PEARL MILLET BIOLOGY

Project Coordinator, AICRP- Pearl millet (ICAR), ARS, Mandor, Jodhpur

Origin of pearl millet

Pearl millet originated in tropical Western Africa some 4000 years ago. It is a member of
the grass family, originally a wild plant in Africa where largest members of both wild and
cultivated forms of this species occur. From there, it differentiated into two races;
globosum race that moved to the western side and the typhoides race that reached
Eastern Africa and spread to India and southern Africa some 2000–3000 years ago. The
evolution of pearl millet under the pressures of drought and high temperatures imparted
the ability to tolerate drought, nutrient deprived soil and high temperatures of Indian
and African hot deserts more effectively than other cereals like wheat and rice.

Taxonomy
Pennisetum is largest genera in the tribe Paniceae with five sections and approximately
140 species that are widely distributed in the tropics and subtropics (Clayton 1972). It is
placed close to the genus Cenchrus (Stapf and Hubbards, 1934 and Bor, 1960) on the
basis of spikelets that are arranged in groups surrounded by involucre. Inter-generic
hybrids between pearl millet and Cenchrus ciliaris L have also been reported (Read and
Bashaw, 1974). According to Brunken (1977), Pennisetum includes two reproductively
isolated species viz. P purpurium Schumach, a tetrapolid (2n = 28) perennial species
which occurs throughout the wet tropics of the world; and P. americanum (syn. P.
glaucum) (L) Leeke, a diploid annual species, native to the semi-arid tropics of Africa
and India. P. glaucum (P americanum according to Brunken 1977) contains three sub
species:
a) Sub-sp glaucum- cultivated, involucres persistent at maturity, distinctly stalked
b) Plants wild or weedy, involucres readily deciduous at maturity, short stalked or
sub-sessile.
i. Subsp, violaceum (monodii) - Involucres sub-sessile, stalks less than 0.25
mm long; mature grains 0.6-1.0 mm deep. Considered to be wild progenitor
of cultivated pearl millets.
ii. Subsp, stenostachyum – Involucre short stalked; stalk more than 0.25 mm
long, mature grain 1.0-2.0 mm deep.

Pennisetum glaucum ssp glaucum is the only cultivated species for seed purpose, P.
purpurium and its hybrids with bajra in both directions (bajra x napier and napier x
bajra) are cultivated for fodder purposes in some areas of world and India.

The taxonomical classification presently adopted for pearl millet is based on Clayton
1972. However, de Wet (1977) accepted Pennisetum glaucum for annual pearl millet
species instead of P. americanum suggested by Brunken (1977), present classification
accepted is –

Family : Poaceae
Subfamily : Panicoideae
Tribe : Paniceae
Subtribe : Panicinae
Section : Panicillaria
Genus : Pennisetum
Species : Glaucum

Biology
Pearl millet (Pennisetum glaucum L.) being a C4 species is endowed with a very high
photosynthetic efficiency and ability for dry matter production. It is an important coarse

1
grain cereal and forage crop of the arid and semi-arid tropics of the Indian subcontinent
and several African regions. Pearl millet grows on a wide variety of soils including sandy,
light textured soils, and may sustain growth on acidic and very infertile soils. At the
same time, wide variations displayed by pearl millet for vegetative, reproductive and
physiological features are advantageous in development of cultivars suiting to various
climates and cropping systems. A sound knowledge of pearl millet biology is essential to
realize the real potential of this crop for varied agro-ecological conditions and to enhance
the breeder's ability to develop requirement specific cultivars (Begg, 1965; Khairwal et
al. 1990).

Pearl millet biology includes all growth and developmental features from germination to
seed formation (Begg, 1965, Maiti and Bisen, 1976, Gregory and Squire, 1979, Maiti,
1979, Fussel et. al., 1980, Powers et. al., 1980, Maiti and Bidinger, 1981, Vanangamudi,
1987, Khairwal et. al., 1990, and Mangat et. al., 1999). These features include three well
defined growth phases i.e. vegetative, reproductive and grain filling which shall be
discussed in relation to crop improvement (Fig. 1, Table 1).

Fig. 1: Major growth and morphologically distinct stages of pearl millet

2
Table 1: Three major growth phases and morphologically distinct development
stages of pearl millet plant1

Growth Identifying characteristics Approximate days

Phases after emergence*
GP I Vegetative phase 0-21
Emergence stage 2-3
Three leaf stage 3-7
Five leaf stage 7-14
Panicle initiation stage 14-21
GP II Reproductive/Panicle development phase 21-42
Flag leaf stage 21-28
Boot stage 28-35
Half bloom stage 35-42
GP III Grain filling phase 42-77
Milk stage 42-49
Dough stage 49-56
Black layer formation or physiological maturity 56-63
1
Main shot
* May vary with environmental conditions, locations and varieties

Vegetative Phase
Upon germination radicle starts emerging from the hilar region approximately within 16
hours of the initiation of the germination followed by development of plumule and
coleoptile sheath approximately two hours later. Radicle grows downwards rapidly and
produces fine root hairs. Coleoptile grows upwards slowly through soil until it emerges
from the soil surface. Emergence of coleoptile from soil surface depends upon the depth
of sowing, soil texture, moisture and temperature. It takes 2-3 days under favorable
conditions (Photo 1-2).

