Theoretical and Applied Genetics 46, 401-406 (1975) 9 by Springer-Verlag

1975

Breeding Implications from a Diallel Analysis in two Environments of Yield and Component Characters in Upland Cotton (Gossypiurn hirsuturn) H. E. Gridley Cotton

Research

Corporation,

Namulonge

Research

Station,

Kampala

(Uganda)

Summary. In a full diallel set of crosses involving nine Albar 51 lines the genetic control of yield and yield components were largely accounted for by additive effects, although non-additive effects were evident. Significant differences in correlation coefficients between sites and the interaction of genetic effects indicate the effect of the environment on development and gene action. The failure to achieve yield improvements appears to stem from a combination of testing at the early selection stage in a single environment and of a narrow genetic base. The implications for future breeding programmes are discussed.

I. Introduction BPA

is the present

commercial

land cotton in southern Church,

1968).

racters,

Uganda

Pedigree

that differ from

variety of African (Arnold,

selection

the first BPA

but no line has been obtained attempts

other characteristics

of successive

therefore,

issues

(Arnold,

Gridley

BPA

stocks

selected

equatorial to assess

lines.

regions

Innes,

genetic variability

within

1973).

made

on the genetic

characters,

in 1969

Environmental

of Africa

diallel set were

between

are extremely

parents

and

con-

a full dialnine pedi-

conditions

the effect of the environment

the genetic parameters,

Buganda,

issues

19711

trol of yield and its component

gree

1970,

and obtain information

was

seed

on the use of multiline seed

to increase

le set of crosses

cha-

the yield and

BPA

Innes and Gridley

and Busuulwa,

and lines

that is superior

to improve

been based

In an attempt

Costelloe

has produced

seed issue in some

to it in yield. Recent

have,

Up-

in the

variable,

on estimates F I hybrids

one at Mubuku,

Analysis

In the Hayman (1954) analysis, variation is divided into components which measure additive and non-additive genetic effects and reciprocal differences. Item 'a' refers to additive parental effects and "o' to non-additive effects. Where an additive-dominance model is adequate the latter can be partitioned into terms providing tests of mean dominance (b• of variation of dominance from array to array (b2) and of dominance deviation that is unique to each Fi (b3). If tests of the value of Wr (covariance) and Vr(variance) do not show the expected relationship, then non-additive effects can not be interpreted in terms of simple dominance. Item 'c' deA 474

b

5eoson

5 5eneroti0ns

1957

A (57)5 I

I

of the

Western

Region. 2. Materials

Genetic

and of

included in a rain fed trial at Namulonge,

and in an irrigated

Mubuku with two plants per stand in two row plots at a spacing of 0.9m X0.3m. The yield components, seed cotton per boll, loculi per boll, seeds per loculus, seeds per boll, seedweight and lint per seed were determined from fifteen boll samples at Namulonge and ten boll samples at Mubuku taken at the time of second pick. Ginning percentage and lint yield were estimated from the produce of whole plots.

and Methods

The nine parents used in the crossing programme, which included reciprocals, were the pedigree selections BPA 68, A(66)22, 29, 36, 102, 131, 134 and A(67)8, 12 from Albar 51 ; all nine selections stemmed from A ( 57 ) 5 (Innes and Jones, 1972). The derivation of these lines is shown in Fig. 1 where A(57)5 is the sixth generation inbred from A474, with single plant selection in each generation. The parents were therefore highly inbred and closely related. In 1970-71 parents, hybrids and reciprocals were grown in a 9 x 9 balanced lattice square with five replicates at Namulonge and Mubuku. Plot size was 4.3 m 2 in theNamulonge trial and 3.2 m e in the one at

I

1958

A(58)14

1959

A(59)Z8

1960

A(60) 6

1951

I

I t

/b A (51) 6

1966

BPA 65

1967

BPA 67

A(66)2Z 29 35 102 131 134

I

1968 BPA 68 F i g . 1. P e d i g r e e o f A l b a r 51 s e l e c t i o n s f u l l d i a l t e l set o f c r o s s e s

A(57} 8

12

used in the 9 x 9

402

H.E.

