T h e o r e t i c a l a n d A p p l i e d G e n e t i c s 48, 1 - 8 ( 1 9 7 6 ) 9 by Springer-Verlag 1976
A Quantitative Evaluation of Inter-varietal Hybrids of Brassica campestris L.* A.A.
Devarathinam,
V. Arunachalam
D i v i s i o n of G e n e t i c s ,
Summary.
andB.R.
Indian Agricultural
Murty
Research
Institute,
New Delhi (India)
A quantitative
e v a l u a t i o n of y i e l d c o m p o n e n t s w a s c a r r i e d out in 150 i n t e r - v a r i e t a l h y b r i d s of Brasbrown sarson (BS), yellow sarson (YS) and toria (TR). The r e s u l t s s h o w e d b o t h a d d i t i v e a n d n o n - a d d i t i v e g e n e a c t i o n f o r p l a n t h e i g h t , n u m b e r of p r i m a r y a n d s e c o n d a r y b r a n c h e s a n d n u m b e r of s i l i q u a e o n t h e m a i n a x i s in a l l t h e s i x c r o s s c o m b i n a t i o n s - B S - Y S , Y S - B S , B S - T R , T R - B S , Y S - T R a n d T R - Y S . T h e g e n e r a l c o m b i n i n g a b i l i t i e s of B S , YS a n d TR i n d i c a t e d t h a t t h e i r n a t u r e a n d m a g n i t u d e d e p e n d e d l a r g e l y o n t h e o t h e r p a r e n t s w h i c h e n t e r e d t h e h y b r i d s . T h e r e w e r e d i f f e r e n c e s in c o m b i n i n g a b i l i t y b e t w e e n d i r e c t a n d r e c i p r o c a l c o m b i n a t i o n s . G B S t I , K a n p u r Lotni 17, K a n p u r L o t n i 27 a n d DS 17D in B S , IB 3, IB 5, I B 6 a n d E P 12 in YS, a n d T 1 6 5 , T 244 a n d T 1842 in TR w e r e i d e n t i f i e d a s p o t e n t i a l p a r e n t s f o r i n t e r - v a r i e t a l h y b r i d i s a t i o n . R e c i p r o c a l e f f e c t s w e r e f o u n d w h e n BS o r YS w a s u s e d a s p a r e n t a n d t h e y w e r e l e a s t w h e n TR w a s u s e d a s a p a r e n t . T h e a m o u n t a n d d e g r e e of h e t e r o s i s w a s s u b s t a n t i a l in i n t e r - v a r i e t a l c r o s ses. Based on the heterosis-combining a b i l i t y r e l a t i o n s h i p , t h e r o l e of i n t e r - v a r i e t a l h y b r i d i s a t i o n in p o p u l a t i o n i m p r o v e m e n t of Brassica carapestrgs i s d i s c u s s e d . A n u m b e r of m e t h o d s of u t i l i s i n g t h e i n t e r - v a r i e t a l h y b r i d s i n m u l t i p l e c r o s s e s a n d s y n t h e t i c c o m p l e x e s i s s u g g e s t e d a s p o t e n t i a l s u p p l e m e n t s to p o p u l a t i o n b r e e d i n g in this crop.
sica campestris u s i n g f i v e e a c h of t h e t h r e e v a r i e t i e s ,
Introduction
w h i c h c o u l d p r o f i t a b l y b e u s e d in m a r k i n g
The o l e i f e r o u s
ful s h o r t t e r m
varieties,
Brassica campestris L. i n c l u d e s
brown sarson(BS),
yellow sarson(YS)
toria(TR),
with distinct genetic
differences
but i n t e r m a t i n g
1958).
W h i l e YS a n d T R
compatible
forms.
changes in flowering changes,
time,
have restricted
ation within each variety
isolation
de-
due to
down the vari-
( R a j a h 1 9 5 8 ) . D u r i n g at l e a s t
t h r e e d e c a d e s no s i g n i f i c a n t a c h i e v e m e n t s
have been
m a d e i n i m p r o v i n g t h e y i e l d of t h e s e p o p u l a t i o n s . tempts
BS
and also chromosomal
and narrowed
At-
w e r e c o n f i n e d m a i n l y to i m p r o v i n g t h e y i e l d of
the three varieties
separately
proved breeding procedures. procedures
by c o n v e n t i o n a l
and im-
T h o u g h t h e m e r i t s of t h e s e
c a n not b e u n d e r r a t e d ,
it w o u l d b e d e s i r -
a b l e to p l a n a s t u d y t o e x a m i n e t h e b a s i c p o t e n t i a l i t i e s of i n t e r - v a r i e t a l to f u r t h e r
hybridisation
reported
conducted in this
Material
and Methods
W e w o u l d l i k e to u s e t h e t e r m , v a r i e t y , w h e n r e f e r r i n g to BS, YS a n d T B a n d t h e t e r m , p o p u l a t i o n s , to r e f e r to d i f f e r e n t s u b - v a r i e t i e s of BS, YS a n d TR. F i f t e e n p o p u l a t i o n s , t h r e e s e t s of f i v e e a c h , i n t h e three varieties as detailed below, were utilised as p a r e n t s in t h i s s t u d y . B__.SS: 1 - G B S II ( I N T ) ; 2 - K a n p u r Lotni 17 ( S I ) ; 3 - K a n p u r Lotni 27 ( S I ) ; 4 - D S 1 7 D ( S C ) ; 5- A s s a m L o c a l ( S C ) YS: 6 - IB 3; 7 - IB 5; 8 - IB 6; 9- IB 7 1 ; 1 0 - E P 12 TR: 1 1 - T 165; 1 2 - T 2 1 7 ; 1 3 T 244; 14- T 267; 15- T 1842. The p a r e n t s 2 a n d 3 i n BS w e r e s e l f - i n c o m p a t i b l e (SI), 4 and 5 were self-compatible (SC) and 1intermediate-compatible (INT). InYS, the exotic parent 10 h a d b i v a l v e d , w h i l e t h e o t h e r f o u r h a d m u l t i v a l v e d s i l i q u a e . The TR p a r e n t s d i f f e r e d i n f l o w e r i n g d u r a tion, besides other characters. E a c h s e t of p a r e n t s served both as lines and as testers for the other and t h i s p r o c e s s r e s u l t e d i n 150 h y b r i d s , i n c l u d i n g 75 c r o s s e s a n d 75 r e c i p r o c a l s a s s h o w n b e l o w :
among these varieties,
boost population improvement
Tester
with regard
to s e e d a n d oil y i e l d . S u c h a s t u d y w o u l d a l s o p r o v i d e a w e a l t h of b a s i c i n f o r m a t i o n
crosses, are
of s u c h
paper.
and self-in-
of s u c h v a r i a b l e
temporal
out p o w e r -
The results
self-com-
groups respectively,
intermediate
an investigation on inter-varietal in t h i s c r o p f o r t h e f i r s t t i m e ,
and morphological
belong to strictly
The p r e s e n c e
g r e e s of i n c o m p a t i b i l i t y ,
and
with one another (Singh
patible and self-incompatible includes self-compatible,
three
breeding methods.
on the yield components
"" P a r t of a P h . D . t h e s i s s u b m i t t e d by s e n i o r a u t h o r to P . G . S c h o o l , I n d i a n A g r i c u l t u r a l R e s e a r c h I n s t i tute, New Delhi
Li n e BS YS TR
9 Crosses o Crosses
made not m a d e
BS
YS
TR
o
9
9 9
o
9 9
9
o
A. Table
Amirtha
I. ANOVA
Devarathinam
et al. : A Quantitative
for four yield components
in inter-varietal
Mean Source
d.f.
Parents
(P)
BS YS TR BS vs YS TR vs Rest (H)
149
BS • YS • a vs b BS • TR TR x BS e vs d YS xTR TR XYS e vs f (BS-YS) (YS-TR)
(a) (b)
24 24 1 24 24 1 24 24 1 1 1
(c) (d) (e) (f) vs (BS-TR) vs BS crosses
P vs H Error ~" N o t s i g n i f i c a n t ;
all other
9.5 6 . 8 -,'~ 3 . 0 ~''" 28.9 129.0
91.6 3.2
among
of variance the parents,
for X3,
those
(Table
A further
showed
was
The TR
nations
fered
from
s.s.
due to lines, study
The hybrids corresponding
s.s.
followed
did not show IBS vs YS mean
significant and YS
of squares
BS-YS
for X2
and X3,
for all characters was
to various
further
significant sum
of squares
while
except
partitioned
inter-varietal
TR
cross
difX3.
into those combi-
Each
one of the cross
subdivided
and line • tester
components
between
(Table
into that
interaction.
A
2 ) brings out the fol-
Significant
the 50 hybrids
differences
(25 crosses
•
BS
where
ter and line x tester
were
differences whereas
for X3
between
asline,
were
Inthe line, tes-
significant
inthereciprocals,
and line • tester (Table
BS • YS
interaction
2). The
and YS
for
YSXBS,
x BS
reciprocal werenot
sig-
for any character. combinations: were
characters
for X3
were
non-significant (TR)
differences
in the crosses
differences
(Table
no significant (BS)
except
BS
for X2
were
2). When
the 50
TR
as line
and X3,
was
differences
were
Line x tester
but
and for X4
significant
line differences
X3.
among
for all the characters,
not significant.
differences
but the tester characters
The
significant
the differences
were
served
not significant
differences
tester
BS
were and 25
crosses,
reciprocals var-
6807.6 86.1
for all the four yield components.
(s.s.) IBS
"" -,',~
reciprocals)
hybrids
popula-
~' ""
further
combinations:
observed
were
among
by YS
562.6 401.8 2 . 5 ~'~ 284.3 78.5 r 1382.3 150.8 157.7 2329.7 8738.5 1232.2
lowing :
BS-TR of the sum
was
testers
of these
nificant
vs Hybrids
that the variation
populations
BS
except
402.3
and their reciprocals.
combinations
for X2
differences
the hybrids,
Parents
the highest,
iation for any character. was
The
459.2 8 6 . 8 ~'~ 6 6 . 3 ~'17.2 ~ 373.8
at 5 % l e v e l
1 ).
partitioning
due to parents populations
(m.s.)
among
and the comparison
all significant
tions.
of plot means:
5 2 . 5 -~ 145.1 9 . 4 "" 972.2 1 . 3 ~,~
1722.0 21.3
line differences
Analysis
203.0
182.1 79.7 19.3 29.7 38.1 183.9 82.2 61.0 251.8 13.8 0.1
all the characters, Results
128.7
1 6 . 8 ~,'-
23.1 15.6 2 . 1 ~,~ 9.7 6.0 12.6 ~ 10.1 6.1 119.5 341.6 117.7
955.0 99.0
All the hybrids were raised in a complete randomised block design replicated four times. Each hybrid was grown in a single row three metres long. Spacing was 75crn between and 10cm within rows. Manuring was at 80kg N, 40kg P~0~ and 30kg K20 per hectare. Observations on four direct yield components - plant height (XI), number of primary branches (X2), number of secondary branches (X3) and number of siliquae on the main axis (X4) - were recorded on five plants at random in each row at the time of harvest. Combining ability analysis was carried out on the line )< tester designs (Kempthorne 1957) and the magnitude of heterosis over the better parent was assessed for each yield component.
