UNIT 7 MODERN TRENDS IN PLANT

TAXONOMY .
Structure
7.1 Introduction
Objectives
7.2 Alpha and Omega Taxonomy
7.3 Morphology in Relation to Taxonomy
7.4 Anatomy in Relation to Taxonomy
7.5 Embryology in Relation to Taxonomy
7.6 Cytology and Biosystematics
7.7 Chemotaxono~ny
7.8 Numerical Taxononly
7.9 Summary
7.10 Terminal Questions
7.11 Answcrs

7.1 INTRODUCTION
You have already read about 'Tools of a taxonomist'. In this unit, you will study
about rccent trends in plant taxonomy. The unit deals with how different bl.anches
of biology helps taxono~nistin synthesising the classification. Each ant1 every
classification collected by taxonomists never becomes obslete and dispensable. It is
retained'pl.ogressively refined and continually added to by successive waves of new
information which accumulate rapidly in the wake of important biological discoveries.
In addition, ncw tools and techniques for studying various organisms provide
interesting observations which are successively exploited by taxonomists. Thus,
taxonomists are synthcsisers of this information and the process of taxonomy and
systematic botany is an unending synthesis.

'Today, the modern taxonomist realises that the ultimate taxonomy of higher plants
rrlust be bascd o n itn understanding of' the morphology, anatomy, embryology,
cytology ancl breeding bchaviour, hesidcs clicmistry and othcr. fcaturcs. This rcqui~.cs
a multi-disciplinery approach and the taxonornist'also uses co~nputersto he111 him
;ln;llyse ciat;~from different nspccts of plant biology.

Thus, taxonomy includes both the latest de\clopmeno and also the present stiite of
the conventional approaches. The next unit deals about modern trends in animal
taxonoln y.

Objectives 1
After studying this unit you will he able to:
0 describe various approaches in the study of plant taxonomy, -. 1

differenti~~te between classical or alpha taxonomy and modern or omega taxonomy, I

e explain the importance of morphology, anatomy and e~nbryologyin relation t o
taxonomy,
e appreciate the use of chromosomal information in cytotaxonomy and biosysternatics,
list the significance of plant chemistry in taxonomy,
0 define numerical taxonomy with suitable examples.
Before we begin our study of modern trends in plant taxonomy, it is necessary to
know about aIpha and omega taxonomy.
I
 Modern Trends La Plant Tovonomy
7.2 ALPHA TAXONOMY AND OMEGA TAXONOMY
7.2.1 Alpha Taxonomy
This relates to the basic or preliminary classification based almost entirely on external
morphology. This can be considered as the empirical approach in taxonomy where
the classification is synthesised from the observed facts. Alpha taxonomy has also
been called Classical or Orthodox or Formal taxonomy. This has been practised since
.ancient times and different aspects of this approach are in use even today. The name
'Alpha taxonomy' was given by Turrill (1935), while two phases of development in
taxonomy come under this, approach according to other taxonomists such as
~ d l e n t i n eand Love (1958), Davis and Heywood (1963). These phases are:
i) the exploratory phase involving collection and subsequent classification; and
ii) the systematic or consolidation phase involving extensive herbarium collections
and field studies for preparing floras, monographs and detailed systems of
classification.
7.2.2 Omega Taxonomy
After synthesising a basic classification, the taxonomist can attempt to improve upon
it. Therefore, the observed facts are interpreted in Omega taxonomy to provide an
interpretive classification. Evolutionary and phylogenetic approaches are applied to
understand taxonomic and evolutionary relationships of plants at all levels. The name
Omega taxonomy was also provided by Turrill (1935). However, this has also been
called Beta taxonomy or Neotaxonomy o r ~ o d e r taxonomy.
n According to Valentine
and Love (1958), this represents the biosystematic or experimental phase in the
development of taxonomy involving detailed cytological and genetical studies. Davis
and Heywood (1963) suggest that Omega taxonomy represents the biosystematic
phase as well as the encyclopedic or holotaxonomic phase in which the taxonomist
analyses and synthesises all kinds of information.

7.3 MORPHOLOGY IN RELATION TO TAXONOMY

Morphology is the basic tool of taxonomy, because identification is primarily based
on the characters of the plant. The morphological characters are easily observable in
both living plants and in herbarium specimens. They have provided the basic
information for a majority of the classification systems in plant taxonomy. In recent
years, electron microscopy has provided a valuable tool to the modern taxonomist to
study different morphological characters at high magnification and this information
is used for purposes of identification, classification and for establishing relationships.
Heywpod and Dakshini used the scanning electron microscope (SEM) to study the
surface patterns of the fruit (called mericarp) in 40 species from 12 genera of the
family Umbelliferae. They found that many microcharacters on the fruit wall observed
with the help of the SEM are of great value in clarifying the relationships of the
different genera. They also observed a great diversity in the structure of the fruit in
these species and this provides new information of significant practical value for
taxonomic purposes.
Most taxonomists have traditionally separated the genus Glinus Linn. from the genus
Mollugo Linn. on the basis of seed characters; but sometimes there were difficulties
in proper identification of the two genera. With the help of the SEM, the seed surface
patterns have been examined in detail to establish the importance of seed surface
microcharacters in the identification of the genera. Also, within the genus Mollugo,
seed-coal micromorphology has helped in identifying different species.
These and other similar jtudies show that even in the present era of specialised and
sophisticated botany, morphological characters continue to provide valuable
taxonomic information.

7.4 ANATOMY IN RELATION TO TAXONOMY

'The use of anatomical*characters in taxonomy began with the development of the
;nicroscope which provided the biologist a new tool to oh~ervethe internal structure
TWIS and Trends in Taxonomy of organs and tissues. it was realised that anatomicai characters are just as valuable
as morphological ones. All parts of a plant provide numerou5 features which have
been used for taxonomic purposes. Some anatomical features are very diagnostic and
are commonly used in routine identification. We also know that this subject is of great
importance to scientisfs who are called upon to ident~fysmall samplestscraps of plant
material for particular purposes such as pharrnacogonosist in the determination of the
source of a drug, or by a forensic expert who may be able to provide clues to a crime
investigation, besides others. These and other similar observations have firmly
established the role of anatomy in plant identification and classification.

The leaf is perhaps the most varied organ of the angiosperms and provides many
anatomical characters of potential taxonomic significance. In your Plant Physiology
Course (Block 3, Unit 13) you have read about Cgand C4pathwaysin photosynthesis.
Investigation of the anatomy of leaves from plants following these pathways has
brought out several significant features associated with the two types. The most
In angiosperms. leaves of distinct character obscrved in the leaves, is the presence of prominent
dicotyledons show reticulate chlorenchymatous sheath surrounding the vascular bundles in the leaves of plants
venation while leaves of showing the Cq pathway and them absence in the leaves of plants showing the C,
mOnocOty'edonssho'v par;'''c'
pathway. Thus, the leaf anatomy also provides information about the photosynthetic
vcnation efficiency of a plant.

