Growth and Development of

the Pearl Millet Plant

Research Bulletin No. 6

International Crops Research Institute for the Semi-Arid

Tropics
Abstract R6sum0

Maiti, R.K., and Bidinger, F.R. 1981. Growth and Maiti, R.K., and Bidinger, F.R. 1981. Growth and
development of the pearl mllet plant. Research development of the pearl millet plant. (DMve-
Bulletin no.6. Patancheru, A.P., India: Inter-
Zoppement du muid chndelle.) Research Bulletin
national Crops Research Institute for the Semi-
No.6. Patancheru, A.P., India: International
Arid Tropics.
Crops Research Institute for the Semi-Arid Tropics.
Pearl millet (Pennisetwn aericanwn [L.] Leeke) Le mil a chandelle (Pennisetin americanzn [L.]
is one of the most important food crops in the Leeke) est l'une des plus importantes cultures
drier semi-arid tropics. The bulletin describes vivri~res des parties les plus s~ches des zones
its three major growth phases: the vegetative tropicales semi-arides. Ce bulletin dcrit ses
phase, from emergence to panicle initiation of trois phases de croissance principales: la phase
the main stem; the panicle development phase, v6g~tative, allant de l'6mergence a la formation
from panicle initiation to flowering in the main de la panicule sur la tige principala; la phase
stem; and the grain-filling phase, from flower- du d~veloppement de la panicule, allant de sa
ing to physiological maturity. Each phase has formation a la floraison sur la tige principale;
been subdivided to make - total of nine morpho- la phase du remplissage des grains allant de la
logically distinct and recognizable growth floraison a la maturit physiologique. Chacune
stages. Descriptions and characteristics to de ces phases a 6t6 subdiviste et l'on retrouve
identify each of these stages are supported en tout neuf stades de d~veloppement morpholo­
with 21 illustrations, some of them in color.
giquement distincts et identifiables. Vingt-et­
The bulletin also describes the growth and
une illustrations, dont quelques-unes en couleurs,
development of the major plant parts: root,
accompagnent les descriptions et les caract6ris­
tiller, leaf, stem, panicle, and grain. The
tiques de chacun de ces stades et en facilite

data given are derived from several years of

l'identification. Ce bulletin d6crit aussi le

research on the physiology of the crop, and the

d6veloppement des principales parties de la plante:

publication should thus serve nct only as a

racine, talle, feuille, tige, panicule et graine.

source of basic information on the growth and

Plusieurs ann~es de recherches sur la physiologie

development of pearl millet, but as a reference

de cette plante ont fourni les donnes de ce

for comparing millet with other cereal crops

bulletin. II peut servir A la fois comme source

based on a standard system for recording the

d'information de base sur le d~veloppement du mil

developmental stages of the crop.

a chandelle et permettre de comparer cette plante

A d'autres cultures c~r~ali~res en utilisant un

systme uniforme pour enregistrer les stades de

d6veloppement.

 Growth and Development

of the

Pearl Millet Plant

R.K. Maiti and F.R. Bidinger

9
ICRISAT

Research Bulletin No.6

International Crops Research Institute for the Semi-Arid Tropics

ICRISAT Patancheru P.O., Andhra Pradesh 502 324, India

October 1981

 CONTENTS

Introduction 2

General outline of crop growth (growth phases) 2

GS, Vegetative phase 2

GS2 • Panicle development phase 3

GS3 Grain-filling phase 3

Detailed stages of growth 3

Introduction 3

Stage 0 Emergence 4

Stage 1 • Three-leaf stage 4

Stage 2 Five-leaf stage 4

Stage 3 • Panicle initiation 4

Stage 4 Flag leaf stage 5

Stage 5 Boot stage 6

Stage 6 • 50% flowering 6

Stage 7 Milk stage 7

Stage 8 ' Dough stage 7

Stage 9 • Physiological maturity 7

Growth and development of individual organs 7

Root development 8

Tiller development 9

Leaf area development 10

Stem elongation 10

Panicle development 11

Grain growth 12

Distribution of plant dry weight 13

Acknowledgements 14

Bibliography 14

 Introduction growth and development of pearl millet for students

and trainees, as a reference for comparing pearl
In the drier semi-arid tropics of the world, pearl millet with other cereal crops for scientists and
millet (Penniseturn americanum jL.] teachers, and as a standard system of recording the
Leeke = Penniseturn typhoides Stapf. and Hubbard) developmental stages of the crop for agronomists,
is one of the most important crops of the small plant protection scientists, and others working with
farmer. It is particularly adapted to conditions of this crop.
nutrient-poor soils and low rainfall, yet is capab!e of
rapid and vigorous growth under favorable
conditions. It is grown both as a grain and aforage General Outline of Crop Growth