X4
16.0
1033.5 924.7 2 6 6 . 3 -~,'638,6 273.0 2353.0 473.8 572.7 12207.6 59682.2 29530.3
significant
X3
16.7
1340.2
m.s.s,
Brassica campestrisL.
of
crosses
X2
1360.4 416.0 2 3 1 . 7 ~'~ 44.9"x 26653.1
1 492
of Hybrids
of Squares
2480.7
4 4 4 1 1
Hybrids
Sum
X1
14
Evaluation
the
while for
inthe
showed the
all the
line,
there
for any character, significant for all interaction
was
A. A m i r t h a
e t a l . : A Q u a n t i t a t i v e E v a l u a t i o n of H y b r i d s o f Brassica campes~ris L.
Devarathinam
Table 2. A n a l y s i s of c o m b i n i n g ability in i n t e r - v a r i e t a l BS • YS
YS • BS
BS x TIR
TR • BS
YS • TR
TR x YS
X1
a b c d e
3223. I 295.6 670.5 54.5 142.8
629.2 3632.6 321.6 90.5 55.6
2084.5 579.9 291.8 52.0 48.2
207. I r 430.8 262.4 2.8 41.1
561.4 407.3 468.5 0.8 92.4
734.4 366.7 583.8 + 121.2
X2
a b c d e
60.3 41.6 9.2 2.1 1.4
X3
a b c d e
X4
a b c d e
a= d = are for
significant ciprocal
I0. I r 21.5 6.7 0.7 +
5.9 ~' 11.5 4 . 6 ~'" 0.2 0.3
475.2 146.0 117.9 9.6 24.1
79.7 11,1 65.8 + 19.7
4 5 . 0 ~64.5 1 6 . 0 "~'~ 2.0 +
21.1 r 2 2 . 1 "'~ 46.3 + 6.2
1971.9 288.1 278.9 42.6 48.2
401.8 220.1 1567.0 36.9 6.1
for XI
and X3.
There
combinations:
There
When
was
used
and line x tester characters
for X2
where
TR
differences XI
only.
were
X4
were was
were
were
(Table
differences
for X2.
Significant
dif-
amonglines for all
2). The tester
not significant.
The line • tester
(TR)
significant line
differences were
reciprocal
for
also
Combining
tion o f p a r e n t s ,
Ability:
An
of the parents
not
differences
overall
G B S II, K a n p u r L o t n i 17, K a n p u r L o t n i 12 i n Y S
and
T 1 6 5 , T 2 4 4 a n d T 1842 i n TIR, a s p o t e n t i a l p a r e n t s f o r inter-varietal
hybridisation
1 9 7 . 0 ~'" 35.7 r 178.4 + 23.1
II >( IB 6) a n d 2
•
w e r e f ound to be t h e b e s t , ning ability (g.c.a.) high specific
having high general
for the parents
combining
consistently
crosses
1•
GBS
( K a n p u r L o t n i 17 • IB 3) i n B S •
ability
combi-
2 and 8 and also
(s.c.a.).
showed high s.c.a,
These two for the yield
but did not p r o v e e q u a l l y good in t h e r e -
YS • BS c o m b i n a t i o n . 6) a n d 3 x 8
2 x 8 (Kanpur
Lotni
( K a n p u r Lotni 27 • IB 6) w e r e t w o with superior cross
g . c . a. f o r all t h e
characters.
The r e c i p r o c a l
8 • 3 was as good
as the cross
3 • 8, i n d i c a t i n g p r a c t i c a l l y
no m a t e r n a l
effects. In the BS
xTR
combination,
5 x 12 and 5 • 13, which ents,
the
involved
showing
yield attributes.
superior BS
g.c.a.,
were
of the TR
parents
to produce
In t h e Y S ing cross,
•
g.c.a,
parents
superior
crosses
the two
4 and 5, with 12 and 13 of TR,
reliable,
SC
were
found
ands.c.a,
2 and 3, which
found
to combine good
combination,
4 • 13, BS
parto be
for some exhibited
well with each
hybrids.
8 • 11wasanoutstand-
showing superior s.c.a, for XI, X 2 and
X3, c o m b i n e d with high g.c.a, for the parent 11 only. This cross, with its reciprocal II • 8, w a s found to be a good specific cross combination. Heterosis and its relation to combining ability: A
( T a b l e 3 ) . T a k i n g all f o u r
into c o n s i d e r a t i o n ,
yield components
components,
39.6 r 2 3 . 1 ~'75.9 + 13.7
1 8 9 . 5 ~'" 282.9 120.1 r 5.8 8.5
other reliable crosses
assessment
led to the identifica-
27 a n d DS 17D i n B S , IB 5, IB 6 a n d E P
ciprocal
dif-
In the recipro-
for all the four characters.
of the performance
crosses
20.6 2 . 2 ~'" 3 . 5 ''~ 0.4 0.1
148.0 52.8 73.1 1.4 12.8
67.0 r 228.1 4 4 . 0 ~'" 5.2 +
17 x l B
hybrids.
significant
and tester
General
the
re-
X2.
significant
among
and X2
observed
except
the line, there were
for XI
significant
significant
as line, the differences interaction
except
ferences
were
for all characters
for all the characters
YS
708.1 263.4 183.5 15.1 24.4
18.7 6 . 9 ~'" 8.8 0.2 0.4
line m.s. (4 d.f.); b = tester m.s. (4 d.f.); c = line• m.s. (16 d.f.); Var(g.c.a.); e = Var(s.c.a.); ''~ = not significant; + = not estimable; all m.s. significant at 5 % level; d,e approximate estimates only (see Arunachalam, 1974 details )
differences
YS-TR
15.2 58.6 4.9 ~ 1,6 0.4
ferences
cal,
crosses
study of the crosses
showing heterosis over better
parent for X2, X 3 and X 4 revealed that 13, 12 and 9 each f r o m B S - Y S ,
BS-TR
and YS-TIR combinations
w e r e heterotic (Table 4). Surprisingly only one cross, I •
s h o w e d s.c.a, effects for all the three charac-
ters, and only three did so for two
crosses,
characters.
2•
7 • 12 andl0•
18 out of the 34 heterotic
A.
Amirtha
Table
Devarathinam
3. Means
et al. : A Quantitative
and g.c.a,
effects in inter-varietal
1
m glf X1
X2
X3
X4
2
120.1 4.0
1.5 3.2 0.9
- 2 2 . 6 '" - 1 6 . 6 '`. - 7.2'"
6.4 0.5 1.4'`. 0.2
-
0.3
- 0.6
m glf glm g2f
8.9 0.3 1.4'`. 0.7
10.2 2 . 3 .> 0.8'`. 1.1':"
g2m
0.6
0.2
m glf glm g2f g2m
20.7 0.8 1.2 1.4 1.9'`.
21.1 4.8'" 0.0 1.9"`. 0.6
m gif glm g2f
44.7 1.8
-
-
g2m
-
-
-
83.1 -20.0""
12.6 4 . 3 ''~ 1.7 1.3 0.3
53.1 1 0 . 7 '`. 5 . 9 ~`. 3.1 - 1.1
38.7
7.3 2.4" 2.2'`. 0.5
-
15.6 5.7'`. 2 . 3 ~`. 0.2 - 0.4 -
5 . 9 ~'" 2.5
0.4
- 3.6
crosses X2,
X3
did not show or X4.
fects were
It was
-
in general
It was
found
intermediate
was
1.7 6 . 5 "~" 0.4
7.2 0 . 8 ~''" 1.4'`. 0.1
9.5 0.6 0.1 1.5'`.
1.2 ~`.
0.3
ent with regard
andX4
in which
parent
(Table
5.4'`.
could
found
13 crosses
out of 34 showed 9 over
showing
between
60 to 70.
for X3
(Table
parent),
9 to YS-TR
combinations. and X4
pectively.
Using
cy of crosses
qual to or above will be named ally for X2, (= 41%)
were
X3
The
these
values
showed
the norms
for X3
hybrids
mean
as norms,
was
found
res-
the frequenof heterosis
worked
of which
and
per cent
and 34,
for 2 or 3 characters
It was
hybrids,
(over su-
heterosis
an amount
5.6 3.2"" 1.6 3 . 5 .> 1.4
-
47.2 2.5 0.8 3.5
-
'`.
0.2
g.c.a,
0.2
effect
combinations
(Table
5). BS-YS
rank
X2,
which X3
also contributed
were
rank
andX4.
11 • 8 were
hybrid
and the rest by BS-TR
hybrids
four of
for all the
The cross8x11andits
both rank
3 • 12 belonging
hybrids
s. c. a. effect for any of the three
re-
in YS-TR.
to BS-TR
did not
The have
characters.
Discussion For
the first time,
in this study,
were
chosen
diversity
attempt
(which
that 14 out of 34 con-
to represent
was
and the primary
interest
ance
of inter-varietal
ter-varietal
hybrids
among
described form
lower
a diallel system,
entries, the BS,
but the system YS
and TR
ents in a series
Since
parents
for populathe intra-var-
than the inter-varietal
was
to evaluate
hybrids,
The
made,
of genetic
BS-TR
a set of possible
and YS-TR
set of crosses
populations
of line x tester
in-
distributed was
made,
as
did not exactly
due to the absence adopted
one
the perform-
and their reciprocals
BS-YS,
earlier.
whose
degree
important
selection.
been
of combining
crosses
a good
for yield components and human
has
the components
ietal divergence
equally e-
out individu-
50 % were
a serious
to analyse
ability in a set of inter-varietal and
was ob-
12 to BS-TR
to be 28,49
Hybrids)
and X4. rank
3
mostofthecros-
to BS-YS,
found
Rank
20to40 cent,
highest heterosis
inter-varietal
was
which
and YS-TR
tion fitness
13 belonged
for X2,
X3
The
X3
4).
Of the 34 heterotic perior
per
50 per cent heterosis
20 to 50 per cent for X4. served
Similarly,
40
0.0
of heter-
for X2
less than
by BS-YS
ciprocal
better
for X2,
heterosis
and
were
were
a range
be
10.7 1.0~' 0.2 0.6
-
inferior par-
percent
ses showed
4).
thanthe
However,
parent
crosses
with the parents.
the hybrids
or worse
to height.
superior
tributed
characters
compared
38.8 2.2 1.1 4.3
-
3.0 3.7 1.5
3.9 2 . 6 ~`. 1.5 3.8'`. 0.8
46.8 3.1 1.6 1.8
the eight hybrids
4).
when
noted
ef-
inter-varietal
that the inter-varietal
than the superior
osis over
(Table
in height
instance
that the s.c.a,
in heterotic
123.4 1.4
2 : BS-TB
effects for any character,
thus apparent
not pronounced
crosses
No
s. c.a.
