~~~~:~
Theor. Appl. Genet. 50, 185-191 (1977)
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: ;. . . .
R
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9 by Springer-Verlag 1977
An Analysis of Association of Components of Yield and Oil in Safflower (Carthamustinctorius L.) V. Ranga Rao and M. Ramachandram Soil Conservation Research Demonstration V. Arunachalam Division of Genetics, Indian Agricultural
& Training Research
Centre,
Institute,
Bellary New
(India
Delhi
(India)
Summary. Inter-relationships of various component characters with yield and oil content were analysed using 215 entries of safflower from India and U.S.A. Correlation of capsule number per plant and capsule weight with yield per plant was pronounced and they showed large direct effects on yield. All other components influenced seed yield mainly through these two components. Seed size had little effect on yield while seed number exerted a positive influence. The proportion of hull expressed as per cent of the whole seed revealed a highly significant and inverse relationship with oil content and was mainly responsible for the observed variability in oil content in the material. Although negative association was indicated between seed size and oil content, it was observed to be due to the indirect effect of hull content and not due to direct effect of seed size. Interestingly, yield per plant and its major components, number of capsules and capsule weight, revealed a negligible relationship with oil content. Both direct as well as indirect effects of hull percent and yield per plant were responsible for the favourable effect of seed number on oil content. The correlation of plant height, days to first flowering and total crop growth period with yield and oil content was either negligible or low, offering scope for developing early maturing and dwarf varieties with high yield and oil content. Spine index showed a non-significant association with yield and oil content. Capsule number, capsule weight and hull per cent were observed to be the most important components in breeding for higher yield and oil content.
Key w o r d s : Safflower -
Carthamus tinctorius-
Oil Content - Seed N u m b e r - P h e n o t y p i c C o r r e l a t i o n
Introduction
hampered
Although
safflower
(Knowles
1958,
seed crop,
is a crop
1969a)
particularly
under
ditions,
has been
moorthy
et al. 1966 ; Rege
De
et al.
be more crops
recognised
1974;
Ramachandram
of antiquity in India
its vast
limiting moisture only recently
Safflower
remunerative
like cotton,
19751
wheat
and Ramachandram grown
stands
and on borders
during
a main
crop
The success traditional presently depend
(Knowles
is reported
1975;
The crop
with cereals winter 1958,
of safflower areas
season 1967;
and oil content.
Direct
selection
to
in dif-
Ranga
Rao
is predominantly and
chick
rather Chavan
into new
by less efficient crops
on the extent of improvement
and
dryland
as a commercial
and its expansion
occupied
Rao
and chick pea grown
1977).
in mixed
1966 ;
Ranga
than the traditional
ferent parts of India (Anonymous
con-
(Krishna-
and Chattopadhyay
Anonymous 1977).
potential as an oil
made
inter-
apart from
their complex
inheritance.
on character
association
(Claassen al. 1968; aspects
Investigations et al. 1950;
Abel
to provide
new
it is essential
and direct forces ter association gramme.
Ashri
et al. 1957;
et al. 1974),
related to yield improvement
inadequate Further,
1969;
Argikar
An
that influence
to formulate experiment
obtain information are reported
directions
to understand
was
Urie et
and other
in this cropare in breeding. the indirect
a particular
charac-
a fruitful breeding hence
on these aspects
formulated
proto
and the results
in this paper.
pea
than as 1961). crop
due to large genotype-environmental
action and low heritability,
in
areas will largely in its yield
for yield and oil is
Materials
and Methods
The experimental material comprised of 210 entries, 189 of which were drawn from important areas of safflower cultivation in India representing a broad spectrum of available genetic variability for different characters, and21 from U.S.A. In addition, 5 standard Indian varieties were used as checks. The material was grown in an augmented randomisedblock design (Federer 1956) with 5 replications during the
186
T h e o r . A p p l . G e n e t . 50 ( 1 9 7 7 )
T a b l e 1. P h e n o t y p i c c o r r e l a t i o n 1 1 2 3 4 5 6 7 8 9 10 11 12 13 14
2 0.30
3
4
5
0.46 0.46 -0.27 0.29 0.25
0.37 0.33 0.18 0.70
coefficients among component characters 6
7
8
9
-0.06 -0.06 0.07 0.06
0.45 -0.26 0.58 0.32 0.18 0.05
0.30 -0.26 0.58 0.23 0.00 0.06 0.56
0.38 -0.29 0.64 0.28 0.26 0.07 0.78 0.95
10 0.07 0.18 0.06 0.04 0.28 0.16 -0.10 -0.02 -0.05
11 0.04 -0.17 0.19 0.07 0.07 -0.12 0.09 0.11 0.12 0.64
in safflower 12
0.07 0.45 -0.20 0.03 0.29 0.28 -0.22 -0.14 -0.19 0.32 -0.24
13
14
0.17 0.28 0.01 0.18 0.28 0.15 -0.04 0.01 -0.01 0.26 -0.16 0.51
-0.13 -0.30 0.08 -0.11 -0.21 -0.10 0.07 0.04 0.05 -0.12 0.29 -0.51 -0.83
15 0.34 -0.14 0.55 0.24 0.36 0.18 0.58 0.79 0.80 0.48 0.43 0.05 0.18 -0.03
1. P l a n t h e i g h t at m a t u r i t y ( c m ) 2. H e i g h t of b r a n c h i n g f r o m g r o u n d l e v e l ( c m ) 3. L e n g t h of b r a n c h e s ( c m ) 4. D a y s t o f i r s t f l o w e r i n g 5. D a y s to m a t u r i t y 6. S p i n e i n d e x 7. N u m b e r of s e c o n d a r y b r a n c h e s 8. N u m b e r of t e r t i a r y b r a n c h e s 9. N u m b e r of h e a d s p e r p l a n t 10. C a p s u l e w e i g h t ( g m ) 11. N u m b e r of s e e d s p e r h e a d 12. S e e d s i z e ( g m ) 13. H u l l p e r c e n t 14. Oil p e r c e n t 15. Y i e l d p e r p l a n t ( g m ) C o r r e l a t i o n c o e f f i c i e n t s at 1 Y0 a n d 5 7~ l e v e l of s i g n i f i c a n c e a r e O. 18 a n d 0 . 1 3 r e s p e c t i v e l y f o r n=215.
w i n t e r s e a s o n of 1 9 7 2 - 7 3 a t t h e Soil C o n s e r v a t i o n Research Demonstration & Training Centre, Bellary (Longitude 77~ Latitude 15~ i n p l o t s of 1 . 3 5 m • 4 . 5 m (3 r o w s ) w i t h a n i n t r a r o w s p a c i n g of 30 c m . The s o i l s a r e ' v e r t i s o l ' w i t h d e p t h v a r y i n g f r o m 6 0 90 c m . T h e y a r e a l k a l i n e i n r e a c t i o n a n d p o o r i n a v a i l able N and P. Fertilizers were band-placed by the s i d e of t h e s e e d r o w s a t t h e r a t e of 60 kg N a n d 60 kg P 2 O s / h a a s b a s a l d o s e . The c r o p w a s i r r i g a t e d w h e n a n d a s r e q u i r e d to s u p p l e m e n t s e a s o n a l r a i n f a l l a n d stored moisture. Observations were recorded on 5 p l a n t s s e l e c t e d a t r a n d o m f r o m t h e c e n t r a l r o w . The n u m b e r of d a y s to f i r s t f l o w e r i n g a n d c o m p l e t e m a turity were computed for individual plants as the peri o d f r o m t h e d a t e of p l a n t i n g t o t h e d a t e o n w h i c h blooming commenced in the primary head for the form e r , a n d f r o m t h e d a t e of p l a n t i n g to t h e d a t e of c o m p l e t e m a t u r i t y f o r t h e l a t t e r . H e i g h t of b r a n c h i n g w a s m e a s u r e d a s t h e h e i g h t of t h e p l a n t f r o m g r o u n d l e v e l to t h e p o i n t a t w h i c h b r a n c h i n g c o m m e n c e d o n t h e m a i n a x i s . O b s e r v a t i o n s o n t h e l e n g t h a n d a n g l e of branches were based on measurements taken from the longest two branches originating on the main stem. H e i g h t of p l a n t at m a t u r i t y w a s m e a s u r e d f r o m t h e g r o u n d l e v e l t o t h e a p e x of t h e c e n t r a l s t e m . N u m b e r of s e c o n d a r i e s i n c l u d e d p r i m a r y h e a d a n d f i r s t o r d e r branches on the central axis, while effective number of t e r t i a r i e s a n d h i g h e r o r d e r b r a n c h e s c o n s t i t u t e d n u m b e r of t e r t i a r y b r a n c h e s p e r p l a n t . S p i n e i n d e x f o r i n d i v i d u a l p l a n t s w a s c a l c u l a t e d a s a p r o d u c t of n u m b e r of s p i n e s a n d l e n g t h of t h e l o n g e s t s p i n e o n t h e o u t e r i n v o l u c r a l b r a c t s of t h r e e s e c o n d a r y h e a d s u s i n g t h e f o r m u l a s u g g e s t e d b y C l a a s s e n et a l . ( 1 9 5 0 ) . C a p s u l e w e i g h t w a s c a l c u l a t e d by d i v i d i n g t o t a l y i e l d p e r p l a n t by t h e t o t a l n u m b e r of e f f e c t i v e h e a d s p e r plant. Seed size (100-seed weight) for individual plants was obtained from 3 random determinations p e r p l a n t . Total s e e d n u m b e r p e r p l a n t w a s d i v i d e d by t h e n u m b e r of e f f e c t i v e h e a d s p e r p l a n t f o r o b t a i n i n g n u m b e r of s e e d s p e r c a p s u l e . To e s t i m a t e h u l l p e r c e n t , f u l l y a i r - d r i e d s e e d s of k n o w n w e i g h t were germinated for 24-30 hours in water. The hulls
including seed coat were separated from the kernels a n d w e i g h e d a f t e r d r y i n g i n a n o v e n at 6 0 ~ f o r 24 h o u r s . Oil c o n t e n t in t h e s e e d w a s e s t i m a t e d u s i n g a n N M R s p e c t r o m e t e r at t h e N u c l e a r R e s e a r c h L a b o r a t o r y , N e w D e l h i . Oil a n d h u l l e s t i m a t i o n s w e r e done on three samples per plant. Statistical analysis was based on the means of five plants for the various characters. A l l p o s s i b l e c o r r e l a t i o n s a m o n g 17 c h a r acters were calculated using the variance covariance matrix obtained after eliminating block effects. Since the correlations exhibited by the characters, capsule d i a m e t e r a n d a n g l e of b r a n c h e s w i t h a l l o t h e r s w e r e negligible, they were subsequently deleted from the correlation table. Path coefficient analysis (Dewey a n d Lu 1959) w a s u s e d t o p a r t i t i o n t h e c o m p l e x c o r r e l a t i o n s of c o m p o n e n t c h a r a c t e r s w i t h oil a n d y i e l d i n t o direct and indirect contributions.
