Anda di halaman 1dari 122

Training Manual

Development of Cultivars and


Seed Production Techniques
in Sorghum and Pearl Millet

Edited by

Faujdar Singh, K N Rai, Belum V S Reddy, and B Diwakar

ICRISAT
Training and Fellowships Program and Genetic Enhancement Division
International Crops Research Institute tor the Semi-Arid Tropics
Patancheru 502 3 2 4 , A n d h r a P r a d e s h , India
1997
Citation: S i n g h , Faujdar, Rai, K.N., Reddy, B e l u m V.S., a n d Diwakar, B. (eds.). 1997. D e v e l o p m e n t of
cultivars a n d s e e d production techniques in s o r g h u m a n d pearl millet. Training m a n u a l . Training a n d
Fellowships P r o g r a m a n d Genetic E n h a n c e m e n t Division, I C R I S A T Asia Center, India. P a t a n c h e r u 5 0 2
3 2 4 , A n d h r a P r a d e s h , India: International Crops Research Institute for the S e m i - A r i d T r o p i c s . 118 p p .
(Semi-formal publication).

An I C R I S A T semi-formal publication issued for limited distribution without f o r m a l review.

T h e opinions in this publication are those of the authors a n d not necessarily those of I C R I S A T . the
designations e m p l o y e d a n d the presentation of the material in this publication do not imply the e x p r e s s i o n
of any opinion w h a t s o e v e r on the part of I C R I S A T c o n c e r n i n g the legal status of any c o u n t r y , territory,
city, or area, or of its authorities, or concerning the delimitation of its frontiers or b o u n d a r i e s . W h e r e trade
n a m e s are u s e d this does not constitute e n d o r s e m e n t of or discrimination against a n y p r o d u c t by the
Institute.
Contributors

Belum V S Reddy Senior Scientist ( S o r g h u m Breeder), I C R I S A T Asia Center, P a t a n c h e r u 5 0 2 3 2 4 ,


A n d h r a P r a d e s h , India.

B S Talukdar S e n i o r Scientist (Pearl Millet Breeder), I C R I S A T Asia Center P a t a n c h e r u 5 0 2 3 2 4 ,


A n d h r a P r a d e s h , India.

C T Hash Principal Scientist (Pearl Millet Breeder), I C R I S A T A s i a Center, P a t a n c h e r u 5 0 2


3 2 4 , A n d h r a P r a d e s h , India.

Faujdar S i n g h Senior Training Officer, I C R I S A T Asia Center, P at anc heru 5 0 2 3 2 4 , A n d h r a


P r a d e s h , India.

G Harinarayana Director, G a n g a Agri S e e d s Ltd., 140B Babu Khan Estate, Basheerbagh,


H y d e r a b a d 5 0 0 0 0 1 , India.

J W Stenhouse Principal Scientist ( S o r g h u m Breeder), I C R I S A T A s i a Center, P a t a n c h e r u 5 0 2 3 2 4 ,


A n d h r a P r a d e s h , India.

K N Rai Senior Scientist (Pearl Millet Breeder), I C R I S A T Asia Center, P a t a n c h e r u 5 0 2 3 2 4 ,


A n d h r a P r a d e s h , India.

V Jaya Mohan Rao Chief Scientist, A n d h r a P r a d e s h State S e e d s D e v e l o p m e n t C o r p o r a t i o n , H A C A


B h a w a n , H y d e r a b a d 5 0 0 0 0 4 , India.
About this manual

S o r g h u m a n d pearl millet are important coarse-grain cereals in the dry agricultural areas of the arid a n d
semi-arid tropics. In both crops, varieties (open-pollinated a n d highly h e t e r o g e n o u s / h e t e r o z y g o u s types
in pearl millet a n d h o m o z y g o u s / h o m o g e n o u s types in sorghum) a n d hybrids ( b a s e d on cytoplasmic-genic
m a l e sterility) h a v e b e e n p r o d u c e d a n d a d o p t e d by farmers. T h e d e v e l o p m e n t of parental materials and
their use in cultivar d e v e l o p m e n t a n d s e e d production of new varieties requires specific skills. With the
aim of imparting s u c h skills, a training course w a s organized for scientists a n d technicians f r o m Asia a n d
Africa at the International Crops Research Institute for the Semi -Arid Tropics ( I C R I S A T ) , India, from 6
to 17 Mar 1995.

T h e c o u r s e c o v e r e d both theoretical a n d practical aspects of d e v e l o p m e n t a n d m a i n t e n a n c e of


cultivars (hybrids a n d varieties) a n d their parental lines. T h e participants w e r e given o v e r v i e w s of cultivar
d e v e l o p m e n t of both crops. T h e course broadly c o v e r e d topics such as the reproductive biology of both
the crops, selfing a n d crossing techniques, production and m a i n t e n a n c e of male-sterile lines, restorer
parents a n d hybrids, a n d varieties in s o r g h u m a n d pearl millet. Production of f o u n d a t i o n a n d certified
s e e d s , p o s t h a r v e st handling of breeder seeds, a n d s e e d testing procedures w e r e also d i s c u s s e d . A
chronology of events relating to the d e v e l o p m e n t of the s e e d industry in India w a s also included in the
c o u r s e . T h e participants w e r e taken to s e e d p r o c e s s i n g plants near H y d e r a b a d , India.

T h e r e w e r e eight participants (An n e x u r e l) f r om Egypt, K e n y a , Mali, M y a n m a r , N a m i b i a , Niger,


Nigeria, a n d Syria. T h e resource persons for the course w e r e from I C R I S A T Asia Center, a n d public a n d
private sector s e e d - p r o d u c i n g organisations in H y d e r a b a d , India. This m a n u a l is a c o mpilation of lectures
delivered by the resource persons. Although s o m e editing has m inimized a great deal of duplication
across the chapters, s o m e duplication of content has b e en purposely retained to avoid any discontinuity
within the chapter.

It is h o p e d that this m anual will be of use to scientists a n d technicians involved in the


d e v e l o p m e n t of s o r g h u m and pearl millet cultivars a n d their s e e d pr oduction.

Information has b e e n t a k e n from published a n d unpublished reports. Efforts h a v e b e e n m a d e to


indicate t r a d e n a m e s ® w h e r e available. O m i s s i o n or inclusion of any trade n a m e does not constitute
e n d o r s e m e n t or discrimination of product by I C R I S A T or the contributors to this publication.

T h e administrative support received f r o m the I C R I S A T m a n a g e m e n t a n d staff is gratefully


a c k n o w l e d g e d . O u r special thanks to Smitha S i t a r a m a n for editorial assistance a n d to S V P r a s a d Rao
a n d K S Vijaya S e k h a r for c o m p u t e r i z e d processing of the d o c u m e n t .
Contents
Sorghum

An O v e r v i e w of S o r g h u m Cultivar D e v e l o p m e n t J W Stenhouse 1

Reproductive Biology of S o r g h u m Faujdar S i n g h 5

S o r g h u m Plant a n d Flower Parts B e l u m V S Reddy 12

Selfing a n d C r o s s i n g T e c h n i q u e s in S o r g h u m B e l u m V S Reddy 17

D e v e l o p m e n t , P r o d u c t i o n , a n d M a i n t e n a n c e of Male-Sterile Lines
in S o r g h u m B e l u m V S R e d d y 22

D e v e l o p m e n t , P r o d u c t i o n , a n d M a i n t e n a n c e of Restorer Parents a n d
Pure-Line Varieties in S o r g h u m J W S t e n h o u s e 28

Production of S o r g h u m Hybrids B e l u m V S Reddy 31

Pearl Millet

An O v e r v i e w of Pearl Millet Cultivar D e v e l o p m e n t a n d M a i n t e n a n c e C T Hash 34

Reproductive B iology of Pearl Millet Faujdar S i n g h 37

Selfing a n d C r o s s i n g T e c h n i q u e s in Pearl Millet K N Rai 45

D e v e l o p m e n t , P r o d u c t i o n , a n d M a i n t e n a n c e of Male-Sterile Lines in Pearl Millet K N Rai 49

D e v e l o p m e n t , P r o d u c t i o n , a n d M a i n t e n a n c e of Pollinators in Pearl Millet B S Talukdar 61

Production of Pearl Millet Hybrid S e e d B S Talukdar 65

S e e d Production of Pearl Millet O pen- P ollinat e d Cultivars C T Hash 68

Seed-Processing Techniques and Chronology of Indian Seed Industry

P r o d u c t i o n of F o u n d a t i o n a n d Certified S e e d of S o r g h u m a n d Pearl Millet V J a y a M o h a n R a o 79

Harvest a n d Postharvest H a n d l i n g of Breeder S e e d of S o r g h u m a n d Pearl Millet C T Hash 88

Purity T e s t of H y b r i d s , Parental Lines, a n d Open-Pollinated Varieties of S o r g h u m


a n d Pearl Millet V J a y a M o h a n R a o 94
D e v e l o p m e n t of S o r g h u m a n d Pearl Millet S e e d Industry in India G Harinarayana 102

References 108

Appendix l List of participants 112


Sorghum

An Overview of S o r g h u m Cultivar Development


J W Stenhouse

S o r g h u m ( S o r g h u m bicolor (L.) M o e n c h ) is the fifth m o s t important cereal crop in the w o r l d after w h e a t ,


rice, m a i z e , a n d barley. It is cultivated annually over approximately 45 million h a , p r o d u c i n g a p p r o x imately
60 million t of g r a i n . S o r g h u m grain is a major f o o d in m u c h of Africa, S o u t h A s i a , a n d Central A m e r i c a ,
a n d an important a n i m a l f e e d in the U S A , Australia, a n d S o u t h A m e r i c a . In addition to t h e s e uses of the
grain, s o r g h u m crop residues a n d green plants also provide sources of a n i m a l f e e d , building materials,
a n d f u e l , particularly in the semi-arid tropics (SAT).

S o r g h u m is a d a p t e d to tropical a n d subtropical climates but the greater part of the area of the
crop falls in d r o u g h t - p r o n e , semi-arid tropical regions of the w o r l d . In these a r e a s , it is usually g r o w n with
limited inputs in conditions of sparse rainfall a n d low soil fertility, a n d e x p o s e d to a range of d i s e a s e s a n d
pests. As a result, the yields are poor.

In fact, the yield potential of s o r g h u m is quite high; grain yields c a n e x c e e d 10 t ha - 1 under


favorable conditions. However, a majority of the subsistence farmers w h o typically cultivate this crop are
unable to take a d v a n t a g e of this potential b e c a u s e they have limited options for improving their
m a n a g e m e n t practices. H e n c e , i m p r o v e m e n t s in s o r g h u m production in the S A T are m o r e readily
a c h i e v e d t h r o u g h s e e d - b a s e d tec hnologies such as cultivars with i m p r o v e d tolerance to d r o u g h t a n d low
soil fertility, a n d resistance to pests a n d diseases.

Breeding Behavior of Sorghum and Types of Cultivars

T h e b r e e d i n g b e h a v i o r of a crop influences the m e t h o d s u s e d by plant b r e e d e r s to p r o d u c e i m p r o v e d


cultivars a n d the types of cultivars that can be d e v e l o p e d .

S o r g h u m is a predominantly self-pollinating crop w h i c h s h o w s little inbreeding d e p r e s s i o n .


H o w e v e r , significant levels of natural outcrossing also occur. T h e level of o u t c r o s s i n g varies accor ding
to the panicle type of t h e cultivar. It can be as m u c h as 3 0 - 6 0 % in loose-panicled grassy s o r g h u m , but
a more typical figure w o u l d be < 1 0 % in c o m p a c t - p a n i c l e d types ( H o u s e 1985). As a result of this
c o m b i n a t i o n of self-pollination a n d outcrossing, most of the landraces of s o r g h u m that are normally g r o w n
by s u b s i s t e n c e f armers are mixtures of inbred a n d partially inbred lines. I m p r o v e m e n t c a n be a c h i e v e d
by purification of the more productive lines in the mixture by selfing to develop pure-line varieties.
H o w e v e r , b e c a u s e of the t e n d e n c y for further outcrossing to occur, these pure-line varieties require
deliberate m a i n t e n a n c e by selfing to k e ep t h e m pure.

S e v e r a l m a l e sterility s y s t e m s are f o u n d in s o r g h u m w h i c h can be u s e d by plant breeders to


w i d e n the range of possibilities for improving the crop a n d to d e v e l o p different cultivar t y p e s . G enetic
m a l e sterility, w h i c h is controlled by recessive g e n e s at any one of several loci (Doggett 1988), can be
u s e d t o p r o m o t e o u t c r o s s i n g a n d g e n e r a t e r a n d o m - m a t i n g populations for i m p r o v e m e n t by various
recurrent selection m e t h o d s . In particular, male sterility c a u s e d by ms3, ms7, a n d a / g e n e s has b e en u s e d
for this p u r p o s e (Murty et al. 1994). T h e s e r a n d o m - m a t i n g populations of s o r g h u m , unlike other crops,
are rarely suitable for direct use by the farmer b e c a u s e of their r a g g e d a p p e a r a n c e ( H o u s e 1985).

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S e v e r a l c y t o p l a s m i c genetic male sterility (CMS) systems occur in s o r g h u m (Schertz a n d Pring
1982). T h e s e c a n be u s e d to capture heterosis in hybrid cultivars by crossing t w o inbred lines, o n e male-
sterile a n d t h e other male-fertile. T h e Milo-Kafir C M S s y s t e m d i s c o v e r e d by S t e p h e n a n d H o l l a n d (1954),
also k n o w n as the A, C M S s y s t e m , is the most c o m m o n l y u s e d s y s t e m for c o m m e r c i a l pr oduction of
hybrids. Like t h e pure-line varieties m e n t i o n e d a b o v e , the inbred line parents of hybrids require deliberate
a n d s y s t e m a t ic m a i n t e n a n ce t o keep t h e m pure, particularly the C M S parent. T h e s e e d o f the hybrid
cultivars t h e m s e l v e s must also be regenerated afresh e a c h time it is required b e c a u s e the hybrids do
not b r e e d true following self-pollination.

Breeding Objectives

S o r g h u m is g r o w n in a wide range of physical conditions in locations ranging f r o m t h e equator to over


5 0 ° N a n d 3 0 ° S . T h e crop is therefore subject to a wide variety of t e m p e r a t u r e , d a y - l e n g t h , a n d m o i s t u r e
r e g i m e s . T h u s b r e e d i n g i m p r o v e d s o r g h u m cultivars for a particular e n v i r o n m e n t a l w a y s involves b r e e d i n g
for adaptation to the specific climatic conditions f o u n d there. This is usually indicated by the appropriate
crop duration for that environment a n d by acceptable a n d stable yield levels a n d appropriate grain
qualities.

T h e type of cultivar required for a target location also influences the objectives of the plant
breeder. For e x a m p l e , the height of a pure-line variety for a specific e n v i r o n m e n t a n d the heights of the
parental lines of a hybrid for t h e s a m e environment are likely to be different.

In addition, i m p r o v e d cultivars for specific locations must p o s s e s s resistances to the major


constraints to production e n co u n t e r e d a n d grain- a n d stover-quality factors appropriate for s o r g h u m there.
T h e s e constraints include biotic — s u c h as d i s e a s e s , insects, a n d parasitic w e e d s — a n d abiotic
stresses, the requirements for which are usually quite different from o n e location to another. Resistances
to these constraints are deliberately bred into cultivars by crossing resistant types with cultivars
p o s s e s s i n g other desirable traits a n d selecting plants with both resistance a n d desirable traits. S o m e of
the constraints for which resistances are c o m m o n l y required or which otherwise influence the type of
cultivar required are outlined below.

Biotic Constraints

Diseases

C h a r c o a l rot. A c o m m o n fungal disease of the drier s o r g h u m gro w i ng a r e a s , particularly those with


terminal drought stress, charcoal rot ( M a c r o p h o m i n a phaseolina) is characterized by lodging a n d
p r e s e n c e of black sclerotia on the vascular tissues of the lower s t e m . It c a u s e s r e d u c e d grain size a n d
loss of grain d u e to lodging.

A n t h r a c n o s e . T h e m o s t important fungal leaf disease in W e s t Africa, a n t h r a c n o s e (Colletotrichum


graminicola) c a n also attack the stalk a n d panicle, c a u s i n g severe loss of grain a n d affecting t h e quantity
a n d quality of stover.

Leaf blight. Leaf blight (Exserohilum turcicum) is c o m m o n in the cooler s o r g h u m p r o d u c i n g e n v i r o n m e n t s


s u c h as s o u t h e r n Africa. An attack by this fungal p a t h o g e n p r e d i sposes plants to stalk rots.

G r a i n m o l d s . C a u s e d by a c o m p l e x of p a t h o g e n s ( Curvulaha, Fusarium, Phoma spp) w h i c h attack the


d e v e l o p i n g a n d m a t u r i n g grain in hot a n d h u m i d c onditions. Grain molds c a n c a u s e s e v e r e loss of

2
quantity a n d quality of g r a i n . Severely infected grain a p p e a r d i scolored to the n a k e d e y e . G r a i n molds
a r e often a c c o m p a n i e d by sprouting of the s e e d while still on the plant with s e v e r e c o n s e q u e n c e s for
viability a n d s e e d quality.

S o r g h u m d o w n y m i l d e w . C h a r a c t e r i z e d b y chlorotic, s t u n t e d plants with white d o w n y g r o w t h o n the


abaxial surface of infected leaves. S o r g h u m d o w n y mildew (Peronosclerospora sorghi) m a y c a u s e the
d e a t h of the plant w h e n infection occurs early. In later stages, leaves turn yellow a n d often s h o w
s h r e d d i n g at t h e tip. D i s e a s e d plants are often barren or partially s o .

O t h e r d i s e a s e s . S o r g h u m is subject to a large n u m b e r of other f u n g a l , bacterial, a n d viral d i s e a s e s , m o s t


of w h i c h are of local or sporadic significance. T h e s e include: seedling rots (Pythium, Fusarium,
Aspergillus, Rhizoctonia, Phoma spp), foliar diseases (gray leaf spot, z o n a t e leaf spot, r ough leaf spot,
oval leaf spot, s o o t y stripe, rust, a n d crazy-top), a n d panicle a n d s e e d d i s e a s e s (head s m u t , loos e kernel
s m u t , c o v e r e d s m u t , long s m u t , h e a d blight, a n d ergot). Several of these d i s e a s e s are of significance for
s e e d multiplication as they c a n be s e e d b o r n e .

Insects

S h o o t fly. F o u n d only in Asia a n d Africa. Shoot fly (Atherigona soccata) attacks the s e e d l i n g b e f o re the
sixth leaf stage a n d c a u s e s drying of the central leaf a n d d e a d h e a d s y m p t o m s , particularly in late
s o w i n g s . Smaller plants m a y be killed but larger ones usually survive by p r o d u c i n g tillers. Tillers often
flower later t h a n the m a i n s t e m , resulting in crop losses.

S t e m b o r e r s . Four species of stem borers are of significance in s o r g h u m in various parts of the w o r l d :


the s p o t t e d s t e m borer (Chilo partellus), maize stalk borer (Busseola fusca), African pink borer (Sesamia
calamistis), a n d African s u g a r c a n e borer (Eldana saccharina) are the m a i n c o n c e r n s . T h e y attack the
crop at a n y s t a g e , c a u s i n g d e a d h e a r t s y m p t o m s in y o u n g plants, shot holes in t h e leaves, a n d tunneling
of the s t e m a n d p e d u n c l e . Late attack can caus e breaking of the p e d u n c l e a n d r e d u c e d grain filling.

S o r g h u m m i d g e . Larvae of s o r g h u m midge (Calocoris sorghicola) f e e d on d e v e l o p i n g grain a n d c a u s e


empty/chaffy spikelets. Grain yield is directly affected as infested spikelets invariably set no s e e d .

H e a d b u g s . T w o m a i n s p e c i e s , Calocoris angustatus in India a n d Eurystylus immaculatus in W e s t Africa,


suck the d e v e l o p i n g grains. This c a u s e s direct losses a n d also predisposes p u n c t u r e d grain to m o l d
attack. G r a i n a t t a c k e d early fail to d e v e l o p , while older grain are r e d u c e d in size, thus affecting grain yield
a n d quality.

O t h e r i n s e c t s . A n u m b e r of other insects attack s o r g h u m at various stages of g r o w t h a n d c a n c a u s e


losses. M o s t a r e sporadic in o c c u r r e n c e a n d of limited general importance. T h e y include: seedling pests
( w i r e w o r m s a n d white grubs), foliage feeders (spittle b u g s , a p h i d s , s h o o t b u g , a r m y w o r m s , g r a s s h o p p e r s ,
a n d spider mites), a n d st orage pests. As with diseases, several of these insects are of significance for
s e e d p r o d u c t i on as t h e y c a n c a u se r e d u c e d s e e d size or quality resulting in impaired viability.

Parasitic Weeds

Striga. S e v e r a l s p e c i e s of Striga (S. hermonthica, S. asiatica, a n d S. forbesii) attack s o r g h u m plants by


sticking to the root a n d d r a w i n g water and nutrients f r o m the host plant, particularly under dry a n d low-
fertility conditions. Striga attack can severely stunt the s o r g h u m plant a n d c a u s e drying a n d failure to
produce seed heads.

3
Abiotic Constraints

T h e m a i n abiotic constraint to s o r g h u m production is drought. This follows f r o m the nature of the


e n v i r o n m e n t s in w h i c h s o r g h u m is c o m m o n l y g r o w n , where rainfall is often low a n d erratic. T h e crop has
natural a d a p t a t i o n to dry e n v i r o n m e n t s but there is variability a m o n g cultivars for dr ought tolerance , w h i c h
c a n be c a p t u r e d for particularly dry conditions.

S o r g h u m is also g r o w n in environments far from the equator w h e r e t e m p e r a t u r e s , particularly at


the b e g i n n i n g of the growing s e a s o n , are often low. Specially selected s o r g h u m with adaptation to
ger m inat ion in low t e m p e r a t u r e s are required for such areas. Similarly, w h e r e s o r g h u m is g r o w n at high
altitude, s u c h as in east ern Africa a n d Mexico, adaptation to low t e m p e r a t u r e t h r o u g h o u t the growing
period is required.

Other abiotic stresses for which tolerance is often required in s o r g h u m include high a n d low
t e m p e r a t u r e s at f l o w e r i n g , both of w h i c h c a n c a u s e sterility problems; soil factors such as low fertility a n d
acidity; high t e m p e r a t u r e s , drought, a n d soil crusting during germination; a n d terminal drought stress.

Quality Considerations

S o r g h u m grain a n d stover are put to m a n y different uses in different parts of the w o r l d . T h e s e uses
influence t h e types of cultivars g r o w n by farmers. F o o d uses vary t r e m e n d o u s l y from location to location,
a n d require different grain textures a n d colors. For e x a m p l e , s o r g h u m injera eaters in Ethiopia prefer a
soft white grain as it gives the preferred texture a n d color to the injera. S o r g h u m beer drinkers in eastern
Africa prefer high-tannin brown-grain s o r g h u m w h i c h gives their beer the bitter flavor they like. S o r g h u m
eaters in W e s t Africa prefer a hard-grain s o r g h u m which gives their porridge the right consistency.
Similarly, f o r a g e s o r g h u m for northern India must c o m b i n e high yields of g r e e n matter with the
appropriate quality of f o r a g e for animal c o n s u m p t i o n .

Seed Production and Maintenance of Improved Cultivars

D e v e l o p i n g i m p r o v e d cultivars of s o r g h u m requires considerable expenditure of time a n d effort to put


together the specific combinations of traits n e e d e d to ac hieve high a n d stable production of grain and/or
stover in a particular environment. T h e s e traits include the correct p h e n o l o g y for the e n v i r o n m e n t , the
n e c e s s a r y resistances to the biotic a n d abiotic constraints prevalent there, a n d the quality traits preferred
by farmers.

If i m p r o v e d cultivars are not maintained systematically, they are likely to deteriorate in yield a n d
quality d u e to o u t c r o s s i n g with the u n a d a p t e d cultivars lacking one or m o r e of the c o m p o n e n t traits.
Deliberate a n d s y s t e m a t i c m a i n t e n a n c e of cultivars a n d multiplication of their s e e d is, therefore, required
to e n s u r e that the genetic p a c k a g e a s s e m b l e d by the plant breeder is kept together a n d delivered to
f a r m e r s . Similarly, attention s h o u l d be given to the crop health of s eed-production plots to e n s u r e that
t h e s e e d d e l i v e r e d to f a r m e r s is in g o o d condition to germinate a n d establish the i n t e n d e d c r o p .

4
Reproductive Biology of Sorghum
Faujdar Singh

S o r g h u m (Sorghum bicolor (L.) Moench.) is a major f o o d a n d f e e d c r o p , g r o w n extensively in the


marginal rainfall a r e a s of the tropics a n d semi-arid regions of the w o r l d . T h e origin of cultivated s o r g h u m
has b e e n t r a c e d to Africa, particularly Ethiopia, S u d a n , a n d the East African region (Doggett 1965).
Cultivated s o r g h u m e v o l v e d f r o m the wild Sorghum bicolor s u b s p arundinaceum.

Germination and Seedling Development

A t o p t i m u m t e m p e r a t u r e ( 2 5 - 3 0 ° C ) a n d moisture, the s o r g h u m s e e d g e r m i n a t e s i n 3 - 5 d a y s . T h e s e e d
a b s o r b s water a n d swells, thereby breaking the s e e d coat. A small coleoptile a n d radicle (primary root)
e m e r g e ( H o u s e 1985). T h e coleoptile (Fig. 1) begins to e m e r g e f r o m the g r o u n d a n d t h e first leaf breaks
t h r o u g h the tip. As the y o u n g plant begins to g r o w , it bears m o r e leaves. T h e m e s o c o t y l g r o w s during
this period a n d a n o de is f o r m e d at the b a s e of the coleoptile, just b e l o w g r o u n d level. T h e y o u n g
seedling takes its nutrients f r o m the e n d o s p e r m . S e c o n d a r y roots d e v e l o p in 3-7 d a y s . G r adually, the
m e s o c o t y l dies a n d the seedling's nutritional requirements are m e t t h r o u g h the n e w roots. S o r g h u m
remains in vegetative p h a s e for 3 0 - 4 0 days.

Root System

T h e s o r g h u m root s y s t e m consists of three types of roots (Fig. 2).

P r i m a r y r o o t s . T h e s e roots d e v e l o p from the radicle a n d die subs equently, leaving a rudiment of t h e m


in the plant.

S e c o n d a r y or a d v e n t i t i o u s roots. T h e s e d e v e l o p f r o m the first internode o n t h e m e s o c o t y l . T h e y o c c u p y


a 5-15 cm a r e a in the soil a r o u n d the b a s e of the s t e m . Adventitious roots are s m a l l , u n i f o r m , a n d f o r m
only a s m a l l portion of the root s y s t e m .

A n o t h e r type of p e r m a n e n t adventitious roots d e v e l o ps f r o m the s e c o n d internode a n d a b o v e .


T h e s e roots a re b r a n c h e d laterally (about 1 m 2 ), interlacing the soil vertically. T h e y mainly supply
nutrients to t h e plant.

B r a c e ( b u t t r e s s ) r o o t s . T h e s e roots (Fig. 2) develop f r o m the root primordia of the b a s a l n o d e s a b o v e


g r o u n d level. T h e y are s t u n t e d a n d thick a b o v e g r o u n d level, but in t h e soil they are thin. Brace roots
provide a n c h o r a g e to the plant.

Shoot System

T h e shoot s y s t e m includes the s t e m , leaves, a n d nodes a n d internodes during the vegetative stage (Fig.
2).

S t e m . T h e s t e m or c u lm of s o r g h u m consists of m a n y alternating n o d e s a n d internodes. It ranges from


slender to v e r y s t r o n g , 0 . 5 - 5 cm in diameter near the b a s e , a n d 0 . 5 - 4 m in length ( House 1985).

5
E a c h n o d e a p p e a r s as a ring at the b a s e of the leaf s h e a t h . This is the point at w h i c h t h e leaf is
a t t a c h e d to the s t e m . A b u d is f o r m e d at each node except the one bearing the flag leaf. At t i m e s , these
b u d s d e v e l o p tillers.

P e d u n c l e . T h e t o p m o s t internode bearing the panicle is the p e d u n c l e . T h e larger the d i a m e t e r of the


p e d u n c l e , the larger is the panicle size. T h e peduncle m a y be straight or c u r v e d .

B l o o m . T h e w a x y coating on the surface of the internodes is b l o o m . It prevents transpirat ion.

Tillers. Tillers d e v e l o p f r o m the axillary buds situated at the n o d e s . Basal tillers d e v e l o p f r o m the axillary
b u d s of the lower n o d e s , a n d nodal tillers f r o m the axillary buds of the upper n o d e s .

L e a v e s . T h e leaves are borne alternately in two ranks (Fig. 2) along the s t e m . A leaf consists of a sheath
a n d a blade or lamina. T h e sheath is attached to the node a n d surrounds the internode, a n d frequently,
the n o de a b o v e it. T h e outer surface of the s heath is c o v e r e d with b l o o m . T h e blades are b r o a d at the
b a s e a n d taper u p w a r d to a fine point. T h e y are glabrous, except on the inside just a b o v e t h e ligule a n d
on the outside near the j u n c t i o n with t h e s h e a t h . Leaf-blade margins are s m o o t h or s c a rb i d . T h e midrib
is p r o m i n e n t , g r e e n i s h , b r o w n or white. T h e blades are thicker at the base than at the tip a n d along the
midrib t h a n a l o n g the m a r g i n s . T h e r e is a short (1-3 m m ) , triangular, m e m b r a n o u s ligule at the junction
of the leaf blade a n d the s h e a t h . T h e ligule deflects the lamina f r o m the stem at an a n g l e .

In cultivated s o r g h u m the n u m b e r of leaves is usually 1 4 - 1 7 . T h e leaves in the middle of the plant


are slightly longer t h a n those in the upper part. T h e t o p m o s t leaf, which is short a n d b r o a d , is called boot
leaf or flag leaf. T h e leaves m a y be as long as 1 m a n d m a y vary in width f r o m 10 to 15 cm ( House
1985).

T h e white e a r - s h a p e d structures on both sides of the base of the lamina are the auricles. T h e y
act as hinges to facilitate the m o v e m e n t of the lamina.

Reproductive System

I n f l o r e s c e n c e . T h e inflorescence of s o r g h u m is a panicle. T h e vegetative p r i m o r d i u m (growing tip)


differentiates into the reproductive primordium (Fig. 3). T h e shoot apex elongates into the main axis of
the inflorescence, w h i c h is called the rachis. T h e rachis tapers off t owards the top a n d is g r o o v e d
longitudinally. It p o s s e s s e s three types of hairs: (a) the fine types s p r e a d in the rachel f u r r o w s ; (b) the
long hairs on the rachel ridges; a n d (c) the scarbid hairs on the ridges. T h e rachis elongates a n d forms
b r a n c h e s a n d b r a n c h l e t s w i t h a rapid increase in d i m e n s i o n s . This results in the f o r m a t i o n of primary-,
secondary-, a n d tertiary-branch primordia. T h e tips of the tertiary-branch primordia d e v e l o p into two
paired spikelet primordia, o n e hermaphrodite a n d the other staminate. In s o m e c a s e s , o n e h e r m a p h r o d i t e
a n d t w o s t a m i n a t e spikelets are s e e n . T h e d e v e l o p m e n t of spikelets a n d florets in the inflorescence
continues c o v e r e d by the flag leaf. T h e primordial differentiation into floral parts m a y take a b o u t 30 days
after s o w i n g ( H o u s e 1985).

S o r g h u m panicles vary morphologically, ranging f r o m c o m p a c t to o p e n . After the c o m p l e t e


unfolding of the flag leaf, the p e d u n c l e elongation forces the panicle out of the leaf s h e a t h (boot). T h e
part of t h e p e d u n c l e b e t w e e n the b a s e of the lamina of the flag leaf a n d the b a s e of the p e d u n c l e is
exertion.

6
Figure 1. Sorghum germination.

Figure 2. Stylised sorghum plant and parts.


(Source: Vanderlip 1979)

7
T h e variation in s h a p e , size, a n d length of s o r g h u m panicles is due to variation in rachis length,
b r a n c h l e n g t h , distance b e t w e e n whorls, a n d the angle of b r a n c h i n g . S o r g h u m spikelet d e v e l o p m e n t is
b a s i p e t a l : t h o s e in the upper region of the panicle develop earlier than those in the lower.

R a c e m e . A r a c e m e consists of one or several spikelets. O n e spikelet of a r a c e m e is a l w a y s sessile a n d


the other pedicellate, e x c e p t the terminal sessile spikelet, w h i c h is a c c o m p a n i e d by t w o terminal
pedicellate spikelets. T h e length of the raceme varies according to the n u m b e r of n o d e s a n d the length
of the internodes. S o m e species have one to four nodes a n d others four to eight. Internode length,
t h i c k n e s s , a n d hairiness also vary f r o m g e n o t y p e t o g e n o t y p e .

S e s s i l e s p i k e l e t s . T h e s h a p e of sessile spikelets ranges from lanceolate to a l m o s t r o u n d or o v a t e .


S o m e t i m e s t h e y are d e p r e s s e d in the middle. At flowering they are green but then c h a n g e s h a d e s ,
b e c o m i n g straw- or c r e a m - c o l o r e d buff, yellow, red, b r o w n , purple, or a l m o s t black at maturity. T h e r e are
t w o g l u m e s w h i c h vary f r o m hairy to nonhairy. T h e g l u m e s are hard a n d t o u g h with n e r v e s , a n d are
o b s c u r e e x c e p t near the tip. In s o m e species, the g l u m e s are thin a n d brittle, while in others they are thin
a n d papery. T h e lower g l u m e is e n c l o s e d by the upper g l u m e with its m e m b r a n o u s m a r g i n . T h e lower
g l u m e is usually flat a n d c o n f o r m s more or less to the s h a p e of the spikelet. T h e upper g l u m e is m o r e
c o n v e x or b o a t - s h a p e d . T h e s e e d m a y be e n c l o s e d by the g l u m e or m a y protrude f r o m it either partially
or completely. T h e n u m b e r of sessile spikelets in a single inflorescence of cultivated s o r g h u m varies from
2 0 0 0 t o 4 0 0 0 ( H o u s e 1985).

T h e r e are t w o l e m m a e , c o m p o s e d of a delicate, white, thin, a n d papery tissue. T h e lower l e m m a


is elliptical or o b l o n g a n d equal in length to the g l u m e . T h e upper l e m m a is short, ovate a n d m a y be
a w n e d . T h e r e a re t w o lodicules a n d a palea. T h e spikelet has two pistils a n d three s t a m e n s . T h e stigma
is fluffy, a t t a c h e d to a short style extending to the ovary. T h e anthers are attached to long, thread-like
filaments.

Pedicellate s p i k e l e t s . T h e s e are narrower t h a n the sessile spikelets a n d are lanceolate. T h e y m a y be


smaller or longer or of t h e size of the sessile spikelets. T h e y are male or neutral, or m a y rarely have a
rudimentary ovary. T h e l e m m a e are short a n d the upper l e m m a rarely has an a w n . T h ree stamens and
t w o lodicules a r e f o u n d b e t w e e n the l e m m a a n d the palea. T h e lodicules at the b a s e of the floret are
truncate, fleshy, a n d ciliate.

Flowering

Floral initiation in cultivated s o r g h u m starts 3 0 - 4 0 d a y s after germination. T h e initial flower d e v e l o p s into


an inflorescence. A b o u t 6-10 d a y s before f l o w e r i n g , the boot f o r m s a bulge in the s h e a t h of t h e flag leaf.
S o r g h u m usually flowers in 55 to 70 days in w a r m climates (House 1985), d e p e n d i n g on the g e n o t y p e .

T w o d a y s after t h e e m e r g e n c e of the inflorescence f r o m the boot, the flowers begin to o p e n . T h e


f l o w e r i n g starts in t h e sessile spikelets at the tip of the inflorescence a n d p r o g r e s s es t o w a r d the bot t om
over 4 or 5 d a y s . It t a k e s 6 d a y s for the w h o l e inflorescence to c o m p l e t e f l o w e r i n g . T h e m a x i m u m
f l o w e r i n g t a k e s p l a c e o n t h e third o r fourth day. A t f l o w e r i n g , the g l u m e s o p e n , a n d the three anthers fall
free, while t h e t w o s t i g m a protrude, e a c h on a stiff style ( H o u s e 1985). As the s t a m e n s e m e r g e out of
the o p e n i n g g l u m e s , t h e y rotate a n d s p r e a d o u t w a r d . T h e filaments elongate rapidly a n d t h e anthers
b e c o m e p e n d e n t . W h e n f l o w e r i n g of t h e sessile spikelets is halfway through on the inflorescence, the
pedicellate spikelets start o p e n i n g f r o m the tip a n d p r o c e e d d o w n w a r d s , c o m p l e t i n g f l o w e r i n g earlier t h a n
the sessile spikelets in the inflorescence. T h e time taken f r o m the c o m m e n c e m e n t of g l u m e - o p e n i n g to
c o m p l e t i o n of its c l o s i n g is a b o u t 1 -2 h o u r s , w h i c h varies f r o m cultivar to cultivar.

8
F i g u r e 3 . G r a d u a l primordial differentiation into s o r g h u m i n f l o r e s c e n c e .

Figure 4. S t a g e s of s o r g h u m s e e d d e v e l o p m e n t .

9
F l o w e r i n g starts at midnight a n d continues up to 1000 d e p e n d i n g on the g e n o t y p e a n d climate.
T h e m a x i m u m a n t h e s i s i s b e t w e e n 0 6 0 0 a n d 0 8 0 0 . W e t a n d cool w e a t h e r delays flowering. T h e anthers
d e h i s c e w h e n they are dry a n d the pollen grains are ejected into the air a n d onto the s t i g m a . S o r g h u m
is primarily self-pollinated (cross-pollination is only 2 to 10 percent). T h e florets of s o m e of t h e v e r y - l o n g -
g l u m e d t y p e s do not o p e n for outcrossing to take place, a p h e n o m e n o n called c leistogamy. T h e
d i s c o v e r y of c y t o p l a s m i c m ale sterility in s o r g h u m has m a d e it possible to p r o d u c e c o m m e r c i a l hybrid
s e e d s . A g o o d male-sterile plant d o e s not develop anthers, or the anthers r e m a i n shrivelled without
pollen ( H o u s e 1985).

Pollination and Fertilization

F l o w e r - o p e n i n g is facilitated by the swelling of the lodicules. W h e n s t i g m a b e c o m e s visible, the s t a m e n


filaments elongate a n d the anthers b e c o m e pendent. This process takes about 10 m i n . T h e flower
remains o p e n for 30 to 90 m i n . After the anther d e h i s c e n c e , the pollen s h e d d i n g is t h r o u g h the apical
pore. T h e s t i g m a is pollinated before the e m e r g e n c e of the anthers f r o m the spikelets. W h e n pollen
grains l a n d o n t h e s t i g m a , they germinate immediately a n d develop pollen t u b e s , e a c h w i t h t w o nuclei,
o n e vegetative nucleus a n d two s p e r m nuclei. O n e s p e r m nucleus fertilizes the e g g t o f o r m a n e m b r y o
(2n) a n d t h e other nucleus f u s e s with the polar nuclei to f o r m the e n d o s p e r m (3n). S o r g h u m has a 2 0 -
c h r o m o s o m e c o m p l e m e n t . After pollination the g l u m e s close, t h o u g h the e m p t y anthers a n d stigmas still
protrude. T h e pollen retain their viability for 5 h at room t e m p e r a t u r e . In refrigerator the pollen retain their
viability for 3 to 4 d a y s ( S a n c h e z a n d Smeltzer 1965). T h e pollen require light to germinate (Artschwager
a n d M c G u i r e 1949). T h e s t i g m a remains receptive for 10 d a y s .

U n d e r n o r m a l conditions on receptive stigma fertilization takes place in 2 h. O r g a n differentiation


o c c u r s the following 12 d a y s , a n d the e m b r y o continues to grow until the s e e d is m a t u r e (Schertz a n d
Dalton 1980).