107.5 3.3
16.1 4.2'`. 1.8 1 . 9 "~`. 0.5
-
7
5.2'`. 2.9 2.2
g I = BS-YS;
Brassica campestrisL.
6
129.7 0.0
24.8 -16.3'`. - 1 5 . 4 "~ - 1 0 . 6 '`.
0 . 8
of
5
122.0 0.8
13.5" 5 . 0 ~`. 5.2"
3.1 1.2
BS 4
122.1 15.2 '`.
2.5 5 . 5 "`. 0.8
of Hybrids
hybrids
3
glm g2f g2m
2.0
Evaluation
enabled as male designs.
of diagonal us to evaluate
andfemalepar-
A.
Amirtha
Devarathinam
et al. : A Quantitative
Evaluation
of Brassica
of Hybrids
YS 8 107.1 3.5 4.4
9
10
11
118.9 9.9 r 5.2 4.8'`.
130.6 1.7 0.6 5.1~"
75.9 0.2 2.6
2.6
-
2.8
-
4.7'`.
7.5 0.3 2.4'`. 0.3 0.0
-
10.1 0.9"`. 1.4 0.5 0.3
-
-
8.3 0.8 1.8 1.4
-
-
1.8 2.0'" 2 . 4 '~ 1.1
-
0.6
-
1.7
-
r
-
38.1 3.2 5.1'`.
-
2.4
-
1 = BS-YS;
The
-
0.7
0.4
-
2.1
• YS,
were
high for X2 in YS
crop
was
plants
both additive
(see
for
example,
1970,
YS
proved
As
used
gene
ac-
1973,
in
on YS;
parents suitable
Zu-
et al. 1973,
on Sorgh~).
of the combining
on
Hence
ability ef-
would
be
breeding
The
YS
a pre-
procedures
to be a success
as a female
a large
of promisinghy-
should
divergent
especially
crop, show
when
YS
used
chosen
should
in the cross
as females.
also recorded or
par-
also comparatively low within-line
divergence
to find that BS
as lines against
number
genotypes
their source
= =
as
BS-TR;
2
of the BS
It was success
estimates
importance
TIR as testers.
cross
33.2 3.0 4.0'`.
0.6
0.7
3 . 7 ''~
combinations
during
of maize
I, 2 and 3 could
could
types
bine well with TR
of gene
as testers
duce
with
hybrids was
limited
highly cross-pollinated,
a result.
population
was
der to have
stressed
for example)
other
example.
The
associated
study
before
high
for
of intra-
for the genetic effects crosses
(1966).
in inter-varietal cross-pollinated,
the parents making
was
with
be responsible
and Cockerham
were
pro-
parent.
parent
of inter-population
to improve
hybrids.
could
other
a closer
success
both the parents
be necessary tion,
FI
by Stuber greater
for
in BS-YS
good
as the
to account
in the
This ap-
foundtocom-
of BS
the
might
for
action.
5 were
usually
populations The need
variability
have
where
TR,
variability
open-pollinated such
YS
when
be-
relative
in that only parents
and produce
populations
heterotic
markedly the
as testers
4 and
In fact, the SI parental
The success
vary and
of BS
with YS
parents
that var-
as lines and that the
variations
combine
while
on open-pol-
and their hybrids
variance
very
components.
a study
in their potential
frequency
combination,
showed
for the yield
observed
of different
combi-
In fact,
1.0
2.6
to be true in the case
and variances
in-
11.5 1.3 1.3 1.3
32.4 0.1 3.4
-
of genetic
of gene
peared
5`.
ability
(1964)
vari-
when
14.0 0.6 1.6 0.4
-
• YS
ieties differed
cause
-
'~"
= YS-TIR
combining
Sprague
5.4 0.5 1.4 0.1 0.7
-
male
intra-population
a self pollinated
late flowering,
nation,
ac-
components
Giriraj 1965,
and TR
ent and has produced
teresting
gene
varieties/hybrids.
The SC YS
m
linated varieties
combinations,
additive
Singh
on TR;
for formulating
While
Brassicacampestris;
on
understanding
fects of the BS,
varietal
all yield
and Webster
reflected
6.3
variance
and non-additive
1973,
Kambal
ability.
- 5.2 '"
1.1
high for all thechar-
over
for almost
beri andAhmad
brids.
2.1
and
-
6.3 0.5 0.4 1.4'`. 0.1
3.3
-
2.1""
1.0
female
4.1'`.
0.0
0.7 as
0.3
-
-
32.6 0.8 4 . 0 '~
high
In the other
14.6 0.6 1.0 0.2
-
40.5 1.8 2.9
f =
of genetic
5.3 0.4 0.2 0.9" 0.4
1.0
some
they were
castor;
to evolve
"~"
s.c.a.
predominant
and Eunus
requisite
level
11.4 1.0 3.0" 1.0
80.3 4.6'`. 8.7'"
-
7.2"
39.8 0.5 0.5 -
of g.c.a,
components
x BS.
Usually
a detailed
~
9.5'"
-
15
69.1 0.1 - 1.7
2.4
-
5
14
60.3 4.5','4 . 5 '~-
6.9 0.7 1.4" 1.1"" 0.1
both additive andnon-ad-
estimates
only,
tion are present
Joarder
5
73.1 0.3 3.3 1.8 1.7
I
2) revealed
additive
non-additive tion.
at
'~"
2.0
action for all the yield components.
in BS
acters
-
-
2 = YS-TR
(Table
ditive gene
0.0
-
approximate
variances
0.1
11.6 0.9 - 1.1 1.7
0.8
-
4.7 0.3 0.5 0.4 0.9
17.3 0.7 3.7'`.
35.1 1.7 1.9
r = significant
6.7""
10.6 1 . 0 "~`. 0.8':" 1.2'" 0.6
-
42.4 4 . 5 ':" 5 . 1 ~`.
1.7
0.2
6.4"
TR 13
12
-
cg~pestris L.
the
(through hybrids.
In or-
crosses, it would selecThis
A. Amirtha T a b l e 4. M e a n s ,
Devarathinam
heterosis
Cross
and s.c.a,
BS-YS
1 x 8 2 • 6 2 x 7 2 X 8 3 X 8 3 • 9 6 X 2 6 • 3 6 • 5 8 • 2 8 • 3 8 X 4 8 X 5 3 xll 3 X 12 3 X 13 3>(14 3 • 15 4 • 4 x 13 4 • 14 4 • 15 5 • 12 5 • 13 11 • 4 7 • 8 • 8 • 12 8 • 14 8• 10X14 11 • 8 14 X 8 14 X l 0
YS-BS
BS-TR
TR-BS YS-TR
TR-YS
increase
16.3 14.0 11.5 15.3 12.2 10.4 11.1 13.3 11.9 11.4 12.6 9.6 10.7 9.5 10.6 7.5 8.9 8.2 9.2 9.0 7.8 7.4 11.3 8.9 8.3 12.5 12.3 8.5 9.1 9,1 13.0 9.5 8.5 11.4
the base
dify his programme
single
binations, 2 •
possible
cross
with high g.c.a. the case
to locate
and 3 x 9, each
hand,
superior
heterosis
h
9.1"" 8 . 8 ~,," -2.1 -1.7 - 4 . 1 "*'9.9~ 4.7 ~ 3.5 -2.2 1.5 1.0 4 . 0 "~:-2.6 -1.4 2.9 0.3 -0.9 -0.8 0.6 0.9 -1.2 -0.5 -0.5 -3.0 1.5 6.4~' 3.9 -3.3 -1.5 3.5 5.1~ 2.5 -0.3 3,0
of hy-
served
for sev-
yield
five crosses, of which
superior
in those
the crosses for three
2 x6
yield
68 58 25
parent;
-
Sim-
even ob-
-
4.7
27
-
1.4 7.4 1.1 5.7
57 40 24 79
-
9.7,',"
7
5.2 -
32 49 23 34 28 40 20 33 39 4 24 10 30 31 34 5 27 47 28
7 . 5 ~
3.0 2.2 1.3 1.8
-
-
9.4
~
3.0 5.8 0.2 2.3 0.4 8 . 4 "~ 7.2 7.6 ~ 0.7 5.8 4.8 1.2
-
-
3.2
32
2.5 0.6 8 . 9 "' 4.7 6.3
26 71
42 17 47
~- = significant
two crosses
was,
at
therefore,
of the high g. c.a.
due
to
of the parents
(of
2 and 8, in particular). The
results
with the reciprocal
did not fall in line with those Two
crosses,
X3
and X4;
as observed 8 as female found
against
other
of the yield components, of parent
The heterosis sistently
to show
each
were
marked
as high as in BS-YS
heter-
involving
of the BS
out the
programme. was
combinations
not con-
for the ma-
the crosses
4 x 12 and 4 x 15, which
would
deserve
as
for one or
bringing
combinations
the
parents
heterosis thus
he-
point
in manifesting crosses
Nevertheless,
3 x 13, 3 x 14,
bited high heterosis,
perhaps
8 in the hybridisalion
in BS-TR
jor yield components. 3 x 12,
substantial
this would
The
male
superiority
combinations.
6 x 3 and 8 x 3, showed
for X2,
parent
YS-BScombinations
of BS-YS
earlier.
and 3 x 9. On
heterosis
12.1 x~ 15.9 ~ 1.2
of the parents
combined
components
75.2 83.8 66.5 67.3 60.7 59.2 65.6 69.6 49.9 70.2 57 5 46 7 62 9 51 1 56 6 46 5 51 5 53 7 41 9 40.4 35.7 43.1 61.3 62.7 41.7 59.9 58.3 51.9 50.3 48.0 59.9 56.6 44.5 51.7
osis,
2 x 8 and 3 x 8 had shown
of high s. c. a. The high
h
to the superiority
in perform-
s.c.a,
s
I •
Of
8, in particular).
superior
m
com-
expressed
components.