crop and is the major cereal for the people in the (Growth Phases)

drier areas of the Indian subcontinent and of both

West and East Africa. Yet until very recent times, The growth cycle of pearl millet may be divided into
pearl millet received comparatively little attention three major developmental phases: the vegetative
from the scientific community. phase (GS 1)-from emergence to panicle (floral)
This research bulletin provides a description of initiation of the main stem; the panicle development
the general growth and development of pearl millet phase (GS)-from panicle initiation to flowering of
based on three major growth phases, followed by a the main stem; and the grain-filling phase (GS)­
more detailed description ot a series of nine stages from flowering to the end of the grain-filling period
into which the growth cycle can be divided for easy (physiological maturity) of the crnp (Fig. 1).
reference and description. A detailed account of the
development of each of the major plant parts, based GS, : Vegetative phase
on information developed at ICRISAT, is presented This phase starts with the emergence of the
in the concluding section. seedlings and continues up to the point of panicle
The information in this bulletin shouid find initiation. During this phase, the seedlings establish
several uses: as a source of basic knowledge of the their primary root system (seminal roots) and

4 5 6 7 8 9

A>

Vegetative phase (GS) Panicle development phase (GS2 ) Grain-filling phase (GS3 )

Figure 1. Schematic diagram of the major growth phases of pearl millet: GS, GS , GS,. The numbers 0 to
2 9 illustrate thp
detailed stages of growth listed in Table 1,and described on pp. 2-6. The encircled enlargement of stage
3
shows the dome-iike shape of the apex, and the constriction at its base, that may be observed at tlis stage
of
change from the vegetative to 'he reproductive phase.

2
 produce adventitious roots. All leaves are initiated grain-.filling phase. These tillers have a shorter
during GS, and, in early varieties, six orseven leaves developmental cycle than the basal tillers,
(including the embryonic leaves) are fully expanded producing only a few leaves and a small panicle.
by the end of this phase. Yiller buds are formed, their The end of the grain-filling phase (physiological
leaf primordia initiated, and several tillers emerge maturity) is marked by the development of a small
by the end of the phase. There is little internode dark layer of tissue in the hilar region of the grain
elongation, however, and the apical meristem (Fig. 9). This occurs in an individual panicle about
remains at or below the soil surface. Dry-matter 20-25 days after flowering. The grain-filling period
accumulation is almost entirely confined to leaves for the entire plant (i.e., from flowering of the main
and roots. shoot to the end of grain filling of the tillers) is
Floral or panicle initiation is marked by the longer where tillers flower after the main panicle.
elongation of the apical dome and the formation of a
constriction at the base of the apex (Fig. 4). The size
of the apex at floral initiation ranges from as little as Detailed Stages of Growth
0.5 mm in early varieties to as much as 1.0 mm in :ate
varieties in which floral initiation may not occur until Introduction
50-80 days after sowing.
The following detailed description of the growth
GS 2 : Panicle development phase 3tages of a pearl millet plant divides the life span of
the plant into nine morphologically distinct stages
During this phase all the remaining leaves expand following Vanderlip. 1 The stages and the
fully and the earliest expanded leaves at the base of characteristics to identify them are presented in
the stem begin to senesce. Stem elongation occurs Table 1. The der.criptions refer to the development
by sequential elongation of internodes beginning at of the main shoot only, which is used to identify the
the base of the stem. Tillers emerge, undergo floral
initiation, leaf expansion, etc., in patterns similar to .......