Table

I. Parent and hybrid performance (N) and Mubuku (M)

Character Seed

per boll (g)

Breeding

Implications

for the eight yield components

Parental mean

Site

cotton

Gridley:

increase F~ mean over parental mean

FI mean

4.8 • 0.08 5.3 -+ 0.05

4.9 -+0.02 5.4 -+0.02

2.8 ~ 1.9

Loculi

per boll

N M

4.2 • 4.3 •

0.02 0.02

4.2 • 4.3 -+0.01

Seeds

per loculus

N M

6.8 7.9

0.08 0.04

6.8 7.9

Seeds

per boll

N M

Seed weight (mg/seed)

N M

Lint p e r s e e d ( r a g )

N M

Lint y i e l d ( k g / h a ) Ginning percentage

28.7 • 0.37 33.7 + 0.34 119 108

+ •

1.1 1.0

0.5 -0.3

-+0.02 •

0.3 0.4

29.0 -+0.12 33.7 •

0.9 0.1

119 109

a Diallel Analysis

at Namulonge

N M

• +

from

r 0.30 -+ 0 . 4 0

0.5 1.2

4.8 + 0.05 5.2 • 0.05

4.9 • 5.3 + 0.02

1.2 1.7

N M

651.0 • 12.17 531.0 • 14.00

710.6 • 4.50 560.4 • 5.74

N M

3 1 . 2 -+ 0 . 1 5 32.2 • 0.09

31.3 +0.05 32.2 -+0.05

9.1 ~ 5.5 0.2 0.3

~,H~ p < O. O01 ~,~ P < 0 . 0 1 ~ p < 0.05

tects average maternal effects and item 'd' the reciprocal differences not ascribable to c. If there are differences between reciprocal crosses, then the mean squares for items c and d become the appropriate error terms for the a and b items, respectively (Wearden, 1964). A joint Hayman (1954) analysis of the data from the two sites was also carried out. The presence of an interaction of genetic effects with sites was then testedby comparing the item x sites with the item x blocks mean square. Adjusted data from the lattice square analysis were used to calculate the phenotypic correlation coefficients but for the Hayman (1954) analysis rows and columns of the lattice square were ignored. All statistical analyses were carried out at Rothamsted Experimental Station on an I.C.L. System 490 Computer using Genstat.

ces in sample

size which

was

9 in the case

of parents.

The

ulations

at both sites of seed

per boll, seeds erally number

pressed above

I indicate

that significant

as the percentage the average

increase

of the parents,

heterosis,

ex-

or high.

contributes

The

more

hybrid

boll with loculi

per boll were

This suggests

weight

the latter are highly

show

gen-

that total seed

to boll size than seed

although

populations

per

seeds

but only

in the two pop-

population

a positive

corre-

at Mubuku.

effect of increas-

ing boll size on lint yield. Correlations indicated

hybrid

in Table

and

lated with boll size in the hybrid

seed Data

cotton

loculus

and lint per seed,

positive

Heterosis

per

significant

significantly

by a bracket

differences 3. Results

72 for the hybrids correlations

in Tables

in degree

relationship.

are seed

for the former

between

2a and

of relationship

to a negative populations

different

2b and range to a switch

Examples

cotton per

and seed

sites are

weight

from

from

from

a

the

boll and lint per

and ginning

percen-

tage for the latter.

of the F I hybrids is shown

only by seed

Diallel Analysis

cotton per boll and lint yield at Namulonge. Analyses model, Correlations Phenotypic components given

were correlation for parents

in Tables

2a and

coefficients and hybrids

among

2b. The greater

cally significant in the hybrid

the eight yield

at the two sites are

populations

number

statisti-

reflects differen-

terms.

