
Although Australia and New Zealand have escaped most of the diseases that
are found in the northern hemisphere, they have suffered considerable
problems with black root rot caused by Phytophthora citricola. No effective
control measures are available against this disease and it, too, has been
countered by breeding programmes that have produced resistant cultivars.
Considerable confusion surrounds the history of the virus diseases of hops
and this is not surprising since the very existence of viruses was for so long
unrecognized while the understanding of individual virus problems has
frequently involved years of research.
The best example of this confusion is to be found in nettlehead disease which
was first recognizably described by Percival (1895) who gave it the alternative
name of'skinkly disease' and attributed the cause to damage by eelworms. It is
considered by some people that Reynolde Scot's 'unkindly Hoppe' that
'commeth up greene and small in stalke, thicke and rough in leaves, very like
unto a Nettle.. .' is a description of nettlehead disease but it could also be a
description of shoots that had gone dormant (see section 1.6), especially as
Scot added that 'the heate of the Summer will reform this matter. . .'. In 1925
Duffield found that eelworm was not the cause of nettlehead but it was not
until some 40 years later that it was shown to be associated with infection by
arabis mosaic virus (Bock, 1966).
112 Pests and diseases: historical review
There was further confusion between nettlehead and the 'curling disease'
(called Krauselkrankheit in Germany and kaderavost in Czechoslovakia)
which were at one time thought to be the same. More recent work has
indicated that the curling disease is not due to a virus but is a symptom of zinc
deficiency (Schmidt et al., 1973).
Nettlehead causes severe damage - reducing the yield of infected plants by
over 75% (Legg, 1959b) - and it became so widespread that it was difficult to
find any gardens that were free of it (Keyworth, 1945) and some growers found
it necessary to abandon hop growing completely (Ogilvie, 1939). It must
therefore have had a considerable effect upon the national yield figures. The
spread of Verticillium wilt was of some benefit to this problem since the
extensive replanting was mostly done with virus-free stocks and this con-
tributed to a decline in the incidence of nettlehead.
There was also some confusion between nettlehead and hop mosaic which
was first described by Salmon (1923). The symptoms of mosaic and nettlehead
are not dissimilar and Salmon's old records can now be interpreted to show
that as far back as 1906 he was describing mosaic-infected plants as suffering
from nettlehead. MacKenzie et al. (1929) recorded that cuttings imported from
Germany by a grower between 1904-8 appeared to have been carriers of the
virus and to have caused widespread infection in the Golding hops amongst
which they were planted. It is probable that Salmon introduced the virus to
Wye in a similar way but because the infected plants were grown in a garden
containing both susceptible and 'carrier' varieties the source was not as
apparent as it was in the grower's garden. It seems certain that the virus was
not present in England previously but it is not possible to estimate how widely
distributed it was on the continent because few of the cultivars grown there are
as susceptible as the English Goldings.
The methods used to control the virus diseases are described in section 8.4.5
but the most important precaution has been to ensure that only virus-free
material is used when planting up a new hop garden. To achieve this several
countries have organized supervised propagation of virus-tested stocks. The
earliest was the 'A-Plus' Certificate set up in England in 1954. At that time the
Ministry of Agriculture was operating, for other crops, an 'SS' (Special Stock)
Certificate which indicated that the stocks had been tested and shown to be
virus-free. In 1954 there was insufficient knowledge for such a standard to be
applied to hops. Only nettlehead and mosaic could be tested for with any
confidence so the A-Plus Certificate was introduced as a compromise that
ensured that growers could obtain the best material that could be produced in
the existing state of knowledge (Jary, 1955). In America, Skotland organized
the propagation of selected clones of the Cluster hops and more recently a
similar scheme has been set up in the Federal Republic of Germany. Research
into hop viruses has also been actively pursued in the German Democratic
Republic in order to ensure that planting material is of the highest quality.
Pests and diseases: historical review 113
In view of the new pest and disease problems with which growers have been
faced it is a tribute to the efforts of pathologists, hop breeders and the chemical
industry that it has been possible not only to maintain but to see increases in
hop yields. The weight of hops harvested is, of course, only one part of the
story. When the enhanced content of brewing material in modern cultivars is
taken into account it is clear that there has, as described in Chapter 8, been a
massive increase in production.
CHAPTER 6

Pests

6.1 DAMSON-HOP APHID: Phorodon humuli

Description
This pest is a problem in all the hop growing districts of the Northern
Hemisphere except, according to reports, some areas in China. If uncontrolled
it is capable of completely destroying the crop and even when control measures
are taken they may not be entirely successful. The aphids feed by inserting their
long stylets into the phloem strands of the leaf veins and can severely weaken
the plant and cause defoliation. The main problem arises after cone formation
because at that stage the insects feed within the cone and present a much more
difficult target for spraying. It is at this time, also, that they cause the most
damage. Even quite light infestations can seriously affect the commercial value
of the hops because sooty mould grows on the honeydew produced by the
aphids. This shows up when the dried hops are valued and causes them to be
down-graded. More heavily invested cones turn brown and limp and many of
them will be lost on the picking machine. Those that survive into the final
sample can make it unsaleable.
The aphid is also of importance as a vector of virus diseases. It has been
especially so in England where the Golding cultivars are unusual in being
susceptible to Hop Mosaic Virus whereas other cultivars are symptomless
carriers. It is important that the Goldings are grown sufficiently far-removed
from other hops to prevent the aphids transmitting the disease from one to the
other. The aphid is also reported to be a vector of American Hop Latent Virus.
The aphid overwinters on various species of Prunus, principally on sloe (P.
spinosa), but also on damsons (P. insititia) and plums (P. domestica). The eggs
are laid in the axils of the buds in the autumn (Figure 6.1) and hatch out in the
spring to produce wingless females (virginoparae) that feed on the Prunus
leaves and reproduce asexually, giving birth to living young. After one or two
generations they then produce winged females (alatae, Figure 6.2) which
migrate to the hop only when the flight threshold temperature of 13° C is
exceeded (Muir, 1968). Soon after arrival the migrants commence producing
the first of several generations of wingless, asexual aphids (apterae) which
116 Pests

Figure 6.1 Eggs of the damson-hop aphid overwintering on buds of Prunus sp. (IRR,
East Mailing).

build up in large numbers throughout the summer unless controlled (Figure

6.3).
Most aphids are attracted more to objects that are yellow than to those that
are green and aphid traps are, for that reason, usually coloured yellow. Muir
(unpublished) found that P. humuli was an apparent exception as he trapped
more of them in green than in yellow traps. Campbell (1984) compared the
numbers of aphids alighting on the normal green form of the cultivar Brewer's
Gold and a yellow mutant of the same cultivar. Contrary to expectations,
about 1.5 times more aphids settled on the yellow-leaved form and laboratory
experiments showed that this was not due to differences in palatability.
A report from East MaIling (Anon, 1969a) describes how practically all the
aphids which settle to feed and reproduce are found on the rapidly expanding
leaves at the top of the bine. They may alight directly on these leaves or else on
the older ones lower down and walk up to the apex. When a barrier was placed
below the fifth node, the numbers of alatae removed from the tip each day was
50-70% less than on control plants. The numbers were further reduced by half
on plants from which all leaves below the fifth node were removed.
Campbell (1977a) found that 100% more migrants were found on strings
with two bines than on those only having one bine and that there was a positive
Damson-hop aphid: Phorodon humuli 117

Figure 6.2 Winged migrant female of damson-hop aphid (Crown copyright).

correlation between bine height and aphid numbers. Plants next to poles were
more heavily infested than those not at poles and migrants were more
abundant on leeward than windward orientated strings. By comparing the
numbers of alatae on hops with the numbers caught in a nearby aphid trap he
estimated that most infestation was by aphids from sources within one hour's
flying time.
The migrant alatae may not settle on the hop plant on which they first alight
but any subsequent flights are short and will only be to plants close to the
original one. At East MaIling, workers marked newly arrived alatae and found
that half a day later only 20-40% remained on the original plant (Anon, 1968).
This report made no reference to the cultivar involved and it is probable that
varietal preference is important in relation to the amount of post-arrival
movement that takes place. Campbell (l977a) found that fewer migrants
settled on the variety Tolhurst than on Northern Brewer and that Fuggle was
intermediate. In this experiment the aphids had the possibility to move from an
unpopular cultivar to one that they preferred but most commercial gardens are
planted with a single variety and though the amount of movement from plant
to plant may reflect the aphids' preference they will finally be forced to settle
on that variety and in such circumstances preference may have little effect
upon the migrant numbers.
118 Pests

Figure 6.3 Leaf of unsprayed hop plant showing heavy infestation of apterae of the
damson-hop aphid.

Differences in post-arrival movement are probably important in relation to

the role of aphids as vectors of virus diseases since movements are likely to be
more frequent in gardens planted with unpopular cultivars and this would
increase the rate of spread of viruses (section 8.1). Most workers agree that the
aphids on the hop produce no winged forms (alienicolae) that will move to
other hops and Campbell (1984/5) concluded that, even if the few claims that
such a morph has been observed could be confirmed, it would be such an
infrequent event that it was of no economic significance. It is, therefore, only
the aphids migrating from Prunus that are important as virus vectors.
Although preference may not, in practice, affect the size of the migrant
population that settles on a cultivar it would be related to the subsequent
multiplication rate if preference reflected the suitability of each cultivar as a
food source for the aphids. Such a relationship is indicated by the results of
Campbell (1983) who raised apterae on the same three cultivars and found that
their capacity to increase was greatest on Northern Brewer and least on
Tolhurst except at the lowest temperature when the positions of Fuggle and
Tolhurst were reversed.
The migration to the hop commences in late Mayor early June but some
wingless forms continue to be produced on the Prunus and these remain as a
Damson-hop aphid: Phorodon humuli 119
source of winged migrants until as late as July or early August. Campbell et al.,
(1987) found that aphid populations persisted longer on cultivated plums than
on hedgerow blackthorn or myrobalan. In September and October migration
back to the Prunus winter host takes place. The wingless viviparous females
switch from producing more of their kind to producing winged migratory
females when the light period falls below about 13.5 hr / day although the
critical daylength varies inversely with the ambient temperature (Campbell,
personal communication).
The first winged forms to move from the hops (gynoparae) produce wingless
females (oviparae) on the Prunus and these are then ready to mate with winged
males which follow from the hops rather later. The males locate the oviparae
by means of a pheromone released from glands in their hind legs. The fertilized
females then lay the overwintering eggs on the Prunus.
It is of considerable importance to growers to know when the migration
onto the hops will start and, more importantly, will end since the effective
timing of control measures depends upon this. Extensive data on the migration
periods have become available from a series of suction traps sited in different
parts of Britain for the Rothamsted Insect Survey. Thomas et. al. (1983)
investigated the relationship between the beginning and end of migration over
a number of years with various weather parameters. They found that the
commencement of migration was earlier when the maximum temperatures in
late March and early April were higher and using this single parameter they
were able to predict the starting date with considerable accuracy. Although it
appeared that there was some association between dry weather in mid-January
and, more particularly, early April, the prediction was not improved by
including this factor. The end of migration was accelerated by high temper-
atures and sunshine hours in June but predictions based on this were only
moderately successful.
According to Kriz (1966) the end of the migration does not depend upon
the completion of a particular number of generations on Prunus since the
number varies from year to year. It is affected by the ratio of total nitrogen to
carbohydrate in the leaves; only the young ones being suitable in this re-
spect for the continued production of apterae. As the production of young
leaves diminishes and the diet changes, alatae are formed which migrate to
the hop.
The annual migration to hops varies, not only in the time at which it starts
and finishes, but also in the numbers of aphids that migrate, and the intensity
of the infestation makes a great deal of difference to the control measures that
are required. Control depends upon the use of chemical pesticides but in recent
years there has been considerable interest in developing biological control.
Although this has been primarily a response to difficulties with purely chemical
methods, it has also been encouraged by increasingly strict regulations about
pesticide residues in the hop cones.
120 Pests

6.1.2 Chemical control

The earliest materials used to control the aphids by spraying were soft soap,
quassia and nicotine. Of these nicotine was the most effective and became the
standard treatment until TEPP and schradan were introduced in 1946 followed
by parathion in 1950 and demeton-S-methyl in 1952. There was then a period
of about ten years during which control was based almost exclusively upon the
organo-phosphorus (OP) insecticides demeton, demeton-S-methyl, parathion
and thiometon applied as foliar sprays, and dimefox applied as a drench to the
base of the bine (its high mammalian toxicity made it unsafe to use as a spray).
These materials were so effective that for several years control of the aphids did
not present a problem.
In the early 1960s, however, the first signs appeared of aphids that were
resistant to these chemicals. Hrdy and Zeleny (1966) reported resistance to
thiometon in Czechoslovakia while Muir (1979) found that by 1966 aphids
from hop gardens in Kent were ten-times more resistant to demeton-S-methyl
than those collected from the north of England where they were remote from
spraying. By 1974 this level of resistance had doubled and by 1976 it had
increased to 50 times that of the susceptible stock. Since the first signs of
resistance to these aphicides, there has been a continuing problem of new
chemicals being introduced and new, resistant strains appearing.
Initially it was possible to find new organo-phosphorus materials that were
more effective than their predecessors. These included acephate, diazinon,
dichlorvos, dicrotophos, disulfoton, mephosfolan, methidathion, metha-
midophos, mevinphos, omethoate, orthen, phorate and phosdrin but most of
these are now of little value due to resistant strains developing. The rather
surprising exception is mephosfolan which has been used for many years as a
soil-applied material in England where it is still proving to be the most effective
chemical available although increasing reliance upon it does mean that there is
more danger of resistant strains arising.
There was hope of overcoming the resistance problem with the introduction
of other groups of chemicals such as the carbamates (propoxur and meth-
omyl), an organochlorine (endosulfan) and most recently the synthetic pyr-
ethroids. Although the pyrethroids have been in use for only a short time there
are already very high levels of resistance developing. Kremheller (1988a), for
example, reported that resistance to cypermethrin increased 40-fold between
1980 and 1985. Numerous other accounts ofthe resistance problem are given in
the Proceedings of the Working Group Integrated Pest and Disease Control in
Hops (Hrdy and Hrdlickova, 1981 and 1984; Campbell and Hrdy, 1988) and
this development leaves growers in a very serious situation.
The onset of resistance has been most rapid in European countries where the
aphids are a very serious problem and where spray applications are frequent.
Selection pressure is greater than in the USA, for example, where the aphids
Damson-hop aphid: Phorodon humuli 121
are less of a problem and some chemicals are consequently still effective in the
USA although they have had to be abandoned in Europe.
There are differences, too, in the reported effectiveness of the same materials
on different farms and it is widely believed that these reflect the efficiency with
which the spraying operations are carried out. It is most important that
spraying machines are checked to ensure that everything is in good order and
that the nozzles and settings of the machines are correct.

6.1.3 Biological control

Mohl (1924) described how in Czechoslovakia ladybirds, lacewings, hover flies
and birds were able to save growers from the ravages of the aphid if
temperatures were high and there was plenty of sunshine. If, however, the
nights were cool and high temperatures only lasted for a few days the aphids
were dominant.
The important effect that predators had upon aphid populations there was
stressed by Blattny and Osvald (1950) who pointed out that there were never
two consecutive years of severe aphid damage because after one such year a
large predator population carried over to the next season. Little attention was
paid to predators for some years after that account, presumably because the
newly developed insecticides gave such effective control. As the aphids have
become resistant to many of these chemicals there has been renewed interest in
predator activity.
Zeleny et al. (1981) studied the occurrence of predators and parasitoids over
a 15-year period. Ladybirds (Coccinellidae) were the most common predator
but many more species were recorded, some in significant numbers. An
important observation was that it was only when aphid populations reached 50
or more per leaf that permanent colonization and reproduction by the
predators occurred. With smaller aphid popUlations the predators only
occurred sporadically.
Ruzicka et al. (1986) also stated that ladybirds are the most important
predator. They reported a trial in which an attempt was made to control a
population of aphids in a hop garden without using insecticides. The 0.9 ha
hop garden was surrounded by a 52 ha pea field and several rows of mustard
were planted inside the plot to attract predators. Conditions that season were
favourable for the development of ladybird populations but not so good for
the aphids. The aphid popUlation declined sharply in July when large numbers
of adult ladybirds migrated into the garden from the surrounding field and it
may only have been the combination of such a migration and the favourable
weather conditions that led to this successful result.
Hrdy (1981) reviewed the problem of resistance to pesticides in hops and
drew attention to the potential effect on the environment of the increased
quantities of chemicals being used. In the Bohemian hop-growing region this
122 Pests
had increased in the period of 1967-77 from 3.5-8.3 kg active ingredient per
hectare. He suggested that the problem could be reduced by, among other
measures, ceasing to use an insecticide as soon as resistance to it developed and
changing to one with a different mode of action. In addition only using them
when absolutely necessary and using selective materials which would not be
toxic to predators. Unfortunately there has rarely been an adequate choice of
products to put these measures into effect.
Knan and Kremheller (1984) found that average populations of 200 aphids
per hop leaf did not cause any damage before flowering and that it was only
necessary to commence control measures when this threshold was reached. In
England an infestation of this magnitude would be considered to be out of
control.
The increasing problems with strains of aphids resistant to chemical
control have focused attention upon non-chemical ways of controlling the
pest. There had been early reports of differing susceptibility of hop cultivars
to aphid attack (Gross, 1900; Hampp and Jehl, 1937) but these were of no
importance at a time when chemical control was very effective. When
evidence of resistance to OP insecticides began to emerge however, a start
was made at Wye in 1965 to look for hop selections that showed resistance to
the attacks of the aphids (Neve, 1966) but not until 1968 did this work show
any promise. In that year the aphid migration started slowly and the most
resistant cultivar in the trial, Tolhurst, survived the whole season without any
insecticide treatment and was almost free from any infestation at harvest
time. This was attributed to predators mUltiplying sufficiently quickly to
control the smaller population of aphids that had built up on this cultivar
(Neve, 1969).
It was thought from this experience that the predators were probably of
greater importance than the rather low level of resistance that had been found
within the hop population and for some years the research effort was directed
towards methods of promoting the effectiveness of the predators. Some
selection of resistant plants continued, however, the results of field trials being
confirmed by laboratory experiments. Darby and Campbell (1988) identified
one female and five male genotypes which were consistently less heavily
infested than the standard varieties. A laboratory feeding experiment with
these six genotypes and three control cultivars showed that four of the
selections had marked antibiotic effects on the reproductivity of two strains of
aphids although the OP resistant strain had a greater reproductive capacity
than the other strain on all genotypes (Campbell et al., 1987).
These resistant genotypes have been used as parents in a breeding pro-
gramme, and testing of the progeny in 1988 indicated that the resistance is
hereditary and not, apparently, sex-linked. The level of resistance in some of
the progeny was better than that of either parent, especially when both parents
were resistant (Darby, personal communication).
Damson-hop aphid: Phorodon humuli 123
In the USA a similar search for resistant genotypes has been carried out by
Dorschner and Baird (1988) by measuring the rate of increase of aphids caged
on leaves of a wide range of material from the US Germ Plasm Collection
growing under growth-room conditions. The most susceptible and the most
resistant genotypes were plants collected from the wild, while less wide-ranging
differences were recorded between commercial varieties, the aphids having low
rates of increase on the German cultivar Perle and the American Chinook.
In Germany a collection of 125 male or female hops, collected from the wild,
have been planted in a garden that receives no chemical control and are being
recorded for the level of infestation with both aphids and spider mites. In 1987,
11 of the wild hops and the Yugoslav cultivar Atlas were recorded as aphid free
(Kremheller, 1988b). Although Perle was recorded by Dorschner and Baird as
showing resistance it is interesting that this was not mentioned in Kremheller's
report although it is likely to have been tested. This might be a result of the
different methods of testing since aphid preference for different genotypes
could affect the size of the initial population on a mixture of plants in the field
but not in the controlled experiments in the growth-room where the varieties
were grown in isolation.
Romanenko (1985) reported that the polyphenol content of the leaves of hop
plants was an objective criterion for measuring resistance to the aphid.
Campbell (1977b, 1978) reported that Anthocoris neh'lOrum, A. nemoralis
and the earwig Forficula auricularia were the most abundant predators of
aphids on hops in East Kent and found in laboratory trials that each anthocorid
would kill about 200 aphids during its nymphal development and about 35 per
day during its adult life. During the latter part of the season he compared the
populations of aphids that developed on bines where the aphids were exposed to
predation with those on bines from which predators were excluded by sleeve-
cages. In the absence of predators, aphid numbers increased rapidly but where
predators had access they were very effective in reducing the numbers.
Aveling (1981a) extended this work by setting up plot trials to test
Campbell's technique of controlling aphids during the early part of the season,
before predators arrived in the hop gardens, by a soil drench of mephosfolan
and observing the effectiveness of predators thereafter. Anthocorid bugs,
especially A. nemoralis, were the most abundant predators and in the first two
years aphids in the cones were controlled. In the third year aphids did build up
in the cones but not to damaging numbers. When predators were excluded the
cones were heavily infested.
If this was to be a practical technique for achieving integrated control it was
important to know the extent to which the mephosfolan which was systemic
within the hop plant would be toxic to the anthocorids. Aveling (1981b)
reported that there was no evidence of toxic effects on anthocorid larvae or
adults fed on aphids that had been killed by feeding on mephosfolan-treated
hop plants. He did record increased mortality of eggs laid in the leaves of
124 Pests
treated plants and this was greatest when the eggs were laid near the leaf
margins where the concentration of mephosfolan was thought to be highest.
Because the aphids migrate to the hop crop before predators are present in
any numbers, it is essential to provide some chemical control, at least during
the early part of the season. Ideally, a selective insecticide should be used which
is toxic to aphids but not to predators. Unfortunately materials that can be
used in this way against other aphids are not effective against the damson-hop
aphid. For this reason selective action has had to rely upon the use of soil
applied materials such as mephosfolan that are taken up systemically by the
hop plants. In this way the predators do not come into direct contact with the
pesticide. Winfield (1981 and 1984) carried out field trials from 1977-83 to
determine whether predators could control the aphids following an early-
season mephosfolan drench but with only limited success. There were six hop
gardens in the trials but in only two did the system work successfully in some
years. Although anthocorids were the key predators in these trials, hover flies
(Syrphidae) , lacewings (Chrysopidae), earwigs (Forficula spp) and, in some
years, ladybird beetles (Coccinellidae) were also common (Figure 6.4)
Although .predator activity following mephosfolan treatment did not pro-
vide adequate commercial control of the aphids, it may well have contributed
to the fact that this chemical, alone amongst the organophosphate materials, is
still effective since resistant aphids that survive the chemical are liable to be
destroyed by predators.
At present there does not appear to be an integrated control programme that
will operate successfully but there are possible developments that could help to
achieve this. Most promising amongst these is the breeding programme to
develop the resistance of the hop plant to the aphid. It appears unlikely, at
present, that this would provide complete protection by itself but it should
greatly increase the possibility of predators providing an effective control.
Another possibility is that of artificially rearing predators that can be
released into the crop to supplement those arriving naturally. Parker (1981)
has published a method for artificially rearing anthocorids but it appears that,
unless an artificial diet can be developed, the feeding requirements with live
prey are likely to make the technique too labour-intensive for it to be
commercially viable. The artificial rearing of lacewings, however, is more
feasible and their release into a low trellis garden at Wye in 1990 gave good
control with or without an early mephosfolan treatment. An alternative
approach has been an attempt to attract egg-laying female lacewings to hops
early in the season by spraying the plants with artificial honeydew just before
the aphids were due to arrive from their winter host.This failed to work in 1987
because prolonged rain delayed spraying until the middle of June but it showed
more promise in 1988 (Campbell, personal communication).
Whereas predators have been shown to be of considerable importance in
controlling aphid populations it does not appear that parasites are likely to
Damson-:hop aphid: Phorodon humuli 125
have a significant effect, even when natural populations are supplemented
artificially. Campbell et al. (1987) released 10000 adult Aphidius matricariae
into each of two 48 hill hop gardens three weeks after the start of aphid
migration and a further release of the same number about three weeks later.
These releases had no detectable influence on the development of the aphid
popUlations and very few parasitized aphids were found and the trials were
abandoned.
There have been trials with the fungus Cephalosporium (Verticillium) lecanii
as a pathogen of the aphid (Figure 6.5) but although it worked well in pots
under glass it gave poor results in the field (Cranham and Firth, 1984).
Campbell et al. (1985) reported on trials with the aphid alarm pheromone
(E)-~-farnesene which, when released by the aphids causes dispersal of other
aphids feeding nearby. This compound substantially reduces the numbers of
aphids settling on other crops but it did not noticeably affect the settling of
damson-hop aphids when applied to hops. It did, however, cause the aphids
to stop feeding and walk over the leaf surfaces. This activity would increase
the likelihood of aphids coming into contact with any insecticide on the leaf
surface and when leaves were sprayed with the pheromone, immediately
followed by a spray of Thiodan (endosulfan) the mortality of adult apterae
was 87% compared with 76% when Thiodan was sprayed without the
pheromone.
Dawson et al. (1984) reported that hops are readily colonized by aphids even
though the plants themselves contain this compound and they found that
another sesquiterpene component of the essential oils, (- )-~-caryophyllene,
inhibited the effect of the farnesene in the hops so that it did not act as a
deterrent to colonization.
A possible method of reducing the aphid popUlation would be to spray the
sex-attractant pheromone, by which males are attracted to oviparae, in places
where there were none, thereby reducing the chances of the oviparae ever being
fertilized. The pheromone has recently been shown to be a nepetalactol and an
effective attractant (Campbell et al., 1990/91).
The prospects for continuing control of the aphid by purely chemical
means are not good. Resistance to all groups of aphicides is already either
well-established or showing clear signs of developing and no new types of
aphicide appear to be forthcoming at present. The problem is being
aggravated further by the refusal of some countries to permit the import of
hops containing residues of pesticides not registered there. Because different
countries have different pest and disease problems, the pesticides that they
need to use (and which are registered) also differ and this is creating major
problems for the international trade of hops. There is, therefore, likely to be
great emphasis given in the near future to any technique which will reduce the
dependence of hop growers upon chemicals for the control of this extremely
damaging pest.
126 Pests

(b)

(e) (d)

Figure 6.4 Predators of the damson-hop aphid: (a) anthocorid adult; (b) ladybird eggs;
(c) ladybird pupa; (d) lacewing egg; (e) lacewing larva; (f) lacewing adult; (g) hover fly
larva; (a and d IHR, Wye. b, c, e, f, g, Crown copyright).

6.2 SPIDER MITE : Tetranychus urticae

6.2.1 Description
This pest, formerly named Tetranychus telarius (L.) or Epitetranychus
althaeae (var. Hanst), is commonly called the two-spotted mite or the red
spider mite but the latter name is at times misleading since it is only the
overwintering females that are a bright red colour. These shelter during the
winter in the soil, under leaves, in cracks in hop poles or in the strands of the
wirework (Darling, 1958). These adults leave their hiding places in the spring,
climb the bines and feed on the lower surfaces of the leaves by sucking the sap
from the epidermal and sub-epidermal cells. They lay small translucent eggs
Spider mite: Tetranychus urticae 127

(f)

(g)

and the mites that emerge are greenish-yellow with black markings and there
are five to nine generations of these during the summer. They are small and,
although just visible to the naked eye, a lens is very helpful in identifying them.
The immature larval stages are six-legged while the adults are eight-legged.
Blattny and Osvald (1948) described them as one of the most important pests
in Czechoslovakia and most serious in warm dry weather. Oviposition began
when the mean temperature reached l2°C (54°F) and at that temperature the
egg stage lasted for six days. At 18°C (64°F) the egg stage only lasted for three
days. The mites stopped feeding when the relative humidity reached 80% and
stopped reproducing when it reached 90%.
Cone et al. (1986) have reported from Washington State that overwintered
females made a very rapid recovery once weather conditions were favourable
and that they commenced egg laying within 24 hours of starting to feed. The
first eggs to be laid had pigment similar to the female but many of these did not
hatch. The subsequent eggs were normal (pearly-white) and hatched. On
128 Pests

Figure 6.5 Damson-hop aphid infected by Verticillium Jecanii (Crown copyright).

average, the females laid 20.8 eggs over a 20 day period. Of 2390 eggs laid in
this study, 1036 hatched into females and 320 into males.
As the hop plants develop, the mites move up to feed on the younger leaves.
Sites and Cone (l985) found that from May to early July the mites were mainly
on the lower half of the plants but that by mid-August they were mainly on the
upper half. The first sign of mite attack is a silvery speckling of the leaves and
their presence is also indicated by the fine web which they spin as a protection.
This can be shown up by sprinkling soil on the underside of the leaf where it is
caught in the webbing. In small numbers the mites do not cause much damage
but under favourable conditions they can multiply very rapidly and , if an
infestation is allowed to develop, it will result in severe browning of the leaves
and the cones. In severe cases the crop may be destroyed. In September the red
hibernating females are produced, reproduction ceasing when daylight is less
than fourteen hours (Moreton, 1964).
Unlike most pests and diseases of the hop, spider mites are not specific to
that crop but have a wide host range outdoors and are also a serious pest of
glasshouse crops. Their geographic distribution is not, therefore, restricted to
areas where hops are native and they are to be found wherever hops are grown
including countries in the southern hemisphere where they are the only major
pest or disease of the crop. Since they flourish under hot dry conditions they
are a serious problem in Australia and the Yakima Valley in the USA.
Spider mite: Tetranychus urticae 129
In cooler countries, they are usually only a problem in hot, dry summers and
in Germany they are reported to be more of a problem on lighter soils
(Kohlmann and Kastner, 1975). They are probably least important in England
where conditions are not so frequently favourable for their development but
even there they have become an increasingly serious problem as a result of the
extensive use of mist propagation during which the cuttings spend some time in
glasshouses where the mites flourish.