Seed and Seed Development

After fertilization, the e n d o s p e r m nuclei f o r m a small n u m b e r of free nuclei near the zygote. T h e s e f o r m
a cellular tissue by w h i c h t i m e the zygote will u n d e r g o the s e c o n d nuc lear division. T h e d e v e l o p m e n t of
the e m b r y o is g r a d u a l . T h e deposition of starch grains begin about 10 days after fertilization.

S e e d d e v e l o p m e n t is in three stages (Fig. 4): milk stage, early or soft- dough s t a g e , a n d late or
h a r d - d o u g h s t a g e (Murty et a l . 1994).

T h e pericarp d e v e l o p s with the g r o w t h of the e m b r y o . T h e m e s o c a r p a n d the h y p o d e r m i s of the


pericarp are chlorophyllous in the developing ovary. As the starch grains d e v e l o p in the e n d o s p e r m , the
g r e e n color d i s a p p e a r s (Sundararaj a n d Thulasidas 1980).

T h e s o r g h u m s e e d is a free caryopsis, also called grain. S e e d s are spherical in s h a p e , but


s o m e w h a t flat on o n e side with t h e e m b r y o at the b a s e . T h e y are r e d , b r o w n , white, yellow or c r e a m -
colored with a dull or pearly lustre. T h e e n d o s p e r m is usually white, s o m e t i m e s yellow. T h e nucellus is
hyaline or b r o w n . B r o w n nucelli d e v e l o p s e e d s that are poor in a p p e a r a n c e . T h e testa w h e n present, is
c o l o r e d a n d c o n t a i n s t a n n i n ( H o u s e 1985). T h e s e e d s h a p e varies from o v a t e , o b o v a t e , elliptical t o
orbicular. T h e s e e d size also varies f r o m s m a l l , m e d i u m , to bold (1-6 g 100' 1 s e e d ) .

T h e s e e d consists o f three parts (Fig. 5): pericarp (outer coat, 6 % b y w e i g h t ) , e n d o s p e r m


(storage tissue, 8 4 % ) , a n d e m b r y o ( g e r m , 10%). T h e pericarp is thin unlike the m e s o c a r p w h i c h has

10
m a n y layers. T h e grain is c o m p o s e d of the embryonic axis a n d the scutellum.

E m b r y o is m a d e up of 7 0 % fat a n d 1 3 % protein in the grain. T h e e n d o s p e r m varies f r o m


c o m p r i s i n g 1 0 0 % soft tissue with a little c o r n e o u s portion to a solid c o r n e o u s s e e d . It contains a layer
of a l e u r o n e cells, the outer c o r n e o u s e n d o s p e r m surr ounding a central floury e n d o s p e r m .

Cuticle
Epicarp
Mesocarp
Cross cells
Tube cells
Pericarp

Stylar area

Scutellum

Shoot

Germ

Hila
Testae

Aleurone layer

Peripheral endosperm

Corneous endosperm

Floury endosperm

Figure 5. Longitudinal view of the structure of a sorghum seed.


( S o u r c e : Murty et al 1994)

Physiological Maturity

T h e grain m a t u r e s in 30 to 35 days after fertilization. At physiological maturity a dark b r o w n callus tissue


is f o r m e d at the b a s e w h e r e the s e e d is a t t a c h e d to the spikelet. This callus tissue stops the translocation
of nutrients f r o m the plant to the s e e d . At physiological maturity, the s e e d contains 25 to 3 0 % moisture
a n d is fully viable. For safe storage, s e e d moisture should be brought d o w n to 1 0 - 1 2 % .

11
S o r g h u m Plant a n d Flower Parts
Belum V S Reddy

T h e p r e v i o u s c h a p t e r dealt with the theoretical aspects of s o r g h u m growth a n d d e v e l o p m e n t . This chapter


will d e m o n s t r a t e t h e g r o w t h stages of s o r g h u m , a n d its floral parts.

Germination, Seedling, and Panicle Development Stages

• W h e n a s e e d is s o w n in moist soil, it swells d u e to moisture a b s o r p t i o n . T h e s e e d coat breaks


a n d a small coleoptile a n d a primary root (radicle) e m e r g e . T h e coleoptile first a p p e a r s a b o v e the
g r o u n d in 3-10 d a y s , d e p e n d i n g on temperature (Fig. 1).

• As the y o u n g plant begins to g r o w , adding more leaves, the coleoptile remains as a s h e a t h at the
b a s e of t h e plant.

• T h e m e s o c o t y l g r o w s , a n d a n o d e is f o r m e d at the base of the coleoptile.

• S e c o n d a r y roots b e g i n to grow f r om this n o d e , 3-7 d a y s after the plant e m e r g e s f r o m t h e soil. T h e


primary roots die, a n d branch roots m a y d e v e l o p o n the l o w e r m o s t n o d e s .

• T h e m a j o r root s y s t e m d e v e l o p s f r o m the s e c o n d a r y or adventitious roots.

• L e a v e s / n o d e s d e v e l o p at the rate of o n e in 3-6 d a y s ; the plant remains in a vegetative p h a s e for


3 0 - 4 0 d a y s , during w h i c h all leaves (12-18) are f o r m e d . T h e s e e d has 6-7 e m b r y onic leaves.

- T h e c u l m or s t e m is m a d e up of a series of alternating n o d e s a n d internodes.


- T h e s t e m is solid, with a hard cortex or rind, with a softer pith at the center.
- T h e leaves are attached to the n o d e , a n d vary in length, width, a n d angle of attachment.
- L e a v e s are a r r a n g e d alternately on the s t e m a n d h a v e a s h e a t h a n d a lamina with a ligule
at the b a s e .

• T h e g r a n d g r o w t h stage is due to cell elongation.

• Floral initiation leading to inflorescence d e v e l o p m e n t involves:

- e l o n g a t i o n of the apical m e r i s t e m
- differentiation of primary-branch primordia on the floral apex
- differentiation of s e c o n d a r y - b r a n c h primordia
- d e v e l o p m e n t of s e c o n d a r y - a n d tertiary-branch primordia
- e l o n g a t i o n of the panicle
- f o r m a t i o n of t h e panicle b r a n c h (raceme)
- f o r m a t i o n of fertile (sessile) a n d sterile (pedicellate) spikelets on the r a c e m e s .

• T h e panicle has a central axis. There are m a n y s e c o n d a r y a n d tertiary rachises, w h i c h b e a r the


r a c e m e s (Fig. 2).

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• E a c h r a c e m e a l w a y s consists of o n e or several spikelets; o n e spikelet is a l w a y s sessile a n d the
o t h er pedicellate, e x c e p t for the terminal sessile spikelet, w h i c h is a c c o m p a n i e d by t w o pedicellate
spikelets.

• Sessile spikelets (seed-bearing) (Fig. 3) consist of:

- an outer g l u m e (convex or boat-shaped) a n d an inner g l u m e (flattened); g l u m e texture,


size, a n d thickness are variable. T h e s e e d is e n c l o s e d by the glume or protrudes from it
d e p e n d i n g on t h e race of s o r g h u m .
- t w o l e m m a s consisting of delicate white tissue; the lower o n e is o b l o n g or elliptical, equal
in size to t h e g l u m e ; the upper o n e is shorter a n d o v a t e , a n d m a y be a w n e d .
- t w o lodicules, m u c h r e d u c e d , useful in flower o p e n i n g .
- a p a l e a , m u c h r e d u c e d , of little interest.
- t w o pistils/stigmas, attached to a short, stout style extending to the ovary.
- three thread-like s t a m e n s , attached to the anthers.

• Pedicellate spikelets:

- m u c h narrower than the sessile spikelets, lanceolate in s h a p e .


- m a l e or neuter, but m a y very rarely have a rudimentary ovary.
- l e m m a s are m u c h r e d u c e d in size.
- the upper l e m m a rarely has a n a w n , d e p e n d i n g o n t h e g e n o t y p e .

• F l o w e r i ng ( o p e n i ng or protruding of anthers a n d stigmas) begins at the tip of the panicle a n d


p r o c e e d s t o w a r d the b o t t o m . It usually takes 4 - 1 2 d a y s for all the florets to flower.

• S e e d or caryopsis:

- spherical in s h a p e or s o m e w h a t flattened on one side, d e p e n d i n g on the race (Fig. 4).


- pericarp color varies (red, b r o w n , white, yellow, c r e a m - c o l o r e d ) .
- testa m a y also be colored (dark red to brown).
- e n d o s p e r m m a y be white or yellow (carotenoid pigments that have a relatively low vitamin
A activity).

Flower Opening/Anthesis—A Discussion

A n t h e s i s refers to t h e period of flower o p e n i n g during w h i c h the spikelets are o p e n , the anthers are
e x t e n d e d , a n d t h e pollen s a c s burst to release pollen.

F l o w e r i n g of s o r g h u m inflorescence normally starts as s o o n as the p e d u n c l e c o m p l e t e s its


e l o n g a t i o n ( H o u s e 1985). B l o o m i n g of the spikelets progresses f r o m the tip to the bot t om of the panicle
in a fairly regular m a n n e r , with t h e spikelets in the s a m e horizontal plane across t h e panicle o p e n i n g at
the same time.

F l o w e r i n g of the inflorescence m a y be s p r e a d over a period of 4 - 1 3 d a y s d e p e n d i n g on the


cultivar, panicle s i z e , t e m p e r a t u r e , a n d humidity. In cooler weather ( D e c / J a n at I C R I S A T Asia Center,
P a t a n c h e r u ) , c o m p l e t i o n of panicle flowering takes about 10-13 d a y s . T h e t w o lodicules at the b a s e of
the floret first swell a n d exert pressure on the g l u m e s , forcing t h e m to o p e n in about 10 m i n . This usually
happens between 0200 and 0800.

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Figure 1. Germination and seedling growth in sorghum (1-7 days).

Figure 2. Sorghum racemes.

14
Figure 3. Parts of sessile florets in sorghum.
(Source: Murty et al. 1994)

15
Usually, the anthers protrude first a n d then the stigmas. This condition is called protandry.
H o w e v e r , t h e r e are cultivar differences in which the stigmas a n d anthers e m e r g e together or one
p r e c e d e s or f ollows the other. For e x a m p l e , in the zera-zera-derived cultivar ICSV 112, the anthers c o m e
out first, while in the line 2 9 6 B, the stigmas protrude first s o o n after anthesis. In Bulk-Y, the stigmas
c o m e out m u c h earlier than the anthers. T h e filaments enlarge rapidly as the g l u m e s o p e n , a n d the
anthers become pendent.

Figure 4. Variation in the size and shape of sorghum seed.

T h e time f r o m the c o m m e n c e m e n t of glume opening to completion of closing is about 1 -2 h, whic h


varies f r o m cultivar to cultivar. A n t h e r o p e n i n g a n d s h e d d i n g of p o l l e n , called d e h i s c e n c e , a n d the time
of its o c c u r r e n c e d e p e n d s greatly on the w e a t h e r conditions; in the tropics, d e h i s c e n c e o c c u r s at a r o u n d
sunrise on clear d a y s . If the w e a t h e r is d a m p or cool early in the m o r n i n g , it can be d e l a y e d until 1000
h. Usually, the sessile (fertile) spikelets o p e n first; the pedicellate spikelets start o p e n i n g a f e w d a y s later,
but the anthesis of pedicellate spikelets is c o m p l e t e d in the panicle m u c h earlier t h a n the sessile
spikelets.

16
Selfing and Crossing Techniques in Sorghum
Belum V S Reddy

Biological variation is the basis of evolution. Plant breeders use this variation to direct a n d control
evolutionary p r o c e s s e s in order to develop new cultivars. Selfing a n d crossing are essential tools in the
regulation of variability in plant breeding p r o g r a m s . T h e breeder must therefore k n o w these techniques.

Importance of Selfing and Crossing

W h e n a flowering panicle is t a p p e d with a finger, a cloud of yellow pollen grains can be s e e n . T h e w i n d


carries the pollen grains to the stigmas, a n d pollination is a c h i e v e d . Pollen is normally viable for 3-6 h
in the anther, a n d 10-20 min outside. In nature, occurrence of outcrossing varies from 1 to 1 0 % ; in wild
t y p e s w i t h l o o s e / o p e n panicles, it m a y be up to 3 0 % . In normal c o m p a c t or s e m i c o m p a c t panicles in
i m p r o v e d cultivars, selfing can be up to 9 0 - 9 5 % , with 5 - 1 0 % outcrossing, occurring m o r e frequently at
the tips of the panicles.

Selfing a n d crossing or outcrossing are processes with opposite effects. Selfing promotes
h o m o z y g o s i t y a n d p r e s e r v e s the linked-gene c o m p l e x e s , w h i c h helps to maintain pure stocks of cultivars.
O u t c r o s s i n g or crossing p r o m o t e s recombination a n d creation of n e w linkage g e n e c o m p l e x e s , leading
to variability, w h i c h provides an opportunity to the breeder to select u p o n .

Techniques of Maintaining Genetic Purity of Seed Stocks

Various techniques are followed in breeding programs to maintain/enhance the purity of s e e d stocks. T h e
choice of the m e t h o d s d e p e n d s on the quantity of s e e d required, the resources available, a n d the extent
of genetic purity a n d trueness-to-type required for the given material. T h e s e m e t h o d s are d e s c r i b e d
briefly here.

R o g u i n g . This m e t h o d is followed w h e n complete purity is not a must, a n d w h e n the br eeder is f a c e d


with a large n u m b e r of lines with limited resources a n d pers onnel. Before flowering, all o d d plants or
'rogues' are r e m o v e d . T h e panicles, after t r i m m i n g the tips (top quarter of the panicle), are h a r v e s t e d a n d
b u l k e d f r o m the plots in b r e e d i n g blocks. This m e t h o d does not assure genuine purity of the stocks.
T h e r e f o r e , w h e n a crop is raised from s e e d multiplied in this manner, roguing should be r e p e a t e d .

Isolation. W h e n one or t w o stocks n e e d to be multiplied in large quantities, s o w i n g in isolation—field


plots a w a y f r o m other s o r g h u m plots, with or without variable s o w i ng t i m e s — i s desirable to maintain a
high level of genetic purity. Isolation with variable s o w i n g times is risky as there is a c h a n c e of
overlapping of flowering b e t w e e n different plots. Isolation that physically separates plots is c o n s i d e r e d
m o r e useful. A m i n i m u m distance of 2 0 0 m b e t w e e n s o r g h u m plots is r e c o m m e n d e d for multiplying
varieties or pure lines with the required standards of genetic purity. Generally, in India, s e e d certification
standards call for 9 9 . 5 % genetic purity. R o g u i n g of 'odds' or 'rogues' before a n d during flowering is
essential, a n d plants with trueness-to-type s h o u l d b e retained/harvested.

Selfing by b a g g i n g . S o r g h u m is a perfect-flowered plant (i.e., it has both sexes in the s a m e floret). Self-
pollination or selfing is the process of ensuring the transfer of pollen of a floret to the stigma of the s a m e
floret or of another floret within the s a m e panicle. This is usually a c c o m p l i s h e d in a b r e e d i n g p r o g r a m
by using kraft paper bags. T h e s e c o m e in varying sizes (14 cm x 6 cm x 37 cm or 14 cm x 6 cm x 42

17
c m ) . T h e c h o i c e of the size d e p e n d s on the potential size of the panicle w h e n it starts d e v e l o p i n g grain
after f l o w e r i n g .

In addition to kraft p a p er b a g s , the other things required for selfing are a pair of scissors, paper
clips or a stapler, w a t c h m a k e r t a g s, a n d a ma rker. T h e steps involved in selfing are:

1. R e m o v e o d d or off-type or rogue plants from the plot before they reach t h e boot leaf s t a g e .

2. W h e n a few florets h a v e o p e n e d at the tip of the panicle, snip off the f l o w e r e d florets.

3. Cut the flag leaf at the b a s e .

4. R e c o r d the date of selfing on t h e selfing b a g .

5. Put t h e b a g (with t h e date) over the panicle, taking care to s e e that the w h o l e of the panicle is
c o v e r e d by t h e b a g , a n d that the b a g also covers about 5-8 cm of the p e d u n c l e .

6. M a k e s u r e the p e d u n c l e stays in the center of the m o u t h of the b a g w r a p p e d over by the f o l d e d


corners of the paper b a g on either side.

7. Either staple the folded corners of the paper b a g or put a paper clip, taking care to s e e that the
b a g holds the p e d u n c l e tightly.

T e n to 15 d a y s after b a g g i n g , the b a g s c a n be r e m o v e d from the panicle. T h e s a m e bags or


w a t c h m a k e r tags c a n be stapled a r o u n d the peduncles to mark selfing. The n u m b e r of plants to be
bagged in a plot to e n s u r e selfing d e p e n d s on the purpose of developing materials (in c a s e of
s e g r e g a t i n g lines), a n d the quantity of s e e d n e e d e d (in c a s e of near uniform lines). Care s h o u l d be t a k e n
to harvest individual plants separately in c a s e of segregating generations, a n d bulk harvest the true-to-
type panicles only in c a s e of uniform lines.

P r e c a u t i o n s . B a g s c a n be blown a w a y by the wind or c a n be d a m a g e d by rain. C a r e must be taken to


replace t h e m immediately, recording the information originally written on the b a g s . Periodic inspection
of the self ed plots is essential to detect d a m a g e d b a g s .

Techniques of Hybridization

O u t c r o s s i n g , referred to as crossing here, is a process of transferring pollen grains f r o m a floret of o n e


panicle to the s t i g m a of a floret of another panicle. In nature, it is usually effected by the w i n d as stigmas
remain receptive up to a w e e k or more after b l o o m i n g , d e p e n d i n g on the t e m p e r a t u r e a n d humidity.
L o w e r t e m p e r a t u r e s (as in D e c e m b e r at IAC) favor a longer period of receptivity of s t i g m a s . H o w e v e r ,
stigmas are m o s t receptive during the first 3-5 d a y s after their e m e r g e n c e .

B r e e d i n g , as m e n t i o n e d earlier, is directed evolut ion, w h i c h allows variability. T h e gain from


b r e e d i n g is directly proportional to the a m o u n t of useful variability c r e a t e d by crossing t w o or m o r e
parents.

Hybridization of s o r g h u m on a field scale is m a d e feasible t h r o u g h the u s e of the genetic a n d


c y t o p l a s m i c - g e n i c m a l e sterile ( C M S ) s y s t e m . T h e r e are a n u m b e r of g e n e s (ms 1 , ms 2 , ms 3 , ms 4 , ms 5 ,
m s 6 , m s 7 , a n d ms 8 ) w h i c h individually contribute in h o m o z y g o u s condition to m a l e sterility. A l s o , another

18
s y s t e m , i n d e p e n d e n t of the genetic s y s t e m , called the C M S s y s t e m , creates m a l e sterility b e c a u s e of t h e
interaction of g e n e s in the c y t o p l a s m with those in the g e n o m e . This s y s t e m will be d i s c u s s e d in the next
chapter.

In addition to this, there are other techniques of hybridization w h i c h do not involve the m a l e sterile
s y s t e m . T h e s e are e x p l a i n e d b e l o w in detail.

Emasculation with Hot Water and Plastic-Bag Technique

Items r e q u i r e d . Plastic s l e e v e s , a pair of scissors, string, butter-paper b a g s , paper clips or a stapler with
staples, hot water in a t h e r m o s flask, a n d a t h e r m o m e t e r .

P r o c e d u r e . T h e steps involved a r e :

1. Cut a n d trim the florets at the tip of the panicle. T a k e a b a g m a d e out of a plastic s leeve a n d tie
it closely a r o u n d the peduncle to surr ound the panicle.

2. Pour hot w a t e r (42°C) into the closed plastic sleeve a n d leave it for 10 m i n , s o a k i n g t h e panicle
in hot water.

3. T h e water is d r a i n e d after 10 min, a n d the sleeve is tied over the panicle.

4. T h e florets at the top of the panicle o p e n after 2-3 d a y s a n d anthers e m e r g e but do not dehisce
a n d do not s h e d pollen; knock these anthers free f r o m the panicle by t apping it.

5. R e m o v e the remaining u n o p e n e d florets from the lower portion of the panicle.

6. Get pollen f r o m another panicle of the desired line in a butter- paper b a g a n d put it over the
e m a s c u l a t e d panicle, tying it a r o u n d the peduncle as in the selfing p r o c e s s .

7. Before collecting the pollen, write the name of the pollen parent and the date of
crossing/pollination on the b a g .

8. On the fourth d a y after pollination, check for selfed florets; these c a n be r e c o g n i z e d by their
distinctively superior size c o m p a r e d with the rest. R e m o v e the selfed florets a n d r e b a g .

(Note: T h i s is a c u m b e r s o m e m e t h o d a n d requires a lot of pr eparation. It leaves s o m e selfs in t h e F 1


w h i c h n e e d to be thor oughly c h e c k e d a n d rogued out. It is a l w a y s safer to follow the h a n d - e m a s c u l a t i o n
m e t h o d w h i c h can be easily d o n e by unskilled staff with s o m e training.)

Hand Emasculation and Pollination

C o n v e n t i o n a l c r o s s i n g or hybridization of different s o r g h u m varieties is carried out by s i m p l e h a n d


e m a s c u l a t i o n of normal bisexual florets, a n d then transferring the pollen f r o m the c h o s e n m a l e parent
(which usually is a pure line but not necessarily always) to t h e stigmas of t h e e m a s c u l a t e d florets.

I t e m s n e e d e d . A pair of scissors, a secateur or a manicuring clipper, a blunt needle or a pencil, a pair


of f o r c e p s , 7 cm x 3 cm x 15 cm butter-paper b a g , p a p e r clips or a stapler, a n d a m a r k i n g p e n .

19
P r o c e d u r e . T h e steps involved a r e :

1. F r o m the d e s i r e d parental line, c h o o s e a panicle that has just started anthesis.

2. Clip off the florets w h i c h h a v e c o m p l e t e d anthesis, with a secateur or scissors.

3. R e m o v e primary- or secondary- a n d tertiary-branch rachises in the lower portion of the panicle,


l e a v i n g a b o u t 2 0 0 - 3 0 0 florets in the central portion of the panicle just below the clipped florets.

4. Clip off all the pedicellate (sterile) florets f r o m the central portion, leaving only the sessile (fertile)
florets.

5. T h i n o u t the sessile florets by clipping off s o m e of the tertiary rachises to m a k e it easier to hold
t h e sessile florets during e m a s c u l a t i o n .

6. G r a s p the sessile floret to be e m a s c u l a t e d b e t w e e n the t h u m b a n d forefinger.

7. Insert a blunt n e e d l e b e t w e e n the g l u m e s below the middle portion of the floret, a n d m o v e it


s l o w l y a r o u n d the inner surface of the glumes so as to break the s t a m e n filaments.

8. Lift t h e n e e d l e out a n d u p w a r d s , slowly pus hing the d e t a c h e d anthers out of the floret.

9. After e m a s c u l a t i n g the florets as d e s c r i b ed a b o v e , cover t h e m with a butter-paper b a g a n d clip


or staple it as e x p l a i n e d earlier in the selfing process. T h e s e bags s h o u l d h a v e the date of
emasculation recorded on them.

10. Inspect t h e e m a s c u l a t e d panicles on the following day for any remaining anthers that might h a v e
e m e r g e d f r o m t h e florets. R e m o v e these florets a l o n g with the a n t h e r s , a n d o n c e a g a i n cover the
e m a s c u l a t e d panicles with b a g s .

11. O n t h e third/fourth day after e m a s c u l a t i o n , take the pollen from the c h o s e n parent into another
butter-paper b a g . Slowly insert the e m a s c u l a t e d panicle into the b a g with t h e pollen, a n d with a
h a n d over t h e b a g clasping the peduncle at the b a s e of the panicle, s h a k e the panicle so that the
pollen in t h e b a g stick to the stigmas that w o u l d have e m e r g e d f r o m the e m a s c u l a t e d florets.

12. Staple/clip as in selfing with t h e f olded corners of the m o u t h of the b a g clasping the peduncle.
M a k e s u r e that the b a g carries information on the date of e m a s c u l a t i o n , date of pollination, a n d
t h e m a l e p a r e n t u s e d in crossing. It is useful to pollinate a s e c o n d time on the f ollowing d a y to
e n s u r e pollination of all the florets.

W h e n t h e s e e d i s h a r v e s t e d f r o m the pollinated panicle, m a k e sure that the b a g containing the


s e e d is properly labelled a n d the male a n d f e m a l e parents a n d the crosses are clearly i n d i c a t e d . T h e
cross is usually d e n o t e d as f ollows:

N a m e of t h e p a r e n t f e m a l e x m a l e , w h e r e 'x' d e n o t e s the cross. For e x a m p l e , IS 3 5 4 1 x


IS 1 0 5 2 m e a n s that IS 3 5 4 1 as f e m a l e has b e e n c r o s s e d with IS 1052 as m a l e .

P r e c a u t i o n s . C a r e s h o u l d be t a k e n that the g l u m e closest to the pedicellate spikelet is h e l d f a c i n g a w a y


f r o m t h e w o r k e r . T r i m m i n g s h o u l d be d o n e so that the individual sessile florets r e m a i n uniformly s p r e a d

20
a l o n g the panicle b r a n c h .

Fertilization

Pollen grains g e r m i n a t e as s o o n as they c o m e in contact with a receptive s t i g m a ; the pollen t u b e s g r o w


through the stigmatic papillae d o w n to the ovary through the stylar region. O n l y o n e pollen t u b e s u c c e e d s
in reaching the m i c r o p y l e . T h e s p e r m nucleus divides into t w o , one of w h i c h fertilizes the e g g cell to give
rise to the e m b r y o (2n = 20 c h r o m o s o m e s in s o r g h u m ) , the other joins the two polar nuclei to f o r m the
e n d o s p e r m (3n = 30 c h r o m o s o m e s , i.e., 20 f r o m the female parent a n d 10 f r o m the m a l e parent). This
process of e m b r y o f o r m a t i o n by the union of e g g nucleus a n d s p e r m nucleus is called fertilization. T h e
g l u m e s close shortly after pollination. T h e ovule begins to develop as a light g r e e n , a l m o s t c r e a m - c o l o r e d
s p h e r e , a n d a b o u t 10 days after pollination, it b e c o m e s bigger a n d turns dark g r e e n . T h e d e v e l o p m e n t
of the e m b r y o a n d e n d o s p e r m continues for another 30 d a y s w h e n t h e s e e d reaches physiological
maturity with the hilum region (a spot on the s e e d through w h i c h the s e e d receives n o u r i s h m e n t of the
plant) b e c o m i n g black. During the d e v e l o p m e n t , t h e s e e d p a s s e s t h r o u g h milk, e a r l y - d o u g h , a n d late-
d o u g h s t a g e s . T h e dried-up style m a y persist i n s o m e s e e d s u p t o physiological maturity ( H o u s e 1985).

21
Development, Production, and Maintenance of Male-Sterile Lines
in Sorghum
Belum V S Reddy

T h e e m a s c u l a t i o n a n d pollination techniques described in the previous chapter cannot be u s e d for


production of hybrid s e e d on a c o m m e r c i a l scale. Male-sterile plants are more suitable for s u c h purposes
( H o u s e 1985). Male-sterile plants are characterized by nonproduction of functional pollen, or if pollen
grains are p r o d u c e d , they are either ill-formed or the anthers do not dehisce (Fig. 1). Usually, the former
type of male-sterile plants are recognisable in the field by their whitish, thin, a n d scaly anthers in contrast
to the p l u m p a n d yellowish male-fertile anthers. T h e latter group, with nondehiscent anthers, are usually
difficult to differentiate on the basis of anther a p p e a r a n c e or m o r phology. As a result, male-sterile
s y s t e m s c h a r a c t e r i z e d by nonfunctional pollen have b e en exploited widely in s o r g h u m . This male-sterile
s y s t e m is attributed to t w o c a u s e s : male-sterile g e n e s present entirely in the g e n o m e , or male-sterile
g e n es p r e s e n t in the g e n o m e as well as the c y toplasm. The former system is called genetic m a l e sterility
a n d the latter cytoplasmic-genic male sterility. Genic male sterility is d u e to single recessive alleles in a
h o m o z y g o u s condition. This system is not used for large-scale hybrid s e e d production, primarily b e c a u s e
it requires r oguing out of male-fertile plants from the male-sterile stocks. Cyt oplasmic-genic male sterility
is used widely in hybrid p r o g r a m s , a n d its principles are given below.

Figure 1. Male fertile (left) and sterile (right) sorghum anthers.

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Cytoplasmic-Genic Male Sterility

In s o r g h u m , S t e p h e n a n d Holland (1954) discovered cytoplasmic-genetic m a l e sterility ( C M S ) in t h e


p r o g e n i e s of a cross b e t w e e n t w o cultivars, Milo a n d C o m b i n e Kafir, with Milo as the f e m a l e a n d Kafir
as the m a l e . Male-sterile plants to the extent of 2 5 % w e r e o b s e r v e d in t h e F 2 generation of the a b o v e
cross w h e n Milo w a s u s e d a s f e m a l e a n d not a s m a l e . T h e male-sterile s e g r e g a n t s f r o m this cross
p r o d u c e d male-sterile hybrids w h e n crossed with the Kafir parent a n d fully fertile hybrids w h e n c r o s s e d
with the Milo parent. T h u s , it w a s recognized that Kafir c o u l d be u s e d as a maintainer of this source of
cytoplasmic-genetic male sterility. Since the pro g e n y received the c y t o p l a s m f r o m t h e f e m a l e , it w a s
h y p o t h e s i z e d that the Milo parent h a d a male sterility-inducing c y t o p l a s m a n d d o m i n a n t g e n e s for pollen
fertility, w h e r e a s the C o m b i n e Kafir parent contained a n o r m a l (fertile) c y t o p l a s m but the r e c e s s i v e m a l e -
sterile g e n e s . All pr ogenies of the Milo x C o m b i n e Kafir cross c o n t a i n e d Milo (sterility-inducing)
c y t o p l a s m , but t h o s e that also inherited the h o m o z y g o u s recessive g e n e s f r o m t h e Kafir parent w e r e
male-sterile. T h e male-sterile plants in the Milo x C o m b i n e Kafir cross w e r e u s e d as f e m a l e s in r e p e a t e d
b a c k c r o s s i n g with Kafir as the male parent. At the e n d of s e v e n b a c k c r o s s e s , t h e entire g e n o m e of Kafir
w a s transferred into the Milo c y t o p l a s m . This resulted in two morphologically similar v e r s i o n s of the
C o m b i n e Kafir (CK 60) parent: a male-sterile C o m b i n e Kafir (CK 60A) a n d a male-fertile C o m b i n e Kafir
(CK 60B). T h e male-sterile lines are designated as A-lines a n d their maintainer lines as B-lines.

Development of New Hybrid Parents (A-, B-, and R-Lines)

T h e parents that p r o d u c e fertile F 1 s w h e n c r o s s e d with A-lines are called restorers or restorer lines or
R-lines. T h e d e v e l o p m e n t of parents involves t w o steps: (1) identification of potential B- a n d R-lines; a n d
(2) d e v e l o p m e n t of A-lines a n d R-lines.

Identification of B- a n d R-lines. I m p r o v e d breeding lines, n a m e d / r e l e a s e d varieties, a n d l a n d r a c e s f o r m


the pollinator collection, the sources that can be u s e d as pollen parents or pollinators. T h e hybrids
o b t a i n e d by c r o s s i n g these pollinators with a male-sterile line, the testcrosses, a r e e v a l u a t e d for t h e
sterility m a i n t e n a n c e or fertility restoration in t h e m (Murty et al. 1994). This ev aluation is usually s o w n
in small plots (one or t w o rows of 2 m length). Examination of anther m o r p h o l o g y m a y be a basis for
classifying the hybrids as male-sterile or male-fertile; but it is not a sure w a y . A m o r e reliable m e t h o d is
the b a g g i n g test, i.e., c o v e r i n g 4-6 panicles with a p a p e r b a g before anthesis, a n d o b s e r v i n g the s e e d - s e t
after 2-3 w e e k s . (The items n e e d e d for b a g g i n g tests are the s a m e as in selfing.) T h e testcrosses are
of the following four types:

1. T e s t c r o s s e s exhibiting absolutely no seed-set on all the b a g g e d panicles, i.e., m a l e sterility w a s


m a i n t a i n e d in t h e s e hybrids. T h e c o r r e s p o n d i n g pollinator is classified as a maintainer or
nonrestorer or B-line. This c o u l d s e r v e as a s o u rce of a n e w A-line.

2. T e s t c r o s s e s with c o m p l e t e seed-set on all the b a g g e d panicles. T h e c o r r e s p o n d i n g pollen parents


a r e classified as potential restorer or R-lines. T h e y c a n serve as m a l e parents to p r o d u c e hybrids.

3. T e s t c r o s s e s with a partial seed-set on all the b a g g e d panicles. T h e c o r r e s p o n d i n g m a l e parents


a r e rejected f r o m the p r o g r a m as they serve neither as restorers nor as m a i n t a i n e r s .

4. T e s t c r o s s e s with a full seed-set on s o m e b a g g e d panicles a n d no seed-set in others. T h e


c o r r e s p o n d i n g pollen parent of such a hybrid is said to be segregating for fertility-restoration or
sterility-maintainer g e n e s . Usually, s u c h parents are not p u r s u e d further in a hybridization
p r o g r a m , as t h e y involve additional work of fixing the g e n e s for fertility restoration/sterility

23
maintenance.

D e v e l o p m e n t of n e w A- a n d R-lines. T h r e e criteria are used in the selection of parents for this p u r p o s e :


genetic diversity, the per se p e r f o r m a n c e of the lines, a n d the a v e r a g e p e r f o r m a n c e of a line in crosses
w i t h other lines [called general c o m b i n i n g ability ( G C A ) ] . Experience in s o r g h u m has s h o w n that parents
of diverse origin p r o d u c e highly heterotic hybrids. It has also been f o u n d that the per se p e r f o r m a n c e of
p a r e n t s is positively correlated with the p e r f o r m a n c e of the hybrids (Murty et a l . 1994). Further, the
g e n e r a l c o m b i n i n g ability is m o r e important t h a n specific c o m b i n i n g ability (the deviation f r o m
p e r f o r m a n c e p r e d i c t e d on t h e basis of general c o m b i n i n g ability) in s o r g h u m . Further, shorter (usually
1.25-1.75 m) a n d high-yielding lines with sterility-maintenance ability are c h o s e n for c o n v e r s i o n into m a l e -
sterile lines. Taller lines (usually 1.75-2.50 m) with restorer reaction are c h o s e n as R-lines.

T h e maintainors identified through the b a g g i n g test p o s s e s s recessive g e n e s for fertility


restoration/sterility m a i n t e n a n c e but h a v e a n o r m a l c y t o p l a s m . T h e selected B-lines c a n be c r o s s e d with
a n y r e c o g n i z e d male-sterile line. T h e resulting F,s a n d the c o r r e s p o n d i ng maintainers are s o w n
alternately in small plots, a n d the hybrids are b a c k c r o s s e d repeatedly with the respective maintainer lines
for six or s e v e n g e n e r a t i o n s u s i n g t he c o r r e s p o n d i n g maintainer lines as recurrent parents until m ale-
sterile lines with a p p e a r a n c e identical to the recurrent B-line parent are o b t a i n e d . It is important that
plant-to-plant c r o s s i n g s h o u l d be a t t e m p t e d in the ba ckcrossing phase. This involves crossing individual
male-sterile plants with individual plants of the recurrent parent that are morphologically similar to e a c h
other. T h i s plant-to-plant m e t h o d is useful to select out the partial sterility maintainers f r o m the p r o g r a m .
A l s o , it e n a b l e s faster realization of A-lines with morphological traits similar to the maintainer line.

T h e A - l i n es t h u s o b t a i n e d m a y be s o w n alternately with t he respective B-lines, a n d the pollen


(bulk) f r o m t h e respective B-lines collected in separate bags m a y be put over the male-sterile panicles
with e m e r g e d s t i g m a s . T h e b a g s s h o u l d b e s h a k e n thoroughly a s i n the outcrossing p r o g r a m explained
in the previous chapter. Before pollination, these male-sterile panicles s h o u l d be b a g g e d as in selfing to
prevent o u t c r o s s i n g with pollen f r o m u n w a n t e d parents. Similarly, the B-lines s h o u l d be s e l f e d . T h e s e e d
b u l k e d within t h e A-lines will f o r m the A-line s e e d . T h e B-line s e e d b u l k e d within the line will f o r m the B-
line s e e d . T h u s A- a n d B-lines a r e m a i n t a i n e d . It s h o u l d be r e m e m b e r e d that roguing s h o u l d be carried
out before selfing/pollination of A-/B-lines.

O n c e u n i f o r m A- a n d B-lines are p r o d u c e d , the stability of the m a l e sterility in t h e A-lines m a y be


e v a l u a t e d by s o w i n g t h e m in areas w h e r e t h e temperature at flowering reaches 4 2 ° C or m o r e . Unstable
A-lines b e c o m e fertile at this t e m p e r a t u r e .

Production of A-, B-, and R-Lines

S m a l l - s c a l e p r o d u c t i o n . R-line s e e d (identified through the b a g g i n g test o f testcrosses) m a y b e


p r o d u c e d by s o w i n g t h e s e e d in a plot of the d e s i r e d size a n d selfing the plants after r oguing out the off-
t y p e s before a n d at f l o w e r i n g . Bulk harvesting of true-to-type panicles m a y be d o n e . A plot of t w o rows
of 4 m l e n g t h , if m a i n t a i n e d properly, m a y give about 2.0-2.5 kg s e e d .

P r o d u c t i o n of A - a n d B-lines involves several operations:

1. S o w A- a n d B-lines in t h e plot side by s ide. Usually, for every four rows of A-line, t w o rows of B-
line a r e s o w n .

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2. C a r r y out roguing regularly in the A-lines a n d B-lines before a n d during anthesis. A p a r t f r o m off-
t y p e s , pollen 'sh e d d e r s ' c a n be a p r o b l e m in the A-lines [a pollen s h e d d e r is a fertile plant in the
A-line that results f r o m a b r e a k d o w n of male sterility; in practice, however, B-line (fertile) plants
w h i c h a p p e a r in t h e A-line plot d u e to m e c h a n i c a l mixing are also referred to as s h e d d e r s ] . T h e s e
s h o u l d be r e m o v e d by inspecting the field e v e r y d a y during anthesis.

3. P r u n e the florets of A-line with protruded anthers/stigmas at the tip of the panicles, a n d pull kraft
p a p e r b a g s o v e r the panicles with the date of b a g g i n g recorded on t h e m . C a r r y out a similar
o p e r a t i on on the B-line.

4. After 4 - 6 d a y s , collect pollen f r o m the B-line panicles into the s a m e bags u s e d for selfing, a n d
put t h e s e b a g s carefully over the respective A-line panicles, by slightly b e n d i n g t h e A - line, a n d
s h a k e the panicles along with the pollen bags by holding the m o u t h of the b a g tightly w r a p p e d
a r o u n d the p e d u n c l e . Each pollen b a g m a y be u s e d to pollinate 2-3 panicles of t h e s a m e A-line.

5. C o v e r t h e pollinated panicles with t h e s a m e pollen b a g o r with a n e w o n e . T h e b a g s h o u l d carry


information on the date of t h e first b a g g i n g a n d pollination, a n d an A x B m a r k indicating that it
w a s pollinated by a B-line.

6. Pollination of A-lines with B-lines m a y be repeated again after the 6th or 7th d a y in order to
pollinate all the florets in the entire panicle.

7. B-line panicles should be selfed by bagging after using their pollen to pollinate t h e A-line panicles.

8. T a k e out t h e b a g s 15-20 d a y s after pollination/selfing, a n d staple t h e m over t h e p e d u n c l e below


t h e b a s e of the panicles, as in selfing.

9. R o g u e out plants at the time of harvest, a n d bulk harvest the panicles in A-lines a n d B-lines
separately a n d label t h e m clearly.

P r e c a u t i o n s . Periodic replacement of d a m a g e d bags is essential.

L a r g e - s c a l e p r o d u c t i o n . Large-scale production of A-, B-, a n d R-lines is usually t a k e n up in isolation


plots ( C h o p r a 1982).