of variance are

shown

calculated The

using

relevant

for interaction

of lint yield, using Hayman's in Table

components

variance

the associated mean

squares

were

shown

( 1954 ) (VR)

interaction

and variance

in Table

calculated

ratios

block

of additive or non-additive

sites for lint yield are other

3. The

ratios

effects between

4. Results

similarly

and

for the are in-

H . E . Gridley: Breeding

Implications f r o m a Diallel Analysis

403

Table 2a. P h e n o t y p i c c o r r e l a t i o n c o e f f i c i e n t s b e t w e e n the eight y i e l d c o m p o n e n t s f o r p a r e n t s at N a m u l o n g e (upper

value)

Character

and Mubuku

(lower

value)

S e e d cotton

Loculi p e r

Seeds per

Seeds per

Seed

per boll

boll

loeulus

boll

weight

Lint p e r seed

Lint

yield

0.56

Loculi per boll

0.75+~

0 962

Seeds per loculus

-0.07 0.53

0.19

Seeds per boll

Seed weight

0 985 ~9+~

0.47

0 9 53

0 . 8 6 +~+~"

0 944 0.35

Lint p e r seed

0.06

0.37

-0.23

0.32

-0.31 0.43

-0.60 O. 10

-0.24 -0.23

-0.29 -0.57

-0.41 -0.47

Seed

Lint p e r

Lint

weight

seed

yield

Lint per seed

0.30+~-

Lint yield

O. 2949 O. 4 8 4 ~

- 0 . 5 0 +~+9+91+9 -0.50 +9+~+91e -0.12 l -0.13 l 1

in a summarised

form,

effects were

for lint per were

and

-0.09

appropriate.

Non-

agreement

of the ~oint regression

suggests

that simple

planation

of the non-additivity

E. However,

statistical tests des-

(1971)

using

None

for individual

Wr

and

Vr

an additive-domiof the three charac(a) the consistenbetweenthe blocks

and

re~

(c) the

0 9 56 +9+~*~]

- 0 . 2 8 +9 ]e~9 - 0 . 2 5 ~ ]~9~*~ 0.47+9~9'9 0 39 +~+91 0 58~9+Hq 0.48+9~

]~9

seed and lint yield at both

over arrays , (b) agreement Vr

0 53 +9~+~]

mation

ters satisfied all three tests, involving

on

0.30 0.00

for seed cotton per boll at

inbred,

Jinks

at both sites

at Namulonge.

0 5049~+~1r

0.19 -0.20

-0.17

with lint yield, in Table 5.

to investigate whether

of Wr

0.08

highly significant

evident

sites. As the parents

O. Z5~ 0.05

0.24

except seed weight

additive effects were

-0.23 +9 -0.20

-0.08 -0.18

O. 19 ~0.0 2

]~ 0 . 3 3 +~+9 ]

gression

O. 69+9~4~ J

]~9~9 -0.33 0.69~+~ ] -0. I0

Ginning percentage -0 902

cy of Wr-Vr

at Namulonge

Seeds per

-0.0 2 ] +9+9+9 - 0 . 2 0 O. 74~+9+~] -0.05

was

for hybrids

boll

Seed weight

model

different with the level of signif-

the eight yield components

loculus

0.80+~+~91~9~9 0.58 ~+~9 0 48~+~+91 0.70 ~*~+~

nance

they are significantly

0.34 0.59

-0.35 0.64

0.57

Loculi p e r S e e d s p e r

Seeds per boll

were performed

-0.27

boll

0.71~9~+914~9 0.10 0.30 +~ J -0.02

by Mather

I

0.591

Seed cotton

Seeds p e r l o c u l u s

cribed

indicates

-0.50]~

per boll 0.43 +9+9 0.36 +9~

and

O. 74+~ O. 73 +~

-0.50

Loculi per boll

Namulonge

-0.65

0.32

2b. Phenotypic correlation coefficients between (upper value) and Mubuku (lower value)