6.2.2 Chemical control

Before the introduction of the organo-phosphorous insecticides the spider mite
could be a serious problem. Brown (1980) wrote that the reports on the Guinness
hop farm at Bodiam made no mention of red spider from 1905, when the farm
was established, until 1930 but that for some years after that it was a serious
problem, especially in 1934. He comments that there was no effective insecticide
and using water to drench the webs was reckoned to be the best that could be
done. However, he also stated that flowers of sulphur were being used to control
powdery mildew and this material was a reasonably effective miticide.
Dinocap, another fungicide used against powdery mildew, also gave some
control of the mites and in many seasons these two products were probably
sufficient to prevent the pest building up to the stage where specific measures
became necessary. It was noted in one season at Wye that the only variety to be
affected by spider mites was Wye Challenger which, because it was at that time
resistant to powdery mildew, had received neither sulphur nor dinocap sprays.
The organo-phosphorous pesticides, when introduced, were as effective
against the spider mites as they were against hop aphids and for some years the
problem appeared to be solved. As with damson-hop aphids, however, resistance
soon began to develop and alternative materials had to be sought. Control is a
somewhat different problem with the mites because of the egg-laying stage
between each generation. Some materials are toxic to the mites but not to the eggs
and treatments need to be repeated to ensure that mites that subsequently emerge
are sprayed before they have time to lay a further batch of eggs.
Several alternative products have been introduced as miticides but all seem to
be subject to the risk of resistance developing while some have been found to be
phytotoxic. Resistance does not develop everywhere at the same time. Muir and
Cranham (1979) reported resistance to dicofol in England yet five years later
Gesner (1984) found no evidence of resistance to this material in Czechoslovakia.
In view of his results, Gesner said that the increasing damage being reported
could not be attributed to resistance and in England dicofol is still proving of
some use against the mites. The lack of agreement between laboratory tests for
resistance and experience in the field may be due to variations within the
popUlation. One material for which there were no reports of resistance was
cyhexatin but this was withdrawn from use in 1988 for health reasons.
130 Pests
6.2.3 Biological control

On fruit crops it has been possible to achieve considerable success in limiting

the damage done by the fruit tree red spider with an integrated control
programme based upon the activity of the predacious mite Typhlodromus pyri.
One such programme relied upon using selective pesticides such as diflu-
benzuron or pirimicarb that are relatively harmless to any strains of this mite.
An alternative approach was to introduce a strain of T. pyri from New Zealand
that is resistant to organo-phosphorous insecticides and to use such materials
in the control programme (Solomon and Cranham, 1980). Pesticide appli-
cations are reserved for occasions when the predacious mites have failed to
keep the pests below a predetermined level.
T. pyri is however less likely to be effective against the two-spotted mite on
hops because it dislikes the webbing which that mite produces. Some success in
preliminary trials, however, has been achieved using the South American
species Phytoseiulus persimilis.
In the USA the native species Typhlodromus occidentalis is an effective
predator but it continues feeding on the two-spotted mite during the winter
and reduces the popUlation to a level where it will no longer support the
predator. The surviving mites are then able to build up again. An alternative
food source for the predator is being sought to bridge this gap so that it would
be present in sufficient numbers to prevent renewed build-up of the mite
(Cone, personal communication).
Blattny and Osvald (1948) reported that natural predators, of which
Stethorus punctillium was the most important, destroyed large numbers of the
mite at all stages and they commenced a programme of releasing them in areas
where they were insufficiently numerous. In their book (1950) they state that
the level of mite infestation depends, above all, on the numbers of natural
predators that survived from the previous year and that is the reason why it is
nearly impossible for it to be a problem in the same garden two years running.
There is little hope, however, of using T. pyri or other predators to control
spider mites while aphids have to be controlled by chemicals, none of which are
sufficiently selective to avoid destroying the predators. But if it is possible to
develop an integrated control programme for the aphid, this is likely to help
with the control of spider mites also since it is known that anthocorids, at least,
will feed on mites as well as on aphids. The release of artificially-reared
Phytoseiulus persimilis to control mites, combined with the release of lace-
wings for aphid control, was remarkably successful in a low trellis garden at
Wye in 1990 - a difficult year for control of spider mites. On standard
wirework they failed to control the mites on the upper parts of the bines.
In the southern hemisphere, where there is no problem with aphids, there
should be far better prospects of success with biological control and recent
work by the CSIRO, Canberra has shown promising results. With reduced
Nematodes 131

Figure 6.6 Predatory mite, Typhlodromus occidentalis (right) attacking red spider mite,
Tetranychus urticae (left). (Division of Entomology, CSIRO, Canberra).

applications of miticides, several species of predatory mites have appeared at

levels sufficient to give control (Figure 6.6). Phytoseiulus persimilis has also
been introduced into hop gardens where it has established well and given good
control although its ability to overwinter has not yet been established (personal
communication).
There are some reports of differences in susceptibility of hop cultivars to the
mite. Peters and Berry (1980a) found that in greenhouse experiments mite
densities were greater on Comet and Fuggle than on L- 8, L- I or Cascade.
There were no differences in oviposition, sex ratio or survival but highly
significant differences in developmental rates. The pre-adult stages developed
slower on L-16 and Talisman than on Cascade, Fuggle or Comet. In another
paper (1980b) they report that oviposition increased on leaves with high
density of hairs and there was a greater proportion of females on leaves with
dense pubescence. The duration of development of pre-adult stages was longer
when the hairs were dense.

6.3 NEMATODES
6.3.1 Dagger nematode: Xiphinema diversicaudatum

Although this eelworm species probably causes no direct damage to hop plants
132 Pests
on which it feeds, it must be the next most serious pest of the crop, after aphids
and spider mites, because it is the vector of Arabis Mosaic Virus (AMV) which
can cause serious losses (section 8.4).
The role of X. diversicaudatum as a vector of AMV in other crops was first
recognized by Harrison and Cadman (1959) and its occurrence in Britain was
not recorded until 1959 when it was found in association with clover (Peacock,
1959) and also with strawberries infected with AMV (Jha and Posnette, 1959).
Taylor and Brown (1976) later showed that it is widespread throughout
England but it is most commonly found in soils with a high proportion of sand.
They also comment that it is widespread in continental Europe although it
appears to be restricted to the cool humid areas such as those along river
banks, the Channel and Atlantic coasts. A survey in the Federal Republic of
Germany found X. diversicaudatum only rarely in hop gardens and then only
in one of the regions (McNamara and Flegg, 1984).
The identification of this nematode as the vector of AMV in hops, which
causes nettle head disease, was reported by Thresh et al. (1972) and Valdez et a1.
(1974). Subsequently there were various experiments to determine how to
prevent it from spreading the disease. This work is described in section 8.4, but
it is of interest to note here that a two-year fallow was found to eliminate the
virus from infected land although it did not destroy the X. diversicaudatum
population that was carrying it (Pitcher and McNamara, 1976). Taylor and
Robertson (1970) demonstrated that this is because the virus particles are
retained by the nematode on the cuticular lining of the gut. During the moult
the cuticular lining is shed, together with the virus particles, and these are
ingested into the intestine so that after moulting the nematodes are no longer
infective.

6.3.2 Hop-root eelworm: Heterodera humul;

Percival (1895) attributed the cause of nettlehead to the eelworm Heterodera
schacht;; having found its cysts (Figure 6.7) on the roots of hops suffering from
that disease but Duffield (1925) found no correlation between its presence and
the incidence of nettlehead. Filipjev (1934) considered that the nematode found
on hops differed morphologically from those found on other plants and
renamed it as a separate species, Heterodera humuli.
It has been suggested in Germany that there may be an association between
the presence of this eelworm in hop gardens and the incidence of verticillium
wilt. This has been supported by von Mende and McNamara (1985) who found
a strong indication of an interaction between Heterodera and Verticillium in
that the bine length of N orthdown plants was much reduced when both
pathogens were present together; this did not occur in the wilt tolerant variety
Wye Target. They had insufficient data to reach definite conclusions about
other damage in the field but concluded that the nematode is a well-adapted
Other pests 133

Figure 6.7 Cysts of hop-root eelworm Heterodora humuli extracted from hop garden
soil (Crown copyright).

parasite that causes little or no growth reduction unless the root system is put
under stress when it may become of major significance.

6.4 OTHER PESTS

There are several other pests that occasionally attack hops but they are
generally not of great importance.

6.4.1 Clay-coloured weevils: Otiorrhynchus singularis

These wingless weevils (Figure 6.8), which are about 0.6 cm long, shelter under
clods of earth during the day where they are difficult to see since their colour
blends with the soil.They emerge at night to climb up the sterns where they will
gnaw at the bark or eat irregular holes in the leaves. If they eat enough of the
bark they can cause the shoots to collapse. They are active in the spring until
early summer when they lay eggs in the soil from which the legless larvae
develop and feed on the roots before pupating.
Kohlmann and Kastner (1975) described the wingless weevil in Germany, the
LiebstOckelrtissler (0. ligusticl), as being 1.0-1.5 cm long and having a two-
year life cycle. The larvae which emerge from the eggs pupate in the autumn
134 Pests

Figure 6.S Clay-coloured weevil (Otiorrhynchus singularis) (Crown copyright).

and overwinter, and the young weevils which develop in the following July
overwinter without leaving the soil but emerge as adults in the second spring to
feed and lay eggs to complete the cycle. It is recommended that control
measures, using organo-phosphorous or carbamate insecticides, should be
undertaken when there are one or more weevils to three plants.
Mohl (1924) recorded that in the years 1880-2, in some regions of what is
now Czechoslovakia, there were clusters of 5-25 beetles per plant and that they
destroyed all the shoots that were produced between March and mid-May.
A recent report from Germany discusses the possibility of using parasitic
nematodes to control this pest (Arndt, 1989).

6.4.2 Rosy rustic moth: Hydroecia m;cacea

These moths are on the wing from late July to early October with a peak in
September. The eggs are laid in the autumn and hatch in the following spring.
The caterpillars are pinkish with a dark central stripe with brown spots on each
side, each with a bristle. French et al. (1973) described the results of a 6-year
survey of this pest in Kent. In early to mid-May the larvae emerged from the
soil (slightly later than reported in Germany) and bored into the stems, usually
near soil level but sometimes up to a height of 30 cm (12 in), and tunnelled the
pith. In June they emerged from the bine usually within 30 cm of soil level and
Other pests 135
then fed on the base of the bine or tunnelled into the crown or roots, pupating
in July. Some larvae never fed on the bines but went immediately to feed on the
crown and roots - usually when bine growth was weak or retarded.
The report says that the tunnelled bines did not wilt or show reduced vigour
but that there was some leaf yellowing of the leaves in July or August. The
tunnelled bines were more susceptible to infection with Fusarium canker and
the larval exit holes sometimes caused the bines to split. More damage resulted
from the subsequent feeding around the join between bine and rootstock which
could cause the bine to collapse. In their survey they found fewer bines
surviving on infested plots and, of those, from 8-12% were damaged. They
obtained the best control using DDT which they applied twice, with a lO-day
interval, commencing the first week in May and they recorded yield increases
in both seasons of the trial. Damage was less if training was left until after mid-
May.
Damage was said to be more severe in weedy gardens or next to the
headlands where the moths were more likely to lay eggs. In recent years there
have been few reports of damage and the pest may have diminished in
importance since non-cultivation made conditions less suitable for it.
In Germany, where the insect is called the Kartoffelbohrer, it still appears to
be of some importance and carbofuran is the recommended control treatment.

6.4.3 Flea beetle: Psylliodes attenuata

This was formerly a common pest in most hop gardens but is not as prevalent
now. It is a small metallic-looking jumping beetle that feeds on young shoots in
April and May, eating holes in the leaves. They lay their eggs in the soil from
early April until early July. The larvae feed on the roots of the plants and then
pupate. The beetles emerge in the following spring.
It is probably the introduction of more effective, systemic insecticides,
coupled with the widespread adoption of non-cultivation in England, that has
reduced the numbers of this insect in hop gardens.

6.4.4 Earwigs: Forficula auricularia

These are common and may, on occasions, cause noticeable damage by eating
holes in the leaves but this is rarely sufficient to affect the crop. This damage
may, however, be outweighed by the beneficial effect that they have as
predators of the damson-hop aphid (Buxton and Madge, 1976; Campbell,
1978).
The eggs are laid in the soil during winter and again in May-June. During
the day they shelter in cracks and crevices and emerge during the night to feed.
Hop poles are a common place for them to shelter and plants next to poles are
more likely to be visited by them.
136 Pests
6.4.5 Wireworm: Agriotes spp.
Young hops may sometimes be damaged by wireworm when they are planted
in sites where the pest is to be found, most commonly land recently ploughed
out of old grassland. They kill young shoots by biting into them below ground
level in the spring. Where they are a problem the recommended control is
currently by soil application of y-HCH.

6.4.6 Slugs: Agriolimax reticulatus and Arion hortensis

These can cause so much loss of bine in certain seasons that control by poison
bait such as methiocarb is necessary. They feed at night so may not be noticed
but their slime trails are evidence of their activity.
CHAPTER 7

Fungal diseases

7.1 DOWNY MILDEW: Pseudoperonospora humuli

A recent comprehensive review of this disease is given by Royle and

Kremheller (1981).
The spread of downy mildew in the 1920s throughout all the hop growing
countries of the northern hemisphere and later into South America, as
described in Chapter 5, must make this the most important disease overall
affecting the crop even though other diseases may be locally more serious.
Australia, New Zealand and South Africa have been able to exclude it by strict
quarantine precautions.
The first appearance of the disease in the spring is when some of the many
shoots that the plants produce develop into 'basal spikes' with a characteristic
stunted form and pale downcurled leaves with a silvery upper surface (Figure
7.1). The undersides of these leaves turn black as dense masses of sporangia are
formed on branched sporangiophores. Once a shoot has developed into a spike
it does not grow any further and, if not removed or destroyed by chemical
treatment, will quite soon die. These spikes are the primary source of infection
each year and, as the sporangia spread the disease, other shoots may develop
into secondary basal spikes. Lateral or terminal buds on the elongating bines
may also develop into spikes. If this happens to the terminal bud there will be
no further development of the main axis of that bine but, because apical
dominance is lost, there is increased development of lateral shoots. If one of
these laterals is trained in place of the main bine it may develop sufficiently
well to give a satisfactory yield but otherwise there can be a serious loss of
crop.
Leaves also become infected and once the diseased areas start sporulating
they appear on the underside as black spots which are bounded by the veins of
the leaf so that they have a characteristic angular shape (Figure 7.2). The black
colour of these 'angular leaf spots' is due to the profuse development of
sporangiophores. Infections of buds and leaves can occur at any time in the
growing season, depending very largely upon weather conditions and the
amount of inoculum being released.
138 Fungal diseases

Figure 7.1 Downy mildew 'spike' (right) and healthy shoot (left) (Crown copyright).

When the hops come into flower the developing burr and cones may also
become infected. If the burr is attacked its development may be completely
checked while diseased cones turn brown, especially the bracteoles, the bracts
frequently being less severely discoloured (Figure 7.3).
Infection during the vegetative phase of growth does not often directly affect
Downy mildew: Pseudoperonospora humuli 139

Figure 7.2 Downy mildew: Angular leaf spot (Crown copyright).

the final yield. If so many shoots developed into spikes prior to training that
there were insufficient healthy ones to train, this could affect the final yield
although this is an unusual situation, especially on mature hills which have an
abundant supply of shoots. When shoots that have already been trained are
attacked the situation is more serious because at that stage there may be no
replacements available. Terminal spikes will certainly check the vegetative
development of the bines even if laterals are trained to take over the leading
role and infection of lateral buds would result in the loss of lateral shoots that
could have borne a crop of hops. Losses from such causes should, however, be
minimal if normal control measures have been carried out.
140 Fungal diseases

Figure 7.3 Downy mildew infected cones (Crown copyright).

Infection during the vegetative growth is much more important as a source

of inoculum that can infect the burr and cones when the greatest losses are
likely to arise. The sudden flush of young tissue in the developing cones is very
susceptible to infection and this is the time when the growth of the plant is at its
most dense, making it difficult to achieve good spray cover with fungicides.
There may be little reduction in yield if infection does not occur until after the
cones are formed but the discoloration of the cones will usually lead to a
serious reduction in the value of the hops.
Even though diseased rootstocks usually produce sufficient healthy shoots
to furnish all the strings, the subsequent growth may still be weaker than that
from uninfected rootstocks. Williams et al. (1961) showed that during the
winter the carbohydrate reserves declined much more rapidly in infected than
in healthy rootstocks and this adversely affected their growth in the following
season. Coley-Smith and Beard (1962) noted which hills of a number of
Golding clones showed evidence of infection when they were dressed in the
spring and at harvest recorded a reduction in yield of 27.7% compared with
those with healthy rootstocks.
Downy mildew: Pseudoperonospora humuli 141

Figure 7.4 Growth of Pseudoperonospora humuli in artificially infected pot plants of

Eastwell Golding. A-D diagrammatic representations of rootstocks, AI-DI vertical
sections through dormant buds. A and Alone week after inoculation (September), B
and BI two weeks after inoculation (early October), C and Clone month after
inoculation (mid-October), D and DI two months after inoculation (November). The
diseased area is shown in black. The dotted line indicates the soil surface.

7.1.1 Overwintering

The fungus is an obligate parasite that is specific to hops. Ware (1926) showed
that it overwinters as mycelium within the rootstock. Coley-Smith (1960)
studied the way it progressed during the winter in plants that were inoculated
in September by pouring a zoospore suspension onto the soil surface around
the base of the bine. The fungus entered the stem at or near the soil surface and
grew both upwards and downwards so that by October the rootstock was
extensively colonized. No buds were infected by that time although hyphae
were present in many of the cushions of tissue from which the buds arose.
From November onwards, however, increasing numbers of buds had been
invaded by hyphae passing through these cushions. Over 40% of all buds
became infected and by March it was obvious that those invaded early in the
winter were already dead (Figure 7.4).
Coley-Smith (1965) demonstrated that rootstocks could become infected by
inoculating the bases of the bine at any time during the growing season or by
inoculating the tips of bines that were 15 cm or less in length. If the bines were
more than 17.5 cm long when the tips were inoculated the infection did not
penetrate into the stock.
142 Fungal diseases

Skotland (1961), on the other hand, observed diseased crowns following the
inoculation of the tips of shoots up to 90 cm (3 ft) tall although at least some of
this could have been the result of secondary infection since this occurred in
some of the uninoculated controls.
Skotland recorded a field plot of 368 plants and found that, in three
consecutive seasons, 14%, 18% and 6% of them produced basal spikes but that
only two plants produced them in all three seasons. All plants that produced
basal spikes had infected rootstocks but not all infected stocks produced
spikes.
When Coley-Smith (1964a) recorded field plots, he found infection persist-
ing in rootstocks of Golding hops for four years and, unlike Skotland, he
found that most of these plants produced basal spikes each year. It was on this
evidence that he recommended the grubbing and replanting of diseased stocks
as a means of reducing the level of infection (Coley-Smith, 1963).
Dormant buds that are heavily infected will normally die whereas those in
which the fungus is only recently established can develop into shoots although
many of these will turn into spikes after they have emerged. It appears that in
some other cases infected shoots can grow away normally but that isolated
pockets of mycelium may be carried upwards in the developing bines and then
infect buds high up the bine to produce lateral or terminal spikes (Ware, 1926
and 1929).
Rootstock infection can have an important effect on yield by causing the
death of the rootstock although cultivars vary in their susceptibility to such
losses. Although in some cases a correlation has been found between numbers
of basal spikes and the degree of rootstock infection within varieties, this is not
true in all cases. The variety WGV is especially sensitive to rotting of the
rootstock yet produces few basal spikes (Coley-Smith, 1964a). This is,
presumably, because when the rootstocks become infected the disease spreads
so rapidly that few buds survive to develop into spikes. On this basis an inverse
relationship would be expected between rootstock susceptibility of a cultivar
and the incidence of primary basal spikes.
Such an inverse relationship has been noted at Wye where Northern Brewer
and Bullion were grown in an experimental garden in which no control of
downy mildew was practised. Bullion produced few spikes but many hills died
out while Northern Brewer produced very many spikes but no hills were lost.
There is some doubt as to whether the oospores, which are produced
abundantly in diseased leaves, shoots and cones in the autumn, are a further
means by which the fungus can overwinter. Early investigators, including
Magie (1942), observed oospores germinating and quoted circumstantial
evidence that they were the source of infection in the spring but more recent
workers have been unable to substantiate these findings (Royle and
Kremheller, 1981). There is no doubt that, even if oospores are capable
occasionally of germinating, they are not an important source of infection.
Downy mildew: Pseudoperonospora humuli 143
7.1.2 Secondary infection

The primary spikes are normally too few to be a direct cause of loss but they
are of the greatest importance as the source from which all secondary infection
develops.
Secondary spread occurs by means of the sporangia produced on stems and
the undersides of infected leaves. They are produced in a daily rhythm on the
ends of each branch of the sporangiophores which emerge through the stomata
of infected tissue. The sporangiophores were found by Yarwood (1937) to
emerge by midnight and to produce small sporangia by 0300 h. By 0600 h the
sporangia were full sized and release commenced by 0900 h. This rhythm was
not disturbed, at least for 24 h, by submitting infected leaves to altered light
routines.
Royle (1968) found that the production of sporangia occurred over a
broad temperature range (approx. 8-25°C) but was absolutely dependent on
high relative humidity, being most profuse between 96-100% and failing to
occur below 90% RH. The release of the spores in static air conditions,
however, increased with lowering humidity, being almost nil at 100% and
greatest at 50% RH. The daily concentration of air-borne sporangia in an
unsprayed hop garden was studied by means of a spore trap and the catches
were more closely related to the amount of sporulating tissue than to any
other factor.
Diurnally spore release also proceeds in a rhythmic fashion. On rainless days
the maximum release of spores was at 10 am GMT but rain caused temporary
increases in numbers as a result of the mechanical disturbance of the leaves.
The sporangia do not reinfect directly but under wet conditions each
sporangium releases 4-8 zoospores which swim to the stomata, which only
occur on the underside of hop leaves. After encysting, zoospores penetrate the
stomata with a germ tube. It has been shown that within 2 min of being
sprayed onto leaves that are in the light, zoospores commence settling
selectively on the stomata which will be open, whereas on leaves in the dark,
with closed stomata, the zoospores remain motile much longer and finally
settle at random. The rapidity with which they locate open stomata is
apparently due to a chemical attraction related to photosynthesis but they
settle there, at least in part, as a physical response to the open stomata since
they also settle on the open 'stomata' of perspex replicas made from leaves that
were in the light but not on those made from leaves in the dark (Royle and
Thomas, 1971a, band 1973).
The zoospores are motile with two flagella and require surface moisture on
the plant tissue in order to reach the stomata. The sporangia require wetness
for a period of at least 1 h at 20-22° C to 10 h at 2° C before they release the
zoospores, so there is generally sufficient surface moisture at that time for the
zoospores' requirements.
144 Fungal diseases
A feature of this fungus recorded by Royle and Thomas (1973) is that the
zoospores settle singly on the stomata whereas in the vine mildew, Plasmopora
viticola, the zoospores settle on the stomata in groups of up to 30.
As sporangia age there is at first an increase, to a minimum of 3-4 h, in the
period of wetness required before zoospores are released (Zattler, 1931; Magie,
1942) and with increasing age they quite rapidly become non-viable. Magie
found that sporangia lived only a few hours when separated from the host
plant and dried on glass slides in the laboratory. When sprayed onto leaves
that were then exposed to sunlight on a greenhouse bench none were viable
after 24 h. Sonoda and Ogawa (1972) also found that only 1% lived longer
than 14 h under Californian conditions while, in Bavaria, Kremheller (1979)
reported that 50% died within 1Yz days although a few survived for 8 days.
Since infection takes place through stomata, and these are absent from
juvenile bud leaves, the likely entry sites for shoot infection are the young bud
stipules in which stomata are well developed. When looking for signs of
secondary infection in the field the stipules should be carefully examined as
they are frequently the first place at which it can be recognized.
After germ tubes have penetrated the stomata, an intercellular hyphal system
develops which spreads quite widely between the cells of the plant. The cells are
invaded by lobed haustoria which are a useful means of identifying the pathogen.

7.1.3 Hygiene
Since diseased rootstocks are the initial source of infection each season a
programme of identifying and grubbing these as recommended by Coley-Smith
(1963) can help to reduce the amount of primary infection in the garden. This
can be done by marking hills that produce primary spikes in the spring so that
they can be grubbed in the autumn. Alternatively the rootstocks can be
inspected when being dressed in the winter and any that cut brown can then be
grubbed.
Such grubbing is unlikely to be completely successful so even if it is carried
out there would be some spikes developing in the spring. The next stage in field
hygiene is carefully to inspect the young plants and remove, by cutting or
pulling, any spikes that are seen. After the bines have been trained, and surplus
shoots pulled out, the lower leaves should be stripped off by hand as soon as
the bines are high enough. Initially they should be stripped to a height of about
0.6 m (2 ft) and then to twice that height or in particularly wet seasons, to as
much as 1.5 m (5 ft) (Glasscock, 1956). This stripping of the base of the bine
removes the leaves that are closest to the basal spikes and therefore the ones
most likely to become infected. It is also helpful in promoting more movement
of air through the garden, thereby reducing the period for which the foliage
remains wet after rain.
The pulling of excess bines and the stripping of the lower leaves by hand has
Downy mildew: Pseudoperonospora humuli 145
now been replaced, very largely, by the use of defoliating chemicals. This has
the advantage that the chemicals will kill any spikes in situ whereas hand work
carried the risk that it would, unintentionally, help to distribute the spores as
diseased material was removed from the garden.
When severe outbreaks of the disease have occurred there have sometimes
been rather desperate attempts to check further spread by removing infected
leaves and side shoots by hand but it is doubtful if this was able significantly to
affect the course of the disease once it was so well-established.

7.1.4 Chemical control

When hop downy mildew was first recorded in 1905 in Japan, the use of copper
fungicides for the control of vine mildew was already well-established so
spraying hops with Bordeaux mixture was immediately adopted as a control
measure, combined with the removal by hand of infected shoots and leaves
(Salmon and Wormald, 1923). Bordeaux and Burgundy mixtures continued to
be used for very many years but, because they could cause a certain amount of
damage to young leaves they were then replaced by less phytotoxic copper
products such as copper oxychloride.
Copper fungicides were the only materials available until the 1950s when the
organic dithiocarbamate compounds such as zineb were introduced. These
were favoured because they did not harden the leaves so much and this became
important with the introduction of machine picking since the harder, copper-
sprayed leaves broke up more on the machine and so gave rise to more broken
leaf in the final hop sample.
Typical spray programmes did not start until the bines reached the top wire
although in wet seasons, which favoured the disease, it was suggested that they
might start as early as April (Burgess, 1964). Some growers, however, believed
that dusting the hills and young shoots very early in the season was beneficial
and trials by Coley-Smith (1965) confirmed that this reduced secondary spread
to leaves and shoots as well as reducing rootstock infection.
These fungicide treatments were only protective and did nothing to cure
infection that had already occurred so it was essential to apply them in time to
prevent further infection occurring.
A significant advance came with the discovery that spray applications of
streptomycin were taken up systemically in the plant and so had some effect
upon infections that were already established. Its use was first advocated by
Horner and Maier (1957) who found that it could transform infected shoots
into healthy ones. Coley-Smith (1966) found that two applications were
necessary in most seasons and that its main effect was to reduce sporulation.
There was always some unease about using this material for non-medical
purposes. It was never approved in the Federal Republic of Germany and after
being used for some 15 years it was withdrawn everywhere.
146 Fungal diseases
Even more effective systemic fungicides were introduced into trials in the
late 1970s and two of these, metalaxyl (Ridomil) and fosetyl-aluminium
(Aliette), were widely approved for use. Royle (1980) showed that metalaxyl
residues could be detected in hop leaves for at least ten weeks after the
application of a soil drench which gave complete control for four weeks and
erratic control thereafter. When applied to the soil these systemic materials
were capable of eliminating the fungus from rootstocks that were already
infected.
Experience with pathogens of other crops showed that there was a risk of the
hop pathogen developing resistance to metalaxyl and so its use as a soil drench
was discontinued in the UK where it is now applied only as a foliar spray in
mixture with copper oxychloride. It was hoped by this means that the risk of
resistant strains becoming established would be reduced. In Germany and the
USA, however, it has been used in granular formulations for soil application
and unconfirmed reports from both these countries that resistant strains have
developed suggest that the policy adopted by the UK distributors was justified.
Metalaxyl was so effective that it would even cure established infections and
by removing sources of inoculum it could effectively eliminate the disease from
a garden beyond the period when the concentration of the fungicide in the
leaves was high enough to give direct control (Darby, 1984a). Writing at the
time when metalaxyl had been in use commercially for only two years, Royle
and Kremheller (1981) suggested that it might reduce the sources of downy
mildew to such an extent that the disease would no longer pose a threat to the
health of the crop. It has, however, never been used extensively enough by all
the growers in a country to achieve that situation and it cannot be said whether
it was even possible. In view of the rapidity with which the disease spread
throughout Europe in the 1920s it is likely that there would always have been
enough foci of infection on wild hops to re-establish the disease as soon as the
spray programme was relaxed.

