A N E ST IM A T IO N O F G E N E T IC C O M P O N E N T S O F V A R IA N C E
IN B U L K E D F , P R O G E N I E S O F B A R L E Y
6

By
B a lw a n t S in g h J o g i

AN ABSTRACT
S u b m i t t e d to th e S c h o o l of G r a d u a t e S t u d i e s of M i c h i g a n
S ta te C o l l e g e of A g r i c u l t u r e a n d A p p l ie d S c i e n c e
in p a r t i a l f u l f i l l m e n t of th e r e q u i r e m e n t s
f o r th e d e g r e e of
D O C T O R O F P H IL O S O P H Y

D ep artm en t of F a r m

Y ear

A pproved

1954

C rops

1
BA.LWANT SINGH JOGI

An estim a te

ABSTRACT

of th e g e n e t i c c o m p o n e n t s

of v a r i a n c e in t h e F ^

g e n e r a t i o n w a s m a d e o n t w o b a r l e y c r o s s e s „ S t e w a r t x B a y , an d
K in d red x B a y .

Seed w as draw n fro m

t h e tw o c r o s s e s .

a b u l k e d F ^ at r a n d o m f r o m

S t e w a r t an d K in d r e d a r e

and B a y i s a s m o o th aw ned v a r ie ty .

rough aw ned v a r ie tie s

C r o s s S tew art x B ay w as d e s ­

i g n a t e d a s A. 2 7 2 a n d K i n d r e d x B a y w a s i d e n t i f i e d a s A. 2 6 7 .
T w en ty f a m ilie s fr o m
seed s from

e a c h f a m i l y w e r e t a k e n at r a n d o m .

w ere grow n for in c r e a se
s p r i n g of 1 9 5 3 , th e F
A.

e a c h c r o s s w e r e s e l e c t e d a n d 18 F „
4

g r o u p of F

5

D

The s e e d so tak en

in th e g r e e n h o u s e in f a l l ,

1952.

In th e

s e e d s w e r e p l a n t e d f o r i n c r e a s e in th e f i e l d .

ro w s tr a c in g to a d e sig n a te d F

3

row w a s c a lle d a

fa m ily .
T h e y i e l d t r i a l s w e r e c o n d u c t e d in t h e | | g e n e r a t i o n at tw o l o ­
c a t i o n s in I n g h a m an d T u s c o l a c o u n t i e s .

In a l l , t h e r e w e r e t h r e e

e x p e r i m e n t s , tw o in I n g h a m C o u n ty a n d o n e i n T u s c o l a .
d e sig n w as u sed .

A la ttic e

I n g h a m E x p e r i m e n t s I a n d II c o n s i s t e d of S t e w a r t

x B a y an d K i n d r e d x B a y c r o s s e s , r e s p e c t i v e l y .

T u sc o la E x p e r i­

m e n t III i n c l u d e d o n ly t h e K i n d r e d x B a y c r o s s .

Ingham E x p e r im e n ts

I a n d II c o n s i s t e d o f 100 p l o t s e a c h , a n d T u s c o l a E x p e r i m e n t III
c o n t a i n e d 90 p l o t s .

E n v iro n m en ta l v a r ia n c e w as d eterm in e d fr o m

2
BA.LWANT SINGH JOGI

p a ren ta l ch eck p lo ts.
b a sis.

ABSTRACT

The t e s t s w e r e on a fo u r -r o w , s in g le -p lo t

It w a s i m p o s s i b l e t o r e p l i c a t e s u b f a m i l i e s u s i n g t h i s p l o t

s i z e b e c a u s e of l i m i t e d s e e d .
w ere treated as

T he fo u r s u b fa m ilie s in e a c h fa m ily

r e p lic a te s.

N o t e s w e r e ta k en on d is e a s e
a cters.

The d is e a s e s

r e a c t i o n s an d a g r o n o m i c c h a r ­

c o n c e r n e d w e r e p o w d ery m ild e w

g r a m i n i s h o r d e i) a n d s p o t b l o t c h (H e l m i n t h o s p o r i u m
a g ro n o m ic c h a r a c te r s

(E r y s i p h e

sa tiv u m ) .

The

s t u d i e d w e r e d a te of h e a d i n g , h e ig h t of th e

p l a n t s , p e r c e n t o f l o d g e d p l a n t s , l o d g i n g r e s i s t a n c e , t e s t w e i g h t , and
aw n ty p e.

E f f o r t s w e r e m a d e to o b ta in a d d i t i o n a l i n f o r m a t i o n a b ou t

t h e s e c h a r a c t e r s to d e t e r m i n e th e e s t i m a t i o n of c o m p o n e n t s
ance.

The com p on en ts

of v a r i a n c e w e r e a l s o e s t i m a t e d f r o m th e

y i e l d d a t a , a n d th e e f f e c t s
w ere

c a lc u la te d .

of v a r i ­

of e a c h c h a r a c t e r o n s e e d y i e l d p e r p lo t

V a r io u s c l a s s

r a t i n g s w e r e g i v e n to th e e f f e c t s

of e a c h c h a r a c t e r a n d e a c h c l a s s w a s c o m p a r e d w it h t h e o t h e r f o r
sta tistic a lly

sig n ific a n t d iffe r e n c e s .