Additive

-0.07

-0.86J -0.53

Table

for all characters

-0.13 ]~

O. 11

Bracketing of two correlation coefficients icance shown at the top right hand corner

cluded

-0.06 -0.52

-0.33

Note:

Character

-0.30 -0.7049

O. 18

O. 11

Ginning percentage

0 . 8 8 +~+~"

0.40 -0.17

0.09

Lint y i e l d

0.85 ~+~

dominance

of the components

sociated

is not an adequate

of variation

significant

ex-

and does not allowthe

the statistically

with

slope with unity. This

DI, HI, H2,

significant

heterosis

estiF and

b I item is as-

for two

out of thl-ee

components. Differences were

between

evident for seed

Namulonge. reciprocals dications

For

arrays

of reciprocal

both characters

wei-e small. that hybrids

crosses

weight and ginning percentage

For

differences

the former

involving

BPA

at

between

there were

in-

68 as the female

404

H.E.

Gridley:

Breeding

Implications

from

a Diallel Analysis

Table 3. Hayman analysis of variance for lint yield in the 9 • 9 full diallel set of crosses at Namulonge and Mubuku Site

Namulonge

Mub~u

Item

DF

MS

VR

MS

VR

a b

8 36 1 8 27 8 28 32 144 4 32 108 32 112

30451.20 8732.29 154814.00 4016.80 4719.04 4297.32 4201.41 3491.70 3046.55 1709.97 2303.27 3316.29 1961.61 3709.00

8.72*** 2.87*** 90.54*** 1.74 ns 1.42ns 2.19 ns 1.13 ns

44844.80 9670.71 18112.11 11725.20 8749.33 10729.99 5218.23 10439.30 7061.76 1123.09 9036.86 6696.50 6699.00 8701.30

4.30** 1.37 ns 16.13" 1.30 ns 1.31ns 1.60 ns 0.60 ns

bl b~ b3 c d a • blocks b x blocks b~ • b l o c k s b2 • b l o c k s bs • b l o c k s c x blocks d • blocks

ns non-significant

Table 4. Analysis

Item

C

d

Table

for lint yield in the 9 • 9 full diallel set of crosses

Item • Sites

b~ b= b3

Note:

of item • site interactions

DF

MS

8 36 1 8 27 8 28

8165.00 6723.65 33779.51 8010.63 5340.26 8864.50 3314.04

Item • Blocks

VB

0.78ns 0.95ns 23.85** 0.89ns 0.80ns 1.32ns 0.38ns

Namulonge

Mubuku

Pooled

DF

MS

MS

DF

MS

32 144 4 32 108 32 112

3491.70 3046.55 1709.97 2303.27 3316.29 1961.62 3709.00

10439.30 7061.76 1123.09 9036.86 6696.50 6699.00 8701.30

64 288 8 64 216 64 224

6965.50 5054.16 1416.53 5670.07 5006.40 4330.31 6205.15

The variance ratios for all items except be are based square as tests showed them to be heterogeneous

5. Summary of Hayman analysis at Namulonge and Mubuku

Site

of variance

conservatively

for eight yield characters

Namulonge

Mubuku

on the greater

mean

in a 9 • 9 full diallel set of crosses

Interaction

Character Seed cotton per boll Loculi per boll Seeds per loculus Seeds per boll Seed weight

a*** a*** a*** a***

b~

Lint per seed

a***

b~

Lint yield Ginning percentage

a***

a***

d*

d*

b***

a*** a** a** a***

c**

d**

a*** a***

c**

d***

a***

a***

b~**

d* a***

d***

a***

b~

b~ b~ c*** d***

H.E.

Gridley:

parent

were

Breeding

Implications

contributing,

marginally

heavier

all hybrids

involving

slightly lower

than their reciprocals,

A(66)36

between

and seeds

at Mubuku.