Results
Phenotypic correlations:
N u m b e r of c a p s u l e s p e r
plant exhibited the highest correlation
coefficient
of 0 . 8 0 w i t h y i e l d ( T a b l e 1 ) . T e r t i a r y b r a n c h e s peared
to be more
closely
(r = 0.79) than secondary two v a r i a b l e s ,
correlated branches
ap-
with yield
( r = O. 5 8 ) . T h e s e
in fact, exhibited a highly significant
positive relationship
with capsule number.
r e l a t i o n of l e n g t h of b r a n c h e s ,
The cor-
capsule weight and
n u m b e r of s e e d s p e r c a p s u l e w i t h y i e l d w a s p o s i t i v e and significant.
T h e m a g n i t u d e of t h e a s s o c i a t i o n
l e n g t h of p e r i o d to f i r s t f l o w e r i n g , plant height at maturity,
of
crop growth period,
and per cent hull with yield
,
V. Ranga Rao, V. Arunachalam
and M. Ramachandaram:
Y i e l d a n d Oil i n S a f f l o w e r
T a b l e 2 . D i r e c t a n d i n d i r e c t e f f e c t s of c o m p o n e n t c h a r a c t e r s
1 2 3 4 5 6 9 10 11 12 13 I4
1
2
3
4
5
6
9
-0.02 -0.01 -0.01 -0.01 -0.01 0 -0.01 0 0 0 0 0
0.01 0.03 -0.01 0.01 0.0l 0 -0.01 0 -0.01 0.01 0.01 -0.0I
0 0 -0.01 0 0 0 -0.01 0 0 0 0 0
-0.01 -0.01 -0.01 -0.03 -0.02 0 -0.01 0 0 0 -0.01 0.01
0 0 0 -0.01 0 ~ 0 0 0 0 0 0
0 0 0 0 0 0.04 0 0 -0.01 0.01 0.01 0
0.32 -0.25 0.55 0.24 0.22 0.06 0.85 -0.04 0.10 -0.16 -0.01 0.04
187
influencing yield in safflower
10
0.03 0.08 0.03 0.02 0.12 0.07 -0.02 0.43 0.28 0.14 0.11 -0.05
(Carthamus tinctor4us L.)
11
12
13
0 0 -0-01 0.02 0-01 -0.01 0 0 0 0.02 -0.01 0.01 0.01 -0.01 0.04 0.02 0.07 -0.01 -0.02 0.06 -0.01 0.03 0.02 -0.03
14
0.02 0.04 0 0.03 0.04 0.02 0 0.04 -0.02 0.07 0.14 -0.12
-0.01 -0.03 0.01 -0.01 -0.02 -0.01 0.01 -0.01 0.03 -0.06 -0.09 0.11
r
0.34** -0.14" 0 . 5 5 ~* 0.24** 0.36** 0.18"* 0 . 8 0 ~* 0.48** 0.43** 0.05 0.18"* -0.03
r = Phenotypic correlation coefficient; R e s i d u a l f a c t o r -- 0 . 2 6 ; U n d e r l i n e d f i g u r e s d e n o t e d i r e c t e f f e c t s ; ~ * S i g n i f i c a n t a t 1 7~ l e v e l ; * S i g n i f i c a n t a t 5 g l e v e l
was,
in g e n e r a l ,
low a l b e i t s i g n i f i c a n t .
Surprisingly,
D a y s to f i r s t f l o w e r i n g w a s f o u n d to h a v e a p o s i -
s e e d y i e l d p e r p l a n t s h o w e d a w e a k but p o s i t i v e a s s o -
tive association
ciation with spine index (r = 0.18)
n u m b e r of c a p s u l e s ,
p o s s i b l y due t o i n -
c l u s i o n of a l a r g e n u m b e r of m o d e r a t e
to intense spiny
w i t h l o c a t i o n a n d l e n g t h of b r a n c h e s , hull per cent and plant height.
Early flowering types,
in g e n e r a l ,
forms in the m a t e r i a l . The association of all other
maturing
c h a r a c t e r s with yield was either too low o r negligible.
ters that were associated
A highly significant and negative association was
and vice versa
and consequently all characw i t h t h e l e n g t h of p e r i o d t o
flowering were also correlated
observed between oil and hull content. Oil content de-
t e n d e d to b e e a r l y
with maturity period.
V a r i a b i l i t y i n c a p s u l e n u m b e r w a s i n d e p e n d e n t of t h e
pended considerably on seed size ( r = -0.51) and was
variability in capsule weight and seed number.
inversely
association
turn,
related to the height of branching
was inversely
tertiary branches
associated
and number
While days to first flowering the oil content,
with secondary of capsules
period revealed
weather
effects of warm
and maturation
affecting the synthesis
oil i n t h e s e e d .
This emphasizes
the need to shorten
flowering period or suitable agronomic b o t h to a l l o w t h e c r o p t o m a t u r e
practices
or
before climatic and
negligible association
Hence,
among compo-
path coefficients were compresented
inTable 1
a n d ( b ) d e l e t i n g t h e n u m b e r of e f f e c t i v e s e c o n d a r i e s of
and tertiaries
from the correlation
were identical in both cases,
table. The results
with the exception of the which showed a negligible
c o n t r i b u t i o n t o y i e l d a n d oil i n ( a ) d u e to c o n f o u n d i n g of t h e i r e f f e c t s w i t h t o t a l n u m b e r of h e a d s p e r p l a n t . only the results
cussed in this paper
of path analysis
(b) are dis-
(Table 2).
per plant, showed a
with oil content.
a n d s p i n e i n d e x w e r e not a s s o c i a t e d Among other
nent characters.
Hence,
yield per plant and a principal yield
n u m b e r of c a p s u l e s
that would influence yield or oil
c o n t e n t d u e to t h e m u t u a l a s s o c i a t i o n
d i f f e r e n t o r d e r of b r a n c h e s
soil conditions become unfavourable.
component,
a n a l y s i s m a d e a b o v e c o u l d not p o i n t
puted (a) using all the characters
(January-
and accumulation
t h e t o t a l g r o w t h p e r i o d b y e i t h e r r e d u c t i o n of p r e -
Fortunately,
out t h e c h a r a c t e r s
period on oil con-
with seed number
per capsule. The c o r r e l a t i o n
prevailing in the region during the
period of seed development February)
Capsule weight revealed a
significant and positive correlation
a
The
of seed size with hull per cent was highly
significant and positive.
with it. Such an un-
effect of long crop growth
tent might be attributed to the adverse and dry
and
per plant.
did not greatly influence
total crop growth
significant negative association desirable
which in
Plant height
with oil content.
factors influencing oil content,
Path Analysis
number
of s e e d s p e r h e a d r e v e a l e d a h i g h l y s i g n i f i c a n t p o s i -
S e e d y i e l d p e r p l a n t : A s t u d y of d i r e c t e f f e c t s s h o w e d
tive relationship
t h e n u m b e r of c a p s u l e s
with oil per cent (r = 0.29).
p e r p l a n t f o l l o w e d by c a p s u l e
188
Theor. Appl. Genet. 50 (1977)
weight
were
the only characters
vely low direct effect. Favourable
directly influencing
yield. The indirect effects of these two component characters
through
all other associated
either negligible or very low.