1. P r o d u c t i o n of R-line. R-line is p r o d u c e d in an isolation field s e p a r a t e d f r o m other s o r g h u m fields


by at least 3 0 0 m. Periodic roguing of the off-types is essential. Bulk harvesting is d o n e by taking
true-to-type panicles.

2. A- a n d B- lines. Production of A- a n d B-lines is d o n e by g r o w i n g the A-line in four r ows


alternating w i t h the c o r r e s p o n d i n g B-line in t w o rows. A c r o s s all the rows in the entire field, it is
r e c o m m e n d e d that a strip of 1 m length s h o u l d be s o w n with the B-line. This is useful in providing
pollen to the A-line panicles at the e n d of the rows. R o g u i n g of the off-type plants a n d pollen
s h e d d e r s s h o u l d be d o n e during anthesis everyday. O p e n pollination by w i n d will e n s u r e seed-set
on the A-lines. Self-pollination takes place in the B-lines. Harvesting of A-line a n d B-line s e e d
s h o u l d be d o n e separately. To avoid mechanical m i x i n g , it is r e c o m m e n d e d that they s h o u l d be
h a r v e s t e d at different t i m e s , preferably o n e after t h e other.

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Improving the B-Lines and A-Lines

We h a v e so far dealt with t h e p r o c e d u r e o f d e v e l o p i n g A-lines f r o m t h e B-lines identified f r o m the


pollinator collection t h r o u g h testcrossing. It is important to k n o w the p r o c e d u r e s involved in i m p r o v i n g A-
a n d B-lines in h y b r i d p r o g r a m s . It involves t h e f ollowing steps.

1. Identify t h e B-line(s) for i m p r o v e m e n t a n d t h e resistance source lines for stress factors or high-
yielding lines ( d e p e n d i n g o n the objective) w h i c h m a y be fertility restorers/sterility maintainers.

2. C r o s s the B-line with the selected source line(s) a n d a d v a n c e t h e m to t h e F 2 g e n e r a t i o n .

3. G r o w F 2 u n d e r the d e s i r e d s c r e e n i n g for resistance, a n d select for m o n o g e n i c or oligogenic traits


a p a r t f r o m resistance.

4. G r o w s e l e c t e d F 3 progenies in head-to-r ows under s c r eening for stress factors of interest. Select
plants with the desired combination of traits within the family selected for resistance a n d
uniformity.

5. T e s t c r o s s the s e l e c t e d s e g r e g a n t s onto an A-line s o w n separately near the F 3 nursery under


s c r e e n i n g . A l s o self the selected segregants (pollinator) u s e d in testcrossing.

6. G r o w the testcross a n d the pollinator (F 4 s) in a block near the pollinator s c r e e n i n g block. W i t h


e x p e r i e n c e , o n e c a n usually d e t e r m i n e the male-sterile testcrosses by anther m o r p h o l o g y at
a n t h e s i s . O t h e r w i s e o n e s h o u l d use the b a g g i n g test to identify male-sterile plants. R e p e a t step
4, i.e., select families for resistance a n d select individual plants for crossing on the basis of
a g r o n o m i c desirability.

7. B a c k c r o s s the male-sterile F 1 (A-line) panicles with pollen f r o m the s elected plants' (as above)
individual panicles as per the proc edure outlined for pollination. T h e F 4 families s e l e c t e d for
resistance s h o u l d be u s e d as pollinators. Harvest the b a c k c r o s s e d A-line panicles a n d selfed
pollinators' panicles individually a n d pair t h e m as per the pollination d o n e .

8. R e p e a t steps 6 a n d 7 for six to s e v e n generations. Care should be taken at every stage in the
f o l l o w i n g a r e a s : c h e c k m a l e sterility on the basis of anther m o r p h o l o g y a n d s e e d - s e t on a f e w
panicles under b a g g i n g ; also, a l w a y s m a k e plant-to-plant b a c k c r o s s i n g , i.e., the individual male-
sterile panicles (2-3) selected for b a c k c r o s s i n g s h o u l d be similar in m o r p h o l o g y to t h o s e individual
plants of t h e pollinators s e l e c t e d for pollination.

9. At t h e stage w h e n male-sterile lines resemble the respective maintainer lines a n d are uniform,
t h e y a r e called A- a n d B-lines. T h e B-lines m a y further be s elected on the basis of their per se
p e r f o r m a n c e a n d resistance to the factors of interest.

10. Further selection of A-lines m a y d e p e n d on G C A tests for traits of interest. T h e s e l e c t e d A- a n d


B-lines m a y t h u s be n u m b e r e d with the year, f ollowed by serial n u m b e r a n d letters A or B to
indicate m a l e sterility o r m a i n t e n a n c e . For e x a m p l e , 9 5 0 0 1 A a n d 9 5 0 0 1 B indicate that t h e s e t w o
represent o n e A a n d B pairs, b r e d in the year 1995, a n d the line n u m b e r is 1.

11. M a i n t e n a n c e of t h e s e l e c t e d A-lines is d o n e as per t h e p r o c e d u r e o u t l i n e d earlier.

26
I C R I S A T A s i a C e n t e r is improving C M S lines for resistance to various yield-limiting factors
(diseases: grain m o l d , a n t h r a c n o s e , leaf blight, rust, a n d d o w n y m i l d e w ; p e s t s : s h o o t fly, s t e m borer,
m i d g e , a n d h e a d b u g ; parasite: Striga asiatica). Also, the Center is attempting to diversify C M S lines for
earliness, grain m a s s , s t a y - g r e e n , tillering, etc. ( I CR I S AT 1 9 9 1 , 1992, 1993, a n d 1994).

27
Development, Production, and Maintenance of Restorer Parents
and Pure-Line Varieties in Sorghum
J W Stenhouse

A restorer line, or R-line, p r o d u c e s a male-fertile hybrid w h e n u s e d as pollinator parent for a male-sterile


line. A pure line is o n e w h i c h is h o m o z y g o u s for all loci, usually as a result of c o n t i n u e d self-pollination
f r o m a single plant, a n d w h i c h b r e e d s true for all its important traits.

S o r g h u m is a p r e d o m i n a n t l y self-pollinated crop w h i c h suffers little i n b r e e d i ng d e p r e s s i o n . As


s u c h , it is a m e n a b l e to f o r c e d inbreeding to produce uniform lines that can be u s e d in different w a y s . T h e
lines m a y be u s e d in their o w n right as pure-line varieties to be g r o w n by f a r m e r s , or as restorer lines
in c r o s s e s with male-sterile lines to p r o d u c e hybrids. T h e relative importance of the t w o types of use
varies a c c o r d i n g to t h e e c o n o m i c status of the target farmers a n d the d e g r e e of infrastructure required
to carry out s e e d multiplication. Pure-line varieties are more c o m m o n in areas w h e r e f a r m e r s are unable
to invest in s e e d e a c h s e a s o n a n d w h e r e sophisticated s e e d multiplication a n d p r o c e s s i n g facilities do
not exist. Hybrids are m o r e important w h e r e f a r m e r s are affluent a n d c a n invest in s e e d , a n d w h e r e
there is g o o d s e e d multiplication capability.

O p e n - p o l l i n a t e d varieties rarely exist in s o r g h u m . Landraces grown by farmers are often mixtures


of inbred a n d partially inbred lines, a n d represent the nearest approximation to open-pollinated varieties.
H o w e v e r , t h e s e are usually i m p r o v e d by selection a n d inbreeding to p r o d u c e pure-line varieties before
a n y large-scale s e e d multiplication takes place.

Procedures for Developing Inbred Lines

T h e p r o c e s s of d e v e l o p i n g inbred lines in s o r g h u m usually begins with deliberate crossing of two lines


with c o n t r a s t i n g characters by a n y o n e of a n u m b e r of m e a n s . T h e crosses are norm ally m a d e by h a n d
e m a s c u l a t i o n of o n e parental line a n d pollination by the other. T h e y m a y also be m a d e using a genetic
or c y t o p l a s m i c - g e n e t i c male-sterile line as the f e m a l e parent. In rare c a s e s , they m a y be d u e to natural
o u t c r o s s i n g . T h e F 1 generation of s u c h crosses is normally uniform but the s u b s e q u e n t generations
s e g r e g a t e , a n d selection of single plants with desirable combinations of traits c a n be practised.
Alternatively, s e g r e g a t i n g materials f r o m r a n d o m - m a t i n g populations or f r o m f a r m e r s ' fields m a y f o r m the
basis for t h e selection of individual plants for inbreeding.

It is e a s y to force self-pollination in s o r g h u m simply by placing a b a g over the panicle prior to


anthesis to p r e v e n t o u t c r o s s i n g . In early-generation materials (F 2 a n d F 3 ), s u c h b a g g i n g is not normally
p r a c t i s e d as t h e s e g e n e r a t i o n s are highly variable a n d the vast majority of plants are d i s c a r d e d . It is
therefore m o r e e c o n o m i c a l to tolerate the occas ional outcrossing in the early generations t h a n to b a g all
the plants. In later generations (F 4 or F 5 o n w a r d s ) , all plants are b a g g e d so that individual plants are
treated alike, a n d the effects of b a g g i n g do not interfere with selection for h e a d a n d grain traits.

S e l e c t e d plants f r o m segregating progenies are a d v a n c e d in a head-to-row p r o c e d u r e . Individual


panicles a re t a k e n f r o m the selected plants, a n d s e e d f r o m e a c h is s o w n in a single row in the
s u b s e q u e n t s e a s o n , often in a special nursery w h e r e selection for specific traits c a n be practised. T h e
p r o c e s s of s e l e c t i o n a n d h e a d r o w i n g of individual plants with d e s i r e d c o m b i n a t i o n s of c h a r a c t e r s , with
or without b a g g i n g to e n s u r e self-pollination, is r epeated for several generations until the p r o g e n y that
results is uniform for important traits. T h e line c a n be bulk harvested at this stage, a n d the bulk s e e d is

28
u s e d to establish replicated trials to c o m p a r e individual selections m o r e critically. Replicated testing of
yield or other traits c a n begin at any stage after progenies are reasonably uniform.

T h e criteria u s e d to select plants vary f r o m location to location. S o m e generally desirable traits


include large panicle, g o o d panicle exertion, appropriate time of maturity, plant height, bold grains, g o o d
seed-set, high s e e d n u m b e r , a n d attractive plant t y p e . Specific criteria, w h i c h m a y differ f r o m location
to location, include resistances to insects a n d diseases a n d grain quality for different e n d - u s e r s .

Testing Restoration Ability

Inbred lines with appropriate traits for use as restorer lines must be tested to d e t e r m i n e their fertility-
restoration reaction. Pollen f r o m the test lines is u s e d to pollinate o n e or m o r e h e a d s of a cytoplasmic-
genetic male-sterile line. T h e s e e d p r o d u c e d is harvested a n d s o w n to p r o d u c e a hybrid c r o p . O n e or
m or e h e a d s of t h e hybrid are b a g g e d prior to flowering to d e t e r m i n e w h e t h e r or not t h e hybrid is self-
fertile. O n l y lines w h i c h give fully fertile hybrids under b a g g i ng will be r etained as restorers for w i d e
testing in c o m b i n a t i o n with different male-sterile lines to generate experimental hybrids for a g r o n o m i c a n d
yield t e s t i n g . T h e fertility of hybrids should be tested during the s e a s o n in w h i c h t h e y will be g r o w n , as
fertility restoration c a n be influenced by temperature a n d humidity, a n d c a n differ dramatically f r o m
season to season.

Desirable Traits of G o o d Restorers and Pure-Line Varieties

Restorer lines are normally not intended for direct use as cultivars. Therefore, they do not necessarily
c o m b i n e all the traits that are f a v o r e d by f a r m e r s . T h e y c o m b i n e traits that m a k e t h e m suitable for use
as p a r e n t s of hybrids. T h e s e include fertility-restoration react i o n , g o o d p o l l e n - s h e d d i n g capacity,
appropriate height (usually shorter than the varieties), a n d g o o d grain yield.

T h e ability of a line to produce g o o d hybrids in combination with male-sterile lines is k n o w n as


its c o m b i n i n g ability. A line with g o o d general c o m b i n i n g ability will t e n d to give high-yielding hybrids with
m a n y male-sterile parents. Clearly, this is an a d v a n t a g e o u s trait in a restorer parent. It c a n be quantified
in c o m p a r i s o n with other restorer lines in specially d e s i g n e d testing s c h e m e s . In s o r g h u m , c o m b i n i n g
ability is closely a s s o c i a t e d with per se performance of t h e line. Yield testing of the restorer lines,
therefore, often substitutes for formal combining-ability tests.

Pure-line varieties are intended for use directly by f a r m e r s . T h e y must, therefore, c o m b i n e the
traits d e s i r e d by t h e farmers in the target location or area. T h e s e normally include high grain yield,
stability of yield over a range of conditions, appropriate duration to maturity, plant height, resistances to
e n d e m i c pests a n d d i s e a s e s , a n d suitable grain a n d stover quality. Typically, pure-line varieties are taller
than restorer lines with higher grain a n d stover yields.

M a i n t e n a n c e of Restorer Parents a n d Pure-Line Varieties

T h e p r o c e d u r e s for m a i n t e n a n c e of restorer lines a n d pure-line varieties of s o r g h u m a r e essentially the


s a m e . I n b o t h c a s e s , t h e p r o c e s s begins with planning s e e d requirements a n d projecting b a c k w a r d s .
Using conservative estimates of yields likely to be a c h i e v e d , the size of the m a i n t e n a n c e exercise
required at e a c h stage a n d its timing to e n s u r e prompt delivery of the final pr oduct is p l a n n e d .

29
For b r e e d e r s e e d m a i n t e n a n c e , a head-to-row procedure is usually f o l l o w e d . Individual self-
pollinated panicles that are true to the type of the line are selected, h a r v e s t e d , a n d t h r e s h e d individually.
T h e s e e d of e a c h panicle is u s e d to establish a single row in the s u b s e q u e n t s e a s o n , all t h e plants of
w h i c h a r e s e l f e d by b a g g i n g . T h e rows are scrutinized thoroughly to d e t e r m i n e their t r u e n e s s t o t y p e . A n y
lines w h i c h s h o w variation a n d deviation f r o m the descriptions are d i s c a r d e d . T h o s e lines w h i c h are
uniform a n d true to type are harvested a n d bulked to form the basis for breeder s e e d multiplication. T h e
quantities of s e e d required at this stage are normally small (a few kg) a n d c a n usually be met by
h e a d r o w i n g 1 0 0 - 2 0 0 plants.

T h e h e a d - t o - r o w p r o c e d u r e is often d i s p e n s e d with by e x p e r i e n c e d b r e e d e r s w h o are confident


of the uniformity of their lines a n d t h e a d e q u a c y of their b a g g i n g procedures to prevent cross-pollination.
In this c a s e , s e l e c t e d panicles of the selfed true-to-type plants are bulk ed without going t h r o u g h the h e a d -
t o - r o w s . But this is a d a n g e r o u s practice, as o n e can never be certain that cross-pollination w a s totally
a v o i d e d or that r oguing has r e m o v e d all products of outcrossing. In c a s e s w h e r e h e a d r o w i n g has b e e n
d i s p e n s e d with a n d c o n t a m ination is detected, it is best dealt with by going back to the head-to-row
p r o c e d u r e to repurify t h e line or variety.

W h e n the d e m a n d for breeder s e e d is high a n d is likely to run to hundreds of kilograms or tons,


large-scale multiplication of breeder s e e d for use in basic s e e d production is usually d o n e in isolation by
the b r e e d e r c o n c e r n e d . Usually this n e e d not be d o n e every year but a sufficiently large quantity c a n be
p r o d u c e d to provide a buffer stock as protection against s u d d e n increases in d e m a n d . Isolation distances
must be strictly a d h e r e d to in order to prevent contamination from adjacent s o r g h u m c r o p s . T h e s e are
r e c o m m e n d e d to be at least 3 0 0 m for other fields of grain s o r g h u m a n d 4 0 0 m for f o r a g e s o r g h u m but
c a n be m o d i f i e d in t h e light of factors s u c h as the prevailing w i n d direction a n d velocity, plot size, a n d
p r e s e n c e of w i n d b r e a k s . Strict scrutiny of the crop must be practised a n d rigorous r oguing of off-type
plants m u s t be d o n e before a n d after flowering to achieve the desired level of s e e d purity.

Certification of breeder s e e d production plots by c o m p e t e n t s e e d certification authorities is


required in India but responsibility for the purity of the br eeder s e e d is more typically that of the breeder
or institution c o n c e r n e d . Often there are no guidelines for its production.

Restorer lines a n d varieties of s o r g h u m are typically pure lines that h a v e b e e n p r o d u c e d by


c o n t i n u o u s selfing of individual plants selected f r o m segregating populations. Varieties t e n d to be taller
a n d higher yielding t h a n restorer lines which are typically dwarf. It is important for varieties to h a v e no
major w e a k n e s s e s but this requirement is less stringent for restorer lines w h o s e traits can be
c o m p l e m e n t e d by t h e traits of t h e male-sterile line. M a i n t e n a n c e of restorer lines a n d pure-line varieties
is d o n e by h e a d - t o - r o w s o w i n g of selected true-to-type plants a n d rigorous elimination of variable or off-
type p r o g e n i e s . Large-scale production of breeder s e e d for c o m m e r c i a l s e e d multiplication is normally
d o n e in isolated plots at least 3 0 0 m a w a y from the nearest grain s o r g h u m fields a n d 4 0 0 m a w a y f r o m
forage sorghum.

30
Production of Sorghum Hybrids
Belum V S Reddy

Identification of s o r g h u m hybrids for c o m m e r c i a l multiplication involves several s t e p s :

• D e v e l o p m e n t a n d production of A-, B-, a n d R-lines with high per se p e r f o r m a n c e a n d G C A . This


has b e e n d i s c u s s e d in the previous chapters.

• D e v e l o p m e n t a n d testing of experimental hybrids.

• Large-scale multiplication of the selected hybrid parents (A-, B-, a n d R-lines) for large-scale
production of designated/selected hybrids.

D e v e l o p m e n t a n d T e s t i n g of Experimental Hybrids

Production of A-, B-, a n d R-lines has b e e n dealt with in the previous chapters. Here we deal with the
production a n d testing of experimental hybrids.

1. G r o w A-lines a n d R-lines side by side in a crossing block. To get 5 0 0 g of hybrid s e e d , a 6-row


plot of 4 m length s h o u l d be s o w n with the required R-line to supply pollen for panicles in the A-
line. T h e A-line m a y be s o w n in 2-3 rows in a 4-m long plot.

2. At ant hes is , b a g the A-line a n d R-line panicles after pruning the florets that have protruded
stigmas/anthers.

3. R o g u e off-type plants before a n d during anthesis a n d avoid using t h e m in c r o s s i n g .

4. Pollinate panicles in the A-line with bulk pollen f r o m the R-line a n d b a g both A- a n d R-lines after
pollination.

5. Mark the bags with the following information: date of bagging of both the A-line a n d the R-line,
d a t e of pollination, a n d identity of the R-line used in crossing on the panicles of A-line.

6. H a r v e s t the F 1 hybrid s e e d f r o m the A-line panicles a n d selfed s e e d f r o m the R-line panicles


separately, a n d tag t h e m accordingly. Harvesting should be d o n e on different d a y s to avoid
mechanical mixing.

It is r e c o m m e n d e d that in c a s e of loss or d a m a g e or mutilation of the m a r k e d p a p e r b a g s, the


pollinated male-sterile panicles should be b a g g e d a n d m a r k e d again. So periodic inspection of the
pollinated plots s h o u l d be u n d e r t a k e n to c h e c k for a n d replace d a m a g e d b a g s .

T h e e x p e r i m e n t a l hybrids thus p r o d u c e d are ev aluat ed in replicated trials initially at o n e or t w o


locations, f o l l o w e d by multilocational testing of selected hybrids f r o m this initial evaluation. Multilocational
testing provides an opportunity to select 1 -3 promising hybrid(s) for large-scale o n - f a r m testing a n d finally
for release to f a r m e r s for c o m m e r c i a l cultivation.

31
D u r i n g this p r o c e s s of production a n d testing of experimental hybrids, information on t h e following
should be collected:

• F l o w e r i n g b e h a v i o r of R-lines in relation to the c o r r e s ponding A-lines. Usually, R-lines that flower


4 - 6 d a y s later t h a n the c o r r e s p o n d i n g A-lines are preferred.

• T h e seed-setting ability of A-lines w h e n c r o s s e d with R-lines. A-lines with inferior (< 5 0 % ) s e e d -


setting ( w h e n properly pollinated) s h o u l d be rejected.

• R-lines that a re shorter than t h e A-lines s h o u l d be rejected, as also R-lines that are too tall. In
g e n e r a l , t h e difference in height b e t w e e n an R-line a n d an A-line s h o u l d b e a b o u t 3 0 c m for
higher hybrid s e e d p r o d u c t i o n .

• R-lines w i t h g o o d pollen-shedding ability should be selected a n d poor pollen-shedders should be


rejected.

Large-Scale Multiplication of Selected Hybrids

W h e n large quantities of s e e d are required for o n - f a r m trials or for c o m m e r c i a l cultivation, s e e d of hybrid


parents (A-, B-, a n d R-lines) a n d hybrids are p r o d u c e d on large plots isolated f r o m other s o r g h u m fields.

S e e d p r o d u c t i o n of h y b r i d p arents (A-, B-, a n d R-Lines). This has b e e n dealt with in the previous
chapters.

H y b r i d (A x R) s e e d p r o d u c t i o n . S o r g h u m hybrid s e e d is p r o d u c e d by g r o w i n g the d e s i g n a t e d male-


sterile (A) lines (4 to 6 A-lines) a n d a single restorer line together in e a c h isolation a n d all owing cross-
pollination. T h e s e e d h a r v e s t e d f r o m the A-lines is the hybrid s e e d . T h u s the n u m b e r of hybrids (seed)
o b t a i n e d is e q u a l to the n u m b e r of A-lines, e a c h of the hybrids having a separate A-line as its f e m a l e
parent a n d t h e restorer as the m a l e parent, w h i c h is c o m m o n to all the hybrids thus p r o d u c e d . T h e area
s o w n with e a c h A-line d e p e n d s on t h e quantity of hybrid s e e d required. This m e t h o d of hybrid s e e d
p r o d u c t i o n is u s e d for p r o d u c i n g hybrids for on-farm testing.

On t h e other h a n d , w h e n a single hybrid is selected for c o m m e r c i a l cultivation, the s e e d is


p r o d u c e d in large quantities by growing the designated A-line a n d R-lines together in an isolation field
( C h o p r a 1 9 8 2 ; Murty e t a l . 1994).

In b o t h t h e situations, an isolation distance of more than 3 0 0 m is generally r e c o m m e n d e d for


hybrid s e e d p r o d u c t i o n . A distance of a b o u t 4 0 0 m is n e c e s s a r y if there are wild or forage s o r g h u m fields
in t h e vicinity ( H o u s e 1985).

H y b r i d s e e d p r o d u c t i o n t h r o u g h the time-isolation m e t h o d (to a v o i d ov er lap of f l o w e r i n g with the


adjacent crop) m a y be practised at an experimental station, but is not permissible in c o m m e r c i a l or
certified s e e d p r o d u c t i o n plots.

Male-sterile a n d restorer lines are s o w n in 4A : 2R or 6A : 2R d e p e n d i n g on the local conditions


s u c h as w i n d velocity, t e m p e r a t u r e d u r i n g anthesis, a n d the pollen-shedding ability of t h e R-line involved.
A l s o , t h e b o r d e r s on either side of the field s h o u l d be s o w n with the R-line over a strip of a b o u t 1 m width
to pr ov ide pollen to t h e A-line panicles at the e n d of the rows.

32
It is important, as indicated earlier, that the hybrid parents s hould 'nick' in their flowering w h e n
s o w n simultaneously. Further, information on the flowering behavior of the hybrid parents is useful in
d e c i d i n g t h e s o w i n g dates of the isolation involved for hybrid s e e d pr oduction. Also, care s h o u l d be taken
to e n s u r e that t h e s e e d d e v e l o p s a n d matures during a period w h e n it does not rain a n d w h e n relative
humidity i n the air d o e s not e x c e e d 5 0 % .

It is difficult to predict the flowering time of the parents accurately, b e c a u s e m a n y A- a n d R-lines


interact with t e m p e r a t u r e a n d day length. T h e interactions cannot be predicted (day length c a n be
calculated for a giv en location, but temperature cannot be predicted). So it is always advisable to stagger
the s o w i n g of R-lines, especially w h e n they are k n o w n to flower earlier than the A-lines. For e x a m p l e ,
if t h e m a l e parent is k n o w n to take 62 days to 5 0 % flowering at a given location a n d the A-line 72 d a y s ,
the m a l e p a r e n t h a s to be s o w n 13-15 days after s o w i n g the A-line. Pollen s h e d d i n g f r o m the R-line
usually lasts 10-15 d a y s . Transplanting the plants either in strips in e a c h row or all plants in o n e of the
t w o rows in t h e R-line is k n o w n to delay flowering by 8-10 days a n d thus increase the period of pollen
availability; this leads to higher seed-setting on the A-line.

It is important that roguing of off-types should be d o n e regularly before a n d during anthesis in both
the A- a n d R-lines. A l s o , monitoring for pollen s hedders in the A-line a n d r e m o v i n g t h e m s h o u l d be d o n e
without fail d u r i n g ant hesis. All possible precautions should be taken against s e e d contam ination through
mechanical mixing during harvesting a n d threshing. It is r e c o m m e n d e d that the R-line s h o u l d be
harvested first a n d r e m o v e d f r o m the field a n d yard before starting the harvest of the hybrid s e e d .

Further, the following precautions will enhance seed production:

• Selection of a field with g o o d fertility a n d appropriate previous c r o p , preferably a l e g u m e , will be


useful in ac hiev ing higher productivity of s e e d with g o o d quality.

• Monitor f l o w e r - b u d d e v e l o p m e n t 2 0 - 3 0 days after s o w i n g . Differences in the time of initiation a n d


size of panicle initially in the A- a n d R-lines w o u l d indicate differences in their time of 5 0 %
f l o w e r i n g ; so hasten flowering by selective irrigation and/or application of nitrogenous fertilizers
s u c h as urea or a m m o n i u m sulphate.

• Follow uniform crop m a n a g e m e n t practices to grow a g o o d crop across the entire isolation.

• Follow plant-protection m e a s u r e s a n d control pests a n d diseases b e c a u s e differential


susceptibility in the parents m a y lead to a s y n c h r o n o u s flowering (Murty et al. 1994).

33
Pearl Millet

An Overview of Pearl Millet Cultivar Development and


Maintenance
C T Hash

Pearl millet ( P e n n i s e t u m glaucum (L.) R. Br.) is the staple cereal g r o w n in the hottest a n d driest areas
of the w o r l d for f o r a g e , g r a i n , a n d stover. Since it is a cross-pollinated crop in w h i c h c o m m e r c i a l l y
exploitable c y t oplasmic nuclear male-sterility s y s t e m s are available, both o p e n - p o l l i n a t e d cultivars
(synthetics a n d c o m p o s i t e s , collectively referred to as "varieties") a n d hybrid cultivars (single-cross, three-
w a y , a n d t o p c r o s s hybrids) are practical. This section will cover b r e e d i n g , m a i n t e n a n c e , a n d s e e d
multiplication p r o c e d u r e s applicable to both groups of cultivars.

Breeding Objectives

Pearl millet is a major f o o d a n d f e e d crop of subsistence farmers in hot a n d dry e n v i r o n m e n t s t h r o u g h o u t


the s e m i - a r i d tropics. Yields are low a n d unreliable—but higher a n d m o r e reliable than a n y other available
alternative c r o p g r o w n in t h e s e regions. Breeding objectives for a staple cereal in s u c h e n v i r o n m e n t s
w o u l d be very different f r o m those for irrigated c r o p s , e.g., rice or w h e a t . F a r m e r s require a reliable
s o u r c e of grain for f o o d , a n d stover for construction material, f u e l , and/or livestock f e e d . T h e y require
cultivars that p r o d u c e e n o u g h surplus in the g o o d y e a r s , e v e n with little or no use of c o m m e r c i a l inputs,
to help t h e m t h r o u g h the b a d y e a r s . O v e r a n d a b o v e stable productivity, the cultivar m u s t be of a quality
a c c e p t a b l e for the p u r p o s e s for w h i c h it is i n t e n d e d . In this scenario, the breeder needs to focus on
i m p r o v i n g yield p e r f o r m a n c e o n - f a r m as o p p o s e d to on-station. I m p r o v e m e n t in t olerance or resistance
to relevant biotic a n d abiotic stresses is also an important br eeding objective.

T h e following objectives m a y h a v e relevance to pearl millet i m p r o v e m e n t in different target


environments.

Abiotic s t r e s s e s : D r o u g h t ( e s c a p e , tolerance), heat, soil acidity, soil fertility (response to fertilizer


application, a n d tolerance of poor fertility).

Biotic s t r e s s e s : Birds (escape, nonpreference bristles), diseases (downy mildew, ergot, pyricularia,
rust, s m u t , viruses), insects (head miner, s t e m borer), Striga.

Yield: F o r a ge (potential—plant/animal product, stability), grain (potential, stability), stover.

Quality: F o r a g e (in vitro dry-matter digestibility, lignin ( b r o w n midrib), palatability (visual


a s s e s s m e n t ) , grain [feed a n d f o o d (chemical, milling, organoleptic)].

Market demands: Grain color, grain s h a p e , grain size, s t e m thickness, a n d uniformity.

Seed: Brightness, longevity, vigor.

Stover: Construction material, f e e d , fuel.

34
Certification r e q u i r e m e n t s : Stable m a l e sterility of s e e d parent, unique trait c o m b i n a t i o n s .

Factors to be c o n s i d e r e d w h e n choosing breeding objectives are the i m p o r t a n c e of t h e trait,


availability of genetic variability for the trait (preferably in desirable a g r o n o m i c b a c k g r o u n d s ) , availability
of cost-efficient s c r e e n i n g p r o c e d u r es for the trait, a n d the time available before another cultivar release
is n e e d e d .

O n c e these general breeding objectives are identified, it is necessary to decide u p o n t h e f o r m of


the genetic p a c k a g e to be delivered to f a r m e r s . Only after this decision is m a d e is it really possible to
d e v e l o p r e a s o n a b l e w o r k plans to do the breeding research required to deliver an a d o p t a b l e cultivar to
farmers.

Cultivar T y p e s

Pearl millet cultivars a re of t w o t y p e s : open-pollinated cultivars a n d hybri d cultivars. O p e n - p o l l i n a t e d


cultivars differ f r o m hybrids primarily in their ability to self-perpetuate a n d t h e d e g r e e of variability
available in t h e m . S e e d h a r v e s t e d f r o m an isolated plot of an open-pollinated cultivar will h a v e essentially
the s a m e yield potential a n d uniformity as the crop f r o m w h i c h it w a s h a r v e s t e d . H o w e v e r , in c a s e of
hybrids it is n e c e s s a r y to reconstitute the cultivar by crossing its parents to p r o d u c e fresh s e e d to retain
uniformity a n d yield potential of the hybrid cultivar.

Hybrid cultivars h a v e several a d v a n t a g e s to s e e d s m e n a n d farmers c o m p a r e d to open-pollinated


cultivars. T h e y include higher yield potential within a given maturity, ability to c o m b i n e desirable
characters f r o m different parents into a single cultivar, a n d greater uniformity (facilitating birdscaring a n d
harvest operations). Further, closed pedigrees a n d the necessity of reconstituting hybrids f r o m their
parents allow private industry to protect their intellectual property resulting f r o m their i n v e s t m e n t in
research a n d d e v e l o p m e n t .

T h e a d v a n t a g e s of open-pollinated varieties are of t w o types: (1) their ability to self-replicate


(facilitating f armer-t o-f armer s p r e a d ) , a n d (2) their inherent genetic variability. This variability within o p e n -
pollinated varieties contributes to stability of performance by reducing risks of crop failure d u e to dr ought,
d o w n y m i l d e w , ergot, a n d s m u t .

Open-Pollinated Cultivars

T h e r e are three general types of open-pollinated cultivars of pearl millet:

1. M a s s - s e l e c t e d landraces
2. Products of recurrent selection within a breeding population
(a) C y c l e bulk
(b) R e c o m b i n a t i o n of s e l e c t e d progenies
3. Synthetic varieties b r e d by r a n d o m m a t i n g a set of inbred lines

Broadly, open-pollinated varieties are products of traditional crop i m p r o v e m e n t by f a r m e r s , or bred


by r a n d o m m a t i n g a s e l e c t e d fraction of a variable b r eeding population.

Hybrid Cultivars

Several types of hybrid cultivars are possible, d e p e n d i n g on the genetic structure of their parental lines.

35
These are discussed below.

Single-cross hybrid: B a s e d on two uniform inbred parents (e.g., A1 x R), w h e r e A1 (male


sterile) is maintained by B1 (maintainer line).

T h r e e - w a y hybrid- B a s e d on three nonrelated uniform inbred parents [e.g., (A1 x B2) x R]; B2
i s maintainer t o other than A 1 .

Modified three-way hybrid: B a s e d on t w o related uniform inbred parents as f e m a l e a n d a third


nonrelated uniform inbred parent as m a l e [e.g., (A1a x B1b) x R].

Topcross hybrid: B a s e d on a uniform inbred parent a n d a more variable, less inbred pollen
parent (e.g., A1 x R-population).

T h r e e - w a y t o p c r o s s hybrid: Hybrid b a s e d on t w o uniform inbred parents to p r o d u c e a F 1 f e m a l e


parents, a n d a nonrelated, variable, relatively less inbred u s e d as a m a l e
parent.

Hybrid Types Based on Seed Production Technology

H y b r i d t y p e s c a n a l s o b e d e s c r i b e d b a s e d o n s e e d production technology.

C h a n c e hybrids: B a s e d on r a n d o m crossing b e t w e e n several parents s o w n in bulk.

C M S - b a s e d hybrids: B a s e d o n male-sterile s e e d parents.

P r o t o g y n y - b a s e d hybrids: A l s o k n o w n as pro-hybrids, these are b a s e d on male-fertile s e e d parents.


S e e d production requires uniform flowering of uniculm f e m a l e . M a l e parent
m u s t s h e d ample pollen before anthesis of f e m a l e .

T h e o p e n - p o l l i n a t e d cultivars are also referred to as varieties. T h e s e c a n be m a i n t a i n e d a n d


multiplied by r a n d o m m a t i n g their representative bulk in isolation. T h e s e varieties are a m e n a b l e to
i m p r o v e m e n t by m a s s selection in the breeder s e e d isolation plot itself, provided the isolation is g r o w n
in t h e target e n v i r o n m e n t (location a n d season).

T h e s e t w o types of classification of hybrid cultivars are c o m p l e m e n t a r y , not mut ually exclusive.


T h u s , it is possible to h a v e a C M S - b a s e d three-way topcross hybrid (although m a i n t e n a n c e of all the
n e c e s s a r y parental stocks for s u c h a hybrid w o u l d be com plicated).

36
Reproductive Biology of Pearl Millet
Faujdar Singh

Pearl millet b e l o n g s to t h e family P o a c e a e , subfamily Panicoideae, tribe Paniceae, subtribe Panicinae,


g e n u s P e n n i s e t u m , a n d section Penicillaria (Pernes et al. 1984). Harlan (1971) has r eported that the
center of origin of pearl millet is in the Sahelian z o n e in Africa b e t w e e n w e s t e r n S u d a n a n d S e n e g a l .

Pearl millet has also b e e n d e scribed earlier as Pennisetum typhoides (Burm) Stapf a n d H u b b . ,
and Pennisetum americanum (L.) L e e k e . It is primarily cultivated for grain p u r p o s e s in India, N e w
M e x i c o , S o u t h e a s t A s i a , a n d the W e s t , East, a n d Southern African countries, a n d in the United States
of A m e r i c a for fodder. Pearl millet is an annual plant, w h i c h grows well on light-textured soil under low
moisture conditions. This chapter deals with the pearl millet plant a n d its reproductive s y s t e m .

Germination a n d Seedling Development

Pearl millet s e e d s g e r m i n a t e in 3-5 d a y s w h e n s o w n at a depth of 3-5 cm in the soil with sufficient


moisture a n d at o p t i m u m t e m p e r a t u r e . Germination begins with the absorption of moisture by the s e e d
d u e to w h i c h it swells a n d ruptures the s e e d coat. A small coleoptile a n d a radicle e m e r g e f r o m the s e e d
(Fig. 1). T h e coleoptile e m e r g e s out of the g r o u n d , a n d the first leaf d e v e l o p s f r o m its tip.

P e a r s o n (1975) reported that final germination of pearl millet w a s i n d e p e n d e n t of day/night


t e m p e r a t u r e s w h e n t e s t e d in a range of 15/10°C to 3 3 / 2 8 ° C . Howev er , the rate of g e r m i n a t i o n a n d
e m e r g e n c e , seedling survival, leaf-area e x p a n s i o n , a n d dry-matter a c c u m ulation w e r e highest at 3 3 / 2 8 ° C .

Root S y s t e m

T h e pearl millet root s y s t e m consists of three types of roots: primary, s e c o n d a r y , a n d brace roots (Fig.
2).

P r i m a r y r o o t s . T h e radicle roots or seminal roots after germination develop into the primary roots. T h e s e
roots continue to d e v e l o p for 4 5 - 6 0 d a y s .

S e c o n d a r y r o o t s . T h e s e are also k n o w n as adventitious roots. T h e first pair of s e c o n d a r y roots d e v e l o p s


f r o m the first n o d e of t h e primary root at the two- or three-leaf s t a g e . T h e next n o d e gives rise to the
s e c o n d pair of s e c o n d a r y roots, a n d thereafter, four to six roots are p r o d u c e d at e a c h n o d e . T h e s e roots
c a n penetrate up to a depth of 5 m into the soil.

C r o w n or b r a c e r o o t s . T h e s e roots develop from the lower n o d e s o f the s t e m a t o r a b o v e g r o u n d level.


T h e y p r o v i de a n c h o r a g e to the plant.

T h e m a x i m u m root s p r e a d in pearl millet is confined to 2 5 - 3 0 cm of the soil a r e a a r o u n d the


plant.

Shoot System

T h e pearl millet s h o o t consists of a s t e m , tillers, a n d leaves during the vegetative stage (Fig. 3).

37
F i g u re 1. S e e d l i n g e m e r g e n c e in pearl millet. Figure 2. Parts of a pearl millet
( S o u r c e : Maiti a n d Bidinger 1981) seedling.
( S o u r c e : Maiti a n d Bidinger 1981)

5 6 7 8 9
4

2
1
0

Vegetative phase (GS 1 ) Panicle development phase (GS 2 ) Grain-filling phase (GS 3 )

GS1 = from seedling to panicle initiation of the main stem

GS2 = from panicle initiation to flowering of the main stem

GS3 = from flowering to physiological maturity period of crop

Figure 3. G r o w t h phases of pearl millet.


( S o u r c e : Maiti a n d Bidinger 1981)

38
S t e m . T h e pearl millet s t e m or c u l m consists of nodes a n d internodes. T h e solid s t e m is thick at the
b a s e . T h e n o d e s a r e slightly swollen, a n d axillary b u d s are present in shallow g r o o v e s at t h e n o d e s . T h e
n o d e s are hairy as well as glabrous with a gradual increase in hairiness f r o m the b a s e to the apex. T h e
lower internodes are largely c o v e r e d by the sheaths of the leaves, while t h e upper internodes a re only
partly c o v e r e d . T h e u p p e r internodes are more elongat ed t h a n the lower o n e s . T h e height of the s t e m
r a n g es f r o m 0.5 m to 3.0 m.

Tillers. T h e d e v e l o p m e n t of tillers starts f r o m the lower nodes of the s t e m 3-5 w e e k s after e m e r g e n c e .


T h e s e are called primary tillers. T h e secondary (axillary) tillers develop f r o m the axillary b u d s at the upper
n o d e s , usually after c o m p l e t i o n of flowering on the main s t e m .