Differences

nents appeared

by c, were

involving

Interactions

were

of variance calculated

ferences,

effects except

be-

ments~

seeds

who

of heterogeneity

square

have been

ratio was

only. Parts

or

due to scale dif-

but only for seed cotton per boll and ginning

percentage accounted

was

a significant

proportion

for by a linear regression

rental or array same

may

means,

be caused

them.

variable

of Wr

- Vr

on pa-

site.

material

The present

as the cause

Arnold

(1971)

interplay care

which

must

or si-

one or both of

found genes

de-

blight in Tanzanian

cot-

in certain environ-

variation

after transfer

of genotype

of

but can not dis-

in his review,

data serve

masked

indicates

to bacterial

(1969),

in expression

of previously

had found unexpected

quoted

authors

in phenotypically

to a new

non-

environment.

further to illustrate the dynamic and environment

and emphasise

be taken in interpreting

the

results from

a

single environment.

of the variance

in neither case for both at the

Walker

the interplay

and be non-transmis-

The diallel analysis

resistance

in develop-

from

by a change

ton stocks that are only expressed

the two sites the variance

all of these interactions

purely

and environment

tinguish between

evidence

mean

stem

or by the expression

termining

those switching

reflect a change

the latter two alternatives

if either of the

for all characters

using the larger

genes

lent genes.

is important.

there was

between

of the genotype

of a number

of additive or non-additive

Where

for seed

This may

sible, or it may

in correlation

sites, particularly

pattern.

and for all compo-

it is doubtful

evident

between

ment

at

on the coordinated

but the differences

crosses,

and ginning percentage

small

coefficients

genes

positive to negative,

different parents.

purposes

sites were

per loeulus.

evident

action of many

from

and ginning percentage

per loculus

sets of differences

tween

The yield of a crop plant depends

while

parent had

of reciprocal

to be due to the behaviour

practical

above

as the female

arrays

cotton per boll, seed weight Namulonge

405

in that six out of eight had

seeds

other than that measured

For

a Diallel Analysis

ginning percentages.

Differences

of crosses

from

In a breeding tion stages,

programme,

owing

testing at the early selec-

to limited

seed supplies,

stricted to a single environment ricultural produce

practices.

Selection

lines adapted

is often re-

employing

advanced

for improved

yield

to these conditions

agmay

but lacking the

4. Discussion stability required An understanding

of the genetic control of agronomical-

ly desirable

characters

ment

genetic variability

where

lection.

The

control

largely

determined

sites although interaction seeds

in crop

is exploited

of the eight yield by genes

non-additive

of genetic

per loculus

between

is necessary

shows

Wimble

having

has

a similar

that parental pattern

shown

shaped

environments, lations.

in this material

in press).

only to large climatic season

to season

cultivated variation

(Arnold

of soils and management, tal combinations. that under between

Arnold

such conditions

genotypes

reach

hybrids optima

factor and re-

will vary

in changes

in different

in dominance

re-

cotton is subject not from

ample,

reselection

produced sampling

ley,

1970,

was

year the environment grown

use of the environment

(St. Pierre,

thy of further evaluation

place to place and

may

selection

characteristically variability

Klinck

and

and might

prove

Short term multiline

(1975)

have

striking interactions for yield.

shown

Tanzania

(Lee,

occur

ly such

mixtures

if

although

advances

from

a programme

inbreeding,

which

by a rapid decrease

is in

1971).

inprovements

might

comprising

over

(Arnold,

1967).

aid is worbeneficial

stem

from

a number

related lines. In district trials they have

in Uganda

environmen-

and intensive

seed issues

yield advantages

in many

stem

each

material

Gauthier,

as a practical

accompanied

(Arnold,

with wide

by alternating

the segregating

be limited as the parents

of pedigree

Innes andGrid-

of barleys

in which

was

ex-

of the diallel cross has

in Canada

Such

resulting

and environment

The selection

achieved

For

yield advantage in trials but fail to maintain this

district trials (Arnold,

1971).

adaptation

1971) but also to a wide range

and Innes

in the parents

lines which exhibit a few environments,

in wide ranging

cultivation.