Days
variables
showed
a positive
with yield only as a consequence
high and positive indirect effects through
capsule
gible. The indirect positive contributions number
combined
sitive direct effect was
mainly
responsible
served capsule
positive correlation and yield.
both through effect was
Per
between
through
the contribution
weight
hull and oil. The combined oil content through outweighed
to
between
indirect effects of
hull per cent and capsule
in a negligible
negative
it was number
a major
variabels
An analysis
owing
found that the indirect effects of capsules
influence. were
per plant and
The direct effects of
small.
phenotypic
of phenotypic
correlation
of component
characters
with yield and oil content and the paths of
association
clearly brought
ence of capsule
number,
out the important
capsule
weight
influ-
on yield, and
hull per cent along with these components
on oil con-
there-
tent. The influence
on yield
asso-
and oil content
weight
its positive and direct contribution,
by resulting
the above
total crop
Discussion
seedyield
relationship
negative
partitioned,
of hull per cent, yield exerted
to flowering,
plant height and height of branching
per
and oil per cent,
of a high inverse
were
period,
the association
The indirect
of the latter being negative
the occurrence
yield and hull per cent. When
for the obnumber
cent hull influenced
capsule
cap-
with the po-
seed
direct and indirect paths.
num-
influence of seed to be the result of a
positive direct effect and positive indirect effects through
growth
being neglithrough
and capsule
on oil content appeared
of oil content with the duration
of the
ber per plant, their own direct contributions
sule weight
were
to first flowering,
plant height and length of branches association
number
of all other characters
were
found to be either negligible or
manifestation
of the indirect effects of these
ciation of oil content with seed yield. All other char-
a mere
acters
principal components on t h e i r association with yield
that exhibited a significant phenotypic
tion with yield influenced rect effects exerted capsule
weight
yields mainly
through
associa-
by their indi-
either capsule
number
or
and oil. But mutual compensation at the component level appeared to be an important factor influencing c h a r a c t e r associations in safflower. This casts doubts
or both.
on the m e r i t s of these component c h a r a c t e r s in breedOil Content:
Oil content was
found to be influenced
by the direct effects of three components, cent (-0.80), capsules
yield per plant (0.40)
per plant (-0.37).
of hull per cent was phenotypic
and number
equal in magnitude
coefficient
were
plant which (r = -0.03
negligible.
was
however,
indirect effects of number and capsule the principal for the very
weight.
Such
components weak
with oil content
by the negative
of capsules, a mutual
hull per cent
cancellation
of oil was
among
also responsible
and non-significant
phenotypic
number
and weight
tent. The significant
inverse
association
corded
size and oil content
seed
positive
direct effect of yield
masked
sociation of capsule
between
yield per
had a substantial
direct effect. Such a favourable on oil content was,
ponents in determining yield and oil content in the
to its
on the above
Interestingly,
slightly associated
not significant)
of relevance to discuss the role of the principal com-
light of published work.
with oil content.
The indirect effects of other characters association
higher oil content in the seed. It would therefore be
of
The direct contribution
nearly
correlation
ing for superior production levels combined with
hull per
as-
with oil conthat was
re-
(r = -0.51)
Capsule Number per plant: This component c h a r a c t e r was positively c o r r e l a t e d with yield and also posseed a marked direct effect. Other components did not influence indirectly the association of capsule number with yield. The c h a r a c t e r also exercised pronounced indirect influence on the association of a host of e t h e r component c h a r a c t e r s with yield. This would mean that it would be possible to breed for a simultaneous improvement in various components of yield by proper breeding methods and selection p r o c e s s e s . However, the success would depend on the stability of the association of this c h a r a c t e r with yield, in addition to the c h o i c e of p a r e n t s and the b r e e d i n g p r o -
negative
c e d u r e . It was e n c o u r a g i n g to o b s e r v e that n u m b e r
indirect effect of hull per cent in addition to a relati-
of c a p s u l e s p e r plant a c c o u n t e d f o r a m a j o r p o r t i o n
was
found to be a consequence
of the sizeable
V. Ranga Rao, V. Arunachalam
and M. Ramachandram
: Y i e l d a n d Oil in S a f f l o w e r
Table 3. Direct and indirect effects of component
1 2 3 4 5 6 9 10 11 12 13 15
characters
2
3
4
5
6
9
0 ~ 0 0 0 0 0 0 0 0 0 0
-0.01 -0.03 0.01 -0.01 -0.01 0 0.01 -0.01 0 -0.01 -0.01 0
0.02 -0.01 0.05 0.01 0.01 0 0.03 0 0.01 -0.01 0 0.03
0.01 0 0.01 0.02 0.01 0 0.01 0 0 0 0 0
0.01 0.01 0.01 0.01 0.02 0 0.01 0.01 0 0.01 0.01 0.01
0 0 0 0 0 0.04 0 0.01 0 0.01 0.01 0.01
-0.14 0.11 -0.24 -0.11 -0.10 -0.03 -0.37 0.02 -0.04 0.07 0 -0.30
-0.01 -0.03 -0.01 -0.01 -0.04 -0.02 0.01 -0.15 -0.09 -0.05 -0.04 -0.07
189
influencing oil content in safflower
I0
1
(Carthamus tinctoriu~ L.)
11
12
0 -0.01 0.02 0.01 0.01 -0.01 0.01 0.05 0.08 -0.02 -0.01 0.04
-0.01 -0.05 0.02 0 -0.03 -0.03 0.02 -0.04 0.03 -0.12 -0.06 -0.01
13
15
-0.14 -0.23 -0.01 -0.14 -0.22 -0.12 0 -0.21 0.13 -0.41 -0.80 -0.14
r
0.14 -0.06 0.22 0.10 0.14 0.07 0.32 0.20 0.17 0.02 0.07 0.40
-0.13 -0.30 0.08 -0.11 -0.21 -0.10 0.05 -0.12 0.29 -0.51 -0.83 -0.03
~v ~
~
~ ~,~ ~
r : Phenotypic correlation coefficient; Residual factor = 0.51; Underlindes figures denote direct effects; ~*~Signifieant a t 1% l e v e l ; ~ S i g n i f i c a n t at 5 % l e v e l
of t h e t o t a l v a r i a n c e
i n y i e l d i n I n d i a n ( 5 8 . 3 70), I r a -
nian (42.87~) and the whole germplasm cessions
e v a l u a t e d at U t a h , U . S . A .
1974). Argikar
et a l .
(1957),
However,
( A s h r i et a l .
(1974) reported head diameter
a weak positive association between and yield, while they were independent
of e a c h o t h e r i n o u r s t u d i e s .
a h i g h d e g r e e of p o s i -
between the two variables
the association
ac-
Leininger (1968) and
Abel (1969) had also reported tive association
(48.6~)
in question.
of c a p s u l e n u m b e r p e r
The a s s o c i a t i o n
of c a p s u l e w e i g h t w i t h oil c o n t e n t
was non-significant;
however,
it h a d a n e g a t i v e d i r e c t
e f f e c t . The p o s i t i v e i n d i r e c t e f f e c t t h r o u g h y i e l d ( = 0 . 2 0 ) was offset by the negative indirect effect through hull
p l a n t w i t h o i l c o n t e n t w a s not i n t e n s e ( T a b l e 3 ) . The
per cent ( = -0.21 ). Again we noticed compensating
path coefficient analysis has provided the underlying
associations
a t t h e c o m p o n e n t l e v e l w h i c h n e e d to b e
causes;
streamlined
in the desired
i t s d i r e c t e f f e c t w a s n e g a t i v e o n oil c o n t e n t
( = -0.37)
but it w a s c o m p e n s a t e d
f e c t of y i e l d (= 0 . 3 2 ) . w a s not a s s o c i a t e d Similarly,
Interestingly,
yield per plant and its principal compof a i l e d to s h o w a n y s i g n i f i c a n t
Portugal and in the world safflower col-
l e c t i o n s ( A s h r i et a l . 1 9 7 4 ) . T h e r e s u l t s selection pressure
suggest that
f o r c a p s u l e n u m b e r d o e s not l e a d
to a n e g a t i v e r e s p o n s e
in yield and oil.
Seed size and seed number were thetwo othercomponents that were reported to be both important
not associated association
w i t h it i n o u r s t u d y . The m a g n i t u d e of
of s e e d s i z e a n d n u m b e r
was very low in our material
germplasm
weight:
The positive and significant corre-
mainly through
due to its direct effect, the indirect effects other characters
being negligible.
sociation was thus as important number
of capsules
This as-
as the one between
and yield. The authors could not
find any published work
on the inter-relationship
of
of y i e l d , n a m e l y c a p -
s u l e n u m b e r a n d w e i g h t , d i d not e x h i b i t a n y a s s o c i a t i o n p e r s e. H e n c e ,
it is essential
to give proper
w e i g h t a g e to b o t h of t h e s e c o m p o n e n t s f e w e r but l a r g e r
et al.
the role
sould also be properly
The t w o p r i n c i p a l c o m p o n e n t s
Argikar
(1969) and Ashri
Both
positively corre-
taken into account in breeding programmes.
higher yields.
Abel
however,
lated with capsule weight (Table 1). Hence,
capsule weight with yield and oil content. However, et al. (1957),
were,
of s e e d s i z e a n d n u m b e r
lation of capsule weight with yield was found to be
with capsule and in the world
c o l l e c t i o n s of A s h r i e t a l . ( 1 9 7 4 ) .
these components Capsule
Though seed num-
ber had a positive association with yield, seed size was
number
f o r oil c o n t e n t .
(Uric
et a l . 1 9 6 8 ; L e i n i n g e r 1 9 6 8 ; A b e l 1 9 6 9 ) a n d n o n i m p o r t a n t ( A s h r i et a l . 1 9 7 4 ) f o r y i e l d s .
w i t h oil c o n t e n t i n t h e g e n e p o o l s f r o m
India, Egypt,
improvement
direction for achieving
yield per plant
w i t h oil c o n t e n t i n o u r m a t e r i a l .
nent, capsule number, association
by the indirect ef-
in breeding for
On t h e o t h e r h a n d , p l a n t t y p e s w i t h heads having more or heavier seed
190
Theor. Appl. Genet.