As a student of botany, from an early age, we learn that the basic pattern of venation
differs in the two major divisions of the angiosperms. Within each division there are
numerous leaf venation patterns and this feature has beer) used by taxonomist for
understanding taxonomic and phylogenetic relationships in various plant groups.

D r Lalitha Sehgal surveyed the leaf venation patterns in 150 species of the genus ,
Euphorbia. She was able to recognise 12 major patterns and used this information
along with other characters to identify and classify these species. She also correlated
the leaf venation pattern with the habitat of the species and showed that xeropl~ytic
members of the genus had 'accumulated groups of tracheidal elements', while the
herbaceous species having prostrate and ascending habit possessed a sheath around
the veins. Several other features of the venation pattern were also found t o be of
taxonomic significance (table 7.1).

Table 7.1 : Classification of Euphorbja species on the basis of leaf venation patterns.*

Category 1.1 Uni-veined 6 species, e.g. E. incisa

Category 1.2 Bi-veined 4 species, e.g. E. polygonifolia
Category .1.3 Tii-v~lned a majority of the species
Group 1.3.1 Veins Ornamented
Type 1.3.1.1 e.g. E. indivisa
Type 1.3.1.2 e.g. E. hirta
Type 1.3.1.3 e.g. E. granulata
Group 1.3.2 Veins Ornamented
Subgroup 1.3.2.1 Three-stranded midrib
Type 1:3.2.1.1e.g. E. milii
Type 1.3.2.1.2 e.g. E. pulvinofa
Type 1.3.2.1.3 e.g. E.rirucalii
Subgroup 1.3.2.2 Single-stranded midrib
Type 1.3.2,2.1C.g. E. gorgorlis
Type 1.3.2.2.2e.g. E. glareosn
, Type 1.3.2.2.3 e.g. &. peplur
Category 1.4 Special 18 species e.g. E. neriifolia

"Data courtesy Dr Lalita Sehgal

 Modern TmndsinRan'Texonom'
In anothcr interesting study, Biesber and Mahlberg (1981) examined thc laticifer cclls
and scanning elcctron micrographs of the starch grains for interpreting the evolution
within the genus Euphorbia.

Several other features of the leaf anatomy have been used for taxonomic purposes.
Some of these include the nature of the epidermis, stomata1 types, the type of
mesophyll, presence and type of sclereids and crystals etc. Foliar sclerids vary in their
kind and distribution amongst various groups of angiosperms.

Another important anatomical feature used in taxonomy and phylogeny, i s the

structure of the secondary wood. Wood anatomy has been used at all taxonomic
levels. This data along with other characters provides useful evidence for determining
the taxonomic position of taxa whenever two or more possibilities are suggestcd. The
classification of a genus o r a family in its appropriate highel, category can be
determined in this manner. Similarly, petiole vascularisatictn pattern and nodal
anatomy provide increasingly useful taxonomic evidences.

Behnke and his asgociates have investigated more than 1500 species from 380 families
to understand the ultrastructure of the sieve tube plastids. There arc broadly two
types of sieve tube plastids: one' is c a ~ ~ S-type
eb that accumulates starch; while the
other type is called P-type that accumulates protein. This anatomical information
obtained by using the transmission electron n~icroscope(TEM) has been used for
understanding relationships amongst different groups of both dicotyledons and
monocotyledons. P I '

Figure 7.1 : TEM niicrugrnplis shuwing A. S-typc pl~stidsand I%,1'-type plastlds in sieve tubes.

I t is, howrvc~.,important to rr~ncnlbert1i:it a~>atotnic:~l

Fcaturo\ Ilavc proviclctl
outstanding data for taxonomy. Thcse characters whcri used alone miiv not p~ovide
vi lble system of classilici~tion,b u ~when they are combincd with knowlcdgc Srom
other clisciplincs, they hnvc solved Inany taxonomic ~ ~ r o b l e m
Taxono~nists
. should
not separate anatomical data from morphological ch:lrnctcrs and thcy should
incol-porate this information i n heir Corrnal t;lxonomic descriptions.

SAQ 1
a) In the following statcmcnts, put tick ( d ) mark on corrcct oncs and a cross ( x )
on the wrong ones in tlic givcn boxcs.
i) Modern taxonnmis~scan use knowledge from different fields of
inv~sti~atio". a
ii) Alpha taxonomy follows an intcrprctive approach, 1
iii) Omcga taxonomy is also called formal taxonomy. 0
iv) Morphological characters are not used in omega taxonomy.
rods and Trends In Taxonomy v) Plant anatomy provides characters for both alpha and omega
taxonomy. D
vi) Morphology of foliar sclerids in different varieties of the tea plant
can be correlated with the presence or absence of certain chemical
substances in the leaf. 0
Classical taxonomy and modern taxonomy follow distinct methods in solving
taxonomic problems. Name the significant difference in the approach of these
methods.

........................................................................................................
c) Plants showing the Cgand C4photosynthetic pathways can be identified by their
leaf anatomy. Which is the most significant anatomical feature?
In Thallophytes, the zygote
directly develops into the new
plant, or it divides into spores
which then develop inta new
plants. But in Embryophytes
(including Bryophytes,
Pteridophytes, Gymnosperms.
,4ngiosperms) the zygote first d) In what ways does information from sieve-tube plastids help taxonomists?
develops into an e m b ~ y o from
.
which a n adult plant is formed.

Embryological information has been used for taxonomic purposes at various levels of
classification. You know of a very basic division of the plant kingdom into 2 units;
the Thallophytes and the Embryophyta where plants are recognised on the basis of
the behaviour of the zygote in addition to other characters (Table 7.2).
Table 7.2 : Embryological characters used in taxonomy.
1. Anther
2. Quadripartition of the microspore mother cell.
3. Pollen grain.
4. Development and structure of the ovule.
5. Origin and extent ,ofthe sporogenous tissue in the ovule.
6. Megasporogenesis and development of the embryo sac.
7. Form and organisation of the mature embryo sac.
8. Fertilisation.
9 .' Endosperm.
10. Embryo.
11. Seed-coat.
12. Special features.