that of the main stem. The first-formed tillers follow Table 1. Developmental timetahle by growth stages of
the main stem closely in their development, whereas the main CLIIno in HB-3 and MI Z ongo. and
the development oi the late tillers frequently ceases duration of major growth phases
due to competition and/or suppression by the mcre Approximate days after
advanced main stem and early tillers. Dry-matter Growth Identifying emergence
accumulation takes place in roots, leaves, and stem. stage characteristic
During stem elongation the panicle undergoes a HB-3 Mil Zongo,
series of distinct morphological and developmental 0. coeoptne visible at 0 0
changes, described below. These include the soil surface
development of spikelets, florets, glumes, stigmas, 1. Third leaf visible 6 6
anthers, and finally stigma emergence (flowering) 2. Fifth leaf visible 14 15
and pollination, which marks the end of the GS, 3. Panicle initiation 22 28
phase. 4. Final or flag leaf visible 33 43
5. Panicle extended in flag 36 47
leaf sheath
GS3 : Grain-filling phase 6. 50% stigma emergence 40 53
7. Milk stage 49 61
This phase begins with the fertilization of florets in 8. Dough stage 58 69
the panicle of the main shoot and continues to 9. Black layer formation 65 75
maturity of the plant (main stem and tillers).
Increases in total plant dry weight during this period Major growth phase Approximate duration (days)
are largely in the grain but, as tillers in many HB-3 Mil Zongo
varieties elongate and flower after the main shoot,
there is also some increase in nongrain GS, 18 25
components, mainly tiller stems. G 25 22
Senescence of the lower leaves continues and,
by the end of the grain-filling phase, normally only
the upper two to four leaves remain green. Some VANDERLIP, R.L. 1972. How a sorghum plant develops.
varieties develop small tillers in the upper nodes of Kansas State Un'versity, Cooperative Experimental Services,
the stem, particularly towards the end of the Report C-447.

3
stage of development of the pla t. The description
and illustrations relate to HB-3 (a commercial
hybrid of the All India Coordinated Millet
Improvement Program) and to an early flowering
selection from Mil Zongo (aWestAfrican landrace).
Developmental rates are a function of the
environmental conditions in which the plant is
grown as well as the variety grown. Day length, for
example, greatly affects the time to panicle initiation
in many varieties, and temperature affects rates of
leaf emergence, length of the grain-filling period,
etc. The number of days for the attainment of the
various growth stages indicated in Table 1 are the
averages for environmental conditions during June-
September at Hyderabad, India (170 N latitude).
Durations of growth stages may vary considerably
for other locations and other varieties, pqrticularly
for West African landraces which typicaly do not
reach panicle initiation in their West African
environments until 50-80 days from emergence.
These varieties thus flower in 80-120 days after
emergence rather than in 40-50 days as Indian Figure 2. Growth stage O: Germination and emergence
varieties do. As a consequence, the West African of the seedling.
types produce many more leaves, stem internodes,
and much more total dry weight than do the Indian
varieties, two adventitious root initials are visible. The leaves
are small in size and light green in color (Fig. 3).
Stage 0 : Emergence
Stage 2 : Five-leaf stage
This stage begins with the emergence of the
coleoptile from the soil surface. Before this occurs, About 13-15 days after emergence the lamina of the
a number of changes have taken place fifth leaf is visible. The first and second leaves are
underground. Germination begins with the uptake fully expanded. The third leaf is still slightly rolled
of water by the seed, which activates metabolism in (Fig. 3). The growing point remains below ground
the cells. Within approximately 16 hours after the level, surrounded by the developing leaf primordia.
initiation of germination, the radicle emerges from The seminal root is now well developed and has a
near the hilar region, followed by the plumu'e with number of branches. Adventitious roots start
the coleoptile sheath approximately 2 hours later. appearing (Fig. 3). Tiller leaves may be seen
The radicle grows downwards rapidly and produces emerging from inside the sheaths of the basal leaves
fine root hairs (Fig. 2). The coleoptile grows at this stage. The plant now appears dark green and
upwards slowly through the soil until it emerges more sturdy.
from the soil surface. The time required from
germination to emergence depends on the depth of Stage 3 : Panicle initiation (growing point
planting, soil moisture, and temperature; under dihferentiation)
favorable conditions it takes 2-3 days.
At this stage the growing point changes from the
Stage 1: Three-leaf stage vegetative to the reproductive stage, that is, from the
development of leaf primordia to the development
Approximately 5 days after emergence of the of spikelet primordia. This change can be
coleoptile, the lamina of the third leaf can just be recognized by the fact that the apex becomes
seen in the whorl of the second leaf without dome-like and a constriction develops at its base
separating the first and second leaves. The first leaf (Fig. 4).
is fully expanded and the second leaf is still slightly All leaves have been initiated at this stage; six to
rolled at the base. At this stage the seminal root seven leaves are fully expanded in early varieties
grows rapidly and develops fine branches. One or and the remaining are in various stages of

4
 Figure 4. Growth stage 3 : Panicle initiation. The
primordium on the left has just reached
initiation; the one on the right is
approximately 2 days past initiation.