7.1.5 Epidemiology

When controlling downy mildew with protectant fungicides, the programme

can be operated much more efficiently if it is known when infection is likely to
occur. It has long been realized that the disease is worse in wet weather but
further work was required to establish more precisely the factors that lead to
the development of epidemics.
Magie (1942) examined the effect of temperature by inoculating leaves at 10,
13, 18 and 24°C and found that the shortest periods of wetness required for
infection to occur at those temperatures were 6, 4, 2 and l.5 h respectively.
Royle (1970), working under controlled conditions in growth rooms, used the
wider temperature range of 5-30°C and recorded minimum wetness require-
Downy mildew: Pseudoperonospora humuli 147
ments ranging from 1.5-24 h. These results agreed with those of Magie at 18
and 24° C but slightly shorter periods were recorded by Magie at 10 and 13° C.
In Royle's experiments, inoculated shoots needed to be exposed to wetness
periods approximately twice as long as those required for leaf infection before
they would eventually develop into spikes and no spikes developed from the
treatments at the two extremes of the temperature range. He suggested that the
reason for this longer period was that spikes only develop when the fungus
reaches the growing point and to achieve this it must grow faster than the
shoot. This may only happen when large numbers of fungal penetrations check
shoot growth sufficiently for the growing point to be invaded.
Royle also examined the effect of temperature upon the incubation period
(the time between inoculation and symptoms appearing) over the slightly
narrower temperature range 7-28°C. The higher the temperature the shorter
the incubation time which, for leaves, ranged from three to seven days. Again it
took approximately twice as long as this for shoots to develop into spikes and
none developed when incubated at the highest or lowest temperatures.
Skotland (1962) exposed plants that had been dusted with sporangia in a
hop yard overnight and dews formed on each occasion. In four out of six trials
some infection occurred but there was no correlation between infection and the
temperature or the duration of the dew. Other workers who have similarly
exposed plants have found that little infection occurred as a result of dew
unless there were high levels of inoculum (Royle, 1973; Dolinar, 1976;
Kremheller and Diercks, 1983). Three reasons suggested by Royle and
Kremheller (1981) for this were: (a) dew starts to form in the dark when
zoospores are not attracted to stomata; (b) rain is more favourable for
infection than dew because inoculum that is splash-dispersed by rain augments
that which is deposited dry after the daily release; and (c) the longevity of dry-
deposited inoculum is reduced during the interval between release and wetting
by dew, especially because dew is often associated with dry weather.
Pejml and Petrlik (1967) studied the incidence of the disease over many years
and developed a points system based on weather parameters. A score of 0-500
meant conditions were unfavourable for infection, from 500-1000 infection
could occur while over 1000 conditions were optimal. The number of sprays
applied was adjusted on this basis.
Royle (1973) exposed healthy pot plants in an unsprayed hop garden for
48 h periods, incubated them for three days in a growth room and then for a
further seven days in a glasshouse where infected leaves and shoots were
recorded. Environmental conditions in the hop garden were recorded through-
out the trial. A Hirst spore trap was used to record airborne inoculum which
was computed hourly and a funnel trap recorded spores deposited by rain.
The level of infection was profoundly influenced by the amount of young,
highly susceptible leaf and shoot available and allowance had to be made for
the time lag between infection and the appearance of disease symptoms.
148 Fungal diseases
Periods when conditions of moisture and temperature were judged to be
suitable for infection to occur were in good agreement with the subsequent
disease recorded provided that wetness periods resulting from dew were
excluded.
The infection records were related by inspection and by multiple regression
analysis to the conditions when the plants were exposed. The spore catches in
the funnel trap, which depended on rain, were closely related to infection while
airborne counts showed only a weak correlation and only when all three years
records were combined. There was no correlation with temperature. Variables
such as temperature, vapour pressure deficit and airborne spores, which in
their own were poorly correlated with infection, often significantly improved
regression equations which were based on variables expressing wetness. The
mUltiple regression equations that were developed accounted for 70-90% of the
variation in infection and when used to predict infection periods there was
good agreement between the predicted and observed results.
As a result two alternative formulae were developed for assessing when
infection periods had occurred. If a spore trap was in operation to record
airborne inoculum the formula was:
Y = 37 + 23 wet + 66 rain + 2 spores
and if no spore count was available the alternative formula was:
y = - 63 + 22 RH + 84 rain

where
wet = hours of surface wetness in 48 h
rain = mm rain in 48 h
RH = hours of relative humidity above 80% in 48 h
spores = number of spores/ m3 air
When Y was greater than 500, infection periods were judged to have occurred.
A warning system was initiated in England based on this work and, although
operated for several years, did not get much support from growers and has
now been abandoned in favour of routine treatments.
In the BRD on the other hand, a similar warning system is being operated
very successfully (Kremheller, 1983; Kremheller and Diercks, 1983). The model
used there is based upon the concentration of airborne spores, as recorded by
spore traps set up in a number of hop gardens, and the duration of rain
wetness. The threshold values set allow for burr and cones being more
susceptible than leaves. In the period 1976-82 sprays could be timed more
effectively and the quantities of fungicides used were reduced on average by
50%.
It is not clear why the German system has been much more successful than
the one in England but a possible explanation is that hop farms in England are
Downy mildew: Pseudoperonospora humuli 149
more scattered. With the highly concentrated hop cultivation in Germany,
each recording station can effectively assess the conditions for the hop gardens
in the neighbourhood. Royle and Shaw (1988) have also pointed out that a
routine programme in England only involves 5-8 sprays whereas in Bavaria
15-18 applications are typical and a potential annual saving of D M 11-15 m by
use of the warning programme has been claimed (Kremheller, 1984).

7.1.6 Resistant cultivars

Soon after downy mildew had spread through the main hop growing areas
varietal differences in susceptibility were noted. Salmon and Ware (1927) first
considered that Fuggle was immune but later (1932) they reported that in the
exceptionally wet season of 1931 it had become infected although not to the
same extent as other cultivars. In the USA also, Fuggle was found to be one of
the most resistant cultivars (Magie, 1942). In the Hallertau district of Germany
it was noted that hops of the Saaz type (Saaz, Spalt, Tettnang and Schwetz-
inger) were less susceptible than the local cultivars but were not so well-
adapted to the growing conditions there (Zattler, 1951).
Little effort was made to develop these differences in susceptibility except in
Germany where a hop research institute was established at Hull in 1926, largely
in response to the problem of downy mildew, and a breeding programme
commenced to develop acceptable cultivars with high levels of resistance to the
disease. This work, which is described in Chapter 9, took many years to come
to a successful result but it has finally led to very high levels of resistance being
bred into new varieties in Germany and England. Since the resistance appears
to be very stable, with little risk of breaking down because of more virulent
strains of the fungus, it offers the possibility of a long-term solution to a very
important disease.
In order to screen young seedlings for resistance it is necessary to have a
plentiful supply of sporangia early in the season before field sources are
available. Derbyshire and Dixon (1957) developed a successful method of
growing hop plants throughout the winter in a heated glasshouse with artificial
light to extend the daylength. Leaves detached from such plants are inoculated
with mildew spores in the laboratory. The disease can be kept in an active state
throughout the winter and then be bulked up in time for large-scale testing of
seedlings when these are at the two to four leaf stage.
Griffin and Coley-Smith (1968) successfully cultured the fungus on hop
callus cells on a solid medium. It produced intercellular hyphae and haustoria
as well as sterile aerial hyphae and sporangiophores but this procedure has not
been put to any practical use.
150 Fungal diseases

Figure 7.S Powdery mildew 'blisters' on a resistant variety. Similar blisters occur in the
early stages of infection of susceptible cultivars (IRR, Wye).

7.2 POWDERY MILDEW: Sphaerotheca humuli

There was a major review of this disease by Royle (1978).
Powdery mildew is the oldest of the fungal diseases of the hop and in
Germany is called 'Echter Mehltau' (true mildew) to distinguish it from the
much more recent downy mildew which is called 'Falscher Mehltau' (false
mildew). In England it is commonly called 'mould'. Although it is so much
older it is more restricted in range than downy mildew. It is a serious problem
in England and Belgium but of less importance in the rest of Europe. In
Germany it was a disease of little significance until Northern Brewer was
planted extensively. This variety is particularly susceptible and the disease has
increased as a consequence. There are recent reports that powdery mildew is
proving a problem in Bulgaria.
It caused great damage in the USA when hops were grown on the east coast
and was one of the problems that led to the American hop industry moving to
the west coast where powdery mildew does not occur in commercial hop yards.
When a collection of wild hops was grown in a glasshouse in Oregon, however,
there was an outbreak of the mildew on them which had to be eradicated
(Hampton, personal communication).
Quarantine restrictions have prevented this disease becoming established in
South Africa or Australasia although on at least one occasion it is known to
have occurred on imported material which had to be destroyed.
Powdery mildew: Sphaerotheca humuli 151

Figure 7.6 Powdery mildew sporulating on a susceptible cultivar (IHR, Wye).

The first sign of the infection on young leaves, especially on plants growing
under glass, is the appearance of raised humps or blisters (Figure 7.5) on which
the white sporulating mycelium (Figure 7.6) then appears (Salmon, 1917a).
The blisters appear to result from hypertrophy of the cells around the infection
site and as their expansion is restricted by the surrounding tissue they are
forced to bulge outwards. These blisters are not as noticeable on the tougher
leaves that are produced in the field or on leaves that are older when infection
occurs. On plants in a glasshouse the sporulating pustules may be profuse on
both the upper and lower surfaces of the leaf. On plants in the field they are
mostly confined to the underside of the lower leaves, though their presence is
indicated by pale spots on the upper surface. Higher up the bine they develop
on both upper and lower surfaces. The poor development on the upper surface
of the lower leaves is probably because these are more exposed to spray
deposits (Royle and Liyanage, 1973).
The effect of the disease on the cones depends very much upon their stage of
development when infected since further growth of infected tissue is almost
completely inhibited. When the burr or very young cones are infected they
remain as hard white knobs but later attacks may be localized and so cause a
one-sided distortion of the cones (Figure 7.7). When hops are grown seedless
they remain in the burr stage much longer than they do when pollinated and
this is the stage at which infection can cause the greatest damage. However,
152 Fungal diseases

Figure 7.7 Cones infected with powdery mildew at various stages in their development,
(Crown copyright).

growers in England are increasingly changing over to seedless production and

there have been no strong indications so far that this has led to greater
problems than those experienced by seeded producers.
In the early stages of infection the disease pustules appear white because of
the prolific chains of asexual conidia that grow up from the surface mycelium.
In the latter part of the season - from July onwards - there is usually a change
to the development of cleistocarps. At this stage mycelium production is
suppressed but the cleistocarps, which are dark coloured, can be recognized
quite easily with the help of a hand lens. Cleistocarps may be produced on
leaves but they are usually much more abundant on cones where chemical
control at the end of the season is most difficult.
The cones on which cleistocarps are produced frequently develop a rusty-red
colour and this stage of the disease is referred to as 'red mould' in contrast to
the 'white mould' of the conidial stage. In some seasons growers have
complained that their hops have ripened prematurely and it has been shown
that this is often associated with the development of the cleistocarp stage on
the cones although there may have previously been no obvious sign of white
mould in the crop (Coley-Smith, 1964b). It should be noted, however, that the
Powdery mildew: Sphaerotheca humuli 153
same conclusion was reached over 60 years earlier by Hammond (1900) who
wrote:
When I have asked further, What is Red Mould? I have never been able to get a direct
specific answer, but have been persuaded to believe that it was a mysterious something,
which, like charity, covered a multitude of sins.
All this seemed so splendidly vague, that I resolved by the aid of the microscope to
attempt the unravelling of the mystery, and I have succeeded so far that I am convinced
absolutely that the author of all the mischief is none other than our old enemy the
ordinary White Hop Mould.
This is not only an interesting example of how scientific discoveries tend to
be forgotten and then rediscovered some 50-60 years later, but also of the more
informal and more readable style of reporting that once existed.
Royle (1976) has demonstrated that the fungus is heterothallic and that
cleistocarp production is preceded by fusion of the two mating types which
exist. The different pathogenic strains, which have been identified from their
reaction to resistance genes in host varieties, may each contain both mating
types so fusion can be either between or within strains.
The white mould that develops on the leaves can usually be controlled
sufficiently well for it not to affect the yield and it is infection of the cones that
is the chief cause of loss. Cones infected at an early stage do not develop at all
and those infected later will have their growth at least partly checked. This
results in a reduction in yield but even more serious losses may result from loss
of quality. Infected cones not only affect the appearance of the hop sample but
they can also give it an unpleasant mushroom-like aroma. There have been
many cases of a hop garden being so badly attacked that it has not been
worthwhile to pick it. Salmon (1917b) described a severe outbreak in 1916
when 'some hundreds of acres had to be left unpicked' while many that were
harvested were badly diseased and downvalued in consequence.

7.2.1 Overwintering
The commonest way for the fungus to overwinter is by means of the
cleistocarps that are produced during the latter part of the growing season.
When mature, the cleistocarps contain eight ascospores which are released in
the spring to reinfect hop leaves. Liyanage and Royle (1976) showed that there
are two peak periods for maturation, one in November and the other in March,
but they were unable to induce any cleistocarps to dehisce and release the
ascospores before April. This coincides with the time when hops are
recommencing growth and when naturally dehisced cleistocarps could be
found in the field.
The temperature at which the cleistocarp-infected material was kept during
the winter months had little effect upon the proportion that finally discharged
ascospores but there was an effect of temperature at the time of discharge.
154 Fungal diseases
When subjected to temperatures of 4,8, 18 and 24°C (39, 46, 64 and 75°p) in
April, more c1eistocarps released ascospores at 18°C than at other temper-
atures. Even under the most favourable conditions, however, only a very small
proportion (<2%) released spores. The germination of the ascospores was also
best at 18°C when nearly 10% of the spores germinated while only 1-2% did so
at the other temperatures. Ascospores were only released from the c1eistocarps
in the presence of moisture.
In the same paper, Liyanage and Royle reported the distribution, in the
spring, of powdery mildew in a hop garden, part of which had been left
unpicked the previous September because of a severe attack of the disease.
Reinfection in the spring was almost entirely restricted to the unpicked portion
of the garden which suggested that it was caused by ascospores released from
infected cone debris in that area. They produced confirmatory evidence by
spreading hop cones bearing many cleistocarps over healthy hop cuttings. Over
10% of the leaves that were examined on the young shoots were infected while
no infection was found on the leaves of control plants, which had no infected
cones spread over them.
During the last 20 or so years in England there have been reports of shoots
emerging in the spring which were almost entirely covered by the white mildew,
instead of showing the usual scattering of mildew pustules on the leaves. This
phenomenon coincided with the adoption of non-cultivation techniques and
Liyanage and Royle showed that this type of infection arose from buds that
had overwintered with mycelium, sometimes accompanied by cleistocarps,
between the bud scales.
It is only buds at or above the soil surface that can become infected with the
mildew and in cultivated gardens all of these are normally removed when the
hills are dressed in the winter. Under non-cultivation this is not done and the
buds at soil level are left undisturbed. It is from such buds that the infected
shoots develop.
Non-cultivation also favours the disease because leaf and cone material that
falls to the ground at the end of one season remains on the surface whereas in
cultivated gardens a high proportion of it would be buried and unable
therefore to release ascospores into the air where they could reinfect the plants.
For these reasons, powdery mildew has presented English growers with
increasingly severe problems since non-cultivation was widely adopted.

7.2.2 Secondary infection

Once primary infections have become established, the disease is spread by
means of the conidia. Whereas the ascospores require water for germination,
conidia can produce germ tubes which will develop on dry leaf surfaces in
various atmospheric humidities.
The subsequent development of the germ tube, as described by Liyanage
Powdery mildew: Sphaerotheca humuli 155
(PhD thesis, 1973) and quoted by Royle (1978), depends upon the genotype of
the hop plant on which it has landed. On susceptible hosts the first germ tube
emerges from the conidium 6 h after inoculation, produces no obvious
appressorium but penetrates the host through the cuticle and establishes an
haustorium in an epidermal cell within 12-15 h. After 48 h the initial germ tube
becomes a branching hypha and up to three more germ tubes develop. The
colony begins to abstrict conidiophore initials after 96 h and the first conidia
are apparent soon afterwards. Sporulation intensifies from 5-7 days after
inoculation. Godwin et al., (1987) give a similar account of the infection
process.
Only the epidermal cells of the host are invaded. The haustoria have a
central nucleate body with lobed outgrowths. Up to three haustoria per cell
have been recorded but one is most common.

7.2.3 Hygiene

In the case of downy mildew there is a very discrete initial source of infection in
the primary basal spikes so that removal of these is a feasible method of
minimizing secondary spread. In the case of powdery mildew the same
technique can be applied to the heavily infected shoots that develop from bud
infection but most of the primary sources are the scattered pustules arising
from ascospore infection and the removal of these by hand is not practicable.
Stripping the lowest leaves from the bine, as for downy mildew, will remove
some of the primary sources and also those leaves most liable to secondary
infection. The use of chemicals to replace hand stripping is probably beneficial,
as with downy mildew, since it destroys the spores in situ and avoids the risk of
further spread as the diseased material is removed from the garden.
A return to cultivation after a period of non-cultivation should help to
reduce the level of infection but the advantages of the non-cultivation
technique are sufficient to outweigh, for most growers, the value of this as a
hygiene method.
Perhaps the most important precaution that should be taken is to ensure
that the minimum of diseased debris, especially infected cones, is left in the
garden at the end of the season since this provides the starting point for the
disease the next season. The greatest danger arises when a crop is so badly
diseased that the grower decides not to harvest it. One virtue of machine
picking is that only the lowest part of the bine, most of which has been stripped
of any leaves, is left in the hop garden, and so most diseased material is
removed. When a badly diseased crop is not harvested, therefore, every effort
should be made to cut the bines down and burn them as quickly as possible.
The longer they are left the greater the risk of the cones shattering and
spreading diseased material.
156 Fungal diseases
7.2.4 Chemical control

The earliest chemical used to control powdery mildew was sulphur and it is still
used to some extent today, either as powder or as a spray. Dinocap became
available later and for many years control was based on these two materials.
Newer products, some with limited systemic activity, include pyrazophos
(Afugan) which was tested by Royle and Liyanage (1973) and found to be
much more effective than dinocap and, even though it caused some phytotox-
icity, the yield was better.
More recently approved chemicals in England include bupirimate (Nimrod),
triadimefon (Bayleton), triforine (Saprol) and penconazole (Topas). Of these
triforine was shown to be the most effective (Royle, 1976) and this has been the
general experience of growers. It can however cause a reduction in yield and it
is recommended that its use should be limited to four applications during the
season. When a crop is threatened with severe damage it is better to risk some
phytotoxicity if the disease can be checked by further use of this material. One
disadvantage of the newer chemicals is that they do not have the same
acaricidal effect as sulphur and dinocap.
Although copper fungicides do not directly affect powdery mildew it has
been noted that they do contribute towards its control (Royle and Griffin,
1968). This is almost certainly an indirect effect due to hardening of the leaves
which are consequently less susceptible to infection.
Since the mycelium of this fungus grows entirely on the plant surface it
should, on the leaves, be an easier target for chemical control than downy
mildew which proliferates within the plant tissues. In practice, however, it can
be an extremely difficult disease to control and this is probably because it is
tolerant of a wide range of environmental conditions.

7.2.5 Epidemiology

Whereas epidemiological studies on downy mildew were very successful in

relating the development of infection to climatic conditions, similar studies
with powdery mildew have proved much more unsatisfactory. One major
reason for this difference is probably that powdery mildew is not dependent
upon free water for secondary infection to occur. Prolonged spells of dry
weather can reduce downy mildew infection in a garden to such a low level that
there is a considerable delay before it builds up again with the return of wet
conditions but mould infections are not so susceptible to adverse weather.
Powdery mildew conidia are able to germinate under dry conditions and in
general these mildews are reputed to be inhibited by water. Royle (1978) found,
however, that high levels of infection could be obtained in hops by spraying
leaves with aqueous suspensions of conidia.
Although secondary infection can therefore occur on either wet or dry leaves
Powdery mildew: Sphaerotheca humuli 157
it appears that the fungus is susceptible to very high temperatures and low
atmospheric humidity. The summer of 1976 was one of the driest on record
and Royle (1977) concluded that this was the reason why infection in that year
was extremely low. It has frequently been noted at Wye that although mildew
will spread on plants in the greenhouse with great speed during the early part
of the season, it practically dies out in the height of the summer. It is probably
such hot dry conditions that account for its absence from American hop yards.
The main conclusion from Royle's investigations was that the amount of the
disease increased most rapidly and inoculum production, as recorded in spore
traps, was greatest when there were flushes of growth of highly susceptible,
young material. There were three main danger periods during the season:
before training, when lateral shoots appeared and when burr was produced
(Royle, 1979).
This gives some guidance to growers for the timing of their chemical control
programme. An early application should be made regardless of whether or not
the disease has been observed since it will not only help to eradicate any
primary infection that has been overlooked but will also protect the plants
during the first of the risk periods. The timing of further sprays may be judged
by the development of the disease, although many growers rely on a routine
application every 10-14 days, but in any event they should include an
application at each of the other risk periods.

7.2.6 Resistant cultivars

In England powdery mildew is proving very difficult to control by chemical
means thus there has been much interest in the development of resistant
cultivars. The first to be widely grown was Wye Challenger which also had a
very high level of resistance to downy mildew. It was first distributed to
growers in the spring of 1972 but in 1974 the first record of the resistance
breaking down was reported. The way in which the outbreaks of the disease
were distributed suggested the possibility that the pathogenic race of the
fungus was already established, possibly on wild hops.
Wye Target was released to growers in 1973 and it has since been so widely
planted that it now occupies 30% of the English hop area. It has a different
resistance mechanism to Wye Challenger and so far this resistance has proved
stable under field conditions except for three reports of infected plants in 1979.
Only one of these locations was notified in time for it to be investigated and
there only one plant was found to be infected. The fungus from that plant was
inoculated onto other plants of Wye Target under laboratory conditions and
confirmed as weakly pathogenic (Neve and Lewis, 1980). On none of the sites
did the disease recur. Isolates which sporulate on Wye Target have also been
recovered from screening programmes under glass (Godwin et al., 1987) but
these do not appear to be viable under field conditions.
158 Fungal diseases

The resistance of Wye Challenger is characterized, on plants grown under

glass, by the development of blisters very similar to the early stages of infection
in susceptible plants. There is then only very sparse development of mycelium
which soon dies. The blistering is rarely seen on plants growing in the field but
since most of the testing for resistance is carried out under glass it is the
diagnostic feature of the genotype and is referred to as the 'blister reaction'. It
is almost certainly the type of resistance described by Salmon (1919) as 'semi-
immunity'. The segregation of this character in seedling families indicates that
it is controlled by a dominant major gene.
When Wye Target is inoculated there are either no visible symptoms or tiny
necrotic flecks which suggest a hypersensitive reaction. The segregation of this
type of resistance also indicates control by a dominant major gene. Other
resistance genes have also been identified, and the original designations
suggested for those controlling the blister reaction of Wye Challenger and the
immune reaction of Wye Target were Band 12 (Liyanage et al., 1973). Four
different resistance genes have now been identified by the reactions between
hop genotypes and fungal isolates and the designations were amended to R B,
R 1, R2 and R3 by Liyanage (1973). He designated the corresponding virulence
genes of the pathogen VB' V h V2 and V3.
Strains of the fungus have arisen that are pathogenic under field conditions
to all commercial varieties except Wye Target but other, non-commercial,
immune host genotypes have been identified. These come from various sources
including Russia and the USA and some, at least, are probably unrelated to the
source of the Wye Target resistance. It is not known whether these carry the
same resistance gene since there are no fungal isolates that will distinguish
between them.
Godwin et al. (1987) reported no consistent differences in the frequency of
conidial germination on susceptible Northern Brewer and resistant Wye Target
but germ tube and hyphallengths were much less on Wye Target. Nor was
there a significant difference in the frequency of haustorium initiation but on
the resistant plants fungal growth was usually restricted soon afterwards. On
Wye Target, the penetrated epidermal cells showed a hypersensitive reaction
with granulation of the contents and death of the haustorial initials.
There have been three instances when sectorial chimaeras have been
discovered on hop seedlings in which one sector has been susceptible to the
disease and the other immune (Figure 7.8) (Neve, unpublished; Darby and
Gunn, 1987). In at least one instance the mutation must have been from a
susceptible to an immune genotype but again it is not known whether the
resistant gene is one already in the gene bank.
The varieties which were resistant when originally selected have frequently
proved to be highly susceptible when strains have appeared that could
overcome their major gene resistance. Wye Challenger was in commercial
production before a pathogenic strain appeared but other seedlings which also
Powdery mildew: Sphaerotheca humuli 159

Figure 7.S Chimaerical hop plant with sectors resistant or susceptible to powdery
mildew (IHR, Wye).

carried the blister gene were at an advanced stage of selection. One of these was
subsequently released under the name Yeoman and this has proved to be
particularly susceptible. This stimulated renewed interest in the development
of breeding lines with high levels of polygenic resistance. On its own such
resistance would yield varieties on which the control of mould by chemicals
would be relatively easy. Combined with major gene resistance it would reduce
the likelihood of races pathogenic to the major gene developing and even if this
did happen the polygenic resistance would still be operative (Neve and Darby,
1982).
The initial screening at Wye of seedling populations for their reaction to
powdery mildew infection is carried out in the glasshouse when the plants are
stilI quite small and large numbers can be tested quickly. Darby et af. (1989)
compared the reaction of seedlings in such a screen with their subsequent
performance in the field where a natural epidemic was encouraged. They found
a significant positive association between the level of disease on male seedlings
in the glasshouse and on their leaves in the field. There was no corresponding
agreement in female plants for leaf infection in the field but a weak association
with cone infection. They suggested that the difference between male and
female plants was due to the plant producing flushes of young, susceptible
leaves at different times during the growing season.
160 Fungal diseases
Although the correlation between glasshouse screening and field
performance was not strong, they found that the initial screen was more
efficient at identifying the highly susceptible seedlings and that by discarding
these they could achieve worthwhile benefits to the selection process.
The continued breeding of resistant cultivars is being given a very high
priority in England as described in more detail in Chapter 9 but in other
countries the disease is not of sufficient importance to require similar activities.

7.3 VERTICILLIUM WILT: Verticillium albo-atrum

7.3.1 Description

The first record of this disease was in England in 1924 at Penshurst, Kent on
the cultivars Fuggle and Tolhurst (Harris, 1927). No further cases were
recorded until 1930 when more severe outbreaks began to appear and by 1937
about 12 had been identified. It continued to spread rapidly both on farms and
between farms. Keyworth (1942) described how, on one farm, it was first
recorded in 1934 with a group of 20 wilted plants in one field. By 1938 there
were 2000 diseased plants in that field and a year later 26 out of 31 fields on the
farm were affected.
The leaves of affected plants develop yellow patches followed by irregular
black necrotic areas between the main veins giving a characteristic black and
yellow pattern described as 'tiger striping' (Figure 7.9). It.is a dry wilt and the
affected leaves drop off very easily. This is used as a help towards diagnosis but
the most reliable symptom is on the lower part of an infected bine, the woody
core of which turns a coffee-brown colour. In some cases the lower 1.2-1.5 m
(4-5 ft) of the bines become swollen but, unlike plants affected by Fusarium
canker, they do not develop a restricted 'neck' at the base.
Keyworth (1942) noted that there appeared to be two different types of
outbreaks which he described as 'fluctuating' and 'progressive'. In fluctuating
cases thickening of the bines was more common, the browning of the wood was
often limited to the centre and plants which had shown wilt symptoms
commonly recovered. The severity of fluctuating outbreaks would vary from
season to season but there was little increase in the number of plants showing
symptoms in succeeding years.
In progressive outbreaks the symptoms appeared earlier in the season
(sometimes as early as May), diseased plants usually died and the infection
spread rapidly through the garden. It was noted that in gardens that were
cultivated in both directions there was a general spread in all directions but in
gardens that could only be cultivated one way the spread was very much
restricted to that direction.
It was originally thought that the differences in severity of the disease were
the result of different soil conditions, especially soil water (Harris, 1936).
Verticillium wilt: Verticillium alba-atrum 161

Figure 7.9 Male plant infected with verticillium wilt showing typical 'tiger striping' of
the leaves (Crown copyright).