48 59 9 14 68 145 20 105 25 2 79 39 19 39 105 45 25 42 44 33 3 10 43 15 21 163 72 109 27 78 51 76 16 17
the reinforcement
also mo-
: In BS-YS
1 • 8 was outstanding showed
s
per cent over
than modifying
crosses
in the crosses
the other
in the absence
or
should
rather
(of the parent
crosses X4
terosis
for all major
five crosses,
ilar was
30.7 33.6 22.9 24.0 21.1 30.9 25.3 25.8 20.1 21.5 22.6 21.6 19.1 17.5 25.8 21.2 17.5 17.9 22.4 20.7 16.1 17.1 23.0 18.5 18.8 30.0 19.9 15.7 18.6 20.5 26.2 20.4 16.2 20.2
separately.
it was
This
60 37 8 50 63 3 9 40 25 12 68 28 43 48 54 17 4 28 26 23 7 1 57 24 14 17 64 23 21 21 23 27 13 8
of the hy-
improvement
in inter-varietal
2 • 8, 3 x8
ance.
m
sampling
inbreeding
breeder
simultaneously
high heterosis those
The to achieve
attributes
Heterosis
performance
in an orthodox
programme.
eral attributes
-1.1 -0.5 0.3 2 . 4 ~'" 1.4 -0.6 1 . 7 ~,'~
extent than the repeated
population
bridisation
3.3 r 1.3 -0.9 0.3 -0.9 1.1 -0.7 0.9 -0,5 -0.1 0.2 1.1 -1.4 0.1 0.3 -1.3 0.3 0.6 -0.4 1.0 -0.1 0.5 1.3 0.5 0.4 2 . 6 ~" 1 . 9 -'>
h
effect:, h = heterosis t-test only)
the expected
brid to a greater
inter-varietal
X3 s
m : mean; s = s.c.a, 5 % level (approximate
should
e f f e c t s of h e t e r o t i c
X2 m
Brassica campestris L.
et a l . : A Q u a n t i t a t i v e E v a l u a t i o n of H y b r i d s of
consideration.
3 X 11, exhiIt
A.
Amirtha
Devarathinam
et al. : A Quantitative
Table 5. Distribution of heterotic ses for major yield components Per Cent Heterosis
inter-varietal
Number
cros-
12 13 6 3
X4
10 7 7 5 3 1 1
-
6 19 6 3 -
-
coner
(1964)
needs
modifying
of using
a
b
Total
n
p
n
p
n
p
Total
7 4 3
50 29 21
6 8 6
30 40 30
13 12 9
14
41
20
59
34
and TR
in a marked
38 35 27
because
in the way
in vigour,
be
several
should
such
effects
sacrifice
1967).
and
in
The
crosses
dif-
effected
of bulking
plants). very
the cros-
The heterosis
greatly
in Brassica
as observed
the harmful
set with some
as also observed
(instead
not be
result
if a representative
are identified
from
hybrids
found
would
(Thomas
overcome
plants only
hy-
was
of the rigid sys-
Inbreeding
for example
partly
those
Thus,
stand
for inter-varietal
populations,
and female
general,
therefore
application.
could
parents
decrease
ficulty can
sed seeds
7
A high level of heterozygosity
Loli~m perenne,
such BS-YS BS-TR YS-TR
difficulties
tern of self-incompatibility.
male
L.
diallelic model,
to be of general
highly inbred
bridisation. in the BS
cmmpest~s
on a one-locus
practical
between
Combination
of Brc~sica
of Hybrids
Some
of Crosses X3
X2
1- 20 2 1 - 40 41- 60 6 1 - 80 81-100 101-120 121-160 161 a n d a b o v e
Evaluation
of
affected
in
o leracea (Watts 1970).
of inbreeding
could
of homogeneity
be off-
in the FI
hy-
brids. A number Crosses
heterotic
for X2,
BS-YS YS-BS BS-TR YS-TR
lX 8 • 3 • 8 •
X3
andX4
8 ~H~', 2 X 6 "~, 3 X 3 12 + 1 1 , 11 • 8
8, 3 X 9
hybridisation
brid progenies (whatever double
as males 1963,
(Welsh
superior
combining
Similar were
results
also found
reciprocal
was
7 x 12 which
parents
which
ability produced to those
and achieve
for YS-TR.
The cross
produced
lines could
8 x 11 and its
neously
Another
heterosis
was
of 163 %
for X3. Heterosis ence
in the gene
character ents
(Falconer
ability.
result found
1964).
to show
non-additive
heterosis components
tion of the expression
hybrids
inthe par-
combining
to be true.
A even
number
of crosses
in the
of gene of heterosis,
absence
action.
The high did not was of high
The implica-
as based
by Fal-
hy-
through ( Ram, Doloiand high andlow
forward
simulta-
would
Comstock
Brassica campestr~s,
sed of as many possible.
of open be
and Ro1957,
good
combining indicates
be built using
pollination
possible
synthetic
This study can even
of
and shuffle the de(Comstock
and Robinson
it should through
effects
and
1963).
of the high degree
yield improvement
of the inter-
the harmful
disequilibrium
Gates,
inter-mated, (c) Bi-parental
generations
obviate
and Cockerham
Because
stage,
in one recombinant
1952;
Matzinger
complex
ex-
where
populations.
and advanced
genes
9 • 12 and 10 x 3 for X1
in high heterosis.
in F2
and linkage
not always
in the crosses
matings
binson
would
Arunachalam,
selection,
and at a later
sirable
to
time
yield improvement
and carried
productive
linkage
with high specific
This appeared
provide
a quantitative
customary
and Dudley
Programmeslike
in the inter-varietal
Murty, way
be selected
controlling of dominance
varieties
on both the right and left tails of the phenoty-
of the differ-
It is, therefore,
high heterosis
1969;
of the square
frequencies
hybrids
in a set of yield components
(b) Two
pic distribution, would
hy-
be - single,
Levings
1966).
for flowering
further
varietal
and also the amount
associate
s.c.a,
is a function
and Doloi 1972).
1973;
variation
response
combinations
combination.
the maximum
Murty Ram
high heterosis.
for BS-TR
an excellent
had shown
selection
may
or high yielding
and Atkins
would
the inter-varietal
hybrids
and Robinson
(a) disruptive
be triedin
and either superior
crosses)
ploit the intra-level
can
One such system
using
as females
and Moll
correlated that only those
systems
the level of these
or multiple
brids
observed
mating
a bid to boost the performance. be multiple
a = crosses showing high heterosis for 2 or 3 characters out of X2, X3, X4 b ..... do ............... one or none .... do .... n = number; p = per cent; + = s.c.a, effects absent; ',,~= s.c.a, effect for X3, X4; ~,~r = s.c.a, effects for all characters, other crosses had s. c. a. effect for one character only.
was
of useful
BS,
to effect
varieties
parental that
such YS
in
compo-
genotypes
as
a synthetic and TR,
es-
8
A. Amirtha
pecially in view
Devarathinam
of the complex
et al. : A Quantitative
level of cross
compa-
tibility found in them. This study on inter-varietal cates that the differences mainly ferences
in flowering
sion of characters. we considered show
direction
inter-varietal
capable
clearly
shown
by the range
population
improvement
were
found
crosses.
and nature
progenies.
of brown
joint
Nevertheless,
by inter-varietal
to the background
the
of the s. c. a. of the
of the varieties
that breeders
ford to relegate
if
could
cases,
out the desirable
iability in the F2 and biparental thus conclude
hybrids
g. c. a. effects a-
of the parents.
of recombination
This was
on pure
cancelled
attributes
gain ground
could not. In some
and magnitude
cross
effects of the g.c.a, the desirable
ideas would
No cryptic
in the expres-
inter-varietal
based
others
indi-
and dif-
of the varieties.
These
good heterosis
system
could be detected
that some
lone, while some opposing
pollinating
time
differences
clearly
in the three varieties were
due to the imposed
structural
crosses
One
sarson
of varcan
can ill af-
the potentialities of
by inter-varietal
hybridisa-
tion.
the use of a moJ. Genet. 3_~4,
280-287 (1974) C o m s t o c k , R . E . , R o b i n s o n , H . F . : E s t i m a t e s of a v e r a g e d o m i n a n c e o f g e n e s . In: H e t e r o s i s , 494-516. A m e s : Iowa State College P r e s s 1952 F a l c o n e r , D.S. : I n t r o d u c t i o n to Q u a n t i t a t i v e G e n e t i c s . E d i n b u r g h and London: O l i v e r and Boyd 1964 Gates, C . E . ; C o m s t o c k , R . E . ~ R o b i n s o n , H . F . : G e n e r a l i s e d g e n e t i c v a r i a n c e and c o v a r i a n c e f o r m u l a e for s e l f - f e r t i l i s e d c r o p s a s s u m i n g l i n k a g e . G e n e t i c s 42, 749-763 (1957) G i r i r a j , K . , M e n s i n k a i , S . W . , Sindagi, S . S . : C o m bining ability of s o m e q u a n t i t a t i v e c h a r a c t e r s in 6 • 6 diallel cross of castor (Ricinus communis L. ). Indian J. Agric. Sci. 43, 319-322 (1973) Joarder, O.I., Eunus, A.M.: Genetic studies of two crosses of Brassica cat,pest,s. Exptl. Agric. 6,
Received December 26, 1975 Communicated by H. Stubbe
of Hybrids
of Brassica campestris L.
351-357 (1970) Kambal, A.E.; Webster, O.J.: Estimates of general and specific combining ability in grain sorghum, Sorghum vulg~e Pers. Crop Sci. 5, 521-523 (1965) Kempthorne, O. : An introduction to Genetic Statistics. New York: John Wiley and Sons 1957 Levings Ill, C.S. ; Dudley, J.W. : Evaluation of certain mating designs for estimation of genetic variance in auto-tetraploid alfalfa. Crop Sci. 3, 532-
535 (1963) Matzinger, D.F.; Cockerham, C.C.: Simultaneous s e l f i n g and p a r t i a l diallel set c r o s s i n g . I. E s t i m a tion of g e n e t i c and e n v i r o n m e n t a l p a r a m e t e r s . C r o p Sci. 3, 309-314 (1963) Moll, R . H . , R o b i n s o n , H . F ~ : O b s e r v e d and expected r e s p o n s e in four s e l e c t i o n e x p e r i m e n t s in m a i z e . C r o p Sci. 6, 319-324 (1966) Murty, B . R . , A r u n a c h a l a m , V.I Doloi, P . C . , R a m , J. : Effects of d i s r u p t i v e s e l e c t i o n for f l o w e r i n g t i m e in Brassica campestria v a r . brown s a r s o n . Her e d i t y 2._~8, 287-295 {1972) R a j a n , S.S. : The e v o l u t i o n of yellow s a r s o n ( B r a a s i c a car~pest~s v a t . s a r s o n ) , a s e l f - c o m p a t i b l e r a c e in a s e l f - i n c o m p a t i b l e group. P r o c . X Int. Cong. G e n e t . , C a n a d a , 336-337 (1958) R a m , J . ; Murty, B . R . , Doloi, P . C . : I m p r o v e m e n t in s e e d y i e l d by d i s r u p t i v e s e l e c t i o n for flowering t i m e in brown s a r s o n . Indian J. Genet. 2__99, 192201 (1969) Singh, D. : I m p r o v e m e n t of r a p e and m u s t a r d . In: Rape and M u s t a r d , I . C . O . C . , H y d e r a b a d (1958) Singh, J . N . : Diallel a n a l y s i s of c o m b i n i n g a b i l i t y and its i n t e r a c t i o n s with e n v i r o n m e n t s for s o m e y i e l d c o m p o n e n t s in ~rassica campestris var. yellow sarson. Ph. D. Thesis, Benares Hindu University, India (1973)
Literature Arunachalam, V. : The fallacy behind dified line • tester design. Indian
Evaluation
Spra~ue, G . F . : E s t i m a t e s of g e n e t i c v a r i a t i o n s i n t w o open p o l l i n a t e d v a r i e t i e s of m a i z e and t h e i r r e c i p r o c a l F1 h y b r i d s . C r o p S c i . 4, 332-334 (1964) S t u b e r , C . W . ; C o c k e r h a m , C . C . : Gene e f f e c t s and v a r i a n c e s in h y b r i d p o p u l a t i o n s . G e n e t i c s 5__44,12791286 (1966) Thomas, R.L. : I n t e r - p o p u l a t i o n v a r i a t i o n in p e r e n n i a l r y e - g r a s s . I. P o p u l a t i o n m e a n s . H e r e d i t y 2_~2, 481498 (1967) Watts, L . E . : C o m p a r a t i v e r e s p o n s e of botanical v a r i e t i e s of Brassica aleracea L. to i n b r e e d i n g a n d h y bridisation, Brussel sprouts, sprouting broccoli and n a r r o w s t e m k a l e . E u p h y t i c a 19, 78-90 (1970) Welsh, E . J . , A t k i n s , R . E . : P e r f o r m a n c e and within h y b r i d v a r i a b i l i t y of t h r e e way and s i n g l e c r o s s e s of g r a i n s o r g h u m . C r o p Sci. 1__33, 267-271 (1973) Zuberi, M~ Abroad, S . V . : G e n e t i c study of y i e l d and s o m e of its c o m p o n e n t s in Braasica campestris L. v a r . t a r i a . C r o p Sci. 1__33, 13-15 (1973)
Dr. A. Amirtha Devarathinam Deputy Agricultural Officer Seed Testing Laboratory Thirunagar Madural 625006 (India) Dr. V. Arunachalam Division of Genetics Dr. IB.R. Murty Project Director Nuclear Research Laboratory Indian Agricultural Research New Delhi 110012 (India)
Institute
A Quantitative Evaluation of Inter-varietal Hybrids of Brassica campestris L.* A.A.