L e a v e s . T h e leaves are a r r a n g e d alternately in two vertical rows on the c u l m . T h e y a r e linear in s h a p e


c o m p r i s i n g of a leaf s h e a t h a n d a blade. T h e leaf sheaths are light green or purple a n d e n v e l o p the s t e m
slightly a b o v e the n o d e s . T h e leaf sheaths overlap at the b a s e of t h e s t e m but are o p e n in the upper
portion. T h e leaf s h e a t h is g l a b r o u s a n d g r o o v e d . Ligules are present at t h e junction of the leaf blade a n d
the leaf s h e a t h . T h e y are m e m b r a n o u s , hairy, 4-5 mm long, a n d clasp the s t e m .

T h e leaf blade is lanceolate, 9 0 - 1 0 0 cm long a n d 5-8 cm w i d e , with a pointed tip. T h e midrib is


dull g r e e n . Auricles a r e present at the b a s e of the leaf blade. T h e upper leaf surface is s c a r b i d a n d hairy,
while the lower o n e is s m o o t h a n d glabrous.

T h e leaf a r e a of pearl millet is calculated by measuring the length a n d the width (at the broadest
point) as s u g g e s t e d by S i n g h et a l . (1970).

Leaf a r e a = Leaf length (cm) x Leaf width (cm) x 0.7236

Reproductive System

T h e s h o o t a p e x is t r a n s f o r m e d into a reproductive a p e x (Fig. 4) with the f o r m a t i o n of t h e panicle


m e r i s t e m . T h e panicle m e r i s t e m is b u l b o u s , with a constriction at its b a s e . It d e v e l o p s t u n i c a a n d c o r p u s
layers in t h e inflorescence axis. T h e s e cells (tunica a n d corpus) are eumeristematic. T h e different floral
o r g a n s d e v e l o p f r o m t h e a p e x . The organogenesis a n d maturation of organs in the spikelet a p e x are
acropetal (Gill 1991).

I n f l o r e s c e n c e . T h e pearl millet inflorescence is a false spike, ranging f r o m 5 to 150 cm in length a n d


1 to 5 cm in diameter. T h e panicle is terminal, varying in s h a p e f r o m cylindrical to c a n d l e - s h a p e d . T h e
spike consists of a central rachis w h i c h is closely p a c k e d with fascicles. E a c h fascicle consists of o n e
or m o r e spikelets a n d a whorl of 7 0 - 8 0 bristles. T h e tip of the spike has only a single spikelet. T h e
bristles m a y be free or united to f o r m an involucre. A b o u t 10-15 bristles in t h e upper whorl s are longer
t h a n t h o s e in t h e lower o n e s . T h e bristles are b r oader at the base a n d narr ower at the tip. T h e y are
scarbid a n d g r e e n or purple. An inflorescence m a y contain 8 7 0 - 3 0 0 0 spikelets with an a v e r a g e of 1600
(Khairwal et a l . 1990).

S p i k e l e t s . T h e spikelets a re s m all, lanceolate, a n d acute. Each spikelet consists of t w o g l u m e s , one


outer a n d o n e inner. T h e outer g l u m e is b r o a d , short, m e m b r a n o u s , a n d truncate. T h e inner g l u m e is
b r o a d a n d half t h e size of t h e spikelet. B e t w e e n t h e t w o g l u m e s , there are (two) florets. T h e lower floret
is s t a m i n a t e a n d t h e u p p e r floret is h e r m a p h r o d i t e (Fig. 5). Spikelets are generally bifloret but c a n
s o m e t i m e s be trifloret or tetrafloret; in rare instances, m o r e t h a n four florets a r e present (Maiti a n d Bisen
1979).

39
Figure 4. Primordial initiation (1), panicle d e v e l o p m e n t (2), a n d
p r e b o o t s t a g e (3) in pearl millet.

F i g u r e 5 . S e e d d e v e l o p m e n t , b i s e x u a l , a n d m a l e f l o w e r s i n pearl millet.

40
T h e staminate flower has o n e l e m m a a n d o n e palea, a n d e n c l o s e d b e t w e e n t h e m i s the
a n d r o e c i u m with three s t a m e n s . T h e upper hermaphrodite floret has a b r o a d , pointed l e m m a a n d a thin,
oval palea, a n d the a n d r o e c i u m a n d g y n o e c i u m are e n c l o s e d b e t w e e n t h e m .

A n d r o e c i u m . T h e pearl millet a n d r o e c i u m consists of three anthers (Fig. 5), e a c h a t t a c h e d to a long


filament. T h e r e is a t w o - l a y e r e d epidermis, a t a p e t u m , a n d pollen grains. T h e anthers are y ellow or purple
with a tuft of fine hairs at the a p e x .

G y n o e c i u m . T h e g y n o e c i u m consists of a monocarpellary a n d superior ovary with t w o styles a n d a


feathery s t i g m a . T h e pistil in its y o u n g stage s h o w s t w o carpels, one larger t h a n the other. T h e larger
o n e contains the primordium of the ovule. T h e growth of the two carpels is u n e q u a l . T h e thicker o n e ,
b e a r i n g t h e p r i m o r d i u m , g r o w s in girth, while the thinner one g r o w s in length a n d s o o n overtakes the
other t o f o r m t h e style a n d s t i g m a at the t o p . T h e mature ovary has an a n a t r o p o u s o v u l e . T h e inner
i n t e g u m e n t c o m p l e t e l y c o v e r s the ovule a n d forms the micropyle, a n d the other stops its g r o w t h at a very
early s t a g e . T h u s the e m b r y o has a scutellum, plumule, coleoptile, primary axis, coleorhiza, a n d radicle
(Rachie a n d M a j u m d a r 1980). T h e styles are c onnate at the b a s e , bearing the stigma at the tip.

Panicle E m e r g e n c e

T h e e m e r g e n c e of t h e panicle f r o m the sheath takes about 4-6 d a y s (Bhatnagar a n d K u m a r 1960).


Flowering starts after the e m e r g e n c e of the panicle out of the boot (Fig. 6), but in s o m e g e n o t y p e s style
exertion c o m m e n c e s before completion of panicle e m e r g e n c e . Stylar exertion begins first in the florets
in the central upper portion of the panicle a n d then progresses u p w a r d as well as d o w n w a r d . T h e
m a x i m u m exertion of styles is on the third day of flowering.

T h e flowering of pearl millet is protogynous, i.e., the stigma e m e r g e s earlier t h a n the anthers (Fig.
7). T h e s t i g m a r e m a i n s receptive for 12-16 h. Protogyny is e x p r e s s e d in v a r y i n g d e g r e e s in pearl millet,
d e p e n d i n g o n t h e g e n o t y p e a n d the e n v i r o n m e n t (Rachie a n d M a j m u d a r 1980).

A n t h e s i s a n d Pollination

A n t h e r e m e r g e n c e begins o n e d a y after the e m e r g e n c e of the stigmas is c o m p l e t e d on the panicle. It first


starts in t h e h e r m a p h r o d i t e florets followed by the staminate florets. Anther e m e r g e n c e is facilitated by
the p r o t o g y n o u s styles a n d the tufts of hair on the tips of the anthers. Anthesis continues t h r o u g h o u t the
day. T h e m a x i m u m anthesis i s b e t w e e n 2 0 0 0 a n d 0 2 0 0 (Sundararaj a n d T h u l a s i d a s 1980).

Exertion a n d e m e r g e n c e of anthers takes about 60 min if it h a p p e ns during the d a y ; during the


night it m a y t a k e twice as m u c h time or m o r e .

A n t h e r e m e r g e n c e starts in the upper portion (at about the two-thirds point) of t h e panicle a n d
p r o c e e d s in both directions. T h e first flush of anthesis is c o m p l e t e d in a w e e k ' s time under irrigated
conditions. Panicles e m e r g i n g f r o m the tillers start flowering later a n d the process m a y continue up to
t h r e e w e e k s . In rainfed conditions, first-flush anthesis of a plant m a y take place over 12 d a y s , a n d it m a y
continue on the tillers till s e e d formation ( C h a l a m a n d V e n k a t e s w a r l u 1965).

T h e pollen r e m a i n viable for 5 h at room t e m p e r a t u r e . Burton (1965) not ed that w h e n the pollen
w e r e s t o r e d in glassine b a g s at 2 7 ° C , they r e m a i n e d 5 9 % as effective as fresh pollen after 1 day, 1 0 %
as effective after 2 d a y s , a n d only 3% as effective after 3 days. H o w e v e r , w h e n stored at low temperature
(4 or 5 ° C ) , they r e m a i n e d viable for 3 w e e k s (Cooper a n d Burton 1965). H a n n a a n d Y o u n g (1974)

41
F i g u r e 6. P e a r l millet boot stage a n d panicle
emergence stages.

F i g u r e 7. P r o t o g y n o u s s t a g e in selfed
panicle.

42
s u g g e s t e d that pollen with a moisture level of 7 . 5 % c a n be stored in a plastic zip lock b a g tor 185 days
a t - 7 3 ° C , r e m a i n i n g 1 0 0 % viable.

Fertilization and Grain Formation

After pollination, the stigmas dry up in 24 h. Seed-set can be seen in the panicle a b o u t a w e e k after
fertilization (Burton a n d Powell 1968). There is a gradual increase in t h e dry m a s s of grains f r o m the milk
to d o u g h s t age, reaching m a x i m u m at maturity. Physiological maturity of the grain is indicated by the
a p p e a r a n c e of a black layer just a b o v e the hylar region on the abgerminal side of the grain opposite the
e m b r y o (Fussel a n d P e a r s o n 1978). A t physiological maturity, the s e e d contains 3 0 % moist ure.

T h e r e are large variations in s e e d s h a p e (Fig. 8) — obovate, oblanceolate, c u n e i f o r m , pyriform,


p y r a m i d a l , elliptical, h e x a g o n a l or globular — size, a n d color. T h e grain m a s s varies f r o m 3 to 5 g 1000' 1
s e e d s for s m a l l - s e e d e d g e n o t y p e s a n d 10 to 12 g 1000 - 1 seeds for b o l d - s e e d e d g e n o t y p e s . T h e s e e d s
are yellow, w h i t e , light b r o w n or gray.

Grain Structure

T h e pearl millet grain is a caryopsis with three m a i n parts: pericarp, e n d o s p e r m , a n d g e r m (Fig. 9). T h e
pericarp contains three layers of tissues: epicarp, m e s o c a r p , a n d e n d o c a r p . T h e t e r m bran refers to the
pericarp, the s e e d coat, a n d the aleurone layer of the s e e d . T h e e n d o s p e r m contains s i m p l e starch
granules a n d protein bodies. T h e g e r m contains about 2 5 % lipids, 2 0 % protein, a n d phytin, v i t a m i n s , a n d
e n z y m e s . T h e g e r m of pearl millet constitutes about 1 7 % of the total s e e d m a s s ( R o o n e y a n d
M c D o n o u g h 1987).

Factors Influencing Flowering and Seed Formation

Photoperiod a n d t e m p e r a t u r e affect flowering in pearl millet. S o m e genotypes fail to flower w h e n the d a y


length e x c e e d s 12 h. M a n y g e n o t y p e s flower under long d a y length conditions (16 h photoperiod), but
flowering t a k e s place earlier under short-day conditions (Burton 1965).

Increasing t h e t e m p e r a t u r e up to 3 2 ° C a n d providing an appropriate p h o t o p e r i o d r e d u c e s the


n u m b e r of d a y s f r o m s o w i n g to flowering (Hellmars a n d Burton 1972).

Drought m a y hasten flowering in s o m e g e n o t y p e s but delay it in o t h e r s . A c o m b i n a t i o n of high


t e m p e r a t u r e a n d d r o u g h t affect pollen viability, stigma receptivity, a n d seed-setting.
Obovate Lanceolate Elliptical

Hexagonal Globular
F i g u r e 8. G r a i n s h a p e s in pearl millet.
( S o u r c e : Descriptors for pearl millet)

Style Pericarp
Starchy Cutin
Pericarp
Endosperm Epicarp
Corneous Mesocarp
Floury Cross cells
Peripheral Tube cells
Seed coat
Aleurone
Peripheral
endosperm
Germ
Scutellar Aleurone
epithelium Starch granules
Scutellum Protein bodies in
Embryonic axis protein matrix
Black layer Endosperm cells
Hilum

F i g u r e 9. Pearl millet grain structure.


( S o u r c e : R o o n e y a n d M c D o n o u g h 1987)

44
Selling and Crossing Techniques in Pearl Millet
K N Rai

T h e p r o t o g y n o u s flowering behavior of pearl millet m a k es it a highly cross-pollinated crop. E m e r g e n c e


of stigmas 2-3 d a y s before anther e m e r g e n c e m a k e s crossing possible without h a v i n g to resort to
e m a s c u l a t i o n . At the s a m e time, the presence of hermaphr odite flowers a n d stigma receptivity lasting 3-4
days m a k e selfing also easy. However, if a pearl millet panicle is not protected f r o m outside pollen, more
than 9 0 % of its s e e d are likely to be the products of cross-pollination. Selfing is done for producing the
s e e d of F 2 populations, S 1 /S 2 progenies, a n d inbred lines. Crossing is d o ne for pr oducing the s e e d of F 1
hybrids, full-sib/half-sib p r o g e n i e s, a n d open-pollinated varieties in a breeding nursery. A l t hough both
selfing a n d crossing in pearl millet require ordinary skills, careful planning is essential to e n s u r e that the
intended quantity of s e e d is p r o d u c e d a n d quality is maintained.

Selfing Plan

T h e most critical part of a selfing p r o g r a m in pearl millet is to plan in a d v a n c e for the requirement of
selfing b a g s as a n y panicle not b a g g e d before stigma e m e r g e n c e will be c o n t a m i n a t e d with outside
pollen a n d lose its identity. T h u s , selfing bags should be procured a n d kept in reserve o n e s e a s o n
(preferably two) before the s c h e d u l e d selfing s e a s o n . At I C R I S A T Asia Center (IAC), t w o types of selfing
bags, white p a r c h m e n t paper bags a n d brown kraft paper bags, are u s e d . T h e choice of bags d e p e n d s
on the cost a n d w e a t h e r considerations. For instance, kraft paper bags of 8 cm x 36 cm size m a d e in
India cost the equivalent of US $ 3.6 per 1000 bags. T h e cost of p a r c h m e n t paper bags of the s a m e size
m a d e in the UK is US $ 2 1 . 8 per 1000 bags. In the dry s e a s o n , kraft paper bags can be u s e d reliably
a n d cost-efficiently. In the rainy s e a s o n , however, they are not suitable as they a b s o r b m o r e water, take
longer to dry, a n d h e n c e are likely to fall apart, leading to contamination. At IAC, we use p a r c h m e n t
p a p e r selfing b a g s of three s t a n d a r d s i z e s — 8 cm x 36 c m , 8 cm x 46 c m , a n d 9 cm x 51 cm
— d e p e n d i n g on the size of the panicles to be selfed. For most purposes, the ideal size is 8 cm x 36 c m .

T h e best plant stage for selfing (bagging) is before stigma e m e r g e n c e . This is generally w h e n
about one-third of the panicle has e m e r g e d out of the boot. In s o m e genotypes, by this stage of panicle
e m e r g e n c e , s t i g m a e m e r g e n c e can already have occu rred. Care should be taken to b a g s u c h genotypes
as s o o n as the tip of the panicle is visible out of the boot. T h e b a g g e d panicles are stapled or clipped
in the p e d u n c l e r e g i o n . W i t h e x p e r i e n c e , o n e w o u l d be able to know h o w m u c h of the glued e n d of the
b a g s h o u l d r e m a i n a b o v e the tip of the panicle to ensure that the e m e r g i n g panicle does not pierce
t h r o u g h the b a g a n d create an o p e n i n g for outside pollen to contaminate it.

Generally, it is not necessary to have a selfing s y m b o l on the selfing b a g : a panicle under a


n o n m a r k e d b a g is c o n s i d e r e d as selfed. Selection for g o o d selfed seed-set is o n e of the selection criteria
in t h e inbred line d e v e l o p m e n t of pearl millet. T h u s , considering that the selfed seed-set is g o o d , selfing
of o n e panicle per plant will usually provide e n o u g h s e e d for use in p e d i g r e e b r e e d i n g , provided that the
panicle is > 1 0 cm l o n g . Selfing of 6-8 panicles of an F 1 hybrid can be e x p e c t e d to provide e n o u g h s e e d
to g r o w an F 2 population of 1000-2000 plants a n d still leave e n o u g h s e e d for replanting, if required.

C r o s s i n g Plan

For c r o s s i n g p u r p o s e s , white p a r c h m e n t paper bags are most convenient. T h e s e bags allow easy
detection of panicles that are ready to be u s e d as males a n d females without having to r e m o v e the bags.

45
T h e b a g g i n g p r o c e d u r e is the s a m e as in selfing, except that t h e b a g s s h o u l d be c l i p p e d , not s t a p l e d .
It is essential to c h o o s e the full-stigma-emergence stage of a b a g g e d panicle to be u s e d as a f e m a l e .
This e n s u r e s that o n c e s u c h panicles are c r o s s e d , no n e w stigmas will e m e r g e later that m a y get self-
pollinated.

F o r e n o o n is the best time for c r o s s i n g , for t w o reasons: (1) the d e w - s o a k e d bags b e c o m e more
transparent a n d h e n c e allow e a s y detection of the right panicle stage for use as f e m a l e s (whitish stigmas)
a n d m a l e s (yellowish pollen a c c u m u l a t e d at the b a s e of t h e panicles); a n d (2) t h e pollen viability is high
at this time of the day, especially before 1100 h. Howev er , experience s h o w s that if the m a x i m u m day
t e m p e r a t u r e d o e s not e x c e e d 3 8 ° C , crossing can be d o n e in the afternoon as well, although it results in
a small reduction in seed-set.

T h e a m o u n t of pollen required to be collected for crossing d e p e n d s on the n u m b e r of f e m a l e panicles


available. T h u s , the first task in a crossing operation is to find out the n u m b e r of f e m a l e panicles with fully
e m e r g e d s t i g m a s . This is most convenient to detect in d e w - s o a k e d panicles, usually before 0 9 0 0 h w h e n
the p a r c h m e n t p a p e r b a g s are m o s t transparent. This can usually be a c c o m p l i s h e d without r e m o v i n g the
b a g s . H o w e v e r , after a panicle has b e en tentatively identified as ready to be u s e d as a f e m a l e , its
flowering s t a g e must be c o n f i r m e d by quickly pushing up the b a g , observing t h e entire panicle for full
stigma e m e r g e n c e for no m o r e than a s e c o n d , a n d covering the panicle a g a i n . O n c e a f e m a l e panicle
has b e e n s e l e c t e d , the clip s hould be placed on the glued upper e n d of the b a g to distinguish it f r o m
others for u s e in cross-pollination.

As the d e w dries up, the situation improves for pollen collection. A c c u m u l a t i o n of pollen (yellowish
p o w d e r ) at t h e b a s e of t h e b a g g e d panicles indicates that they are ready for pollen collection. First, the
b a g is held tightly a b o v e the point of pollen a c c u m u l a t i o n , a n d the clip is r e m o v e d to release the
a c c u m u l a t e d pollen at the b a s e . T h e n , by still holding the bag tightly, the panicle is t a p p e d with the other
h a n d , c a u s i n g the s h e d d i n g a n d a c c u m u l a t i o n of fresh pollen at the b a s e . This pollen is m a d e to slip
t o w a r d s the g l u e d e n d of the b a g by b e n d i n g the panicle a n d peduncle d o w n w a r d s . Thereafter, the
panicle is r e b a g g e d with the s a m e or another selfing b a g if more pollen needs to be collected f r om it on
the following d a y or if the selfed s e e d needs to be harvested from it to maintain the line. W h e r e selfed
s e e d is not required f r o m the b a g g e d panicles, t h e s e c a n be b a g g e d at the beginning of anthesis, a day
prior to pollen collection. T h e s e b a g s should not be clipped, to distinguish t h e m f r o m those that have
b e e n c l i p p e d for production of selfed s e e d . Pollen f r o m several b a g g e d panicles of an entry m a y be
collected in o n e b a g , w h i c h shall bear the plot or entry number.

T h e pollen collected is d u s t e d onto the female panicles after quickly r e m o v i n g the selfing bags
f r o m the latter. O n c e crossing is c o m p l e t e d , the bags are m a r k e d with (x) plot or entry n u m b e r of the
pollen parent (Table 1), a n d p l a c e d b a c k over the c r o s s e d panicle a n d s t a p l e d . T h i s c o m p l e t e s the
crossing p r o c e d u r e .

O c c a s i o n a l l y , a portion of t h e panicle (usually the lower one) m a y not be fully pollinated d u e to


insufficient pollen or the stigmas in this portion might not have fully e m e r g e d , giving s c o p e for self-
pollination at t h e later stage of flowering. Spikelets from s u c h portions of panicles s h o u l d be r e m o v e d
before r e b a g g i n g t h e c r o s s e d panicles. If bulk pollen of the s a m e line or population has b e e n u s e d for
crossing to p r o d u c e the s e e d , t h e n a simple (#) mark on the c r o s s e d f e m a l e panicle will be sufficient
(Table 1). If o n e plant is c r o s s e d with another plant, as in the c a s e of full-sibs, then t h e plants u s e d as
m a l e s a s w e l l a s t h o s e u s e d a s f e m a l e s s h o u l d b e n u m b e r e d , a n d these n u m b e r s s h o u l d b e r e c o r d e d
on the c r o s s e d f e m a l e panicle (e.g., P 5 x P 107).

46
In a selfing p r o g r a m , it is possible to do a quick visual a s s e s s m e n t of the a p p r o x i m a t e n u m b e r
of selfed panicles to determine if the program is on target. This is difficult to do w h e n c r o s s e d s e e d is
to be p r o d u c e d , especially w h e n the s a m e line is u s e d as a f e m a l e for crossing with a n u m b e r of m a l e
lines. T h e s a m e thing applies w h e n a large n u m b e r of crosses are to be m a d e within a population to
p r o d u c e full-sibs or half-sibs. In s u c h situations, maintaining a crossing record b e c o m e s necessary. This
record is essentially a list of the f e m a l e plots against which are written all the male plots with w h i c h they
s h o u l d be c r o s s e d (Fig. 1). During the crossing p r o g r a m , the n u m b e r of panicles c r o s s e d of e a c h
c o m b i n a t i o n s h o u l d be recorded in the appropriate c o l u m n to monitor the progress of crossing a n d
a c h i e v e m e n t of t h e target. This helps e n s u r e that only the required crosses are m a d e .

Harvesting of Selfed/Crossed Panicles

S e l f e d / c r o s s e d panicles c a n be harvested at physiological maturity after the formation of the black layer.
It is, h o w e v e r , a d v i s a b le to harvest w h e n the s e e d s are dry as this necessitates little post harvest d r y i n g .
S o m e d e g r ee of p o s t h a r v est drying is unavoidable to ensure that even late-maturing panicles, which
might not h a v e d r i e d at harvest time, b e c o m e dry for threshing. Wherever possible, about a w e e k before
harvesting, the upper glued ends of the bags are torn off a n d p u s h e d d o w n the selfed panicle so that it
is e x p o s e d for s o m e time to facilitate natural clearing of the sticking anthers. T h i s permits harvesting of
clean panicles, a n d also m a k e s it possible to evaluate t h e m for s e e d characters (including seed-set).

In the c a s e of c r o s s e d panicles, the bags are left intact. Harvested panicles f r o m a plot (line or
population) are collected in a harvesting b a g , w h i c h s h o u l d bear the plot number. Both self ed a n d
c r o s s e d panicles h a r v e s t e d f r o m a plot can be collected in the s a m e b a g , w h i c h s h o u l d be s t a p l e d .

T h r e s h i n g a n d Storage

After d r y i n g , selfed panicles are separated f r o m the c r o s s e d panicles a n d t h r e s h e d individually (in


pedigree selection) or as a bulk (in bulk pedigree selection). T h e c r o s s e d panicles of a plot a re t h r e s h e d
as a bulk if they w e r e all c r o s s e d with only o n e male parent. T h e y n e e d sorting out if t h e y w e r e c r o s s e d
with m o r e t h a n o n e m a l e parent. T h e s e panicles are t h r e s h e d by h a n d or by a m a c h i n e . T h e s e e d s are
then c l e a n e d of g l u m e s in a tray a n d transferred to s e e d packets, bearing the plot n u m b e r for s e e d s f r o m
selfed panicles a n d "plot x plot" n u m b e r for s e e d s f r o m crossed panicles. Pieces of n a p h t h a l e n e balls
are put in t h e s e e d packets to avoid d a m a g e f r om pests. T h e s e s e e d s c a n be s t o r e d at r o o m
t e m p e r a t u r e for a y e a r without a n y serious loss of viability.

47
T a b l e 1 . A n illustration o f t h e s y m b o l s u s e d for v a r i o u s t y p e s o f c r o s s e s i n p earl m illet.

Cross type Symbols

Line x line 81 B x J 104 (for entries)


2 0 6 x 701 (for plots)

Plant x p o p u l a t i o n P... x E B C (different populations)


# ( s a m e population)

Line x p o p u l a t i o n 81 B x E B C (entry x population)


2 0 6 x E B C (plot x population)

Population x population NC x E B C (for entries)


805 x 9 2 0 (for plots)

Plant x p l a n t P 5 x P 107 ( s a m e population)


NC (P 5) x E B C (P 210) (different populations)

N u m b e r of c r o s s e d panicles

Female M a l e plot (number)


plot
(number) 86 180 210 275 380 400

5 i ii iii ii iiii iiii

17 iiii ii ii

210 i

285 iiii

362 iiii

390 iiii iiii iiii iiii iiii iiii

451 iii iiii

480 iiii iiii iiii iiii iiii iiii

Figure 1. Recording progress in crossing work with a target


o f four c r o s s e d p a n i c l e s per c o m b i n a t i o n .
Development, Production, and Maintenance of Male-Sterile Lines
in Pearl Millet
K N Rai

T h e w i d e s p r e a d cultivation of single-cross hybrids of pearl millet in India reflects the great s u c c e s s of


cultivar d e v e l o p m e n t a n d s e e d production technology. T h e s e hybrids are p r o d u c e d by c r o s s i n g restorer
lines o n t o cytoplasmic-genic male-sterile lines, also called cytoplasmic male-sterile ( C M S ) lines. T h e C M S
s y s t e m theoretically provides a m e c h a n i s m to p r oduce pure single-cross hybrid s e e d on a c o m m e r c i a l
scale (under o p e n pollination in isolation). In practice, however, several genetic a n d nongenetic factors
d e t e r m i n e t h e s u c c e s s of this s y s t e m . T h e t w o most important genetic factors are: (1) the stability of male
sterility under various environmental conditions, a n d (2) the availability of marker(s) for the detection of
off-types. T h e nongenetic factors are various aspects of s e e d production a n d s e e d m a i n t e n a n c e ,
including purification of s e e d stock. Besides the question of the genetic purity of hybrid s e e d s , there is,
of c o u r s e , the related question of the e c o n o m i c feasibility of hybrid s e e d production.

This paper provides a brief description of the nature a n d sources of C M S a n d d e v e l o p m e n t of


male-sterile lines, a n d a s o m e w h a t detailed account of production a n d m aintenance of nucleus/breeder
s e e d of male-sterile lines a n d their respective maintainer lines.

C y t o p l a s m i c - G e n i c Male Sterility

C y t o p l a s m i c - g e n i c m a l e sterility results f r o m an interaction b e t w e e n cytoplasmic factors a n d nuclear


g e n e s (Fig. 1). It is n o w k n o w n that male-sterility-inducing cytoplasmic factors are mitochondrial g e n e s ;
h e n c e it w o u l d be m o r e appropriate to call this type of male sterility cytoplasmic-nuclear m a l e sterility
( H a n n a 1989).

Identification of C M S . Male-sterile plants are easily identifiable at the time of anthesis by their shrivelled
anthers, w h i c h do not s h e d pollen. T h e sterility c ould be due to genetic factor(s) in the nucleus alone,
in w h i c h c a s e it is called genetic male sterility ( G M S ) . W h e n G M S plants are c r o s s e d with a w i d e range
of inbreds, they never p r o d u c e hybrids in w h i c h all or most of the plants are male sterile. In contrast,
w h e n C M S plants are c r o s s e d with a set of s u c h inbreds, they produce s o m e hybrids in w h i c h all or m o s t
of the plants are m a l e sterile. This is the sole criterion to distinguish C M S from G M S .

C M S s o u r c e s . Several well-classified C M S sources are available in pearl millet. T h e s e are called A 1 , A 2 ,


A 3 (Burton a n d A t h w a l 1967), a n d A 4 ( H a n n a 1989). Besides these, there are others that h a v e not yet
b e e n classified well (Rai a n d S i n g h 1987). At present, however, only the A, system is under c o m m e r c i a l
utilization. T h e r e are three m a i n reasons for it: (1) the male sterility of this s y s t e m is m o r e stable than
others e x c e p t A 4 ; (2) several inbred lines identified as restorers of the A, source produce sterile hybrids
on the A 4 s o u r c e a n d h e n c e are unsuitable for the production of grain hybrids on the A 4 source; a n d (3)
most of the breeders are presently contented with the male-sterile lines b a s e d on the A, source.

G e n e t i c s of C M S . An oversimplified m o d e l a s s u m e s that the cytoplasmic factor interacts with a single


recessive nuclear g e n e in the h o m o z y g o u s (ms 1 ms 1 ) condition (Fig. 1) to p r o d u c e the A, type of male
sterility (Burton a n d A t h w a l 1967). T h e r e are indications, however, that more than o n e major g e n e might
be involved (Siebert 1982) a n d in fact, s o m e modifiers might be involved as well (Rai a n d Singh 1987;
Rai a n d H a s h 1990).

49
s

1. Male-sterile plant/line

or
F F

2. Male-fertile plant/line

S =Sterile cytoplasm, F=Fertile cytoplasm;


MS=Dominant nuclear gene for male fertility;
ms=Recessive nuclear gene for male sterility.

Figure 1 . Interaction b e t w e e n c y t o p l a s m a n d nuclear


g e n e s c a u s i n g m a l e sterility in pearl millet.

50
T h e nuclear g e n e s that interact with the cytoplasmic factors to produce male sterility in the A 2 , a n d
A 3 c y t o p l a s m s are s u g g e s t e d to be different from that of the A, cytoplasm (Burton a n d Athwal 1967). T h e
genetics of the other C M S systems has not yet been investigated.

D e v e l o p m e n t of Male-Sterile Lines (A-lines)

T h e first step in the d e v e l o p m e n t of an A-line is to breed a g o o d maintainer line (B-line). O n c e this has
b e e n a c h i e v e d , an A-line is d e v e l o p e d simply by backcross transfer of the nuclear g e n o m e of the B-line
into the sterility-inducing cytoplasm of an A-line.

Maintainer line (B-line). A maintainer line is an inbred line that produces a sterile hybrid w h e n c r o s s e d
on a male-sterile line (A-line). T h u s , B-lines are simply identified by crossing t h e m on an A-line a n d
identifying sterile hybrids. N e w A-lines are dev eloped by backcrossing these B-lines into the cytoplasm
of an A-line.

H o w e v e r , just b e c a u s e an inbred line has proved to be a B-line, it does not b e c o m e a candidate


entry for c o n v e r s i o n into a commercially viable A-line. The B-line should have several traits to m a k e it
worthy of c o n v e r s i o n into an A-line. S o m e of the essential traits include: high s e e d y ield, g o o d lodging
resistance, g o o d pollen production a n d stigma receptivity, maturity appropriate to the target environment,
resistance to pests a n d diseases (e.g., d o w n y mildew), a n d g o o d c o m b i n i n g ability. S o m e of the desirable
traits include: short to m e d i u m plant height, synchronous tillering, m e d i u m to large s e e d size, grain color
as per c o n s u m e r s ' preference, a n d morphological marker(s). O n c e the requirement of all the essential
traits is satisfied, a n d s o m e or m a n y of the desirable traits are also present in a B-line, it m a k e s a g o o d
candidate entry for conversion into an A-line.

C o n v e r s i o n of a B-line into an A-line. A highly inbred B-line, w h e n c r o s s e d o n t o an A-line, will produce


an F 1 hybrid in w h i c h all the plants will be fully male sterile. T h e backcrossing of this sterile F 1 to the B-
line will p r o d u c e a BC 1 p r o g e n y in w h i ch all the plants will again be fully male sterile. In s u c h a situation,
the s u b s e q u e n t b a ckcrosses will produce a d v a n c e d BC progenies w h i c h will carry only male-sterile
plants. If the B-line is highly uniform for all morphological traits, bulk pollen c a n be u s e d f r o m the B-line
during the b a c k c r o s s i n g p r o g r a m (Fig. 2).

If t h e B-line is variable for a trait (including genetic variation for sterility-maintenance ability),
selection of individual plants for (Plant x Plant) crossing during the ba ckcross program is resorted to (Fig.
3). Generally 3-5 plants f r o m e a c h of the B-lines a n d the corres p o n d i n g BC p r o g e n y are a d e q u a t e . This
requires s o w i n g of e a c h selfed progeny of the B-line a n d its corresponding BC progeny in paired plots,
e a c h in a single row of 2 m, consisting of 15-20 plants. At e a c h BC generation, selection is m a d e both
b e t w e e n - a n d w i t h i n - t h e B C p r o g e n y rows a s well a s b e t w e e n - a n d w i t h i n - t h e B-line rows t o ensure that
the n u m b e r s do not e x p l o d e a n d that the B-line is further i m p r o v e d .

Production a n d Maintenance of Nucleus Breeder Seed

An efficient n u c l e u s / b r e e d e r - s e e d production technology has four major c o m p o n e n t s :

• m a x i m i z a t i o n of s e e d yield per unit a r e a ;

• reduction of s e e d cost;

• timely availability of s e e d ; a n d

51
A-line B-line D o n o r genes ( % )

F1 B-line 50.0

BC1 B-line 75.0

BC2 B-line 87.5

BC3 B-line 93.8

BC4 B-line 96.9

BC5 B-line 98.4

( ) % B-line 99.2

BC7 B-line 99.6

BC8 B-line 99.8

() 9 B-line 99.9

Figure 2. Conversion of a highly inbred B-line into an A-line.

52
A-Iine Partial inbred B-Iine
Partial inbred B-line
F1
S P1
S P2
F
S P3

S P4
F

B C 1 (P1) P1 B C (P2) P2 BC1 ( P 4 ) P4

S P1 S S P1
S P2 F S P2
F F S P3
S P3 S S P4

S P4 F S P5

Rejected

BC2(P1) P1 BC2 ( P 4 ) P4 BC2 ( P 1 ) P1 BC2 ( P 5 ) P5


S S S P1 S P1
F S S
S P2
S S P2
S S P3
S S
S S P4
S S P3
S S P5
F S
Rejected Rejected

BC3(P1) P1 BC3(P3) P3 BC3 (P1) P1 BC3(P4) P4

S S S S
S S S S
S S S S
S S S S
S S S S
- Proceed f u r t h e r w i t h any selected pair.
- D r o p P x P crossing p r o g r a m
- R e s o r t to L x L crossing/backcrossing
(S = Sterile; F = Fertile)

F i g u r e 3. C o n v e r s i o n of a partial inbred B-line into an A-line.

53
• highest quality of s e e d in terms of genetic purity, s e e d vigor, a n d germinability, free of u n w a n t e d
materials.

T h e s e objectives c a n be achieved by observing the following practices.

Site s e l e c t i o n . Pearl millet is highly sensitive to w a t e r l o g g i n g . T h e r e f o r e , s e e d p r o d u c t i o n of this crop


requires well-levelled l a n d with g o o d d r ainage. Improved genotypes of pearl millet have b e e n s h o w n to
r e s p o n d to a p p l i e d nitrogen levels of up to 100 kg ha - 1 . T h u s , selecting a fertile field or g r o w i n g the s e e d
crop under high levels of applied fertilizer is r e c o m m e n d e d . However, growing the s e e d crop on d e e p
soils s h o u l d be a v o i d e d , particularly in the rainy s e a s o n . T h e field s h o u l d h a v e a c c e s s to irrigation
facilities e v e n in the rainy s e a s o n , a n d s h o u l d not h a v e b e e n s o w n with pearl millet in the prec eding
season.

S e l e c t i o n of s e a s o n . Pearl millet is basically a rainy-season crop. It c a n , however, be successfully


g r o w n in t h e dry s e a s o n with irrigation, provided the m i n i m u m temperature does not fall below 12°C a n d
the m a x i m u m t e m p e r a t u r e d o e s not e x c e e d 3 8 ° C . In fact, higher grain yields h a v e b e e n reported from
d r y - s e a s o n c r o p s t h a n f r o m rainy-season crops in India. This is due to longer s u n s h i n e hours, less
lodging, less risk of pollen w a s h a n d hence better seed-set, better uptake of nutrients d u e to g o o d
m o i s t u r e control t h r o u gh irrigation, a n d fewer disease a n d insect pest p r o b l e m s . Moreover, dry-season
crops p r o d u c e better quality s e e d that is disease-free a n d has an attractive lustre.

It is a d v i s a b l e , w h e r e v e r possible, to produce both nucleus a n d breeder s e e d in the dry s e a s o n


b e c a u s e b e i n g t h e off-season, there is greater likelihood of getting g o o d isolation. T h e rain-free period
in this s e a s o n facilitates m u c h better control over roguing operations. Both g o o d isolation distance a n d
roguing are essential to maint ain the genetic purity of male-sterile lines. Also , the longevity of s e e d s
p r o d u c e d in the dry s e a s o n is better than those p r o d u c e d in the rainy s e a s o n .

Isolation d i s t a n c e . T h e p r o t o g y n o u s flowering behavior of pearl millet puts it at a greater risk of


c o n t a m i n a t i o n f r o m w i n d b o r n e alien pollen than maize a n d s o r g h u m . Experimental data for det erm ining
t h e isolation distance for s e e d production plots of pearl millet continue to be a major c o n c e r n (Chopra
1982). For pearl millet, the distance b e t w e e n the s e e d production plot of the intended genot y p e a n d t hose
of others has b e e n arbitrarily f i x e d ; it varies greatly for various s e e d classes. T h e isolation distance
r e c o m m e n d e d for n u c leus-seed production plots is at least 2 k m , a n d for b r eeder-seed production plots
it is m o r e t h a n 1 km (Andrews a n d Harinarayana 1984). S e e d s of A- a n d B-lines can be p r o d u c e d in the
s a m e s e e d production plot, provided utmost care is taken in harvesting these lines. Alternatively, they
c a n also be p r o d u c e d in different plots with an isolation distance of 5 m b e t w e e n the t w o .

S e v e r a l factors n e e d to be taken into account while fixing the isolation distance. T h e s e include
w i n d direction a n d velocity, humidity, windbreaks (including the intervening fields s o w n with other
species), a n d the size of the plots serving as the source of alien pollen load. S o m e t i m e s it is a r g u e d that
if the s e e d p r o d u c t i on plot of the i n t e n d e d genotype is u p w i n d , the isolation distance requirement for it
m a y be slightly relaxed. It is, however, not always certain before s o w i n g w h a t the w i n d direction will be
at the time of flowering a n d whether or not it will always be unidirectional.

T h e r e c o m m e n d e d isolation requirements given above apply not only to field s o w i n g s of other


g e n o t y p e s but also to v o l u n t eers, although the alien pollen load f r o m the latter will be of m u c h less
c o n s e q u e n c e . T h e p r o b l e m of isolation distance can be s o l v e d to a great d e g r e e by resorting to the "seed
village" a p p r o a c h as practised in India (Chopra 1982). This a p p r o a c h w o u l d require all farmers of a
village t o p r o d u c e s e e d o f t h e s a m e g e n o t y p e . T h e " s e e d village" a p p r o a c h also m a k e s the s e e d

54
certification e x e r c i s e m o r e convenient a n d less expensive.

Direct s o w i n g vs t r a n s p l a n t i n g . Direct sowing is a more widespread practice for the production of all
classes of s e e d s . However, in certain areas, transplanting of 18-20-day-old seedlings f r o m nurseries is
preferred to direct s o w i n g b e c a u s e :

• the field might be o c c u p i e d by another crop at the appropriate s o w i n g time;

• transplanting saves expenditure on weeding a n d irrigation water w h i c h w o u l d be incurred for the


initial 18-20 d a y s in the direct-sown crop;

• transplanting requires only about 3 0 - 4 0 % of the seed quantity required for direct s o w i n g ;

• a t r a n s p l a n t e d crop has a m u c h higher establishment success rate than the direct-sown c r o p ,


without a n y n e e d for thinning; a n d

• a t r a n s p l a n t e d crop is believed to yield more than the direct-sown crop b e c a u s e unhealthy


seedlings are discarded a n d there is a m u c h better control over plant s p a c i n g .

