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Phenotyping For Plant Breeding - Applications of Phenotyping Methods For Crop Improvement-Springer-Verl
Phenotyping For Plant Breeding - Applications of Phenotyping Methods For Crop Improvement-Springer-Verl
Phenotyping For Plant Breeding - Applications of Phenotyping Methods For Crop Improvement-Springer-Verl
Edited by
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).
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
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.
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
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
Pearl Millet
References 108
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.
1
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
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.
Biotic Constraints
Diseases
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.
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.
Parasitic Weeds
3
Abiotic Constraints
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 .
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
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
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).
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.
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 .
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
6
Figure 1. Sorghum germination.
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.
Flowering
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).
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).
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.
Cuticle
Epicarp
Mesocarp
Cross cells
Tube cells
Pericarp
Stylar area
Scutellum
Shoot
Germ
Hila
Testae
Aleurone layer
Peripheral endosperm
Corneous endosperm
Floury endosperm
Physiological Maturity
11
S o r g h u m Plant a n d Flower Parts
Belum V S Reddy
• 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.
- 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 .
12
• 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.
• Pedicellate spikelets:
• S e e d or caryopsis:
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.
13
Figure 1. Germination and seedling growth in sorghum (1-7 days).
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.
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.
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 .
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.
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.
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 .
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.
Techniques of Hybridization
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.
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.
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.
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 .
19
P r o c e d u r e . T h e steps involved a r e :
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 .
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.
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.
20
a l o n g the panicle b r a n c h .
Fertilization
21
Development, Production, and Maintenance of Male-Sterile Lines
in Sorghum
Belum V S Reddy
22
Cytoplasmic-Genic Male Sterility
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.
23
maintenance.
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 .
24
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.
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.
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.
25
Improving the B-Lines and A-Lines
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.
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.
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.
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
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.
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.
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.
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.
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.
30
Production of Sorghum Hybrids
Belum V S Reddy
• 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.
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.
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.
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:
• 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 .
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.
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 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 .
• 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.
33
Pearl Millet
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
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.
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 .
Cultivar T y p e s
Open-Pollinated Cultivars
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
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.
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 .
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).
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.
36
Reproductive Biology of Pearl Millet
Faujdar Singh
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 .
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 .
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 )
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.
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).
Reproductive System
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 .
Panicle E m e r g e n c e
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 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.
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.
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).
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
44
Selling and Crossing Techniques in Pearl Millet
K N Rai
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.
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.
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 .
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 .
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
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.
N u m b e r of c r o s s e d panicles
17 iiii ii ii
210 i
285 iiii
362 iiii
C y t o p l a s m i c - G e n i c Male Sterility
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 .
49
s
1. Male-sterile plant/line
or
F F
2. Male-fertile plant/line
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.
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.
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 .
• 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
( ) % B-line 99.2
() 9 B-line 99.9
52
A-Iine Partial inbred B-Iine
Partial inbred B-line
F1
S P1
S P2
F
S P3
S P4
F
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
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)
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.
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.
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 :
• transplanting requires only about 3 0 - 4 0 % of the seed quantity required for direct s o w i n g ;
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.
55
A-line B-line
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
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.
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 B R
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
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
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
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.
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).
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.
61
RMB/F1
Self
S1/F2
Self
S 2 /F 3
Self
KVS
PMIHT,P(S4/F5) PRLN 1
S4/F5
Self, G H D M S N
D M S N , KVS
Self-multilocational
NT, Isolation plot evaluation
A I C P M I P trial SCA, U T
G C A / S C A , UT
ICRISAT
Pollinator Collection
(IPC)
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.
T e s t i n g of Fertility Restoration
• D o w n y m i l d e w resistance;
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
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.
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
64
Production of Pearl Millet Hybrid Seed
B S Talukdar
Choice of Location
Choice of Field
Cropping History
65
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-Line Border
Labelling
Roguing
66
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
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 .
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.
67
Seed Production of Pearl Millet Open-Pollinated Cultivars
C T Hash
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.
68
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:
• 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).
• 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
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.
70
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 .
Characters
71
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 .
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
72
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.
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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 .
• 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.
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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 .
Foundation Seed
Certified Seed
75
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
S e e d Multiplication by Farmers
• 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.
Reserve Stocks
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 .
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:
• 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 ;
• 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 ,
Conclusion
78
Seed-Processing Techniques and Chronology of
Indian Seed Industry
Organizational Setup
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 .
Seed Subsidy
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Import and Export of Seed
Planning
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:
Procurement of Seed
Buffer Stock
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 .
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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 .
Land Requirements
Field Inspections
Field standards
82
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 .
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
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 .
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Pearl Millet
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
Field Standards
M i n i m u m distance (m)
M a x i m u m permitted (%)
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
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
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.
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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.
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.
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Harvest and Postharvest Handling of Breeder Seed of Sorghum
and Pearl Millet
C T Hash
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 .
Drying
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• L o n g - t e r m storage of the s e e d ;
• Production of better-quality s e e d .
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 :
• 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
• 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;
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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.
• weed 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 .
• density,
• shape, and
• surface texture.
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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
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 .
Storage
Packaging
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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
• 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
Classes of Seed
• B S P - 5 : B r e e d e r s e e d history sheet.
94
organizations).
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 .
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
Causes of Contamination
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Category Producing agency
Breeder s e e d (Stage-I)
-do-
Breeder s e e d (Stage-ll)
Foundation s e e d (Stage-I)
Certified seed
seed corporations (public/private)
(First generation)
Commercial group
Farmer
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
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Methods of Genetic Purity Assessment
Genotype
Gray - 842A,841A
and 841B
T a b l e 2 b . S e e d s h a p e i n pearl millet.
Genotype
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Phenol Reaction Test
Genotype
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
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
99
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.
Sorghum 6 10 60 90 2
Pearl millet 6 10 45 100 2
Certified s e e d
Varieties 98.0
Hybrids 95.0
Leaf s h e a t h Hairy/nonhairy/pubescent/nonpubescent
100
Bristling Present/absent
Certification Procedures
• Establishing a s e e d s o u r c e
• Postharvest supervision
• G r a n t of certificate a n d tagging
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Development of Sorghum and Pearl Millet Seed Industry in India
G Harinarayana
P r e - l n d e p e n d e n c e Era (1900-1946)
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.
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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.
S e e d - t e s t i n g laboratories w e r e started.
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 .
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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.
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 .
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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).
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.
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
References
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the Semi-Arid Tropics.
B u r t o n , G.W., and A t h w a l , D.S. 1967. T w o additional sources of cytoplasmic m a l e sterility in pearl millet
a n d their relationship to Tift 23 A , . C r o p Science 7 : 2 0 9 - 2 1 1 .
C o o p e r , R.B., and B u r t o n , G.W. 1965. Effect of pollen storage on s e e d set in pearl millet, Pennisetum
typhoides. C r o p S c i e n c e 5:18-20.
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in pearl millet. C r o p S c ience 12: 198-200.
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Tropics.
R a i , K.N., and C.T. H a s h . 1990. Fertility restoration in male sterile x maintainer hybrids of pearl millet.
Crop Science 30:889-892.
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S i n g h , D., S i n g h , V., and C h a n d e i , S.R.S. 1970. Determination of leaf area in bajra (Penn/sefum
typhoides B u r m S. & H.). Science a n d Culture 36:46-47.
W i t c o m b e , J.R., R a o , M.N.V.R., and Lechner, W . R . 1995. Registration of 'ICMV 8 8 9 0 8 ' pearl millet.
C r op S c i e n c e - 3 5 : 1 2 1 6 - 1 2 1 7 .
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Annexure I
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 .
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