Figure 5. Growth stage 4: Flag leaf emergence. The tip

of the flag leaf lamina is visible in the whorl of
the second leaf.

Figure 3. Growth stages 1and 2: Three-leaf stage (left)

and Five-leave stage (right). In both cases the
tip of the lamina of the new leaf is just visible in
the whorl of previous leaves.

development. Following initiation, the growing

point is above the soil surface as the first two or
three internodes begin to elongate. The seminal
root has produced a network of lateral roots and the
adventitious roots are growing rapidly. A number of
tillors have now emerged and are undergoing the
same developmental pattern as that of the main
shoot, although the main shoot is more advanced
than the tillers.

Stage 4 : Flag leaf stage

The lamina of the final leaf is visible in the rolled

lamina of the preceding leaf. The final leaf is easily
distinguished from the preceding leaves as there are
no other l9avas within the rolled lamina of this leaf
as it emerges from the whorl (Fig. 5). Between
panicle initiation and the appearance of the flag leaf,

5
there are numerous changs which occur in the completion. The panicle rapidly increases in length
plant. The unexpanded leaves that were present as and width.
initials at the time of panicle initiation, sequentially Following the boot stage, the panicle emerges
develop, emerge from the whorl, and expand to full from the collar of the flag leaf as the peduncle (the
size. The stern rnternodes elongate in sequence, uppermost internode) begins to elongate.
beginning from the base, with each successive
internode longer than the previous one. Branch, Stage 6 : 50% flowering (h.lf bloom)
spikelet and floret primordia are initiated in
sequence, the process beginning at the base of the Pearl millet is protogynous, i.e., the stigmas appear
panicle meristem and proceeding towards the top. first. The stigmas begin emerging about 3-5 days
By the flag leaf stage, the florets are undergoing after panicle emergence, though this varies with
rapid development. The panicle is enclosed by the genotype. Stigma emergence starts generally in the
sheaths of the flag and penultimate leaves, and is florets several centimeters below the tip of the
raised well above the soil surface by the elongation panicle and then proceeds upwards and downwards
of the lower internodes. simultaneously. 50% flowering is attained by the
time tlie stigmas emerge in the middle region of the
Stage 5 : Boot stage panicle (Fig. 7). It takes 2-3 days for the completion
of stigma emergence, and unpollinated stigmas
The panicle at this stage is enclosed within the may remain fresh for several days. However, on
sheath of the flag leaf but has not yet emerged from pollination stigmas shrivel within a few hours.
the collar (Fig. 6). Panicle development is nearing On completion of stigma emergence, or in some

Figure 6. Growth stage 5 : Boot stage. The panicle is Figure 7. Growth stage 6 : 50% stigma emergence (left)
enclosed in the sheath of the flag leaf. and 50% anther emergence (right).
cases slightly before completion, the emergence of Stage 8: Dough stage
the first flush of anthers begins from near the top of
the panicle and proceeds toward the base of the This stage is identified by the change in the
panicle. The process takes 2-3 days for completion endosperm from the mainly liquid milk stage to a
(Fig. 7). Anthers emerge in two flushes-the first first semisolid and then a solid state. It is a gradual
flush occurs in perfect flowers and the second from change and not a distinct stage, occurring as starch
the male flowers; thus anther emergence from one content in the endosperm increases and the
head may extend over 5-6 days. Depending on the moisture percentage declines. Within the dough
rate of tiller development, different heads on a plant stage itself there is agradual change from asoft to a
may flower at the same time, or sequentially. hard dough consistency as grain filling approaches
completion (Fig. 8).
Stage 7: Milk stage
Stage 9 : Physiological maturity
Within 6-7 days after fertilization the grains grow
sufficiently to become visible within the floret. At Physiological maturity is marked by the formation
this stage they consist of the seed coat filled with of a small black layer in the hilar region of the seed
first a watery and later a milky liquid. This marks the (Fig. 9). The formation of this layer coincides with
beginning of the period of rapid starch deposition in the cessation of movement of materials into the
the endosperm cells, and the period of rapid grain, and hence with the cessation oT grain growth.
increase in the dry weight of the grains (Fig. 8). Black layer formation begins from the upper part of
the panicle (as does stigma emergence) and
proceeds down the panicle. By this stage the grain
has achieved its maximum dry weight, has partly
Figure 8. Growth stages 7 and 8: Milk stage (left) and dried, and the endosperm is becoming hard.
dough stage (right).
Growth and Development of
Individual Organs
The previous section described the development of
the pearl millet plant as a whole. This section
describes the growth and development of individual
plant parts (rcots, leaves, etc). The data are taken
from studies on Mil Zongo and BK-560, a now
hybrid released by the All India Coordinated Millet
Improvement Program. (BK-560 was substituted for