In the same way you are also aware that the characters of the embryo along with
several other features provide the basis for the division of the angiosperms into 2
major groups, the monocotyledons and the dicotyledons.
In the following paragraphs weshall elaborate on the value of embryological data as
a modirn trend in taxonomy of plants, By studying this section, you shall be able to
know the various kinds of embryological characters used by taxonomists; understand
the importance of embryological characters in taxonomy; and apply this knowledge
from embryology for solving taxonomic problems.
 Embryology strictly refers to the study of the development of the embryo and the
structure of the mature embryo. However, Professor P. Maheshwarl ar~dmany other
famous botanists included, all events whi'ch led to the process of fertilisation besides
the study of the embryo, under the term embryology. This enlarged concept has
proved very useful for providing a large number of characters which can be used for
taxonomic purposes, and during the last 50 years or so, a vast amount of knowledge
has been accumulated which has been used for taxonomic purposes. These features
from sporogenesis, gametogenesis, fertilisation and embryogenesis in flowering plants
have been recognised as less prone to adaptive stress and therefore relatively stable.
They are, therefore, of great siglxificancein plant taxonomy, especially when external
morphology has suggested two or more possibilities concerning taxonomic
relationships.
There are several aspects which favour the use of embryological characteis in
taxonomy. The most significant and important feature is the high degree of correlation
amongst embryological characters. There are, for example, as many as 10
embryological characters which always present themselves together in all plants
classified under the order Ericales (Table 7.3).This highly significant correlation of
characters is very Important for identification of-this order, and no other group of
angiosperms shows all these characters together.

Table 7.3 : Embryological characters of the order Ericales

1. Undifferentiated endothecium
2. Glandular tapetum with multinucleate cells
3. Pollen in permanent tetrads
4. Unitegrnic, tenuinucellate ovules
5. Endothelium present
6. Monosporic &nucleate megagametophyte
7. Elongated zygote
8. Cellular endosperm with the first 4 cells in a linear row
9. Straight embryo
10. Single layered seed-coat

Similarly, reproductive organs show less variability in different climatic conditions

providing stable embryological characters for the purposes of identification. 'The);
also do not show ecotypic variation and remain unchanged .at different ploidy levels
in a polyploid series. These aspects and the fact that most biologists are of the opinion
that embryological characters are conservative, increase the value of embryology as
an important trend in m o d e ~ ntaxonomy.
The value of embryology in solving taxonomic problems can be appreciated by
studying some specific cases. Amongst the classical examples cited by embryologists
is the taxonomic position of the genus Paeonia. In a majority of the classical systems
of angiosperm classification, this genus is considered as a member of the family
Ranunculaceae. However, several botanists have been recognising several characters
of Paeonia which distinguish it from other members of the family Ranuncuiaccae.
These include vascular and floral anatomy, basic chromosome number as well as-the
size and morphology of the chromosomes. The most significant difference bitwee3
the genus Paeonia and other members of the family Ranunculaceae concerns several
embryological characters and this study was carried out independently by Russian
botanists Yakovlev and Yoffe as well as by the hdian botanist Prem Murgai. The
most significant observation relates to' the embryogeny in Paeonia. Unlike other
angiosperms, this genus exhibits a unique type of embryogeny. According to the
Russian botanists, the zygote nucleus undergoes repeated nuclear divisions forming
a coenocytic structure. Later, the nuclei lodge themselves in a peripheral layer of the
-cytoplasm, and this is followed by wall formation so that the peripheral region
becomes cellular. Some of the peripheral cells then function as embryo initials but
only one of these develops into an adult embryo. Murgai on the other hand observed
a division of the zygote into a Zcelled proembryo and the basal cell of this proembryo
develops into the coenocytic structure leading to the development of the adult embryo
as described by the Russian botanists. The other embryological characters which 47
Tau11s and Trrnds in Tnmonnmy
differe'ntiate Poeoniu from the rest of the Ranunculaceae are tabulated below. his
justifies the separation of the genus Paeonia from family Ranunculaceae and its
classification in the family Paeoniaceae (Table 7.4).
Table 7.4 : comparison of embryologicalcharacters of Paconia and Ranunculaceae
Feature Paeonia Ranunculace'ae
Stamen Spirally arranged, centrifugal Spirally arranged. centripetal
Anther Multilayered endothecium, One-layered endothecium and
mostly 2-layered tapetum one-layered tapctum
Pollen Reticulately pitted exine, large Granular, papillate or smooth exine,
and elongate generative cell small lenticular generative cell
Female Multicclled, many megaspore Uni- or multicelled, one cell
archesporium mother cells function functions
Antipodal cells Persistent, not polyploid Persistent (ephemeral in Adonis),
nuclei one or more than one,
polyploid
Embryogeny Unique Onagrad or, rarely, Solanad type
Seed Arillate Non-arillate
Fruit Follicle Achene

W e a r e familiar with the 'water chestnut' commonly called 'Singhara' and botanically
known as Trapa bispinosa. The classification of this plant has seen many changes.
Bentham and Hooker classified the genus Trapa in the family Onagraceae, but other
taxonomists including Engler and Hutchinson moved the genus Trapa t o the family
Trapaceae ,listing several morphological characters as evidence for this separation.
D r Manasi Ram undertook a detailed embryological study of Trapa bispinosa and
listed several characters by which Trapa differs from Onagraceae thus supporting thc
classification of Trapa in a separate family Trapaceae (Table 7.5).
Table 7.5 : Comparison of embryological characlers.of Trapa and Onagraceae

Feature Trapa Onagraceae

Pollen grain Pyramidal with 3 much folded Bluntly triangular, and basin-shaped
rneridional crests
Ovary Semi-inferior, biolocular with a Inferior, mostly tritocular with many
single pendulous anatropous ovule ovules per chamber on an axile
in each chamber placenta
Embryo sac Polygonurn type Oenothera type
Endosperm Absent Present and Nuclear
Embryo Solanad type Onagrad type
Suspensor Well developed suspensor Short and inconspicuous
haustorium
Cotyledons One cotyledon extremely Cotyledons equal
reduced
Fruit Large, one-seeded drupgwith Loculicidal capsule
prominent spines

Besides the examples discussed above, there are many other interesting observations
on t h e genera Exocarpus, Pentaphragma, Butomus, Daphniphyllum, a n d the families
Podosteknaceae, Onagraceae, and Loranthaceae where embryological characters have
been used for understand taxonomic relationships.
~
I

SAQ 2 1

.a) Using embryological information, differentiate between: i

i) Thallophytes and Embryophytes I
 ii) Monocotyledons and Dicotyledons Modern Trends in Plant Taxonomy 1

....................................................................................................
0

b) List six embryological characters used for taxonomic purposes:

iii) .................................................................................................
iv) ..................... . . . .... ..................................................................

I
vi) ................................................................................................. ' i
c ) Mention 3 features which render embryalogical characters significant in solving
taxonomic pmblems.

ii) ............................... ..................................................................

iii) ...................... ........................................................................... :II
d) List 3,genera and 3 families of flowepng plants in which embryological characters
have helped in understanding taxonomic relationships.
Genera Families

ii)
iii)
....................................... .
.
.
.
....
..................................................... .........................a,..........+....