Keyworth was unable to confirm this in the fields that he examined and made
many isolations of the pathogen from outbreaks of all types.
A study of such isolates by Isaac and Keyworth (1948) established that the
difference between fluctuating and progressive outbreaks was due to differ-
ences in the virulence of the pathogen which appeared to exist in two distinct
forms. The terms 'fluctuating' and 'progressive' which were used to differen-
tiate between the strains of the fungus really describe symptoms and these
depend not only on the virulence of the pathogen but also upon the
susceptibility of the host. For this reason the strains would be better identified
as 'mild' or 'severe' but the original designations have been incorporated into
legislation in The Progressive Wilt of Hops Order introduced in 1947, which
made it compulsory to notify outbreaks of progressive wilt while fluctuating
wilt was not notifiable.
The purpose of this legislation was to prevent the spread of the progressive
form of the disease to areas other than the Weald where it was firmly
162 Fungal diseases
established. When outbreaks were notified in other areas steps were taken to
eradicate them and this approach was very successful for many years, even in
East Kent which was so close to the Weald that isolated outbreaks occurred
quite frequently. Within the Weald itself the disease was so firmly established
by the time its true cause was identified that any attempt at eradication was
impractical.
The legal distinction between the two forms of the pathogen made it
necessary to be able to distinguish between them. Talboys and Wilson (1954)
described a method for doing this by inoculating susceptible plants growing in
concrete troughs in the open and this method was adopted for the official tests
on one of the Ministry of Agriculture experimental stations. The facilities there
became inadequate to cope with the numbers that required testing and a new
system was developed in which plants grown in growth chambers were used for
the test.
The results of the outdoor test could be seriously affected by weather
conditions and Clarkson and Heale (1985a) found that tests over more than
one season would be necessary to reliably distinguish fluctuating and progres-
sive isolates. The outdoor tests had the further disadvantage that no results
were available until at least 12 months after the outbreak was notified. During
that time the uncertainty could lead to further spread of the disease since
control measures could not be enforced until the fungus had been proved to be
a progressive strain.
Growth rooms were not affected by weather conditions and more than one
series of tests could be carried out during the year so that the earliest results
were obtained quicker than with the outdoor test. This was, however, only a
partial improvement since even the earliest results were not available for some
months while the latest were as long delayed as with the outdoor test. A great
deal of research has therefore been devoted to attempts to find a rapid method
of characterizing the two forms of the pathogen.
Sewell and Wilson (1980) describe how the 'progressive' strain spread
rapidly through Kent destroying all the commercial varieties. By 1965 the
disease was largely controlled by the use of resistant varieties, accompanied by
restricted nitrogen fertilizers, non-cultivation and high standards of hygiene.
But between 1968 and 1971 severe wilt developed in previously resistant
cultivars and pathogenicity testing confirmed that this was due to two 'super-
virulent' strains, V2 and V3.
In a later paper, Sewell and Wilson (1984) reported that the levels of
resistance of the test varieties retained the same relative ranking to the 'super-
virulent' V2 and V3 strains as they did to other isolates and that there was no
evidence of specificity. The new strains were initially isolated from nine farms
all in the same area and they concluded that they had spread from a single
focus as did the original progressive (VI) strain.
Although the progressive strains of wilt spread very fast through the Weald
Verticillium wilt: Verticillium albo-atrum 163
of Kent, it was possible for very many years to eliminate the outbreaks that
occurred in other districts. The situation in the West Midlands was made more
confusing by the fairly widespread incidence of mild, fluctuating outbreaks. In
the end, however, a new outbreak of a severe strain could not be contained and
it spread rapidly to many farms in the area. In east Kent, where fluctuating wilt
was rare, there have been frequent outbreaks of the severe strains but most
farms have been able to contain these.
Isolates of the fungus can be grouped by comparing their effects, in tank
tests, on the varieties Bramling Cross and Wye Target. VI is of low virulence
on both, V2 causes severe wilt on Bramling Cross but not on Wye Target while
V3 is virulent on both although less so on Wye Target. Wye Challenger, which
had been judged to be fully susceptible when released, is moderately resistant
to VI (Sewell et al., 1979).
There have been various attempts to develop a faster and more reliable
method of distinguishing between the fluctuating and progressive strains of the
fungus. Webb et al. (1972) used gel electrophoresis to determine total protein
and specific enzyme patterns of different isolates but found no association with
their virulence. Mohan and Ride (1984) studied the morphological and
biochemical characteristics of three serotypes of the fungus but also failed to
establish such a correlation. Nor did the assay of polygalacturanase by workers
at East Malling using electrophoresis, in collaboration with the Ministry of
Agriculture laboratory at Harpenden, consistently correlate with the virulence
of the isolates.
Hignett et al. (1983) grew a number of mild and virulent isolates on medium
augmented with hop cell wall as the sole carbon source on which enzyme and
pH changes were monitored. By statistically combining four characters 90% of
the isolates could be correctly classified. The use of the ELISA technique to
distinguish between isolates serologically has also proved unreliable. The
results supported previous data indi,cating that there is no absolute disconti-
nuity between the progressive and fluctuating types (Swinburne et aI., 1985).
The most recent attempt to develop a rapid test to distinguish between mild
and severe strains is adapting the technique of 'genetic finger-printing' to this
purpose (Heale, personal communication).
It is becoming increasingly clear, as a result of these various procedures, that
there is now a continuum of strains ranging in pathogenicity from very mild to
very severe with no clear-cut division into two distinct classes.
The increased range of pathogenicity now encountered may be the result of
genetic recombination between strains. Hastie (1962) first demonstrated that
Verticillium albo-atrum from hops could undergo such recombination.
Clarkson and Heale (1985b) studied heterokaryon compatability between
isolates and concluded that there was greater genetic homology between
fluctuating isolates than between progressive strains and that V2 and V3
isolates showed the greatest divergence from other isolates.
164 Fungal diseases
Because the distinction between fluctuating and progressive strains has
apparently been eroded, Talboys (1985) proposed that the time had come for
all cases of wilt to be treated in the same way. He suggested that the risk of
even more virulent strains arising is not necessarily greatest amongst the
present range of 'super-virulents' but might equally arise from the mild types.
He suggested that the West Midland outbreak might have originated in this
way and not from the movement of an existing strain from Kent.
Observations in the field have indicated that, although the level of infection
in Fuggle hops is not affected by temperature or rainfall, disease expression in
other cultivars is greatest in years when soil temperatures in the spring are low
(Talboys and Wilson, 1970). This was confirmed by inoculating hop plants that
were growing in temperature-controlled tanks (Sewell et al., 1978).
Although English hop growing suffered the first, and the most virulent
outbreaks of wilt, it has since become a problem elsewhere. In the Federal
Republic of Germany the first report of wilt was by Zattler (1960b) who stated
that there had been 55-60 ha affected in 1956 which had increased to 120 ha by
1959. Of the traditional varieties being grown, Spalter was the least and
Hallertauer the most susceptible, Hersbrucker and Rottenburger being
intermediate. The examination of a number of isolates indicated that these
were all of equal virulence (Zattler and Chrometzka, 1960) and there has since
then been no indication of increasing virulence similar to that in England. The
disease has, however, spread very widely through the Hallertau district, the
main hop growing area, and has led to a drastic reduction of the local
Hallertauer variety.
The English cultivar Northern Brewer is highly susceptible to progressive
wilt in England but highly resistant in Germany suggesting that the strain there
is equivalent to a severe fluctuating type in England to which the cultivar has
also shown resistance.
In the DDR the disease was first recorded in 1966 on an 11 ha holding in
which there were three main foci of infection. There have also been reports of
the disease in Poland, Belgium, Yugoslavia and France.
In New Zealand, Christie (1956) isolated the fungus from hops of the
Californian variety that were planted to follow a potato crop. He compared his
isolate with a fluctuating strain obtained from England and found his to be
more pathogenic to Californian and less pathogenic to Fuggle than the English
isolate. In the USA, Zehsazian (1968) found that the strain isolated from hops
in Oregon was not a virulent pathogen to that crop, being better adapted to
other species. In none of these countries does the disease appear to be as
serious a problem as it is in England or Germany.
Verticillium wilt: Verticillium albo-atrum 165
7.3.2 Hygiene
Keyworth (1942) demonstrated that diseased bine or leaves and the soil from
around diseased plants were all potential sources of infection and he described
many of the farming operations, such as cultivations and movement of plant
material, that could spread the disease. Sewell et al. (1962) studied the effect of
composting on the infectivity of the waste material from a picking machine by
collecting it in an enclosure that was formed by brick walls on two sides and
straw bales on the other two. They tested waste material from different
positions in the heap 21 weeks after harvest and found 99% of the samples to
be non-infective, the exception coming from the outside of the heap. On most
farms the waste is left in an unenclosed heap so there is likely to be a greater
volume of material that has not reached a sufficiently high temperature for the
fungus to be destroyed.
Because there is no chemical control available and because the disease can
very easily be carried in mud or plant remains, scrupulous attention to hygiene
has been the only available method of minimizing its spread. Where the disease
is endemic there has been little incentive for farmers to undertake tedious and
time-consuming control measures, but in the wilt-free areas growers have gone
to great lengths to prevent the disease appearing on their farms and this has
been reinforced by government action.
The legislation for the control of wilt restricts the movement of plants, used
hop poles, etc., from areas where wilt is endemic to eradication areas. Every
care should be taken to ensure that vehicles are not driven from infected areas
onto land in clean districts. Visitors to uninfected farms should be made to
wear clean footwear before entering any hop gardens.
On a farm where only some areas are affected separate implements should, if
possible, be kept for the diseased and healthy gardens. Otherwise each round
of operations should start in the clean areas and on leaving the infected areas
the tractor and implements should be carefully washed down.
Diseased hills should be carefully cut down and all the bines and as much of
the leaf material as possible bagged up before removing it for burning. Infected
bines may not always have developed wilt symptoms by the time they are
harvested so all the waste collected at the picking machine should be regarded
as highly dangerous. It should not be allowed to blow around nor be returned
to hop gardens though after composting it could be used in orchards or arable
land.

7.3.3 Cultural methods

Sewell and Wilson (1974) found that, compared with normal tillage, non-
cultivation reduced wilt by some 28% on sites where the incidence was high.
They attributed the reduction to the lack of root damage and the action of the
166 Fungal diseases

simazine herbicide treatment which eliminated weed hosts of the fungus and
inhibited surface rooting by the hops. On sites where the initial wilt incidence
was low, however, it increased slightly under non-cultivation and they
suggested that this was due to differential effects on soil nitrogen.
Since cultivations also spread infected material, non-cultivation was very
widely adopted in England as a means of reducing the incidence of wilt. The
firmer soil surface and better drainage under this system also made it far easier
to clean tractors and sprayers which might otherwise spread diseased material
from infected gardens.
The importance of the level of nitrogen treatments to the incidence of wilt
was demonstrated by Sewell and Wilson (1967) who compared rates of 235,
157 and 78 kg/ha (210, 140 and 70 lb/ acre). The 157 and 78 kg/ha treatments
reduced the incidence of wilt by 25% and 60% respectively compared with the
heaviest application.
There have been many similar observations on the influence of nitrogen
levels. In Germany, for example, Kamm (1970) reported that wilt was
increased by high rates of nitrogen fertilizers but was suppressed by rich
humus. This conflicts with observations in England where some severe
infections of resistant varieties have followed heavy applications of farmyard
manure. In France Marocke et al. (1977) found that wilt was most severe when
nitrogen was applied as calcium nitrate and less if ammonium sulphate was
used. The level of infection was least when urea was used and, in one instance,
it even gave an apparent cure.
On farms where wilt is established there is little more that the grower can do
by cultural management to minimize the disease. On farms where efforts are
being made to eradicate it the management of infected areas, after the diseased
plants and those close to them have been grubbed, is most important.
Trials conducted by Sewell and Wilson (1966) showed that the most effective
way of eliminating the pathogen from infected soil was to eliminate all the
broad-leaved plants that could act as hosts. When tomatoes were planted in
test sites after a four year bare fallow there was a very low level of infection
while on plots that had been under grass for the same period there was none.
When hops were used to detect residual infection this was found to be very low
after two years and nil after three to five years under a weed-free grass sward.
In eradication areas, plant health regulations in England required, for many
years, that infected areas should be grassed down, fenced and left in that state
indefinitely. More recently, growers have been allowed to bring such areas
back into production after a suitable period under grass. It is most important
for grassing down to be effective that the sward be kept free of broad-leaved
weeds, if necessary by the use of selective herbicides.
Verticillium wilt: Verticillium albo-atrum 167
7.3.4 Chemical control
Keyworth (1942) applied a range of soil disinfectants to infected soil which was
then left for two to three weeks before planting Fuggle hops to test for residual
infection. None of the treatments were completely effective though 2%
formalin at 361/m2 (8 gal/yd 2) gave the best result. Formalin has proved very
effective in the treatment of 'apple replant disease' and this success may be
related to the rapid recolonization of the soil by Trichoderma spp. (Freeman,
1980) which can be antagonistic to some pathogens. It has been suggested that
formalin treatment might be worth looking at again in the light of the apple
experience.
Sewell et al. (1971) detected benomyl at very low levels in xylem sap of pot
plants to which it had been applied, but not in mature field-grown plants. The
treatment controlled wilt in the pot plants but not in the field. Other fungicides
that have been assessed for their effectiveness against wilt have included
metalaxyl, fosetyl-aluminium, thiabendazole and some experimental mater-
ials. Although some treatments have reduced wilt infection when applied to
pot plants they have been ineffective in the field and some have, in addition,
proved phytotoxic (Chambers et aI., 1982, 1983).
A major problem with such treatments is that the hop is so deep rooted that
it is difficult to incorporate the fungicide to sufficient depth to reach all the
infected layer. A systemic fungicide that was effective against the pathogen and
would translocate downwards into the root system might give control but most
systemic materials only move upwards with the sap flow.
Because high soil nitrogen levels favour wilt development trials have been
carried out with nitrification-inhibiting compounds. No difference was noted
with ammonium nitrate but when urea was used as the source of nitrogen there
was a reduction in infection in the presence of the inhibitor (Chambers, 1987).

7.3.5 Biological control

Wilderspin et al. (1983) initiated a programme to investigate three possible
approaches to biological control of wilt. One was to elicit resistance of hops by
treating the roots with heat-killed conidia or with culture filtrates. Another was
to increase the antagonism of other organisms by organic soil amendments
while the third was an investigation of the 'wilt-suppressive' effect observed in
some gardens. In their next report (1984) it was said that the addition of 1%
chitin to hop garden soil reduced the popUlation of Verticillium propagules by
10% after 48 days and that hops and antirrhinums grown in amended soil
showed 5-8% infection compared with 40% in the controls. Similar results
were obtained with a crude preparation of Laminaria seaweed.
Their next report (1985), on the other hand, stated that adding Laminaria
stimulated the germination of dark resting mycelium and sporulation so that
168 Fungal diseases
there were more propagules than in unamended soil although it remained to be
seen whether the rapid germination would lead to reduced viability after 12
months or more. Chitin, at 1% addition, was less stimulatory and inhibition of
germination occurred after two months while three months after a 5% addition
no propagules were detectable. The addition of chitin, Laminaria or yeast
waste as soil amendments three months prior to planting in Verticillium
infected soil reduced the number of wilted plants to about 20% of the controls.

7.3.6 Resistant cultivars

Once wilt has become established on a farm there is no control method that
makes it possible to grow susceptible cultivars economically and hop cultiva-
tion has only been able to continue in such areas as the Weald of Kent or the
Hallertau district of Bavaria by planting resistant varieties, the development of
which is described in Chapter 9.
Keyworth (1953) demonstrated by grafting experiments that resistance to
wilt was located in the root and not the bine and this was supported by the
results of stem inoculation. When a small volume of spore suspension was
injected into bines of the susceptible cultivar, Fuggle, they developed mild
symptoms, but when a large volume was injected the symptoms were severe
and the bines died above the injection point. The reaction of the grafted bines
presumably reflected the quantity of inoculum that was able to reach the bines
through the root system of the susceptible or resistant varieties on which they
were growing.
Talboys (1958a) found that the initial reaction of the hop to infection is a
lignification of epidermal and cortical cells in contact with the fungus. Where a
hypha penetrates a cell wall, lignin-like deposits build up on the inside of the
wall, extending the distance that the hypha has to penetrate. These deposits
form peg-like lignitubers through which the hyphae sometimes emerge but,
more frequently, are complete occluded. This mechanism is most pronounced
in sensitive hosts invaded by a mild pathogen and does not appear to be an
effective defence mechanism against more virulent strains. The more effective
defence mechanism is located in the endodermis, the cells of which, in resistant
cultivars, are strongly suberized. Rudolph (1968) reported that varietal
resistance to wilt is related to the distance between the root tip and the first
lateral root because the xylem above the lateral root is protected by the
endodermal suberin layer.
If the fungus penetrates to the vascular system this stimulates the production
of tyloses in the xylem vessels which help to limit the development of the
infection. Leaf necrosis is due to fungal toxins and not to tyloses restricting sap
flow (Talboys, 1958b).
No hop genotypes have been discovered that are immune to wilt and even
highly resistant cultivars can become infected, often without showing any
Verticillium wilt: Verticillium albo-atrum 169
symptoms. For this reason it was part of the eradication policy in England to
ban the planting of resistant cultivars in wilt-free areas for fear that they might
become infected and provide an unidentified focus of infection for other hops
in the neighbourhood.
This policy underwent some modification when a few growers were unable
to eradicate the disease from their holdings so that, in spite of grubbing and
grassing down each outbreak that occurred, further infections continued to
develop. Such gardens continued to represent a risk to neighbouring farms
because fresh diseased material was produced each year and it also became
uneconomic for them to continue in that way. These farms were, therefore,
licensed to plant resistant varieties and it was thought that by reducing the
amount of infected material that would result from this the risk to other farms
might also be reduced.
This judgement was supported by observations on the levels of wilt in Wye
Target, the most resistant cultivar so far developed in England (Chambers,
1985). It was found that in gardens of this variety that initially had high levels
of wilt symptoms, there was a rapid and continuous decline in the succeeding
years that could not be attributed to the effects of temperature or nitrogen
levels. It was suggested that this indicated that with a cultivar of such high
resistance, the amount of inoculum returned to the soil from new infections
was less than the amount destroyed by the activity of soil organisms.
This report was accompanied by a warning that planting highly resistant
hops in soils with a high level of infection creates strong selection pressure for
the development of yet more virulent strains of the fungus. It was
recommended that such varieties should only be planted into 'clean' land or
heavily infected land that is first 'cleaned up' by at least two years under grass.

7.3.4 Verticil/ium dahliae

Although this is a common pathogen of other crops such as potatoes, tomatoes
or strawberries it rarely attacks hops .. It is more likely to do so in warmer
conditions where it is much more widespread and it is occasionally reported on
hops from the USA and also from New Zealand.
It can occur elsewhere and since the symptoms are similar to those of V.
albo-atrum it can be a cause of alarm where steps need to be taken to eradicate
that disease. For this reason it is advisable not to plant hops immediately
following crops such as potatoes and strawberries which are liable to provide
sources of infection.
A particular problem occurred in the USA where soil had been treated with
the insecticide heptachlor in past years when other crops were grown. Skotland
and Romanko (1987) record that in hop fields in which there were heptachlor
residues of 0.02 and 0.03 ppm, nearly 100% of the hop plants became infected
with V. dahliae whether the cultivar was normally susceptible or resistant to
170 Fungal diseases

the pathogen. In Idaho, the insecticide is used as an aid in the routine testing of
selections for resistance to Verticillium by growing them in a field that had
been treated with it.

7.4 OTHER FUNGAL DISEASES

7.4.1 Fusarium canker: Fusarium sambucinum

The commonest form of damage from this disease is found at the base of the
bine which is girdled so that only the innermost core is left attached. This
constriction causes the bine to wilt and the reason for this is readily identified
by the way the narrow neck can be broken by quite a gentle pull.
The fungus usually invades through a damaged point and this may be where
the bine joins the rootstock. If there is much movement of the bine at this point
a crack can develop providing a point of entry. In gardens that are cultivated
there is usually some mounding of soil about the base of the bine which is
supported so that cracking is less likely to occur than on non-cultivated hops
which lack this support. The mounded-up earth may also cover any cankered
area that has developed and roots will then grow from above the canker which
will help to keep the bine from wilting.
The bine above the canker will usually become thickened because the
downward flow of carbohydrates to the rootstock is interrupted. Although
Verticillium wilt can also cause the bine to fatten, the two diseases can quite
easily be distinguished. With canker the wilted leaves become flaccid whereas
on Verticillium-infected bines they are stiffer and drop off easily while the
pulling-off of the bine at the base is typical of canker.
Normally no special control measures are required other than good hygiene,
any diseased material being cut off and removed. In the early 1970s there was a
major replanting programme in England during the course of which there were
a number of complaints about rotting of setts between their delivery to the
farm in the autumn and planting in the spring. The setts arrived looking
healthy and were bedded in until required but when lifted for planting many
were found to be rotten. Fusarium sambucinum was identified as the cause of
the problem but investigations by Royle and Liyanage (1976) could find no
evidence of the plants on the propagators' holdings being infected. Their
experiments were rather inconclusive because the problem only occurred
sporadically but it did appear that the setts were particularly vulnerable if they
were allowed to dry out. On the basis of information on hop canker from
Germany and of the disease on other crops, they suggested that dips or
drenches with fentin acetate and maneb or benomyl might be useful.
A few years later a problem arose on the cultivar Yeoman, when hill deaths
became frequent after the setts had been planted, most deaths occurring during
Other fungal diseases 171
the next winter. This cultivar also developed a lot of bine canker and it is
suspected that F. sambucinum was involved in both these problems.
Trials were carried out to attempt chemical control of the problem and there
were encouraging results from basal sprays with a formulation of
thiabendazole (Darby, 1984a).

7.4.2 Black root rot: Phytophthora citricola

A black root rot of hops caused by Phytophthora citricola has been a serious
problem in New Zealand for many years although the development of resistant
cultivars has reduced its significance there (section 9.2). In Australia the same
pathogen was reputed to be the cause of a black root rot which became much
less common following the establishment of Pride of Ringwood as the
principal cultivar. In a recent study, Skotland (personal communication)
consistently isolated Pythium intermedium from rotted crowns of Pride of
Ringwood whose storage roots had typical 'black rot' symptoms. He did not
isolate Phytophthora citricola and it is possible that, in the past, this was
erroneously assumed to be the causal organism in Australia because it had
been identified as such in New Zealand. P. citricola has, however, been
identified as the cause of black root rot in South Africa, where it has been a
serious problem, and in England, where it is only rarely serious (Royle, 1966).
It is a soil-borne fungus that invades the rootstock at or near the crown. The
water-conducting vessels of the rootstock become blocked with fungal hyphae
and tyloses and the infected area becomes watersoaked and blackened and
initially is clearly delimited from neighbouring healthy tissue. The first sign of
infection of the rootstock appears late in the season when the bines suddenly
wilt in a soft, flagging or drooping manner. This type of wilt can be confused
with canker but bines infected with black root rot do not pull-off like cankered
bines.
The disease is too infrequent in Europe for any chemical control measures to
have been adopted. In New Zealand and South Africa there were trials with
soil disinfecting agents but the development of resistant cultivars has largely
solved the problem.

7.4.3 Grey mould: Botrytis cinerea

This is a disease of hop cones that is reported to have become increasingly
serious in Japan since 1952 (Sakai, 1976) but elsewhere it is only a problem if
conditions are very favourable to its development. Although reports of it in
Germany go back to 1953 it was not of importance until around 1970 when it
forced some growers in the Hallertau district to harvest their hops early before
the disease caused too much damage. It was originally thought that the disease
only developed after the cones had suffered from insect damage but in more
172 Fungal diseases
recent outbreaks insects have not been involved (Kohlmann and Kastner,
1975).
It is favoured by wet conditions during cone development and appears as a
grey, fuzzy mould, starting at the tip of the cone where moisture persists for the
longest period of time, and causing the cones to turn brown. The bracteoles are
more susceptible to damage than the bracts and so the cones often have a
striped appearance.
Only in Germany are any fungicide treatments officially approved for the
disease on hops, the materials recommended there being dichlofluanid,
procymidon and vinclozolin.

7.4.4 Alternaria alternata

This fungus is not normally recognized as a pathogen of hops so there is no
common name to describe it. In September 1982 there was a widespread
discoloration of hop cones in England which was at first attributed to late
infection by powdery mildew. Since cultivars resistant to the mildew were also
attacked, this diagnosis was unlikely and, on examination of affected cones, A.
alternata was the only pathogen to be consistently recovered. Inoculation of
healthy cones with the fungal isolates produced similar symptoms and it was
found that infection was favoured by high humidity and by injury to the cones
such as would be caused by high winds. As with Botrytis, it was the bracteoles
which were most prone to infection and so the cones developed a similar
striped appearance (Darby, 1988). In the case of A. alternata infection,
however, there was none of the grey mould typical of Botrytis.

7.4.5 Black mould: Cladosporium

This is another relatively unimportant disease first described by Salmon and
Ware (1936). It also causes a brown discoloration of the cones but in this case it
is the bracts that are discoloured and the bracteoles that remain green so the
striped effect is the reverse of that seen on hops infected with Botrytis or
Alternaria.

7.4.6 Armillaria root rot: Armillaria mellea

Hops are only liable to be attacked by this fungus when there is a source of
infected wood nearby since it is primarily a disease that affects trees. The
fungus grows out of tree roots with long round rhizomorphic strands, from
which it gets its name of the 'bootlace fungus', and these spread through the
soil below the surface.
Salmon and Ware (1937) described an outbreak in which the infection came
from an oak stump some 7 m from the nearest hop hill to be infected.
Other fungal diseases 173
Hedgerows can be another source of infection while one instance was noted at
Wye where a wooden stump used to mark an experimental plot was the cause.
The problem is an infrequent one but hops planted on old orchards are a
special risk since this is one of the rare occasions when they will be planted
after trees.
CHAPTER 8

Virus diseases

Because commercial hop gardens are invariably planted with vegetatively

propagated material the stocks are liable to accumulate virus infections. The
traditional hop varieties have been in cultivation for a very long time, possibly
for as long as 200-300 years. In England the Fuggle variety, which is a
comparative newcomer, has been grown for over 100 years. In view of their
long history it is surprising that virus problems have not been more severe.
The first account of a virus disease in hops is probably the description by
Percival in 1895 of what he called the eelworm disease of hops although the
plants were also described as 'nettle-headed' or 'skinkly'. The problem is now
called nettlehead and is known to be a virus disease. Since it was not until 1898
that the first virus (tobacco mosaic virus) was isolated it is not surprising that
Percival failed to identify the true cause of the problem. It was not until 1966
that Bock was able to identify arabis mosaic virus in hops and suggest that it
was implicated in the disease.
The identification of the viruses infecting hops has been one of the most
difficult problems facing hop research workers and only recently has a fairly
complete picture of the situation emerged. The work has been greatly facilitated
by the development of the enzyme-linked immunosorbent assay (ELISA)
method of determining the presence of viruses in plants which was very rapidly
adopted for use with hops (Thresh et al., 1977; Barbara et al., 1978).
Just as important for the control of the diseases, once they had been
identified, has been the elimination of the viruses from infected plants by heat
therapy and meristem-tip culture (Vine and Jones, 1969; Schmidt et al., 1973;
Adams, 1975). The use of these two techniques combined with the Ministry of
Agriculture A plus certification scheme for hop propagators has transformed
the health of the English hop crop and similar measures have been adopted
elsewhere.
When the very tip of an apical shoot is dissected out and grown on under
sterile conditions, plants often can be regenerated that are virus-free. Virus
mUltiplication is inhibited by maintaining plants at a high temperature and a
combination of heat treatment and meristem-tip culture has made it possible to
obtain virus-free clones of all the important cultivars.
176 Virus diseases
The ELISA technique uses an antiserum for the particular virus under
investigation and a quick and reliable colour reaction determines within two
days whether or not the virus is present in the sample. The method is much
quicker and more reliable than the older method of inoculating indicator
plants of other species that are virus sensitive and many more samples can be
tested.

8.1 HOP MOSAIC VIRUS (HMV)

Hop varieties are either sensitive to this virus or else, when infected, the
plants show no symptoms but carry the virus and can be foci for further
spread. Whether infected or not such varieties are referred to as 'carriers'. The
typical symptoms on sensitive hops first appear as pale vein-banding of the
leaves (Figure 8.1) which then become mottled and strongly down-curled.
The down-curling is normally a good diagnostic feature to distinguish HMV
from nettlehead in which the leaves curl upwards, but the Australian cultivar,
J78, is an exception. It very rapidly became infected with HMV when grown
at Wye although, because the leaves curled upwards, nettlehead was first
suspected.
Diseased hills can survive for several years during which time they are
stunted with short internodes and the bines fall away from the strings.
Production from such plants is minimal and they should be grubbed as soon as
possible to prevent them acting as sources of infection.
Adams and Barbara (1980) purified the virus and found it to be a member of
the carlavirus group with filamentous particles c. 14 x 650 nm composed of
c. 6% single-stranded RNA of mol. wt c. 3.0 x 106 and a single protein species
of mol. wt c. 34 000. It is a distinct member of the group though distantly
related to hop latent virus and others.
Although the damson-hop aphid (Phorodon humull) was thought to be the
most likely vector of the virus, Massee (1943) failed to demonstrate trans-
mission while Paine and Legg (1953) obtained transmission with the spring
alatae but failed to do so with apterae. Bock (1967) recorded the spread into
650 seedlings from a mosaic-sensitive cross that were exposed in batches of 50
for 24 h periods during the spring migration of P. humuli with yellow traps
distributed amongst the seedlings. Only 22 of the seedlings developed mosaic
but from correlations with the aphid species caught in the traps Bock
considered that Macrosiphum gei was the probable vector. Adams and
Barbara (1980) showed that the apterae of the hop aphid can transmit the
virus, as can those of Myzus persicae and Macrosiphum euphorbiae. It seems
therefore that HMV is similar to other carlaviruses in having low vector
specificity, being transmissible by numerous aphid species.
Transmission can only occur when the aphids move from one hop plant to
another and apterae seldom do this. Even though the alates of P. humuli are
Hop mosaic virus (HMV) 177

Figure 8.1 Hop mosaic (Crown copyright).

evidently inefficient vectors they are very numerous and it is possible that they
are the cause of most of the spread although other species which rarely settle
after alighting on hop plants might be important. Once hop aphids have
alighted on a hop plant they are more likely to move if they find the host
unsatisfactory. They may then fly to another plant but such flights are short so
the isolation required to prevent spread between susceptible and carrier
178 Virus diseases
varieties is only about 100 m (Thresh and Adams, 1983). It is much more
important to ensure that none of the plants in a garden of a sensitive cultivar
are carriers. Problems were sometimes experienced because males that were
infected carriers were planted amongst sensitive Golding hops, leading to a
conspicuous patch of diseased plants around the male (Keyworth, 1947).
Field trials of mosaic-free and infected material of carrier varieties have been
carried out at Wye College and Rosemaund Experimental Husbandry Farm in
Hereford and no consistent effects on yield or a-acid found (Thresh and
Adams, 1983) although in one of the trials with Wye Saxon there was some
evidence of a fall in production in the year in which inf~ction was detectable
followed by a recovery in subsequent years (Neve and Lewis, 1979). In
Australia, preliminary results indicate there may be severe yield reduction in
Pride of Ringwood infected with HMV (Lewis, 1988).
The striking feature of two of these trials was the difference between the two
cultivars, Wye Saxon and Wye Northdown, in the rate at which reinfection
occurred. In the Saxon trial about half the meristem plants were infected
between planting in 1975 and harvest in 1976 and by 1978 most were infected
and the trial had to be abandoned (Neve and Lewis, 1977, 1979). The Wye
Northdown was planted in 1980 and seven plants were found to be infected by
1981 but there was only one additional case in 1982 and none in 1983 (Neve
and Darby, 1983, 1984a). It is suggested that the differences reflect aphid
preference for the two cultivars, with Wye Northdown being so acceptable that
aphids remain on the plant on which they first alight.
Hop mosaic was first described by Salmon in 1923 but examination of his
field records indicates that for some years he had mistakenly diagnosed the
problem as nettlehead and cases of mosaic infection in his breeding material at
Wye can be traced back as far as 1906.
Mackenzie et aZ. (1929) described how a grower in Kent obtained cuttings of
the Hallertauer variety from Germany between 1904 and 1908. These were
planted with the English variety Cobbs on either side and, although the
Hallertauer hops remained healthy, the Cobbs adjoining them suffered from
mosaic disease. Similar events followed the distribution from Wye of two
seedling varieties to farms where they were planted amongst Golding hops. An
extensive series of intervarietal grafts was carried out to determine which
varieties were sensitive to the virus and which were carriers or potential
carriers. All the clones of the true Goldings were sensitive as was the 'Golding
Variety' Tutsham. Although the disease had not been reported from Germany
they found that some hops of German origin, though not exactly identified,
were sensitive, as were hops obtained from France labelled 'Alsace' and 'Spalt'.
The identity of these two sorts must be suspect since there is no evidence that
genuine stocks of these varieties are sensitive.
Since sensitive cultivars formed a large part of the English hop area at the
time of these reports and the disease had not previously been reported, it must
Hop mosaic virus (HMV) 179
be concluded that it was introduced early this century. Apart from the
importation of Hallertauer hops mentioned above, Mackenzie et a1. (1929)
stated that a garden of German varieties existed at Wye College before 1900
and that other hop growers had also introduced them. They found that most
introduced hops were either carriers or potential carriers but were unable to
say whether the carriers were infected when introduced or had become infected
subsequently. All the female hops from the USA were carriers although the
Red Vine obtained from Canada (but which had originated from New York)
sometimes showed atypical virus symptoms that they termed 'masked mosaic'.
On the other hand all the male hops they obtained from the USA were
extremely sensitive so that there was great difficulty in maintaining the stocks
at Wye.
An atypically mild strain of hop mosaic was described by Legg (1959a) in
which symptoms, consisting of a light green or yellow mottle, did not appear
on young leaves until they were fully expanded. The bines continued to climb
but vigour was impaired. Flowers were produced normally but the cones were
small and loosely formed. Infection with the mild form provided cross-
protection against infection with the severe strain.
Hop mosaic has, until recently, attracted little attention outside England,
but the spread of verticillium wilt in Germany has led to the replanting of large
areas of the Hallertauer Mittelfruh variety with Hersbrucker Spat hops. This
has been accompanied by reports of mosaic infection in the Hersbrucker
variety. Kremheller et a1. (1989) recorded the incidence of HMV in plants that
were virus-free when planted 3-5 years earlier and expressed surprise that only
1.3% had become infected but they gave no indication of the proximity of
sources of infection. They found no effect of infection upon yield or a-acid
contents.
They contrasted the situation in 1956, when Zattler found mosaic symptoms
in 3% of plants of Hersbrucker Spat in the Hersbruck region but none in
Hallertauer Mittelfruh, with what had happened in the 1970s when such
symptoms began to appear in the Hallertau region in Northern Brewer,
Brewer's Gold and Huller Bitterer. Subsequently the symptoms were observed
in Perle and Orion. All the plants with these symptoms that were tested were
found to be infected with HMV, but other infected plants were symptomless.
Tests showed that the symptoms were not associated with infection with Hop
Stunt Viroid (see section 8.7.1) and they were unable to establish the cause.
In Australia, Munro (1987) carried out a survey on the main cultivar, Pride
of Ringwood and found levels of infection ranging from 1-68%. Since the
damson-hop aphid is not present in Australia there must be some other vector
operating there since Pride of Ringwood was bred in Australia and there is no
evidence that the virus is seed transmitted.
180 Virus diseases
8.1.1 Control
Where mosaic-sensitive varieties of hops are grown it is important to ensure
that the planting material comes from a mosaic-free stock and that there are no
admixtures of plants of a carrier variety. When replanting a garden that
previously grew a carrier variety, great care must be taken that none of those
plants survive to act as sources of infection. The garden should be isolated
from others containing carrier varieties but the distance between them need not
be very great - it would appear that 100 m is probably sufficient to avoid
infection being transmitted from one to the other. Any male plants to be
included in the garden should come from a known mosaic-sensitive source.
If any plants do become infected they are easily recognized and should be
grubbed as soon as possible but their position should be noted. If there are
rogue carrier plants in the garden they will tend to infect the plants in their
immediate neighbourhood and a record of where infection has occurred can
help to identify the source.
Since there is no clear evidence of any reduction in yield in carrier varieties
infected with the virus there is no justification in going to the expense that
would be involved in raising mosaic-free stocks and ensuring that they could be
mUltiplied without becoming reinfected. It is accepted in the UK, therefore,
that material of carrier varieties within the A plus certification scheme are
infected with mosaic.