50 ( 1 9 7 7 )
w e r e t h o u g h t to b e m o r e s u i t a b l e t h a n l i n e s w i t h l a r g e
i n t h e I n d i a n l i n e s ( A s h r i et al 1 9 7 4 ) . The s t a b i l i t y of
n u m b e r of h e a d s f o r l i m i t i n g m o i s t u r e
association
c o n d i t i o n s of
arid and semi arid regions in safflower
(Ashri et al.
1974), castor and wheat (Swaminathan
1969). Further,
head number cannot be wholly reliably used as a crit e r i o n f o r s e l e c t i o n b e c a u s e o f i t s low h e r i t a b i l i t y a n d high genotype environmental
interactions.
Hence, these
of s e e d s i z e a n d n u m b e r w i t h h u l l p e r c e n t ,
y i e l d a n d oil c o n t e n t w o u l d n e e d f u r t h e r s t u d y t o d e termine
t h e r o l e of t h e s e t w o c o m p o n e n t s
in breeding
programmes. Nevertheless,
t h e e x i s t a n c e of c o n s i d e r a b l e
nega-
t i v e d i r e c t a n d i n d i r e c t e f f e c t s of h u l l p e r c e n t o n oil
a s p e c t s n e e d to b e e x a m i n e d i n d e t a i l b e f o r e p l a n n i n g
content would suggest breeding for reduced hull per
a strategy for a simultaneous
c e n t to i n c r e a s e
improvement
of y i e l d
lead to marked
a n d oil i n s a f f l o w e r . Hull per cent:
This character
was negatively corre-
l a t e d w i t h oil c o n t e n t ( r = - 0 . 8 3 ~ ) with several other studies
as was the case
(Claassen
et a l .
1950;
L e i n i n g e r a n d U r i e 1 9 6 4 ; U r i e et a l . 1 9 6 8 ) a n d p o s s e s s e d a s u b s t a n t i a l d i r e c t e f f e c t (= - 0 . 8 0 ) .
(Argikar
Seed
et al. 1957;
L e i n i n g e r 1 9 6 8 ; A s h r i et a l . 1 9 7 4 ) s h o w e d a n i n v e r s e association
of c o n s i d e r a b l e
in the material
m a g n i t u d e w i t h oil c o n t e n t
(see also Urie et al. 1968). Such a ne-
gative correlation
w a s a l s o o b s e r v e d by A s h r i e t a l .
(1974) in Egyptian (r = -0.32) germplasm
and Iranian (r = -0.33)
c o l l e c t i o n s but not i n t h e l i n e s f r o m I n d i a .
The n e g a t i v e a s s o c i a t i o n
The incorporation
of s e e d s i z e w i t h oil c o n t e n t
w a s f o u n d t o b e d u e m o r e to a n i n d i r e c t e f f e c t t h r o u g h h u l l p e r c e n t ( - 0 . 4 1 ) t h a n d u e to a d i r e c t e f f e c t ( - 0 . 1 2 ).
i n p r o t e i n c o n t e n t of of stripped,
reduced and
t h i n h u l l m u t a n t s o f f e r s a n e w s o u r c e of m a t e r i a l
for
i m p r o v i n g t h e oil c o n t e n t in t h e k e r n e l a n d p r o t e i n level in the meal.
D e s p i t e t h e low p o s i t i v e d i r e c t e f -
fect and weak association
s i z e w h i c h h a d l i t t l e o r no a s s o c i a t i o n w i t h y i e l d s i n o u r s t u d i e s a n d t h o s e of o t h e r s
the meal.
oil c o n t e n t . S u c h a s t e p c o u l d a l s o improvements
selection pressure
of h u l l p e r c e n t w i t h y i e l d ,
for low hull might impose restric-
t i o n s o n t h e e x t e n t to w h i c h i m p r o v e m e n t s could be made.
in y i e l d
T h i s c o u l d b e p a r t l y o f f s e t by t h e i n -
c l u s i o n of c a p s u l e w e i g h t a s o n e of t h e c o m p o n e n t s the breeding programmes.
However,
in
other sources
of v a r i a b i l i t y f o r oil c o n t e n t will h a v e t o b e e x p l o i t e d i n v i e w of t h e s e l i m i t a t i o n s a n d o t h e r d r a w b a c k s available hull mutants
(Ebert and Knowles 1966).
A c c o r d i n g to C l a a s s e n
et a l .
(1950),
of
the character,
hull per cent, can be very useful for initial screening w i t h a h i g h d e g r e e of a c c u r a c y
in a large population
f o r oil c o n t e n t a n d w o u l d h e n c e e l i m i n a t e t h e n e e d Claassen
e t a l . ( 1 9 5 0 ) c o u l d not o b t a i n a n y s i g n i f i -
cant association
between seed size and hull, while
seed size and oil content exhibited a weak phenotypic positive correlation
i n t h e F 2 p o p u l a t i o n of a c r o s s .
Obviously, the increase was associated
in seed size in our material
more with an increase
t i o n of h u l l t h a n w i t h a n i n c r e a s e
in the propor-
in kernel size.
This
apparently resulted in a high inverse association
bet-
ween seed size and oil content. chymatous pericarp
In f a c t , t h e s c l e r e n -
of t h e s e e d a n d s e e d c o a t , n e i t h e r
of w h i c h h a s a n y f o o d o r f e e d v a l u e ( K n o w l e s 1 9 6 9 b ) , f o r m e d 45 to 50 7~ of t h e w h o l e s e e d by w e i g h t a n d w a s mainly responsible
f o r t h e l o w oil c o n t e n t of t h e s e e d
a n d t h e p r o t e i n c o n t e n t of t h e m e a l i n a m a j o r i t y of Indian collections.
for large oil estimations laboratory
by c o m p l e x a n d e x p e n s i v e
procedures.
The o t h e r c o m p o n e n t s
of y i e l d , n a m e l y p l a n t h e i g h t ,
l e n g t h a n d h e i g h t of b r a n c h e s ,
days to flowering,
days
to m a t u r i t y a n d s p i n e i n d e x , h a d no p r o n o u n c e d i n f l u e n c e e i t h e r o n y i e l d o r oil c o n t e n t . The f i n d i n g s of Argikar
et a l .
Claassen
et a l .
( 1 9 5 7 ) , A s h r e t a l . ( 1 9 7 4 , 1975) a n d (1950) confirm these results.
The results would suggest that the major components in question can be altered in breeding programrues freely to bring about concomittant associations process
and desirable
among the component characters
in the
of g e n e t i c u p g r a d a t i o n f o r y i e l d a n d oil c o n -
tent.
The s i g n i f i c a n t a n d p o s i t i v e r e l a -
t i o n s h i p of s e e d n u m b e r w i t h o i l c o n t e n t ( r = 0 . 2 9 ) w a s d u e to i t s d i r e c t e f f e c t a n d t h e i n d i r e c t e f f e c t through hull per cent and yield. However,
the asso-
c i a t i o n of s e e d n u m b e r w i t h oil c o n t e n t w a s f o u n d to be significant and positive only in the Iranian and Egyptian lines,
while it was weak and non-significant
Literature A n o n y m o u s : P r o c . of C r o p M a n a g e m e n t S e m i n a r o n Dryland Agriculture, All India Co-ordinated Research Project for Dryland Agriculture, Hyderabad, (1975).
V. R a n g a R a o , V. A r u n a c h a l a m and M. R a m a c h a n d r a m : Y i e l d a n d O i l i n Safflower (Garthamus tinctorius L.)