7.6 CYTOTAXONOMU AND BIOSUSTEMATICS

Towards the end of the 19th century and in the early years of the 20th century,
botanists were faced with a problem of analysing variations occurring naturally in
plants. This led to a shift in the emphasis from descriptive classical methods in plant
taxonomy to a new approach called experimental taxonomy whereby attempts were
made to understand the cause of these variations. This new approach, also called
biosystematics, depended largely on establishing the cytological basis of variation.
This use of cytological data in taxonomy is now referred to as cytotaxonomy.
For the purposes of discussion, this trend in plant taxonomy may be considered under
three broad headings:
i) chromosome number,
ii) chromosome morphulogy, and
iii) chromosome behaviour at meiosis.

7.6.1 Chromosome Number

We are generally aware that the number of chromosomes in each cell of all individuals
of a single species is constant. ,It is also established that the more closely related
species are likely to have similar chromosome numbers while the more distantly . 2n is equal to the diploid or
related ones shall have different numbers. Due to this'relative conservativeness, somatic chromosome number:
chromosome number becomes an important and frequently used taxonomic character. while n is the haploid or the
In addition, there is a very wide range of chromosome numbers. in the angiosperms gametic nurnber.
from as low as 2n = 4 (in Haplopappus gracilis) ('Asteraceae) to as high as 2n = 530
(in Poo litterosa) (Poaceae). A large number of angiosperms have been analysed for
their chromosome numbers, providing useful taxonomic information.
Tools and Trends In Taxonomy Many interesting ideas have developed from knowledge of chromosome numbers.
For example, in the genus Festuca, different species have different chromosome
x =n = diploid numbers forming a mathematical series. The chromosome numbers are 2n = 14,28,
2x = n = tctraploid 42, 56, 70, etc. From this information, a generalisation can be made, that different
3x .= n = hexaloid species may have some common basis. If we assume that these chromosome numbers
4x = n = octaploid
are based on a common denominator called x (and x = 7), then we can consider the
5x = n = dccaplo~a
different species to have multiples of this number. This denominator or base number
(x = 7) can be considered as the basic set of genetic information carried by a plant,
and due to the multiplication of this basic genetic set, the eiiolution of different species
has occurred. Such a series is said t o be polyploid in which the basic number (x) is
equivalent to the haploid number of chromosomes in a diploid species (i.e. x = n = 7).
The other species would then be tetraploid, hexaploid, octaploid, decaploid, etc.
respectively.

7.6.2 Chromosome Structure

Cytologists have studied chromosome morphology and have pointed out that the most
interesting feature about the chromosome structure is the position of the centromere.
The centromere or constriction in the length of the chromosome provides information
about the relationship of the 2 arms of the chromosome. Thus, depending on the
position of the centromere, chromosomes are described as metacentric, acrocentric,
and telocentric (Fig. 7.2).

Fig. 7.2 : Chromosome structure-a) acrocentric, b) mctacentrlc, c) telocentric.

The appearance of the basic chromosome set in a dividing cell is known as the
karyotype of the cell. This can be analysed to provide information not only of the
chromosome number, bit also about chromosome size, chromosome volume, and
type of chromosomes in the cell. This information is used by taxonomists for
identifying plants and understanding relationships. The karyotype can be represented
diagrammatically as an ideogram or karyogram, and these diagrams can be compared
for raxonomic purposes. Another interesting observation is that the absolute size of
the chromosomes of a karyotype is fairly constant since it is controlled by the
genotype. Taxonomists have found that rnonocots generally have larger cl~romosomes
than dicots, and that smaller chromosomes are found in hardwood plants in
comparison to their herbaceous relatives.

7.6.3 Chromosome Behaviour

+When we study meiosis. we not only observe the regularity of pairing which is
important'for'the fertility of the plants, we are also able to make a chromosome to
chromosome comparison. This provides valuable information about the role of
chromosomes in heredity. Taxonomists use this information to understand
relationships amongst different species of plants. We can also determine the nature
of the genome to find out if it is homozygous or heterozygous. Genome analysis in
plant taxonomy has been particularly useful in understanding polyploidy add for
establishing the parentage of polyploids. A very significant study in this regard is the
 case of the common hexaploid bread-wheat, Triticum;aestivum. The geri&meof this Modern Trends in plant Ta.;nnnmy
economically important plant has been designated as AABBDD wiih 2n = 42
chromosomes. Detailed genome analyses have established that 3 diploid species have
confiibuted to the evolution of this hexaploid wheat. The A genome is from the
diploid Triticum monococcum (2n = 14), the B genome is from a wild grass Aegilops
speltiodes (2n = 14), and the D genome has been contributed by Aegilopssquarrosa
(2n = 18).

FinaUy, we must remember that as with any other character, the value of
'~ytotaxonomi~ data depends upon the group or category,under irfvestigation. A
combinatton of cytological information with data from other disciplines will provide
'a more useful tool to the taxonomist.

SAQ 3
a) Define
i) Basic chromosome number

ii) Haploid chromosome number

iii) Diploid chromosome number

b) By consulting books in your study centre, find out the diploid chromosome
number in the following plants:

i) Rice (Oryza sntir~u).................................

ii) Potato (Solanurn tubrrosum) .................................

iii) Tea (Then Sinervis) ....................

...................................
iv) Coffee (Coffea arabica) ...................4.,.,..
.

v) Mango ( M a ~ l ~ i f eindica
ra ......;. .........................
vi) Onion (Allium cepn) ...............................
C) Where is thc centromere located in a
i) acrocentric chromosome
....................................................................................................
ii) metacentric chromosome
Tools and Trends in Taxonomy I I ~ ) Telocentric chromosome,

d) What is a karyotype? How does it provide information for cytotaxonomic

purposes?