U n d e r both s o w i n g m e t h o d s , it is r e c o m m e n d e d to maintain an interrow spacing of 60-75 cm a n d


an intrarow s p a c i n g of 15-20 cm in the nucleus- and breeder-seed production plots. T h e lower plant
population attained with these spacings will allow full plant expression and facilitate easy identification
of off-type plants a n d deviant phenotypes, and leave adequate room for undertaking intensive roguing.
T h e wider s p a c i n g also w o r k s in favor of achieving a high seed-multiplication rate (i.e., ratio of s e e d
s o w n to s e e d harvested).

To a v o i d s o w i n g errors in the s e e d production of all s e e d classes, it is r e c o m m e n d e d that the


direct s o w i n g or transplanting of all rows of the A-line is completed before beginning t h e s o w i n g or
transplanting of the B-line. In case of machine s o w i n g , both the A-line a n d B-line have to be a n d c a n be
s o w n simultaneously with a negligible c hance of errors.

A-line/B-line ratio. There is little literature on the o p t i m um A-line/B-line ratio for pearl millet. Of c o u r s e ,
m u c h w o u l d d e p e n d on the pollen-producing ability of the B-line a n d the row-to-row s p a c i n g . T h e
r e c o m m e n d e d ratio is four r o w s of the A-line to two rows of the B-line (Khairwal et a l . 1990), but it c a n
be six rows of the A-line to two rows of the B-line, if the latter is a prolific pollen producer. T h e s e e d -
production s e a s o n m a y also affect the o p t i m u m A-line/B-line ratio. For instance, the greater risk of pollen
w a s h a n d ergot (a panicle disease) in the rainy s e a s o n w o u l d be a c a s e for a closer A-line/B-line ratio
t h a n for a d r y - s e a s o n c r o p . It is a c o m m o n practice to s o w 4-8 rows of the B-line on all four sides of the
s e e d - p r o d u c t i o n plot (Fig. 4). T h e s e borders serve not only as an additional source of pollen for better
s e e d - s e t in the A-line rows but also as a trap for alien pollen f r o m outside sources. For easy identification
of A-line r o w s f r o m B-line rows which are morphologically similar (except for the difference in pollen
s h e d d i n g , w h i c h is discernible only during flowering), it is advisable to maintain a record of t h e s o w i n g
plan of A-line a n d B-line rows (Fig. 4) till the e n d of harvesting. Alternatively, all B-line rows are m a r k e d
with stakes at o n e or b o t h e n d s , or live-marked with plants of other species (e.g., s u n n h e m p a n d
sunflower) with s o m e convenient spacing within the rows.

A-line/B-line s y n c h r o n i z a t i o n . In pearl millet, A-lines flower about 2 days earlier than t h e c o r r e s p o n d i n g


B-lines. If the A-line has a longer duration of stigma receptivity (>3 days), no synchronization problem
is likely to occur in t h e s e e d production of A-lines.

55
A-line B-line

F i g u r e 4. S o w i n g pattern of A- a n d B-lines for s e e d p r o d u c t i o n


in pearl millet.
will essentially be identical to the maintainer line, w h i c h w o u l d have no effect on the purity of the seeds
harvested f r o m the A-line, provided they are not included in the harvest. Fortunately, the majority of the
pollen s h e d d e r s f o u n d in pearl millet have b e en reported to arise from mutations in the c y t o p l a s m (Burton
1977).

Type of Cytoplasmic-nuclear
pollen shedder constitution Origin

Type 1
* Cytoplasmic mutation
Identical to F
B-line * Mixture of B-line s e e d

Type 2

Different from S
B-line * Nuclear mutation

Figure 5. Origin of pollen s h e d d e r s in t h e A-line.

Pollen s h e d d e r s of nuclear origin have a different implication in maintaining the genetic purity of
both the A-line a n d B-line. T h e only w a y to remove this type of impurity from the nucleus s e e d is to
undertake plant x plant (P x P) crossing b e t w e e n the A-line and the B-line, followed by one backcrossing
to the progenies of the B-line, with a view to discarding those crosses a n d backcrosses that are fertile,
a n d bulk the s e e d of those backcross families which are uniformly sterile to reconstitute the A-line. T h e
bulking of the c o r r e sp o n d i n g B-line families w o u l d reconstitute the B-line. Our experience with two diverse
male-sterile lines having a low proportion of pollen s hedders indicates that the fr equency of pollen
shedders is higher in the rainy season than in the postrainy s e a s o n . T h u s , the evaluation of P x P
crosses a n d b a c k c r o s s e s for fertility/sterility will be most effective in the rainy s e a s o n .

Since anthesis starts in the night, morning hours are the best time for roguing pollen shedders
in the A-line. C o o p e r a n d Burton (1965) have s h o w n that w h e n stored at 26.7°C for up to 24 hours, the
loss of pollen viability in pearl millet is less than 5 0 % . T h u s , for this reason too, m o r n i n g is the safest time
for pearl millet field activity not only for roguing pollen shedders a n d off-type plants but also for avoiding

57
contamination of seed-production plots with pollen carried inadvertently f r o m other br eeding fields visited
earlier.

In the case of nucleus- and breeder-seed production, roguing is required to be m o r e stringent.


It involves the removal of e v e n those plants appearing to be phenotypic deviants of the target g e n o t y p e .
It s h o u l d be n o t e d that the highly cross-pollinated nature of pearl millet provides a potential m e c h a n i s m
for small c h a n g e s in various plants to c o m e together a n d c a u s e , after recombination, genetic shifts in the
parental lines, if the pressure of roguing at the nucleus- and b r eeder-seed level is relaxed.

H a r v e s t i n g , t h r e s h i n g , a n d g r a d i n g . The B-line sh o u l d b e harvested first, a n d the b a g s / h e a p s s h o u l d


be double-labelled (one label inside the bag a n d another outside) a n d stored, if possible, separately. T h e
labels s h o u l d h a v e on t h e m the n a m e of the line, the field, a n d the crop s e a s o n . T h e field s h o u l d be
c h e c k e d carefully to e n s u r e that no panicles of the B-line are left in the s e e d plot. After c o m p l e t i n g these
practices, the harvest of the A-line c a n be taken up. Its panicle b a g s / h e a p s s h o u l d also be d o u b l e -
labelled a n d stored separately. T h e panicles s hould be dried well d o w n to 1 2 % moisture before they are
t a k e n up for t h r e s h i n g .

T h r e s h i n g a n d processing of the A-line a n d B-line seeds should be d o ne separately. Processing


includes r e m o v i n g s e e d s of other crops a n d w e e d s , a n d inert matter f r o m the seed-lot; a n d gr ading it to
the s t a n d a r d size typical of the g e n o t y p e , b a s e d mainly on the s e e d size a n d s h a p e .

S e e d t r e a t m e n t a n d s t o r a g e . T h e seeds should be treated with a mixture of fungicide a n d insecticide


to disinfect a n d protect t h e m f r o m s e e d b o r ne a n d soilborne pathogenic organi s m s a n d storage pests
(Khairwal et al. 1990). Application of thiram 7 5 % W D P @ 85 g for 100 kg - 1 s e e d is a practice followed
by the National S e e d s Corporation in India. T h e treated s e e d is stored in standard containers that are
s e a l e d a n d labelled both inside a n d outside. The label inside should have on it the n a m e of the crop, the
n a m e of the variety, the class of s e e d , the s e a s o n , and the field/place w h e r e the s e e d w a s p r o d u c e d .
T h e label outside s h o u l d h a v e additional details s u c h as s e e d purity, germ ination p e r c e n t a g e , etc. Sealed
containers of the nucleus s e e d are stored at 4 ° C under 2 0 % relative humidity. T h e breeder s e e d is stored
at 10°C under 2 0 % relative humidity.

F o r e c a s t i n g l a n d a n d s e e d r e q u i r e m e n t s . T h e a m o u n t o f s e e d available in a s e e d class is the principal


factor d e t e r m i n i n g the a m o u n t of s e e d one c a n expect to produce in a s u b s e q u e n t s e e d class. T h u s , an
efficient s e e d - p r o d u c t i o n t e c hnology should be able to forecast the s e e d a n d land requirements for
various s e e d classes, a n d plan the seed-production program accordingly.

B a s e d on 3 kg ha - 1 s e e d rate, 6:2 ratio of A-line a n d B-line (R-line) rows, a n d s e e d yields of


certified-, f o u n d a t i o n - , a n d breeder-seed-production plots @ 8 0 0 , 6 0 0 , a n d 4 0 0 kg ha - 1 respectively, the
land a n d s e e d requirements for a target area of 100 0 0 0 ha under hybrid are given in Table 1. Even with
the c o n s e r v a t i v e s e e d yield a s s u m e d in this calculation, just 45 g of A-line s e e d , 15 g of B-line s e e d , a n d
about 2 0 0 m 2 land for the A-line breeder-seed production plot is sufficient to m e e t the requirement of 100
0 0 0 ha under hybrid (Table 1). This extraordinarily explosive multiplication rate of v a r i o us classes of
s e e d s of pearl millet m e a n s that 1 kg of A-line nucleus s e e d (adequate for s o w i n g 0.3 ha of a breeder-
s e e d plot) is e n o u g h to p r o d u c e m o r e than 7 0 0 0 tons of certified hybrid s e e d , w h i c h is a d e q u a t e for
s o w i n g as m u c h as 2.3 million ha (i.e., about 2 0 % of the total pearl millet a c r e a g e in India).

N u c l e u s s e e d is the basis for all s e e d classes; h e n c e , it is e x p e c t e d to be t h e purest s e e d class.


This s e e d class is the only o n e that regenerates f r o m itself a n d needs utmost personal attention f r o m the
b r e e d e r ( A n d r e w s a n d H a r i n a r a y a na 1984). It is r e c o m m e n d e d that e a c h multiplication of the nucleus

58
s e e d be d o n e with e n o u g h quantity to meet the d e m a n d for breeder s e e d for five y e a r s , retaining a
portion of the s e e d f r o m e a c h multiplication for long-term storage as b a c k u p stock (Fig. 6). B a c k u p stocks
c a n be u s e d for monitoring genetic c hanges in the nucleus s e e d so that timely purification m e a s u r e s can
be t a k e n , if required. It is also advisable to keep backup stocks in a different store (perhaps a long-term
store, if available) to a v o i d loss due to natural or h u m a n factors.

A n d r e w s a n d Har inar ay an a (1984) have s u g g e s t e d that a nucleus s e e d - p r o d u c t i o n p r o g r a m


s h o u l d not be t a k e n up on an a r e a e x c e e d i n g 0.2 ha b e c a u s e this class of s e e d requires t h e closest
attention of the breeder a n d his staff, w h o need to scrutinize individual plants at various stages of growth
a n d d e v e l o p m e n t . T h e extent of nuc leus-seed production, however, w o u l d d e p e n d on the size of the
breeder's t e a m . C h o p r a (1982) has s u g g e s t e d planning for production of e n o u g h br eeder s e e d in one
attempt to m e e t the f o u n d a t i o n - s e e d r equirement for 4-5 years. S u c h a strategy w o u l d d e p e n d on the
availability of st orage a n d isolation facilities. In any c a s e , the breeder s h o u l d p r o d u c e e n o u g h breeder
s e e d in o n e attempt to m e e t the foundation-seed requirements for at least two y e a r s .

T a b l e 1 . L a n d a n d s e e d r e q u i r e m e n t s for v a r i o u s s e e d classes for a target area of > 1 0 0 0 0 0 ha


under pearl millet h y b r i d . Certified h y b r i d -seed requirement: 100 0 0 0 ha x 3 kg ha - 1 = 3 0 0 0 0 0 k g .

A B R

Certified hybrid-seed p r o d u c t i o n plot (6A:2R)

S e e d yield: 8 0 0 kg ha - 1
L a n d requirement (ha) 400 - 140
S e e d requirement (kg) 1200 - 420

Foundation-seed p r o d u c t i o n plot (6A:2B)

S e e d yield: 6 0 0 kg ha - 1
L a n d r equirement (ha) 2 0.7 0.7
S e e d requirement (kg) 6 2 2

Breeder-seed production plot (6A:2B)

S e e d yield: 4 0 0 k g ha - 1
L a n d requirement (ha) 0.015 0.005 0.005
S e e d r e q u i r e m e n t (g) 45 15 15

A = M a l e s t e r i l e , B = M a i n t a i n o r , a n d R = R e s t o r e r to p a r e n t .

59
A,B,R Lines A,B,R Lines A,R Lines Hybrid

Nucleus

Nucleus Yr = year

Backup Stock

F i g u r e 6. S e e d - p r o d u c t i o n s t a g e s for parental lines a n d


h y b r i d in p ear l millet.
(Modified f r o m A n d r e w s a n d Harinarayana 1984)

60
Development, Production, and Maintenance of Pollinators in Pearl
Millet
B S Talukdar

Pollinators a r e m a l e parents of hybrids. They are bred to impart to the hybrid high grain yield, resistance
to d o w n y mildew, a n d adaptability to a wide range of environments.

Procedure for Breeding Pearl Millet Pollinators

Pearl millet, being a cross-pollinated crop ( > 8 5 % outcrossing), is available in the natural condition as a
r a n d o m - m a t i n g population. Inbred lines can be derived by selfing. Pollinator lines are derived thr ough
repeated selfing of plants in an open-pollinated population, random-mating bulk or F 1 s d e r i v e d t h r o u gh
planned crossing.

B r e e d i n g the inbred pollinator for a single-cross hybrid involves seven to ten generations of
selfing. In the p r o c e s s , most inbred lines b e c o m e uniform in terms of time to 5 0 % flowering, plant height,
panicle size, panicle s h a p e , etc. Evaluation a n d selection for downy-mildew resistance begins as early
as at F 3 (Fig. 1).

T h r e e types of breeding s c h e m e s are generally followed:

1. Early-generation testing
2. A d v a n c e d - g e n e r a t i o n testing
3. C o m b i n a t i o n of 1 a n d 2

In early-generation testing, fertility restoration and combining ability are ev aluat ed at the F 4 stage.
S e l e c t e d F 5 families are evaluated in the field for downy mildew (DM) resistance a n d are also selfed to
p r o d u c e F 6 lines in the DM nursery.

Superior specific combiners are identified at the F 6 stage in the pearl millet a d v a n c e d hybrid trial
( b o r d e r e d r o w plot, multilocational trials). Nicking a n d uniformity tests are c o n d u c t e d at this stage in the
dry s e a s o n at I C R I S A T Asia Center, Patancheru.

Hybrids multiplied f r o m the F 7 lines are entered in the All India Coordinated Pearl Millet
I m p r o v e m e n t Project ( A I C P M I P ) trial a n d pollinators are entered in the Pollinator Collection.

In a d v a n c e d - g e n e r a t i o n testing, individual F 3 s are bulk a d v a n c e d to F 4 . Individual plant progenies


are g e n e r a t e d in F 5 . F 5 progenies are evaluated in the greenhouse DM screening nursery. Selected F 6
lines are e v a l u a t e d multilocationally for plant height, time to flowering, a n d pollen s h e d d i n g . C o m b i n i n g
ability, uniformity, a n d nicking tests are d o ne at the F 7 stage. F 7 lines are included in t h e Pollinator
Collection.

In c a s e of certain pollinators, an additional two to three generations of selfing is d o ne to achieve


an a c c e p t a b l e level or uniformity before the progeny are included in the Pollinator Collection.

Pollinators f r o m t h e I C R I S A T Pollinator Collection are distributed widely, in r es p o n s e to s e e d


requests. Pollinators s upplied to the Indian national p r o g r a m during the last 10 years have p r o v i d e d wide

61
RMB/F1

Self

S1/F2

Self

S 2 /F 3

DMSN, KVS, TXT Self

TXT, P(S 3 /F 4 ) S3/F4

Self
KVS

PMIHT,P(S4/F5) PRLN 1
S4/F5

Self, G H D M S N
D M S N , KVS

P M A H T , ICMR ( S 5 / F 6 ) S5/F6 PRLN 2

Self-multilocational
NT, Isolation plot evaluation

IHT, ICMR ( S 6 / F 7 ) RLCAT, S 6 / F 7 PRLN 3

A I C P M I P trial SCA, U T
G C A / S C A , UT

ICRISAT
Pollinator Collection
(IPC)

AHT = A d v a n c e d hybrid trial KVS = Kharif visual season


P M A H T = Pearl millet a d v a n c e d hybrid trial
AICPMIP = All India Coordinated Pearl
P M I H T = P e a r l millet initial h y b r i d trial
Millet Improvement Project, India P R L N = P o t e n t i a l R-line n u r s e r y
DMSN = D o w n y mildew screening nursery RMB = R a n d o m m a t e d bulk
GCA = General combining ability R L C A T = R-line c o m b i n i n g ability trial
ICMR = I C R I S A T millet restorer SCA = S p e c i f i c c o m b i n i n g ability trial
IHT = Initial hybrid trial TXT = T e s t c r o s s trial
IPC = I C R I S A T pollinator collection UT = Uniformity test

F i g u r e 1 . I n b r e e d i n g a n d e v a l u a t i o n d u r i n g p earl millet pollinator d e v e l o p m e n t .

62
diversity. C e r t a i n pollinators a r e preferred m o r e t h a n others by I C R I S A T c o o p e r a t o r s in India.

C o m b i n i n g ability studies h a v e indicated that m o s t pollinators are g o o d specific c o m b i n e r s . Grain


yield e v a l u a t i o n of r a n d o m s a m p l e s of pollinators h a v e indicated relatively low grain yield f r o m inbred
pollinators.

Currently, the c o n c e p t of topcrossing is being applied to breeding of topcross hybrids. T o p c r o s s


hybrids are p r o d u c e d by crossing inbred male-sterile lines with variety pollinators (Fig. 2 ) . Variety
pollinators are open-pollinated varieties derived through recurrent selection. S u c h pollinators are generally
high yielding with relatively m o r e stable resistance to d o w n y mildew t h a n inbred pollinators.

T e s t i n g of Fertility Restoration

T h i s is d o n e by c r o s s i n g pollinators with male-sterile lines (A-lines) a n d g r o w i n g the F 1 . Fertility


restoration c a n be j u d g e d on the basis of visual observation of pollen s h e d d i n g a n d by s c o r i n g it on a
1-9 scale ( w h e r e 1 = highly effective restorer, a n d 9 = nonrestorer). It c a n also be tested by s c o r i n g the
s e e d - s e t u n d e r setting b a g s on a 1-9 scale (where 1 = best seed-set or highly effective restorer, a n d 9
= no s e e d - s e t or nonrestorer).

Desirable Traits of a G o o d Restorer Line

T h e desirable traits of a g o o d restorer line are

• G o o d fertility restoration against a large n u m b e r of male-sterile lines with different s y s t e m s of


sterility;

• D o w n y m i l d e w resistance;

• Capability of p r o d u c i n g high-yielding hybrids (combining ability); a n d

• G o o d pollen s h e d d i n g in s e e d production plots in a wide range of e n v i r o n m e n t s .

Production of Restorer Parent

Restorer lines of pearl millet c a n be p r o d u c e d on a large scale in isolation plots. Isolation plots s h o u l d
be a p p r o x i m a t e l y 1 km a w a y f r o m other pearl millet fields. T h r e e s t a g e s of pr oduction are practised in
India: nucleus s e e d , b r e e d e r s e e d , a n d f o u n d a t i o n s e e d s t a g e s . Production of nucleus s e e d is the
responsibility of the breeder; no external inspection is r e q u i r e d . B r e e d e r s e e d is p r o d u c e d by t h e br eeder
u n d e r inspection by s e e d certification a g e n c i e s . B r e e d e r s are not i n v o l v e d in f o u n d a t i o n s e e d p r o d u c t i o n .

M a i n t e n a n c e of Restorer Parent

Restorer lines c a n be m a i n t a i n e d either by sib-mating (by hand) or by r a n d o m m a t i n g in an isolation plot.


T h e y c a n also be m a i n t a i n e d by selfing.

Summary and Conclusion

Pearl millet pollinators in c o m b i n a t i o n with male-sterile lines p r oduce high yielding, d o w n y m ildew

63
resistant, a n d w i d e l y a d a p t e d single-cross hybrids in India. T h r e e major types of b r e e d i n g s c h e m e s are
f o l l o w e d to b r e e d pollinators: (1) early-generation testing; (2) a d v a n c e d - g e n e r a t i o n t e s t i n g ; a n d (3)
c o m b i n a t i o n of (1) a n d (2). M o s t of the pollinators b r e d so far have b e e n g o o d specific c o m b i n e r s . Inbred
pollinators are relatively low yielding. T o p c r o s s or variety pollinators are b e i n g e v a l u a t e d for the
p r o d u c t i o n of high-yielding hybrids. G o o d restorers s h o u l d be able to restore fertility against a w i d e range
of male-sterile lines, p o s s e s s d o w n y mildew resistance, a n d be g o o d pollen p r o d u c e r s in s e e d p r o d u c t i o n
plots. Pollinators c a n be m a i n t a i n e d by sib-mating or by selfing.

Population for OPVC

Mass selection (DMN, EDLN)

OPVC with high yield (TCPC )O A-line TCPC O= TCH


TCHT
Self

S1 s
Self S1s and cross selected
S1,s on A-lines

S 2 ,TXs
Evaluate S for DM resistance and
TXs for fertility restoration

Selected S 2

Random mate and cross on A-line

TCPC 1 A-line x TCPC 1 =TCH


AICPMIP

TCPC 2 A-line x TCPC 2 = TCH


AICPMIP

TCPC 3 A-line x TCPC 3 =TCH


AICPMIP

AICPMIP = All India Coordinated Pearl Millet


Improvement Project
C1...C3 = Cycles of selection
DMN = Downy mildew nursery
EDLN = Extended-daylength nursery
OPVC = Open-pollinated variety from composite
S1...S2 = Self generations 1 and 2
TCH = Inbred % variety topcross hybrid
TCP = Topcross pollinator
TCHT = Topcross hybrid trial.

Figure 2. Breeding of topcross pollinator.

64
Production of Pearl Millet Hybrid Seed
B S Talukdar

In pearl millet, c y t o p l a s m i c - g e n i c m a l e sterility has b e e n exploited in c o m m e r c i a l s e e d p r o d u c t i o n . Details


of hybrid s e e d production h a v e b e e n p r e s e n t e d in Khairwal et al. (1990). M o s t of the s e e d producers in
India prefer not to rely on carry-over stocks b e c a u s e they generally s h o w poor g e r m i n a t i o n if they are
not stored w e l l . M o r e o v e r , st orage also adds to the production cost. T h e r e f o r e , m o s t of the s e e d
p r o d u c e r s in India prefer to t a k e up s e e d pr oduction in t h e postrainy s e a s o n . In s u c h a s c e n a r i o , it is
n e c e s s a r y to calculate the a m o u n t of hybrid s e e d required before the s e e d production p r o g r a m is t a k e n
up. H o w e v e r , s o m e p r o d u c e r s m a y not be able to exercise the p o s t r a i n y - s e a s o n option d u e to
nonavailability of irrigation w a t e r a n d unfavorable weat her conditions.

S t e p s Involved in C o m m e r c i a l Hybrid Seed Production

T h e major factors involved in c o m m e r c i a l hybrid s e e d production are choice of location, c h o i c e of field,


c r o p p i n g history, isolation distance, m ale:female ratio, male-line border, nicking of m a l e a n d f e m a l e
plants, labelling, r o g u i n g , crop m a n a g e m e n t , a n d certification of s e e d .

Choice of Location

In India, a c o n s i d e r a b l e a m o u n t of pearl millet hybrid-seed production is d o n e in the p r e m o n s o o n or


postrainy s e a s o n in non-pearl millet g r o w i n g a r e a s using irrigation. S u c h locations are generally a w a y
f r o m fields w h e r e pearl millet is g r o w n in the rainy s e a s o n by f a r m e r s . Hybrid s e e d is p r o d u c e d ,
p r o c e s s e d , a n d b a g g e d in the s a m e locations that favor raising a g o o d crop of pearl millet u n d e r g o o d
m a n a g e m e n t . S u c h locations are preferred by the highly o r g a n i z e d s e e d sector, s u c h as large public or
private s e e d c o m p a n i e s . In s u c h a situation, the transportation cost is high a n d the time available for
m a r k e t i n g is very short.

S e e d p r o d u c t i o n is also d o n e in the rainy s e a s o n , w h i c h is the main c r o p p i n g s e a s o n . In this


scenario, s e e d p r o d u c e d in the previous year is used for s o w i n g . This requires a d e q u a t e st orage
facilities, a n d the capital invested is locked up for a cons iderable period of time. H o w e v e r , t r a n s portation
cost is m i n i m i z e d . T h e r e is also a d e q u a t e time for m a r k e t i n g . G e n e r a l l y , small local s e e d p r o d u c e r s
prefer this o p t i o n . S e e d p r o d u c e d in the rainy s e a s o n is also u s e d for s o w i n g in the postrainy s e a s o n in
s o m e pearl millet g r o w i n g areas w h e r e the postrainy-season crop contributes significantly to pr oduction
(e.g. Gujarat).

Choice of Field

S e e d p r o d u c t i o n fields s h o u l d be of superior quality c o m p a r e d to fields in w h i c h the F 1 hybrid is g r o w n


by f a r m e r s for grain p r o d u c t i o n . T h e y s h o u l d be adequately isolated f r o m other pearl millet fields. T h e y
s h o u l d be free f r o m w a t e r l o g g i n g , a n d preferably have a light s a n d y soil. T h e length of t h e field s h o u l d
not be too m u c h m o r e t h a n its w i d t h .

Cropping History

C r o p p i n g history is an important consideration for a s e e d production plot. This e n a b l e s the s e e d p r o d u c e r


to plan his fertilizer application. If the previous crop w a s pearl millet, then the field s h o u l d not be selected

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s i n c e t h e r e m i g h t be a possibility of volunteer millet plants f r o m the previous crop.

Isolation Distance

Pearl millet is a highly cross-pollinated crop. In a still environment , a majority of the pollen grains c a n
travel only a f e w m e t e r s , but under m o d e r a t e w i n d velocity (>30 kph) they c a n travel m o r e t h a n 1 km in
t h e direction of t h e w i n d . D u r i n g the s e e d production s e a s o n in India, the w i n d direction generally
c h a n g e s . T h e d i s t a n c e travelled by pollen also d e p e n d s on the p r e s e n c e or a b s e n c e of physical barriers
s u c h as tall, d e n s e v e g e t a t i o n , h e d g e s , buildings, etc. T h u s , for the production of hybrid s e e d , isolation
distances r a n g e f r o m 5 0 0 m to 1000 m d e p e n d i n g on the conditions. In m a n y c a s e s , a large n u m b e r of
s e e d g r o w e r s multiply the s a m e hybrid in a locality (village) isolated f r o m ot hers. In s u c h a situation,
careful p l a n n i n g of s e e d production helps in maintaining the isolation required for large-scale p r o d u c t i o n .

Male: Female Ratio

T h e m a l e : f e m a l e ratio for a hybrid s e e d production field d e p e n d s primarily on t h e p o l l e n - s h e d d i n g


b e h a v i o r of t h e m a l e p a r e n t a n d t he e n v i r o n m e n t al conditions. Usually, a 1:3 ratio provides g o o d s e e d - s e t
in diverse e n v i r o n m e n t s , ranging f r o m the rainy s e a s o n in northern India to the postrainy s e a s o n in
s o u t h e r n India. T h i s ratio c a n range f r o m 1:2 to 1:7.

Male-Line Border

S o w i n g t h e m a l e p a r e n t (pollinator) on all sides of the hybrid s e e d production plot e n s u r e s a d e q u a t e


pollen for g o o d s e e d - s e t on the male-sterile line, a n d eliminates the possibility of c o n t a m i n a t i o n by foreign
p o l l e n . T h i s practice is v e r y simple but is rarely f ollowed in hybrid (certified or truthfully-labelled) s e e d
p r o d u c t i o n p r o g r a m s . Probably, s u c h a practice is c o n s i d e r e d u n n e c e s s a r y b e c a u s e the turbulent w i n d
m o v e m e n t that is often prevalent over large production plots e n s u r e s an a d e q u a t e pollen load within a n d
a r o u n d the plot. W h e r e pollen s h e d d i n g is not a d e q u a t e a n d the production plot is not sufficiently isolated,
a male-line b o r d e r is useful.

Nicking of Male and Female Lines

T h e event of s h e d d i n g a b u n d a n t pollen by the m a l e line (pollinator) w h e n the s t i g m a s of the f e m a l e


(male-sterile) line are m a t u r e a n d receptive is t e r m e d "nicking" in hybrid s e e d p r oduction. G o o d "nicking"
is essential for g o o d s e e d - s e t on the male-sterile line. W h e n "nick ing" is not g o o d , it c a n be i m p r o v e d
by s o w i n g the m a l e a n d f e m a l e lines on different d a t e s . In A n d h r a P r a d e s h , pearl millet hybrid s e e d
p r o d u c t i o n is d o n e by raising nurseries of the parental lines by s o w i n g t h e m on different d a t e s a n d t h e n
transplanting both the parents at the s a m e time in the main production field. This practice e n s u r e s
nicking.

Labelling

In c a s e m o r e t h a n o n e h y b r i d is b e i n g multiplied at t h e s a m e t i m e , proper labelling of the parental lines


is n e c e s s a r y to a v o i d transplanting or s o w i n g w r o n g parental c o m b i n a t i o n s . It is safer to s o w o n e hybrid
p r o d u c t i o n plot at a t i m e .

Roguing

R o g u i n g is an import ant operation in the production of quality hybrid s e e d . T h e major a s p e c t s of roguing

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a r e : (1) r e m o v a l of off-type plants f r o m b o t h t h e pollinator a n d t h e male-sterile lines; a n d (2) r e m o v a l of
pollen s h e d d e r s f r o m the male-sterile line. R o g u i n g s h o u l d be d o n e before anthesis to a v o i d
contamination.

Crop Management

In a hybrid s e e d production plot, crop m a n a g e m e n t involves:

• Proper land preparation;


• Timely sowing;
• A d e q u a t e fertilizer a n d irrigation applications;
• Proper w e e d control; a n d
• C o n t r o l of p e s t s a n d d i s e a s e s , if any.

T h e major objective here is to obtain a high s e e d yield per unit a r e a . T h u s , s e e d p r o d u c t i o n plots are
often g o o d plots a n d a re s e p a r a t e d f r o m fields in w h i c h F 1 hybrids or other pearl millet c r o p s are b e i n g
g r o w n by f a r m e r s . S e e d production is generally d o n e by the o r g a n i z e d sector s u c h a s s e e d c o m p a n i e s
or public sector u n d e r t a k i n g s , w h i c h h a v e a d e q u a t e m e a n s for g o o d crop m a n a g e m e n t .

Certification of Hybrid Seed

In India, certification of hybrid s e e d is d o n e by s e e d certification agencies. Pr oduction plots of pearl millet


hybrids are i n s p e c t e d a n d certified on fulfillment of the conditions laid d o w n by the a g e n c i e s in
consultation with r e s e a r c h e r s . Certification of hybrid s e e d is optional in India; so both certified a n d
truthfully-labelled s e e d s are sold in the market.

Summary and Conclusion

Pearl millet hybrid s e e d pr oduction at present is entirely d e p e n d e n t on the cyt oplasmic-genic male-sterility
s y s t e m in India. T h e p r e d o m i n a n t hybrid type p r o d u c e d is the single-cross hybrid. S e e d production of
pearl millet hybrid is, therefore, straightforward. T i m e l y roguing of off-type plants a n d pollen s h e d d e r s is
very important to p r o d u c e quality s e e d . C r o p - m a n a g e m e n t practices are of critical i m p o r t a n c e in obt aining
g o o d returns.

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Seed Production of Pearl Millet Open-Pollinated Cultivars
C T Hash

T h i s p a p e r d e s c r i b e s p r o c e d u r e s appropriate for the m a i n t e n a n c e of o p e n - p o l l i n a t e d cultivars a n d


t o p c r o s s pollinators of pearl millet, regardless of the m e t h o d s u s e d in d e v e l o p i n g t h e s e genetic materials.
T h e p a p e r c o n c e n t r a t e s on aspects of genetic purity of open-pollinated cultivars a n d t o p c r o s s pollinators.
It is a s s u m e d that a d e q u a t e precautions are taken against physical admixtures during s o w i n g (e.g., s e e d
carry-over in m a c h i n e r y or v o l u n t e er s e e d in the soil), harvest, storage, a n d p r o c e s s i ng or c o n d i t i o n i n g .

Open-Pollinated Cultivars and Topcross Hybrids

T h e principal r e a s o n s for the continuing use of open-pollinated cultivars a n d potential use of topcross
hybrids in pearl millet lie in the c h e c k e r e d history of single-cross grain hybrids in India. T h e s e hybrids
b e c a m e s u s c e p t i b l e to d o w n y m i l d e w , ergot, a n d s m u t , a n d the cost o f their s e e d multiplication w a s h i g h .
T h e visible variability in pearl millet open-pollinated cultivars a n d topcross pollinators is a c o n s e q u e n c e
of their variability for m a n y useful traits, including variability for resistance to d o w n y m i l d e w, w h i c h is t h e
key to durability of resistance. This range of variability for resistance to d o w n y m ildew is very difficult a n d
costly to b r e e d into pearl millet hybrid parental lines. I n d e e d , m a r k e r - a s s i s t e d b a c k c r o s s p r o g r a m s to
attempt this h a v e only recently b e e n initiated at I C R I S A T Asia Center. H o w e v e r , genetic variability for
d o w n y m i l d e w resistance c a n easily be carried in br eeding populations that serve as the s o u r c e for
open-pollinated cultivars a n d topcross pollinators. Additionally, the variability in flowering time within an
open-pollinated cultivar provides normal protection against ergot a n d s m u t infection, d u e to earlier a n d
m o r e prof use pollen s h e d d i n g over a longer period than in single-cross hybrids. This variability in
f l o w e r i n g t i m e facilitates s e e d production of topcross hybrids b a s e d on variable ("topcross") pollinators,
as it p r o v i d e s a w i d e r w i n d o w of nicking. Finally, variability in flowering time within the pearl millet
o p e n - p o l l i n a t e d cultivar or t o p c r o s s hybrid cultivar contributes to superior stability of grain a n d stover yield
across e n v i r o n m e n t s . This provides greater opportunity for e s c a p e and/or recovery f r o m b o t h drought a n d
pollen w a s h t h a n is offered by the uniform single-cross hybrid cultivars.

T h e s e c o n d reason for continuing with i m p r o v e d open-pollinated cultivars of pearl millet is that


their s e e d multiplication is technically simpler t h a n that of hybrids. This facilitates s e c o n d a r y s p r e a d of
s u c h cultivars via local s e e d s y s t e m s , a n d provides a m e c h a n i s m for c o n t i n u e d evolution of the crop in
f a r m e r s ' fields. T h e role of local s e e d multiplication a n d the n e e d for c o n t i n u e d involvement of farmers
in the d e v e l o p m e n t of their o w n cultivars is especially important in remote a n d h e t e r o g e n e o u s regions
that a r e poorly s e r v e d by the formal s e e d s y s t e m (Almekinders et al. 1994). It is p r o b a b l y critical to
m a i n t e n a n c e of local adaptation a n d yield stability in m a n y of the m o r e marginal pearl millet production
e n v i r o n m e n t s . H e t e r o g e n e o u s open-pollinated cultivars a n d topcross hybrid cultivars are appropriate for
s u c h h a r s h a n d isolated a r e a s . Millions of hectares in southern Asia, a n d in w e s t e r n , central, a n d
s o u t h e r n Africa are annually s o w n to open-pollinated cultivars. T h e s e cultivars are often f armers'
l a n d r a c e s , but increasingly, i m p r o v e d open-pollinated cultivars are being a d o p t e d . T h e s e open-pollinated
cultivars a r e g r o w n b e c a u s e they are well suited to the vast regions w h e r e traditional agricultural
practices a re still the rule. In t h e s e a r e a s , hybrid s e e d distribution c h a n n e ls are not sufficiently flexible
to a c c o m m o d a t e t h e s e a s o n - t o - s e a s o n variation in s e e d d e m a n d . In the arid a n d s e m i - a r i d e n v i r o n m e n t s
w h e r e pearl millet is traditionally g r o w n , these g e n o t y p e s m e e t f a r m e r s ' n e e d s better t h a n any other
available alternative.

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I m p r o v e d open-pollinated cultivars a n d topcross pollinators have several features that m a k e t h e m
more appropriate:

• M a i n t e n a n c e a n d s e e d multiplication of i m p r o v e d open-pollinated cultivars, topcross pollinators,


a n d to a lesser extent topcross hybrids, is relatively simple c o m p a r e d to that of single-cross
hybrids b a s e d on inbred parental lines. S e e d production targets for open-pollinated cultivars c a n
be a c h i e v e d easily a n d rapidly on relatively small production a r e a s .

• O p e n - p o l l i n a t e d cultivars d e r i v e d f r o m an o n g o i n g p o p u l a t i o n - i m p r o v e m e nt p r o g r a m c a n replace
older varieties, either as n e w cultivars or as i m p r o v e d versions of the existing cultivar. An e x a m p l e
of this in pearl millet is the r eplacement of the original O k a s h a n a I in N a m i b i a (ICTP 8203) with
I C M V 8 8 9 0 8 released in 1989 ( W i t c o m b e et al. 1995).

• S e e d production costs for open-pollinated cultivars (all s e e d classes) a n d t opc r o s s pollinators


(nucleus-, breeder-, a n d f o u n d a t i o n - s e e d classes) are relatively low as smaller a r e a s are required
to multiply a given quantity of s e e d on their v igorous, high-yielding plants than w o u l d be required
for multiplication of single-cross hybrids on their relatively w e a k e r inbred s e e d p a r e n t s . T h u s , s e e d
quantities c a n be built up rapidly a n d economically.

• O p e n - p o l l i n a t e d cultivars have a distinct a d v a n t a g e in areas w h e r e s e e d distribution is difficult a n d


costly. T h e s e e d of open-pollinated cultivars can m o v e from farmer to farmer a n d c a n be s a v e d
by the f a r m e r f r o m s e a s o n to s e a s o n . Both of these factors have a multiplicative effect on the
a r e a c o v e r e d by a particular cultivar.

• E x c h a n g e of i m p r o v e d g e r m p l a s m a m o n g national p r o g r a m s , a n d b e t w e e n the public a n d private


sectors of these national p r o g r a m s , is easier with open-pollinated materials than with c l o s e d -
pedigree materials, which might involve proprietary rights.

• T h e r e are greater opportunities to exploit genetic variability within the introduced open-pollinated
cultivars a n d t o p c r o s s pollinators than exist within the inbred parental lines of single-cross,
t h r e e - w a y, a n d d o u b l e - c r o s s hybrids. This facilitates reselection to m e e t r e q u i r e m e n t s for local
adaptation and adoption.

Open-Pollinated Cultivars

T h e t e r m "open-pollinated cultivar", or "variety", for a cross-pollinated crop originally m e a n t a


self-reproducing population of plants that, although not genetically identical, exhibit unique, recognizable,
a n d stable associations of traits. T h e t e r m has more recently b e e n redefined as a superior fraction of a
population that is different, relatively uniform, a n d stable. S u c h a variety is different b e c a u s e it p o s s e s s e s
a c o m b i n a t i o n of traits that distinguish it f r o m other k n o w n cultivars a n d define its identity. In pearl millet,
the following types of b a s e populations of i m p r o v e d open-pollinated cultivars are possible: landrace
cultivars, synthetic varieties initially p r o d u c e d by intermating inbred parents, varieties resulting f r o m
long-term m a s s selection, experimental varieties constituted f r o m elite progenies identified during a
p r o g r a m o f recurrent selection, a n d the various intermediates possible a m o n g these four g r o u p s . A n
i m p r o v e d o p e n - p o l l i n a t e d cultivar c a n be readily identified a n d m a i n t a i n e d , a n d is m o r e likely to be
multiplied a n d w i d e l y a d o p t e d by f a r m e r s . A successful open-pollinated cultivar of pearl millet is likely to
h a v e r e d u c e d variation for significant a g r o n o m ic traits ( 5 0 % f l o w e r i n g , tiller n u m b e r , panicle d i m e n s i o n s ,
plant height, a n d grain m a s s ) , a n d s h o u l d be relatively stable in terms of e x p r e s s i o n of t h e s e traits over
time. It s h o u l d not exhibit variation b e y o n d the acceptable st andards fixed for different traits. Experimental

69
varieties that are constituted by r e c o m b i n i n g 8-15 s e l e c t e d families f r o m a famil y - s t r u c t u r e d p o p u l a t i o n
c a n be sufficiently uniform in a p p e a r a n c e prov ided care is e x e r c i s e d in s electing families that a r e similar
in maturity, plant height, tillering, a n d panicle characters.