Devarathinam,
V. Arunachalam
D i v i s i o n of G e n e t i c s ,
Summary.
andB.R.
Indian Agricultural
Murty
Research
Institute,
New Delhi (India)
A quantitative
e v a l u a t i o n of y i e l d c o m p o n e n t s w a s c a r r i e d out in 150 i n t e r - v a r i e t a l h y b r i d s of Brasbrown sarson (BS), yellow sarson (YS) and toria (TR). The r e s u l t s s h o w e d b o t h a d d i t i v e a n d n o n - a d d i t i v e g e n e a c t i o n f o r p l a n t h e i g h t , n u m b e r of p r i m a r y a n d s e c o n d a r y b r a n c h e s a n d n u m b e r of s i l i q u a e o n t h e m a i n a x i s in a l l t h e s i x c r o s s c o m b i n a t i o n s - B S - Y S , Y S - B S , B S - T R , T R - B S , Y S - T R a n d T R - Y S . T h e g e n e r a l c o m b i n i n g a b i l i t i e s of B S , YS a n d TR i n d i c a t e d t h a t t h e i r n a t u r e a n d m a g n i t u d e d e p e n d e d l a r g e l y o n t h e o t h e r p a r e n t s w h i c h e n t e r e d t h e h y b r i d s . T h e r e w e r e d i f f e r e n c e s in c o m b i n i n g a b i l i t y b e t w e e n d i r e c t a n d r e c i p r o c a l c o m b i n a t i o n s . G B S t I , K a n p u r Lotni 17, K a n p u r L o t n i 27 a n d DS 17D in B S , IB 3, IB 5, I B 6 a n d E P 12 in YS, a n d T 1 6 5 , T 244 a n d T 1842 in TR w e r e i d e n t i f i e d a s p o t e n t i a l p a r e n t s f o r i n t e r - v a r i e t a l h y b r i d i s a t i o n . R e c i p r o c a l e f f e c t s w e r e f o u n d w h e n BS o r YS w a s u s e d a s p a r e n t a n d t h e y w e r e l e a s t w h e n TR w a s u s e d a s a p a r e n t . T h e a m o u n t a n d d e g r e e of h e t e r o s i s w a s s u b s t a n t i a l in i n t e r - v a r i e t a l c r o s ses. Based on the heterosis-combining a b i l i t y r e l a t i o n s h i p , t h e r o l e of i n t e r - v a r i e t a l h y b r i d i s a t i o n in p o p u l a t i o n i m p r o v e m e n t of Brassica carapestrgs i s d i s c u s s e d . A n u m b e r of m e t h o d s of u t i l i s i n g t h e i n t e r - v a r i e t a l h y b r i d s i n m u l t i p l e c r o s s e s a n d s y n t h e t i c c o m p l e x e s i s s u g g e s t e d a s p o t e n t i a l s u p p l e m e n t s to p o p u l a t i o n b r e e d i n g in this crop.
sica campestris u s i n g f i v e e a c h of t h e t h r e e v a r i e t i e s ,
Introduction
w h i c h c o u l d p r o f i t a b l y b e u s e d in m a r k i n g
The o l e i f e r o u s
ful s h o r t t e r m
varieties,
Brassica campestris L. i n c l u d e s
brown sarson(BS),
yellow sarson(YS)
toria(TR),
with distinct genetic
differences
but i n t e r m a t i n g
1958).
W h i l e YS a n d T R
compatible
forms.
changes in flowering changes,
time,
have restricted
ation within each variety
isolation
de-
due to
down the vari-
( R a j a h 1 9 5 8 ) . D u r i n g at l e a s t
t h r e e d e c a d e s no s i g n i f i c a n t a c h i e v e m e n t s
have been
m a d e i n i m p r o v i n g t h e y i e l d of t h e s e p o p u l a t i o n s . tempts
BS
and also chromosomal
and narrowed
At-
w e r e c o n f i n e d m a i n l y to i m p r o v i n g t h e y i e l d of
the three varieties
separately
proved breeding procedures. procedures
by c o n v e n t i o n a l
and im-
T h o u g h t h e m e r i t s of t h e s e
c a n not b e u n d e r r a t e d ,
it w o u l d b e d e s i r -
a b l e to p l a n a s t u d y t o e x a m i n e t h e b a s i c p o t e n t i a l i t i e s of i n t e r - v a r i e t a l to f u r t h e r
hybridisation
reported
conducted in this
Material
and Methods
W e w o u l d l i k e to u s e t h e t e r m , v a r i e t y , w h e n r e f e r r i n g to BS, YS a n d T B a n d t h e t e r m , p o p u l a t i o n s , to r e f e r to d i f f e r e n t s u b - v a r i e t i e s of BS, YS a n d TR. F i f t e e n p o p u l a t i o n s , t h r e e s e t s of f i v e e a c h , i n t h e three varieties as detailed below, were utilised as p a r e n t s in t h i s s t u d y . B__.SS: 1 - G B S II ( I N T ) ; 2 - K a n p u r Lotni 17 ( S I ) ; 3 - K a n p u r Lotni 27 ( S I ) ; 4 - D S 1 7 D ( S C ) ; 5- A s s a m L o c a l ( S C ) YS: 6 - IB 3; 7 - IB 5; 8 - IB 6; 9- IB 7 1 ; 1 0 - E P 12 TR: 1 1 - T 165; 1 2 - T 2 1 7 ; 1 3 T 244; 14- T 267; 15- T 1842. The p a r e n t s 2 a n d 3 i n BS w e r e s e l f - i n c o m p a t i b l e (SI), 4 and 5 were self-compatible (SC) and 1intermediate-compatible (INT). InYS, the exotic parent 10 h a d b i v a l v e d , w h i l e t h e o t h e r f o u r h a d m u l t i v a l v e d s i l i q u a e . The TR p a r e n t s d i f f e r e d i n f l o w e r i n g d u r a tion, besides other characters. E a c h s e t of p a r e n t s served both as lines and as testers for the other and t h i s p r o c e s s r e s u l t e d i n 150 h y b r i d s , i n c l u d i n g 75 c r o s s e s a n d 75 r e c i p r o c a l s a s s h o w n b e l o w :
among these varieties,
boost population improvement
Tester
with regard
to s e e d a n d oil y i e l d . S u c h a s t u d y w o u l d a l s o p r o v i d e a w e a l t h of b a s i c i n f o r m a t i o n
crosses, are
of s u c h
paper.
and self-in-
of s u c h v a r i a b l e
temporal
out p o w e r -
The results
self-com-
groups respectively,
intermediate
an investigation on inter-varietal in t h i s c r o p f o r t h e f i r s t t i m e ,
and morphological
belong to strictly
The p r e s e n c e
g r e e s of i n c o m p a t i b i l i t y ,
and
with one another (Singh
patible and self-incompatible includes self-compatible,
three
breeding methods.
on the yield components
"" P a r t of a P h . D . t h e s i s s u b m i t t e d by s e n i o r a u t h o r to P . G . S c h o o l , I n d i a n A g r i c u l t u r a l R e s e a r c h I n s t i tute, New Delhi
Li n e BS YS TR
9 Crosses o Crosses
made not m a d e
BS
YS
TR
o
9
9 9
o
9 9
9
o
A. Table
Amirtha
I. ANOVA
Devarathinam
et al. : A Quantitative
for four yield components
in inter-varietal
Mean Source
d.f.
Parents
(P)
BS YS TR BS vs YS TR vs Rest (H)
149
BS • YS • a vs b BS • TR TR x BS e vs d YS xTR TR XYS e vs f (BS-YS) (YS-TR)
(a) (b)
24 24 1 24 24 1 24 24 1 1 1
(c) (d) (e) (f) vs (BS-TR) vs BS crosses
P vs H Error ~" N o t s i g n i f i c a n t ;
all other
9.5 6 . 8 -,'~ 3 . 0 ~''" 28.9 129.0
91.6 3.2
among
of variance the parents,
for X3,
those
(Table
A further
showed
was
The TR
nations
fered
from
s.s.
due to lines, study
The hybrids corresponding
s.s.
followed
did not show IBS vs YS mean
significant and YS
of squares
BS-YS
for X2
and X3,
for all characters was
to various
further
significant sum
of squares
while
except
partitioned
inter-varietal
TR
cross
difX3.
into those combi-
Each
one of the cross
subdivided
and line • tester
components
between
(Table
into that
interaction.
A
2 ) brings out the fol-
Significant
the 50 hybrids
differences
(25 crosses
•
BS
where
ter and line x tester
were
differences whereas
for X3
between
asline,
were
Inthe line, tes-
significant
inthereciprocals,
and line • tester (Table
BS • YS
interaction
2). The
and YS
for
YSXBS,
x BS
reciprocal werenot
sig-
for any character. combinations: were
characters
for X3
were
non-significant (TR)
differences
in the crosses
differences
(Table
no significant (BS)
except
BS
for X2
were
2). When
the 50
TR
as line
and X3,
was
differences
were
Line x tester
but
and for X4
significant
line differences
X3.
among
for all the characters,
not significant.
differences
but the tester characters
The
significant
the differences
were
served
not significant
differences
tester
BS
were and 25
crosses,
reciprocals var-
6807.6 86.1
for all the four yield components.