Figure 9. Growth stage 9 : Physiological maturity. The

small black layer is visible in the hilar region at
the base of each grain.
HB-3 in these studies because of the disease forming the main support of the plant during the
susceptibility of the latter. BK-560 is of equivalent seedling stage. The seminal root system generally
maturity and has a similar plant type to HB-3.) remains active up to 45-60 days, after which it
decays.
Root development The adventitious roots begin to appear 6-7 days
after seedling emergence in the nodal region at the
The root system in pearl mnillet has three base of the seedling (Fig. 10). They can be
components: (1) the seminal root, or the primary distinguished from the seminal root by their
root, derived directly from the radicle; (2) the generally greater diameter, and by '.heir point of
adventitious roots, which develop from nodes atthe initiation at the base of the stem. They vary in
base of the stem; and (3) the crown (or collar) roots, number from three to five per shoot. They rapidly
which originate from several lower nodes of the develop a very extensive system of secondary and
stem at or above the soil surface. tertiary branches and are the main pathway for
The seminal root system can be easily identified supplying water and nutrients to the plant during
because its main axis (the radicle) is thinner than most of its life.
the adventitious roots and it produces an extensive The crown (or collar) roots develop in the lower
systemoffinelaterals.Theselateralsdevelopwithin nodes of the stem near the soil surface
3-4 days after the radicle emerges from the seed and approximately 30 days after the emergence of the
are visible as fine branches along the length of the seedling (Fig. 11). They are considered to serve
axis (Fig.10). They elongate and develop rapidly, ma;nly as support for the stem, but have been

Figure 10. Root development 1 : The seminal and Figure 11. Root development 2: The crown, seminaland
adventitious roots at approximately 15 days adventitious roots at approximately 30 days
after emergence. after emergence.

II
ADVENTITIOUUS ROOT-*

SEMINAL ROOT*

observed to develop many lateral branches by the

time of flowering. These appear to be active in the
uptake of water and nutrients. The development of
the plant's root systems 60 days after emergence is
shown in Figure 12.

COLLAR ROOT-00
Tiller development
The tiller initials or buds develop in the axils of the
lower leaves and are initially enclosed by the leaf
sheath. The first tiller leaf appears about 12 days
after emergence in the axil of the coleoptile.
Subsequent tillers develop on alternate sides of the
main shoot following the alternate arrangement of
leaves on the shoot (Fig. 13). The development and
growth of the tillers follows a pattern identical with ADVENTITIOUS ROOT+
that of the main shoot. The tiller development may
be either nearly synchronous with the development
of the main shoot or may be considerably delayed,
or even suppressed by the main snoot. The number
of tillers reaching flowering is a function of both the
variety and the environmental conditions,
particularly the spacing between the plants.
Some varieties produce tillers (called nodal
tillers) from the upper nodes of the main stem after
grain set in the main panicle. These have a short
developmental cycle, producing only a few leaves
and usually a small panicle. Nodal tillers are
common when grain set on the main panicle is poor Figure 12. Root development 3 The crown and
or the main panicle is damaged in some way. adventitious roots at approximately 60 days
Tillers in both BK-560 and Mil Zongo started to after emergence.