An o p e n - p o l l i n a t e d cultivar s h o u l d be an a s s e m b l a g e of plants h a v i n g relatively uniform


p h e n o t y p e s a n d representing a superior fraction of a population that has u n d e r g o n e s o m e d e g r e e of
i m p r o v e m e n t . Selection of superior families for constituting an e x p e r i m e n t a l variety m a y be n e c e s s a r y
e v e n in p o p u l a t i o n s that h a v e b e e n s u b j e c t e d to several cycles of i m p r o v e m e n t , especially if t h e gross
plant architecture of t h e b a s e population is similar to that of an existing a d a p t e d o p e n - p o l l i n a t e d cultivar.

Characterizing an Open-Pollinated Cultivar or T o p c r o s s Pollinator

O n c e an o p e n - p o l l i n a t e d cultivar has r e a c h e d the release stage, it s h o u l d be d e s c r i b e d for salient


f e a t u r e s in t h e a r e a of its a d a p t a t i o n . Similarly, o n c e a topcross hybrid has r e a c h e d the release sta g e ,
it a n d its parents s h o u l d be d e s c r i b e d . For topcross pollinators, the description required is essentially
identical to that required for an open-pollinated cultivar. If the s e e d production a n d seed-purity evaluation
e n v i r o n m e n t s a r e likely to be m a r k e d l y different f r o m the e n v i r o n m e n t s in w h i c h the grain crop is to be
p r o d u c e d , it m a y be helpful to d e v e l o p a s e p a r a t e varietal description of t h e o p e n - p o l l i n a t e d cultivar or
h y b r i d p a r e n t al p o p u l a t i o n for t h e se e n v i r o n m e n t s as w e l l . This will reduce the likelihood of a genetically
g e n u i n e s eed-lot b e i n g rejected by seed-certification officials d u e to an e n v i r o n m e n t a l l y - i n d u c e d departure
f r o m t h e varietal d e s c r i p t i o n .

An import ant attribute of a g o o d open-pollinated cultivar or topcross pollinator is uniformity.


H o w e v e r , o p e n - p o l l i n a t e d cultivars a n d topcross pollinators will s e l d o m be as uniform as e v e n the most
variable of single-cross hybrids. E v e n t h o u g h an elite fraction of the population is r e c o m b i n e d to p r o d u c e
the o p e n - p o l l i n a t e d cultivar or topcross pollinator, it will s h o w s o m e variation for several important
a g r o n o m i c c h a r a c t e r s . T h e morphological traits a n d range of variation e x p e c t e d within a n o p e n - p o l l i n a t e d
variety or t o p c r o s s pollinator m u s t be a d e q u a t e ly d e s c r i b ed to s e r v e as a guide for its m a i n t e n a n c e a n d
s e e d certification.

Seed-certification s t a n d a r d s s h o u l d be f i x e d carefully for various s t a g e s of s e e d multiplication to


provide quality c o n t r o l — n o t to h a m p e r s e e d production a n d distribution. T h e s t a n d a r d s set for pearl millet
o p e n - p o l l i n a t e d cultivars a n d t o p c r o s s pollinators s h o u l d not be too s t r i n g e n t — i n d e e d , they s h o u l d be
c o n s i d e r a b l y r e l a x e d c o m p a r e d with t h o s e set for the genetically uniform, inbred parental lines of hybrids.
T h e y s h o u l d be realistic (not idealistic), enforc eable, a n d appropriate for the conditions prevailing in a
given country.

C h a r a c t e r s s u c h as a d a p t a t i o n , plant height, panicle length a n d girth, crop d u r a t i o n , grain color,


grain texture, grain size, panicle pigment a t i o n a n d bristling, plant architecture, a n d tolerance or resistance
to pests a n d p a t h o g e n s s h o u l d be c o n s i d e r e d in varietal descriptions.

E a c h o p e n - p o l l i n a t e d cultivar o r topcross pollinator r e l e a s e d s h o u l d h a v e s o m e distinct genetic


feature(s) that c a n be u s e d to distinguish it f r o m others. Attributes s u c h as p i g m e n t a t i o n a n d p u b e s c e n c e
of v a r i o us plant parts, d e g r e e of panicle bristling, panicle length a n d girth, s e e d s h a p e a n d color, a n d
d e g r e e of i n t e r n o d e c o n d e n s a t i o n c a n be c o n s i d e r e d for characterization. A list of traits that c a n be u s e d
in d e s c r i b i n g an o p e n - p o l l i n a t e d cultivar or topcross pollinator is given in Table 1. W h e n quantitative
c h a r a c t e r s a r e u s e d a s varietal descriptors, the e x p e c t e d m e a n , r a n g e , a n d s t a n d a r d d e v i a t i o n f r o m the
m e a n of t h e v a r i o u s c l a s s e s s h o u l d be giv en to indicate t h e a c c e p t able variation within the cultivar or
t o p c r o s s pollinator. For qualitative characters, the e x p e c t e d variants m a y be given in p e r c e n t a g e s (e.g.,

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leaf b l a d e s m o o t h , hairy leaf blade not e x c e e d i n g 5%). It is also useful to include in t h e varietal
description a short list of off-type trait combinations that are not e x p e c t e d to occur at all e x c e p t as a
result of physical a d m i x t u r e , the p r e s e n c e of volunteer seedlings, or pollen c o n t a m i n a t i o n .

T a b l e 1. P l a n t c h a r a c t e r s for potential inclusion in t h e varietal d e s c r i p t i o n of a p e a r l millet


o p e n - p o l l i n a t e d cultivar o r t o p c r o s s pollinator.

Characters

Plant part Qualitative Quantitative

Seedling Coleoptile color Vigor


S e e d l i n g color

Stem Dwarf/tall Height


Internode color N u m b e r of n o d e s
I n t e m o d e hairiness N u m b e r of tillers -
N o d e color Primary, S e c o n d a r y ,
N o d a l ring of hairs Tertiary

Leaf Auricle color M a i n s t e m leaf n u m b e r


Blade color Flag leaf length
M a r g i n color Flag leaf w i d t h
Midrib color Fourth leaf length
S h e a t h color Fourth leaf w i d t h
Blade hairiness Leaf angle
M a r g i n hairiness
S h e a t h hairiness

Panicle S t i g m a color Diameter


Fresh anther color Length
Dry anther color N u m b e r plant - 1
G l u m e color Ratio of d i a m e t er to length
G l u m e c u p color
Bristling -
Color, S h a p e , C o m p a c t n e s s

Seed Pericarp color Length


E n d o s p e r m color Width
E n d o s p e r m texture 1000-seed mass
Shape
Color in F e S O 4 test
Color in phenol test
Color in modified
P h e n o l tests
CuSO4, 0.4%
Na2CO3, 0.6%

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Quantitative descriptors will be of value in the m a i n t e n a n c e of o p e n - p o l l i n a t e d cultivars a n d
t o p c r o s s pollinators, a n d in the production of breeder s e e d . In s u b s e q u e n t s t a g e s of s e e d multiplication
a n d for certification s t a n d a r d s , it is the qualitative characteristics, especially t h o s e listed for off-type
plants, that s h o u l d be u s e d as guidelines. A pearl millet open-pollinated cultivar or t o p c r o s s pollinator
s h o u l d s h o w little c h a n g e in its phenotypic attributes relative to those of other released materials w h e n
r e p r o d u c e d in its a r e a of a d a p t a t i o n for m a i n t e n a n c e and/or s e e d increase f r o m o n e stage to another.
P h e n o t y p i c stability is highly d e s i r e d , b e c a u s e only a stable material will perform in a c c o r d a n c e with
e x p e c t a t i o n s f r o m y e a r t o y ear a n d s e a s o n t o s e a s o n .

Maintenance and Seed Production

P r o c e d u r e s for the m a i n t e n a n c e a n d s e e d pr oduction of open-pollinated cultivars a n d t o p c r o s s pollinators


vary slightly with the s t a g e of multiplication—reflecting the principle that extra care is n e e d e d to maintain
the identity of t h e material during the early st a g e s , a n d that there s h o u ld a l w a y s be an o n w a r d f l ow f r o m
stage to s t a g e . O n l y n u c l e u s s e e d , w h i c h is u s e d tor s o w i n g b r e e d e r - s e e d plots, s h o u l d r egenerate itself,
a n d that s h o u l d b e u n d e r rigorously controlled conditions.

Pearl millet is m o r e easily c o n t a m i n a t e d by outside pollen b e c a u s e of its p r o t o g y n o u s f lowering


habit. Further, the pollen of pearl millet is more robust. It can survive a greater d e g r e e of desiccation a n d
c a n travel farther with the w i n d , on insects, or e v e n on the clothing or bodies of p e o p l e . T h e r e f o r e , dur in g
the first or s e c o n d d a y of flowering, a pearl millet field is most vulnerable to a n y external pollen,
especially w i n d b o r n e p o l l e n , b e c a u s e little pollen is being p r o d u c e d within the field.

T h e n e e d for an appropriate isolation distance for multiplication of s e e d of o p e n - p o l l i n a t e d


cultivars a n d t o p c r o s s pollinators is r e c ognized. T h e permissible levels of off-type plants in s u c h materials
c a n be higher t h a n in cultivars of self-pollinated crops or hybrid parental lines. This is b e c a u s e (a)
cultivars of cross-pollinated crops h a v e a higher "buffering capacity" for off-type effects as a natural
c o n s e q u e n c e of their h e t e r o z y g o u s a n d h e t e r o g e n e o u s population structure; a n d (b) if g o o d isolation
distance h a s b e e n m a i n t a i n e d , m o s t of the off-types o b s e r v e d will h a v e arisen t h r o u g h r e c o m b i n a t i o n of
g e n e s alr e a d y p r e s e n t a n d not f r o m the introduction of alien g e n e s .

Multiplication rates in pearl millet are high, particularly with open-pollinated cultivars a n d topcross
pollinators. T h r e e to four kg of s e e d a re more t h a n sufficient for s o w i n g o n e hectare, a n d for p r o d u c i n g
1 0 0 0 kg of c l e a n s e e d , e v e n u n d e r c onditions of short d a y s , cool t e m p e r a t u r e s , a n d m o d e r a t e soil fertility.
T h u s , t h r e e s t a g e s of s e e d multiplication (breeder, f o u n d a t i o n , a n d certified) a r e generally sufficient for
multiplication of o p e n - p o l l i n a t e d cultivars. Note that in the case of t o p c r o s s pollinators, the certified s e e d
class d o e s not exist as the f o u n d a tion s e e d class of these materials is u s e d in the pr oduction of the
certified s e e d of t o p c r o s s hybrids.

Nucleus Seed

T h e n u c l e u s s e e d m a i n t a i n e d by the breeder is the s e e d stock f r o m w h i c h all other s e e d classes are


d e r i v e d . It is not subject to certification. Howev er , it is the basic stock a n d requires special attent ion.
N u c l e u s s e e d s h o u l d be the only stock that regenerates itself. It s h o u l d not be r e g e n e r a t e d e v e r y year
unless i n a d e q u a t e s e e d - s t o r a g e conditions necessitate this. Regeneration of nucleus s e e d o n c e in four
y e a r s is r e c o m m e n d e d . T h e p r o d u c e of a n u c l e u s - s e e d plot s hould be thoroughly dried, treated against
insects ( n a p h t h a l e n e at t h e rate of 100 g per 10 kg of s e e d will render the s e e d unattractive to storage
pests), a n d d i v i d e d into six lots of 5-10 kg e a c h . T h e s e lots s h o u l d be kept in hermetic ally se a l e d
containers in a cool store, if available. Lots 1 , 2 , 3 , a n d 4 are u s e d to s o w b r e e d e r - s e e d plots for t h e next

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four y e a r s ; lot 5 is u s e d to s o w the next n u c l e u s - s e e d plot four years h e n c e ; a n d lot 6 is a b a c k u p or
insurance lot that s h o u l d be kept in a different building than the other lots.

M a s s s e l e c t i o n in isolation. This is the preferred m e t h o d of production a n d m a i n t e n a n c e of nucleus


s e e d of o p e n - p o l l i n a t e d cultivars a n d topcross pollinators. N u c l e u s - s e e d plots s h o u l d be g r o w n in
isolation at least 1 5 0 0 m a w a y f r o m a n y other plot of pearl millet, wild pearl millet, or e l e p h a n t grass
( P e n n i s e t u m purpureum), preferably in the off-season w h e n crop g r o w t h a n d s e e d quality will be
excellent. T h e c r o p s h o u l d be t h i n n e d at an early stage to e v e n l y - s p a c e d single plants (or s o w n in hills
a n d t h i n n e d to single plants), using only one-third of the n o r m a l c o m m e r c i a l plant population so that
differences in individual plant e x p r e s s i o n are m a x i m i z e d . Several inspections prior to f l o w e r i n g are
r e c o m m e n d e d . Off-type plants s h o u l d be r e m o v e d by uprooting (roguing). Cutting s u c h off-type plants
at g r o u n d level is not a c c e p t a b l e as m a n y will produce ratoon tillers that c a n c o n t a m i n a t e t h e p r o d u c t i o n
plot. D u r i n g f l o w e r i n g , daily inspections are n e e d e d to identify a n d r e m o v e all plants s u s p e c t e d of being
not true to type before they s h e d pollen. Care should be t a k e n during inspections at flowering to visit the
n u c l e u s - s e e d plot b e f o r e — n o t after—visiting other pearl millet plots in order to m i n i m i z e c o n t a m i n a t i o n
by pollen b o r n e on the bodies a n d clothing of the roguing crew. A final r o g u i n g s h o u l d be c o n d u c t e d in
the s t a n d i n g c r o p before harvest to eliminate a n y remaining off-type plants. To e n s u r e that the varietal
n o r m s do not c h a n g e d u e to genetic drift, n u c l e u s - s e e d plots of open-pollinated cultivars a n d topcross
pollinators s h o u l d not be less than 0.1 ha, a n d s h o u l d contain at least 3 0 0 0 plants. On t h e other h a n d ,
s u c h plots s h o u l d not e x c e e d 0.2 ha b e c a u s e it is difficult to scrutinize all plants daily in large plots.

B u l k p o l l i n a t i o n . T h i s m e t h o d of maintaining a n d p r o d u c i n g pearl millet nucleus s e e d m a y be


u n d e r t a k e n w h e n an isolation distance of 1500 m is not possible. Howev er, in this m e t h o d it is preferable
to maintain an isolation distance of several h u n d r e d meters d u r i n g flowering in order to minimize the risk
of pollen c o n t a m i n a t i o n . T h e cost of labor a n d supplies is m u c h greater in this m e t h o d than in an isolated
m a s s - s e l e c t i o n plot. S o w the n u c leus-seed bulk to obtain about 8 0 0 0 s p a c e d plants. Select a b o u t 3 0 0 0
plants that fit the p h e n o t y p ic varietal description, a n d cover all panicles possible on these plants with
setting b a g s . Collect bulk pollen f r o m the selected plants a n d pollinate t h e m using this bulk. To r educe
inbreeding, this m a y be d o n e by collecting pollen from the o d d - n u m b e r e d rows a n d pollinating the
e v e n - n u m b e r e d rows, a n d reversing this procedure every alternate d a y during flowering. At harvest,
select h a n d - c r o s s e d panicles f r o m about 1000 plants that have the d e s i r e d panicle a n d grain charac-
teristics. Bulk e q u a l quantities of h a n d - c r o s s e d s e e d f r o m e a c h of these plants to p r o d u c e t h e nucleus
s e e d . T h e quantity of nucleus s e e d p r o d u c e d by this m e t h o d can be m a n i p u l a t e d by v a r y i n g the n u m b e r
of h a r v e s t e d plants a n d t h e quantity of h a n d - c r o s s e d s e e d s a m p l e d f r o m e a c h . B r e e d e r s e e d m a y b e
obtained by bulk i n g the h a n d - c r o s s e d s e e d p r o d u c e d on the remaining plants not h a r v e s t e d for nucleus
seed.

Isolated bulk s o w i n g c o n v e r t e d into half-sib c r o s s i n g block. Establish a b o u t 12 0 0 0 plants in rows


in an isolated plot s o w n with bulk nucleus s e e d . Prior to first panicle e m e r g e n c e , arbitrarily des ignate
rows as m a l e or f e m a l e in the ratio of 1 male row to 2-4 f e m a l e rows. C o v e r the panicles in t h e f e m a l e
rows with setting bags. Undesirable a n d off-type plants in the male rows (as m a n y as 2 0 - 4 0 % ) should
be u p r o o t e d . Collect bulk pollen daily f r o m the m a l e rows a n d use it to cross o n t o receptive panicles in
the f e m a l e r o w s . In this m a n n e r , better control c a n be e x e r c i s e d over the pollen source in the m a l e r o w s
u s e d as pollinators. T h e m a l e rows provide a g o o d indication of the e n v i r o n m e n t a l variation in the field,
w h i c h in turn facilitates selection within the f e m a l e rows of plants c o n f o r m i n g to the varietal description.

Prior to harvest, treat e a c h 5 m of a f e m a l e row as a g r i d . Select 3-5 plants that m e e t t h e varietal


description, a n d harvest t w o or three hand-crossed panicles f r o m e a c h grid. A b o u t 1000 panicles c a n be
s e l e c t e d in this m a n n e r to serve as sources of the next generation of nucleus s e e d . A bulk s a m p l e of

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s e e d f r o m t h e s e panicles and/or additional h a n d - c r o s s e d panicles f r o m t h e selected true-to-type f e m a l e
plants c o u l d p r o v i de t h e b r e e d e r s e e d .

I s o l a t e d half-sib p a n i c l e - t o - r o w plot. This is a simple a n d effective s y s t e m for m a i n t a i n i n g an


o p e n - p o l l i n a t e d cultivar or t o p c r o s s pollinator of pearl millet a n d for p r o d u c i n g b o t h n u c l e u s s e e d a n d
b r e e d e r s e e d . It requires isolation a n d c a n be initiated with s e e d f r o m individual panicles h a r v e s t e d for
nucleus s e e d following any of the m e t h o d s d i s c u s s e d a b o v e . It involves the following s t e p s :

• S e l e c t e d half-sib panicles (500-1000) are individually t h r e s h e d . T h e following s e a s o n , these are


s o w n as half-sib f e m a l e r o w s in a well-isolated plot. T h e f e m a l e rows are s o w n alternately with
r o w s of a b a l a n c e d m a l e bulk constituted by mixing equal s e e d quantities f r o m e a c h half-sib
panicle.

• Prior to f l o w e r i n g , a n y f e m a l e half-sib rows having off-type plants are r e m o v e d . Similarly, off-type


p l a n t s in t h e m a l e bulk a r e r o g u e d out as they a r e identified.

• Prior to harvesting, about 5 0 % of the f e m a l e rows that best c o n f o r m to the varietal description are
s e l e c t e d . T h e m a l e r o w s serve as controls for the selection of f e m a l e rows. F r o m e a c h of the
s e l e c t e d f e m a l e r o w s , panicles f r o m 2-4 plants are harvested to serve as nucleus s e e d for future
i n c r e a s e a n d m a i n t e n a n c e . If the r equirement for breeder s e e d is s m a l l , it m a y be possible to take
a bulk s a m p l e f r o m these s elected panicles to provide breeder s e e d . If larger quantities are
r e q u i r e d , b r e e d er s e e d m a y b e h a r v e s t e d f r o m the remaining true-to-type plants in the selected
female rows.

Breeder Seed

T h e responsibility for m aintaining the purity of breeder s e e d , as long as the open-pollinated cultivar or
t o p c r o s s hybrid is in p r o d u c t i o n , s h o u l d rest with the br eeder. W h e n a released cultivar is replaced by
a superior o n e , a n d w h e n r e p l a c e m e n t is well underway, t h e n breeder s e e d m a i n t e n a n c e of the replaced
cultivar or its p a r e n t a l s t o c k s c a n be d i s c o n t i n u e d . H o w e v e r , the s e e d s h o u l d be contributed to a suitable
g e r m p l a s m collection, if that h a d not b e e n d o n e earlier, in order to e n s u r e that viable s e e d is available
in future, s h o u l d the n e e d for it arise.

T h e quantity o f b r e e d e r s e e d p r o d u c e d c a n b e regulated b y increasing the n u m b e r a n d length


of t h e r o w s in t h e b r e e d e r s e e d p r o d u c t i o n plot. However, in order to maint ain the highest level of genetic
identity, the b r e e d e r - s e e d plot s h o u l d be small a n d m a n a g e a b l e . T h e s e plots s h o u l d be g r o w n in the off-
s e a s o n at less t h a n n o r m a l plant population, a n d in extremely g o o d isolation (more t h a n 7 5 0 m f r o m
other f l o w e r i n g pearl millet plots f r o m the time of first panicle e m e r g e n c e until harvest). B o t h isolated
m a s s - s e l e c t i o n a n d isolated half-sib panicle-to-row m e t h o d s d e s c r i b ed a b o v e for nucleus s e e d production
are a p p r o p r i a te for b r e e d er s e e d p r o d u c t i o n . T h e plots s h o u l d b e t h i n n e d t o s p a c e d plants a n d r o g u e d
daily d u r i n g f l o w e r i n g . G o o d r o g u i n g at this stage c a n greatly reduce the n e e d to do further roguing in
the later s t a g e s . N o t m o r e t h a n 1 % of off-types s h o u l d be permitted at final inspection. T h e plot size c a n
be 0.1-0.5 h a , d e p e n d i n g on t h e quantity of breeder s e e d required to p ro d u c e t h e t a r g e t e d quantity of
certified s e e d . A s s u m i n g a v e r y conservative multiplication factor of 2 0 0 p e r g e n e r a t i o n , o n e c a n m a k e
the n e c e s s a r y calculations a s g i v e n i n T a b l e 2 .

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T a b l e 2 . E s t i m a t e d a r e a s re q u i red for s e e d multiplication o f different s e e d c l a s s e s o f
a pearl millet o p e n - p o l l i n a t e d variety ne c e s s a r y to s o w 1 m h a .

Breeder Foundation Certified


seed seed seed

Area 0.1 ha 25 ha 5000 ha

Quantity 100 kg 20 t 4000 t

T h u s , 1 0 0 kg of b r e e d e r s e e d of an open-pollinated cultivar, p r o d u c e d f r o m 0.5 kg of n u c l e u s s e e d


of t h e s a m e cultivar, c a n easily m e e t the requirement for the production of certified s e e d n e c e s s a r y to
s o w 1 m h a . In fact, w i t h v i g o r o u s cultivars g r o w n under g o o d m a n a g e m e n t conditions, it is possible to
achieve a seed-multiplication rate t w o or three times higher.

Foundation Seed

T h e first increase f r o m breeder s e e d is referred to as foundation s e e d . T h e responsibility for p r o d u c i n g


foundation s e e d often rests with a s e e d production agency, but it m a y call for a s s i s t a n c e f r o m the
breeder(s) responsible for maintaining the identity of the cultivar or hybrid parent. Only certified b r e e d e r
s e e d s h o u l d be u s e d to s o w f o u n d a t i o n s e e d plots. Breeder s e e d can be u s e d to cover a larger a r e a for
f o u n d a t i o n s e e d p r o d u c t i o n if it is first s o w n to a small nursery a n d t h e seedlings are t h e n t r a n s p l a n t e d
at appropriate s p a c i n g into the s e e d multiplication plot. T h e foundation s e e d plot s h o u l d be s o w n on
uniform land at least 1 km f r o m the nearest pearl millet plot, preferably e v e n farther if s o w n in the
direction of the w i n d . W i t h t h e s a m e s e e d , first s o w a border 4 m w i d e a r o u n d t h e per im et er of t h e plot
4-7 d a y s before s o w i n g the center. T h e purpose of this perimeter s o w i n g (from w h i c h the s e e d will be
separately h a r v e s t e d a n d sold as grain) is to provide pollen by the time the plants in t h e center b e g i n to
c o m e to flower. This provides protection, b a s e d on dilution, against i n c o m i n g w i n d b o r n e pollen f r o m other
fields. Before f l o w e r i n g , s e a r c h for a n d destroy a n y volunteer pearl millet plants on t h e field b o r d e r s a n d
in ditches, a n d in nearby crop land. Rogue the obvious off-type a n d d o w n y mildew plants as t h e y are
d e t e c t e d , preferably before flowering, to m e e t foundation s e e d standards. As m a n y as 1 0 - 1 5 % of the
plants m a y h a v e to be r o g u e d out, but this fraction will be lower in f o u n d a t i o n s e e d plots s o w n with
b r e e d e r s e e d f r o m w e l l - r o g u e d plots. In foundation s e e d plots, m a i n t e n a n c e of the genetic identity of the
o p e n - p o l l i n a t e d cultivar or t o p c r o s s pollinator s h o u l d be e m p h a s i z e d . T h e s e plots s h o u l d be closely m o n i -
t o r e d by p e r s o n s responsible for the production of foundation s e e d .

Certified Seed

Certified s e e d is the last stage of s e e d multiplication. It s h o u l d be p r o d u c e d by select s e e d g r o w e r s or


at s e e d f a r m s , a n d its production s h o u l d b e c o o r d i n a t e d a n d s u p e r v i s e d b y public or private s e e d
a g e n c i e s r e s p o n s i b l e for s e e d multiplication a n d distribution. T h e s e e d specialists a n d s e e d certification
a g e n c y s h o u l d assist g r o w e r s i n t h e p r oduction of g o o d quality certified s e e d . Slightly l o w e r - t h a n - o p t i m u m
plant density for t h e s e e d crop m a y be u s e d as this helps achieve better s e e d quality.

Certified s e e d of open-pollinated cultivars should be p r o d u c e d only f r o m officially a p p r o v e d


f o u n d a t i o n s e e d , a n d s h o u l d b e s o w n i n large c o n t i g u o u s b l o c k s — p r e f e r a b l y i n a n a r e a w h e r e other pearl
millet plots h a v e not b e e n s o w n . T h e r e s h o u l d be an isolation distance of at least 4 0 0 m. It m a y be

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possible to e n s u r e t h a t pearl millet-producing farmers in the vicinity of the certified s e e d production plots
a r e g i v e n — a n d a g r e e t o g r o w — t h e s a m e cultivar a s that being multiplied. R o g u e off-type a n d d i s e a s e d
plants if possible, inspect ( m a k i n g s u r e that plants not c o n f o r m i n g to the varietal description do not
e x c e e d 5% at t h e final inspection), certify, condition, a n d treat with insecticides a n d fungicides before
b a g g i n g for sale to f a r m e r s .

Truthfully-Labelled Seed

In India, "truthfully-labelled s e e d " is s o l d on the strength of the cultivar n a m e a n d the b r a n d n a m e of the


p r o d u c e r rather t h a n h a v i n g its identity g u a r a n t e e d by the g o v e r n m e n t s e e d certification a g e n c y . T h e
legal r e q u i r e m e n t is that it be g r o w n with an isolation distance of at least 10 m f r o m pearl millet plots of
the s a m e cultivar that for s o m e r e a s on fail certified s e e d standards. T h e truthful label provides a m e c h a -
n i s m by w h i c h private a n d public s e e d a g e n c i e s c a n m a r k e t p r o d u c e that fails to m e e t the strict s t a n d a r d s
for certified s e e d , a n d t h u s limit losses f r o m s u c h s e e d pr oduction plots. It also provides these
o r g a n i z a t i o n s a m e a n s to m a r k e t s e e d of cultivars t h a t have not c o m p l e t e d r e q u i r e m e n t s for f o r m a l
release by g o v e r n m e n t authorities. This is w e l c o m e w h e n it permits a b r o a d array of g o o d proprietary
products to r e a c h f a r m e r s earlier t h a n w o u l d be possible following official pro c e d u r e s for cultivar testing
a n d release.

Standards for Maintaining Varietal Uniformity

As an o p e n - p o l l i n a t e d cultivar or t o p c r o s s pollinator g o e s t h r o u g h different s t a g e s of s e e d multiplication,


it is likely to b e c o m e m o r e a n d m o r e variable. In open-pollinated cultivars of m a i z e , C I M M Y T (1984)
s u g g e s t s that in m a i n t e n a n c e of b r e e d e r s e e d the s elected families s h o u l d fall within ±0 . 7 s t a n d a r d
deviations of the varietal m e a n . In f o u n d a t i o n s e e d plots, plants differing f r o m t h e varietal m e a n by m o r e
t h a n ±1 . 5 s t a n d a r d deviat i o n s s h o u l d be r e m o v e d . If it is possible to carry out roguing in the certified
s e e d p r o d u c t i on plots, plants falling m o r e t h a n ±2 s t a n d a r d deviations f r o m the varietal m e a n s h o u l d b e
r e m o v e d . T h e s e s t a n d a r d s a p p e a r t o b e appropriate for pearl millet a s w e l l , a s s u m i n g "eyeball" estimates
of the deviations of plants f r o m the varietal m e a n . T a b l e 1 indicates suitable plant characters for inclusion
in t h e varietal d e s c r i p t i o n of pearl millet open-pollinated cultivars. For e x a m p l e , a n y plant that is clearly
shorter a n d later or earlier f l o w e r i n g t h a n a v e r a g e c a n safely be r o g u e d out. H o w e v e r , strict numerical
a d h e r e n c e to t h e s e s t a n d a r d s for t h e p u r p o s e of pearl millet s e e d certification w o u l d not be a p p r o p r i a t e .

S e e d Multiplication by Farmers

W i t h o p e n - p o l l i n a t e d cultivars, it is possi b l e for f a r m e r s to p r o d u c e their o w n s e e d . It is prefer able, for


m a i n t e n a n c e of varietal identity, that it is d o n e only f r o m crops g r o w n directly f r o m certified s e e d .
A l t h o u g h s o m e o u t c r o s s i n g with other pearl millet is likely to occur w h e n f a r m e r s multiply their o w n s e e d ,
multiplication by t h e m helps very m u c h in p r o m o t i n g the wider use of i m p r o v e d cultivars. B e c a u s e m a n y
f a r m e r s will multiply their o w n s e e d a n y w a y , it is helpful to provide advice to t h e m — i n their local
l a n g u a g e , if p o s s i b l e — t o assist t h e m in obtaining s e e d that is as g e n u i n e a n d as v igorous as possible.
Points that s h o u l d b e n o t e d include:

• W h e r e irrigation is poss ible, a small off-season s e e d multiplication plot will give excellent isolation
f r o m c o n t a m i n a t i n g pollen of other pearl millet. In t h e n o r m a l crop s e a s o n , v e r y early s o w i n g is
d e s i r a b l e , in fields at least 4 0 0 m f r o m fields of other pearl millet cultivars. In either c a s e ,
multiplication s h o u l d only be t a k e n up in fields that did not h a v e a pearl millet c r o p the previous
season.

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• Select panicles for s e e d f r o m standing plants in the field, before the general harvest.

• Collect panicles for s e e d only f r o m the middle portion of the field, providing at least 10 border
rows on all sides. This d o e s not apply to small off-season irrigated plots.

• C h o o s e panicles with c l e a n , well-formed grain f r o m typical, a v e r a g e plants. Do not h a r v e s t s e e d


f r o m early, late, extra-tall, or d i s e a s e d plants. T h e best plants a re likely to be o u t c r o s s e s , so it
is best to leave s e e d f r o m t h e m in the field if y o u are to retain t h e typical p e r f o r m a n c e of the
cultivar.

• C h o o s e at least 5 0 0 panicles, a n d dry t h e m thoroughly. T h r e s h these panicles in a specially


c l e a n e d place to avoid contamination with other s e e d . Treat the s e e d with insecticide a n d store
it in s e a l e d c o n t a i n e r s , with labels inside a n d outside.

• If the p e r f o r m a n c e of the i m p r o v e d cultivar is to be m a i n t a i n e d , it is a d v i s a b l e that f a r m e r s only


g r o w their o w n s e e d for o n e generation after p u r c h a s e of certified s e e d . T h u s village-level
f a r m e r - s e e d multipliers s h o u l d buy a n d s o w 1 -2 kg of certified s e e d every y e a r to g e n e r a t e s e e d
for their o w n use a n d for sale to their neighbors.

Other Considerations in Planning Seed Production

If g o o d quality s e e d is to reach farmers on time a n d in the quantities n e e d e d , it is n e c e s s a r y to do m o r e


t h a n s i m p l y follow the pro p e r p r o c e d u r e s for m a i n t e n a n c e a n d s e e d p r o d u c t i o n . It is also n e c e s s a r y to
m a i n t a i n reserve s t o c k s , to multiply the s e e d in appropriate e n v i r o n m e n t s , a n d to d e v e l o p guidelines to
d e t e r m i n e h o w m u c h s e e d o f e a c h cultivar s h o u l d b e p r o d u c e d .

Reserve Stocks

In a ny s e e d p r o d u c t i on p r o g r a m , the n e e d to s a v e a n d store e n o u g h reserve s e e d is well r e c o g n i z e d .


This g u a r d s a g a i n s t losses resulting f r o m crop failures or o t h e r disasters. R e s e r v e s t o c k s h e l p e n s u r e
continuity of a s e e d multiplication p r o g r a m . T h e r e s h o u l d be e n o u g h n u c l e u s s e e d in c o l d st orage
(preferably at t w o different places) to s o w at least t w o nucleus s e e d multiplication plots of a p p r o p r i a t e
size. Similarly, there s h o u l d be reserves of b r e e d er s e e d a n d f o u n d a t i o n s e e d a d e q u a t e for s o w i n g at
least t w o g e n e r a t i o n s .

Location of Seed Multiplication Fields

If open-pollinated cultivars a n d topcross pollinators are multiplied in a r e a s outside their z o n e of


a d a p t a t i o n , rapid shifts c a n occur in their genetic m a k e u p , phenotypic characteristics, a n d p e r f o r m a n c e .
T h u s , m a i n t e n a n c e of their nucleus s e e d , a n d multiplication of breeder s e e d a n d f o u n d a t i o n s e e d , is best
t a k e n up in e n v i r o n m e n t s that permit reproduction of all plants. Plots of these s e e d multiplication s t a g e s
s h o u l d be located in an a r e a to w h i c h the pearl millet g e n o t y p e is a d a p t e d . Since pearl millet is generally
g r o w n u n d e r d r y l a n d conditions during the rainy s e a s o n , rain at harvest is an ever-present risk w h e n this
p r o c e d u r e is a d o p t e d in order to maintain the genetic identity of these o p e n - p o l l i n a t e d g e n o t y p e s .
A r r a n g e m e n t s s h o u l d be m a d e for artificial drying of panicles h a r v e s t e d f r o m nucleus s e e d , breeder s e e d ,
a n d f o u n d a t i o n s e e d plots, w h e r e possible, in order to e n s u r e that the physical quality of the h a r v e s t e d
s e e d is also m a i n t a i n e d . M a i n t e n a n c e of local adaptation is generally well t a k e n c a r e of w h e n s e e d is
multiplied locally for f armer-t o-f armer sale or e x c h a n g e , outside t h e formal s e e d s y s t e m . T h i s restriction
on t h e s e e d multiplication e n v i r o n m e n t is less important for certified s e e d . T h e m a i n t e n a n c e a n d

77
multiplication of different classes of s e e d in their a r e a of a d a p t a t i o n , a n d u n d e r a p p r o p r i a te m a n a g e m e n t ,
also helps p r o v i de larger quantities of s e e d .

Guidelines for Determining Seed Requirements

To a v o i d s h o r t a g e s or s u r p l u s e s of s e e d , it is important to plan r e a s o n a b l e a n d a d e q u a t e s e e d
multiplication. T h e r e q u i r e m e n t s of different s e e d classes s h o u l d be k n o w n in a d v a n c e for e a c h cultivar
a n d p a r e n t . It c a n be c a l c u l a t e d as per details given in T a b l e 3. T h e f o l l o w i n g factors s h o u l d also be
considered:

• A r e a to be c o v e r e d by a released cultivar b a s e d on its adaptation a n d available alternatives;

• S e e d - r e p l a c e m e n t i n t e r v a l — o n e year for hybrids a n d o n e to three years for o p e n - p o l l i n a t e d


cultivars;

• S e e d g e n e r a t i o n s required to p r o d u c e certified s e e d ;

• Pdtential r e q u i r e m e n t s for certified s e e d — t h i s d e t e r m i n e s r e q u i r e m e n t s for f o u n d a t i o n s e e d a n d


breeder seed;

• L a n d r e q u i r e m e n t s for p r o d u c t i o n of different s e e d c l a s s e s ; a n d ,

• S e e d i n g rate (lower t h a n that u s e d in c o m m e r c i a l grain production) for production of breeder


s e e d , f o u n d a t i o n s e e d , a n d certified s e e d .

Conclusion

Multiplication of s e e d of o p e n - p o l l i n a t e d cultivars a n d t o p c r o s s pollinators of pearl millet is relatively


simple as the cultivars are vigorous a n d the s e e d multiplication rates are h i g h . M a i n t e n a n c e of a d e q u a t e
isolation d i s t a n c e , a n d m a n a g e m e n t of labor operations to minimize pollen m o v e m e n t b e t w e e n s e e d
multiplication plots of different cultivars is critical if true-to-the type s e e d is to be p r o d u c e d . W h e n b r e e d e r
s e e d is u s e d for multiplication of foundation s e e d , or its equivalent, prior to certified s e e d multiplication,
pearl millet b r e e d e r s e e d r equirements are small, a n d the nucleus s e e d class is not necessary.
T h e r e f o r e , p r o c e d u r e s indicated for nucleus s e e d production can be f o l l o w e d for p r o d u c t ion of breeder
s e e d itself. If sufficient care is t a k e n in the nucleus s e e d , breeder s e e d , a n d f o u n d a t i o n s e e d s t a g e s of
multiplication, little r o g u i n g will be required in t h e certified s e e d production plots. F a r m e r s c a n multiply
their o w n s e e d of o p e n - p o l l i n a t e d cultivars, a n d s h o u l d be e n c o u r a g e d to do so in a w a y that t h e genetic
integrity of t h e cultivar is m a i n t a i n e d .

78
Seed-Processing Techniques and Chronology of
Indian Seed Industry

Production of Foundation and Certified Seeds of Sorghum and


Pearl Millet
V Jaya Mohan Rao

T h e strength of crop production p r o g r a m s d e p e n d s on the success of s e e d production p r o g r a m s . In the


a b s e n c e of good-quality s e e d s , the investment on other inputs m a y not bring a b o u t the d e s i r e d results.
T h i s chapter deals with the organizational structure a n d planning required for production of s e e d , s e e d
certification s t a n d a r d s , a n d s e e d - p r o c e s s i n g p r o c e d u r e s .

Organizational Setup

It is n e c e s s a r y to h a v e an a p e x s e e d organization at the national level w h i c h monitors the s e e d p r o g r a m s


in t h e country, t a k e s care of training n e e d s , a n d functions as a nodal a g e n c y for s e e d buffer s t o c k i n g a n d
other special s c h e m e s .

E a c h state s h o u l d h a v e a s e e d corporation that looks after the s e e d r e q u i r e m e n ts of f a r m e r s in


that state. Initially, s u c h corporations c a n be set up in a f e w important states, a n d after w a t c h i n g their
p e r f o r m a n c e , similar corporations c a n be established in other states, incorporating suitable modifications,
if n e c e s s a r y . T h e s e c o r p o r a t i o n s m u s t not only h a v e a service function but must also be self-s upporting.
T h e s e state corporations s h o u l d p r o d u c e a n d supply s e e ds to f a r m e r s in the state. S e e d p r o d u c e d in
other states c a n be o b t a i n e d t h r o u g h the national organization or by direct a r r a n g e m e n t with other state
corporations.