e) In what ways in genome analysis useful in cytotaxonomy2

- - -

7.7 CHEMOTAXONOMY a

Chemotaxonomy is a science which uses chemical information as a character for

taxonomic purposes. Before we analyse the basis of this modern trend in plant
taxonomy, let us for a moment think about the different kinds of plants in our daily
lives. When we drink tea or coffee, we appreciate the flavour or aroma and
differentiate the two by this character. Similarly, when we eat fruits such as the
mango, the banana or the apple, we find that they taste differently. This difference
is due to the chemical constituents of these foods and this forms the basis of
chemotaxonomy where the chemical features or chemical constituents serve as the
evidence for taxonomy. The potential importance of chemical evidence in plant
taxonomy has been suggested by both botanists and chemists and this has become an
I
important recent trend especially because newer techniques for quick analysis of plant I
material have been developed. Chemotaxonomists suggest that chemical characters I
have a particularly high taxonomic value because they are i) stable, ii) unambiguous,
and iii) not easily (if at all) changeable. Further, chemical characters will show
chemical relationships amongst plants in the same way as morphological characters
show morphological relationships.
Although chemotaxonomy is considered to be a relatively recent development in
modem taxonomy, its origin can be traced to very early classical taxonomy. You will
recall that the spice plants were identified on the basis of their aromatic properties,
or the medicinal plants by their curative value. These aromatic properties or the
curative value was largely based on the chemical constituents of the plants and
taxonomists have classified them since ancient times using these chemical features
along with morphological characters. However, it is only in recent years that
chemotaxonomy as an important field of study has been established.
A review of the large amount of literature published in this field reveals that chemical
data may be obtained from any part of the plant. Secondly, depending on the purpose
of the investigation, the chemical information may be used for description or
identification of plants, or for establishing relationships. This evidence assumes
greater significance when it is used to sort out differences in taxonomic relationships
when 2 or more possibilities are suggested on the basis of morphological characters.
Although theoretically, all chemical constituents of a plant are potentially valuable
to a taxonomist, in practice some kinds of molecules are more useful than others.
Thus we can use directly visible chemical constituents such as crystals, raphides, or
starch grains occurring in different plants as chemical characters.
Alternatively, we can chemically analyse plant material for different chemical
constituents and use this information for taxonomic purposes. Most
chemotaxonomists recognise three broad categories of chemical compounds, primary
metabolites, secondary metabolites, and semantides, as important taxonomically.

7.7.1 Directly Visible Chemical Characters

Very few chemical substances in plants can be observed directly, but the few
substances such,s.starch gtains whicfi are present in most green plants as food
reserves have been used for chemotaxonomic purposes. The type of starch grains Modern Trends In Plant Taxonomy'
present in different plants are very specific and this information can be used without
any ambiguity. Reichert (1913) examined the starch grains in 350 species of plants
and established their differentiation and specificity of occurrence in relation to genera
and species, thus providing chemotaxonomic information. Tateoka (1962) reviewed
the starch grain form in the grass family (Gramineae) and used this as additional
information to classify the fainily into tribes. For example, in the Tribe Hordeae, the
typical members such as Hordeum have compound starch grains, while other genera
like Lolium, Nardus, and Papapholis have simple starch grains.

Fig. 7.3 : Types of starch (After Kochaer 1981).

Raphides are the crystals of calcium oxalate which are present in large cells in different
plant tissues and can be observed directly. They are long needle shaped crystals,
pointed at both ends and usually occur in bundles, thus being easily identified. They
have been observed in as many as 35 families of angiosperms. Several families of the
Order Centrospermae and the Family Cactaceae show the presence of raphides. This
feature along with other chemical characters and the similarity in embryological
characters strengthens the suggestion that the Family Cactaceae shows some
relationship with the Order Centrospermae.

7.7.2 Primary Metabolites

As the name indicates, primary metabolites are molecules involved in vital metabolic
pathways. They are of universal occurrence and not very significant in
chemotaxonomy. However, these molecules become useful as chemotaxonomic
features when the quantity of such molecules varies condiderably between taxa. For
example, the sugar containing carbohydrate 'sedoheptulose' is stored in large
quantities as a reserve food in the genus Sedum. Thus members of this genus can be
easily identified by the presence of this primary metabolite. Interestingly,
sedoheptulose diphosphate is a part of the photosynthetic carbon cycle and in a
majority of the plants sedoheptulose does not accumulate at all. In the same way, the
22 amino acids are of universal occurrence. They serve as the building blocks of
proteins. They can provide useful macromolecular data for chemotaxonomy. The
amino acid sequence of different proteins can be investigated and the degree of
similarity is presumably proportional to the degree of genetic relationship. However,
only a few out of about 3 lakh species of angiosperms have been analysed for amino
acid sequences. For example, the amino acid data on wheat and barley confirms the
relationship of these genera as-suggested by classical taxonomists.

7.7.3 Secondary Metabolites

Secondary metabolites or secondary plant products are those macromolecules that
lhck rlilrogen anu arc of restricted occurrcncc and therefore of greater taxonomic
T~mlsand Trends in Taxtm ..,.!
~mportancethan primary metabolites. This group includes different kinds of
compounds such as phenolics, alkaloids, terpenoids, etc. They are usually not
involved in vital functions and are largely storage products or pigments.
Amongst the secondary metabolites, flavonoids, which are the commonest phenolic
compounds of leaves, have been very useful for chemotaxonomic purposes. Both
monocots and dicots have been extensively surveyed for these compounds which show
structural variability and chemical stability besides widespread distribution. They can
be rapidly and easily identified and provide important chemical characters for
taxonomic purposes. For example, 80 species of plants from the family Ulmaceae
were investigated for their flavonoid chemistry by Ciannasi (1978). A majority of the
species contain flavonols, but a few species have glyco-flavonols and these two types
of flavonoid compounds are never present together in any species. Interestingly,
enough, in most classical systems of classification, the family Ulmaceae is divided,
into two subfamilies called Ulmoideae and Celtoideae which are aiso distinguishable
by the flavonoid chemistry. Therefore, morphological criteria combined with
flavonoid dichotomy can be used to divide the family Ulmaceae (sensu lato) into two
distinct families: family Ulmaceae (sensu stricto) characterised by the presence of
flavonols, and family Celtaceae characterised by the presence of glucoflavonols.
. ,

Several other studies have used flavonoid chemistty for faxonomic purposes in
families such as Arilidaceae, Cornaceae, Labiatae (Lamiaceae), Leguminosae
(Fabaceae), Orchidaceae, Rutaceae, Lemnaceae and others.
A second group of secondary metabolites commonly examined by chemotaxonomists
are the terpenes. Chemically speaking, these compbunds can be classified on the basis
of their molecular {tructure into monoterpenes, diterpenes, triterpenes, ,
sesquiterpenes, etc., and each group can be used for taxonomic purposes. For
examp1e;;n'the genus Salvia, 19 species could be distinctly identified and classified
on the basis of their monoterpenes. The terpene composition was as 'useful as the
morphological'characteristics in the analysis of introgression and hybridisation within
the genus. Similarly, triterpenes and sesquiterpenes have been particularly important
and useful in the classification of the families Cucurbitaceae and Coinpositae
(Asteraceae) respectively.
Other secondary metabolites used in chemotaxonomy include the iridoid compounds,
the alkaloids, and the ellagitannins.