1. Emergence 2. Three leaf, five leaf

Photo 1-2. Vegetative Phase

Pearl millet has a typical monocotyledonous type of root system consisting of seminal or
primary root, adventitious roots, and crown or collar roots (Table 2, Photo 3-5).

3
Table 2: Types of root, initiation and effectiveness

Types Initiation time Effective up to

Primary within 4 days 45-60 days
Adventious 8-12 days Maturity
Crown or collar 30-40 days Maturity

3. Primary 4. Adventious 5. Crown or collar

Photo 3-5. Types of root, initiation and effectiveness

Pearl millet root can penetrate up to 180 cm and attain a total length of 1500 m per m2
of the cultivated area and touch 3000 m per m2 at the harvest with a mass of 35 g per
m2. In sandy soils, a mature pearl millet plant can spread its roots to about 3 m to
absorb water. In heavily tillering plants, roots have more horizontal spread than deep
penetration. It has been observed that genotypes tolerant to early season drought (3-15
days after sowing) have 30 % more root length than the susceptible one.

Pearl millet is an erect annual with a good tillering habit (Fig. 2). The main stem attains
a height of 2 - 4 m with 0.5 - 1.5 cm diameter. It is generally round to oval in shape.
Nodes are slightly swollen, may be pubescent with ring of adventitious root primordia at
the basal end and its inter-nodes are glabrous. The internodal length increases from the
base of the stem upwards. Single leaf appears on each node in alternate orientation. An
axillary bud is borne on the node at the base of the groove, which may remain dormant
or develop in to a nodal tiller. Soft and thin palatable stem having very low fiber content
is desirable for forage. Dry matter accumulation is increased in stem in the long duration
pearl millet which limit the increase in grain yield in such types. Genotypic variation
exists for stem growth, thickness, internode length and colour (Photo 6-8).

4
 Fig. 2. The pearl millet plant

6 7 8
Photo 6-8. Genotypic variation for stem colour

The tiller leaf appears ~ 12 days after seedling emergence from the base of the main
stem on both sides, following the alternate arrangement of leaves (Fig. 3). Pearl millet
has the potential to produce many effective tillers that may increase under wide spacing.
Synchrony in flowering coupled with a higher number of effective tillers enhances the
probability of producing more seeds from the same plant.

Fig. 3: Development of tillers in pearl millet

5
Leaves are linear, 20-100 cm long and 0.5-5.0 cm wide, usually sparse to densely hairy
or glabrous. The leaf consists of a leaf sheath and lamina or blade (Fig. 4).

Fig 4. Pearl millet leaf

The midrib may or may not be prominent. The leaf margins bear small saw teeth (see
photo 9–13). Stomata are present on both leaf surfaces in equal numbers, varying
approximately 45-96 stomata per mm2. Early maturing genotypes have less number of
leaves with faster growth and emergence that helps to escape early drought. The Kranz
anatomy, a characteristic of C4 pathway in pearl millet makes it adaptive to drought and
palatable for grazing cattle.

9 10 11 12 13
Photo 9-13. Midrib and leaf margins

Reproductive Phase
A change from vegetative to reproductive phase is marked by the formation of an apical
dome - like structure and a constriction at the base of the shoot meristem (Fig. 5)
leading to a change from leaf primordia to spikelet primordia, development of spikelets,
florets, glumes, stigmas and anthers. The period between panicle initiation to anthesis is
critical in determining the grain number.

Fig. 5. Panicle initiation with primordia in pearl millet

Panicle emerges in ~35-70 days from the day of sowing depending upon the early or late
genotypes. Panicle emergence is marked by the rapid elongation of the peduncle and the
inter-node below it, and by the appearance of the flag leaf or boot leaf (Photo 14).
Panicle takes nearly six days to emerge from the leaf sheath (Photo 15), with the

6
maximum emergence on the fourth or fifth day (Photo 16). Hybrids generally take less
time for panicle emergence than their parents indicating heterosis for the early panicle
emergence.

14 15 16
Photo 14-16. Panicle emergence stages

Pearl millet inflorescence is a compound terminal spike called panicle and its length
generally varies between 20-25 cm with a circumference of 7-9 cm (see photo 16). The
panicle shape varies considerably as shown in Fig. 6, but, the common shapes are either
cylindrical or conical.