Abel, G.H. : An analysis of yield components in safflower. Proc. of 3rd Safflower Res. Conf., Univ. of California, Davis. pp. 18-22 (1969) Argikar, G.P.; Morbad, I.R.; Thobbi, V.V.: The range of variation of some quantitative characters in Carthamus tinctorius L. Ind. O i l s e e d s J o u r . 1 ( 4 ) , 232-234 (1957) Ashri, A . ; Z i m m e r , D . E . I Urie, A . L . ; Cahaner, A, ; M a r a n i , A. : E v a l u a t i o n of the world c o l l e c tion of s a f f l o w e r Carthamus tinctorius L. IV. Yield and y i e l d c o m p o n e n t s and t h e i r r e l a t i o n s h i p s . C r o p . Sci. 14, 799-802 (1974) A s h r i , A . ; Z i m m e r , D . E . ; U r i e , A . L . ; Knowles, P. F . : E v a l u a t i o n of the G e r m p l a s m c o l l e c t i o n s of s a f f l o w e r Carthamus tinctorius L. VI. Length of p l a n t i n g to f l o w e r i n g p e r i o d and plant height in I s r a e l , Utah and Washington. Theor. Appl. Genet. 4_66, 359-364 (1975) Chavan, V.M. : Niger and s a f f l o w e r . Indian C e n t r a l O i l s e e d s C o m m i t t e e , H y d e r a b a d , India, pp. 57-150
(1961)
Claassen, C.E.; Ekdahl, W.G.; Severson, G.A.: The estimation of oil percentage in safflower seed and the association of oil percentage with hull and nitrogen percentages, seed size and degree of spininess of the plant. Agron. J. 42, 478-482 (1950) De, Rajat; Turkhede, B.B.; Gangasaran: Safflower a good Rabi crop for Dryland. Indian Fmg. 24(2) , 26 (1974) Dewey, D . R . ; Lu, K . H . : A c o r r e l a t i o n and path c o efficient a n a l y s i s of c o m p o n e n t s of c r e s t e d wheat g r a s s s e e d p r o d u c t i o n . A g r o n . J. 5_!1, 515-518 (1959) E b e r t , W . W . ; Knowles, P . F . : I n h e r i t a n c e of p e r i c a r p t y p e s , s t e r i l i t y and d w a r f n e s s in s e v e r a l safflower c r o s s e s . C r o p . Sci. 6, 579-582 (1966)
Received March 15, 1977 Communicated by B.R. Murty
191
Federer, W.T. : Augmented designs. Hawaiian Planters Record 5_~5, 191-207 (1956) Knowles, P.F. : Safflower. Adv. Agron. I_0_0,289323 (1958) Knowles, P.F. : Processing seeds for oil in towns and villages of Turkey, India and Egypt. Econ. Bot. 21, 156-162 (1967) Knowles, P.F. : Centres of plant diversity and conservation of crop germ plasm. Econ. Bot. 23, 324-329 (1969a) Knowles, P.F. : Modification of quantity and quality of safflower oil through plant breeding. J. Amer. Oil C h e m . Soc. 4_.66, 130-132 (1969b) K r i s h n a m o o r t h y , Ch. ; Chattopadhyay, S. ; Rao, S. B. P. : Safflower for b e t t e r u t i l i z a t i o n of shallow and m e d i u m black cotton s o i l s of B e l l a r y a r e a . Ann. A r i d Z o n e . 5, 44-48 (1966) L e i n i n g e r , L.N. 1968: In: C a s t o r , S e s a m e and s a f flower, ( W e i s s , E . A . ) pp. 529-744. London: Leonard Hill 1974 Leininger, L.N. ; Urie, A . L . : D e v e l o p m e n t of s a f flower s e e d from flowering to m a t u r i t y . C r o p Sci. 4, 83-87 (1964) Ranga R~o, V. ; Ramachandram, M. : Substitute safflower for rainfed cotton in the rabi black soils of Bellary region for higher and stable returns. IndianFmg. (in press) (1977) Rege, N.D. ; Chattopadhyay, S. : Safflower for shallow and salty lands. Indian Frog. XIV, p. 39
(1966) Swaminathan, M.S. : Role of mutation breeding in a changing agriculture. In: Induced mutations in plants, pp. 719-734. Vienna: I.A.E.A. 1969 Urie, A.L.; Leininger, L.N.; Zimmer, D.E.: Effects of degree and time of defoliation on yield and related attributes of safflower. Crop Sci. 8, 747-750
(1968)
V. Ranga Rao and M. Ramachandram Soil Conservation Research Demonstration & Training Centre Bellary (India) V. A runachalam Division of Genetics Indian Agricultural Research New Delhi-il00i2 (India)
Institute
Theor. Appl. Genet. 50, 185-191 (1977)
~"
: ;. . . .
R
~ ....
9 by Springer-Verlag 1977
An Analysis of Association of Components of Yield and Oil in Safflower (Carthamustinctorius L.) V. Ranga Rao and M. Ramachandram Soil Conservation Research Demonstration V. Arunachalam Division of Genetics, Indian Agricultural
& Training Research
Centre,
Institute,
Bellary New
(India
Delhi
(India)
Summary. Inter-relationships of various component characters with yield and oil content were analysed using 215 entries of safflower from India and U.S.A. Correlation of capsule number per plant and capsule weight with yield per plant was pronounced and they showed large direct effects on yield. All other components influenced seed yield mainly through these two components. Seed size had little effect on yield while seed number exerted a positive influence. The proportion of hull expressed as per cent of the whole seed revealed a highly significant and inverse relationship with oil content and was mainly responsible for the observed variability in oil content in the material. Although negative association was indicated between seed size and oil content, it was observed to be due to the indirect effect of hull content and not due to direct effect of seed size. Interestingly, yield per plant and its major components, number of capsules and capsule weight, revealed a negligible relationship with oil content. Both direct as well as indirect effects of hull percent and yield per plant were responsible for the favourable effect of seed number on oil content. The correlation of plant height, days to first flowering and total crop growth period with yield and oil content was either negligible or low, offering scope for developing early maturing and dwarf varieties with high yield and oil content. Spine index showed a non-significant association with yield and oil content. Capsule number, capsule weight and hull per cent were observed to be the most important components in breeding for higher yield and oil content.
Key w o r d s : Safflower -
Carthamus tinctorius-
Oil Content - Seed N u m b e r - P h e n o t y p i c C o r r e l a t i o n
Introduction
hampered
Although
safflower
(Knowles
1958,
seed crop,
is a crop
1969a)
particularly
under
ditions,
has been
moorthy
et al. 1966 ; Rege
De
et al.
be more crops
recognised
1974;
Ramachandram
of antiquity in India
its vast
limiting moisture only recently
Safflower
remunerative
like cotton,
19751
wheat
and Ramachandram grown
stands
and on borders
during
a main
crop
The success traditional presently depend
(Knowles
is reported
1975;
The crop
with cereals winter 1958,
of safflower areas
season 1967;
and oil content.
Direct
selection
to
in dif-
Ranga
Rao
is predominantly and
chick
rather Chavan
into new
by less efficient crops
on the extent of improvement
and
dryland
as a commercial
and its expansion
occupied
Rao
and chick pea grown
1977).
in mixed
1966 ;
Ranga
than the traditional
ferent parts of India (Anonymous
con-
(Krishna-
and Chattopadhyay
Anonymous 1977).
potential as an oil
made
inter-
apart from
their complex
inheritance.
on character
association
(Claassen al. 1968; aspects
Investigations et al. 1950;
Abel
to provide
new
it is essential
and direct forces ter association gramme.
Ashri
et al. 1957;
et al. 1974),
related to yield improvement
inadequate Further,
1969;
Argikar
An
that influence
to formulate experiment
obtain information are reported
directions
to understand
was
Urie et
and other
in this cropare in breeding. the indirect
a particular
charac-
a fruitful breeding hence
on these aspects
formulated
proto
and the results
in this paper.
pea
than as 1961). crop
due to large genotype-environmental
action and low heritability,
in
areas will largely in its yield
for yield and oil is
Materials
and Methods
The experimental material comprised of 210 entries, 189 of which were drawn from important areas of safflower cultivation in India representing a broad spectrum of available genetic variability for different characters, and21 from U.S.A. In addition, 5 standard Indian varieties were used as checks. The material was grown in an augmented randomisedblock design (Federer 1956) with 5 replications during the
186
T h e o r . A p p l . G e n e t . 50 ( 1 9 7 7 )
T a b l e 1. P h e n o t y p i c c o r r e l a t i o n 1 1 2 3 4 5 6 7 8 9 10 11 12 13 14
2 0.30
3
4
5
0.46 0.46 -0.27 0.29 0.25
0.37 0.33 0.18 0.70
coefficients among component characters 6
7
8
9
-0.06 -0.06 0.07 0.06
0.45 -0.26 0.58 0.32 0.18 0.05
0.30 -0.26 0.58 0.23 0.00 0.06 0.56
0.38 -0.29 0.64 0.28 0.26 0.07 0.78 0.95
10 0.07 0.18 0.06 0.04 0.28 0.16 -0.10 -0.02 -0.05
11 0.04 -0.17 0.19 0.07 0.07 -0.12 0.09 0.11 0.12 0.64
in safflower 12
0.07 0.45 -0.20 0.03 0.29 0.28 -0.22 -0.14 -0.19 0.32 -0.24
13
14
0.17 0.28 0.01 0.18 0.28 0.15 -0.04 0.01 -0.01 0.26 -0.16 0.51
-0.13 -0.30 0.08 -0.11 -0.21 -0.10 0.07 0.04 0.05 -0.12 0.29 -0.51 -0.83
15 0.34 -0.14 0.55 0.24 0.36 0.18 0.58 0.79 0.80 0.48 0.43 0.05 0.18 -0.03
1. P l a n t h e i g h t at m a t u r i t y ( c m ) 2. H e i g h t of b r a n c h i n g f r o m g r o u n d l e v e l ( c m ) 3. L e n g t h of b r a n c h e s ( c m ) 4. D a y s t o f i r s t f l o w e r i n g 5. D a y s to m a t u r i t y 6. S p i n e i n d e x 7. N u m b e r of s e c o n d a r y b r a n c h e s 8. N u m b e r of t e r t i a r y b r a n c h e s 9. N u m b e r of h e a d s p e r p l a n t 10. C a p s u l e w e i g h t ( g m ) 11. N u m b e r of s e e d s p e r h e a d 12. S e e d s i z e ( g m ) 13. H u l l p e r c e n t 14. Oil p e r c e n t 15. Y i e l d p e r p l a n t ( g m ) C o r r e l a t i o n c o e f f i c i e n t s at 1 Y0 a n d 5 7~ l e v e l of s i g n i f i c a n c e a r e O. 18 a n d 0 . 1 3 r e s p e c t i v e l y f o r n=215.