2T IT

, I . .

!I /

Ai
K.'
, , -..

Figure 13. The sequential development of tillers 1T= first tiller; 2T= second tiller; etc.
 emerge 10 days after the emergence of the Following flowering there is a steady decline in
coleoptile and the maximum tiller number was leaf area as the older leaves begin to senesce. By the
reached by 25-30 days (Fig. 14). Of the six tillers time of physiological maturity there generally
produced by BK-560, three to four formed heads; remain on!y three to four green leaves per shoot.
but only one or two of the five tillers produced by
Mil Zongo formed heads. Stem elongation

Leaf area development Elongation of the stem internodes begins shortly

after panicle initiation, starting with the short basal
The five embryonic leaves (those which were within internodes, followed by the longer upper
the embryo itself at the time of germination) emerge internodes, and finally the peduncle. This produces
at the rate of approximately one leaf per day. Rates
of emergence of the subsequent leaves (which are
initiated after germination) decline slowly, reaching 2500- FL
a seasonal average rate of 0.40-0.45 leaves per day , 4'
by the time of final leaf emergence. The rate of E
a2000-
emergence and the final number of leaves varies
somewhat among varieties, with earlier varieties M

generally having fewer leaves and a faster rate of Pi

lant
leaf emergence.
The rate of development of the total leaf area per 1000­
plant is a product of the rate of leaf expansion and
the size and longevity of the individual leaves for 500
both the main shoot and the tillers. Rate of leaf area M
development is slow early in the season (Fig. 15a ssh,,

and b) because of the small size of the embryonic 20 40 60 80

leaves, but increases rapidly approximately 15-20 Days from emergence
days after emergence. as the size of individual Figure 15a. Time course of leaf area development for the
leaves increases and as the tillers begin to expand main shoot and the whole plant in BK-560. PI
their leaves. Maximum leaf area is attained at indicates panicle initiation and FL 50% stigma
approximately 50% flowering, by which time the emergence.
majority of the tillers have expanded all their leaves.
The contribution of the tillers to the total leaf area
varies among varieties, depending on the number of 3500- FL
tillers that complete development. F,
3000­
6~
­
.- 2500 P\

5 // PI
5- Whole
it a,0 plant
c P1500

=2- o.to
a-- /

/ 500" Main
shoot
/P
, ", ,,2'0 40 6'0 80
5 10 15 20 25 30 35 Days from emergence
Days from emergence Figure 15b. Time course of leaf area development in the
Figure 14. Time course of tiller emergence in BK-560 main shoot and the whole plant in Mil Zongo.
(broken line) and Mil Zongo (solid line). PI PI indicates panicle intiation and FL 50%
indicates the time of panicle initiation. stigma emergence.

10
 the common sigmoid pattern of stem elongation, The floret primordia are then formed by a
with the max;mum rate of elongation occurring division of the spikelet primordia, which begins
around flag leaf stage (Fig. 16). There may be a about 6 days after panicle initiation (Fig. 17). This Is
further increase in stem length following flowering
in some varieties due to continued elongation of the
peduncle. Taller and/or later varieties undergo a Table 2. Panicle development stages in pearl millet
longer period of stem elongation, due to either variety HB-3 (from the All India Coordinated
longer internodes or a larger number of internodes Millet Improvement Proiectl.
or both, and reach a greater height, but the general Time interval
pattern and rate of elongation are similar. Develop- Developmental (days after
menta I process emergence of
Panicle development stages seedlings)

The process of panicle development consists of a 1. Initiation of panicle meristem 22

of developmental processes, one foreach 2. Initiation of branch primordia at the 23-24
sequence obase of panicle meristem
of various panicle structures, which proceed from 3. Completion of branch primordia initiation 25
the base of the apex to the tip in regularsuccession. (at the apex of the panicle meristem)
These processes are listed with their approximate 4. Formation of floret primordia at the 28
time intervals, for HB-3, in Table 2. base of the panicle
5. Initiation of glumes at the base of 28
The change from the vegetative to the the panicle
reproductive apex is marked by the formation of a 6. Initiation of the lemma and palea at 30
constriction at the base of the apex (Fig. 4). Branch the base of the panicle
primordia initiation begins at the base of the panicle 7. Stamen initiation at the base of 30
within 1-2 days after floral initiation and proceeds the panicle

8. Completion of stamen development (at 31

rapidly up the panicle, reaching the apex in about 3 the tip of the panicle)
days. Each branch primordium rapidly subdivides 9. All floral parts developed 31-32
to form two spikelet and several bristle primordia.
This second stage of differentiation follows an
acropetal pattern similar to the branch primordia
differentiation.