applied to the hybrid populations, and

surface

Fl'S and parents

In East Africa

vary

in the analy-

that parents

response

resulting

The

except

differences

emerged

at different levels of one environmental lations between

is

effects are also evident.

and Gridley,

(1973)

se-

effect at both

effects for all characters

sites. A similar

Knight

through

components

of additive

sis of three fibre quality characters (Innes,

improve-

for commercial

the mean

and Jackson,

of genotypes

against the environment

shown

small

of their component

Innes and Gridley,

Walton

the use of

of closely

1970)

1973).

lines

and

in

Theoretical-

will be better buffered

than a single genotype.

Although

406

H.E. Gridley: Breeding Implications from a Diallel Analysis

c o n c l u s i v e e v i d e n c e is l a c k i n g , the l a r g e a m o u n t of a d d i t i v e g e n e t i c v a r i a t i o n d e t e c t e d s u g g e s t s the f e a s i b i l i t y of s u c h an a p p r o a c h w h e r e a s e e d i s s u e i s g r o w n c o m m e r c i a l l y for s e v e r a l y e a r s . The y i e l d a d v a n t a g e of the F 1 h y b r i d s at both s i t e s i s not s u f f i c i e n t f o r the e c o n o m i c p r o d u c t i o n of h y b r i d s e e d , a s c r o s s i n g has to be a c h i e v e d by t i m e c o n s u m i n g m e t h ods of e m a s c u l a t i o n and hand p o l l i n a t i o n . S e v e r a l g e n e s c a u s i n g m a l e s t e r i l i t y h a v e b e e n found ( W e a v e r and A s h l e y , 1971) but no s a t i s f a c t o r y s y s t e m of r e s t o r i n g f e r t i l i t y has yet b e e n d i s c o v e r e d . Long t e r m i m p r o v e m e n t s in y i e l d will h a v e to s t e m f r o m s e l e c t i o n in g e n e t i c a l l y w i d e - b a s e d p o p u l a t i o n s s y n t h e s i s e d f r o m the c r o s s i n g of e x o t i c m a t e r i a l to l o c a l l y adapted s t o c k s . T h e s e r e s u l t s , and t h o s e on lint quality (Innes et a l . , in p r e s s ) ,

e m p h a s i s e the n e e d to t e s t

e a r l y g e n e r a t i o n s e l e c t i o n s o v e r as wide a r a n g e of e c o l o g i c a l c o n d i t i o n s as is p r a c t i c a b l e . The s y n t h e s i s of a r a n d o m m a t i n g pool of the b e s t l i n e s in the e a r l y s t a g e s of a p r o g r a m m e

might p r e s e r v e g e n e t i c v a r i a b i l i t y to

which a b r e e d e r c o u l d l a t e r r e t u r n .

Acknowledgement I would l i k e to thank D r . N . L . Innes f o r help in the d e s i g n of the e x p e r i m e n t and d i s c u s s i o n of the r e s u l t s and D r . R . H . W i m b l e for a s s i s t a n c e in the s t a t i s t i c a l a n a l y sis.

Literature A r n o l d , M . H . : Cotton i m p r o v e m e n t in E a s t A f r i c a . In: C r o p I m p r o v e m e n t in E a s t A f r i c a , ed. L e a k e y , C . L . A .