w i n t e r s e a s o n of 1 9 7 2 - 7 3 a t t h e Soil C o n s e r v a t i o n Research Demonstration & Training Centre, Bellary (Longitude 77~ Latitude 15~ i n p l o t s of 1 . 3 5 m • 4 . 5 m (3 r o w s ) w i t h a n i n t r a r o w s p a c i n g of 30 c m . The s o i l s a r e ' v e r t i s o l ' w i t h d e p t h v a r y i n g f r o m 6 0 90 c m . T h e y a r e a l k a l i n e i n r e a c t i o n a n d p o o r i n a v a i l able N and P. Fertilizers were band-placed by the s i d e of t h e s e e d r o w s a t t h e r a t e of 60 kg N a n d 60 kg P 2 O s / h a a s b a s a l d o s e . The c r o p w a s i r r i g a t e d w h e n a n d a s r e q u i r e d to s u p p l e m e n t s e a s o n a l r a i n f a l l a n d stored moisture. Observations were recorded on 5 p l a n t s s e l e c t e d a t r a n d o m f r o m t h e c e n t r a l r o w . The n u m b e r of d a y s to f i r s t f l o w e r i n g a n d c o m p l e t e m a turity were computed for individual plants as the peri o d f r o m t h e d a t e of p l a n t i n g t o t h e d a t e o n w h i c h blooming commenced in the primary head for the form e r , a n d f r o m t h e d a t e of p l a n t i n g to t h e d a t e of c o m p l e t e m a t u r i t y f o r t h e l a t t e r . H e i g h t of b r a n c h i n g w a s m e a s u r e d a s t h e h e i g h t of t h e p l a n t f r o m g r o u n d l e v e l to t h e p o i n t a t w h i c h b r a n c h i n g c o m m e n c e d o n t h e m a i n a x i s . O b s e r v a t i o n s o n t h e l e n g t h a n d a n g l e of branches were based on measurements taken from the longest two branches originating on the main stem. H e i g h t of p l a n t at m a t u r i t y w a s m e a s u r e d f r o m t h e g r o u n d l e v e l t o t h e a p e x of t h e c e n t r a l s t e m . N u m b e r of s e c o n d a r i e s i n c l u d e d p r i m a r y h e a d a n d f i r s t o r d e r branches on the central axis, while effective number of t e r t i a r i e s a n d h i g h e r o r d e r b r a n c h e s c o n s t i t u t e d n u m b e r of t e r t i a r y b r a n c h e s p e r p l a n t . S p i n e i n d e x f o r i n d i v i d u a l p l a n t s w a s c a l c u l a t e d a s a p r o d u c t of n u m b e r of s p i n e s a n d l e n g t h of t h e l o n g e s t s p i n e o n t h e o u t e r i n v o l u c r a l b r a c t s of t h r e e s e c o n d a r y h e a d s u s i n g t h e f o r m u l a s u g g e s t e d b y C l a a s s e n et a l . ( 1 9 5 0 ) . C a p s u l e w e i g h t w a s c a l c u l a t e d by d i v i d i n g t o t a l y i e l d p e r p l a n t by t h e t o t a l n u m b e r of e f f e c t i v e h e a d s p e r plant. Seed size (100-seed weight) for individual plants was obtained from 3 random determinations p e r p l a n t . Total s e e d n u m b e r p e r p l a n t w a s d i v i d e d by t h e n u m b e r of e f f e c t i v e h e a d s p e r p l a n t f o r o b t a i n i n g n u m b e r of s e e d s p e r c a p s u l e . To e s t i m a t e h u l l p e r c e n t , f u l l y a i r - d r i e d s e e d s of k n o w n w e i g h t were germinated for 24-30 hours in water. The hulls
including seed coat were separated from the kernels a n d w e i g h e d a f t e r d r y i n g i n a n o v e n at 6 0 ~ f o r 24 h o u r s . Oil c o n t e n t in t h e s e e d w a s e s t i m a t e d u s i n g a n N M R s p e c t r o m e t e r at t h e N u c l e a r R e s e a r c h L a b o r a t o r y , N e w D e l h i . Oil a n d h u l l e s t i m a t i o n s w e r e done on three samples per plant. Statistical analysis was based on the means of five plants for the various characters. A l l p o s s i b l e c o r r e l a t i o n s a m o n g 17 c h a r acters were calculated using the variance covariance matrix obtained after eliminating block effects. Since the correlations exhibited by the characters, capsule d i a m e t e r a n d a n g l e of b r a n c h e s w i t h a l l o t h e r s w e r e negligible, they were subsequently deleted from the correlation table. Path coefficient analysis (Dewey a n d Lu 1959) w a s u s e d t o p a r t i t i o n t h e c o m p l e x c o r r e l a t i o n s of c o m p o n e n t c h a r a c t e r s w i t h oil a n d y i e l d i n t o direct and indirect contributions.
Results
Phenotypic correlations:
N u m b e r of c a p s u l e s p e r
plant exhibited the highest correlation
coefficient
of 0 . 8 0 w i t h y i e l d ( T a b l e 1 ) . T e r t i a r y b r a n c h e s peared
to be more
closely
(r = 0.79) than secondary two v a r i a b l e s ,
correlated branches
ap-
with yield
( r = O. 5 8 ) . T h e s e
in fact, exhibited a highly significant
positive relationship
with capsule number.
r e l a t i o n of l e n g t h of b r a n c h e s ,
The cor-
capsule weight and
n u m b e r of s e e d s p e r c a p s u l e w i t h y i e l d w a s p o s i t i v e and significant.
T h e m a g n i t u d e of t h e a s s o c i a t i o n
l e n g t h of p e r i o d to f i r s t f l o w e r i n g , plant height at maturity,
of
crop growth period,
and per cent hull with yield
,
V. Ranga Rao, V. Arunachalam
and M. Ramachandaram:
Y i e l d a n d Oil i n S a f f l o w e r
T a b l e 2 . D i r e c t a n d i n d i r e c t e f f e c t s of c o m p o n e n t c h a r a c t e r s
1 2 3 4 5 6 9 10 11 12 13 I4
1
2
3
4
5
6
9
-0.02 -0.01 -0.01 -0.01 -0.01 0 -0.01 0 0 0 0 0
0.01 0.03 -0.01 0.01 0.0l 0 -0.01 0 -0.01 0.01 0.01 -0.0I
0 0 -0.01 0 0 0 -0.01 0 0 0 0 0
-0.01 -0.01 -0.01 -0.03 -0.02 0 -0.01 0 0 0 -0.01 0.01
0 0 0 -0.01 0 ~ 0 0 0 0 0 0
0 0 0 0 0 0.04 0 0 -0.01 0.01 0.01 0
0.32 -0.25 0.55 0.24 0.22 0.06 0.85 -0.04 0.10 -0.16 -0.01 0.04
187
influencing yield in safflower
10
0.03 0.08 0.03 0.02 0.12 0.07 -0.02 0.43 0.28 0.14 0.11 -0.05
(Carthamus tinctor4us L.)
11
12
13
0 0 -0-01 0.02 0-01 -0.01 0 0 0 0.02 -0.01 0.01 0.01 -0.01 0.04 0.02 0.07 -0.01 -0.02 0.06 -0.01 0.03 0.02 -0.03
14
0.02 0.04 0 0.03 0.04 0.02 0 0.04 -0.02 0.07 0.14 -0.12
-0.01 -0.03 0.01 -0.01 -0.02 -0.01 0.01 -0.01 0.03 -0.06 -0.09 0.11
r
0.34** -0.14" 0 . 5 5 ~* 0.24** 0.36** 0.18"* 0 . 8 0 ~* 0.48** 0.43** 0.05 0.18"* -0.03
r = Phenotypic correlation coefficient; R e s i d u a l f a c t o r -- 0 . 2 6 ; U n d e r l i n e d f i g u r e s d e n o t e d i r e c t e f f e c t s ; ~ * S i g n i f i c a n t a t 1 7~ l e v e l ; * S i g n i f i c a n t a t 5 g l e v e l
was,
in g e n e r a l ,
low a l b e i t s i g n i f i c a n t .
Surprisingly,
D a y s to f i r s t f l o w e r i n g w a s f o u n d to h a v e a p o s i -
s e e d y i e l d p e r p l a n t s h o w e d a w e a k but p o s i t i v e a s s o -
tive association
ciation with spine index (r = 0.18)
n u m b e r of c a p s u l e s ,
p o s s i b l y due t o i n -
c l u s i o n of a l a r g e n u m b e r of m o d e r a t e
to intense spiny
w i t h l o c a t i o n a n d l e n g t h of b r a n c h e s , hull per cent and plant height.
Early flowering types,
in g e n e r a l ,
forms in the m a t e r i a l . The association of all other
maturing
c h a r a c t e r s with yield was either too low o r negligible.
ters that were associated
A highly significant and negative association was
and vice versa
and consequently all characw i t h t h e l e n g t h of p e r i o d t o
flowering were also correlated
observed between oil and hull content. Oil content de-
t e n d e d to b e e a r l y
with maturity period.
V a r i a b i l i t y i n c a p s u l e n u m b e r w a s i n d e p e n d e n t of t h e
pended considerably on seed size ( r = -0.51) and was
variability in capsule weight and seed number.
inversely
association
turn,
related to the height of branching
was inversely
tertiary branches
associated
and number
While days to first flowering the oil content,
with secondary of capsules
period revealed
weather
effects of warm
and maturation
affecting the synthesis
oil i n t h e s e e d .
This emphasizes
the need to shorten
flowering period or suitable agronomic b o t h to a l l o w t h e c r o p t o m a t u r e
practices
or
before climatic and
negligible association
Hence,
among compo-
path coefficients were compresented
inTable 1
a n d ( b ) d e l e t i n g t h e n u m b e r of e f f e c t i v e s e c o n d a r i e s of
and tertiaries
from the correlation
were identical in both cases,
table. The results
with the exception of the which showed a negligible
c o n t r i b u t i o n t o y i e l d a n d oil i n ( a ) d u e to c o n f o u n d i n g of t h e i r e f f e c t s w i t h t o t a l n u m b e r of h e a d s p e r p l a n t . only the results
cussed in this paper
of path analysis
(b) are dis-
(Table 2).
per plant, showed a
with oil content.
a n d s p i n e i n d e x w e r e not a s s o c i a t e d Among other
nent characters.