8.2 HOP LATENT VIRUS (HLV)

Adams and Barbara (1982) found that virtually all commercial varieties in
England are uniformly infected with this carlavirus, the only exceptions being
some of the meristem clones most recently released to propagators. The
filamentous virus particles of this virus (Figure 8.2) were c. 14 x 675 nm
composed of c. 6% single-stranded RNA of mol. wt c. 2.9 x 106 and a single
protein species of mol. wt c. 33 000.
The virus is transmitted by the damson hop aphid and is widespread in
Europe, the USA and Australia but there is no evidence that it has any effect
upon the productivity of infected hops and no control measures are
advocated.
Munro (1987) also checked for this in Pride of Ringwood and recorded
levels ranging from 1-58%.

8.3 AMERICAN HOP LATENT VIRUS (AHLV)

This is the third of the carlaviruses known to infect hop plants. First reported
by Probasco and Skotland (1978) in North America, it was subsequently found
at Wye College in some recent introductions from the USA although it was not
American hop latent virus (AHLV) 181

Figure 8.2 Virus particles of hop mosaic and hop latent viruses distinguished by
'decoration' with hop mosaic antiserum (IHR, East Mailing).

present in any of the older introductions from there. Eppler and Sander (1980)
carried out a survey of hops in the BRD following the report of AHL V having
been found in England but only found it in a hop clone introduced from the
USA and growing in an isolated breeding garden. A check on the neighbouring
seedlings did not disclose any indication of the virus having spread. It has also
been found in New Zealand where 9-10% of the plants tested were infected
(Hay, F . S., unpublished).
The infected material at Wye had been planted out in an experimental
garden but it was grubbed as soon as the infection was identified and, although
this virus is also transmitted by the damson hop aphid, checks of neighbouring
plants failed to detect any evidence that it had spread from the original source.
While very little is known, at this stage, about the effects of the virus on
infected plants, it is obviously desirable that every effort should be made to
prevent it from becoming established outside the USA.
Virus preparations contained filamentous particles c. 15 x 680 nm composed
of c. 6% single-stranded RNA of mol. wt 3.0 x 106 and a single protein species
of mol. wt c. 33 000.
182 Virus diseases
8.4 ARABIS MOSAIC VIRUS (AMV)

Nettlehead disease has been a major problem for hop growers for a very long
time. Percival (1895) attributed the cause to the nematode Heterodora schachtii
but Duffield (1925) eliminated nematodes as the cause by demonstrating that
they occurred as frequently in the soil surrounding healthy plants as those that
were diseased. It was not until much later that Bock (1966) suggested that arabis
mosaic virus (AMV) was the cause. He identified 30 nm polyhedral virus
particles found in hops as AMV and roughly spherical particles of c. 25 nm
diameter as necrotic ringspot virus (NRSV). He found that AMV was always
present in nettlehead-diseased plants but not all plants with AMV showed the
disease symptoms. A rod-shaped virus that he also detected was not associated
with the disease. He suggested that nettlehead was the result of dual infection
with AMV and NRSV and mentioned that the nematode Xiphinema
diversicaudatum was present in .low numbers in most gardens but extremely
numerous in three gardens where nettlehead was prevalent.
Schmidt et af. (1972) similarly identified AMV and apple mosaic, which is a
form of NRSV, in hops suffering from nettlehead and concluded that the
disease was caused by the combination of those two viruses.
Subsequent work confirmed the role of AMV in nettlehead but not Bock's
suggestion that NRSV was also involved although it became evident that there
must be a second element to the disease since all nettlehead plants were
infected with AMV but not all AMV-infected plants developed nettlehead.
Clark and Flegg (1979) found that a satellite consisting of a low molecular
weight species of nucleic acid (SNA) was consistently associated with AMV in
nettlehead plants. Subsequent reports (Clark and Davies, 1980, 1981; Davies
and Clark, 1982, 1983) described a second species (SNA-2) and continued to
find a good correlation between plants with nettlehead or stunting and the
presence of SN A in the AMV preparations.
Nettlehead usually spreads slowly but tends to be particularly serious
alongside hedgerows, where hedges have been grubbed and after orchard crops
or permanent pasture (Keyworth and Davies, 1946; Keyworth and Hitchcock,
1948). These observations suggested that it is soil-borne, but it was many years
before the vector was identified as the dagger nematode Xiphinema divers-
icaudatum (Valdez et al., 1974).
An unusually rapid spread of the disease was recorded on a farm in
Worcestershire where hops were planted in the winter of 1968-9 with locally
produced setts on a site that was formerly an old pasture. Nematodes were
widely distributed and AMV was probably introduced in the planting material.
Infection increased from 2.9% ofthe plants in 1970 to 43.6% by 1977, the newly
infected plants usually being adjacent to those that had earlier shown
symptoms (Adams et al., 1978). By 1984 over 70% of the plants were showing
symptoms (Thresh and Adams, 1985).
Arabis mosaic virus (AMV) 183

Figure 8.3 Nettlehead (IHR, East Mailing).

Thresh et af. (1972) described three distinct diseases, nettlehead, split leaf
blotch and bare-bine that could result from infection with AMV. Subsequently
Adams et al. (1987) identified AMV as the cause of hop chlorotic disease also.

8.4.1 Nettlehead
This is the most serious of . the three diseases associated with AMV and
apparently results from infection with the virus in association with SNA.
Infected plants have weak bines which climb badly, the tops falling away from
the strings. The internodes are shortened, the leaves are small with margins up-
curled, in contrast to mosaic in which they are normally down-curled. The
cones are small and distorted and the crop is reduced by as much as 75% (Legg,
1959b) (Figure 8.3).
184 Virus diseases
Symptoms do not usually appear until the year after the hill becomes
infected and although the symptoms from then on are variable, diminishing in
hot weather, there is a progressive weakening of the plant although they do not
normally die. The centre of the stock tends to become rotten and new buds are
confined to a rim of living tissue.
A severe outbreak of nettlehead was reported in 1983 on the variety Yeoman
which had been planted following Bramling Cross, a hop which has long been
known to have considerable tolerance of infection compared with others such
as Fuggle and W.G.V. The Yeoman plants were severely stunted with very
conspicuous nettlehead symptoms whereas previously there had been no
obvious problem at the site with the Bramling Cross (Thresh, 1984).

8.4.2 Split leaf blotch

This disease was given its very descriptive name by Salmon and Ware (1934) on
account of the translucent, yellow, oily blotches which develop on some leaves
of infected plants. As the leaves expand the blotches frequently split (Figure
8.4). Fuggle is the only variety to suffer from this disease to any noticeable
extent and this may be because infection accentuates the tendency of even
uninfected plants to show some slight blotching. Legg (1959b) recorded
reductions of yield of infected plants of about 50% and Thresh et al. (1972)
suggested that in Fuggle it might cause greater losses than nettlehead because,
although less severe, it was much more common.

8.4.3 Bare bine or spidery hop

The majority of plants infected with AMV show bare bine symptoms in the
spring even though they may subsequently recover and appear to be healthy.
Affected plants are weak when growth commences in the spring and have a
spidery, bare appearance compared with normal hills (Figure 8.5). Few shoots
develop and these have a characteristic curvature, dark colour and small,
retarded leaves. The difference disappears within a few days and is no longer
recognizable after the hills have been trained and the excess shoots removed
(Thresh et al., 1972).
The condition was not recognized until recently because the symptoms are
very transient and they were only noticed after serological tests had made it
possible to compare the growth of large numbers of plants known to be
infected or free from AMV. With experience, however, the symptoms can be
recognized without too much difficulty.
The only evidence about the effect of this type of infection on yield is from a
trial on the variety Bullion in which plants showing bare bine symptoms, and
subsequently confirmed as being infected with AMV, yielded some 25% less
and had a-acid contents about 8% lower than uninfected plants so that their
Arabis mosaic virus (AMV) 185

Figure 8.4 Split leaf blotch in cv Fuggle (IHR, East MaIling).

total a-acid production was reduced by 30% (Thompson and Neve, 1971).
Although bare bine causes far less damage than nettle head to affected plants, it
is much more prevalent so that its overall effect may be at least as serious.

8.4.4 Hop chlorotic disease

This disease was described in detail by Salmon and Ware (1930) but has rarely
been reported since. Leaves produced early in the season are severely distorted
and chlorotic, often with a characteristic parrot-beak shape (Figure 8.6).
Growth is reduced but the lateral shoots and upper parts of the main bine are
usually symptomless. The disease is transmitted by grafting, by sap inoculation
and to seedlings of infected plants. It has also been reported from Germany,
Russia and Poland (Schmidt and Klinkowski, 1965; Eppler and Sander, 1981).
Adams et al. (1987) investigated an outbreak in Kent and found that it was
caused by a distinct isolate of AMV. Although indistinguishable from other
hop isolates serologically or by inoculation to other herbaceous species, it
produced distinctive chlorotic symptoms in hop plants inoculated with purified
virus. The basis of the difference in pathogenicity was not established. Because
the symptoms and effects on growth are much more severe than the bare bine
condition, plants infected with this strain of the virus are unlikely t6 be used
for propagation and this probably accounts for its comparative rarity.
186 Virus diseases

(a)

Figure 8.S Early season growth of: (a) healthy plant; and (b) one showing bare bine
symptoms (IHR, East Mailing).
Arabis mosaic virus (AMV) 187

Figure 8.6 Hop chlorotic disease (IHR, East MaIling).

8.4.5 Control

Percival (1895) advised growers to avoid planting cuttings from infected

localities. This advice had little effect and nettlehead became so widespread
that, according to Keyworth (1945), it was difficult to find any gardens that
were free from it. In spite of the policy of grubbing infected plants, growers
found it necessary to abandon hop growing completely in some gardens
(Ogilvie, 1939).
A certification scheme was introduced in 1943, for which gardens were
checked primarily for freedom from verticillium wilt so that they could be
188 Virus diseases
used as sources of propagating material but HMV and nettlehead were
included in the inspection. Nettlehead was so widespread that it was not
possible to set a nil standard but only 'a limit of very low percentage of virus
infection' (Keyworth, 1945). Since then, however, the A plus certification
scheme has been upgraded to the stage where AMV-free clones of all cultivars
are available.
Arabis mosaic virus will only spread within a garden if Xiphinema eelworms
are present and there are diseased plants to act as sources of infection. The first
precaution, therefore, must be to plant only AMV-free material and when
Xiphinema are not present this is the only precaution that needs to be taken.
Where the eelworm is present it is equally important to ensure that there are no
plants surviving from a previous crop in which the disease was present. This
means that grubbing of the previous crop must be carried out with extreme
care, or that there is a sufficient interval between grubbing and replanting for
any ground-keepers to be seen and removed.
Although Xiphinema are widespread throughout England they are more
common on lighter soils and in perennial crops (Taylor and Brown, 1976). The
likelihood of them being present in a hop garden depends upon the previous
cropping of the land and on the soil type and there is a good chance of a garden
being uninfested. Where they are present they will carry the virus over when a
garden is replanted unless special precautions are taken.
Experiments at Rosemaund Experimental Husbandry Farm and elsewhere
(Pitcher and McNamara, 1976; Adams et aI., 1979) showed that fumigation
with 0-0, Oi-Trapex, dazomet or quintozene gave good control of the
nematodes in infested soils but even the most effective material, 0-0, did not
reduce their numbers sufficiently to stop re-infection of hops with AMV
(McNamara and Pitcher, 1974). This was found to be unimportant when
cultural treatments ensured that the nematodes had lost their infectivity. This
could be achieved by fallowing the land for two years during which time the
nematodes would moult and shed the cuticle lining of the oesophagus where
the virus particles are retained (Taylor and Robertson, 1970). The results of
fallowing were improved if it was combined with an initial fumigation and this
resulted in no reinfection of hop plants following a fallow of only 19 months.
Fallow periods of 19 or 20 months could be achieved by grubbing the previous
crop immediately after harvest and replanting in the months of Mayor June
using setts that had been held in a cold store. In this way only one full cropping
season was lost (Adams et ai., 1979).
In hop gardens where there are no Xiphinema there is no need to carry out
these procedures and virus-free material can safely be planted immediately
following hops that were even heavily infected. Unfortunately, soil tests can
only examine such a small sample, relative to the volume of soil occupied by
the roots of a hop plant, that a negative result does not necessarily mean that
the garden is free of the nematode.
Prunus necrotic ringspot virus (NRSV) 189
8.5 PRUNUS NECROTIC RINGS POT VIRUS (NRSV)

This virus was first detected in hops in the 1960s and it has since been
shown that it was present throughout all the main commercial varieties in
England at that time and no uninfected stocks were discovered. Hops in
Germany are not, at the present time, so totally infected since Eppler and
Sander (1981) found it in only 57% of 2112 plants examined although it was
found in 91 % of the gardens surveyed. The level of infection varied
significantly between certain varieties and this may be because, as in
England, recent plantings have been with NRSV-free material.
Using the ELISA technique, two serotypes of the virus have been detected
in hops. One is similar to apple mosaic virus (ApMV) and is referred to as
such by some workers while others designate it the 'A' type of NRSV. The
other appears to be a strain intermediate between 'A' type and the 'C' type
from cherry or plum and this has been designated 'I' type (Adams et al.,
1978). The authentic 'C' type has not been found in hops although refer-
ences to 'C' strains occur in the literature.
Under English conditions NRSV is normally symptomless in hops but in
1976, following a prolonged period of unusually hot dry weather, a con-
spicuous line pattern appeared in the leaves of several varieties in late July
to early August when temperatures were lower (Figure 8.7). A resurgence of
NRSV was detected in some plants and the incidence of the symptoms
appeared to be correlated with infection with that virus (Adams et al.,
1977).
In other countries NRSV is reported as causing ringspot-type symptoms on
hops. In the DDR, Schmidt (1967) reported that a disease described as hop
necrotic crinkle mosaic was caused by a serotype ofNRSV. In Czechoslovakia,
Albrechtova et al. (1979) detected ApMV in plants with ring and band patterns
on the leaves and similar symptoms are reported from Germany. Smith and
Skotland (1986) in the USA described two serotypes of NRSV, both of which
gave line pattern and ringspot. The symptoms were most prevalent in the
summer when cool periods followed warm periods, and in the fall when
temperatures were decreasing. Sano et al. (1985) isolated ApMV which, when
reinoculated produced chlorotic spots, ringspots and a band pattern accompa-
nied by leaf necrosis, symptoms that are prevalent in the field in Japan.
By 1969 a meristem clone of Northern Brewer and a clone of the new variety
Wye Northdown were available in England that were free of NRSV and a trial
was planted at Wye to compare these with clones infected with the virus. Trials
were also started with other varieties as suitable clones became available. These
trials showed that although the infection usually remains symptomless it does
have a significant effect upon crop production. The virus-free clones out-
performed the infected clones consistently as shown in Table 8.1.
190 Virus diseases

Figure 8.7 Line pattern (IRR, East Malling).

TABLE 8. 1 Increased production/rom NRSV1ree clones.

Yield a-acid a-acid, yield per

(% change) content unit area
(% change) (% change)

Northern Brewer + 3 +16 +21

Wye Northdown +16 +11 +27
Wye Target +10 + 4 +14
Wye Challenger +21 +10 +33
Bullion +18 + 6 +26

Kremheller et of. (1989) found that yield and a-acid contents were not
affected when plants were infected with HMV alone but that infection with
HMV and ApMV reduced yield by 4-38% and a-acid content by 18-26%.
The studies on NRSV in England coincided with a period of extremely rapid
replanting of new varieties and, because it was possible to release these largely
free of NRSV, this contributed a great deal to the increased production of a-
acid that resulted, estimated at the time as being worth an additional £1 million
a year in a crop worth some £16 million.
Other viruses 191
The rate of reinfection of NRSV-free plants has been recorded and usually
the only plants to acquire the virus are those growing adjacent to others
already infected. The vector, if any, has not been identified and spread might
possibly be purely mechanical by stem or root contact.

8.5.1 Control
Because the spread of NRSV is so restricted and there is no evidence of it
surviving in the soil or in a soil-inhabiting vector, it is one ofthe easiest disease
problems to control. All that is required is that the hop garden should be free
from any infected hop plants left after grubbing a previous crop and that the
new plant should be free of the virus. In these circumstances the benefits of
planting healthy material, which does not involve much additional cost, will
continue for a very long time.

8.6 OTHER VIRUSES

The discovery of AHLV in hops introduced to Europe in breeding material
from the USA indicates the care that must be taken to ensure that other
diseases do not become established as a result of such introductions. It is not
yet clear what effect AHLV may have on production but the evidence from
NRSV shows that absence of any symptoms does not necessarily mean that it
is not damaging.
Quarantine precautions are not difficult to operate against diseases that are
known to exist in other countries since methods of identifying them will be
available. The problem is discovering the presence of diseases, particularly
virus diseases, that have not previously been identified. AHL V could very
easily have become established in Europe if it had not been identified in
America shortly before it was exported.
Another recent example of a virus being introduced was the discovery of a
virus in seed of Humulus japonicus imported from China. Little is known
about this virus (that has been code-named humulus japonicus virus) except
that it is an ilarvirus and distantly related to NRSV. Since some other members
of that group are pollen-borne or mite-transmitted, there could be a danger of
it spreading into cultivated hops (Adams and Barbara, 1988; Adams et al.,
1989).
There are various reports of other viruses being found in hops including
alfalfa mosaic virus (Novak and Lanzova, 1976; Yu and Liu, 1987), tomato
bushy stunt virus (Novak and Lanzova, 1976) and tobacco necrosis virus
et
(Albrechtova al., 1979). While none of these may be of any significance, they
do indicate how easily new virus problems could be spread around the hop
growing countries of the world if the greatest care is not taken to check the
health status of imported plants.
192 Virus diseases
8.7 VIROIDS

Viroids are similar to viruses in some ways but consist of only a small piece of
RNA and have no protein coat and so cannot be detected serologically.

8.7.1 Hop stunt viroid (HSV d)

A stunt disease of hops has been reported in which the growth rate of all parts
of an infected plant is retarded and the wax deposits of the leaf cuticle are
modified. It has been shown to be caused by a viroid which has been named
hop stunt viroid (HSVd). Yield is reduced from ~ - 13 of healthy plants and
the lupulin glands are severely shrivelled (Momma and Takahashi, 1984).
Vectors are not known for viroids and transmission appears to be purely
mechanical. Takahashi and Yaguchi (1985) have described chemical and heat
treatments that will successfully decontaminate tools. HSVd has only been
recorded in Japan and in the Japanese variety Kirin 2 grown in South Korea. It
has been very largely eradicated by the rogueing of infected plants and by
avoiding transmission on contaminated tools. Although it has such a limited
distribution in hops it is much more widely distributed in grapes and Japanese
workers have suggested that it originally infected hops from grapes (Barbara et
al., 1988).

8.7.2. Hop latent viroid (HLVd)

A second viroid was identified by Puchta et al. (1988) and found to be
widespread in German hops and in many English varieties, although some of
the English hops tested had been grown in Germany for several years. It
appeared to be common in hops from other countries also. Although these
workers found the viroid in a number of stunted plants, these were also
infected with other viruses and they were unable to link symptoms with the
presence of the viroid and they therefore named it hop latent viroid (HLVd).
Subsequent work by Barbara et al. (1990a) in England showed an interesting
distribution of the viroid. All cultivars susceptible to verticillium wilt (except
for one old variety grown on only one farm) were infected to some extent, the
most heavily infected being Omega, the cultivar most recently released to
growers. No infection was found in commercial samples of the three most
widely grown wilt tolerant cultivars, although some of the less important wilt
tolerants yielded some infected samples.
Four plants of each cultivar are maintained in an insect-proof glasshouse at
Wye to provide the nucleus for commercial propagation stocks and the virus
status ofthese is regularly checked. When tested for HLVd infection, however,
it was found that in many cases all four plants were infected and, with one
exception, the remainder had some infected plants. The exception was the
Viroids 193
Australian variety 178 which was free of the viroid. These workers thought that
the difference in levels of infection between wilt-susceptible and wilt-tolerant
hops was more likely to be a chance effect following the recent introduction of
HL Vd into the nuclear stocks, rather than a direct relationship with levels of
susceptibility to wilt.
Further work (Barbara et al. 1990b) indicated that this viroid could be a far
more serious problem than had at first been thought. During sampling of the
heavily infected cultivar Omega, it was noted that some plants were greener
and more vigorous than surrounding plants and subsequent testing showed
that these were not infected while the less vigorous ones were. In the cultivar
Wye Northdown, however, no visual symptoms were seen that would dis-
tinguish infected plants.
Yields from infected Omega plants were about 35% less than from
uninfected ones and this was due to lower cone weight rather than fewer cone
numbers. Alpha-acid contents were significantly lower in infected plants but {3-
acids, total hop oils and myrcene were higher. These results suggested that
presence of the viroid accelerated the maturation of the hop cones. In Wye
Northdown the differences between healthy and infected plants were not so
great but the results indicated an overall reduction in a-acid production of 15-
20% compared with a 50% reduction in Omega.
Further work is required to confirm these initial results and to determine
differences in susceptibility of the various cultivars, but it has already been
suggested that the spread of this viroid through commercial hop gardens may
be partly responsible for the disappointing yields that have been a feature of
the late 1980s.
These results indicate the importance of establishing HL Vd-free nuclear
stock plants to replace those presently being used and in the UK such plants of
most commercial varieties have already been identified. It will, however, take
much longer before infected stocks in commercial gardens can be replaced by
healthy ones.
CHAPTER 9

Varieties and breeding

9.1 HISTORICAL INTRODUCTION

The origins of hop growing and the sources of the original plants must be a
matter for conjecture but presumably hop cones were first harvested from wild
plants and some of the same plants would have been collected when they were
first taken into cultivation.
The original gardens would have contained a mixture of several different
genotypes from which, with experience, the better plants would be selected.
The first selection would have been on the basis of how successfully the plants
grew and yielded, the best being used as a source of cuttings to either plant new
areas or to replace the inferior plants in an existing garden.
This process of selection would lead to the plants within a hop garden
becoming more uniform, and once this happened, brewers would be able to
notice the relative merits of hops from different sources. This could then lead
to the second stage of selection based upon the suitability of different types for
brewing. A grower whose hops were in demand would be able to sell cuttings
of his plants to growers who had more difficulty in selling the cones of their less
popular sorts.
There is an interesting comment on varieties in England by Mills (1763) who
wrote:
The several kinds and goodness of hops may likewise be known by the colour of the
vines, binds, or stalks. The whitish binds produce the white hop, both the long and the
oval: the grey or greenish binds commonly yield the large square hop: and the red binds
bear the brown hop, which is the least esteemed.
The planter of hops ought to be extremely careful in the choice of his plants, or sets,
particularly in regard to the kind of the hop: for it is a great trouble and loss to him
when his garden proves to be a mixture of several sorts of hops, ripening at different
times. He who plants the three sorts of hops before mentioned, viz. the early, the long
white, and the square hop, in three distinct parts of his ground, will have the
conveniency of picking them successively as they become ripe.

In the days when transport was difficult over any long distance, hops would
have been mostly used close to the area where they were grown and so each
locality would tend to make its own selections, though these would become
196 Varieties and breeding
more widely distributed as transport improved. In England, Percival (1901)
described 20 different varieties, although these did not include some of the
attractively named sorts, such as Golden Tips or Farnham Bell, referred to by
earlier writers. In Germany, Wagner (1905) also listed 20 different varieties, 8
of which were early maturing, 5 mid-early and 7 late, while Braungart (1901)
referred to ' .. .the 60 European hop varieties at Weihenstephan'.
From these many different types, one or two became dominant in each area,
a trend that was probably encouraged by the development of the railways and
the consequent extension of national and international trade since it would be
far easier for merchants to deal in a few standard types. This standardization
resulted in the practice of continental hops being sold by the district of origin
rather than by variety and Thompson (1972) states that the majority of the
'district' hops in Germany, and the hops grown in most east European
countries, were of the Saazer or Hallertauer type. In Yugoslavia, however, the
variety grown in Slovenia, although known there as the Savinja Golding, is the
same as the English Fuggle. It is recorded that hops were introduced into
Yugoslavia from England and it seems likely that the Fuggle variety was
supplied under the misnomer of 'Fuggles Golding' - a practice at one time
resorted to since Goldings were of superior quality to Fuggles.
These English varieties illustrate the common practice of naming varieties
after the growers (Mr Fuggle and Mr Golding) who developed them.
Thompson referred to the same practice on the Continent where the Semsch
hop, selected by a grower of that name from the Altsaazer hop, was long
regarded as the best type of Saaz hop although marketed without references to
the grower's name.
The origins of most of the old varieties are obscure but several of them are
almost certainly clonal selections that differ from one another in only one or
two characters. The English Golding hops are a good example of this since
there are many different types which are scarcely distinguishable, except for
differences in the time of ripening, and it is only the most recently selected
clones whose history is recorded. One distinct variety with a recorded history is
the Fuggle which is reputed to have been selected as a seedling in 1861, the
plant finally being introduced into commerce about 1875 (Parker, 1934).
There is little evidence that the old European cultivars are an improvement on
the hops that were growing in the wild; they would appear to be merely selections
of the best of the existing material that could be found. In the USA, however, the
situation is different. The traditional hops there, the Clusters, do appear to be
hybrids between European cultivated hops and the wild hops of America.
Brooks and Horner (1961) suggest that the Late Cluster originated as a seedling
of English Cluster during the early settlement of the eastern seaboard, where
hops were introduced by the Massachusetts Company in 1629, and was then
taken to the west coast by early settlers. The Early Cluster probably originated
from the Late Cluster in Oregon about 1908 as a bud-sport.
Historical introduction 197
The earliest efforts at improving hops by hybridizing and selecting the
resulting seedlings appear to have been commenced in Germany by Stambach
in 1894 and Remy in 1898 followed in the USA in 1904 by Fairchild (Smith,
1937). These early efforts were soon abandoned and of far greater significance
was the initiation of a breeding programme at Wye College in England, also in
1904, by Howard (1905) who reported that 'During the past year, various
English and German female plants have been crossed with widely different
males, and twenty-three sets of seed have been obtained for growth next
spring'. The collection of the plants used as parents in this programme had
been made during the preceding 10 years, the College having been opened in
1894.
When Howard left Wye his work on hop breeding was taken up by Salmon
who joined the College in 1906. At that time Salmon was aged 35 and an expert
plant pathologist, having published an outstanding work on the powdery
mildews, A Monograph of the Erysiphaceae. Burgess and Glasscock (1960), in
their obituary to Salmon, point out that hop powdery mildew received special
mention in this work and suggest that it may have been his interest in the
disease that led him to make hops the main object of future research. Although
Salmon officially retired in 1937 this made no difference to his enthusiasm for
hop breeding which continued to occupy him until very shortly before his
death in 1959. His successors at Wye have continued to build on his pioneering
efforts.
Salmon's work will be referred to frequently in the sections that follow and
there can be little doubt that he has had more influence on the nature of the
crop than anyone else who has worked on it.
Fairchild's initial work in the USA was followed by a new programme,
commenced by Stockberger in 1908, which was pursued energetically until
1916 when it was abandoned as a consequence of the 1914-18 war and the
introduction of prohibition. Hop research was resumed in the USA in 1931 in
Oregon where most of the American hops were then grown (Haunold, 1981).
Another breeding programme that started very early was one at the
Carlsberg Breweries in Denmark, initiated by Schmidt (1915a) and later
conducted by Winge (Winge, 1963). Their approach was very similar to that of
Salmon and they might well have had a similar influence on the crop if hop
production in their country had not been of so little importance that it was
abandoned after the 1914-18 war. A breeding programme was also carried on
at Svalof in Sweden but this too was abandoned because the crop was not
grown commercially there. One of the Svalof varieties has, however, been used
in the Wye programme because of its exceptional earliness which is the result
of selection under very long-day conditions.
Research on hops in Czechoslovakia was commenced at agricultural schools
in 1894, the same year as at Wye College in England, and a Hop Research
Institute was established in 1925 (Fric, 1985). The approach to varietal
198 Varieties and breeding

improvement there has been to concentrate, until quite recently, on clonal

selection, rather than hybridization, in order to retain the particular quality
characteristics of the Saazer hop.
The spread of downy mildew throughout Europe was one of the principal
reasons for the founding in Germany, in 1926, of the Hans-Pfiilf-Institut for
Hop Research at Hull where a programme was immediately commenced to
develop varieties resistant to that disease (Zattler, 1951). Downy mildew also
provided the stimulus for the recommencement of breeding work in the USA,
in 1930, by the Department of Agriculture in co-operation with Oregon
Agricultural Experiment Station (Smith, 1937).
More recently, breeding work has been carried out in both the Slovenian
and Backa regions of Yugoslavia, in the DDR, Poland, Russia, South Africa,
Australia, New Zealand and Mexico.
The traditional cultivars were originally judged by brewers mainly on their
aroma and appearance, with relatively little importance being attached to their
bittering power, or 'preservative value', although the superiority of American
hops in this respect was recognized by some brewers who imported consider-
able quantities. Salmon (l917c) and Schmidt (l915b) both emphasized that
aroma depended upon the genetic nature of the variety and not the country in
which it was grown. Both of them paid particular attention to the resin content
of the seedlings that they raised but it was Salmon's work which bore fruit and
led, eventually, to a whole class of 'bitter' cultivars as distinct from the old
'aroma' hops.