(s.s.) IBS
"" -,',~
reciprocals)
hybrids
popula-
~' ""
further
combinations:
observed
were
among
by YS
562.6 401.8 2 . 5 ~'~ 284.3 78.5 r 1382.3 150.8 157.7 2329.7 8738.5 1232.2
lowing :
BS-TR of the sum
was
testers
of these
nificant
vs Hybrids
that the variation
populations
BS
except
402.3
and their reciprocals.
combinations
for X2
differences
the hybrids,
Parents
the highest,
iation for any character. was
The
459.2 8 6 . 8 ~'~ 6 6 . 3 ~'17.2 ~ 373.8
at 5 % l e v e l
1 ).
partitioning
due to parents populations
(m.s.)
among
and the comparison
all significant
tions.
of plot means:
5 2 . 5 -~ 145.1 9 . 4 "" 972.2 1 . 3 ~,~
1722.0 21.3
line differences
Analysis
203.0
182.1 79.7 19.3 29.7 38.1 183.9 82.2 61.0 251.8 13.8 0.1
all the characters, Results
128.7
1 6 . 8 ~,'-
23.1 15.6 2 . 1 ~,~ 9.7 6.0 12.6 ~ 10.1 6.1 119.5 341.6 117.7
955.0 99.0
All the hybrids were raised in a complete randomised block design replicated four times. Each hybrid was grown in a single row three metres long. Spacing was 75crn between and 10cm within rows. Manuring was at 80kg N, 40kg P~0~ and 30kg K20 per hectare. Observations on four direct yield components - plant height (XI), number of primary branches (X2), number of secondary branches (X3) and number of siliquae on the main axis (X4) - were recorded on five plants at random in each row at the time of harvest. Combining ability analysis was carried out on the line )< tester designs (Kempthorne 1957) and the magnitude of heterosis over the better parent was assessed for each yield component.
X4
16.0
1033.5 924.7 2 6 6 . 3 -~,'638,6 273.0 2353.0 473.8 572.7 12207.6 59682.2 29530.3
significant
X3
16.7
1340.2
m.s.s,
Brassica campestrisL.
of
crosses
X2
1360.4 416.0 2 3 1 . 7 ~'~ 44.9"x 26653.1
1 492
of Hybrids
of Squares
2480.7
4 4 4 1 1
Hybrids
Sum
X1
14
Evaluation
the
while for
inthe
showed the
all the
line,
there
for any character, significant for all interaction
was
A. A m i r t h a
e t a l . : A Q u a n t i t a t i v e E v a l u a t i o n of H y b r i d s o f Brassica campes~ris L.
Devarathinam
Table 2. A n a l y s i s of c o m b i n i n g ability in i n t e r - v a r i e t a l BS • YS
YS • BS
BS x TIR
TR • BS
YS • TR
TR x YS
X1
a b c d e
3223. I 295.6 670.5 54.5 142.8
629.2 3632.6 321.6 90.5 55.6
2084.5 579.9 291.8 52.0 48.2
207. I r 430.8 262.4 2.8 41.1
561.4 407.3 468.5 0.8 92.4
734.4 366.7 583.8 + 121.2
X2
a b c d e
60.3 41.6 9.2 2.1 1.4
X3
a b c d e
X4
a b c d e
a= d = are for
significant ciprocal
I0. I r 21.5 6.7 0.7 +
5.9 ~' 11.5 4 . 6 ~'" 0.2 0.3
475.2 146.0 117.9 9.6 24.1
79.7 11,1 65.8 + 19.7
4 5 . 0 ~64.5 1 6 . 0 "~'~ 2.0 +
21.1 r 2 2 . 1 "'~ 46.3 + 6.2
1971.9 288.1 278.9 42.6 48.2
401.8 220.1 1567.0 36.9 6.1
for XI
and X3.
There
combinations:
There
When
was
used
and line x tester characters
for X2
where
TR
differences XI
only.
were
X4
were was
were
were
(Table
differences
for X2.
Significant
dif-
amonglines for all
2). The tester
not significant.
The line • tester
(TR)
significant line
differences were
reciprocal
for
also
Combining
tion o f p a r e n t s ,
Ability:
An
of the parents
not
differences
overall
G B S II, K a n p u r L o t n i 17, K a n p u r L o t n i 12 i n Y S
and
T 1 6 5 , T 2 4 4 a n d T 1842 i n TIR, a s p o t e n t i a l p a r e n t s f o r inter-varietal
hybridisation
1 9 7 . 0 ~'" 35.7 r 178.4 + 23.1
II >( IB 6) a n d 2
•
w e r e f ound to be t h e b e s t , ning ability (g.c.a.) high specific
having high general
for the parents
combining
consistently
crosses
1•
GBS
( K a n p u r L o t n i 17 • IB 3) i n B S •
ability
combi-
2 and 8 and also
(s.c.a.).
showed high s.c.a,
These two for the yield
but did not p r o v e e q u a l l y good in t h e r e -
YS • BS c o m b i n a t i o n . 6) a n d 3 x 8
2 x 8 (Kanpur
Lotni
( K a n p u r Lotni 27 • IB 6) w e r e t w o with superior cross
g . c . a. f o r all t h e
characters.
The r e c i p r o c a l
8 • 3 was as good
as the cross
3 • 8, i n d i c a t i n g p r a c t i c a l l y
no m a t e r n a l
effects. In the BS
xTR
combination,
5 x 12 and 5 • 13, which ents,
the
involved
showing
yield attributes.
superior BS
g.c.a.,
were
of the TR
parents
to produce
In t h e Y S ing cross,
•
g.c.a,
parents
superior
crosses
the two
4 and 5, with 12 and 13 of TR,
reliable,
SC
were
found
ands.c.a,
2 and 3, which
found
to combine good
combination,
4 • 13, BS
parto be
for some exhibited
well with each
hybrids.
8 • 11wasanoutstand-
showing superior s.c.a, for XI, X 2 and
X3, c o m b i n e d with high g.c.a, for the parent 11 only. This cross, with its reciprocal II • 8, w a s found to be a good specific cross combination. Heterosis and its relation to combining ability: A
( T a b l e 3 ) . T a k i n g all f o u r
into c o n s i d e r a t i o n ,
yield components
components,
39.6 r 2 3 . 1 ~'75.9 + 13.7
1 8 9 . 5 ~'" 282.9 120.1 r 5.8 8.5
other reliable crosses
assessment
led to the identifica-
27 a n d DS 17D i n B S , IB 5, IB 6 a n d E P
ciprocal
dif-
In the recipro-
for all the four characters.
of the performance
crosses
20.6 2 . 2 ~'" 3 . 5 ''~ 0.4 0.1
148.0 52.8 73.1 1.4 12.8
67.0 r 228.1 4 4 . 0 ~'" 5.2 +
17 x l B
hybrids.
significant
and tester
General
the
re-
X2.
significant
among
and X2
observed
except
the line, there were
for XI
significant
significant
as line, the differences interaction
except
ferences
were
for all characters
for all the characters
YS
708.1 263.4 183.5 15.1 24.4
18.7 6 . 9 ~'" 8.8 0.2 0.4
line m.s. (4 d.f.); b = tester m.s. (4 d.f.); c = line• m.s. (16 d.f.); Var(g.c.a.); e = Var(s.c.a.); ''~ = not significant; + = not estimable; all m.s. significant at 5 % level; d,e approximate estimates only (see Arunachalam, 1974 details )
differences
YS-TR
15.2 58.6 4.9 ~ 1,6 0.4
ferences
cal,
crosses
study of the crosses
showing heterosis over better
parent for X2, X 3 and X 4 revealed that 13, 12 and 9 each f r o m B S - Y S ,
BS-TR
and YS-TIR combinations
w e r e heterotic (Table 4). Surprisingly only one cross, I •
s h o w e d s.c.a, effects for all the three charac-
ters, and only three did so for two
crosses,
characters.
2•
7 • 12 andl0•
18 out of the 34 heterotic
A.
Amirtha
Table
Devarathinam
3. Means
et al. : A Quantitative
and g.c.a,
effects in inter-varietal
1
m glf X1
X2
X3
X4
2
120.1 4.0
1.5 3.2 0.9
- 2 2 . 6 '" - 1 6 . 6 '`. - 7.2'"
6.4 0.5 1.4'`. 0.2
-
0.3
- 0.6
m glf glm g2f
8.9 0.3 1.4'`. 0.7
10.2 2 . 3 .> 0.8'`. 1.1':"
g2m
0.6
0.2
m glf glm g2f g2m
20.7 0.8 1.2 1.4 1.9'`.
21.1 4.8'" 0.0 1.9"`. 0.6
m gif glm g2f
44.7 1.8
-
-
g2m
-
-
-
83.1 -20.0""
12.6 4 . 3 ''~ 1.7 1.3 0.3
53.1 1 0 . 7 '`. 5 . 9 ~`. 3.1 - 1.1
38.7
7.3 2.4" 2.2'`. 0.5
-
15.6 5.7'`. 2 . 3 ~`. 0.2 - 0.4 -
5 . 9 ~'" 2.5
0.4
- 3.6
crosses X2,
X3
did not show or X4.
fects were
It was
-
in general
It was
found
intermediate
was
1.7 6 . 5 "~" 0.4
7.2 0 . 8 ~''" 1.4'`. 0.1
9.5 0.6 0.1 1.5'`.
1.2 ~`.
0.3
ent with regard
andX4
in which
parent
(Table
5.4'`.
could
found
13 crosses
out of 34 showed 9 over
showing
between
60 to 70.
for X3
(Table
parent),
9 to YS-TR
combinations. and X4
pectively.
Using
cy of crosses
qual to or above will be named ally for X2, (= 41%)
were
X3
The
these
values
showed
the norms
for X3
hybrids
mean
as norms,
was
found
res-
the frequenof heterosis
worked
of which
and
per cent
and 34,
for 2 or 3 characters
It was
hybrids,
(over su-
heterosis
an amount
5.6 3.2"" 1.6 3 . 5 .> 1.4
-
47.2 2.5 0.8 3.5
-
'`.
0.2
g.c.a,
0.2
effect
combinations
(Table
5). BS-YS
rank
X2,
which X3
also contributed
were
rank
andX4.
11 • 8 were
hybrid
and the rest by BS-TR
hybrids
four of
for all the
The cross8x11andits
both rank
3 • 12 belonging
hybrids
s. c. a. effect for any of the three
re-
in YS-TR.
to BS-TR
did not
The have
characters.