Hence,
yield per plant and a principal yield
n u m b e r of c a p s u l e s
that would influence yield or oil
c o n t e n t d u e to t h e m u t u a l a s s o c i a t i o n
d i f f e r e n t o r d e r of b r a n c h e s
soil conditions become unfavourable.
component,
a n a l y s i s m a d e a b o v e c o u l d not p o i n t
puted (a) using all the characters
(January-
and accumulation
t h e t o t a l g r o w t h p e r i o d b y e i t h e r r e d u c t i o n of p r e -
Fortunately,
out t h e c h a r a c t e r s
period on oil con-
with seed number
per capsule. The c o r r e l a t i o n
prevailing in the region during the
period of seed development February)
Capsule weight revealed a
significant and positive correlation
a
The
of seed size with hull per cent was highly
significant and positive.
with it. Such an un-
effect of long crop growth
tent might be attributed to the adverse and dry
and
per plant.
did not greatly influence
total crop growth
significant negative association desirable
which in
Plant height
with oil content.
factors influencing oil content,
Path Analysis
number
of s e e d s p e r h e a d r e v e a l e d a h i g h l y s i g n i f i c a n t p o s i -
S e e d y i e l d p e r p l a n t : A s t u d y of d i r e c t e f f e c t s s h o w e d
tive relationship
t h e n u m b e r of c a p s u l e s
with oil per cent (r = 0.29).
p e r p l a n t f o l l o w e d by c a p s u l e
188
Theor. Appl. Genet. 50 (1977)
weight
were
the only characters
vely low direct effect. Favourable
directly influencing
yield. The indirect effects of these two component characters
through
all other associated
either negligible or very low.
Days
variables
showed
a positive
with yield only as a consequence
high and positive indirect effects through
capsule
gible. The indirect positive contributions number
combined
sitive direct effect was
mainly
responsible
served capsule
positive correlation and yield.
both through effect was
Per
between
through
the contribution
weight
hull and oil. The combined oil content through outweighed
to
between
indirect effects of
hull per cent and capsule
in a negligible
negative
it was number
a major
variabels
An analysis
owing
found that the indirect effects of capsules
influence. were
per plant and
The direct effects of
small.
phenotypic
of phenotypic
correlation
of component
characters
with yield and oil content and the paths of
association
clearly brought
ence of capsule
number,
out the important
capsule
weight
influ-
on yield, and
hull per cent along with these components
on oil con-
there-
tent. The influence
on yield
asso-
and oil content
weight
its positive and direct contribution,
by resulting
the above
total crop
Discussion
seedyield
relationship
negative
partitioned,
of hull per cent, yield exerted
to flowering,
plant height and height of branching
per
and oil per cent,
of a high inverse
were
period,
the association
The indirect
of the latter being negative
the occurrence
yield and hull per cent. When
for the obnumber
cent hull influenced
capsule
cap-
with the po-
seed
direct and indirect paths.
num-
influence of seed to be the result of a
positive direct effect and positive indirect effects through
growth
being neglithrough
and capsule
on oil content appeared
of oil content with the duration
of the
ber per plant, their own direct contributions
sule weight
were
to first flowering,
plant height and length of branches association
number
of all other characters
were
found to be either negligible or
manifestation
of the indirect effects of these
ciation of oil content with seed yield. All other char-
a mere
acters
principal components on t h e i r association with yield
that exhibited a significant phenotypic
tion with yield influenced rect effects exerted capsule
weight
yields mainly
through
associa-
by their indi-
either capsule
number
or
and oil. But mutual compensation at the component level appeared to be an important factor influencing c h a r a c t e r associations in safflower. This casts doubts
or both.
on the m e r i t s of these component c h a r a c t e r s in breedOil Content:
Oil content was
found to be influenced
by the direct effects of three components, cent (-0.80), capsules
yield per plant (0.40)
per plant (-0.37).
of hull per cent was phenotypic
and number
equal in magnitude
coefficient
were
plant which (r = -0.03
negligible.
was
however,
indirect effects of number and capsule the principal for the very
weight.
Such
components weak
with oil content
by the negative
of capsules, a mutual
hull per cent
cancellation
of oil was
among
also responsible
and non-significant
phenotypic
number
and weight
tent. The significant
inverse
association
corded
size and oil content
seed
positive
direct effect of yield
masked
sociation of capsule
between
yield per
had a substantial
direct effect. Such a favourable on oil content was,
ponents in determining yield and oil content in the
to its
on the above
Interestingly,
slightly associated
not significant)
of relevance to discuss the role of the principal com-
light of published work.
with oil content.
The indirect effects of other characters association
higher oil content in the seed. It would therefore be
of
The direct contribution
nearly
correlation
ing for superior production levels combined with
hull per
as-
with oil conthat was
re-
(r = -0.51)
Capsule Number per plant: This component c h a r a c t e r was positively c o r r e l a t e d with yield and also posseed a marked direct effect. Other components did not influence indirectly the association of capsule number with yield. The c h a r a c t e r also exercised pronounced indirect influence on the association of a host of e t h e r component c h a r a c t e r s with yield. This would mean that it would be possible to breed for a simultaneous improvement in various components of yield by proper breeding methods and selection p r o c e s s e s . However, the success would depend on the stability of the association of this c h a r a c t e r with yield, in addition to the c h o i c e of p a r e n t s and the b r e e d i n g p r o -
negative
c e d u r e . It was e n c o u r a g i n g to o b s e r v e that n u m b e r
indirect effect of hull per cent in addition to a relati-
of c a p s u l e s p e r plant a c c o u n t e d f o r a m a j o r p o r t i o n
was
found to be a consequence
of the sizeable
V. Ranga Rao, V. Arunachalam
and M. Ramachandram
: Y i e l d a n d Oil in S a f f l o w e r
Table 3. Direct and indirect effects of component
1 2 3 4 5 6 9 10 11 12 13 15
characters
2
3
4
5
6
9
0 ~ 0 0 0 0 0 0 0 0 0 0
-0.01 -0.03 0.01 -0.01 -0.01 0 0.01 -0.01 0 -0.01 -0.01 0
0.02 -0.01 0.05 0.01 0.01 0 0.03 0 0.01 -0.01 0 0.03
0.01 0 0.01 0.02 0.01 0 0.01 0 0 0 0 0
0.01 0.01 0.01 0.01 0.02 0 0.01 0.01 0 0.01 0.01 0.01
0 0 0 0 0 0.04 0 0.01 0 0.01 0.01 0.01
-0.14 0.11 -0.24 -0.11 -0.10 -0.03 -0.37 0.02 -0.04 0.07 0 -0.30
-0.01 -0.03 -0.01 -0.01 -0.04 -0.02 0.01 -0.15 -0.09 -0.05 -0.04 -0.07
189
influencing oil content in safflower
I0
1
(Carthamus tinctoriu~ L.)
11
12
0 -0.01 0.02 0.01 0.01 -0.01 0.01 0.05 0.08 -0.02 -0.01 0.04
-0.01 -0.05 0.02 0 -0.03 -0.03 0.02 -0.04 0.03 -0.12 -0.06 -0.01
13
15
-0.14 -0.23 -0.01 -0.14 -0.22 -0.12 0 -0.21 0.13 -0.41 -0.80 -0.14
r
0.14 -0.06 0.22 0.10 0.14 0.07 0.32 0.20 0.17 0.02 0.07 0.40
-0.13 -0.30 0.08 -0.11 -0.21 -0.10 0.05 -0.12 0.29 -0.51 -0.83 -0.03
~v ~
~
~ ~,~ ~
r : Phenotypic correlation coefficient; Residual factor = 0.51; Underlindes figures denote direct effects; ~*~Signifieant a t 1% l e v e l ; ~ S i g n i f i c a n t at 5 % l e v e l
of t h e t o t a l v a r i a n c e
i n y i e l d i n I n d i a n ( 5 8 . 3 70), I r a -
nian (42.87~) and the whole germplasm cessions
e v a l u a t e d at U t a h , U . S . A .
1974). Argikar
et a l .
(1957),
However,
( A s h r i et a l .
(1974) reported head diameter
a weak positive association between and yield, while they were independent
of e a c h o t h e r i n o u r s t u d i e s .
a h i g h d e g r e e of p o s i -
between the two variables
the association
ac-
Leininger (1968) and
Abel (1969) had also reported tive association
(48.6~)
in question.
of c a p s u l e n u m b e r p e r
The a s s o c i a t i o n
of c a p s u l e w e i g h t w i t h oil c o n t e n t
was non-significant;
however,
it h a d a n e g a t i v e d i r e c t
e f f e c t . The p o s i t i v e i n d i r e c t e f f e c t t h r o u g h y i e l d ( = 0 . 2 0 ) was offset by the negative indirect effect through hull
p l a n t w i t h o i l c o n t e n t w a s not i n t e n s e ( T a b l e 3 ) . The
per cent ( = -0.21 ). Again we noticed compensating
path coefficient analysis has provided the underlying
associations
a t t h e c o m p o n e n t l e v e l w h i c h n e e d to b e
causes;
streamlined
in the desired
i t s d i r e c t e f f e c t w a s n e g a t i v e o n oil c o n t e n t
( = -0.37)
but it w a s c o m p e n s a t e d
f e c t of y i e l d (= 0 . 3 2 ) . w a s not a s s o c i a t e d Similarly,
Interestingly,
yield per plant and its principal compof a i l e d to s h o w a n y s i g n i f i c a n t
Portugal and in the world safflower col-
l e c t i o n s ( A s h r i et a l . 1 9 7 4 ) . T h e r e s u l t s selection pressure
suggest that
f o r c a p s u l e n u m b e r d o e s not l e a d
to a n e g a t i v e r e s p o n s e
in yield and oil.