The information carrying molecules in plants are called semantides, and they have
been recognised to be 3 kinds; deoxyribonucleic acid or DNA (primary semantide),
ribonucleic acid or RNA (secondary semantide) and proteins (tertiary semantide)
following the sequential transfer of the genetic code. Of these, the proteins are the
most favoured molecules for chemotaxonomic purposes. Plai~tproteins can be studied
by different methods; by electrophoresis or by serological methods, and both
processes have been used for obtaining information about the protein chemistry of
different plants.
In the common breadwheat, Triticum aestivum, the storage proteins were analysed
by.electrophoresis. For comparative purposes, the storage proteins of the. --tetraploid
.
wheat, Triticum dicoccum and the diplb'id grass Aegilops Squarrosa were also analysed
electrophoretic~lly.TGSstudy confirmed the conclusion that the hexaploid wheat did
contain a sum of the proteins possessed by the diploid species which have contributed
to the evolution of the hexaploid wheat. This study supports the observations based
on morphology and cytological evidence.
Serological analysis of proteins is based on the immunological reaction shown by
mammals when a foreign protein is introduced into the system. In other words, this
is based on the antibody-antigen reaction, the antibodies being specific to an antigen
bringing about coagulation. This information can then be analysed to understand the
relationships of the different plants on the basis of the serological evaluation of the
plant proteins. Serology has proved a useful taxonomic tool at different levels of
classification. J.G. Hawkes (1960) and his co-workers studied several tuber-producing
species of Solanum to understand the evolution of the cultivated potato Solanum
tuberosum and determine the species of Solanum which could be established as the
ancestors of the common cultivated potato, Similarly, in the family Ranunculaceae,
serological studies supported cytological data for the classification of the family into
 Modern Trend8 In Flant Taxonomy
tribes and genera. Fairbrothers (1959) and his CO-workershave studied several plant
groups serologically particularly the members of grass family. A general conclusion
frqm such studies is that the different amount of serological activity in members o f ,
different plant families may b e interpreted as a reflection of the evolutionary
diffcrcnces in the primary structure of the proteins due to which serological
,\il.fl.r~nces
can be recognised between members of different families.

s/ty 4
a) List 3 reasons for using chemical characters for taxonomic purposes:

j) ....................................................................................................
ii) ...................................................................................................
iii) ...................................................................................................

b) Name 3 kinds of chemical constituents which are'useful to the taxonomists.

i) ...................................................................................................
ii) ......................................... .. . ...............................................
iii) ...................................................................................................

c) Name 2 ways by which proteins can b e analysed for taxonomic purposes and name
one gellus analysed by each metbod. Btiefly indicate the principles on which these
two methods are based.

7.8 NUMERICAL TAXONOMY

Taxonomy today is, in many dctails, different from what it was a generation ago. The The book "Numerical
use of computers by taxonomists has established an interesting modern trend called Taxonomy" by P.H.A. Sneath
Numerical 'I'axonomy o r Taximetrics. Mathematical and statistical evaluation of and R.R. Sokal (1973, W.H.
Frooman, San Francisco)
taxonomic information and computation of this data has provided taxonomists with provides valuable information
new approaches to understand classification, lil this section, you shall learn about the relating to the aims, the
fundamentals of numerical taxonomy; know the principles and important terms used principles, the methods and the
in this science; and u~lderstandthe procedures adopted by numerical taxonomists in results of numerical taxonomy
arriving at a classification.

You had studied in Block I, units 1 and 2, that orgar~ismsare classified on the basis
of evidince obtained from thcircharacters. You have also studied that different kinds
of classifications can b e designed by ,using a few or many characters. M. Adanson
(1763) proposed that a classification should use a vast range of characters covering
all aspects of the plants, and in construction of a classification all characters must be
given equal importance. This idea forms the basis of modern numerical taxonomy,
also callcd Neo-Adansonian Taxonomy. So far as the character number is concerned
there is no limitation but larger the number better is the approach for generalisation
of the taxa. You should remember that numerical taxonomy is not a totally new
approach, but it is an organised method of evaluating data with computers in an
objective and repeatable manner enabling comparison of many characters from many
populations of plants.
Tools and Trends in Taxonomy
7.8.1 Principles of Numerical Taxonomy
Numerical taxonomy is based on the following 7 principles.
1) The greater the content of information in the taxa of a classification is and the
more characters on which it is based, the better a given classification will be.
2) Every character is of equal weight in creating natural taxa.
3) The overall similarity between any two,entities is a function of their individual
similarities in each of the many characters for which they are being compared.
4) Distinct taxa can be recognised because correlations of characters differ jn the
groups of organisms under study.
5) Phylogenetic inferences can be made from the taxoriomic structure of a group
and from character correlations, given certain assumptions about evolutionary
pathways and mechanisms.
6) Taxonomy is viewed and practised as an empirical science. Classifications are
based on phenetic similarity.
7) Classifications are based on phenetic similarity.

7.8.2 Procedures Adopted by Numerical Taxonomists

Since numerical taxonomy is an operational science, the procedure is divided into a
number of repeatable steps, allowing theresults to bc checked at every step.
i) Choiceof units to be studied: The first step is to decide what kind of units to study.
In numerical taxonomy, the basic unit of study is called the "operational
taxonomic unit" (OTU). Thus the OTU can be an individual plant if the
taxonomist is studying a single population of plants to find out the range of
variations in its characters. Similarly, you may treat an entire population of plants
as an OTU if you are studying a single species represented by different
populati'ons existing in nature; or the OTU may be different species wheii genus'
is being evaluated. Therefore, in numerical taxonomy, the O T U varies with the
material being studied, and this' helps the taxonomists in making an objective
study.
ii) Character selection : After selecting the OTU's, it is necessary to select characters
by which they are to be classificd. By experience, you will lCarr1 lhnt characters
which vary greatly amongst the OTU's are clearly more useful in numerical
taxonomy; and we know that as many characlers as possible m a y be used.
Preferably a mininlunl of GO and generally 80 to 100 or more characters arc
needed to produce a fairly stable and reliable classification. T h e selected
e 2
characters have thcn to be coded or given some symbol or ~ n a r k . ' ~ h e rare
methods of coding taxonomic infortnation.

a) Binary coding or two-state coding-This is the simplest form of coding adopted

+
in numerical taxonomy where the characters are divided into and -, or as 1
+
and 0. The positive characters are recorded as or 1 and the negative cl~aractcrs
as - or 0. It is possible to use this method of coding for all characters studied.
In case a particular character is not present in an O T U being examined, the
symbol or code NC is used, indicating that there is no comparison for thnt
characters. However, we find that by using this method of coding, we tend to
increase our workhecause there are large variations in the plant, and very often
a single character such as colour of flowcr can be rcpresentcd in a wide range
We can have white, pink, red, yellow and other colours in roses. If we are to use
this data in a binary coding, thcn we will have to use each colour as a character
and it would be coded as + or - , as the casc lnay be.

b) Multi-state coding-An alternative method would be to use multi-state coding

where a single character can be coded in a number of states, each being
..
represented by a numerical symbol or code (e.g. 1 , 2 , 3 , 4 , 5 , , ..) depending on
the range of variation. Thus, if we again look at the colour of the rose flower, we
can give different codes to different colours such as white = 1 , pink = 2, red = 3.
ycllow = 4, and so on. Bcsidcs qualitative characters sucll as colour of tlowci.
type of placentation, etc., multistate coding is also useful of quantitative
characters such a s plant height, leaf length, leaf breadth, and other characters Modern Trends in PIMI Taxonomy
involving measurements. A code is prepared for the range of variation and
appropriate symbols are allotted to each unit in the. range.