Discussion For
the first time,
in this study,
were
chosen
diversity
attempt
(which
that 14 out of 34 con-
to represent
was
and the primary
interest
ance
of inter-varietal
ter-varietal
hybrids
among
described form
lower
a diallel system,
entries, the BS,
but the system YS
and TR
ents in a series
Since
parents
for populathe intra-var-
than the inter-varietal
was
to evaluate
hybrids,
The
made,
of genetic
BS-TR
a set of possible
and YS-TR
set of crosses
populations
of line x tester
in-
distributed was
made,
as
did not exactly
due to the absence adopted
one
the perform-
and their reciprocals
BS-YS,
earlier.
whose
degree
important
selection.
been
of combining
crosses
a good
for yield components and human
has
the components
ietal divergence
equally e-
out individu-
50 % were
a serious
to analyse
ability in a set of inter-varietal and
was ob-
12 to BS-TR
to be 28,49
Hybrids)
and X4. rank
3
mostofthecros-
to BS-YS,
found
Rank
20to40 cent,
highest heterosis
inter-varietal
was
which
and YS-TR
tion fitness
13 belonged
for X2,
X3
The
X3
4).
Of the 34 heterotic perior
per
50 per cent heterosis
20 to 50 per cent for X4. served
Similarly,
40
0.0
of heter-
for X2
less than
by BS-YS
ciprocal
better
for X2,
heterosis
and
were
were
a range
be
10.7 1.0~' 0.2 0.6
-
inferior par-
percent
ses showed
4).
thanthe
However,
parent
crosses
with the parents.
the hybrids
or worse
to height.
superior
tributed
characters
compared
38.8 2.2 1.1 4.3
-
3.0 3.7 1.5
3.9 2 . 6 ~`. 1.5 3.8'`. 0.8
46.8 3.1 1.6 1.8
the eight hybrids
4).
when
noted
ef-
inter-varietal
that the inter-varietal
than the superior
osis over
(Table
in height
instance
that the s.c.a,
in heterotic
123.4 1.4
2 : BS-TB
effects for any character,
thus apparent
not pronounced
crosses
No
s. c.a.
107.5 3.3
16.1 4.2'`. 1.8 1 . 9 "~`. 0.5
-
7
5.2'`. 2.9 2.2
g I = BS-YS;
Brassica campestrisL.
6
129.7 0.0
24.8 -16.3'`. - 1 5 . 4 "~ - 1 0 . 6 '`.
0 . 8
of
5
122.0 0.8
13.5" 5 . 0 ~`. 5.2"
3.1 1.2
BS 4
122.1 15.2 '`.
2.5 5 . 5 "`. 0.8
of Hybrids
hybrids
3
glm g2f g2m
2.0
Evaluation
enabled as male designs.
of diagonal us to evaluate
andfemalepar-
A.
Amirtha
Devarathinam
et al. : A Quantitative
Evaluation
of Brassica
of Hybrids
YS 8 107.1 3.5 4.4
9
10
11
118.9 9.9 r 5.2 4.8'`.
130.6 1.7 0.6 5.1~"
75.9 0.2 2.6
2.6
-
2.8
-
4.7'`.
7.5 0.3 2.4'`. 0.3 0.0
-
10.1 0.9"`. 1.4 0.5 0.3
-
-
8.3 0.8 1.8 1.4
-
-
1.8 2.0'" 2 . 4 '~ 1.1
-
0.6
-
1.7
-
r
-
38.1 3.2 5.1'`.
-
2.4
-
1 = BS-YS;
The
-
0.7
0.4
-
2.1
• YS,
were
high for X2 in YS
crop
was
plants
both additive
(see
for
example,
1970,
YS
proved
As
used
gene
ac-
1973,
in
on YS;
parents suitable
Zu-
et al. 1973,
on Sorgh~).
of the combining
on
Hence
ability ef-
would
be
breeding
The
YS
a pre-
procedures
to be a success
as a female
a large
of promisinghy-
should
divergent
especially
crop, show
when
YS
used
chosen
should
in the cross
as females.
also recorded or
par-
also comparatively low within-line
divergence
to find that BS
as lines against
number
genotypes
their source
= =
as
BS-TR;
2
of the BS
It was success
estimates
importance
TIR as testers.
cross
33.2 3.0 4.0'`.
0.6
0.7
3 . 7 ''~
combinations
during
of maize
I, 2 and 3 could
could
types
bine well with TR
of gene
as testers
duce
with
hybrids was
limited
highly cross-pollinated,
a result.
population
was
der to have
stressed
for example)
other
example.
The
associated
study
before
high
for
of intra-
for the genetic effects crosses
(1966).
in inter-varietal cross-pollinated,
the parents making
was
with
be responsible
and Cockerham
were
pro-
parent.
parent
of inter-population
to improve
hybrids.
could
other
a closer
success
both the parents
be necessary tion,
FI
by Stuber greater
for
in BS-YS
good
as the
to account
in the
This ap-
foundtocom-
of BS
the
might
for
action.
5 were
usually
populations The need
variability
have
where
TR,
variability
open-pollinated such
YS
when
be-
relative
in that only parents
and produce
populations
heterotic
markedly the
as testers
4 and
In fact, the SI parental
The success
vary and
of BS
with YS
parents
that var-
as lines and that the
variations
combine
while
on open-pol-
and their hybrids
variance
very
components.
a study
in their potential
frequency
combination,
showed
for the yield
observed
of different
combi-
In fact,
1.0
2.6
to be true in the case
and variances
in-
11.5 1.3 1.3 1.3
32.4 0.1 3.4
-
of genetic
of gene
peared
5`.
ability
(1964)
vari-
when
14.0 0.6 1.6 0.4
-
• YS
ieties differed
cause
-
'~"
= YS-TIR
combining
Sprague
5.4 0.5 1.4 0.1 0.7
-
male
intra-population
a self pollinated
late flowering,
nation,
ac-
components
Giriraj 1965,
and TR
ent and has produced
teresting
gene
varieties/hybrids.
The SC YS
m
linated varieties
combinations,
additive
Singh
on TR;
for formulating
While
Brassicacampestris;
on
understanding
fects of the BS,
varietal
all yield
and Webster
reflected
6.3
variance
and non-additive
1973,
Kambal
ability.
- 5.2 '"
1.1
high for all thechar-
over
for almost
beri andAhmad
brids.
2.1
and
-
6.3 0.5 0.4 1.4'`. 0.1
3.3
-
2.1""
1.0
female
4.1'`.
0.0
0.7 as
0.3
-
-
32.6 0.8 4 . 0 '~
high
In the other
14.6 0.6 1.0 0.2
-
40.5 1.8 2.9
f =
of genetic
5.3 0.4 0.2 0.9" 0.4
1.0
some
they were
castor;
to evolve
"~"
s.c.a.
predominant
and Eunus
requisite
level
11.4 1.0 3.0" 1.0
80.3 4.6'`. 8.7'"
-
7.2"
39.8 0.5 0.5 -
of g.c.a,
components
x BS.
Usually
a detailed
~
9.5'"
-
15
69.1 0.1 - 1.7
2.4
-
5
14
60.3 4.5','4 . 5 '~-
6.9 0.7 1.4" 1.1"" 0.1
both additive andnon-ad-
estimates
only,
tion are present
Joarder
5
73.1 0.3 3.3 1.8 1.7
I
2) revealed
additive
non-additive tion.
at
'~"
2.0
action for all the yield components.
in BS
acters
-
-
2 = YS-TR
(Table
ditive gene
0.0
-
approximate
variances
0.1
11.6 0.9 - 1.1 1.7
0.8
-
4.7 0.3 0.5 0.4 0.9
17.3 0.7 3.7'`.
35.1 1.7 1.9
r = significant
6.7""
10.6 1 . 0 "~`. 0.8':" 1.2'" 0.6
-
42.4 4 . 5 ':" 5 . 1 ~`.
1.7
0.2
6.4"
TR 13
12
-
cg~pestris L.
the
(through hybrids.
In or-
crosses, it would selecThis
A. Amirtha T a b l e 4. M e a n s ,
Devarathinam
heterosis
Cross
and s.c.a,
BS-YS
1 x 8 2 • 6 2 x 7 2 X 8 3 X 8 3 • 9 6 X 2 6 • 3 6 • 5 8 • 2 8 • 3 8 X 4 8 X 5 3 xll 3 X 12 3 X 13 3>(14 3 • 15 4 • 4 x 13 4 • 14 4 • 15 5 • 12 5 • 13 11 • 4 7 • 8 • 8 • 12 8 • 14 8• 10X14 11 • 8 14 X 8 14 X l 0
YS-BS
BS-TR
TR-BS YS-TR
TR-YS
increase
16.3 14.0 11.5 15.3 12.2 10.4 11.1 13.3 11.9 11.4 12.6 9.6 10.7 9.5 10.6 7.5 8.9 8.2 9.2 9.0 7.8 7.4 11.3 8.9 8.3 12.5 12.3 8.5 9.1 9,1 13.0 9.5 8.5 11.4
the base
dify his programme
single
binations, 2 •
possible
cross
with high g.c.a. the case
to locate
and 3 x 9, each
hand,
superior
heterosis
h
9.1"" 8 . 8 ~,," -2.1 -1.7 - 4 . 1 "*'9.9~ 4.7 ~ 3.5 -2.2 1.5 1.0 4 . 0 "~:-2.6 -1.4 2.9 0.3 -0.9 -0.8 0.6 0.9 -1.2 -0.5 -0.5 -3.0 1.5 6.4~' 3.9 -3.3 -1.5 3.5 5.1~ 2.5 -0.3 3,0
of hy-
served
for sev-
yield
five crosses, of which
superior
in those
the crosses for three
2 x6
yield
68 58 25
parent;
-
Sim-
even ob-
-
4.7
27
-
1.4 7.4 1.1 5.7
57 40 24 79
-
9.7,',"
7
5.2 -
32 49 23 34 28 40 20 33 39 4 24 10 30 31 34 5 27 47 28
7 . 5 ~
3.0 2.2 1.3 1.8
-
-
9.4
~
3.0 5.8 0.2 2.3 0.4 8 . 4 "~ 7.2 7.6 ~ 0.7 5.8 4.8 1.2
-
-
3.2
32
2.5 0.6 8 . 9 "' 4.7 6.3
26 71
42 17 47
~- = significant
two crosses
was,
at
therefore,
of the high g. c.a.
due
to
of the parents
(of
2 and 8, in particular). The
results
with the reciprocal
did not fall in line with those Two
crosses,
X3
and X4;
as observed 8 as female found
against
other
of the yield components, of parent
The heterosis sistently
to show
each
were
marked
as high as in BS-YS
heter-
involving
of the BS
out the
programme. was
combinations
not con-
for the ma-
the crosses
4 x 12 and 4 x 15, which
would
deserve
as
for one or
bringing
combinations
the
parents
heterosis thus
he-
point
in manifesting crosses
Nevertheless,
3 x 13, 3 x 14,
bited high heterosis,
perhaps
8 in the hybridisalion
in BS-TR
jor yield components. 3 x 12,
substantial
this would
The
male
superiority
combinations.