9.2 AROMA HOPS

One of the major concerns of brewers is that the character of their beers should
remain constant and, for that reason, they are generally reluctant to change the
hops that they use unless there is some special reason for doing so. It is,
therefore, almost a contradiction in terms to suggest improving the aroma of a
variety since that involves change. The improvement of such hops has
consequently been aimed at introducing superior agronomic characters while
retaining an aroma as close as possible to the original.
Aroma is not only a very complex genetic character but its organoleptic
assessment is sUbjective and dependent upon the preferences of individual
buyers. Hop plants are, genetically, extremely heterozygous so that there is
great variability within any seedling family. The chances of a hybrid having the
same aroma characteristics as its female parent are extremely low and
selections raised from seed have, in general, only been accepted as replace-
ments for traditional aroma types when the production of the latter has been
threatened in some way.
The outstanding example of this is the devastation of Fuggles in England by
verticillium wilt. This disease can only be controlled by growing resistant
Aroma hops 199
cultivars and the first ones to be discovered were very different in aroma from
Fuggles but had to be grown for want of anything better. They were
Keyworth's Midseason and Keyworth's Early and were second and third
generation, open-pollinated seedlings, respectively, of a wild American female
hop var. neo-mexicanus (Salmon, 1949).
Bramling-Cross, which followed (Salmon, 1951), came from a cross
between Bramling (one of the true Golding hops) and a first generation male
from a wild Canadian hop from Manitoba. It was quite well-accepted by
brewers and was widely grown for a time but its popularity subsequently
declined. These varieties had been raised before wilt became a problem and
were discovered purely as a result of Keyworth screening a wide range of
Salmon's breeding material. The most successful of all the wilt-resistant
aroma hops in England has been WGV, a hybrid raised by a private grower in
1911 and discovered on his farm after it had been bought by the Whitbread
brewery. The records detailing the parentage of this cultivar have, unfortu-
nately, been lost.
Once such sources of resistance had been discovered, a breeding programme
was initiated by Keyworth in 1944 aimed specifically at developing a wilt-
resistant replacement for the Fuggle. This resulted in the release, in 1958, of
three selections named Density and Defender (seedlings of Keyworth's
Midseason) and Janus (from WGV) (Wilson, 1959). Of these only Janus was
planted to any extent and that quickly declined. Another programme,
commenced jointly by Wye and East Malling in 1950, led to the selection of
Progress and Alliance, both of which were seedlings of WGV but by different
wilt-resistant males (Farrar et al., 1966; Wilson et al., 1967). They were judged
to be acceptable replacements for Fuggle in extensive brewing trials but they
only achieved limited success when grown commercially. This was partly
because they were released at the time when brewers suddenly started to
demand many more bitter hops and fewer of the aroma types that they had
previously required.
Although work in England has concentrated on hybridization, there has also
been some clonal selection of established cultivars. From amongst a range of
Fuggle material grown at East MaIling Research Station, Beard selected three
clones that were planted in a trial in 1931 and, of these, Fuggle N was selected
as a superior clone (Beard, 1943b). A further selection of 78 Fuggle clones was
commenced in 1943 but only two of these were found to match the
performance of Fuggle N (Thompson and Farrar, 1965). Although not in itself
superior to Fuggle N, one of these, Fuggle 37, was selected for distribution
when virus-tested stocks became available so that the different designation
would serve to distinguish between the old virus-infected and the new virus-
tested material.
In 1943 a trial was planted to compare seven different Golding clones
between which Beard (1952) found big differences in the incidence of downy
200 Varieties and breeding
mildew. In the meantime, a much larger trial had been planted at Wye in 1949
to compare 118 Golding clones, selected from a number of growers' farms.
These were reduced to 23 which were replanted in a replicated trial in 1957 and,
from these, four clones with a range of maturation dates were finally selected
on the basis of cropping, market valuation, resin content and susceptibility to
downy mildew (Beard and Thompson, 1961).
The breeding programme at Hull was aimed at developing downy mildew
resistant cultivars that were to be of comparable quality to the traditional
varieties. It required many years of patient work, finding sources of limited
resistance amongst cultivars and wild hops and crossing and back-crossing
these to build up the resistance to an effective level (Zattler, 1951). It was
almost 40 years before the first varieties, Huller Anfang and Huller Start
(1962) and Huller Fortschritt (1966), were released for commercial production
but these had relatively low yields which, combined with the spread of wilt, to
which they were susceptible, meant that they did not succeed (Maier and
N arziss, 1979). Some non-commercial selections that were highly resistant to
downy mildew were made available to English research workers and have
played an important part in the breeding programme at Wye.
Just as Fuggle was the cultivar to be most seriously affected by wilt in
England, so it was Hallertauer Mittelfruh, the principal German cultivar, that
was devastated there. The breeding programme was re-orientated in the early
1960s to deal with this problem and the first selection with resistance to both
downy mildew and verticillium wilt was Huller Bitterer (now known simply as
Huller) which is a spontaneous triploid. By 1974 it occupied one-fifth of the
aroma hop area in the Hallertau district, although an alternative traditional
cultivar, Hersbrucker Spat, that is reasonably resistant to wilt, was even more
widely planted. Both these cultivars suffered from the disadvantage of
unsatisfactory yield, and also susceptibility to viruses, so the release of a
superior new variety, Perle, in 1978 proved very valuable. Not only is it
reported to have aroma characteristics as good as those of Hallertauer
Mittelfruh, but it also yields well and has a higher a-acid content (Maier and
Narziss, 1979).
The revival of hop research in the USA in 1931 was stimulated by the spread
of downy mildew although the disease was, for many years, controlled in
Oregon by growing the English cultivars Fuggle, Bullion and Brewer's Gold
which were more resistant than the American Clusters. The cultivation of
Clusters moved to the drier states of Washington and Idaho where the disease
was much less of a problem. That left Fuggle as the only aroma hop in Oregon
where it was found to be uneconomic to grow it seedless. The first cultivar to
be released as an aroma hop from the Oregon breeding programme was
Cascade (Brooks et aI., 1972) which had much of Fuggle in its parentage and
was quite widely grown for some years but has since declined (Romanko,
1986).
Aroma hops 201
A colchicine-induced tetraploid form of Fuggle was later introduced into the
USA breeding programme and two triploid seedlings, Willamette and Col-
umbia, were released as replacements for Fuggle itself (Haunold et al., 1976a,
b, 1977). The nearly sterile nature of the triploids meant that good yields of
seedless cones were readily achieved. Willamette is very widely grown in
Oregon but Columbia is of little importance.
There is a considerable demand from some American brewers for a supply of
European hops, especially Hallertauer MittelfIiih, and as supplies of it became
limited, because of losses from wilt in Germany, there was increased interest in
attempts to grow it in the USA. The European varieties are, however, not well
adapted to growing conditions there and, as an alternative, triploid seedlings
have been bred from a tetraploid form of Hallertauer (Haunold and Nick-
erson, 1987) and one of them released under the name Mt Hood.
In Czechoslovakia, the Saazer hops are renowned for their good aroma but
they give low yields and research has been concentrated on improving the type
by clonal selection. It is feared that attempts to increase yield by hybridization
would result in a loss of the aroma that is characteristic of the traditional sort.
Improvement has been achieved by clonal selection from within the old land
cultivars. Such selections have been made on populations from different
regions and the relationships between them are not clear. Three Osvald clones
(31, 72 and 114) were selected from the original Zatec (= Saazer) variety and
released in 1952. Zlatan came fr~m the same source and was released in 1976.
Other varieties have been developed by mass selection within land cultivars.
Aromat was selected from hops growing at the village of Lhota while the
variety Sirem came from a village ofthat name (Fric, 1985). Vent and Beranek
(1970a, b) state that Aromat is outstanding for aroma and spiciness while both
cultivars out yield the Osvald clones with which they were compared. All these
selections are marketed as Zatec (Saazer) hops and it is claimed that they are
extremely uniform although there do seem to be noticeable differences in form
between some of the clones.
Such clonal selection is still an important part of the Czech programme.
Between 1971-80 over 700 selections from commercial cultivars have been
evaluated, of which 125 were further tested in 1982-7. Of the 8 clones finally
selected, 4 have been recommended for state trials (Rigr and Beranek, 1988).
In Russia also, the emphasis appears to have been on clonal selection from
the standard variety Serebrianka. Haunold (1981) listed a number of clonal
selections and suggested that the emphasis on 'medium early' and 'red' in the
Russian varietal names indicated a relationship to the Czechoslovakian Saazer
variety. Some hybridization has also been carried out, however, since the
selections 38/19 and Zitomir 8 have been described as hybrids that are earlier
and more productive than clone 18.
The standard hop variety in Japan, Shinshuwase, is reputed to be a hybrid
between Saaz and White Vine but Ono (1959) queried this because, from the
202 Varieties and breeding
morphological point of view, he thought it resembled the American hop more
than the European one. Kirin II is a clonal selection from Shinshuwase, while
Mori et al. (1969) reported the selection of a mutant form, named Golden Star,
that was higher yielding, was easier to pick and more resistant to downy
mildew.
There is, in reality, no very clear distinction between aroma hops and bitter
hops since there appears to be no evidence that good aroma characteristics
cannot be combined with high a-acid contents. The American Clusters occupy
an intermediate position since they were, for many years, the most widely
grown variety in the USA and accepted as the standard kettle hop against
which others would be judged. Their a-acid content of around 7% was higher
than that of the standard European hops, which was generally around 5%, and
their a-acids are very stable in storage. It was for these reasons that they were
bought in considerable quantities by English brewers.
The Early Clusters are thought to be a clonal selection from the Late Cluster
and, apart from the difference in maturation dates, they are very similar.
Because these were the most widely grown hops in the USA, and because it was
realized that the productivity of individual plants was very variable, a clonal
selection programme was initiated in 1957. Forty-one selections were tested for
virus infections by grafting to susceptible hop cultivars but only hop mosaic
was identified. Yields and a-acid contents were recorded in 1962-4 and four
clones selected to give a range of harvest dates. By 1972 these four clones
constituted 80% of the Washington hop acreage (Skotland, 1973).
Another hop that is classed with the Clusters in the USA, but should
perhaps be included with the bitter hops, is Talisman which was raised from
open pollination of Late Clusters by Romanko in Idaho (Romanko, 1964b)
and released in 1965. It has a higher a-acid content than Clusters (about 8-9%)
and gives high yields but it has now been largely superseded.

9.3 BITTER HOPS

In a paper given to a meeting of the Institute of Brewing in London, Salmon
(1917c) described the main objective of his breeding programme as combining
the high resin content of American hops with the European-type aroma. At
that time he had achieved some success by crossing American Clusters with
European males and European cultivars with American males but his real
break-through came almost immediately after he gave the paper with the
introduction from Canada of a wild hop from Manitoba. This plant, which he
designated BB 1, was planted in 1917, and open pollinated seed was collected
from it in 1918, after which it died. From the seedlings raised in 1919 he
selected the varieties Brewer's Gold (Salmon, 1934) and Bullion (Salmon,
1938). These set new standards by combining high resin contents and excellent
yields although their aromas had definite traces of the 'blackcurrant' character
Bitter hops 203
that is a feature of wild American hops. Their a-acid contents are greatly
influenced by environmental conditions but with values around 7% in Europe
and 9% in the USA they were above anything else available at that time.
Salmon later crossed a male seedling of Brewer's Gold with Canterbury
Golding and obtained Northern Brewer (Salmon, 1944) which had a much
better aroma than the other two selections but was not so high yielding. Its a-
acids were frequently higher than those of his other two cultivars and were
more consistent.
Bullion and, to a lesser extent, Brewer's Gold were widely planted in Oregon
in the 1950s and in the Yakima valley in the late 1970s, Bullion being preferred
because it was earlier maturing (Romanko, 1986). Apart from their high
productivity, they were popular with growers because they were, under those
conditions, resistant to crown infection by downy mildew.
In Germany, Brewer's Gold was preferred to Bullion but Northern Brewer
was of even greater importance because it was found to be resistant to the
strain of verticillium wilt that was destroying the hops in the Hallertau district.
The planting of high a-acid cultivars was also stimulated by the development
of hop processing, especially in Germany, and by 1978 they accounted for 47%
of the German hop area. Brewer's Gold and Northern Brewer have also been
widely planted in Belgium and they are the main cultivars in Spain where they
are called Hybrid 3 and Hybrid 7. They are also the principal varieties in
Bulgaria while Northern Brewer is grown extensively in the DDR where a
clonal selection from it, named Braustern, with higher yield and a-acid
content, has been described by Dolzmann and Wilding (1985).
It is ironic that these cultivars bred by Salmon should have been grown so
widely in other countries yet never achieved similar importance in England
where they had been developed. Bullion and Brewer's Gold were only
acceptable to a limited number of UK brewers because of their American-type
aroma and expansion of the Northern Brewer acreage was resisted by growers
because the hop was very susceptible to powdery mildew. It is a further irony
that Salmon became interested in hops through working on their diseases, and
gave the first reports of resistance to powdery mildew, yet his successful
cultivars should have been very susceptible to either downy or powdery mildew.
Before they were superseded by newer selections, Salmon's cultivars
accounted for about one-third of the world hop acreage but their importance
did not stop there since nearly all the new high a-acid cultivars have been bred
from his varieties. In most countries, the objectives of recent breeding
programmes have been to raise the a-acid contents still higher while, at the
same time, improving the aroma so that it came closer to that of the traditional
types. These common objectives have been combined with more local require-
ments such as resistance to diseases or seedlessness.
The first of the new generation cultivars to be developed was Pride of
Ringwood, bred by Nash in Australia. He collected open-pollinated seed of
204 Varieties and breeding
Salmon's variety, Pride of Kent, at Wye and raised the seedlings in Australia.
Pride of Ringwood was a seedling of one of these plants that was also open
pollinated and not, as usually reported, crossed with a Tasmanian male plant
(Nash, personal communication). Released in 1958 it came to dominate the
Australian hop industry, reaching over 90% of the acreage.
In England the increased demand for high a-acid hops that developed in
1966 had been anticipated and a revised breeding programme commenced in
1961. The main objectives were to improve the aroma and resistance to the
mildew diseases and so eliminate the objections that brewers and growers had
to the existing bitter varieties. Northern Brewer was chosen as the female
parent because it had a better aroma than Bullion or Brewer's Gold and was
less susceptible to downy mildew. Additional resistance to downy mildew was
introduced by the use of male plants that had been bred from the female
genotypes generously made available from Hull.
Wye Northdown was selected from the first generation seedlings from this
cross. One of its sisters was crossed with a male which was resistant to downy
mildew but also had the 'blister' (RB) type of resistance to powdery mildew
(section 7.2.6) and this cross yielded Wye Challenger. Another sister of Wye
Northdown was crossed with a male seedling of Eastwell Golding that was
resistant to verticillium wilt and immune to powdery mildew (R2) and Wye
Target was selected from this family (Neve, 1972).
Wye Northdown, which was released to growers in 1971, had a-acid
contents higher than Northern Brewer although subsequent work showed that
this advantage was due to its freedom from prunus necrotic ringspot virus. It
had fairly good resistance to downy mildew and was not quite as susceptible to
powdery mildew as Northern Brewer. Wye Challenger, released in 1972, was
highly resistant to downy mildew and, initially, completely field resistant to
powdery mildew (RB) although this only persisted for a few years until a more
virulent strain of the fungus became widespread. Its a-acids were generally
lower than those of Wye Northdown but it was higher yielding. Both cultivars
were very popular with some brewers and they were widely planted, although
only in wilt-free areas since both of them were susceptible to that disease.
Because of pressing demands from growers in the wilt areas for a high a-acid
hop, Wye Target was released in 1972 on extremely limited evidence and, in
spite of some cultural disadvantages such as very poor climbing ability, it has
become the most widely grown cultivar in England.
The main advantages of Wye Target are a very high level of resistance to
Verticillium, immunity to powdery mildew (R2) and a high a-acid content of
about 11 %, but it is very susceptible to downy mildew. It is unusually infertile
for a diploid hop and, even when heavily pollinated, its seed content rarely
exceeds 7%. It is therefore quite easy to achieve seed contents below the
European limit of 2% for seedless hops without having to eliminate male plants
from the entire neighbourhood of the garden.
Bitter hops 205
Because Wye Target is late maturing, growers required an earlier variety to
spread the picking season and Yeoman was selected as a hop that would meet
this need (Neve and Darby, 1984b). Although it has shown satisfactory
resistance to verticillium wilt, has a-acid contents similar to those of Wye
Target and excellent storage stability, many growers have had difficulty in
growing it. It is susceptible to canker (Fusarium sambucinum) which appears
to be the cause of some heavy losses from hill death during the winter, and it is
also very susceptible to powdery mildew (Sphaerotheca humulz). When first
selected, Yeoman was resistant to powdery mildew because it carried the
'blister' gene but when this resistance was overcome by a pathogenic strain of
the fungus it became apparent that the hop lacked an effective alternative
resistance mechanism.
At about the same time as these cultivars were being released in England, a
series of high a-acid cultivars were put into production in Yugoslavia. In
Slovenia there were four 'A' varieties of which Atlas, Apolon and Ahil were
seedlings of Brewer's Gold while Aurora was a seedling of Northern Brewer.
Aurora and Apolon were the most resistant to downy mildew but Aurora was
the most susceptible to verticillium wilt. All four varieties are sold under the
general description of 'Super Styrians' but Aurora and Atlas are the most
widely grown. The trade name relates solely to the area in which they are
grown and does not imply any genetic relationship to 'Styrian Goldings'.
In the Backa region of Yugoslavia, three cultivars, Neoplanta, Vojvodina
and Dunav with high a-acid contents were selected from seedlings of Northern
Brewer and put into production (Mijavec and Spevak, 1973).
A second, 'B', series of new selections was released in Slovenia in 1980 with
the names Blisk, Bobek and Buket. Bobek and Buket are both seedlings of
Northern Brewer while Blisk is a triploid seedling of a tetraploid form of Atlas.
Buket has an a-acid content of about lO%, with Blisk about 8% and Bobek
about 6.5%. Although they have good characteristics they do not appear to
have been widely planted and this is probably because they are not competitive
as producers of a-acids and are not as yet in demand as aroma hops.
Northern Brewer has been the starting point for breeding programmes at
Hull, on account of its high a-acids and its resistance to the German strain of
Verticillium. Although the emphasis was originally on developing aroma
varieties, more recently bitter hops have become a breeding objective and the
first of these, Orion, is reported to have a-acid contents of about 10% (20%
higher than Northern Brewer), to be resistant to wilt and downy mildew and
high yielding (Gmelch, 1985).
In the USA there has been some very successful breeding of high alpha
varieties. The first, but least successful, was Comet (Zimmermann et al., 1975)
from a cross between a seedling of Salmon's variety Sunshine and a wild male
from Utah. It had a-acid contents of 8-11 % but also a pungent 'Wild
American' aroma and is no longer grown (Romanko, 1986).
206 Varieties and breeding
Much more successful were Galena and Eroica both of which were raised by
Romanko in Idaho from open pollination of Brewer's Gold. Galena
(Romanko et ai., 1979) is early maturing with a-acids from 12-13% and good
storage stability. It is unusually sensitive to day length and yields are reduced if
it is trained too early. Eroica (Romanko et ai., 1982) is later maturing with a-
acids from 11-12% and inferior storage stability. It is more resistant to downy
mildew than Galena but is not so popular with growers and is less widely
planted (Probasco, 1985).
These were followed in 1982 by Nugget, raised by Haunold in Oregon with
Brewer's Gold in the pedigree of both male and female parents so that 5/8 of its
genetic composition came from that variety (Haunold et ai., 1984). It is useful
because it matures after Galena and before Eroica and has the added
advantages of good yield, high a-acids (12-13.5%), good storage stability and
fairly good resistance to downy mildew.
Two further varieties, Olympic and Chinook, were released soon afterwards,
both bred by Zimmermann at Prosser in Washington. Olympic was released in
1983 with 3/4 of its parentage derived from Brewer's Gold and it is. similar in
yield, maturity and a-acid content to Nugget (Kenny and Zimmermann, 1984).
Probasco (1985) suggests that, because it is inferior to Nugget in storage
stability and is more susceptible to downy mildew and insect attack, it offers no
advantage over that variety. Chinook, which was released in 1985, only has 1/4
of its parentage from Brewer's Gold but another 1/4 from a wild hop from
Utah (Kenny and Zimmermann, 1986). Although, according to Romanko
(1986), it has a pronounced 'Wild American' aroma it appears to have been
found more acceptable to growers than Olympic.
It should be noted that the a-acid values quoted for the American cultivars
are from spectrophotometric analysis which generally gives higher readings
than the lead conductance method used in Europe (Chapter 2).
In Japan there is only a limited effort put into hop breeding but there
Salmon's variety Northern Brewer was used also, in a cross with one of his
male selections (OB79), to produce the variety Toyomidori (Mori, 1981).
The above account shows very clearly the extent to which the development
of high a-acid varieties has been based upon Salmon's original efforts. His
success must be attributed in the first place to the fact that he set himself the
very precise objective of increasing the resin content of hops. Any successful
breeding programme depends upon clearly defined objectives, since attempts
to achieve too many improvements in the same programme can cause excessive
delays. Salmon's lack of success with disease resistance has already been
mentioned but if he had added this as an objective he might have been far less
successful. He also showed great initiative in going back to wild plants as a
source of new genetic material, long before this became one of the standard
methods in plant breeding.
The very painstaking work at Hull, which led to the development of plants
Varietal acreages 207
that are so highly resistant to downy mildew, must be given similar acknow-
ledgement. Salmon was able to see the benefits of his work quite quickly; the
workers at Hull had to persevere for many years before they reaped their
reward and again it was the clearly defined objective of the programme which
was essential to ultimate success. The breeding programmes at Wye and Hull
have each benefited from the work of the other. Downy mildew resistant
genotypes from Hull made it possible to introduce such resistance into the Wye
programme very rapidly, whereas the breeding for verticillium wilt resistance
and high a-acids at Hull has been based on Salmon's cultivar Northern
Brewer.
A breeding programme that was virtually independent of any other was that
of Roborgh in New Zealand. Californian Clusters became the standard cultivar
there because they were better adapted to the daylength at that latitude than
European hops, but they suffered considerable loss every year from Phytoph-
thora root rot. By crossing the Californian hop with English males that were
resistant, Roborgh produced a number of resistant varieties, the outstanding
member of which was Smoothcone which also had a somewhat higher a-acid
content than Californian. His major achievement came when he bred three
triploid hops which have much higher yields and a-acid contents than the
diploids from which they were developed. The most successful are Green Bullet
and Roborgh Superalpha, both of which were seedlings of a tetraploid form of
Smoothcone. Less successful is Stickelbract, bred from tetraploid First Choice
which is itself less successful than Smoothcone (Frost, 1980).

9.4 VARIETAL ACREAGES

The increasing demand from brewers for high a-acid hops and the need for
growers to plant cultivars that are resistant to verticillium wilt have led to
major changes in the varieties being grown. Whereas in 1958 production was
almost entirely of traditional varieties, by 1981 new varieties occupied 56% of
the hop area in BRD, 38% in the USA, 81% in the UK over 37% in Yugoslavia,
99% in Australia, 70% in DDR, 55% in Hungary, 74% in France, 77% in
Belgium and 100% in New Zealand.
The changes from aroma to bitter types and wilt susceptible to resistant in
England and BRD are illustrated in Figures 9.1 and 9.2. Verticillium wilt has
not been a factor in the USA but there was a major switch to high a-acid hops
there too. In England from 1972-81 the total area under hops declined by 15%,
the proportion of high a-acid cultivars increased from 17-62% and the overall
a-acid content of the crop increased from about 5.5% to nearly 8%. Production
of a-acid increased from about 550-750 tonnes in spite of the reduced area.
High a-acid hops have been so widely planted that there is, at present, a
surplus of them and the current trend is for a move back to aroma hops,
especially in the USA and BRD.
 UK UK ~
1967 00
1957
Aroma : susc 5415

..... ·.. ···T. 155

!R... ... i . Bitter ; rest N · :...·.·.....
· . ·.·;·... .b*' 81
Bitter: rest
298
Bitter : susc 645
292 Bitter : susc
Aroma : rest

1601 Aroma : rest

UK UK
1977 1987
Aroma : rest 982 Aroma : rest 248

1117
Aroma ; su5C ~
'"1
B itter : susc o·
1123
.....
o·
C/>
~
::3
Q...
t;j'
'"1
(l)
(l)
Bi t ter: res t Q...
5'
OCI
Figure 9.1 Varietal changes in UK, 1957 87.
 BRD BRD 1.0
1972 ~
1968
Aroma : susc -<
9891 ~
0;'
e:.
-
~
'"1
(l)

~en
rest

1597
Aroma : rest
Bitter : res

BRD BRD
1980 1987
Aroma : susc Aroma: rest
4237 8496
2838
Aroma : susc

2460
tv
Bitter: susc o
1.0
Bitter : rest 2452
Figure 9.2 Varietal changes in BRD, 1968-87. Hectares of high bitter/aroma varieties, resistant or susceptible to verticillium.
210 Varieties and breeding
9.5 FUTURE DEVELOPMENTS

Since it normally takes a minimum of about 10 years from the time a hop
breeder makes a cross to having a new selection ready for release, it is essential
to anticipate what will be required in the future. Some of these needs are
already apparent.

9.5.1 Pesticide residues

Increasing concern about the effect of pesticide residues on health and the
environment is already having serious effects upon the international hop trade
because some countries have set limits on the residues that are accepted. These
are based upon the needs of their own hop producers and do not provide for
growers elsewhere having different problems that require different pesticides.
There is little doubt that restrictions on pesticide usage will become more
stringent and alternative methods of control will be required. The most
effective of these is to breed resistance to the pest or disease into the hop plant
itself and considerable progress has already been achieved with this. Complete
field immunity to powdery mildew has survived in Wye Target for some 15
years and other sources of resistance are available. Very high levels of
resistance to downy mildew have been incorporated into several cultivars and
can probably be developed still further.
Aphids and spider mites are less easily dealt with in this way but consider-
able effort is being devoted to finding sources of resistance to the attacks of
these two pests. Any success in significantly reducing pesticide usage will,
however, probably depend upon augmenting such resistance with biological
control by predators.

9.5.2 Dwarf hops

The first published reference to dwarf hops was by Neve (1979) who reported
finding a dwarf at Wye in 1977. However, it has recently been pointed out to
him that Salmon made the following note about one of his seedlings in 1911.
'V. vig growth, laterals of medium length, v. closely placed, hops densely
clustered, v. fruitful, of no direct promise'. This would appear to be a
description of a dwarf-type hop and suggests that Salmon may have envisaged
the possibility of making use of this character. The breeding and selection of
dwarfs is being actively pursued at Wye together with the appropriate cultural
and harvesting techniques. Gunn and Darby (1987) have described the benefits
that can be expected from this development as being cheaper supporting
systems, reduced stringing costs, easier pest and disease control and, most
significantly, much lower harvesting costs.
Inheritance of important characters 211
Much remains to be done before dwarfs can be shown to be a viable
commercial crop but, if successful, there is no reason why dwarf varieties
cannot be developed to meet all needs. Because the criteria for bitter hops are
less stringent than those for aroma hops it is likely that the first commercial
dwarf varieties would be of the bitter type.