m e data obtained by scoring the characters in the OTU's ?re then presented in a
table as a .data matrix giving the OTU's on one side of the table and the codes for
different'charactersagainst each OTU. Thus, if one has studied 25 OTU's and has
scored 75 characters from each, the data matrix will contain 25 x 75 = 1875 units of
information. his kind of l a r ~ unit
e of information in the data matrix necessitates the
use'of computers to help the taxgnomists. to digest the knowledge qurckly. It is also
important to.remember that computer programmes are based on mathematical
equations and-computer language and the data matrix is essential for this purpose.
In addition, 'tlie.next step in numerical taxonomy is entirely dependent ;on the data
matrix.
The information is then presented in a t x n table or data matrix consisting of OTU's
scored for n characters (table 7.6).
Table 7.6 : Coded data (t X n table)
Characters (n) T p ~ pOTU'S (t)
(1-12)

(After Sneath, 1962, in Microbial ~lossificution,edited by Ainsworth crnd Snenth.)

iii) Measurement of Similarity : Overall similarity (s) is calculated by comparing each

OTU with every other and is usually expressed as a percentage, 100 per cent S
for identity and 0 per cent S for no resemblance. A similarity table or matrix is
then constructed by tabulating the S coefficients for each one of the OTU
(Fig. 7.4).

iv) Cluster analysis : After making a similarity table, it is then rearranged so that
OTU's whose members have the highest mutual similarity are brought together.
This can be done by several methods and related taxas or groups are rec~gnised.
These clusters are called phenons and can be arranged hierarchically in a tree
diagram or dendrogram (Fig. 7.5).

The groups or clusters thus recognised may be treated as equivalent to the categories
having ranks in classical taxonomy, such as the genus, family, order, etc. A problem
faced by many taxonomists is whether there is any equivalence between the ranks in
different taxonomic groups of organisms. Is a family of flowering plants, for example
equivalent in any sensz to one of algae, or other organisms? To overcome this
problem, numerical taxonomists have advocated a new terminology. Here, the tern
"phenon" is introduced and the particular phenons are desipated by numerical
prefixes (e.g. 80 similarity) showing the level of. resemblance by which they are
defined. The delimitation of the phenons is done by drawing horizontal lines across
the dendrogram at a chosen similarity value. Such a dendrogram will have reference
to a given study only, and cannot be generalised. Thus, phenons will be arbitrary and
relative t~ groups within the limits of only one ~nalysis.
Tools and Trends in Taxonomy

(a)

A B E I C P G J D H

Percentage similarity Taxa

Fig.7.4 : a) Scbematlediagram ghowing'a matrix of hypothetical slmliarity coefficent between of

group (taxa);the magnitude of the coefficient.isshown hy the depth of shadlag.
b) The same coemcient arranged by placing similar taxa next to each other; this giva trinngle of
hlgh similarity values. Phenons are groups by desired rank (After Sneaths 1962, Microbial
clnsailication)

Taxa

per cent phenon lines

per cent phenon lines

per cent phenon lines

Fig. 7.5 : A dendrogran~representing the hypothetical hierarchy of group (taxa) obtained from Fig 7.4. The
ordinate indicates magnitude of similarity cwfncient at which stems join to form higher ranking
group. H o h n t s l delimit groups of equal rank (per cent phenon lines). (After Sneath, 1962)
Microbial Ciwiflcation, edited by Ainsworth and Sneath. University of Camhri.lge).
 01 the numerous numerical taxonomic studies, the reclassification of the dicotyledbns TrendsInp w ,
by Young arid Watson (1970) serves as a good example. They studied 83 characters
frqm mor~hologyand anatomy in 543 genera and computed this information to
classify'these angiosperms. The computer-based classification was in many ways
similar to other traditional classification, as well as in some ways different. However,
the different genera were clearly distinguishable into distinct categories on the basis
of the nature of the ovule into crassinucellate (ovules with a massive nucellus) and
tenuinucelfate (ovules with .a small amount of nucellus). In view of these, some
taxonomists are of the opinion that numerical taxonomy may never replace traditional
methods as standard procedure. It would, however, be successful where other
methods have failed or are laborious or otherwise difficult to apply. The most
important contribution of numerical taxonomy has been to help taxonomists analyse
their methods, data, and conclusions more logically and objectively.
I

SAQ 5
a) Why is numerical taxonomy also called Neo-Adansonian taxonomy?

b) Briefly describe the procedure adopted by Numerical taxonomists.

c) What is meant by 'opcr:~tioiialtiixonomic unit' (OTU)'?

d) Differentiate hetwecn binary coding and multi-state coding.

c*) Mention 2 uscs of I: dntn ~natrixin numerical taxonomy.

............................................................................................................
 What is 'cluster analysis' and how is this arrived at in numerical taxonomy?

7.9 SUMMARY
In t h i ~ u n i t we
, have briefly discussed various aspects of modern trends in plant
taxonomy. This has helped you to study that:
Taxonomy utilises data from all the other branches of biology.
0 Taxonomic information never becomes obsolete, but it is progressively refined and
successively exploited by taxonomists to synthesise systems of classification.
@ Modern taxonomy follows a multi-disciplinary approach.

Alpha taxonomy or classical taxonomy is based on an empirical approach involving

the exploratory or pioneer phase and the systematic o r consolidation phase of
development of taxonomic knowledge.
0 Modem or omega taxonomy follows an interpretive approach using evolutionary
information to understand taxonomic and phylogenetic relationships. This involves
usage of knowledge from the biosystematic or experimental phase and the
encyclopedic or holotaxonomic phase of taxonomy.
0 Morphological and anatomical characters can be studied with light as well as
electron microscopes, and this information is useful for solving taxonomic
problems.
0 Plant embryology has proved extremely useful as a source of evidence in taxonomy

0 Cytological and biosystematic data uses information from chromosome number,

structure, and behaviour, and this helps taxonomists to understand relationships
amongst plants.
Chemotaxonomy provides useful data from chemical characters which show
chemical relationships amongst plants in the same way as morphological characters
show morphological relationships. Different kinds of chemical compounds can be
evaluated in different ways to provide taxonomically useful information.
Taxonomists can use computers to analyse large amounts of information in
numerical taxonomy.