300­

250-
FIL
200- 2

._9 150­
, FL
o0. 10oo / LM

50- PI / / LP
5 1I P1
I' ? q-PL
20 40 60 80
Days from emergence Figure 17. Photomicrograph of c developing panicle,
Figure 16. Time course of stem elongation in BK-560 approximately 8 days after panicle initiation.
(broken line) and Mil Zongo (solid line). PI SP1, SP2 =Paired spikelets; GL =glume;
indicates panicle initiation and FL 50% stigma LM =lemma; FLP = floret primordium; and
emergence. PL = palea.

11
followed by the development of the glumes and the Grain growth
elongation of the spikelet stalks. Initiation of the
floral parts (lemma, palea, stamens, and stigmas) The increase in dry weight in the grains in an
begins at the base of the panicle about 8 days after individual panicle follows a normal sigmoid pattern.
floral initiation and is completed at the apex of the There is an initial lag period of 5-6 days during
panicle by about 10 days after initiation. In most which there is an active division of cells in the
varieties there are two florets per spikelet-one endosperm,but duringj which there is little increase
perfect (containing both anthers and stigmas) and in dry weight in the grain (Fig. 19). Following this
one male (containing only anthers). The distinction there is a longer period of rapid accumulation of dry
between these is evident as soon as the floret weight until black layer development, after which
primordia become recognizable. The difference in there is no further increase in grain weight. The
flowering of these is illustrated in Figure 18. most rapid rate of dry weight increase in the grain
thus occurs during the milk to dough stages.
There is some variation in both grain size and the
time to physiological maturity among grains in
different locations in the panicle. Typically both are
greater at the base of the panicle than in the center
or the apex, and frequently greater at the center
Anther than at the apex. There is also considerable
4-- emergence variation in grain size among varieties, from as little
as 3-4 g per 1000 grains to as high as 10-12 g.
Mature grains vary somewhat in shape, being
generally roundish at the upper, exposed end and
narrow at the hilar region. Colors vary from white to
gray to brownish. The black layer is located just
above the hilar region on the basal abgerminal side
3 and the embryo is opposite (Fig. 20).

Stigma
emergence

8­

7­

.34­

2 ­
,,/ 3:I

Perec flrtMl foe ' '5 A 2 0 3

122 Perfect~~~~ay ffore 20 flalowertr015

25 30 3

Figure 18. Flowering sequence in the perfect (left) and Figure 19. Time sequence of grain growth in BK-560
male (right) florots. (broken line) and Mil Zongo. (solid line).

12
 filling, if the need for carbohydrates for filling the
-Grain grain is either less than cr greater than the available
supply from photosynthesis in the leaves. Under
such conditions, the stem serves as either astore for
excess carbohydrate or a source of supply of
carbohydrate for grain filling.
There is relatively little dry-matter distribution to
-- Glume the panicle during 3S, as the panicle does not
Black layer undergo rapid growth until the end of this phase
(Fig. 21). During G33 , however, the largest
_ Hilar region dry-matter increase in the plant is in the panicle, as

Grain: abgerminal side

120-
Spikelet
FL

100- Panicle

Endospe-m t"4
0-
Po Stem
._9 o
00-
Em ry
Embryo Embryo-2

1W 20-

Grain: germinal side Grain: lateral section Leaf

Figure 20. Spikelet and grain of Mi Zongo. 20 40 60 80

Days from emergence

Figure 21a. Time course of dry matter accumulation in the

leaf, stem, and panicle of Mil Zongo. P/
Distribution of plant dry weight indicates panicle initiation and FL 50% stigma

frates of dry-matter accumulation during GS 1 are emergence.