9.6 INHERITANCE OF IMPORTANT CHARACTERS

Dark (1951) pointed out that there was no precise information on the
inheritance of any characters in the hop and the situation today, although
better, is still very unsatisfactory. It is important for the breeder to have some
idea about the heritability of the characters that he is trying to improve. Some
of the variation between individual plants is due to the genotype they have
inherited from their two parents, some is due to other causes such as the
environment in which they are growing. Selection of individuals for characters
that are highly heritable can be expected to result in worthwhile genetic
improvement, whereas the same amount of effort put into selection of
characters with low heritability will be far less rewarding. Several attempts
have been made to assess the heritability of some of the most important
characters, particularly resin contents and yield.

9.6.1 Resin contents

Schmidt (l915a) measured the lupulin content of the progeny of several

families and found that the progeny of high-Iupulin females had means lower
than the parent while low-Iupulin females gave progeny with a mean exceeding
that of the parent but in all cases some seedlings had lupulin contents
exceeding those of the parents. Schmidt considered it: ' ... hopeless to attempt,
on the basis of these experiments, any determination of what has, genetically
speaking, taken place in the course of these crossings '. Lewis (1980) was able to
calculate from Schmidt's data that the heritability in that population was
approximately 0.66.
Farrar et al. (1956) crossed a high and a Iowa-resin containing female,
which had similar fJ-resin contents, with two male parents and concluded from
the analyses of the progeny that a- and fJ-resin contents were inherited
independently while the total resin content was merely the sum of the two
quantities, implying that limiting reactions occurred after the synthetic path-
ways of the a- and fJ-resins diverged. Neve (1958b) crossed a tetraploid with six
different males and found significant differences between the mean a-resin
contents of the progenies. There was no segregation into separate classes
within the progenies, indicating that control was polygenic, and in most
families there was no correlation between a- and fJ-resin contents.
212 Varieties and breeding
Keller and Likens (1955) estimated the heritability for a number of characters,
including resin contents, in early and intermediate maturity groups of selected
clones. They calculated heritability values for total soft resin, a-acid and 13-
fraction of 0.87-0.96 for plants grown in replicated plots and estimated the
potential gain from selecting the top 5% of the population to be as high as 50%
for a-acids.
Brooks and Likens (1962) measured the a- and f3-acids in the flowers of male
hops and determined the heritability of resin content, gland number and gland
size. There was a significant correlation between a- and f3-acids but they
considered that the lack of a perfect correlation indicated limiting reactions
after their synthetic pathways diverged.
Tetraploid plants with XXXY sex chromosomes are monoecious, making it
possible to compare the resin contents of male flowers and female cones on the
same plant. Hartley and Neve (1965) found that the mean value for a-acids in the
cones was 13.0 times that for male flowers but that there was a significant
correlation between the two sets of values. The f3-resin content of cones was only
5.9 times that of male flowers so the a-acid:f3-resin ratio was higher in the cones.
Similar results were obtained when comparing male and female seedlings of two
normal, dioecious diploid plants from two families. In a further study on
dioecious families they found no correlation between the a-acid content of the
flowers of male parents and the cones of their female progeny (Hartley and
Neve, 1968) although the males were clearly exercising some genetic control.
They suggested that the breeding value of a male was indicated better by the a-
acid content of its female parent than by analysis of its own flowers.
Whereas the resin contents in the above investigations were determined as a
percentage ofthe whole cones or male flowers, Likens et al. (1978) additionally
measured the a- and f3-acid contents of isolated lupulin glands of 112 female
and 74 male siblings. They found that lupulin from the females had an average
(a + 13) content of 73.3% and from the males 72.5%. Although there was no
significant correlation between the a and f3-acid contents of the cones of these
plants, the a-acid and f3-acid contents of the lupulin were negatively correlated.
This was interpreted as indicating a biochemical and genetic relationship
between the two and that the a:f3 ratio was inherited as such. However,
because the a- and f3-acid values were recorded as a percentage of the weight of
lupulin and their combined value was regularly about 73% of the total, such a
negative correlation is mathematically inevitable and, in the opinion of the
present writer, adds nothing to the attempt to determine whether or not a-
acids and f3-acids compete for a common precursor.
Lewis (1980) distinguished between broad sense heritability, which gives no
indication of the genetic causes, and narrow sense heritability which partitions
them into additive genetic effects versus the rest. Narrow sense heritability has
been described as the fraction of the phenotypic differences between parents
that the breeder can reasonably expect to recover in the progeny. The quality
Inheritance of important characters 213
traits Lewis studied included the lead conductance value (LCV), spectrophoto-
metric determination of a-acid, ,8-acid, a+,8-acids and a:,8 ratio. All were
highly heritable in the broad sense but, in the narrow sense, LCV, a-acid and
a+,8-acids were more highly heritable than either ,8-acid or aj,8 ratio. The
results for storage stability of the resins were inconclusive but it appeared that,
although fairly highly heritable in the broad sense, the genetic variability was
partly non-additive and there were consistent indications of a substantial
maternal effect.

9.6.2 Yield
Keller and Likens (1955) found yield to be highly heritable with heritability
values from replicated plots ranging from 0.67-0.92 and the expected gains from
selecting the top 5% of the population to vary between 15 and 55%. Lewis (1980)
on the other hand found that although flowering date, ripening date, bine
internode length and mean cone weight were highly heritable, cone number and
yield were so much affected by the height of the shoots that were trained that this
masked the genetic variation. He was, however, recording individual seedling
plants whereas Keller and Likens worked with 5-hill plots of each clonallY
propagated variety and this would reduce the non-genetic variation.

9.6.3 Resistance to verticilIium wilt

The source of resistance to wilt is not clear. All the genotypes from within
Salmon's material that were found to be resistant to wilt had North American
hops at some point in their pedigree, suggesting that resistance might have
come from that source. Since most of his selections were of such hybrid origin,
however, this is not very strong evidence. Dark (1952) pointed out that one of
the American parents, a New Mexican hop, was itself susceptible and
suggested that the European and American stocks contained complementary
genes which, when brought together, developed resistance. Some support for
this theory was obtained by Neve (unpublished) who crossed an American
female and European male, both susceptible to wilt, and found some resistance
amongst the progeny.
On the other hand, neither the cultivar WGV nor a male (No. 15) amongst
the Wye breeding material are thought to have any American ancestry yet both
are wilt resistant. Moreover, other wild American introductions have been
found to be resistant (Neve, 1964b; Darby, personal communication) suggest-
ing that complementary gene action is not essential for resistance genes in
either American or European populations to express themselves.
No major genes for resistance have been identified and, as there appears to
be a continuous range of reactions from very susceptible to very resistant, it
would appear that control is polygenic.
214 Varieties and breeding
There is some evidence that resistance is linked with undesirable agronomic
characters. Neve (unpublished) found an association between resistance and
low yield in one family while Darby (personal communication) has found a
similar association with Iowa-acid levels.

9.6.4 Resistance to downy mildew

As with wilt resistance, there is no evidence for major gene action in host
reaction to downy mildew and it is assumed that control is polygenic. There
appear to be some differences between the disease in the USA, where the main
problem is rootstock infection, and in Europe where it is leaves and cones that
are most seriously affected. It is not clear whether this reflects differences in the
pathogen or the environmental conditions.

9.6.5 Resistance to powdery mildew

Salmon (1917b) first reported resistance to this disease in nine plants raised from
seeds of wild Italian hops as well as in an ornamental 'golden hop '. These plants
were immune when grown in the glasshouse and subjected to very heavy levels of
inoculum, but some infection developed on some of the Italian plants at the very
end of the season when they were grown in the field. In another report Salmon
(1919) described a 'semi-immune' response to infection which appears to be
identical with the 'blister' reaction described by Liyanage et al. (1973).
Salmon (1920) raised open-pollinated seedlings from the 'golden hop' and
one of the immune Italian plants. One group of seedlings from the golden hop'
segregated for leaf colour 170 yellow: 178 green while another group, when
subjected to heavy doses of inoculum, segregated 72 immune: 33 susceptible.
Amongst the suceptible seedlings there was wide variation in the level of
infection. There did not appear to be any linkage between leaf colour and
susceptibility. The progeny of the Italian female segregated 24 immune: 9
susceptible.
Since most of the male plants in the breeding garden would have been
susceptible to the disease, it is unlikely that the large excess of immune over
susceptible would be due to the male parents. It is more likely that both the
female parents were heterozygous for two dominant genes, either of which could
confer immunity. In this paper Salmon also described a third source of
immunity in a plant supplied to him from the USA as the cultivar 'Golden
Cluster' but which turned out to be a male, the true origin of which was never
established.
More recent investigations (Liyanage, 1973; Liyanage et al., 1973) have
established that there are at least three dominant major genes that confer
immunity to some strains of the pathogen as well as the 'blister' gene (section
7.4.6). There is also a wide range of reactions amongst susceptible plants
Polyploidy 215
indicating polygenic control. Recent work indicates that the immune reactions
of the major genes are also modified by this polygenic background.

9.6.6 Reaction to viruses

Only in the case of hop mosaic virus is there any evidence of genetic variability
in host reaction to infection although cultivars may vary in the frequency with
which they become infected with other viruses. Plants infected with hop mosaic
are either susceptible or symptomless carriers. Susceptibility appears to be due
to a single major gene for which the English Goldings are heterozygous. Good
1 : 1 or 3 : 1 genetic ratios have been obtained in several crosses but there have
also been some unexplained cases where such ratios have not been found.

9.7 POLYPLOIDY

Ono (1959) referred to an earlier Japanese paper of his (Ono, 1948) which
appears to be the first report of the induction of polyploidy as a breeding
technique in hops. He stated that some economically promising triploid
seedlings had been obtained but this work appears to have been terminated
before any new cultivars were developed.
Dark (1953) considered Ono's work to be of limited value because it
involved the treatment of seedlings with colchicine and he initiated a pro-
gramme based on the treatment of young vegetative buds on shoots of
standard cultivars. He suggested that triploids would have three valuable
properties: they would be stimulated by pollination yet remain largely seedless,
they would be more vigorous than their parents and, because they would
receive two chromosome sets from the tetraploid female parent but only one
set from the diploid male, they would closely resemble the original diploid
female variety.
Dark's work was continued at Wye for several years and tetraploid forms of
Fuggle, Goldings, Saazer and Hallertauer Mittelfruh were produced and used
as parents in the breeding programme. The low seed content of pollinated
triploids was confirmed but it was only rarely below the 2% level necessary to
qualify as seedless by European standards. The greater genetic contribution
from the tetraploid female parents proved to be a disadvantage when trying to
introduce resistance to verticillium wilt from the male parents (Neve and
Farrar, 1961). Although several thousand triploid seedlings have been raised
and tested at Wye, only two ever reached the stage of farm trials and neither of
these proved satisfactory.
Elsewhere the breeding of triploids has been much more successful. In New
Zealand, Roborgh bred three high yielding, high a-acid varieties named
Harley's Fulbright, Green Bullet and Stickelbract which now occupy most of
216 Varieties and breeding
the hop acreage. It was found that successful triploids were invariably later
maturing than their female diploid parent (Frost, 1980).
Haunold (1971) described the objectives of a polyploid breeding programme
in the USA and listed the frequencies of triploid and aneuploid seedlings in the
progeny of tetraploid parents. From the tetraploid Fuggle induced for this
programme, the triploid variety Willamette was raised (Haunold et al., 1976a)
and more recently tetraploid Hallertauer has yielded seedlings that have a
European-type aroma (Haunold and Nickerson, 1987).
In England many triploids have been too late maturing and this is one
reason why none have proved commercially successful. The main reason for
their lack of success, however, is that they represent a dead-end in the breeding
programme. Not only do they produce little seed but the few seedlings that can
be raised from them have very variable chromosome numbers and are rarely of
any practical value. In order to combine adequate levels of wilt resistance with
the other characteristics required in a commercial variety, several generations
of breeding have been required and triploids can only be of use in the last stage
of such a programme.
Neve and Farrar (1961) suggested the possibility of using tetraploid plants
with XXXV sex chromosomes, which are monoecious, as male parents after
selecting them on the basis of their female cones. This has not been pursued
because it would involve too much time, breeding suitable monoecious plants,
before they could be used as parents in the second stage of such a programme.
Haunold et al. (1979) have described triploid male plants that can be used as
pollinators to stimulate yield of fertile diploids while retaining the seedless
nature of the cones (section 3.8).

9.8 MUTATIONS
Spontaneous mutations modifying two different characters have been
recorded. One is a mutation from green to yellow leaf which was first observed
at Wye in Brewer's Gold but has occurred in other varieties since then. Also at
Wye, on three separate occasions when screening Y0l:lng seedlings for their
reaction to powdery mildew infection, a plant has been noted with a leaf, or
leaves, divided into susceptible and resistant sectors. On at least one of these
occasions the mutation was from susceptible to resistant.
Both these mutant forms were immediately recognizable and, by taking
cuttings from the mutant sector, plants were produced that were entirely ofthe
mutant type. There would, however, be no possibility of recovering a mutant
sector that was not immediately recognizable. Many mutations are recessive
and, if it is a character not present in the population, the mutant cells will be
heterozygous for it and the new form will not be expressed. Even dominant
mutations may not be immediately recognizable. Changes in the chemistry or
content of oils or resins, for example, could only be detected by analysis and
Wild hops and gene pools 217
there would be nothing to indicate from which part of the plant samples should
be collected to demonstrate the change. Since most of the characters required
by the hop breeder are available in existing genotypes, combining these with
other commercial qualities is more likely to be achieved by hybridization than
by mutagenesis.
The artificial induction of mutations is not, therefore, a very promising
technique but various workers have attempted it. A report from Cze-
choslovakia (Beranek and Srp, 1976) gives details of radiation dose rates and
claims that seed obtained from irradiated plants yielded seedlings with
improved properties. Since these were of necessity hybrid plants, such
improvement could have been due to normal genetic segregation and not the
result of any mutation.
More reliable methods of producing and selecting mutations would be of
great interest to hop breeders because, unlike hybrids, they offer a means of
introducing a new character, such as disease resistance or different maturation
periods, into the original cultivar without affecting other important features
such as its aroma. There is, therefore, much interest in the use of in vitro
culture techniques aimed at developing somaclonal mutant forms. Mutagenic
agents applied to whole plants or seeds are only likely to give rise to mutant
sectors in the treated plant whereas somaclonal variants regenerated from cell
culture or from callus are much more likely to be non-sectorial.
Tissue culture has been used by Heale and co-workers at King's College,
London, in attempts to select novel sources of resistance to Verticillium albo-
atrum. They screened regenerating green callus of Wye Challenger in liquid
medium against crude culture filtrate of the pathogen for a 4 week period, with
fresh culture filtrate added at weekly intervals. Some green, viable areas of
callus survived from which shoots were excized. Regenerated plants were
screened for resistance against live inoculum in the glasshouse and four clones
showed reduced symptom expression. When subsequently tested under field
conditions at East MaIling, however, they were found to be susceptible (Heale
et ai., 1988).
Tissue culture would be most usefully applied in cases, like this one, where
screening for the desired character can be carried out in culture, but such
situations would seem to be rare. The demand for improved cultivars that
retain the brewing characteristics of established types is such that a start has
been made at Wye to apply the technique on a large scale as an alternative to
hybridization as a means of creating variability in the population.

9.9 WILD HOPS AND GENE POOLS

Wild hops have already proved a source of valuable characters for the plant
breeder. High a-acid and resistance to Verticillium albo-atrum have both been
introduced from wild American plants and resistance to downy mildew
218 Varieties and breeding

has been derived mainly from wild sources. The current search for plants
resistant to aphid attack may also benefit from some of the wild hops
maintained in breeders' collections.
Wagner (1975) made an extensive collection of wild hops from the different
regions of Yugoslavia and some of these were found to have good agronomic
characters. The main value of such material, however, would be when breeders
required a source of some new character that was lacking in the currently
grown cultivars. This could be resistance to some new pest or disease or it
might be that brewing chemists wanted hops with different oils or resin
characteristics. The wider the range of material available for screening, the
greater the chances of finding a hop with the required character.
The importance of maintaining as broad-based a collection of hop genotypes
has been recognized, and Wagner (1978, 1980, 1984, 1986), on behalf of the
Scientific Commission of the International Hop Growers' Convention, has
assembled lists of the various collections maintained by each member country
and it is considered most important that such collections should not only be
maintained but, whenever possible, extended.

9.10 CHARACTERISTICS OF PRINCIPAL CULTIVARS

The characteristics of any cultivar vary considerably from season to season and
from location to location. Table 9.1 is an attempt to detail the more important
features of each of the varieties listed, but it should be emphasized that the
information does not come from hops grown under uniform conditions or
assessed by identical methods.
 TABLE 9.1 Principal features of hop cultivars

~ Susceptability or resistance
o 0p
o c::
?
o:s . -
~ to diseases
- ... "0 :.a o
~ ] ~ 'S ·8 ~
"0 o:s 0 o:s:9
~ !':: o:s ... "0 Co) . -
.~ o ~.... - - 4J >.. ~o
;;>'4-< u0';:: o~ S S'""' S
oD 0 :91§2. I
CQ.
~ ~~gl§2. o o:s ~S ;;>, ::l'""' ::l ;;>, .2
~ ___ ""3S ..c:: ... o ;;>, 4-< t>O
.soo s... :.::::"'0
:-;::::: Q ::::
..9..d oS +-I o:s o ;;>, ... 0 S a
I o !'::
.c~
....!':: Co) o:s
::l 0.. !':: !'::
do
"0 ::l -;::"0 ~~ !':: !':: 0"0
d· 5 0 o ·8 S "0 o:s ~
.C ..c::
'" .... Co) S~"'~ o ~
.... :g s... :g <'
0
_p..!':: ·~.9 ~ ::l ... 0 !':: o:s
o~'u ooD
- ._
S 00.. o "0 ..c:: 0 ... 0
'€~ .... t[/)
0::l .~
... I td 0 ... I .... o:s ... l:J 0 o~ 0;:::J
>-'" Co) 0 o:s Co) [/) .... ~ p..,::?,
<:l ... U ~ ~ 8 p.., <:l en ·c ~ ~ ~8
-<5 ::z: >~ >---- >-'-'

ENGLAND
Goldings 4.5-6.0 2.3 22 1.0 3.5 good mid-late good good susc susc susc
Fuggle 4.5-5.5 1.8 26 1.4 3.3 2.0 0.63 good mid- good mod mod susc susc
early
Northern
Brewer 6.5-10.0 2.0 23 2.0 2.8 3.2 0.35 good mid mod mod susc v.susc susc rest
Bullion 6.0-9.0 1.9 36 3.2 1.5 poor late poor v.good susc susc susc
Brewers
Gold 5.5-8.5 1.9 38 1.5 2.3 2.7 0.48 poor late poor v.good susc susc susc mod -
Wye
Northdown 7.5-10.0 1.6 30 2.7 2.7 2.6 0.30 good mid good mod mod susc susc
Wye Challenger 6.5-8.5 2.0 21 1.7 3.1 2.4 0.30 good late good good rest susc susc
Omega 9.0-11.5 3.0 27 1.3 3.7 good late good good rest susc susc
Progress 5.0-7.5 2.8 27 1.0 3.3 mod mid mod mod susc susc rest
WGV 5.5-7.0 2.8 27 1.1 3.5 good mid good good susc susc rest
Bramling Cross 5.0-7.0 2.2 27 1.0 2.2 mod early mod mod susc susc rest
Wye Target 9.5-13.0 2.2 35 1.4 2.4 poor late mod mod susc rest rest
Yeoman 10.0-12.0 2.0 25 1.1 3.5 v. early good poor mod v.susc rest
good
 TABLE 9.1 Principal features of hop cultivars

OJ)
o o
Susceptability or resistance
o ".;:::: c= C
C<:I .-
~ to diseases
I- "0 "0 v
~
v
~
"0 v E 'a 'a C<:I v C<:I::9
C C C<:I 1-"0 u .-
>-.'-
'u
C<:I o v v >-. ~o 0::= v E E E~ E
.0 0 .~ 252 :; i:: c;:::: u >--.0 u ";:: ::s~ >-.
::s ::s
o >-. 0 0 ' o C':l ~E -5 >-. ;.:"0
cO. ~ _..c..c..- ..c l- v >-. '- OJ) o l- -C
::
C<:I
>-.>-' E C C<:I o >-. I- v ::= C
- I-
¥c "0 ::s o ::s 0.. - C o .:: C C V"O C<:I
:'OS v ~:E 'u E :g ~
.~ c: .:: u E o C v i::"" C<:I . - _ C<:I
v C E "0 o ~ C<:I ~ "';:u eo "';:: l-
..c -
o..C ·~.2 ~ 0:.0 ._
E v
0.. o ..c 0 I- C VI-t::C/l
:'OS 25 .~ _::s C<:I
C I-
~ C0 I-
o ~, I- ~ ::; 0
I ~ 0 0..0.. vI".) v;:J
-~ 0
u (5 f- .;::
> u 0 <:l I- U ..... u 0.. u 0.. c:s C/l -CJ} « >- o..~ C/l ~ >~ >~ >~

GERMANY
Hallertauer 3.5-5.5 1.0 21 1.0 3.7 3.7 0.89 good early- good mod susc susc susc susc
mid
Tettnanger 3.5-5.5 1.0 25 0.8 3.6 good early v.good mod susc susc susc susc
Spalter 4.0-5.5 l.l 24 0.7 3.3 good early- v.good mod susc susc susc mod
mid
Hersbrucker 3.5-6.0 0.9 25 1.2 2.5 1.6 0.33 good late good good susc susc susc rest
Huller Bitterer 4.5-7.0 1.2 30 1.2 1.9 mod mid mod mod rest susc susc rest
Perle 6.0-8.5 1.6 29 1.0 3.3 v. mid good good rest susc susc rest
good
Orion 7.0-10.0 2.0 30 1.4 2.6 good late mod good rest susc rest

BELGIUM
Record 5.5-8.5 1.0 30 1.8 2.5 mod late mod mod susc susc susc susc

FRANCE
Strisselspalt 3.0-5.0 1.0 24 0.7 2.4 good mid good mod susc susc susc
CZECHOSLOVAKIA
Saazer 3.0-4.5 0.9 26 0.4 3.5 3.5 1.46 good early v.good poor susc susc susc susc

POLAND
Lublin 3.5-4.5 1.3 27 1.0 3.7 - good early good mod
Lubelska
(Pulawy) 4.5-6.0 1.4 - good early- good mod
mid
YUGOSLAVIA
Savinja Gold-
ing 4.5-6.0 2.0 28 0.8 3.1 good early- good mod mod susc susc susc
mid
2.2 27 1.0 2.4 mod late good good susc susc S1.!sc -
Ahil} 8.0-10.0
Apolon 'Super 8.0-10.0 2.2 27 1.0 2.5 mod mid-late good good susc susc susc -
Atlas Styrians' 8.0-10.0 2.2 31 0.8 2.3 poor late good good susc susc susc -
Aurora 8.5-10.5 2.1 25 1.0 2.9 - good early- good good susc susc susc -
mid
Backa 3.0-6.0 0.7 20 0.6 3.2 mod mid-late good good susc susc susc -
Dunav 7.0-10.0 1.3 30 0.7 3.4 - good early- mod good susc susc susc -
mid
Neoplanta 6.0-7.5 1.5 36 1.5 2.4 mod early- mod good susc susc susc -
mid
Vojvodina 7.5-10.5 1.6 33 1.0 2.9 - good early- mod good susc susc susc -
mid

USSR
Zitomir 3.0-4.0 early- good poor susc susc susc -
mid
Serebrianka 2.4 mid-late good poor susc susc susc -
TABLE 9.1 Principal features of hop cultivars

01)
g o Susceptability or resistance
".=
~
s::
.- ---
.:: ~ to diseases
... "0 0)
~ --- :9 ~
0) ·a I: ~ 0) ~:9
~ g "8 ~ I: ... "0 u .-
~ 0 ~--C1J>" g;.,o 0:::: 8 8~ 8
>''- u .- ] 8 =' ~ >,
~ 0 :9~ ~ g ~ g ~ ... >, ='
~
___ :;
8 o 'iii '- 01)
~8 o ... ;.:::"0 .- I:
>,>' o..c..c ..... ..c ... 0) >, :::: I: ~
... ... o I: ~ o >, ... 0)
¥c I: I: 0)"0 u ~ ·u
~ 0) o ~
:9 ='
= gg.c~ ~~ 0) 8 "0
= 8~ :g
.~ E·5 ..c ... u 0 ..c u ~ >.. c:u ..... ~ ... ·a o ~ ~ ~
coo.-
0.1: ctt.- I o g o~ ._
8 0)
0. o OJ ..c 0 "E ~ "=... 0)"- t:r/J
~ g.!:!l , ... 0 :::: =';12§ ... , ~ ~ 0 O);:J
... .- 0.0.
r/J __ 0)"
> u ::; -~ 0u
~ ~ U o ::I:uo.. u 0.. <:! r/J t; f- ·c .< >= o..~ >~ >-- >--
USA
Clusters 4.5-5.5 1.4 39 0.6 1.8 4.0 0.67 v. early- mod good susc susc susc
good late
Talisman 7.5-10.0 1.8 52 0.7 1.2 mod late mod mod mod* susc
Eroica 11.0-13.0 1.4 40 1.0 0.1 poor late mod good mod susc
Cascade 4.5-7.0 1.0 37 1.2 2.7 1.6 0.31 poor mid mod mod rest* susc mod rest
Willamette 5.0-7.0 1.6 33 1.2 2.9 4.2 0.81 good early- good mod mod susc susc
mid
Nugget 12.0-14.0 3.3 27 2.0 2.2 good mid good good mod rest
Olympic 11.5-13.5 2.7 31 1.7 1.4 mod mid mod good susc susc
Chinook 12.0-14.0 3.9 32 2.0 2.2 good mid-late mod good mod susc
Mount Hood 5.0-8.0 1.1 23 l.l 2.5 mod mid good mod
Aquila 5.0-8.0 1.1 47 1.4 0.1 mod- mid mod good
poor
Banner 8.0-12.0 1.5 33 2.2 2.4 mod- mid mod good
poor
AUSTRALIA
Pride of
Ringwood 9.0-11.0 1.7 33 2.0 0.1 poor mid-late mod good susc susc susc

NEW
ZEALAND
Gre{.n Bullet 12.5-13.5 1.8 42 0.8 3.2 late good
Stickle bract l3.5-14.5 1. 7 38 1.0 1.8 mid good
Super Alpha 12.5-l3.5 1.5 38 1.5 3.2 eady- good
mid
Pacific Gem 14.0-16.0 1.8 41 1.5 3.2 eady- good -
mid

CHINA
641 5.5-8.0 mod mod

* Resistance to rootstock infection

This information should be treated as an approximate indication of varietal characters since it is drawn from a number of different
sources involving different environmental conditions and experimental techniques.
Resistance to powdery mildew may only be effective against some strains of the fungus.
CHAPTER 10

The hop trade

The hop growers' problems do not end with harvesting and drying the crop
since they still have to worry about selling it at a price that will give them a
reasonable return. Unlike many crops the consumption of hops does not
increase if the price is reduced since the brewers' requirements are determined
solely by the quantity of beer that they sell. If a cheap supply of hops is
available they may buy more than their immediate requirements in order to put
them into storage for future use but that only depresses the market the
following year.
On the other hand, it is essential for the brewer that he has sufficient hops to
produce all the beer that he can sell so that, if hops are in short supply, he is
willing to pay high prices to ensure that his needs are met.
The introduction of cold storage has prolonged the useful life of whole hops,
enabling brewers to hold reserves that reduce their dependency on any
particular year's production. This flexibility has been increased still further by
processing hops into powders or extracts which can be stored for long periods
with very little deterioration. Even so the market is still subject to marked
fluctuations as the supply situation changes and it has frequently been the case
that growers have achieved a better return in years when yields have been bad
because the high price has more than compensated for the small crop.
It is not clear how hops were marketed in the earliest days although there are
references to merchants being involved from an early period and any trade
with brewers living remote from the growing areas must have been handled by
middlemen. There are records to show that large quantities of hops were
imported into England, for example, before they were cultivated there, while
the discovery of what was apparently a cargo of hops in the Graveney boat
(section 2.1), carbon-dated to about 950 AD, indicates that trading must have
started at a very early date. Parker (1934) quoted an act of 1603 entitled 'An
Acte for avoyding of deceit in selling, buying or spending corrupt and
unwholesome Hoppes', the preamble of which commences: 'For so much as of
late great fraudes of deceits are generally practised and used by Forreiners
Merchants Strangers and others in forreine parts beyond the Seas in the false
packing of all forreine Hoppes brought into this Realme of England ... ' This
226 The hop trade
act clearly shows that merchants were very much involved, at least in foreign
trade.
For the English home trade, many hops were sold at fairs that were held in
various parts of the country. Stourbridge, in Cambridgeshire, is mentioned as
the principal outlet in the 17th and 18th centuries and Defoe (1724) said that
prodigious quantities of hops were sold there and that the prices obtained set
the standard for the rest of the country. Stourbridge was later overtaken as the
main fair by Weyhill in Hampshire while, although there were small fairs and
markets in Kent and Sussex, most of the crop from the south-eastern counties
was sold in London.
Parker (1934) recorded that there was a market in Little East Cheap on the
north side of the Thames from the year 1681 but at some stage it moved just
south of the river to the Borough of Southwark (Figure 10.1). There were rules
governing the sale of hops, one being that they should be brought to market
before being sold and in 1800 a Mr Waddington was fined £500 and
imprisoned for a month for forestalling the hop market (Clinch, 1919). The
Borough gradually became the main centre of the trade with most of the
merchants and factors established there although ' ... the hops from Hereford
and Worcester were principally sold at the County Towns' (Lance, 1838).
In an Act of 1710 a duty of 3 pence per pound was imposed on imported
hops and in 1734 a duty of 1 penny per pound was levied on hops grown in
England and this tax continued until 1862. The requirement that hops should
not be sold before being brought to market was presumably to ensure that they
did not escape being recorded for the purpose of this tax.
Hop production was subject to close control to ensure that the tax was
collected and some of the requirements (and penalties) were as follows. Hop
grounds and places for curing and keeping the hops had to be entered in the
books of the excise (40 shillings per acre); defrauding the revenue by using
twice or oftener the same bag, with the officer's mark upon it (£40); the
removal of hops before they had been bagged and weighed (£50); concealment
of hops (£20 and the concealed hops) and privately conveying away hops with
intent to defraud (5 shillings per pound) (Lance, 1838).
Although England has, at times, had a sizeable export trade in hops it has
never played an important part in the world's hop market which, according to
Gross (1900):
.. .is controlled by Germany, Austria and the United States, and is largely in the hands
of merchants who serve as go-betweens for the grower on the one side and the consumer
(brewer) on the other, the latter generally preferring - whether as a matter of
convenience, credit, or for other reasons - to deal with agents rather than direct with the
growers.
The largest hop market in the world is that of Niirnberg, where a large proportion of
the total crop finds its way every year, and where there is a large colony of hop
merchants, agents and dealers. Many of these have their own warehouses and
conditioning houses, and there are also warehouses for provisional storage in the town.
The hop trade 227

Figure 10.1 The hop market in the Borough (London). (From an engraving of 1729.)
228 The hop trade
Conditioning establishments are really necessary at Niirnberg, it being the custom
among a large section of Bavarian growers to leave to the merchant the task of getting
the hops ready for sale; consequently, it frequently happens that the former are careless
over the drying of their produce. Moreover, in many cases the hops are bought by the
merchants as soon as picked, and are sent into the town for further treatment.
The part played in Germany by Niirnberg is performed in Austria by the town of
Saaz, which is the centre of the hop trade in the latter country. Of course the volume of
business done is smaller than in Niirnberg, since the supply is a local one; nevertheless a
large trade is carried on, chiefly in the finer qualities.
Other large hop markets are held in London, New York and Warsaw; and there are
smaller business centres in all producing districts.