7.10 TERMINAL OUESTIONS

1) "Systematic botany is an unending synthesis". Elaborate.
2) a) Differentiate between "Ranunculaceae sensu lato" and "Ranunculaceae
sensu stricto"
b) Name the genus which led to this change in the circumscription of the family.
c) List the important characters responsible for this change.
3) The common breadwheat, Triticum aestivum, is a hexaploid. List evidences from
cytotaxonomy and chemotaxonomy to support the evolution of this hexaploid
wheat from diploid ancestors.
4) Briefly outline the procedures adopted in numerical taxonomy.
5)' Write explanatory notes on:
a) Biosystematics
. b) Secondary metabolites as characters in chemotaxonomy
.c) Cluster analysis in numerical taxonomy
 Modern Trends in Plant Tuxonorny
7.3131 ANSWERS
Self-assessment Questions
1) a) i) d, ii) x iii) x , iv) X, -v) d, vij d
b) Classical taxonomy follows an empirical approach synthesising a basic
classification from observed facts. Modern taxonomy follows an interpretive
approach improving upon the basic classification.
c) The most significant anatomical feature differentiating leaves of C3 plants
from C4 plants is the absence of a chlorenchymatous bundle sheath in the
former and its presence in the latter.
d) sieve tube plastids are basically of two types, called S-type (accumulating
starch) and P-type (accumulating proteins). These are recognisable by
electron micrographic studies and different plant families have been
characterised on the basis of this information. It has also been possible to
understand relationships amongst monocotyledons and dicotyledons using
sieve tube plastid type as evidence.
2) a) i) In Thallophytes, the zygote directly develops into a new plant or it
produces spores which develop into new plants. In Embryophytes, the
zygote first develops into an embryo from which the new individual plant
is then formed.
ii) In nionocytyledons the mature embryo has one cotyledon, while in
dicotyledons it has two.
b) .List any six characters from Table 7.2.
c) i) There is a high degree of correlation amongst embryological characters.
ii) Due to less variability in floral characters, embryological characters,
show stability.
iii) Embryological characters are not affected by ecotypic variation and they
remain unchanged.
d) Genera Families
i) Butornus i) Loranthaceae
ii) Daphrliphyllum ii) Podostemaceae
iii) Exocarpus iii) Ranunculaceae
or any other genus or family in which embryological characters have been used
for taxonomic purposes.
3) a) i) Basic chromosome number refers to the basic set of genetic information
irl the chromosomes of an individual of a species or a polyploid series.

ii) Haploid chromosome number is the gametic chromosome number

present in the sex cclls or the gametes.
iii) Diploid chromosome number IS the somatic chromosome number usually
seen in the vegetative cells of an individual.
b) Diploid chromosome number in:
i) Rice - 2n = 24, ii) Potato - 2n = 48, iii) Tea - 2n = 303
-
iv) Coffee 2n = 44, v) Mango - 2n = 40, vi) Onion - 2n = 16
C) i) The centron~ereis'located in between the short arm and the long arm of
the chromosome.
ii) It is located in the middle of the two equal arms of the chromosome.
iii) It is located at the tip of the arm of the chromosome.
d) The karyotype refers to the appearance of the basic chromosome set
(genome) under the light micr~scope.It tells us about the number of
chromosomes, the diffcsent types of chromo~o!nesand their relationship with
each other. This information can be compared for cytotaxonomic purposes,
e) Genome analysis is the study of chromosome pairing in diploid hybrids. It
helps in the investigation of polyploids for determining the ancestral genomes
t
and this information is used for ascertaining taxonomic relationships
 l s I rends in l a111n11tn?
T ~ ~ rand
4) a) i) Chemical characters are stable
ii) They are unambiguous
iii) Chemical characters are not easily changeable.
b) i) Directly visible chemical constituents such as starch grains.
) Secondary metabolites such as phenolic compounds, alkaloids, etc.
iii) Semantides or information carrying molecules such as DNA, RNA, or
proteins.
I c) Proteins can be analysed by gel-electrophoresis or by serological methods.
The genus Triticurn has been analysed by gel-electrophoresis, while the genus
Solanurn has been analysed by serological methods.
Proteins are separated electrophoretica~l~ on the basis of their arnphoteric
properties in a column of acrylamide gel which is positively or negatively
charged to various extents. In serology, proteins are analysed on the basis of
the immunological reactions. This requires a test animal to evaluate the
antibody-antigen reaction.
Numerical taxonomy is also called Neo-Adansonian taxonomy because it is
based o n the same basic taxonomic principles formulated by M. Adanson in
the 18th century.
Carefully read the Section 7.8.2 and rewrite the procedure in your own words.
'Operational Taxonomic Unit' or OTU is the basic unit of study in numerical
taxonomy. It can vary with the nature of the material being investigated as
well as the purpose o f t h e investigation.
Binary coding is two-state coding according to which every character studied
can be analysed in 2 states viz, present (+) or absent (-); whereas multi-state
coding analyses more than two states in which a particular character exists
and each state is given a definite symbol or code.
The data matrix basically presents taxonomic information in a tabular form
for all the. OTU's examined and all the characters studies. It 'also provides
information for clustering of the OTU's.
Cluster analysis is a process by which the OTU's are sarted out to form groups
or clusters on the basis of their oyerall similarity. Similarity or dissimilarity
coefficients are calculated by comparing each OTU with every other OTU
and this is represented as a percentage. This information is used for cluster
analysis in numerical taxonomy.
Terminal Questions
1) Taxonomists utilise information from many disciplines for synthesising
classifications. This process is a continuous one. The material provided in this
unit has briefly described the developments in modern taxonomy to elaborate
the unending nature of systematic botany. You must carefully study the entire
unit and write out an essay of about 1500 words to answer this question.
2) a) Ranunculaceae 'Sensu lato' refers to the broad concept in the taxonomy of
the family Ranunculaceae. While Ranunculaceae 'sensu stricto' refers to a
restricted concept after one or more genera have been removed from the
family and classi'fied in a separate family, thus changing the circumscription
of the family.
b) The genus Paeonia was first classified in the family Ranunculaceae (broad
circumscription) but later it was separated from this family Ranunculaceae
giving it a restricted circumscription.
 c) See answer 2 d . Modern Trends in Plant Texonomy

'3) The origin of the hexaploid breadwheat Triticum aestivum from diploid ancestors
has been established cytotaxonomically. ~riticam'm~n'ococcum and Aegilops
peltoides both diploid species can hybridise and the hybrid can, by chromosome
doubling give rise to a tetraploid wheat, Triticum dicoccum. This in turn can
hybriaise with another diploid species,, Aegilops squarrosa and by doubling the
chromosomes of the hybrid, the hexaploid wheat, Triticum aestivum is produced.
In chemotaxonomy, protein gel-electrophoresis of the seed proteins from
Triticum dicoccum, Aegilops squarrosa, and Triticum aestivum provide evidence
to show that the hexaploid wheat arose by hybridisation.
4 ) Carefully read Section 7.6 for (a), 7.7.3 for (b) 7.8 for (c).
5). a) See Section 7.4
b) See Section 7..5.3
c) See Section 7.8.2, particularly the paragraphs on cluster analysis as in answer
5 f.