1. Cylindrical 2. Conical 3. Spindle 4. Candle 5. Lanceolate.

6. Dumb-bell 7. Club 8. Oblanceolate 9. Globose

Fig. 6: Panicle shapes

7
17. Conical 18. Spindle 19. Cylindrical 20. Candle
Photo 17-20. Panicle shape

Inflorescence consists of a central rachis covered with soft/short hairs and bears
fascicles on rachillae. The density of fascicles and the length of rachillae determine the
degree of compaction of the panicle. Each fascicle contains spikelets surrounded by a
wall of bristles (i.e., involucre) (Fig. 7). The prolongation of the fascicle axis determines
the length of bristles (Photo 21).

Fig .7: Floral parts of pearl millet Fig. 8: Flowering sequence in perfect and
male floret

8
 • Fascicles
• Spikelets 800-3000 (1600)
• One spikelet – 2 glumes

Photo 21. Bristles in the panicle

A spikelet may contain 2-4 flowers or florets, but generally two. The lower floret is
staminate and the upper floret is bisexual or hermaphrodite (Fig. 7, Photo 22-23, Fig. 8).

22. Staminate 23. Hermaphrodite

Photo 22-23. Staminate and hermaphrodite flowers

Pearl millet is a protogynous species. The styles start protruding two to three days after
the emergence of the panicle. The stylar branches protrude first from the florets in the
upper middle region of the panicle and then proceed both up and downwards. In the
hermaphrodite flowers, the stigmas emerge earlier than the anthers and hence stigmas
receive pollen from inflorescence of other plants. The time required for complete stigma
emergence varies from 2 to 3 days and they remain receptive for next two to three days
(Photo 24-30).

9
 24 25 26 27
Stigma emergence Fresh stigma

28 Withered stigma 29. Protogynous flowering 30. Full protogyne

Photo 24-30. Stigma emergence and receptiveness

By the time anther emergence commences, the stigma would have emerged and also
pollinated, avoiding selfing. The emergence of the first anther usually begins about three
to four days after the first stigma has emerged. The anther emergence occurs in two
phases. The first phase involves solely the hermaphrodite flowers, and the second phase
includes the staminate flowers. When the first phase of the emergence of anthers has
reached the basal spikelets, the second phase begins when the staminate flowers are
functional from the upper part of the panicle (Photo 31-34). A panicle continues
shedding pollen for about 3 days. The anther emergence continues throughout the day
and night. The anthesis occurs between 8 am and 2 pm with a peak at about 10 am. The
increase in humidity and a decrease in temperature have been noted to reduce anther
emergence, while lowering of humidity and a rise in temperature enhances up the
anthesis.

10
 31. Anthers 32. Dehiscence 33. After 34.Complete
emergence pollination pollination
Photo 31-34. Anther emergence and pollination

Pearl millet is a highly cross-pollinated species. Wind is supposed to be the major cross
pollinating agent. However, insects also effect cross pollination. Protogyny and the time
lag between stigma emergence and anther dehiscence favor cross pollination, but
asynchronous flowering prevents its full realization. The protogyny in pearl millet is
exploited for controlled cross pollination without resorting to emasculation.

Selfing and Crossing Techniques

Controlled pollination without resorting to emasculation is a common phenomenon in
pearl millet. Usually, there are 900-3000 spikelets per panicle, averaging 1600 spikelets.
Each spikelet has two florets, one bisexual and the other, staminate. The bisexual or
hermaphrodite floret consists of a pistil and three anthers while the staminate floret has
three anthers. Looking to tender, large number and small structure of florets, it is
difficult to follow emasculation in pearl millet. Protogynous nature is exploited for
effecting cross-pollination; protogyny, is a unique feature in which the stigmas emerge
and become active earlier than anthers or pollen dehiscence and receive pollens from
other plants. This sequence of flowering practically excludes self pollination in the same
inflorescence.

The inflorescence to be used as a female or male is covered with the glassine paper bag
before any stigma is visible. Generally, the safest stage is when about one third of the
inflorescence is out of the flag leaf sheath. When all stigmas have emerged, the panicle
can be considered ready for cross pollination. If selfed seed of the male parent is not
required, fresh pollen from dehiscing anthers, visible as yellow powder in the transparent
selfing bags can be collected by tapping even in the inflorescences in which stigmas have
completely emerged. The pollination is carried out by quickly removing the bag from the
female inflorescence, dusting the pollen collected from the male infloresence, and then
rebagging the pollinated inflorescence (see photo 35-36).