The town of Saaz referred to is now the town of Zatec in Czechoslovakia.

According to Giinzel (1904) there are, unfortunately, no records for the town
from the 10th and succeeding centuries because the town hall was the victim of
fires in 1767 and 1788. He stated, however, that there is no doubt that hop
growing was introduced to Saaz from Germany and especially from Bavaria
where hops had been grown at Freising since the 9th century. Similarly, hop
merchants moved into Saaz from Germany and there is documentary evidence
that they were established there by 1774.
In Niirnberg the hop trade can be traced back to 1400, at which time it was a
town monopoly and brewers purchased hops from the town administration.
Hops became economically important when hop cultivation was increased in
the beginning of the 18th century (Anon, 1969b).
The German merchants have continued to dominate international trade and
Raiser (1987) accounted for this as follows:

By tradition Germany has a high consumption of beer and an important brewing

industry which has been the backbone of German hops. As beer became gradually
popular worldwide, breweries were built in almost every country. This trend has not yet
come to an end. Germany through not having been able to take advantage of this
development to expand its brewing industry beyond its own borders like other
countries, such as the Netherlands, nevertheless was having considerable benefit from it
by becoming a leading exporter of brewing technology and know-how. A majority of
breweries in the world are brewing a German type of lager beer, a great many of them
are using German equipment and are employing German technicians or have their own
people trained in Germany. It is only natural that they are looking into Germany also
for their raw materials and particularly into hops, which are supposed to be of the
greatest importance as to the type and quality of beer. Thus German hops were able to
build up a world-wide image for quality, the industry had a sound base for expansion
and a strong international dealership could evolve. Efficient and large-scale processing
facilities were established to satisfy the needs both of the homemarket and abroad for a
reduced volume and the preservation of brewing value even under strenuous conditions.

A standard arrangement grew up in England whereby the growers' interests

were managed by factors and the brewers' by merchants, each of them
operating on a commission basis. In Germany, on the other hand, there was
only one middleman, the merchant who acted as go-between for the grower on
The hop trade 229
the one side and the brewer on the other. Brewers generally preferred to deal
with agents rather than direct with the growers, either as a matter of
convenience or because the merchant allowed them credit (Gross, 1900).
However, Gross also deplored the number of intermediaries between the
grower and brewer - the local dealers, the buyers and their agents, the
merchants, the consignment house for export trade and the commission agents
- and also the way that dealers resorted to talk of overproduction when buying
and of shortages when selling in order to increase their profits. The English
grower may, therefore, have been well-served by his factor who would have
been far better informed than most growers about the state of the market.
In Germany growers started forming themselves into associations to streng-
then their position, the first being formed at Forrenbach in the Hersbruck
district by a clergyman named Kelber. These associations supervised the
quality of the hops to be marketed and fixed the price and, in order to help
them compete with the dealers, the Bavarian Government assisted them with
subsidies and interest-free loans. Today, each hop growing region has its own
organization under the umbrella of the 'Verband deutscher Hopfenpflanzer
e.V.' (Kohlmann and Kastner, 1975).
Such associations can do little to overcome the fluctuations in price that
result from the unpredictability of hop yields from season to season. In
England there was close control of production during the 1914-18 war by the
Hop Control. When this ended, a duty of £4 per cwt was imposed to protect
the industry but in spite of this there were serious difficulties with marketing in
the subsequent years. From 1925-8 there was an effort to regulate sales by
means of a voluntary co-operative, English Hop Growers Ltd, whose members
represented nearly 90% of the hop acreage. Its efforts to control prices were
hampered by a large surplus left over from 1924 and large crops in 1925 and
1926. Its members agreed to reduce their acreage but non-members under-
mined this action by increasing theirs and the organization collapsed.
The introduction of the Agricultural Marketing Act in 1931 gave legal
support to co-operative action and the hop industry was the first to take
advantage of this legislation. A Hops Marketing Scheme came into operation
in 1932 under which all producers were obliged to sell through the Hops
Marketing Board while the Board was obliged to accept any English hops
consigned to them in fulfilment of a grower's contract. Growers were given a
basic quota, based on their previous production, which was varied each year in
accordance with brewers' requirements. The costs of production were determi-
ned by means of a survey each year and the price based on this. This meant that
prices in the UK bore no relationship to prices in the rest of the world and the
system could only operate because there were restrictions on the importation
of hops from other countries.
This scheme was successful in ensuring that brewers and growers were
protected from violent fluctuations in price and it was possible for them to
230 The hop trade
budget forward with some confidence. On the other hand it severely restricted
the ability of English merchants to deal in foreign markets because they could
never quote a price until some months after the crop was harvested, when all
the calculations of costings and weights had been finalized.
The entry of the UK into the EEC opened the market to all hops from within
the Community and it was no longer realistic to impose an embargo on any
imported supplies so the industry was forced into an internationally com-
petitive situation. Nevertheless, the Board continued to function for some
years with all growers remaining members, even after EEC regulations
required it to become a voluntary organization, and it changed its name to
English Hops Ltd. Many UK brewers continued to purchase hops from it at
prices fixed on a similar basis to the old arrangement but, eventually, conflicts
of interest led to some break-up of the organization and growers have now
formed themselves into several different producer groups although English
Hops Ltd is still the largest. The development of these co-operative grower
organizations reduced the role of the factors in the marketing of the hops
although they continued to serve the growers by representing their interests
when hop samples were being valued.
In the USA there has been a somewhat similar sequence of events. In 1937
their Agricultural Marketing Agreement Act was amended to include hops and
a Marketing Agreement and Order operated, effectively controlling prices,
from the 1938 crop. After 1942 there was no surplus to be controlled and the
Agreement and Order were terminated in 1945. The return of a surplus in 1948
however, led to their renewal in 1949 (Productivity Team Report, 1951) only to
be terminated again in 1952. Once again the market, and American hop
acreages, fluctuated widely and a further renewal of the Order was made in
1966 only to be again terminated in 1986. The difference between the UK and
USA situations was that changes in the USA were at the wish of growers
whereas in the UK the ending of the Board was the result of political changes
and was very unpopular with most of its members.
The marketing schemes in the UK and USA were effective because they had
legal backing, whereas voluntary co-operative schemes failed because of the
activities of non-members within the country. Similar problems faced these
marketing schemes on the international scale because of the activities of other
countries where growers were quick to take advantage of their freedom from
any restrictions. On each occasion that production in the USA was controlled,
there was a very satisfactory increase in the price paid to American growers but
this made hops from other sources more competitive and they captured some
of the American growers' markets. The Americans would doubtless have lost
even more sales were it not that many brewers are reluctant to switch their
sources of supply, partly because they do not want to change the type of hop
that they use but also because they like to feel that supporting their usual
sources helps to guarantee their future supplies. Nevertheless, without inter-
The hop trade 231
national agreements, it is doubtful whether efforts to control production in any
one country can be a permanent solution to the growers' problems.
In Europe, the EEC has adopted various devices to support hop production
in the member countries. Income support has been given to try and ensure that
growers' income does not fall below a reasonable level but the scheme has not
been very satisfactory. If growers in one country could show that a particular
variety warranted support, the same support had to be given to all growers of
that variety, even if it had given better returns elsewhere. Without any legal
means of controlling acreage, the Community has provided financial incentives
to growers for grubbing their hops in times of surplus but has not been able to
prevent them being replanted as soon as there were signs of the market
recovering.
In the communist hop growing countries the marketing arrangements are
unified under government control but whatever the internal arrangements, all
producers have to compete in a free market where international trade is
concerned. To succeed in this it is important that buyers are confident that the
hops will be of the type specified in the contract and of acceptable quality.
In 1884 the town of Saaz established a Hop-Marking Institute (Signirhalle)
which checked that the hops submitted by the producers were ' ...fine red
hops, grown in the district and in an unspoiled condition'. After weighing, the
bags were numbered and sealed. Similar arrangements for certifying and
sealing were commenced in Germany at about the same time (Gross, 1900) and
these continue to this day.
Within the EEC it is now required that all hops harvested there (except those
grown and used by a brewer or those put up in small packages for sale to
private individuals for their own use) are certified in accordance with Council
regulations. These require that, with each consignment, the grower must
provide his name and address, the year of harvest, the variety, the registered
number of the holding where they were produced and the number of packages
in the consignment. The hops must be supplied in closed bags or bales and be
marked as appropriate: 'prepared hops' or 'unprepared hops' and 'seeded' or
'seedless'. In England, the drying and packaging is always completed on the
farm so all hops sent from there are 'prepared hops'. On the continent most
hop preparation is finished at the merchant's warehouse so they are supplied as
'unprepared'.
The consignment must be sampled and the samples examined to ensure that
they meet the following requirements:

1. M oistuTe Content; the moisture content should not exceed 12% but if it lies
between 12-14% the hops may ,be certified and labelled as 'unprepared
hops'.
2. Leaf and Stem; (defined as leaf fragments and pieces of stem, leaf strigs and
cone strigs 2.5 cm or more long).
232 The hop trade
Maximum allowed (by weight) 6%
3. Hop Waste; (defined as small particles of the hop plant resulting from
machine harvesting, varying in colour between dark green and black, which
generally do not come from the cone)
Maximum allowed 3%
4. Seed Content; Seedless hops must not contain more than 2% by weight.
Hops with more than 2% must be certified and labelled as seeded.

Hops which do not meet these requirements may not be sold but they may be
returned to the grower for additional drying and cleaning and resubmitted for
certification.
In the USA, hops are also subject to inspection but there are not such rigid
limits to what may be sold. Instead there is a system whereby the contract price
for the hops is adjusted with premiums for low seed or leaf and stem contents.
Seedless hops are defined as those with less than 3% seed by weight, those
between 3-6% are 'semi seedless' and over 6% are classed as seeded. Only when
the leaf and stem content exceeds 15% are the hops unmerchantable.
The EEC regulations do not permit the blending of hops (other than for the
preparation of powders or extracts) unless they are from the same harvest,
from the same production area and are of the same variety; then the blending
must be carried out under the supervision of a certifying officer. For the
manufacture of powders and extracts different varieties from different produc-
tion areas within the Community may be blended provided the details are
recorded on the certificate.
These restrictions on blending are especially important for the aroma hops
since brewers buying these are generally paying a considerably higher price to
obtain a particular type of hop. Those buying bitter hops are usually much less
concerned about their source and many contracts are simply defined in terms
of a weight of a-acid at a price per kg, usually with a minimum a-acid content
stipulated. Powders and extracts generally have to be supplied at a specified a-
acid content and in order to achieve this blending is essential.
As a result of the increased demand for high a-acid hops in the 1960s the
cultivars then available were planted more widely, replacing aroma sorts, while
the newer, higher a-acid selections that became available were rapidly taken up
by growers in many countries. This change of varieties, combined with the
adoption of brewing techniques which utilize a-acids more efficiently, went too
far and there is at present a surplus of bitter hops, the market for which is very
depressed, while there is a more active demand for aroma types.
Not only is the basic price of the aroma hops higher but their lower a-acid
content means that two to three times the quantity is required to produce the
same level of bitterness in the beer. Although the most conservative brewers
will doubtless continue to insist on using them, the switch towards high a-acid
cultivars seems certain to continue, especially as breeding continues to improve
The hop trade 233
their aroma. Most bitter hops will be used in a processed form and, in order to
achieve a predetermined level of a-acids in the products, the manufacturers
will need to buy hops wherever they can obtain those that will blend to give the
required level.
The brewing trade has already gone through a period of mergers and take-
overs that have concentrated much of the industry in the hands of relatively
few companies and there is little doubt that the process will continue still
further. This is leading to a corresponding concentration of hop buying into
fewer hands which will have a dominant effect upon the market, laying down
the criteria for their large-scale purchases. Growers will have no choice but to
try and meet these requirements and the surviving small brewers may have
little influence and will have to accept the same standards.
The large number of hop varieties being grown at the end of the 19th century
was quickly reduced to five or six dominant types during the first half of this
century. Intensive breeding activity to meet the changing needs of brewers and
growers has led to renewed variability in the hops that are available and it is
certainly desirable that this should continue. Too much uniformity exposes the
industry to the very real threat of another new disease problem creating havoc
in the way that downy mildew in the 1920s and, to a lesser extent, verticillium
wilt in the 1960s have already done. It is to be hoped that the major hop buyers
will be willing to purchase a mixture of alternative varieties from different
countries so that as much diversity as possible can be maintained within the
crop as a safeguard against any future disruption of supply.
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258 References
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260 References
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growth responses in Fuggle hops (Humulus lupulus L.). Crop Sci., 4,310-13.
Zimmermann, C. E., Likens, S. T., Haunold, A., Horner, C. E. and Roberts, D. D.
(1975) Registration of Comet hop. Crop Sci., 15,98.
Index

Acreage of hop varieties, see Varietal Arsenic 90

acreages
Agriolimax reticulatus, see Slugs Bales 45, 100
Agriotes spp., see Wireworm Bare-bine, see Arabis mosaic virus
Air speed 91, 93-5, 98-9 Bedded setts 72-3
Alpha-acid, see Resins Bine-cutting 80, 86-7
Alternaria 172 effect on yield 80, 84
Armillaria root rot 172-3 Bine harvest, see Harvesting
American hop latent virus 1I5, 180-1 Bines 1, 56-8, 160, 168, 183
Analysis Biological control
essential oils 31-2 aphids 1I9, 121-6
resins 35-6, 75, 206 spider mites 130-1
Anthocorids 123-4, 130 verticillium wilt 167-8
Aphid, damson-hop 1I5-26, 176, 180-1 see also Predators
in America 105, 120-1 Bitter hops 35, 69, 199, 202-9, 232-3
biological control1I9, 121-6 Bitterness 2, 30, 35
chemical control1I9-21 Black mould 172
effect on yield 101-3 Black root rot Ill, 171, 207
first records 10 1, 105-6 Boron, see Trace elements
life history 115-19 Botrytis cinerea, see Grey mould
migration 15-20 Bracteoles 2, 22, 96, 138, 172
varietal preference 117-18, 177-8 Bracts 2, 22, 96, 138, 172
Aphidius 125 Breeding 1I0-ll, 124, 149, 159, 195-
A-plus, see Certification 218
Apple mosaic virus (ApMV) 182, 189-90 Buds 4-5, 17,72, 1I5, 141-2, 144, 154,
Arabis mosaic virus (AMV) 132, 175, 215
182-8 Burr 5, 137-8, 148
bare-bine 183-6
control 187-8 Cannabis 1,37-8
hop chlorotic disease 183, 185 Carbohydrates 20-3, 140, 170
SNA 182-3 Centre of origin 16
split-leaf blotch 183-4 Cephalosporium (Verticillium) lecanii
nettlehead 1I1-12, 132, 175, 178, 125
182-5, 188 Certification
Arion hortensis, see Slugs of dried hops 231-2
Aroma hops 33, 69, 198-202,207-9, of planting material1I2, 175, 187-8
232 Chemical analysis, see Analysis
262 Index

Chemical control spikes 137-9, 142-147

aphids 104 sporangia 137, 143-4, 147-9
downy mildew 109 zoospores 141, 143-4, 147
nematodes 188 Drying 89-100
powdery mildew 104-5 thermal efficiency of 92-5
spider mites 129-30 Drying times 91-4
verticillium wilt III Dwarfs 10,68,210-1
Chemical stripping 58, 145, 155
China 115 Earthing-up 54, 72
production methods 53 Earwigs 123-4, 135
Chromosomes, see Cytology Eelworms, see Nematodes
Chrysopids, see Lacewings ELISA technique 163, 175-6
Cladosporium, see Black mould Epidemiology
Clay-coloured weevil 133-4 downy mildew 146-9
Climate 76-8 powdery mildew 157
Climbing, 1,204 Essential oils 2, 31-3, 38-43, 125, 193,
Clonal selection 196, 199-202 219-23
CO2 extract, see Processed hops Exhaust air 92-3,95-6, 99
Coal for drying 90 recirculation 95-6
Coccinelids, see Ladybirds Extracts, see Processed hops
'Cock-hops' 71 Extrusion cooking 46-7
Conditioning of hops 96-9, 226-8
Continuous driers 92, 96 Factors 228-9
Cones 2, 21-2 Fallowing 132, 166, 188
Cooling 96-8 Farmyard manure, see Organic
'Crows nests' 87 manures
Cultivars, see Varieties Fertilizers, see Manuring
Cultivations 53-6 Flavour 2, 30, 38-43, 69
Cytology 11-16 Flea beetle 101-5, 135
aneuploids 15 Flowers
autosomes 15 female 2, 5
polyploids 12-15 male 2, 5
sex chromosomes 12-16,212 initiation 18-20
Fluctuating wilt, see Verticillium wilt
Dagger nematodes, see Nematodes Fluffy tip 62
Damson hop aphid, see Aphid Forficula auricularia, see Earwigs
Daylength 16-20,28 Fuel for drying 90-2
Defoliants 58, 145 Fumigation (against nematodes) 188
Depth of hop bed 91-5 Fusarium canker 170-1, 205
Dormancy
of plants 17, 19-21,29 Gambling on hop production 101
of seeds 18-19 Gene pool 158, 217-8
Downy mildew 57, 107-10, 137-49, 198, Genetics, see Inheritance
203,214,219-23 Germination 10, 18-19
chemical control 145-6 Gas-liquid chromatography 31
hygiene 144-5 Gibberellic acid 20, 70-1
infection periods 140, 147-9 Glands, see Lupulin glands
oospores 142 GLC, see Gas-liquid chromatography
overwintering 141-2 Grafting 37-8, 168, 185
resistant varieties 198,200,219-23 Graveney boat 26, 225
rootstock infection 140-2, 144 Grey mould 171-2
Index 263
Growers associations 229-31 Hydroecia micacea, see Rosy rustic
Growth retardants 71-2 moth
Growth substances 70-2 Hygiene
downy mildew 144-5
Hairs 1,9, 131 powdery mildew 155
Hand picking, see Harvesting verticillium wilt 165
Harvesting viruses 180, 187-8, 191
bine pulling 58
by hand 76, 79-82 Inheritance 122, 158,211-15
by machine 58, 76, 79, 83-9 Inoculum levels 137, 147-8
Haze 43 Integrated control, see Biological
Heat exchangers 91 control
Heat therapy 175 Intersexes 14-16,216
Height of wirework 49, 52-3, 75-6 Infection periods, see Downy mildew;
Hemp, see Cannabis Powdery mildew
Herbal uses 29-30 Irrigation 62-4
Herbicides 54-6 Iso-humulone 33-4
Heterodera spp., see Hop-root Iso-lupulone 33-4
eelworm
High a-acid hops, see Bitter hops Kaderavost 62, 112
High temperature drying 93-5 Kilns, see Drying
Hills 49-50, 54, 79 Krauselkrankeit 62, 112
History
production 25-29 Labour54,57-8,67,79-80, 87,91-2
breeding programmes 195-8 Labour costs 87
diseases 103-13 Lacewings 121, 124
pests 101-7 Ladybirds 105-6, 121, 124
use 29-31 Lateral pickers 67, 85
Honeydew 105, 115, 124 Layering 72-3
Hop chlorotic disease, see Arabis Leaf water potential 64
mosalc VlruS Leaves 1-3,9, 22, 137, 144-5, 148, 160,
Hop latent viroid (HLVd) 192-3 183
Hop latent virus (HLV) 180-1 Light, see Daylength
Hop mosaic virus (HMV) 112, 115, Line pattern 189
176-81,183,190 Loading of kilns 98-9
Hop-root eelworm, see Nematodes Low trellis 53, 66-8
Hop stunt viroid (HSVd) 179, 192 see also Dwarfs
Hover flies 121, 124 Lupulin glands 2, 9,31-2,212
Humidity Lupulones 33, 36
and diseases 143, 148, 157, 172
and drying 95-9 Machine picking, see Harvesting
and growth 64 Magnesium 62
Humulones 33-4, 219-23 Male plants 2-4,10-16,18,27,76,178,
Humulus 1-16 204-5,212,215
H. americanus 6 Manuring 59-62, 75
H. cordifolius 6 Marketing 225-33
H. japonicus 1, 9, 11, 70 Maturation 17-18,70-1,193,205,219-
H. lupulus 1-9, 70 23
H. neomexicanus 6, 199 Mechanical harvesting - effect on yield
H. yunnanensis 1, 10-11 84
Humulus japonicus virus 191 Mechanization of drying 91-2,94,97-9
264 Index
Meiosis 11-12 Plant spacing 50-3, 71, 75-6
Mephosfolan 123-4 Pockets 45, 100
Meristem clones 189 Poles 49, 79-80, 117
Meristem-tip culture 175 Pollen competition 10
Mist propagation 74 Pollen collection 19
Mitosis 12 Pollination 2, 4, 68-70, 216
Mobile harvesters 87-9 Polyphenols 43-5, 123,219-23
Moisture content of dried hops 89, 94, Polyploids 12-16,201,212,215-16
96-7,231 Potassium manures 59, 61
Monoecious plants, see Intersexes Powdery mildew 56-7, 101-4, 107, 109,
Mosaic, see Hop mosaic 150-60,203-5,214-15
Mould, see Powdery mildew chemical control 156
Mutations 158,216-17 cleistocarps 152-4
conidia 152-6
Necrotic ringspot virus (NRSV), see heterothallism 153
Prunus necrotic ringspot virus hygiene 155
Nematodes infection periods 157
hop-root eelworm (Heterodera overwintering 153-4
humulz) 111, 132-3, 182 variety/strain interactions 153, 157-
dagger nematode (Xiphinema 60,204,219-23
diversicaudatum) 131-2, 188 Predators 105-6, 121-5
Nettlehead, see Arabis mosaic virus see also Biological control
Nicotine 106, 119 Prediction
Nitrate content 61 of a-acid content 77-8
Nitrogen manures 59-61, 75, 162 of aphid migration 121-5
Non-cultivation 54-6, 154-5, 162, 165-6 Pre-isomerized extract, see Processed
hops
Oil for drying 91-2 Preservative value 30-1
Organic manures 56,59-62, 166 Proanthocyanidin, see Polyphenols
Otiorrhynchus spp., see Clay-coloured Processed hops 36-7, 45-7, 203, 225,
weevils 233
Overhead sprinkling 64 CO 2 extracts 43, 46
Oxidation of resins 36-7 pellets 45
pre-isomerized extracts 46
Packing dried hops 100 solvent extracts 43, 45
Parasites 121, 124-5 Production methods 49-78
Parthenogenesis 10 in China 53
Pellets, see Processed hops in USA 51, 60, 62-3, 66-7, 69
Peronospora humuli, see Downy Progressive wilt, see Verticillium wilt
mildew Propagation 4, 72-5, 112
pH 59 Prunus spp. 106-7, 115, 118-19
Pheromones 125 Prunus necrotic ringspot virus (NRSV)
Phorodon humu/i, see Aphids 75, 107, 182, 189-91,204
Phosphate manures 59, 61 Pseudoperonospora humu/i, see Downy
Photoperiod, see Daylength mildew
Photosynthesis 20-2 Psylliodes attl:nuata, see Flea beetle
Phytophthora citricola, see Black root rot Pythium intermedium 171
Phytoseiulus persimilis 130-1
Picking, see Harvesting Quality 75-8, 198,202
Planting material 4, 72-5, 175, 188, 191, Quarantine 110, 150
195 Quassia 106, 119
Index 265
Quota 229 Soil-applied insecticides 120, 123-4
Soils 54-6, 59, 129, 132, 188
Recirculation of exhaust air 95-6 Sooty mould 115
Red mould 152-3 Spacing, see Plant spacing
Red spider, see Spider mites Sphaerotheca humuli, see Powdery
Reek 93 mildew
Resin glands, see Lupulin glands Spider mites 57, 127-31
Resins 33-8, 211-13 biological control 130-1
a-acid 2, 33-8, 45-7 chemical control 129-30
a-acid contents 69-71, 75, 178, 184, life history 126-8
190,219-23 Split leaf blotch, see Arabis mosaic
~-acid 33-8, 46, 75 ViruS
hard 33, 36 Spraying 64-6, 115
soft 33, 36, 73 Sterile hops, see Seeded/ seedless hops
Resistance to chemicals Stethorus punctillium 130
of aphids 120-2, 124-5 Stomata 22, 143, 147
of downy mildew 146, 149 Storage material, see Carbohydrates
of spider mites 129-30 Storage organ 20-2
Resistance of varieties Storage stability 35-7, 205-6, 219-23
to aphids 122-4,210,218 Strap cuttings 72-3
to downy mildew 110, 198, 200, 204, Strig 2, 96
206-7,210,214,219-23 Stringing 50-3
to powdery mildew 157-60, 203-4, Stripping 57-8, 84, 144-5, 155
210,214-15,219-23 Sulphur
to spider mites 131,210 as fungicide/ acaricide 106, 129, 156
to verticillium wilt 111, 168, 198-200, in essential oils 40-3
204-5,207,210,213-14,217-23 in hop drying 99-100
Ringspot 189 Sunshine 76
'Ripe-to-flower' 17 Supporting structures 17, 49-53
Rootstock 4, 21-2 low-trellis 66-8, 87
Root system, 4, 54-6, 168 Syrphids, see Hover flies
Rosy rustic moth 134-5 Systemic fungicides 145-6, 156
Row width 50-3 Systemic insecticides 120, 123-4
Runners 56, 72
Tallies 80
Seeds 2, 68-70 Tannins, see Polyphenols
Seeded/seedless hops 2, 68-70, 76, 151- Tax on hops 101,226
2,204,215,231-2 Temperature
Seedlings 10, 185, 148, 159, 198,215 of drying air 93-6
Self-training 57 effect on growth 19-20
Setts, see Bedded setts effect on pests and diseases 127, 129,
Sex chromosomes, see Cytology 146-8, 156, 164, 184
Sex determination 12-16 effect on quality 76-8, 93-4
Sex ratios 10 Tetranychus urticae, see Spider mites
Shoots 4, 29, 56-7, 133, 144-5 Thermal efficiency of drying, see Drying
Sleeves for planting 56 Time of harvest - effect on yield 84
Slope of strings 53 Tissue culture 47, 75, 149,217
Slugs 136 Tokens 80
SNA, see Arabis mosaic virus Trace elements
Soft soap 106, 119 boron 62
Soft-wood cuttings 73-4 zinc 62
266 Index

Training 17,56-7,67 WGV 199

Trellis, see Supporting structures Wye Challenger 129, 157-8, 204
Trickle irrigation, see Irrigation Wye Northdown 132, 178, 193,204
Typhlodromus spp. 130 Wye Saxon 178
Triploids 69~ 70,201,207,215-16 Wye Target 45, 132, 157-8, 169, 204
Yeoman 45, 158
Varietal acreages 207-9 Vectors of viruses 1I5, 1I8, 132, 176-7,
Varietal characteristics 219-23 180-1,191-2
Varieties Vegetative growth 19-21, 139-40
Bramling Cross 199 Verticillium albo-atrum, see
Brewers' Gold 53, 179,200,202-4 Verticillium wilt
Bullion 2, 7, 200, 202-4 Verticillium dahliae 169-70
Cascade 131, 200 Verticillium lecanii, see
Chinook 123,206 Cephalosporium
Clusters 200, 202 Verticillium wilt 56, 60, 72, 110-12, 132,
Comet 131 160-70,179,187,192-3,198-201,
Eroica 206 204-5,213-14,217,219-23
Fuggle 2, 7, 45, 117, 131, 149, 160, effect of nitrogen 166
175, 196, 199-201 effect of temperature 164
Galena 206 in England 11O-1I, 160
Goldings 178, 196, 199-200 fluctuating and progressive forms
Hallertauer 53,164, 178-9, 196,200- 160-4
I in Germany Ill, 164, 166, 179,201,
Hersbrucker 32, 53, 164, 179, 200 205
Hiiller Bitterer 53,179,200 legislation 161-2, 165
Northern Brewer 2,7,45,53, II7, in USA 164,207
150,158,164,179,189,203-5,207 Viroids 192-3
Nugget 206 Viruses 75, llI-12, 175-191,215
Olympic 206 see also American Hop Latent Virus;
Omega 192-3 Arabis Mosaic Virus; Hop Latent
Orion 179,205 Virus; Hop Mosaic Virus;
Perle 123, 179, 200 Humulus japonicus virus; Prunus
Pride of Ringwood 32, 171, 178-9, Necrotic Ringspot Virus; Vectors
203-4 of viruses
Saazer 32, 45, 149, 190,201
Savinja Golding 196 Water requirement 62-3
Spalter 149, 164 Weed control 53-6
'Super Styrians' 205 Wire 49-50
Talisman 202 Wirework, see Supporting structures
Tolhurst 1I7, 122, 160 Wireworm 136