E a c h state corporation must decide about its product mix. For profitability, it can p r o d u c e h i g h -
v a l u e , l o w - v o l u m e s e e d s , e.g., hybrid seeds of different crops (cotton, millets, o i l s e e d s , v e g e t a b l e s , etc.)
besides p r o d u c i n g s e e d s of self-pollinated crops as a social obligation to f a r m e r s .

S e e d m a y be supplied to different a g e n c i e s u p o n full p a y m e n t of c a s h , or alternatively, u p o n


p a y m e n t of 5 0 % c a s h s u p p o r t e d by a bank guarantee for the other 5 0 % of the a m o u n t . S e e d s h o u l d not
be s o l d on a c o n s i g n m e n t basis. Timely positioning of the s e e d in the m ar k et at least 3 0 - 4 0 d a y s a h e a d
of the s o w i n g s e a s o n in the c o n s u m i n g centers is very important. A plan for the s upply of s e e d s a n d an
estimate of s e e d r e q u i r e m e n t b a s e d on the s e e d - r e p l a c e m e n t rate s h o u l d be p r e p a r e d .

Seed Subsidy

To popularize the use of newly-released varieties/hybrids, it is necessary to provide a subsidy on certified


s e e d at s o u r c e . T h i s will p r o m o t e the use of certified s e e d w h i c h will in turn help increase crop
production.

79
Import and Export of Seed

T h e p a r e n t material for p r o d u c i n g varieties/hybrids suitable to a country m a y have to be i m p o r t e d f r o m


other c o u n t r i e s . T o m e e t t h e i m m e d i a t e d e m a n d , t h e finished products o f superior varieties/hybrids c a n
be i m p o r t e d after preliminary testing by research institutions a n d after c o n d u c t i n g test m a r k e t i n g . India
i m p l e m e n t e d a N e w S e e d Policy in October 1988, liberalizing the import p r o c e d u r e s for s e e d s .

S e e d p r o d u c t i o n of p r o m i s i n g varieties c a n be t a ken up by the importing countries if they have


strong national a n d state production p r o g r a m s . Alternatively, s o m e countries c a n take up production in
collaboration with foreign c o m p a n i e s . T h e National S e e d C o r p o r a t i o n , a G o v e r n m e n t of India
o r g a n i z a t i o n , e x t e n d s a s s i s t a n c e to other d e v e l o p i n g countries in formulating s e e d projects.

If t h e climatic conditions permit it, a country c a n take up production of s e e d s of hybrids,


o r n a m e n t a l plants, oilseeds, v e g e t a b l e s , fibre c r o p s , etc. T h e s e s e e d s c a n also be p r o d u c e d for export,
after f o r m u l a t i n g a suitable s e e d policy.

Planning

S e e d p r o d u c t i o n p r o g r a m s s h o u l d be t a k e n up in areas that are suitable for pr oduction of good-quality


s e e d s . T h e s e e d plots m u s t h a v e the required isolation distances in order to maintain the purity of the
s e e d . S e e d p r oduc t i o n projects m a y be taken up in collaboration with enterprising f a r m e r s in a r e a s w h e r e
s e e d yields are high a n d it is m o r e e c o n o m i c a l to do s o .

T h e s e e d p r o d u c t i o n p r o g r a m s h o u l d be c o n c e n t r a t e d in a f e w clusters of 2-3 c o n t i g u o u s villages


w i t h a b o u t 2 0 0 h e c t a r e s . T h e n u m b e r of s u c h clusters to be included in t h e p r o g r a m d e p e n d s on the total
s e e d requirement. U s i n g c o m p a c t blocks for s e e d production involves f e w e r isolation p r o b l e m s a n d
m a k e s s u p e r v i s i o n a n d m a i n t e n a n c e of quality e a s y . S u c h clusters will also solve the nicking p r o b l e m
to a l a r g e e x t e n t , a n d c a n a l s o s e r v e as d e m o n s t r a t ion b l o c k s .

S e e d p r o d u c t i o n p r o g r a m s s h o u l d be so p l a n n e d as to m a k e certified s e e d available at least 35-


45 d a y s b e f o r e t h e b e g i n n i n g of t h e s o w i n g s e a s o n . To e n s u r e this, it is n e c e s s a r y to fix a target date
for e a c h s t e p of t h e s e e d p r o d u c t i o n p r o c e s s , i.e., p r o c u r e m e n t of s e e d f r o m the s e e d p r o d u c i n g f a r m e r s
t o p r o c e s s i n g , certification, a n d p a c k i n g .

D u r i n g p r o c e s s i n g , t he s e e d m u s t b e thoroughly dri e d . E x c e s s m o i s t u r e m a y lead t o pest


infestation, f u n g a l d e v e l o p m e n t , deterioration in quality, a n d loss in g e r m i n a t i o n . T h e s e e d m u s t be
p a c k e d suitably (cloth b a g s , polythene p o u c h e s , cartons, tins, etc.) to hold sufficient s e e d to s o w 0.2 ha
or 0.4 ha as p e r the p r e f e r e n c e of the f a r m e r s .

It is desirable to h a v e m o r e small processing plants of 1000-20 0 0 0 tons capacity rather than o n e


with large-capacity. S u c h plants s h o u l d b e e s tablished near the s e e d - p r o d u c i n g a r e a s t o s a v e o n
transportation a n d facilitate s upply to farmers well before the crop s e a s o n begins. T h e p r o c e s s i n g plant
s h o u l d h a v e e n o u g h s p a c e to store at least 5 0 % of the capacity of t h e plant at a n y giv en t i m e .
M a i n t e n a n c e of s t o c k s a n d f u m i g a t i o n against storage pests will be easier, a n d o v e r h e a d expenditure
will be m i n i m i z e d . T h e p r o c e s s i n g plant must h a v e a quality-control laboratory.

S e e d c a n be s t o r e d in transit g o d o w n s or at the p r o c e s s i n g plant itself before d e s p a t c h . It is


n e c e s s a r y to store t h e s e e d in a dry p l a c e . S e e d stored in a h u m i d a r e a will lose g e r m i n a t i o n faster;
t heref ore, it s h o u l d be shifted to a dry place immediately after p r o c e s s i n g a n d b a g g i n g . S e e d b a g s m u s t

80
be s t a c k e d on w o o d e n pallets. C a r e must be taken to see that the n u m b e r of stacks of s e e d b a g s d o e s
not e x c e e d t h e r e c o m m e n d e d limit so that the germination of s e e d in the lower b a g s is not affected. T h e
s e e d m u s t be p r o t e c t e d by spraying the following insecticides at regular intervals:

M a l a t h i o n ® 50 E C : 1 part:300 parts of water; apply @ 3 L 100 m 2 of surface a r e a . D D V P ®


(0.25%) 2.5 mL in 1 L water; spray 100 m - 2 s p a c e .

In a d d i t i o n , f u m i g a t i o n s are d o n e with a l u m i n u m phosphide with a d o s a g e of 1 tablet of 3 g ton - 1 ,


under airtight conditions. This d o s a g e can be increased to 2 tablets ton' 1 if the g o d o w n s required to be
f u m i g a t e d are r eas onably airtight.

Procurement of Seed

Estimation of s e e d d e m a n d is a c o m p l e x matter. It d e p e n d s on s e a s o n a l conditions, particularly rainfall


p a t t e r n , prices, varietal p r e f e r e n c e , s e e d availability with farmers, marketing a n d distribution c h a n n e l s ,
p a c k i n g s i z e , natural calamities, etc.

To estimate the s e e d requirements for different crops in a state/country, the p r o b a b l e r e p l a c e m e n t


rate of s e e d of e a c h crop s h o u l d be d e t e r m i n e d , i.e., a decision has to be taken by the g o v e r n m e n t with
regard to t h e p e r c e n t a g e of the a r e a under high-yielding varieties of e a c h crop for w h i c h s e e d is to be
s u p p l i e d . For e x a m p l e , in a self-pollinated crop like rice in A n d h r a P r a d e s h (India), f a r m e r s prefer to
replace their s e e d after every three y e a r s . T h u s , the s e e d r eplacement rate for rice w o u l d be 3 3 % i.e.,
sufficient s e e d must be s upplied every year to cover 3 3 % of the total area under high-yielding varieties
of this c r o p . Besides a s s e s s i n g the s e e d requirement on the basis of the s e e d replacement rate, realistic
s e e d d e m a n d c a n b e a s s e s s e d t h r o u g h different channels.

S e e d d e m a n d c a n be a s s e s s e d with the help of the De partment of Agriculture, s e e d dealers,


c o o p e r a t i v e s or allied societies that deal with s e e d . T h e a m o u n t of carry-over stocks available in t h e
country s h o u l d also be e s t i m a t e d . This estimate s h o u l d be d o n e by the s e e d - m o n i t o r i n g division of t h e
central g o v e r n m e n t . T h e a s s e s s m e n t of local requirement of s e e d c a n be d o n e with the help of village-
level a g e n c i e s that distribute other inputs like fertilizers to the f a r m e r s . It will be e a s y for s u c h a g e n c i e s
(e.g., c o o p e r a t i v e society) to find out the required quantity of s e e d , variety-wise. T h e village level
information c a n be sent up to the district level, a n d o n w a r d to the state-level authorities. T h e s e e d
requirements of various crops in e a c h state c a n thus be a s s e s s e d a n d c o m p i l e d by the state
administration.

Buffer Stock

It is n e c e s s a r y to keep a buffer stock of s e e d to meet contingencies s u c h as natural calamities a n d to


s a f e g u a r d a g a i n s t shortfalls in s e e d p r o d u c t i o n .

T h e quantity of s e e d to be kept in buffer should not be less than 1 0 % of the projected requirement
in the s u b s e q u e n t year in the c a s e of certified s e e d , 2 0 % in the c a s e of f o u n d a t i o n s e e d , a n d 5 0 - 1 0 0 %
( d e p e n d i n g u p o n t h e hybrids/varieties) in the c a s e of br eeder s e e d . In the s e e d industry, the quantity of
carry-over stocks m u s t be kept at a m i n i m u m .

T h e operational expenditure (local transportation of s e e d , storage cost a n d interest c h a r g e s , a n d


also t h e difference in cost in c a s e the s e e d is to be sold as grain) is to be b o r n e by t h e central
g o v e r n m e n t . This will help in the successful im plementation of overall crop-production p r o g r a m s during

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national ca l a m i t i e s .

Certification Standards

Sorghum

T h e p r o d u c t i o n a n d h a n d l i ng o f f o u n d a t i on s e e d a n d certified s e e d m u s t b e s u p e r v i s e d a n d a p p r o v e d
by the certification a g e n c y . T h e s e e d must meet the standards prescribed by the certification a g e n c y .

Eligibility Requirement for Certification

• S e e d of varieties a n d hybrids m e a n t for certification s h o u l d be f r o m a s o u r c e that is reliable a n d


a p p r o v e d by the certification a g e n c y .

Land Requirements

• T h e s e e d p r o d u c t i o n field offered for certification s h o u l d be irrigated at least three w e e k s before


s o w i n g , a n d p l o w e d sufficiently a h e a d of s o w i n g to destroy volunteer s e e d of s o r g h u m .

• C a r e s h o u l d be t a k e n to r e m o v e Sorghum helepense s e e d in the field a n d within the isolation


distance.

Field Inspections

• A m i n i m u m of four field inspections s h o u l d be m a d e during the crop p e r i o d :

First i n s p e c t i o n : During preflowering stage


S e c o n d a n d third inspections: During flowering stage
Fourth inspection: During preharvest stage

Field standards

T h e m i n i m u m field s t a n d a r d s a n d a c c e p t a b l e s e e d standards are given in Tables 1-4.

Table 1. M i n i m u m isolation d i s t a n c e re q u i red for f o u n d a t i o n a n d certified s e e d o f s o r g h u m .

M i n i m u m isolation distance (m)

Field Foundation s e e d field Certified s e e d field

O t h e r grain s o r g h u m fields 300 200

Field of s a m e variety not c o n f o r m i n g to


varietal purity 300 25

F o r a g e s o r g h u m a n d J o h n s o n grass 400 400

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Table 2. M a x i m u m p e r m i t t e d level o f pollen s h e d d e r s , off-types, a n d d i s e a s e d h e a d s for
f o u n d a t i o n a n d certified s e e d o f s o r g h u m .

M a x i m u m permitted level (%)

Foundation s e e d field Certified s e e d field

Pollen s h e d d e r s 0.05 0.1

Off-types in b o t h s e e d parent
a n d pollinator 0.01 0.05

D i s e a s e d h e a d s like grain s m u t
a n d h e a d s m u t at final inspection 0.05 0.1

* H o w e v e r , reinspections are a l l o w e d after removal of d i s e a s e d plants.

Table 3. S e e d s t a n d a r d s for f o u n d a t i o n a n d certified s e e d o f s o r g h u m .

Seed standard Foundation s e e d Certified s e e d


(%) (%)

S e e d purity ( m i n i m u m ) 98 98
Inert matter ( m a x i m u m ) 2 2
Other crop s e e d ( m a x i m u m ) 0.05 0.1
Weed seed (maximum) 0.05 0.1
Germination (minimum) 80 80
Moisture ( m a x i m u m ) 12 12

Table 4. S c r e e n s i z e s p r e s c r i b e d for c l e a n i n g o f s o r g h u m s e e d s .

Screen Round Slotted

Top screen 1 2 / 6 4 " or


4.75 m m --

Bottom s c r e e n 9 / 6 4 " or 1/12" x 1/2"


3.55 m m or 2.1 m m

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Pearl Millet

Eligibility Requirements for Certification

• An inbred line, to be eligible for certification, s h o u l d be f r o m a reliable s o u r c e a n d a p p r o v e d by


t h e certification a g e n c y .

• H y b r i d s e e d s h o u l d be the F 1 p r o g e n y of t w o a p p r o v e d inbred lines, o n e of w h i c h s h o u l d be a


male-sterile line.

Classes and Sources of Seed

• An inbred line s h o u l d be a relatively true-breeding strain resulting f r o m self-pollination a n d


selection.

• F o u n d a t i o n s e e d s h o u l d be the p r o g e n y of an a p p r o v e d male-sterile line a n d an a p p r o v e d restorer


line (the m a l e parent) for t h e p u r p o s e of pr oducing hybrid s e e d .

• A male-sterile line s h o u l d be a strain carrying cytoplasmic-genic m a l e sterility.

• T h e certified s e e d class s h o u l d be the hybrid s e e d that can be s o w n for a n y use e x c e p t s e e d


production.

Land Requirements

• T h e land to be u s e d for hybrid s e e d production of pearl millet should be free of volunteer plants.

Field Inspection

• A m i n i m u m of four inspections should be m a d e .

• T h e first inspection s h o u l d be m a d e before flowering, preferably within 30 d a y s after s o w i n g in


or der to c h e c k isolation, volunteer plants, outcrosses, s o w i n g ratio, errors in s o w i n g , incidence
of d o w n y m i l d e w , a n d other relevant factors.

• T h e s e c o n d a n d third inspections s h o u ld be m a d e during f l o w e r i ng to c h e c k pollen s h e d d e r s , off-


t y p e s , d o w n y m i l d e w / g r e e n ear, a n d other relevant factors.

• T h e fourth inspection s h o u l d be m a d e at maturity a n d prior to harvesting in order to detect the


i n c i d e n ce of d o w n y m ildew/green ear, ergot, a n d smut, a n d to verify the true nature of the plant,
a n d other relevant factors.

Field Standards

• S e e d fields s h o u l d be adequately isolated f r o m other fields as listed in T a b l e 5.


T a b l e 5. G e n e r a l field s t a n d a r d s for pearl millet s e e d p r o d u c t i o n .

M i n i m u m distance (m)

Field Foundation s e e d field Certified s e e d field

Fields of other varieties including c o m m e r c i a l


hybrids of the s a m e variety 1000 200

Fields of t h e s a m e hybrid (code designation)


not c o n f o r m i n g to varietal purity
r e q u i r e m e n t s for certification 1000 200

Fields of other hybrids h a v i n g a c o m m o n male


parent a n d c o n f o r m i n g to varietal purity
r e q u i r e m e n t s for certification — 5

Fields of other hybrids h a v i n g a c o m m o n male


parent but not c o n f o r m i n g to varietal
purity r e q u i r e m e n t s for certification — 200

• S e e d fields s h o u l d be free f r o m a n y contaminants as listed in Table 6.

T a b l e 6. Specific r e q u i r e m e n t s for pearl millet s e e d p r o d u c t i o n .

M a x i m u m permitted (%)

Factor Foundation s e e d Certified s e e d

Off-types in s e e d parent at a n y one inspection


a t a n d after f l o w e r i n g 0.050 0.10

Off-types in pollinator at a n y o n e inspection


at a n d after f l o w e r i n g 0.050 0.10

P o l l e n - s h e d d i n g h e a d s in s e e d parent at any
o n e inspection at flowering 0.050 0.10

Plants infected by d o w n y m i l d e w / g r e en ear


d i s e a s e at a n y o n e inspection 0.050 0.10

Ergot e a r h e a d s p a r e n t at final inspection 0.020 0.040

E a r h e a d s infected by grain s m u t
at final inspection 0.050 0.10

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• S e e d f r o m s u c h fields that have b e e n reported to contain ergot i n f e c t i o n — e v e n within the
p r e s c r i b e d limits—at field s t a g e s h o u l d b e s u b j e c t e d t o floatation t r e a t m e n t w i t h brine t o b e c o m e
eligible for certification.

• S e e d fields with incidence of grain s m u t more than the m a x i m u m permissible level c a n , however,
be certified if s u c h s e e d is treated with an a p p r o v e d o r g a n o - m e r c urial fungicide not earlier t h a n
a m o n t h prior to its s o w i n g .

Seed Processing

In t he b r o a d e s t s e n s e , s e e d p r o c e s s i n g e n c o m p a s s e s all the steps involved in the preparation of


h a r v e s t e d s e e d for m a r k e t i n g , i.e., handling, shelling, preconditioning, dry i n g , c l e a n i n g , s i z e - g r a d i n g ,
u p g r a d i n g , treating, a n d p a c k a g i n g . S e e d processing requires e q u i p m e n t a n d m a c h i n e r y , c o n v e y e r s a n d
structures w h i c h require m e c hanical skills a n d engineering principles.

A variety of c o n t a m i n a n t s must be r e m o v e d f r o m the raw s e e d s u c h as inert matter a n d


u n d e r s i z e d s e e d s , w h i c h are not in t h e m s e l v e s harmful but influence s e e d flow ability a n d plantability.
Insect-infested s e e d contribute t o storage p r o b l e m s , a n d w e e d a n d other crop s e e d s seriously affect crop
p r o d u c t i o n if they are not r e m o v e d .

A variety of e q u i p m e n t is available for processing s e e d . It ranges from the simple, indigenous


w i n n o w i n g a n d sieving trays to t h e sophisticated electric sorting m a c h i n e s . T h o u g h differing in type a n d
d e s i g n , all p r o c e s s i n g m a c h i n e r y h a v e o n e thing in c o m m o n , i.e., separation of g o o d s e e d s f r o m ill-filled
a n d u n d e r s i z e d s e e d s a n d other undesirable material (inert matter). D e p e n d i n g on the quality, the s e e d
requires to be p r o c e s s e d in a definite s e q u e n c e by several m a c h i n e s , e a c h r e m o v i n g a certain type of
u n w a n t e d mater i a l . T h e c h o i c e of m a c h i n e s for processing s e e d d e p e n d s on the kind of s e e d being
p r o c e s s e d , t h e nature a n d kind of u n w a n t e d materials ( w e e d s e e d , other crop s e e d , inert matter, etc.)
in it, the quantity of e a c h in the raw s e e d , a n d the quality standards that must be met. T h u s , the
processor m u s t be as familiar with s e e d st andards a n d s e e d characteristics as he is with the processing
equipment.

Steps in Processing

S e e d - p r o c e s s i n g operations c a n be divided into several definite steps that must be carried out in a
specific s e q u e n c e . T h e first step is receiving the s e e d . S e e d arrive at the p r o c e s s i n g plant in bags or
in bulk. F r o m t h e receiving station, the seeds go into bulk storage to be held for later p r o c e s s i n g , or
directly into t h e p r o c e s s i n g line for c l e a n i n g . T h e next step is conditioning a n d preclearing. T h i s includes
r e m o v a l of a p p e n d a g e s a n d large pieces of trash.

T h e first actual cleaning step is basic cleaning. T h e air-screen m a c h i n e is the m o s t c o m m o n basic


cleaner. It m a k e s size separations a n d aspirates the s e e d . Often, it may be n e c e s s a r y to s e n d the s e e d
t h r o u g h o n e or m o r e special separating or upgrading machines to r e m o v e a specific c o n t a m i n a n t . T h e s e
special m a c h i n e s s e p a r a t e c r o p a n d w e e d s e e d by differences in their specific physical characteristics.
W h e n all possible inert matter a n d w e e d or other crop s eeds have b e e n r e m o v e d , the s e e d s a r e ready
for b a g g i n g . A f ungicide or insecticide treatment is applied before p a c k i n g . T h e s e e d s m a y then be
s h i p p e d to other p l a c e s or held in storage.

S e e d p r o c e s s i n g is b a s e d on differences in physical properties b e t w e e n the desirable s e e d a n d


the c o n t a m i n a t i n g s e e d . S e e d s that do not differ in s o m e physical characteristic c a n n o t be s e p a r a t e d .

86
If a difference exists b e t w e e n the s e e d s a n d a m a c h i n e is available w h i c h c a n differentiate b e t w e e n t h e m
in a consistent m a n n e r , t h e n t h e y c a n be s e p a r a t e d . S e e d processors c a n c h o o s e f r o m a w i d e selection
of m a c h i n e s that differentiate b e t w e e n s e e d s differing in size, length, s h a p e , weight, surface, texture,
color, affinity for liquids or conductivity. A single m a c hine cannot separate seeds that differ in all these
characteristics. In m o s t instances, a different machine must be used to m a k e separations b a s e d on e a c h
of these characteristics.

Size. Size i s the m o s t c o m m o n difference a m o n g s eeds, a n d b e t w e e n s e e d a n d undesirable material.


T h e air-screen m a c h i n e e n a b l e s g r ading of the s e e d into different sizes d e p e n d i n g u p o n w i d t h a n d
thickness by t h e use of a series of perforated shutters. O n e or m o r e air-blast separations r e m o v e light
material.

L e n g t h . L e n g t h differences are c o m m o n b e t w e e n different crop s e e d s a n d w e e d s e e d s . L e n g t h


separators are u s e d by m a n y processors to upgrade a n d improve the quality of s e e d . B o t h t h e i n d e n t e d
cylinder a n d disc s eparator m a k e length separations.

S h a p e . S h a p e varies widely a m o n g s e e d s . T h e separations m a d e by the air-screen m a c h i n e are often


related to differences in s h a p e , especially w h e n triangular-hole scr e e n s are u s e d . T h e i n d e n t e d cylinder
a n d t h e disc s e p a r a t o r t a k e a d v a n t a g e of s h a p e differences, particularly w h e n they are a function of
length. T h e r e is another m a c h i n e , k n o w n as spiral separator, specially d e s i g n e d to s e p a r a t e r o u n d f r o m
flattened s e e d s .

M a s s . M a n y s e e d s differ in m a s s , specific gravity or relative density per given unit of v o l u m e . M a s s or


specific gravity is the basis of the air-blast separations in an air-screen m a c h i n e . Howev er, the gravity
separator, the stonier, the aspirator, a n d the pneumatic separator are all d e s i g n e d to m a k e specific
separations by differences in m a s s or specific gravity of s e e d .

S u r f a c e t e x t u r e . T h e relative ro u g h n e s s or s m o o t h n e s s of the s e e d coat, i.e., surface t e x t u r e , is a


c o m m o n difference b e t w e e n s e e d s . T h e roll or dodder mill, the draper belt, the magnetic separator, a n d
the vibrator separator, all effect separation of s e e d s differing in surface texture.

Color. M a n y s e e d s differ in color or reflectivity. Electronic color sorters are u s e d to m a k e color


separations in the larger c r o p s e e d s .

Affinity for liquids. S e e d s also differ in their affinity for liquids, or the rate at w h i c h their surface will
a b s o r b liquids. T h e m agnetic separator, separates the s e e d s on the basis of t h e s e differences.

C o n d u c t i v i t y . S e e d s also differ in their ability to hold or conduct an electrical c h a r g e . T h e m a c h i n e w h i c h


s e p a r a t e s s e e d on the basis of electrical properties is called electrostatic separator, a n d consists
essentially of a c o n v e y o r belt w h i c h carries a single layer of s e e d b e n e a t h an electrode.

87
Harvest and Postharvest Handling of Breeder Seed of Sorghum
and Pearl Millet
C T Hash

Important s t a g e s in the p r o c e s s of safe a n d efficient m o v e m e n t of breeder s e e d from the breeder s e e d


production field to the foundation s e e d producer include selection a n d harvest of panicles in t h e isolation
plot, drying the panicles, threshing a n d cleaning the s e e d , further drying prior to s t o r a g e , st orage in bulk,
p a c k a g i n g a n d d i s p a t c h to f oundation s e e d producers, a n d record k e e p i n g . During all t h e s e s t a g e s , it is
n e c e s s a r y to t a k e precautions to continue to maintain the purity of the individual breeder s e e d stocks a n d
prevent adulteration. Close supervision is necessary to ensure that the breeder s e e d that reaches the
f o u n d a t i o n s e e d p r o d u c e r s is viable, vigorous, a n d contains only few c o n t a m i n a n t s .

Harvest

T h e harvest of panicles f r o m a breeder s e e d production plot is the last but o n e opportunity that the
breeder has to control the f e m a l e parentage of the breeder seed-lot. Harvest is a far m o r e appropriate
stage for this t h a n the last opportunity, i.e., evaluation of harvested panicles prior to t h r e s h i n g . Harvest
s h o u l d be u n d e r t a k e n as s o o n as possible after the crop reaches physiological maturity. Harvest after
physiological maturity d e p e n d s on the facilities available for postharvest drying of the panicles. If only
s u n - d r y i n g facilities are there, then it is better to delay harvesting a w e e k or more after physiological
maturity before harvesting in order to allow the s u n a n d w i n d to reduce the moisture content of the s e e d .

Panicles s h o u l d normally be harvested f r o m the center of the breeder s e e d production plot. T h e


a m o u n t of border to be leaft d e p e n d s on the species a n d type of material being p r o d u c e d : O n e row for
s o r g h u m R-lines a n d open-pollinated cultivars, four rows or more for pearl millet open-pollinated varieties
a n d A / B pairs of b o t h s p e c i e s . Panicles selected for harvest should be standing plants that are typical
of t h e line. T h e best plants a n d the worst plants s hould both be left in the field. Early plants of pearl millet
are m o r e likely to bear s e e d p r o d u c e d by pollen from outside the plot; so these too s h o u l d not be
h a r v e s t e d for s e e d . Late plants, extra-tall plants, a n d diseased plants s h o u l d not be h a r v e s t e d . T h e
h a r v e s t e d panicles s h o u l d be collected in clean (preferably new) gunny b a g s , labelled inside a n d outside
with the n a m e of the material a n d the location w h e r e it w a s p r o d u c e d . H a r v e s t e d panicles s h o u l d be dried
a n d s t o r e d prior to threshing. During drying a n d stor age, care must be taken to k e e p panicles of different
g e n o t y p e s (e. g. , an A-line a n d its B-line maintainer) in separate physical facilities to a v o i d inadvertent
mixing.

Drying

T h e p u r p o s e of drying is to reduce the moisture content of the s e e d to facilitate threshing a n d cleaning,


to m a i n t a i n s e e d viability a n d vigor, a n d to reduce deterioration of the s e e d d u e to growt h of m o l d s , other
m i c r o b e s , a n d s t o r a g e insects. A t w o - s t a g e drying process is preferred. In the first s t a g e , panicles are
dried d o w n to a moisture content of 1 5 - 2 0 % w h e r e the s e e d is easily t h r e s h e d free f r o m t h e a t t a c h e d
floral structures ( l e m m a , palea, g l u m e s , etc.). But the s e e d is still not dry e n o u g h , a n d h e n c e , is easily
d a m a g e d . After t h r e s h i n g a n d c l e a n i n g , the s e e d s h o u l d be dried further—to a moisture content below
1 2 % — t o permit safe long-term storage. Drying offers the following a d v a n t a g e s :

• Early harvest, w h i c h r educes the risk of crop loss or d a m a g e d u e to rains at maturity;

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• L o n g - t e r m storage of the s e e d ;

• M o r e efficient use of land a n d staff/labor;

• U s e of stover as green fodder; a n d

• Production of better-quality s e e d .

S u n - d r y i n g a n d forced-air-drying are the t w o most c o m m o n l y u s e d m e t h o d s . S u n - d r y i n g c a n be


very effective in most s o r g h u m a n d pearl millet production environments, but forced-air-drying is not as
d e p e n d e n t o n t h e weather.

Sun-Drying

For s u n - d r y i n g , it is n e c e s s a r y to allow the standing crop to dry in the field for a w e e k or m o r e after
physiological maturity. T h e panicles are then harvested a n d allowed to dry further for several d a y s in the
sun over dry soil or tarpaulins or, preferably, on a c lean, dry, c e m e n t e d threshing floor. T h e s e e d c r o p
is then t h r e s h e d a n d w i n n o w e d before being s p r e a d in a thin layer on the threshing floor to dry further.
Stirring a n d turning the drying s e e d will s p e e d up the process. C a r e must be taken to a v o i d d a m a g i n g
the s e e d , for it gets m o r e brittle a n d fragile as its moisture content is r e d u c e d . During d r y i n g , o n e n e e d s
to take precautions against d a m a g e by storage insects, especially ants. T h e major a d v a n t a g e of
sun-drying c o m p a r e d with forced-air-drying is that the former requires m u c h less capital ex penditure a n d
limited operational e x p e n s e s . T h e d i s a d v a n t ages are that it delays harvest, w h i c h increases the risk of
w e a t h e r d a m a g e to the standing crop. Since the harvested crop is dried outdoors, the risk of w e a t h e r
d a m a g e continues until the drying process is complete. Finally, c o m p a r e d with forced-air-drying, there
is greater likelihood of physical admixtures w h e n the s e e d is sun-dried. T h e following precaut ions must
be t a k e n with s u n - d r y i n g :

• Use a c l e a n , dry, c e m e n t e d threshing floor;

• Only o n e crop variety, a n d produce from one plot, should be handled on one threshing floor at
any one time.

Forced-Air-Drying

In this p r o c e s s , f o r c e d air p a s s e s through the d a m p s e e d a n d picks up the water. T h e e v a p o r a t i n g


moisture f r o m the s e e d cools both the air a n d the s e e d . As the air cools, its w a t e r - holding capacity is
r e d u c e d , a n d the c o o l , moist air must be carried a w a y f r o m the s e e d . T h e r e f o r e , y o u n e e d a cont inuous
supply of w a r m , dry air to drive a w a y the cool a n d moist air, a n d dry the s e e d . This air n e e d not be
h e a t e d in m o s t s o r g h u m a n d pearl millet s e e d production e n v i r o n m e n t s , although it m a y s o m e t i m e s be
n e c e s s a r y . Forced-air-drying will be most effective w h e n the following factors are t a k e n into
consideration:

• Relative humidity: T h e ratio of the a m b i e n t v a p or density to the saturation vapor density at air
temperature;

• Moisture equilibrium b e t w e e n the s e e d a n d the air; a n d

• T h e t e m p e r a t u r e difference b e t w e e n the a m b i e n t air a n d h e a t e d air.

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Forced-air-drying c a n rely on natural air-drying, or s u p p l e m e n t al heat-drying in w h i c h air t e m p e r a t u r e s
a r e raised 5-10°C to r e d u c e t h e relative humidity a n d facilitate drying. W h e n using natural air-drying, it
is n e c e s s a r y to force f r e s h air t h r o u g h the drying s e e d during the w a r m , dry parts of t h e d a y . At night,
t e m p e r a t u r e s normally fall near the d e w point at w h i c h the saturation v a p o r density is 1 0 0 % . Forcing air
of 1 0 0 % humidity through the s e e d m a y increase rather than d e c r e a s e the moisture content. T h e s e
m e t h o d s will require 1-3 w e e k s to r educe the s e e d moisture content to a safe level ( < 1 2 % ) . A drying
t e m p e r a t u r e of 4 0 ° C is the m a x i m u m r e c o m m e n d e d for drying of s o r g h u m a n d pearl millet b r e e d e r - s e e d
and nucleus-seed stocks.

For c o m m e r c i a l s e e d p r o d u c t i o n , h e a t e d air drying is often u s e d . In this m e t h o d , air is h e a t e d to


a b o u t 4 0 ° C a b o v e the a m b i e n t air t e m p e r a t u r e before b e i n g f o r c e d t h r o u g h the s e e d .

Threshing and Cleaning

O n c e the s e l e c t e d h a r v e s t e d panicles have b e e n dried to a moisture content of a b o u t 1 2 % , t h e s e e d can


easily a n d safely be t h r e s h e d . Various w i n n o w i n g a n d cleaning operations can b e u n d e r t a k e n t o s e p a r a t e
the s e e d f r o m the chaff. T h r e s h i n g a n d w i n n o w i n g can be d o n e m anually or mechanically. If m a c h i n e s
a r e u s e d , t h e y m u s t be c l e a n e d a n d inspected before thres h i n g o f e a c h b r e e d e r seed-lot. C a r e must b e
taken not to t h r e s h the s e e d w h e n it is very dry as it is more easily injured. Similarly, care m u s t be taken
to use the m i n i m u m effective rotor s p e e d of the thresher a n d to m inimize the time that a given portion
of t h e s e e d s p e n d s in t h e t h r e s hing c h a m b e r in order to minimize injury to t h e s e e d .

C l e a n i n g is u n d e r t a k e n to separate undesirable materials f r o m the s e e d . T h e s e undesirable


materials c a n include:

• inert matter (e. g. , dirt, s t o n e s , a n d chaff);

• weed seed,

• other crop seed,

• light a n d chaffy s e e d , a n d

• off-size, d a m a g e d , or deteriorated s e e d .

S e e d c l e a n i n g is b a s e d on the physical properties of the s e e d a n d undesirable materials.


Properties import ant for s o r g h u m a n d pearl millet s e e d include:

• s e e d size (length, w i d t h , a n d thickness),

• density,

• shape, and

• surface texture.

For s o r g h u m a n d pearl millet, cleaning operations c a n be divided into three g r o u p s :

• preconditioning/precleaning using a scalper to r e m o v e large inert matter,

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• basic s e e d c l e a n i n g , using an air-screen cleaner, a n d

• u p g r a d i n g , preferably using a gravity separator.

After threshing a n d cleaning, the s e e d should be dried further, d o w n to a moisture content no


greater t h a n 1 2 % t o permit safe storage (Table 1).

T a b l e 1 . M o i s t u r e c o n t e n t o f s o r g h u m s e e d as a f u n c t i o n of relative h u m i d i t y .

Relative humidity (%) 15 45 75 100


Moisture (%) 6.4 10.5 15.2 21.9

Safe Not safe

Storage

B r e e d e r s e e d a n d n u c l e u s s e e d of pearl millet a n d s o r g h u m n e e d to be stored in a c o o l , dry place. W h e n


no humidity control is possible in the cool place that is available, dry s e e d must be stored in hermetically
s e a l e d c o n t a i n e r s (e.g., n e w paint cans). A g o o d seed-lot c a n be safely held for 20 years or m o r e under
s u c h conditions. For places w h e r e humidity control is possible, Table 1 provides an indication of the
moisture content of s e e d that equilibrates with the air in the store r o o m . It is the relative humidity of the
air in the store r o o m , not outside, that is tabulated. Safe s e e d storage will require that relative humidity
i n t h e store r o o m b e m a i n t a i n e d below 4 5 % .

Place n a p h t h a l e n e balls at the rate of 20 g 10 kg - 1 of breeder s e e d in the storage containers to


serve as a deterrent to storage insects. If insects are already present in the seed-lot, this t r e a t m e n t will
not eliminate t h e m .

T h e s e e d - s t o r a g e area must be kept clean to prevent breeding of insects, rodents, or fungi. S e e d


s h o u l d be d r i e d to a safe moisture level before being placed in the seed-storage area. Finally, it is only
reasonable to store only high-quality s e e d in the seed-storage area.

Packaging

B r e e d e r s e e d of s o r g h u m a n d pearl millet sh o u l d be p a c k a g e d in small lots (1-10 k g , d e p e n d i n g on t h e


material). This s e e d can be p a c k a g e d in a d v a n c e in standardized units, or p r e p a r e d f r o m bulk u p o n
receipt of t h e r equest f r o m a seed-multiplication a g e n c y . It is important that the p a c k a g i n g be c l e a n ,
moisture-proof, well-labelled, a n d not easily d a m a g e d in transit. T h e s e e d packet s h o u l d h a v e a t t a c h e d
(preferably) or e n c l o s e d , a b r e e d e r - s e e d tag indicating the c r o p , label number, variety, quantity, seed-lot
n u m b e r , s e e d class, a n d actual data indicating inert matter, germ ination, a n d genetic purity, with a date
for the germination test. In addition, the producing institution should be indicated, a l o n g w i t h the signature
of the b r e e d e r r e sponsible for multiplication of the seed-lot.

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Despatch and Planning

D e s p a t c h of the breeder s e e d in response to requests should be timely. This is possible only if br eeder
s e e d requirements are accurately forecast well in a d v a n c e , a n d the different cultivars a n d hybrid parental
lines are multiplied well in time. In India, seed-multiplication a g e n c i e s indicate their likely r equirements
over a year in a d v a n c e , a n d m a k e a p a y m e n t to the g o v e r n m e n t b a s e d on t h e quantities required.
P a y m e n t for the breeder s e e d itself is a separate matter. T h e g o v e r n m e n t compiles the requests, a n d
submits t h e totals to t h e institutions responsible for multiplying this breeder s e e d 8-10 m o n t h s before the
s e e d is actually required. This gives e n o u g h time to produce the required breeder s e e d .

Record Keeping

For e a c h s a m p l e of breeder s e e d sent out to seed-multiplication a g e n c i e s , it is n e c e s s a ry to maintain a


record. At I C R I S A T , a s e e d request n u m b e r is a s s i g n e d to e a c h request received. This n u m b e r is then
included in all c o r r e s p o n d e n c e related to the request. A register of requests is m a i n t a i n e d w h i c h includes
information on the date the request w a s received, w h o m the s e e d w a s sent to, what materials w e r e sent
a n d in w h a t quantities, w h e n the s e e d w a s d e s p a t c h e d , a n d h o w it w a s d e s p a t c h e d . In addition, a
b r e e d e r - s e e d register is m a i n t a i n e d in w h i c h e a c h d e s p a t c h of breeder s e e d is r e c o r d e d , in serial order
of the b r e e d e r - s e e d t a g . A cover letter is sent separately, indicating w h a t material w a s sent, b r e e d e r - s e e d
t ag n u m b e r a n d b r e e d e r - s e e d quantities, a n d requesting f e e d b a c k on the utility of the mater ial. Finally,
the cover letter is a c c o m p a n i e d by a b r e e d e r - s e e d certificate, in the f o rmat required by the state in w h i c h
t h e f o u n d a t i o n s e e d will be multiplied. S a m p l e s of b r e e d e r - s e e d tag a n d s e e d r e c o r d are g i v e n in Figure
1 a n d Figure 2.

C r o p — P e a r l Millet (Bajra) Label No. 6 5 3 9


Variety: ICTP 8 2 0 3 Qty.: 2 kg Lot No. BS 193-5
C l a s s of S e e d — B r e e d e r s S e e d

Date of Test: Siqnature

* Pure S e e d 99.8% * Germination 95%

* Inert matter 0.2% * Genetic Purity 100%

• B a s e d on actuals

P r o d u c i n g Institution:
International C r o p s R e s e a r c h Institute for the
S e m i - A r i d Tropics
Patancheru 502 324,
A . P . , India.