slow (Fig. 21) becausa of the small leaf area of the
plant (Fig. 15). The majority of dry matter produced 120­
during this phase goes into the leaves and roots.
Leaf growth continues during the GS2 phase as first - 100-
the main shoot and then the tillers expand their C
remaining leaves. By the time of flowering, however, C8

leaf growth in the majority of the tillers as well as in .9 Panic le

the main shoot is completed, and leaf dry weight . -
remains constant (or decreases slightly as lower 26
leaves senesce) fcr the remainder of the life of the ? e
40- Stem
plant.
Stem elongation begins shortly after floral PI
initiation and an increasing fraction of the total plant 20-
dry matter goes to the stem during the GS2 phase Leaf
(Fig. 21). By the time of flowering in the main shoot, ,0 4 ,
approximately two-thirds of the plant dry weight is 20 4o 60 80
in the stem. Stem dry weight of the whole plant may Days from emergence
continue to increase following flowering on the Figure21b. Time course of dry matter accumulation in the
main culm as many of the tillers are still undergoing leaf,stem, and panicle of BK-560. Pl indicates
rapid stem elongation. There mayalso besomegain panicle initiation and FL 50% stigma
or loss in the dry weight of the stem during grain temergence.

13
 the tillers reach flowering and the grains fill in all RACHIE, K.O., and MAJMUDAR, J.V. 1981. Pearl
panicles. millet. State College: Pennsylvania State Univer­
The final distribution of dry matter in leaf, stem, sity Press.
and panicle varies rather widely in pearl millet,
depending upon the variety. In dwarf, high-yielding
varieties as much as 50% of the dry matter may be in Growth and yield
the panicles, with the remainder divided between
the stems and the leaves in an approximately 3 : 1 BEGG, J.E. 1965. The growth and development of
ratio. In some tall varieties, however, 50% or more of a crop of bulrush millet (Pennisetum typhoides
the dry weight may be in the stems, with no more S. and H.). ,ournal of Agricultural Sciences
than 20-30% in the panicles. Total dry-matter 65:341-349.
production in these types, however, may be EGHAREVBA, P.N. 1977. Tiller number and millet
considerably greater than in the dwarf varieties and grain productivity. Cereal Research
actual dry wfeight in the panicles may be similar to Communication 5:235-247.
that in the dwarf varieties. GREGORY, P.J., and SQUIRE, G.R. 1979.
Irrigation effects on roots and shoots of pearl
millet (Pennisetum typhoides). Experimental
Agriculture 15:161-168.
Acknowledgements GREGORY, P.J. 1979. Uptake of N, P and K by
irrigated and unirrigated pearl millet (Pennise­
The authors are grateful to Drs. G. Alagarswamy tum typhoides). Experimental Agriculture
and L.K. Fussell for reviewing the manuscript, to 15:217-223.
Messrs. G. Anjaiah and P.S. Raju for assistance in KASSAM, A.H., and KOWAL, J.M. 1975. Water use,
the field observations, and to Mr. V. Ramesh and the energy balance and growth of Gero millet at
staff of ICRISAT Information Services for drawings Samaru, Northern Nigeria. Agricultural
and photographs. Meteorology 15:333-342.

Botany
BIBLIOGRAPHY
FUSSELL, L.K., and PEARSON, C.J. 1978. Course
The following is a selected list of references on of grain development and its relationship to
various aspects of the growth and development of black layer appearance in Pennisetum
pearl millet. They will provide the interested reader americanum. Field Crops Research 1:21-32.
with more detailed information on the subjects they MAITI, R.K., and BISEN, S.S. 1979. Pearl millet
cover. anatomy. Information Bulletin No. 6. Patancheru,
A.P., India: International Crops Research Institute
for the Semi-Arid Tropics. 24 pp.
Reviews POWERS, D., KANEMASU, E.T., PIARA SINGH,
and KEITNER, G. 1980. Floral development of
FERRARIS, R. 1973. Pearl millet (Pennisetum pearl millet (Pennisetum americanum). Field
typhoides). Commonwealth Bureau of Pasture Crops Research 3:245-265.
and Field Crops, Review Series No. 1. RAYMOND, C. 1968. Pour une meilleure
KRISHNASWAMY, N. 1962. Bajra (Pennisetum connaisance de la crossance et du d6veloppe­
typhoides S. & H.). Indian Council of Agricultural ment de mils Pennisetum. Agronomie Tropicale
Research, Cereal Crop series No. 2. 23:844-863.

14