6 x 3 and 8 x 3, showed
for X2,
parent
YS-BScombinations
of BS-YS
earlier.
and 3 x 9. On
heterosis
12.1 x~ 15.9 ~ 1.2
of the parents
combined
components
75.2 83.8 66.5 67.3 60.7 59.2 65.6 69.6 49.9 70.2 57 5 46 7 62 9 51 1 56 6 46 5 51 5 53 7 41 9 40.4 35.7 43.1 61.3 62.7 41.7 59.9 58.3 51.9 50.3 48.0 59.9 56.6 44.5 51.7
osis,
2 x 8 and 3 x 8 had shown
of high s. c. a. The high
h
to the superiority
in perform-
s.c.a,
s
I •
Of
8, in particular).
superior
m
com-
expressed
components.
48 59 9 14 68 145 20 105 25 2 79 39 19 39 105 45 25 42 44 33 3 10 43 15 21 163 72 109 27 78 51 76 16 17
the reinforcement
also mo-
: In BS-YS
1 • 8 was outstanding showed
s
per cent over
than modifying
crosses
in the crosses
the other
in the absence
or
should
rather
(of the parent
crosses X4
terosis
for all major
five crosses,
ilar was
30.7 33.6 22.9 24.0 21.1 30.9 25.3 25.8 20.1 21.5 22.6 21.6 19.1 17.5 25.8 21.2 17.5 17.9 22.4 20.7 16.1 17.1 23.0 18.5 18.8 30.0 19.9 15.7 18.6 20.5 26.2 20.4 16.2 20.2
separately.
it was
This
60 37 8 50 63 3 9 40 25 12 68 28 43 48 54 17 4 28 26 23 7 1 57 24 14 17 64 23 21 21 23 27 13 8
of the hy-
improvement
in inter-varietal
2 • 8, 3 x8
ance.
m
sampling
inbreeding
breeder
simultaneously
high heterosis those
The to achieve
attributes
Heterosis
performance
in an orthodox
programme.
eral attributes
-1.1 -0.5 0.3 2 . 4 ~'" 1.4 -0.6 1 . 7 ~,'~
extent than the repeated
population
bridisation
3.3 r 1.3 -0.9 0.3 -0.9 1.1 -0.7 0.9 -0,5 -0.1 0.2 1.1 -1.4 0.1 0.3 -1.3 0.3 0.6 -0.4 1.0 -0.1 0.5 1.3 0.5 0.4 2 . 6 ~" 1 . 9 -'>
h
effect:, h = heterosis t-test only)
the expected
brid to a greater
inter-varietal
X3 s
m : mean; s = s.c.a, 5 % level (approximate
should
e f f e c t s of h e t e r o t i c
X2 m
Brassica campestris L.
et a l . : A Q u a n t i t a t i v e E v a l u a t i o n of H y b r i d s of
consideration.
3 X 11, exhiIt
A.
Amirtha
Devarathinam
et al. : A Quantitative
Table 5. Distribution of heterotic ses for major yield components Per Cent Heterosis
inter-varietal
Number
cros-
12 13 6 3
X4
10 7 7 5 3 1 1
-
6 19 6 3 -
-
coner
(1964)
needs
modifying
of using
a
b
Total
n
p
n
p
n
p
Total
7 4 3
50 29 21
6 8 6
30 40 30
13 12 9
14
41
20
59
34
and TR
in a marked
38 35 27
because
in the way
in vigour,
be
several
should
such
effects
sacrifice
1967).
and
in
The
crosses
dif-
effected
of bulking
plants). very
the cros-
The heterosis
greatly
in Brassica
as observed
the harmful
set with some
as also observed
(instead
not be
result
if a representative
are identified
from
hybrids
found
would
(Thomas
overcome
plants only
hy-
was
of the rigid sys-
Inbreeding
for example
partly
those
Thus,
stand
for inter-varietal
populations,
and female
general,
therefore
application.
could
parents
decrease
ficulty can
sed seeds
7
A high level of heterozygosity
Loli~m perenne,
such BS-YS BS-TR YS-TR
difficulties
tern of self-incompatibility.
male
L.
diallelic model,
to be of general
highly inbred
bridisation. in the BS
cmmpest~s
on a one-locus
practical
between
Combination
of Brc~sica
of Hybrids
Some
of Crosses X3
X2
1- 20 2 1 - 40 41- 60 6 1 - 80 81-100 101-120 121-160 161 a n d a b o v e
Evaluation
of
affected
in
o leracea (Watts 1970).
of inbreeding
could
of homogeneity
be off-
in the FI
hy-
brids. A number Crosses
heterotic
for X2,
BS-YS YS-BS BS-TR YS-TR
lX 8 • 3 • 8 •
X3
andX4
8 ~H~', 2 X 6 "~, 3 X 3 12 + 1 1 , 11 • 8
8, 3 X 9
hybridisation
brid progenies (whatever double
as males 1963,
(Welsh
superior
combining
Similar were
results
also found
reciprocal
was
7 x 12 which
parents
which
ability produced to those
and achieve
for YS-TR.
The cross
produced
lines could
8 x 11 and its
neously
Another
heterosis
was
of 163 %
for X3. Heterosis ence
in the gene
character ents
(Falconer
ability.
result found
1964).
to show
non-additive
heterosis components
tion of the expression
hybrids
inthe par-
combining
to be true.
A even
number
of crosses
in the
of gene of heterosis,
absence
action.
The high did not was of high
The implica-
as based
by Fal-
hy-
through ( Ram, Doloiand high andlow
forward
simulta-
would
Comstock
Brassica campestr~s,
sed of as many possible.
of open be
and Ro1957,
good
combining indicates
be built using
pollination
possible
synthetic
This study can even
of
and shuffle the de(Comstock
and Robinson
it should through
effects
and
1963).
of the high degree
yield improvement
of the inter-
the harmful
disequilibrium
Gates,
inter-mated, (c) Bi-parental
generations
obviate
and Cockerham
Because
stage,
in one recombinant
1952;
Matzinger
complex
ex-
where
populations.
and advanced
genes
9 • 12 and 10 x 3 for X1
in high heterosis.
in F2
and linkage
not always
in the crosses
matings
binson
would
Arunachalam,
selection,
and at a later
sirable
to
time
yield improvement
and carried
productive
linkage
with high specific
This appeared
provide
a quantitative
customary
and Dudley
Programmeslike
in the inter-varietal
Murty, way
be selected
controlling of dominance
varieties
on both the right and left tails of the phenoty-
of the differ-
It is, therefore,
high heterosis
1969;
of the square
frequencies
hybrids
in a set of yield components
(b) Two
pic distribution, would
hy-
be - single,
Levings
1966).
for flowering
further
varietal
and also the amount
associate
s.c.a,
is a function
and Doloi 1972).
1973;
variation
response
combinations
combination.
the maximum
Murty Ram
high heterosis.
for BS-TR
an excellent
had shown
selection
may
or high yielding
and Atkins
would
the inter-varietal
hybrids
and Robinson
(a) disruptive
be triedin
and either superior
crosses)
ploit the intra-level
can
One such system
using
as females
and Moll
correlated that only those
systems
the level of these
or multiple
brids
observed
mating
a bid to boost the performance. be multiple
a = crosses showing high heterosis for 2 or 3 characters out of X2, X3, X4 b ..... do ............... one or none .... do .... n = number; p = per cent; + = s.c.a, effects absent; ',,~= s.c.a, effect for X3, X4; ~,~r = s.c.a, effects for all characters, other crosses had s. c. a. effect for one character only.
was
of useful
BS,
to effect
varieties
parental that
such YS
in
compo-
genotypes
as
a synthetic and TR,
es-
8
A. Amirtha
pecially in view
Devarathinam
of the complex
et al. : A Quantitative
level of cross
compa-
tibility found in them. This study on inter-varietal cates that the differences mainly ferences
in flowering
sion of characters. we considered show
direction
inter-varietal
capable
clearly
shown
by the range
population
improvement
were
found
crosses.
and nature
progenies.
of brown
joint
Nevertheless,
by inter-varietal
to the background
the
of the s. c. a. of the
of the varieties
that breeders
ford to relegate
if
could
cases,
out the desirable
iability in the F2 and biparental thus conclude
hybrids
g. c. a. effects a-
of the parents.
of recombination
This was
on pure
cancelled
attributes
gain ground
could not. In some
and magnitude
cross
effects of the g.c.a, the desirable
ideas would
No cryptic
in the expres-
inter-varietal
based
others
indi-
and dif-
of the varieties.
These
good heterosis
system
could be detected
that some
lone, while some opposing
pollinating
time
differences
clearly
in the three varieties were
due to the imposed
structural
crosses
One
sarson
of varcan
can ill af-
the potentialities of
by inter-varietal
hybridisa-
tion.
the use of a moJ. Genet. 3_~4,
280-287 (1974) C o m s t o c k , R . E . , R o b i n s o n , H . F . : E s t i m a t e s of a v e r a g e d o m i n a n c e o f g e n e s . In: H e t e r o s i s , 494-516. A m e s : Iowa State College P r e s s 1952 F a l c o n e r , D.S. : I n t r o d u c t i o n to Q u a n t i t a t i v e G e n e t i c s . E d i n b u r g h and London: O l i v e r and Boyd 1964 Gates, C . E . ; C o m s t o c k , R . E . ~ R o b i n s o n , H . F . : G e n e r a l i s e d g e n e t i c v a r i a n c e and c o v a r i a n c e f o r m u l a e for s e l f - f e r t i l i s e d c r o p s a s s u m i n g l i n k a g e . G e n e t i c s 42, 749-763 (1957) G i r i r a j , K . , M e n s i n k a i , S . W . , Sindagi, S . S . : C o m bining ability of s o m e q u a n t i t a t i v e c h a r a c t e r s in 6 • 6 diallel cross of castor (Ricinus communis L. ). Indian J. Agric. Sci. 43, 319-322 (1973) Joarder, O.I., Eunus, A.M.: Genetic studies of two crosses of Brassica cat,pest,s. Exptl. Agric. 6,
Received December 26, 1975 Communicated by H. Stubbe
of Hybrids
of Brassica campestris L.
351-357 (1970) Kambal, A.E.; Webster, O.J.: Estimates of general and specific combining ability in grain sorghum, Sorghum vulg~e Pers. Crop Sci. 5, 521-523 (1965) Kempthorne, O. : An introduction to Genetic Statistics. New York: John Wiley and Sons 1957 Levings Ill, C.S. ; Dudley, J.W. : Evaluation of certain mating designs for estimation of genetic variance in auto-tetraploid alfalfa. Crop Sci. 3, 532-
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Dr. A. Amirtha Devarathinam Deputy Agricultural Officer Seed Testing Laboratory Thirunagar Madural 625006 (India) Dr. V. Arunachalam Division of Genetics Dr. IB.R. Murty Project Director Nuclear Research Laboratory Indian Agricultural Research New Delhi 110012 (India)
Institute