R e c e i v e d A p r i l 8, 1975 C o m m u n i c a t e d by R. R i l e y

C o m m o n w e a l t h A g r i c u l t u r a l B u r e a u x Techn. C o m m . 1_~9, 178-208 (1971) Arnold, M.H.; Costelloe, B.E.; Church, J.M.F.: BPA and SATU, Uganda's two n e w cotton v a r i e t i e s . Cotton G r o w . R e v . 45, 162-174 (1968) A r n o l d , M . H . ; Innes, N . L . : P l a n t B r e e d i n g . In: A g r i c u l t u r a l R e s e a r c h f o r D e v e l o p m e n t . The N a m u l o n g e C o n t r i b u t i o n . London: Cotton R e s e a r c h C o r p o r a t i o n and C a m b r i d g e U n i v e r s i t y P r e s s 1975 (in p r e s s ) A r n o l d , M . H . ; Innes, N . L . ; G r i d l e y , H . E . : Cotton B r e e d i n g . P r o g r . R e p . E x p . S t a s . , Uganda, 196970. Cotton R e s . C o r p . , London, 12-34 (1970) A r n o l d , M . H . ; Innes, N . L . ; G r i d l e y , H . E . : Cotton B r e e d i n g . P r o g r . R e p . E x p . S t a s . , Uganda, 1970-71. Cotton R e s . C o r p . London, 17-34 (1971) H a y m a n , B . I . : The a n a l y s i s of v a r i a n c e of d i a l l e l t a b l e s . B i o m e t r i c s 1._.O0, 235-244 (1954) Innes, N . L . ; G r i d l e y , H . E . ; B u s u u l w a , L . N . : Cotton B r e e d i n g . P r o g r . R e p . E x p . S t a s . , Uganda, 1971-72. Cotton R e s . C o r p . London, 13-28 (1973) Innes, N . L . ; W i m b l e , R . H . ; G r i d l e y , H . E . : E s t i m a t e s of g e n e t i c p a r a m e t e r s for lint quality in upland c o t t o n . T h e o r . a n d A p p l . G e n . , 46, 249-256 (1975) Innes, N . L . ; J o n e s , G . B . : A l l e n ; a s o u r c e of s u c c e s s f u l A f r i c a n cotton v a r i e t i e s . Cotton G r o w . R e v . 4_..99, 201215 (1972) Knight, R. : The r e l a t i o n b e t w e e n h y b r i d v i g o u r and g e n o t y p e - e n v i r o n m e n t i n t e r a c t i o n s . T h e o r e t . and Appl. Gen. 4..~3, 311-318 (1973) Lee, B . J . S . ; Walton, I . C . ; J a c k s o n , A . C . : B r e e d i n g and A g r o n o m y . P r o g r . R e p . E x p . S t a s . , Tanzania, 1972-73. Cotton R e s . C o r p . , London, 10-35 (1973) M a t h e r , K. ; J i n k s , J . L . : B i o m e t r i c a l G e n e t i c s ; 2nd E d . London: C h a p m a n and H a l l , 1971 St. P i e r r e , C . A . ; K l i n c k , H . R . ; G a u t h i e r , F . M . : E a r ly g e n e r a t i o n s e l e c t i o n u n d e r d i f f e r e n t e n v i r o n m e n t s as it i n f l u e n c e s a d a p t a t i o n in b a r l e y . Can. J . P l a n t S c i . 47, 507-517 (1967) W a l k e r , J . T . : S e l e c t i o n and q u a n t i t a t i v e c h a r a c t e r s in f i e l d c r o p s . B i o l . R e v . 4..~4, 207-243 (1969) W e a r d e n , S. : A l t e r n a t i v e a n a l y s e s of the d i a l l e l c r o s s . H e r e d i t y 1__.99, 669-680 (1964) W e a v e r , J . B . ; A s h l e y , T. : A n a l y s i s of a d o m i n a n t g e n e f o r m a l e s t e r i l i t y in Upland c o t t o n , G o s s y p i ~ h i z ~'suSw~ L. C r o p S c i . 1._!1, 596-598 (1971)

H. E . G r i d l e y Cotton B r e e d e r I n s t i t u t e of A g r i c u l t u r a l R e s e a r c h Samaru P . M . B . 1044 Zaria (Nigeria)

Breeding implications from a diallel analysis in two environments of yield and component characters in upland cotton (Gossypium hirsutum).

In a full diallel set of crosses involving nine Albar 51 lines the genetic control of yield and yield components were largely accounted for by additiv...
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