11
 35 36
Photo 35-36. Selfing and crossing techniques

Grain filling stage:

Pearl millet seed is a caryopsis (Fig. 9 photo 37-42) and its shape is highly variable,
ranging from globular to conical shape (Fig. 10). The seed colour varies from ivory to
purplish black, with light to deep gray being the most common seed colour. A small
embryo is present on the depressed or flat surface at the tapering end of the seed. The
size of the grain depends on its position in the panicle, being largest at the base,
medium in the middle, and smallest at the apex. Variations exist in grain size among
varieties generally ranging from 4-12 g per 1000 grains.

Fig. 9: Schematic cross section of a pearl millet caryopsis

37 Developing seed 38 Developing seed 39 Developing seed

12
 40 Developing seed 41 Dough stage 42. Ready to harvest
Photo 37-42. Grain filling stages

Obovate Elliptical

Hexagonal Globular
Fig. 10: Seed shape in pearl millet

Seed viability and seedling vigor are dependent upon the extent of grain development.
Selection of varieties with higher percentage of seed filling and larger seed size may lead
to higher yields. The temperature at which the seed developed did not affect seed
viability, but it did affect the vigour. Post-harvest dormancy has been reported for at
least 14 days in pearl millet (Burton, 1969; Khairwal, 1980; Khairwal et al., 1980).
However, pearl millet grains germinated in the inflorescence when harvesting was
delayed in prolonged wet weather.

13
References:

Begg, J.E. 1965. The growth and development of a crop of bulrush millet (Pennisetum
typohoides S&H). Indian Journal of Agricultural Sciences 65: 341-349.
Bor, N.L. 1960. The Grasses of Burma, Ceylon, India and Pakistan. Pergamon Press,
Oxford, London, U.K.
Brunken, J.N. 1977. A systematic study of Pennisetum Sect. Pennisetum (Gramineae).
American Journal of Botany 64: 161-176.
Burton, G.W. 1969. Breaking dormancy seeds of pearl millet, Pennisetum typhoides.
Crop Science 9: 659-664.
Clayton, W.D. 1972. Gramineae. In: Flora of West Tropical Africa. (F.N. Hepper, ed.).
Crown Agents, London, U.K. Pages 170-465.
de Wet, J.M.J. 1977. Domestication of African cereals. African Economic History 3: 15-
32.
Fussell, L.K., Pearson, C.J., Norman, M.J.T. 1980. Effect of temperature during various
growth stages on grain development and yield of Pennisetum americana. Journal
of Experimental Botany. 31: 621-633.
Gregory, P.J. and Squire, G.R.1979. Irrigation effect on roots and shoots of pearl millet
(Penisetum typhoides). Experimental Agriculture. 15: 161-168.
Khairwal, I.S. 1980. Seed dormancy in pearl millet. Food Farming and Agriculture 12:
128-129.
Khairwal, I.S., Kapoor, R.L., Yadav, H.P. 1980. Seed dormancy and its genetic basis in
pearl millet. Seeds and Farms 6: 25-28.
Khairwal, I.S., Ram, C., Chhabra, A.K. 1990. Pearl Millet Seed Production and
Technology. Manohar Publications, New Delhi, India.
Maiti, R.K. 1979. How a pearl millet plant develops. ICRISAT, Hyderabad.
Maiti, R.K. and Bidinger, F.R. 1981. Growth and development of pearl millet plant.
Research Bulletin No. 6. ICRISAT, Hyderabad.
Maiti, R.K. and Bisen, S.S.1976. Studies on growth and development of panicle and
grains in two contrasting genotypes of pearl millet (Pennisetum typhoides Stapf
&Hubbard). First International Symposium on Physiology of Sexual Reproduction
in Flowering Plants. Punjab Agriculture University, Ludhiana, India.
Mangat, B.K., Maiti, R.K., Kairwal, I.S. 1999. Pearl Millet Biology. Pearl Millet Breeding
(eds. I.S. Khairwal, K.N. Rai, D.J. Andrews, G. Harinarayana). Oxford and IBH
Publishing Co. Pvt. Ltd. Pp: 1-27.
Powers, D., Kanemasu, E.T., Singh, P., Kreinter, G. 1980. Floral development of pearl
millet (Pennisetum americanum (L.) K. Schum). Field crops Research 3: 245-266.
Read, J.C. and E.C. Bashaw. 1974. Intergeneric hybrid between pearl millet and
buffelgrass. Crop Science 14: 401-403.
Stapf, O. and C.E. Hubbard. 1934. Pennisetum. The Flora of Tropical Africa. Vol. 9. (D.
Prain, ed.). Crown Agents, London, UK.
Vanangamudi, K. 1987. Studies on seed development and maturation in bajra. Madras
Agriculture Journal. 74: 132-134.

14