Seed size and seed number were thetwo othercomponents that were reported to be both important
not associated association
w i t h it i n o u r s t u d y . The m a g n i t u d e of
of s e e d s i z e a n d n u m b e r
was very low in our material
germplasm
weight:
The positive and significant corre-
mainly through
due to its direct effect, the indirect effects other characters
being negligible.
sociation was thus as important number
of capsules
This as-
as the one between
and yield. The authors could not
find any published work
on the inter-relationship
of
of y i e l d , n a m e l y c a p -
s u l e n u m b e r a n d w e i g h t , d i d not e x h i b i t a n y a s s o c i a t i o n p e r s e. H e n c e ,
it is essential
to give proper
w e i g h t a g e to b o t h of t h e s e c o m p o n e n t s f e w e r but l a r g e r
et al.
the role
sould also be properly
The t w o p r i n c i p a l c o m p o n e n t s
Argikar
(1969) and Ashri
Both
positively corre-
taken into account in breeding programmes.
higher yields.
Abel
however,
lated with capsule weight (Table 1). Hence,
capsule weight with yield and oil content. However, et al. (1957),
were,
of s e e d s i z e a n d n u m b e r
lation of capsule weight with yield was found to be
with capsule and in the world
c o l l e c t i o n s of A s h r i e t a l . ( 1 9 7 4 ) .
these components Capsule
Though seed num-
ber had a positive association with yield, seed size was
number
f o r oil c o n t e n t .
(Uric
et a l . 1 9 6 8 ; L e i n i n g e r 1 9 6 8 ; A b e l 1 9 6 9 ) a n d n o n i m p o r t a n t ( A s h r i et a l . 1 9 7 4 ) f o r y i e l d s .
w i t h oil c o n t e n t i n t h e g e n e p o o l s f r o m
India, Egypt,
improvement
direction for achieving
yield per plant
w i t h oil c o n t e n t i n o u r m a t e r i a l .
nent, capsule number, association
by the indirect ef-
in breeding for
On t h e o t h e r h a n d , p l a n t t y p e s w i t h heads having more or heavier seed
190
Theor. Appl. Genet.
50 ( 1 9 7 7 )
w e r e t h o u g h t to b e m o r e s u i t a b l e t h a n l i n e s w i t h l a r g e
i n t h e I n d i a n l i n e s ( A s h r i et al 1 9 7 4 ) . The s t a b i l i t y of
n u m b e r of h e a d s f o r l i m i t i n g m o i s t u r e
association
c o n d i t i o n s of
arid and semi arid regions in safflower
(Ashri et al.
1974), castor and wheat (Swaminathan
1969). Further,
head number cannot be wholly reliably used as a crit e r i o n f o r s e l e c t i o n b e c a u s e o f i t s low h e r i t a b i l i t y a n d high genotype environmental
interactions.
Hence, these
of s e e d s i z e a n d n u m b e r w i t h h u l l p e r c e n t ,
y i e l d a n d oil c o n t e n t w o u l d n e e d f u r t h e r s t u d y t o d e termine
t h e r o l e of t h e s e t w o c o m p o n e n t s
in breeding
programmes. Nevertheless,
t h e e x i s t a n c e of c o n s i d e r a b l e
nega-
t i v e d i r e c t a n d i n d i r e c t e f f e c t s of h u l l p e r c e n t o n oil
a s p e c t s n e e d to b e e x a m i n e d i n d e t a i l b e f o r e p l a n n i n g
content would suggest breeding for reduced hull per
a strategy for a simultaneous
c e n t to i n c r e a s e
improvement
of y i e l d
lead to marked
a n d oil i n s a f f l o w e r . Hull per cent:
This character
was negatively corre-
l a t e d w i t h oil c o n t e n t ( r = - 0 . 8 3 ~ ) with several other studies
as was the case
(Claassen
et a l .
1950;
L e i n i n g e r a n d U r i e 1 9 6 4 ; U r i e et a l . 1 9 6 8 ) a n d p o s s e s s e d a s u b s t a n t i a l d i r e c t e f f e c t (= - 0 . 8 0 ) .
(Argikar
Seed
et al. 1957;
L e i n i n g e r 1 9 6 8 ; A s h r i et a l . 1 9 7 4 ) s h o w e d a n i n v e r s e association
of c o n s i d e r a b l e
in the material
m a g n i t u d e w i t h oil c o n t e n t
(see also Urie et al. 1968). Such a ne-
gative correlation
w a s a l s o o b s e r v e d by A s h r i e t a l .
(1974) in Egyptian (r = -0.32) germplasm
and Iranian (r = -0.33)
c o l l e c t i o n s but not i n t h e l i n e s f r o m I n d i a .
The n e g a t i v e a s s o c i a t i o n
The incorporation
of s e e d s i z e w i t h oil c o n t e n t
w a s f o u n d t o b e d u e m o r e to a n i n d i r e c t e f f e c t t h r o u g h h u l l p e r c e n t ( - 0 . 4 1 ) t h a n d u e to a d i r e c t e f f e c t ( - 0 . 1 2 ).
i n p r o t e i n c o n t e n t of of stripped,
reduced and
t h i n h u l l m u t a n t s o f f e r s a n e w s o u r c e of m a t e r i a l
for
i m p r o v i n g t h e oil c o n t e n t in t h e k e r n e l a n d p r o t e i n level in the meal.
D e s p i t e t h e low p o s i t i v e d i r e c t e f -
fect and weak association
s i z e w h i c h h a d l i t t l e o r no a s s o c i a t i o n w i t h y i e l d s i n o u r s t u d i e s a n d t h o s e of o t h e r s
the meal.
oil c o n t e n t . S u c h a s t e p c o u l d a l s o improvements
selection pressure
of h u l l p e r c e n t w i t h y i e l d ,
for low hull might impose restric-
t i o n s o n t h e e x t e n t to w h i c h i m p r o v e m e n t s could be made.
in y i e l d
T h i s c o u l d b e p a r t l y o f f s e t by t h e i n -
c l u s i o n of c a p s u l e w e i g h t a s o n e of t h e c o m p o n e n t s the breeding programmes.
However,
in
other sources
of v a r i a b i l i t y f o r oil c o n t e n t will h a v e t o b e e x p l o i t e d i n v i e w of t h e s e l i m i t a t i o n s a n d o t h e r d r a w b a c k s available hull mutants
(Ebert and Knowles 1966).
A c c o r d i n g to C l a a s s e n
et a l .
(1950),
of
the character,
hull per cent, can be very useful for initial screening w i t h a h i g h d e g r e e of a c c u r a c y
in a large population
f o r oil c o n t e n t a n d w o u l d h e n c e e l i m i n a t e t h e n e e d Claassen
e t a l . ( 1 9 5 0 ) c o u l d not o b t a i n a n y s i g n i f i -
cant association
between seed size and hull, while
seed size and oil content exhibited a weak phenotypic positive correlation
i n t h e F 2 p o p u l a t i o n of a c r o s s .
Obviously, the increase was associated
in seed size in our material
more with an increase
t i o n of h u l l t h a n w i t h a n i n c r e a s e
in the propor-
in kernel size.
This
apparently resulted in a high inverse association
bet-
ween seed size and oil content. chymatous pericarp
In f a c t , t h e s c l e r e n -
of t h e s e e d a n d s e e d c o a t , n e i t h e r
of w h i c h h a s a n y f o o d o r f e e d v a l u e ( K n o w l e s 1 9 6 9 b ) , f o r m e d 45 to 50 7~ of t h e w h o l e s e e d by w e i g h t a n d w a s mainly responsible
f o r t h e l o w oil c o n t e n t of t h e s e e d
a n d t h e p r o t e i n c o n t e n t of t h e m e a l i n a m a j o r i t y of Indian collections.
for large oil estimations laboratory
by c o m p l e x a n d e x p e n s i v e
procedures.
The o t h e r c o m p o n e n t s
of y i e l d , n a m e l y p l a n t h e i g h t ,
l e n g t h a n d h e i g h t of b r a n c h e s ,
days to flowering,
days
to m a t u r i t y a n d s p i n e i n d e x , h a d no p r o n o u n c e d i n f l u e n c e e i t h e r o n y i e l d o r oil c o n t e n t . The f i n d i n g s of Argikar
et a l .
Claassen
et a l .
( 1 9 5 7 ) , A s h r e t a l . ( 1 9 7 4 , 1975) a n d (1950) confirm these results.
The results would suggest that the major components in question can be altered in breeding programrues freely to bring about concomittant associations process
and desirable
among the component characters
in the
of g e n e t i c u p g r a d a t i o n f o r y i e l d a n d oil c o n -
tent.
The s i g n i f i c a n t a n d p o s i t i v e r e l a -
t i o n s h i p of s e e d n u m b e r w i t h o i l c o n t e n t ( r = 0 . 2 9 ) w a s d u e to i t s d i r e c t e f f e c t a n d t h e i n d i r e c t e f f e c t through hull per cent and yield. However,
the asso-
c i a t i o n of s e e d n u m b e r w i t h oil c o n t e n t w a s f o u n d to be significant and positive only in the Iranian and Egyptian lines,
while it was weak and non-significant
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V. Ranga Rao and M. Ramachandram Soil Conservation Research Demonstration & Training Centre Bellary (India) V. A runachalam Division of Genetics Indian Agricultural Research New Delhi-il00i2 (India)
Institute