Figure 1. A s a m p l e of a breeder s e e d t a g .

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93
Purity Test of Hybrids, Parental Lines, and Open-Pollinated
Varieties of Sorghum and Pearl Millet
V Jaya Mohan Rao

Purity of s e e d is a v e r y important criterion for the a c c e p t a n c e of a variety. This is e n s u r e d t h r o u g h


various administrative a n d technical m e t h o d s involved in s e e d production.

Classes of Seed

Since the introduction of hybrids of s o r g h u m a n d pearl millet, there h a v e b e e n significant a d v a n c e s in


the t e c h n o l o g y of s e e d p r o d u c t i o n a n d p r o c e s s i n g . In s o r g h u m a n d pearl millet, s e e d - p r o d u c t i o n
p r o g r a m s are m o s t l y b a s e d o n hybrids.

It is n e c e s s a r y to h a v e a clear idea a b o u t the s e e d multiplication p r o c e s s before p l a n n i n g the


production of large quantities of certified s e e d . Certified s e e d is p r o d u c e d in s t a g e s f r o m t h e breeder
s e e d . T h e originating breeder supplies the breeder s e e d to different institutions s u c h as agricultural
universities, central a n d state research institutions, a n d other r e c o g n i z e d / s p o n s o r e d o r g a n i z a t i o n s .

B r e e d e r s e e d c a n also be p r o d u c e d by s p o n s o r e d br eeders. In special cases a n d in selected


c r o p s , p e r m i s s i o n m a y also be given to organizations s u c h as the National S e e d s C o r p o r a t i o n ( N S C ) ,
the State S e e d s D e v e l o p m e n t Corporat ions, a n d National o r State S e e d F a r m s w h i c h p o s s e s s big f a r m s ,
p r o c e s s i n g plants, st orage facilities, a n d qualified p e r s o n n e l .

T h e breeder s e e d required for national varieties in India is a r r a n g e d through the D e p a r t m e n t of


Agriculture a n d the National S e e d s Corporation of the Central G o v e r n m e n t . T h e breeder s e e d for state
varieties c a n be p r o d u c e d by b r e e d e r s of the states c o n c e r n e d . B r e e d e r - s e e d plots are m o n i t o r e d by a
t e a m consisting of b r e e d e r s , representatives of the state s e e d certification a g e n c y ( S S C A ) , a n d the s e e d -
p r o d u c i n g a g e n c y . A set of five p r o f o r m a are required to be filled by the b r eeder undertaking breeder
s e e d p r o d u c t i o n . T h e y are called 'Br e e d e r - S e e d Production Proforma' ( B S P ) .

• B S P - 1 : Allocation of br eeder s e e d production by the c r o p - i m p r o v e m e n t project;

• B S P - 2 : T i m e t a b l e of production a n d availability of breeder s e e d ;

• B S P - 3 : Inspection report of the monitoring t e a m ;

• B S P - 4 : B r e e d e r s e e d final production statement; a n d

• B S P - 5 : B r e e d e r s e e d history sheet.

After receiving the five p r o f o r m a , the S e e d s Division of the D e p a r t m e n t of Agriculture of the


Central G o v e r n m e n t estimates the total production of breeder s e e d for e a c h c r o p a n d variety. E a c h state
is required to submit its indent stating the requirement of br eeder s e e d in that state, 18 months in
a d v a n c e . B a s e d o n t h e s e indents a n d the final production figures, the S e e d s Division allocates t h e
quantity of b r e e d e r s e e d to be s u p p l i e d to e a c h state, a n d indicates the place of availability a n d the dates
by w h i c h to take delivery. T h e b r e e d e r s e e d c o m e s with a golden yellow t a g . T h e state g o v e r n m e n t s in
turn reallocate the b r e e d e r s e e d to the different indenters in the state (private a n d public sector

94
organizations).

F o u n d a t i o n s e e d is p r o d u c e d under the supervision of the S S C A on the f a r m s of agricultural


universities, state g o v e r n m e n t s , corporations, a n d in farmers' fields. T h e S S C A e x a m i n e s the source of
the breeder s e e d , a n d inspects the s e e d plots three or four times during the crop s e a s o n . After the crop
is h a r v e s t e d , t h e n o n p r o c e s s e d s e e d is s ealed by the S S C A and sent to the processing plants. From the
p r o c e s s e d s e e d , t h e S S C A d r a w s s a m p l e s a n d s ends t h e m t o the notified seed-testing laboratories
(STLs) for testing of physical purity, germination, etc. T h e samples are also subjected to a grow-out test,
to d e t e r m i n e genetic purity. S e e d of varieties a n d parents of hybrids are put through this test. T h e s e e d -
lots w h i c h m e e t the certification standards are given the foundation-seed tags (white tags).

T h e p r o c e d u r e for production of certified s e e d is almost the s a m e as for f o u n d a t i o n s e e d .


Howev er, t h e g r o w - o u t test for genetic purity is c o n d u c t e d only in select crops like hybrid c o t t o n , hybrid
castor, a n d doubtful lots of other crops. T h e lots that meet the m i n i m u m s e e d certification st andards are
p a s s e d by the S S C A s a n d certified-seed tags (blue tags) are issued.

A m o d e l of the seed-multiplication chain is given in Figure 1. It is essential that every individual


involved in s e e d production is familiar with the seed-multiplication chain b a s e d on the multiplication ratios
(Table 1) to be able to plan for the production of certified s e e d . Defective planning m a y result in deficit
or surplus production, both of which are harmful to the seed industry.

T a b l e 1 . S e e d multiplication ratio ( M R ) o f I C R I S A T 9s m a n d a t e c r o p s .

Crop S e e d rate (kg ha -1 ) Multiplication ratio

Sorghum 12 1:100
Pearl millet 4 1:200
Pigeonpea 20 1:100
Chickpea 75 1:10
Groundnut 100 1:8
Finger millet 5 1:80

T h e s y s t e m of variety/hybrid d e v e l o p m e n t a n d release in public a n d private sector organizations


is different. D e v e l o p m e n t proc edures are as per breeding principles, w h e r e a s the release s y s t e m is
linearly a d o p t e d by public organizations only.

Field and Laboratory Tests

T h e seed-certification s c h e m e , controlled pedigree s y s t e m s , a n d rules a n d regulations for s e e d gr owing


a n d distribution a re all a i m e d at maintaining cultivar trueness a n d purity of s e e d . In spite of this,
possibilities a l w a y s exist for c o n t a m i n a t i o n of the original seed-lots by u n w a n t e d s e e d s of other cultivars
or other t y p e s .

Causes of Contamination

C o n t a m i n a t i o n c a n occur due to several factors.

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Category Producing agency

Nucleus s e e d Agricultural universities and

(Basic seed) Research institutes

Breeder s e e d (Stage-I)

-do-
Breeder s e e d (Stage-ll)

Foundation s e e d (Stage-I)

Agricultural universities and

Foundation s e e d (Stage-ll) seed corporations (public/private)

Certified seed
seed corporations (public/private)
(First generation)

Commercial group
Farmer

Figure 1. Seed-multiplication chain

T h e total number of generations after breeder seed and before the commercial
crop should not e xceed three. Production of Breeder seed (stage-ll) should be
taken up only in the case of acute shortage.

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• Natural crossing with another cultivar, especially in open-pollinated crops

• Mutation

• U n c l e a n harvesting e q u i p m e n t

• C a r e l e s s n e s s at processing plant

• M i s t a k e s in b a g g i n g a n d tagging

To detect a n d control s u c h contamination, pre- a n d postharvest control tests are c o n d u c t e d by


m e a n s of cultivar identification/purity determination. Intensive crop-improvement p r o g r a m s h a v e resulted
in t h e d e v e l o p m e n t of a large n u m b e r of varieties in all important crop species, including s o r g h u m a n d
pearl millet. Variety identification has therefore attained critical importance in national a n d international
seed programs.

Different cultivars are c o m m o n l y identified on the basis of morphological differences in s e e d ,


s e e d l i n g , a n d m a t u r e plant. T h e Distinctness, Uniformity, a n d Stability (DUS) test or testing a cultivar for
these characters is an essential step for the registration of a variety.

Distinguishing varieties on the basis of s e e d morphology is not always possible. But it is


u n d o u b t e d l y o n e of the m o s t c o m m o n l y used criterion. Cultivar-identification tec hniques to a s s e s s purity
are essential for c o m m e r c i a l s e e d production a n d certification. Conventionally, s u c h identification has
b e e n a c c o m p l i s h e d t h r o u g h the grow-out test. In practice, this is an extremely successful p r o c e d u r e , but
it c a n be t i m e - c o n s u m i n g , requiring large areas of land. Also, m a n y of the mor phological descriptors u s e d
are multigenic a n d quantitative, displaying continuous variation, w h o s e expression is altered by
e n v i r o n m e n t a l factors. T h u s , there is a n e e d to develop alternative tests w h i c h can distinguish varieties
on the basis of stable biochemical properties of s e e d or seedlings.

An a u t h e n t i c s t a n d a r d s a m p l e must be available for comparison which is required to be treated


a n d e x a m i n e d in the s a m e w a y as the sample under test. In other w o r d s , seedlings a n d plants are
c o m p a r e d at the s a m e stage of development. A combination of laboratory a n d field m e t h o d s m a y be
u s e d to d e t e r m i n e the cultivar trueness a n d genetic purity of the s a m p l e . T h e laboratory m e t h o d s u s e d
for the e x a m i n a t i o n of m o r p h o l o g y , s e e d characters, color reaction to certain c h e m i c a l treatments,
properties of s e e d l i n g s , r e s p o n s e of seedlings to controlled environmental factors, a n d g r o w t h stimulants
are u s e d to d e t e r m i n e cultivar trueness. Of the various techniques available, analysis of s e e d a n d
seedling proteins a n d i s o e n z y m e s using electrophoresis techniques are the most widely u s e d b e c a u s e
of their reliability, rapidity, a n d cost-efficiency.

To ascertain the genetic purity of the certified/truthfully-labelled s e e d , d e v e l o p m e n t of quick


laboratory tests for key diagnostic characters has great value for the ev er - ex panding s e e d industry.
T h e s e d e v e l o p m e n t s will go a long w a y in maintaining high s e e d standards a n d will benefit both s e e d
producers and farmers.

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Methods of Genetic Purity Assessment

Examination of Seed in the Laboratory

T h i s test is a c c o m p l i s h e d by m e a n s of other distinctive variety (ODV) test in the laboratory. T h e test is


on t h e basis of the texture, color, a n d s h a p e of the s e e d in c o m p a r i s o n with an aut hentic s a m p l e . T h e
e x a m p l e s of s e e d color a n d s h a p e s in pearl millet are listed in T a b l e s 2a a n d 2 b .

T a b l e 2 a . S e e d color i n pearl millet.

Genotype

S e e d color Hybrids Hybrid parents

Brown HHB 50, HHB 60 H77/833-2, H90/4-5

Brownish gray H H B 5 1 , HHB 67, ICMP 4 5 1 ,


H H B 6 8 , H H B 69 D23

Gray - 842A,841A
and 841B

Light b r o w n ICMH 423 I C M P 4 5 1 , 843A,


843B, ICMP 423

T a b l e 2 b . S e e d s h a p e i n pearl millet.

Genotype

Seed shape Hybrids Hybrid parents

Pyriform H77/833-2, HHB 60 841 A, 8 4 1 B , H 90/4-5

Obovate HHB 60, HHB 68, 81 A, 81B, ICMP 4 5 1 ,


H H B 67 D23

Globular ICMH 4 5 1 , ICMH 423 842A, 842B, 843A,


843B, ICMP 432

Chemical Treatment of Seedlings

C h a r a c t e r s of s eedlings w h i c h m a y be p r o d u c e d in a relatively short time under laboratory conditions are


s o m e t i m e s u s e d in cultivar testing. It is important that seedlings are g r o w n under controlled conditions
a n d that a control s a m p l e is g r o w n simultaneously for c o m p a r i s o n .

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Phenol Reaction Test

To study t h e p h e n o l color reaction (Table 3) the seeds are s o a k e d in a solution of C u S o 4 at 1 a n d 2%


a n d i n a solution o f N a 2 C o 3 a t 0 . 1 % a n d 0 . 2 % prior t o test. T h e n the seeds are s o a k e d i n 1 % p h e n o l . T h e
reaction is r e c o r d e d after incubating the s a m p l e s at 30°C for 24 h. Reaction to p h e n o l with 0 . 2 % N a 2 Co 3
p r e s o a k i n g is better as c o m p a r e d to other treatments.

T a b l e 3. P h e n o l color reactions of various pearl millet g e n o t y p e s .

Genotype

S e e d color Hybrids Hybrid parents

Black (+++) H H B 6 1 , H H B 69 H 77/833-2

Dark b r o w n (+++) HHB 67, HHB 68 D 23, IMMP 451

B r o w n (++) HHB 50, IMCH 423 81 A, 8 1 B , H 90/4-5,


I C M P 4 2 3 , I C M P 451

No reaction (-) - 842A, 842B, 843A,


843B, 841A

+++=High, + + = M e d i u m , a n d +=Light reactions.

Shoot, Root, and Seedling Growth Response to Added Chemicals in Sorghum

To study the effect of D D T , B H C , Folidol, thiram, captan, 200 randomly selected s e e d s (25 s e e d s e a c h
in 8 replications) a r e treated with the powder in a m anner that a layer of the chemical is visible on the
s e e d coat. T h e s e e d s are then placed on two layers of moistened germination paper t o w e l s . T h e y are
c o v e r e d w i t h another m o i s t e n e d paper towel a n d rolled up. At the e n d of the s e v e n t h day, t h e rolled
towels are t a k e n out f r o m the germinator a n d the shoot, root, a n d seedling length are c o m p a r e d with the
control. A similar test c a n be d o n e by using gibberellic acid (GA 3 ) a n d polyethylene glycol ( P E G ) .

Electrophoresis

In c o m m o n u s a g e , electrophoresis refers to the m o v e m e n t of ions through a m e d i u m , w h e t h e r in free


solution, as in m o v i n g - b o u n d a r y electrophoresis or w h e n a supporting m e d i u m s u c h as starch gel,
p o l y a c r y l a m i d e g e l , cellulose, acetate or paper is e m p l o y e d .

E m p l o y i n g this principle, separation of proteins by electrophoresis is utilized for variety


identification.

Field Plot Test

T h e possibility of pr ov ing the g e n u i n e n e s s of a cultivar by the field plot test is b a s e d on the hereditary
characteristics of t h e plants. Usually, cultivar differences are mor e distinct if the g r o w t h conditions are

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f a v o r a b l e . T h e c r o p s h o u l d be so g r o w n that genetic differences e x p r e s s t h e m s e l v e s as clearly as
possible. It is essential to s o w the various s a m p l e s of the s a m p l e cultivar in s u c c e s s i o n a n d to s o w
s t a n d a r d s a m p l e s at suitable intervals, e.g., one s t a n d a r d s a m p l e for every 10 s a m p l e s to be t e s t e d . T h e
d e s i g n of a field plot is given in Table 4.

Table 4. Suggested design of a field plot to determine cultivar purity in sorghum and pearl millet.

Row Plant- Spacing Spacing


length to-plant between between
Crop (cm) spacing rows (cm) plots (cm) Replications

Sorghum 6 10 60 90 2
Pearl millet 6 10 45 100 2

For a s s e s s i n g genetic purity, a m i n i m u m of 4 0 0 plants a re re q u i r e d . T h e test is t e r m i n a t e d after


e x p r e s s i o n of t h e distinct character of a particular cultivar. T h e duration of the test d e p e n d s u p on the
crop/variety. T h e variety is identified on the basis of diagnostic characters.

Genetic Purity Requirements

T h e genetic purity requirements for various classes of s o r g h u m s e e d are given in Table 5.

T a b l e 5. G e n e t i c purity required for different c l a s s e s of s o r g h u m s e e d .

S e e d class Genetic purity (%)

Foundation seed 99.0

Certified s e e d
Varieties 98.0
Hybrids 95.0

Diagnostic Characters for Pearl Millet

Leaf blade Hairy/nonhairy

Leaf s h e a t h Hairy/nonhairy/pubescent/nonpubescent

Midrib color White/dull white

S t e m characters Nodal hair predominant/absent, violet or green nodal pigmentation

Spike characters Spike exertion negative/zero/positive

Spike s h a p e Conical/cylindrical/candle or oblanceolate/lanceolate

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Bristling Present/absent

Certification Procedures

T h e generally a c c e p t e d system of s e e d certification involves field inspections, s a m p l e testing, a n d


e n f o r c e m e n t of m i n i m u m standards, which constitute the m e c h a n i s m of s e e d quality control. T h e f a r m e r
is a s s u r e d of a rich harvest if he uses certified seeds. T h e following definition of the term "seed
certification" will be helpful in understanding the principles involved in the process.

S e e d certification is a scientific a n d systematically d e s i g n e d process to s e c u r e , m a i n t a i n , multiply,


a n d m a k e available s e e d s of superior crop plant varieties to f a r m e r s , to e n s u r e :

- t h e genetic identity of a variety with respect to its distinguishable, u n i f o r m , a n d stability


parameters;

- a high d e g r e e of physical purity;

- a high s t a n d a r d of s e e d health a n d germinability to pr oduce vigorous seedlings a n d g o o d plant


stand; and

- f r e e d o m f r o m all d e s i g n a t e d s eedborne diseases, w e e d s , a n d other crop s eeds ascertainable


under field a n d laboratory conditions.

Phases of Seed Certification

• Application for certification

• Establishing a s e e d s o u r c e

• Field inspection to verify conformity to field st andards

• Postharvest supervision

• G r a n t of certificate a n d tagging

M a i n t e n a n c e of purity/quality at every stage of s e e d production is essential. A n y flaw at a n y stage


results in failure to deliver good-quality s e e d s , resulting in w a s t a ge of e n e r gy a n d m o n e y .

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Development of Sorghum and Pearl Millet Seed Industry in India
G Harinarayana

T h e s e e d industry in India o w e s its d e v e l o p m e n t to several factors: T h e evolution of high-yielding


varieties a n d hybrids, t h e role of s e e d as a vital a n d critical c o m p o n e n t of p r o d u c t i o n , t h e ability of s e e d
to support a productive agricultural s y s t e m , the d e p e n d e n c e on s e e d for sustaining agricultural
productivity a n d p r oduction, a n d the ever-increasing d e m a n d for good-quality s e e d h a v e p r o m p t e d the
d e v e l o p m e n t of a full-fledged s e e d industry c o m p r i s i n g s e e d pr o d u c t i o n , p r o c e s s i n g , certification, storage,
and marketing.

Development of the Indian Seed Industry

T h e b e g i n n i n g of s o r g h u m a n d pearl millet s e e d production a n d distribution c a n be t r a c e d to the s e c o n d


d e c a d e of t h e 20th century. T h e history of the s o r g h u m a n d pearl millet s e e d industry is intertwined with
the birth a n d growth of the Indian s e e d industry in g e n e r a l . T h e r e f o r e , to recapitulate the history in
chronological order it is pr e s e n t e d here as P r e - l n d e p e n d e n c e (1900-1946), Early P o s t - I n d e p e n d e n c e
( 1 9 4 7 - 1 9 5 9 ) , a n d P o s t - I n d e p e n d e n c e (1960-1995) eras. T h e important milestones in the history of the
s o r g h u m a n d pearl millet s e e d industry are the establishment of the All-India C o o r d i n a t e d Project on
S o r g h u m I m p r o v e m e n t in 1960, the All-India C o o r d i n a t e d Pearl Millet I m p r o v e m e n t Project in 1965, the
e s t a b l i s h m e n t of the National S e e d s Corporation in 1963, a n d the establishment of a private s e e d
c o m p a n y , the M a h a r a s h t r a Hyb rid S e e d C o m p a n y ( M A H Y C O ) i n 1964.

P r e - l n d e p e n d e n c e Era (1900-1946)

1922 E s t a b l i s h m e n t of seed-multiplication laboratories by the D e p a r t m e n t of Agriculture of the United


Provinces (now divided into Uttar Pradesh a n d M a d h y a Pradesh). T h e s e laboratories u s e d to
multiply a n d supply s e e d to big landlords for further multiplication. Loans w e r e given for
establishing chains of s e e d stores in e a c h taluk (part of a district).

1925 T h e Royal C o m m i s s i o n on Agriculture e x a m i n e d the introduction, s p r e a d , a n d distribution of s e e d


of i m p r o v e d varieties in India. Following its s uggestions, several state g o v e r n m e n t s established
r e s e a r c h institutes.

1928 T h e D e p a r t m e n t of Agriculture started distribution of seeds of i m p r o v e d varieties. T h e s e seeds


w e r e either multiplied at o n e location a n d distributed over large areas or multiplied by a large
n u m b e r of f a r m e r s for their o w n use and/or distribution.

1944 T h e F a m i n e Enquiry C o m m i s s i o n a n d Foodgrains Policy C o m m i t t e e o b s e r v e d that crop botanists


s h o u l d e v o l v e n e w varieties, test t h e m , a n d hold on-farm d e m o n s t r a t i o n s . Initially, multiplication
of n e w varieties started on the farms of registered s e e d gr owers. S u c h s e e d s w e r e p r o c u r e d at
p r e m i u m prices a n d distributed at concessional rates.

1945 Private v e g e t a b l e s e e d c o m p a n i e s started p r o d u c i n g s e e d s of t e m p e r a t e v e g e t a b l e s in Quetta


( n o w in Pakistan) a n d Kashmir.

1946 T h e All-India S e e d G r o w e r s M e r c h a n t s a n d N u r s e r y m e n ' s Association w a s f o r m e d t o promote


d e v e l o p m e n t of t h e v e g e t a b l e s e e d industry.

102
Early Post-Independence Era (1947-1959)

1951 India's first Five-Year Plan (1951-1956) e m phasized the need to develop a s e e d production a n d
distribution p r o g r a m of i m p r o v e d seeds b a s e d on the food production forecast.

1952 T h e G r o w More F o o d Enquiry Committee noted that s e e d of required purity w a s not available.
An Expert S t a n d i n g C o m m i t t e e constituted by the Indian Council of Agricultural R e s e a r c h (ICAR)
f o r m u l a t e d a concrete s c h e m e for s e e d multiplication and distribution.

1956 During the S e c o n d Five-Year Plan period (1956-1961), a program for multiplication of nucleus
s e e d into f o u n d a t i o n s e e d w a s started at s e e d f a r m s , a n d s e e d stores w e r e e s t a b l i s h e d in e a c h
National Extension Service block. Plans for setting up seed-testing laboratories a n d c ooperative
stores w e r e d r a w n up.

1957 T h e All-India Coordinated Maize Improvement Project w a s established in collaboration with the
Rockefeller F o u n d a t i o n . This w a s a turning point in Indian agriculture, leading to the f o r m a t i o n of
a series of All-India Coordinated Crop Improvement Projects.

1959 T h e first I n d o - A m e r i c a n Agricultural Production T e a m r e c o m m e n d e d the est ablishment of a


private s e e d industry. T h e team s uggested that village-, block-, a n d district-level ext ension
w o r k e r s s h o u l d be m a d e primarily responsible for educating farmers on the use of i m p r o v e d s e e d .
T h e state d e p a r t m e n t s of agriculture are made responsible for s e e d certification, a n d private
g r o w e r s for s e e d supply (Agrawal 1980). T h e team further r e c o m m e n d e d setting up of state s e e d -
testing laboratories, a n d d e v e l o p m e n t of uniform s e e d certification standards a n d s e e d laws. A
c o m m i t t e e of technical experts developed plans for a s o u n d s e e d p r o g r a m for India. T h e y also
f o r m u l a t e d a m o d e l s e e d law which f o r m e d the basis of s u b s e q u e n t s e e d laws.

Post-Independence Era (1960-1995)

1960 T h e s e c o n d Indo-American Agricultural Production T e a m e n d o r s e d the first team's


r e c o m m e n d a t i o n s , a n d e m p h a s i z e d the need to accelerate their implementation.

T h e P r o g r a m Evaluation C o m m i t t e e felt that the norms for setting up 10-ha s e e d f a r m s w e r e not


strictly f o l l o w e d by the states. It also pointed out that seed farms w e r e being run by nonqualified
p e r s o n n e l , a n d that precautions necessary for maintenance of purity of s e e d s w e r e not t a k e n . It
also said that the s e e d s p r o d u c e d by registered growers w e r e not being u s e d .

1961 T h e Seed-Multiplication Review T e a m m a d e significant r e c o m m e n d a t i o n s r egarding production


a n d distribution of all important agricultural crops over a period of five years. T h i s included
varieties of millets, oilseeds, a n d pulses, and production of breeder s e e d under the supervision
of crop specialists. It r e c o m m e n d e d proper distribution of registered s e e d , a n d prescribed
s t a n d a r d s for purity, germination, field and s e e d inspections, harvesting, drying, storage, a n d s e e d
treatment.

T h e first four maize hybrids w e r e released by the All-India C o o r d i n a t e d M a i z e I m p r o v e m e n t


Project.

1962 T h e G o v e r n m e n t of India constituted a committee to formulate the S e e d s Act.

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1963 T h e N a t i o n a l S e e d s C o r p o r a t ion (NSC) (formerly the Central S e e d s Corporation) w a s established
to initiate f oundation a n d certified s e e d p r o g r a m s , e n c o u r a g e production a n d market ing of s e e d s ,
assist in the d e v e l o p m e n t of s e e d certification p r o g r a m s , s e e d law a n d s e e d - l a w e n f o r c e m e n t
p r o g r a m s , train personnel i n s e e d p r o g r a m s , a n d coordinate the i m p r o v e d s e e d p r o g r a m s . This
w a s the b e g i n n i n g of the d e v e l o p m e n t of a strong s e e d industry b a s e d on scientific principles.

1964 T h e first s e e d c o m p a n y in the private sector, the M a h a r a s h t r a Hybrid S e e d C o m p a n y Limited


( M A H Y C O ) w a s established. This c o m p a n y started a research p r o g r a m in 1966.

T h e C r o p I m p r o v e m e n t a n d Certified S e e d Producers Association w a s f o r m e d with active support


f r o m t h e Rockefeller Foundation a n d the G o v e r n m e n t of India.

S e e d - t e s t i n g laboratories w e r e started.

1965 T h e National S e e d s Cor por at ion o r g a n i z e d the first S e e d - i m p r o v e m e n t training c o u r s e .

1966 A H i g h - Y i e l d i n g Varieties P r o g r a m ( H Y V P ) w a s e n v i s a g e d in India, c o v e r i n g an a r e a of 9.2 m ha


in 1 9 6 8 - 6 9 , a n d 25 m ha in 1973-74 with rice, w h e a t , s o r g h u m , pearl millet, a n d m a i z e . This
e n c o u r a g e d strong linkages a m o n g N S C , the state departments of agriculture, a n d private
entrepreneurs.

T h e S e e d s Act w a s p a s s e d to regulate production a n d supply of quality s e e d . T h e law w a s later


a m e n d e d in 1 9 7 2 . This law h a d a provision for voluntary certification of s e e d s , but c o m p u l s o r y
truthful labelling. T h e m i n i m u m st andards for s e e d to be sold w e r e fixed. T h e S e e d Certification
Act c a m e into b e i n g officially.

1968 S e e d Rules w e r e f r a m e d . T h e y w e r e a m e n d e d i n 1973, 1974, a n d 1 9 8 1 .

T h e State R e v i e w T e a m m a d e 101 r e c o m m e n d a t i o n s relating to registration a n d release of


varieties, s e e d p r o d u c t i o n , p r o c e s s i n g , storage, quality control, a n d m a r k e t i n g . T h e
r e c o m m e n d a t i o n s c o v e r e d training, the role of var i o u s a g e n c i e s , f i n a n c e , a n d credit. T h e N S C
w a s to c o n t i n u e p r o d u c i n g f o u n d a t i o n s e e d till the agricultural universities took o v e r the function
under the coordination of a central agency. T h e state g o v e r n m e n t s w e r e to set up their o w n s e e d
certification a g e n c i e s . T h e cooperative a n d private sectors w e r e to be e n c o u r a g e d to replace
g o v e r n m e n t a l a g e n c i e s i n s e e d p r o d u c t i o n , p r o c e s s i n g , a n d m a r k e t i n g . P r o c u r e m e n t a n d sale
prices w e r e to be regulated by market forces.

1969 T h e T e r a i D e v e l o p m e n t Corporation (TDC) Limited w a s est ablished during the fourth Five-Year
Plan (1969-1974) with assistance from the W o r l d Bank. T h e primary objective of T D C w a s to
p r o d u c e one-third of the quality s e e d re q u i r e m e n t e n v i s a g e d b y t h e f o u r t h Five-Year P l a n , a n d
t h e r e b y to p r o m o t e the d e v e l o p m e n t of the Terai a r e a of Uttar P r a d e s h .

T h e F a r m s Corporation of India w a s establishe d with a view to accelerating foundation s e e d


programs.

1971 T h e Indian Society of S e e d T e c h n o l o g y w a s f o r m e d to provide its m e m b e r s a f o r u m to s hare their


s e e d p r o d u c t i o n e x p e r i e n c e . This society also b r o u g h t out t w o publications, S e e d R e s e a r c h a n d
S e e d T e c h n o l o g y N e w s . M i n i m u m s e e d certification standards w e r e f o r m u l a t e d .

104
T h e National C o m m i s s i o n on Agriculture in its interim s e e d review called for maintaining the
highest standards of s e e d purity.

1973 T h e first p o s t g r a d u a t e c o u r s e in s eed technology w a s started at the Indian Agricultural R e s e a r c h


Institute (IARI), N e w Delhi.

1975 T h e National S e e d Project (NSP) w a s initiated to reorganize a n d e x p a n d the s e e d industry. It


d e v e l o p e d a b r o a d - b a s e d network of s e e d production agencies in collaboration with the N S C as
principal coordinator for s e e d testing, s e e d certification, s e e d storage, a n d m a r k e t i n g . Public as
well as private agencies w e r e to be involved in breeder, f o u n d a t i o n , a n d certified s e e d p r o d u c t i o n .

1976 T h e National C o m m i s s i o n on Agriculture in its final report r e c o m m e n d e d that the s e e d industry


s h o u l d be c o m m e r c i a l i z e d . It also r e c o m m e n d e d that c o m p a c t productive a r e a s s h o u l d be
identified for s e e d production. It said seed processing must be m a d e compulsory a n d the grow-
out test s h o u l d be m a d e an integral part of the certification of parental mater ial. T h e c o m m i s s i o n
w a n t e d storage conditions for nucleus, breeder, foundation, a n d certified s e e d s i m p r o v e d . It
w a n t e d incentives to be provided for marketing. It felt that s eed-technology research a n d
e d u c a t i o n o u g h t to receive attention a n d that the S e e d s Act must be i m p l e m e n t e d rigorously.

S e e d corporations w e r e established in four states: A n d h r a P r a d e s h , H a r y a n a , M a h a r a s h t r a , a n d


Punjab under N S P Phase I.

1978 A buffer stock of s e e d s w a s initiated to meet the requirements of s e e d s in c a s e of natural


calamities a n d to bridge shortfalls in production by state s e e d corporations. S u b s e q u e n t l y , the
G o v e r n m e n t of India s u p p o r t e d buffer-stock s e e d production f r o m 1987.

D u r i n g N S P P h a s e II, the states of Bihar, Karnataka, Orissa, R a j a s t h a n , a n d Uttar P r a d e s h


established state seed corporations.

1979 T h e All-India C o o r d i n a t e d Project on S e e d Technology Research w a s initiated with 14 centers.

1980 T h e G o v e r n m e n t of Maharashtra decided to supply breeder s e e d to private s e e d c o m p a n i e s for


p r o d u c i n g their o w n f o u n d a t i o n s e e d for improving the overall quality of s e e d . This w a s
s u b s e q u e n t l y e n d o r s e d by the G o v e r n m e n t of India in 1983.

1982 T h e Indo-Danish Project on S e e d Technology Research a n d Training c a m e into o p e r a t i o n .

1983 T h e S e e d s Act w a s legislated to bring s e e d under the Essential C o m m o d i t i e s Act.

1985 D u r i n g the s e v e n t h Five-Year Plan period (1985-90), public a n d private sector c o m p a n i e s w e r e


s t r e n g t h e n e d for distributing certified/quality s eeds. In addition to agricultural universities a n d the
Indian Council of Agricultural Research (ICAR), N S C , and the State Farms Corporation of India
(SFCI) w e r e a s k e d to produce breeder s e e d . Private s e e d producers w e r e allowed to p r o d u c e
f o u n d a t i o n s e e d . Seed-testing laboratories, s e e d certification a g e n c i e s , s e e d - l a w e n f o r c e m e n t ,
s e e d - t e c h n o l o g i c a l research a n d training facilities were s t r e n g t h e n e d.

T h e S e e d Association of India (SAI) w a s established to provide a platform for s e e d s m e n to


e x c h a n g e v iews, ideas, a n d information.

105
1987 T h e Expert G r o u p o n S e e d s e x a m i n e d the w h o l e g a m u t o f the s e e d sector, s u c h a s p r o d u c t i o n ,
quality control, a n d m a r k e t i n g .

1988 A s e e d s e c t i o n w a s e s t a b l i s h e d at ICAR headquarters to deal with all a s p e c t s of s e e d .

T h e C e n t r a l S e e d T e s t i n g Laboratory w a s established at IARI, N e w Delhi, to d i s c h a r g e statutory


functions.

T h e M i n i m u m S e e d Certification S t andards w e r e modified a n d revised a s the Indian M i n i m u m


S e e d Certification S t a n d a r d s p u b l i s h e d b y the Central S e e d Certification B o a r d .

A n e w s e e d policy for import of i m p r o v e d varieties/hybrids of coarse cereals, pulses, oilseeds, a n d


f o d d e r w a s a n n o u n c e d . V e g e t a b l e a n d flower seeds w e r e p l a c e d u n d e r the O p e n G e n e r a l
Licence.

1989 A C e n t e r f o r E x c e l l e n c e in S e e d T e c h n o l o g y w a s established at the H a r y a n a Agricultural


University, Hisar, with the financial assistance of the United Nations D e v e l o p m e n t P r o g r a m
(UNDP).

T h e E x p e r t G r o u p on S e e d s s u g g e s t e d to the G o v e r n m e n t of India that in-service training be


p r o v i d e d to s e e d inspectors. T h e y m a y be m a d e exclusively responsible for s e e d quality control
a n d that targets b e fixed for s e e d testing.

1990 T h e National S e e d s Project III e n v i s a g e d strengthening of the All-India C o o r d i n a t e d R e s e a r c h


Projects. It w a n t e d breeder s e e d production units to be located in agricultural universities. It
r e c o m m e n d e d that g o v e r n m e n t a l agencies should be involved in quality control of s e e d s a n d that
t h e private sector s h o u l d be in the s e e d bus iness.

1994 T h e G o v e r n m e n t of India's W o r k i n g G r o u p on S e e d s e s t i m a t e d the s e e d r e q u i r e m e n t for the year


1 9 9 4 - 9 5 at 0.86 million tons c o m p a r e d to the 1.28 million tons that w o u l d be required by 2 0 0 0
A.D.

Sorghum and Pearl Millet Seed Industry

1960 T h e A c c e l e r a t e d S o r g h u m a n d Pearl Millet I m p r o v e m e n t Project w a s est ablished in collaboration


with t h e Rockefeller F o u n d a t i o n . A large n u m b e r of g e r m p l a s m collections w e r e a s s e m b l e d f r o m
India a n d a b r o a d , w h i c h later w e r e transferred to the International C r o p s R e s e a r c h Institute for
the S e m i - A r i d T r o p i c s ( I C R I S A T ) .

1963 T h e N S C s h o u l d e r e d t h e responsibility of fo u n d a t i o n a n d certified s e e d p r o d u c t i o n , certification,


a n d m a r k e t i n g . It d e v e l o p e d field a n d s e e d s t a n d a r d s .

1964 M A H Y C O b e g a n s o r g h u m a n d pearl millet s e e d pr oduction.

T h e first grain s o r g h u m hybrid, Coordinated S o r g h u m Hybrid 1 ( C S H 1) w a s released for


c o m m e r c i a l p r o d u c t i o n in India. This w a s f ollowed by the release of 13 m o r e hybrids ( C S H 2 to
C S H 14) until 1995.

106
1965 T h e first pearl millet grain hybrid, Hybrid Bajra 1 (HB 1), w a s released in India. This w a s f ollowed
by hybrids HB 2 to HB 7, BJ 104, CJ 104, I C M H 4 5 1 , MH 182, H H B 6 7 , Pusa 2 3 , a n d m a n y
others in the private sector in due course. HB 1 certified s e e d production w a s initiated in
K a r i m n a g a r a n d R a n g a R e d d y districts of Andhra Pradesh in 1965. Large-scale s e e d production
of HB 1 w a s u n d e r t a k en in N i z a m a b a d district a n d on a limited scale in W a r a n g a l district. In 1995,
N i z a m a b a d h a d nearly 12 0 0 0 ha under hybrid pearl millet, the highest in India f o l l o w e d by 6 0 0 0
ha in Gujarat ( P a n c h m a h a l a n d V a d o d a r a districts). T h e dry climate of N i z a m a b a d promotes
d e v e l o p m e n t of b o l d , lustrous a n d disease-free s e e d .

T h e A c c e l e r a t e d S o r g h u m and Pearl Millet Improvement Project w a s bifurcated into the All India
C o o r d i n a t e d S o r g h u m Improvement Project (AICSIP) and the All India C o o r d i n a t e d Millets
I m p r o v e m e n t Project (AICMIP).

T h e T e l a n g a n a Cooperative Hybrid Seeds Society w a s established to produce m aiz e, s o r g h u m ,


a n d pearl millet hybrids with the help of the United States A g e n c y for International D e v e l o p m e n t
(USAID).

1967 Production of C S H 1 s e e d w a s started in Kurnool, Andhra Pradesh. This district still p r o d u c e s


hybrid s e e d s of s o r g h u m on a large scale.

1972 T w o more districts, Nellore a n d Chittoor, started C S H 1 production.

1973 W e s t G o d a v a r i district in A n d h r a Pradesh started production of C S H 5 a n d C S H 6, but had to


discontinue it due to the problem of discoloration of s e e d .

1978 T h e production of C S H 5 a n d C S H 9 w a s started in Nandyal in Kurnool district, w h i c h p r o v e d to


be the most ideal niche for s o r g h u m hybrid s e e d production in India. It continues to be so e v e n
today.

1982 M A H Y C O - b r e d pearl millet hybrid M B H 110 w a s released by the Central Variety Release
Committee.

K h a m m a m district in A n d h r a Pradesh started pearl millet hybrid s e e d production.

1986 In G u n t u r district in A n d h r a P r a d e s h , production of hybrid pearl millet failed due to high incidence
of d o w n y m ildew. Similarly, s e e d production of HB 1 in 1967 a n d BJ 104 in 1 9 7 8 failed in
Nellore, d u e to d o w n y m ildew disease.

107
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Annexure I

List o f p a r t i c i p a n t s w h o a t t e n d e d t h e t r a i n i n g c o u r s e o n 9M a i n t e n a n c e o f male-sterile lines a n d


o p e n - p o l l i n a t e d v a r i e t i e s o f s o r g h u m a n d pearl millet a n d their u s e i n s e e d m u l t i p l i c a t i o n 9.

Dr C C N w a s i k e I A R / A B U , PO Box 1044, Zaria, Nigeria.

M s G a d a Issa Elias Agricultural R e s e a r c h Center, Horns, Syria.

Mr Gonda Jada C N R A , T a m a , M a r a d i , Niger.

Mr Karim Traore S R C V O / I E R , P.O. Box 2 5 9 , B a m a k o , Mali.

Mr M b u r u C M b o t h u K a k a m e g a Regional R e s e a r c h Center, P O Box 169, K a k a m e g a , K e n y a .

Mr M o h d R A H o v n y S h a n d w e e l Agricultural R e s e a r c h Station, S o h a g , Egypt.

Mr S N A Ipinge O m a h e n e n e R e s e a r c h Station, P O Box 144, O s h a k a t i , N a m i b i a .

Mr U T h a n A u n g Tropical R e s e a r c h F a r m , N y a u n g O o , M y a n m a r .

112