On a s in g le gen e b a s is

if t h e o r i g i n a l

seed chosen w ere

A A o r a a , t h e r e s u l t i n g p lo t m e a n w i l l r e m a i n u n c h a n g e d a f t e r a n y
n u m b e r of g e n e r a t io n s of s e lf in g .

O n th e o t h e r h a n d , if s e e d c h o s e n

w e r e A a , t h e n t h e p l o t m e a n w i l l c h a n g e w it h e a c h g e n e r a t i o n of

BALW ANT SINGH JOGI

ABSTRACT

se lfin g .

L e t th e m e a n

T h i s w a s i l l u s t r a t e d w it h a n a l g e b r a i c m o d e l .

e f f e c t s of A A , A a , a n d a a e q u a l d ' , h*
Aj

an d -d *

a

o n ly t h e a d d i t i v e a c t i o n of g e n e s a n d no e p i s t a s i s .

The m o d e l a s s u m e s
The v a lu e H

e q u a l s d o m i n a n c e d e v i a t i o n f r o m th e a d d i t i v e s c h e m e a n d m a y b e
p o sitiv e

or n e g a tiv e .

ta tio n w a s

In th e F ,

o

g e n e r a t i o n th e m e a n s q u a r e e x p e c -

c a l c u l a t e d b e t w e e n th e f a m i l i e s :

1/20 + is n H

< r£ ........ ,a>

an d m e a n s q u a r e e x p e c t a t i o n f o r w i t h i n th e f a m i l i e s w a s:

3 /80

D is

3 /4

+ W

il H

o

k

(b)

of t h e s i z e f o r w i t h i n t h e f a m i l i e s a s c o m p a r e d t o t h e s i z e

fo r b e tw e e n th e f a m i l i e s .

T h e c o e f f i c i e n t in b o t h t h e c a s e s i s e x ­

t r e m e l y lo w s o th a t it a p p r o a c h e s z e r o .

H a v in g b o t h th e v a l u e s

f o r e n v i r o n m e n t a l e r r o r an d f o r th e a d d it iv e g e n e t i c f r a c t i o n , th e
h e r ita b ility v a lu es can be e a s ily d ete r m in e d .

The fo r m u la u se d h e r e

to c a lc u la te h e r ita b ility p e r c e n t a g e s in e a c h e x p e r im e n t w as a s fo llo w s
2

<7?

H e r i t a b i l i t y = ~ --------

<r*of

x

100

4
BA.LWA.NT SINGH JOGI
2
2

ABSTRACT

i s e q u a l to v a r i a n c e d u e to a d d i t i v e g e n e t i c e f f e c t s .
.

i s t h e e n v i r o n m e n t a l v a r i a n c e c a l c u l a t e d f r o m p a r e n t s o n ly .

The e n v ir o n m e n ta l e r r o r , a d d itiv e g e n e tic f r a c t i o n s , and h e r ita b ilit y
p ercen tages w ere

c a lc u la te d in a ll th r e e e x p e r im e n ts .

b ility p e r c e n ta g e s fo r y ie ld a re
o f 33 p e r c e n t in t h e
th e F ,

31, 3 7 , a n d 32 p e r c e n t , w i t h a m e a n

g en era tio n .

T h e h ig h h e r i t a b i l i t y v a l u e s i n

g e n e r a t i o n i n d i c a t e s th a t h o m o z y g o s i t y h a s

a m a x im u m .

The h e r it a ­

reach ed p r a c tic a lly

D e l a y l a t e r t h a n t h i s w i l l b e a w a s t e of t i m e o n th e

p a r t of t h e p la n t b r e e d e r .
The com p on en ts

of v a r i a n c e w e r e c a l c u l a t e d f o r p o w d e r y
/

m i l d e w , H. s a t i v u m , d a te of h e a d i n g , h e i g h t of t h e p l a n t s , p e r c e n t
of lo d g e d p la n ts , lo d g in g

r e sista n c e

(c L r ), fa c to r and aw n ty p e.

The

h e r i t a b i l i t y v a l u e s f o r e a c h c h a r a c t e r i s g i v e n b e l o w f o r a l l th e t h r e e
ex p er im e n ts.
84%

80%

40%

4%

33%

36%

D a te of H e a d in g

7 6%

83%

-

H e ig h t

49%

40%

35%

L o d g in g

16%

35%

-

L o d g in g R e s is t a n c e

62%

17%

60%

T e s t W e ig h t

87%

55%

78%

M ild ew
H. s a tiv u m

5
BA.LWA.NT SINGH JOGI

ABSTRACT

L o d g in g r e s is t a n c e f a c t o r c L r is

F
e q u a l to — w h e r e F i s
b

e q u a l to f o r c e a p p l i e d an d b i s t h e h e i g h t of th e p la n t .
T h e s e h e r ita b ility v a lu e s a re

r e l a t i v e l y h ig h and p r o v i d e

i n f o r m a t i o n a s to p o s s i b l e g e n e t i c p r o g r e s s t h r o u g h s e l e c t i o n .

AN ESTIMATION OF GENETIC COMPONENTS OF VARIANCE
IN BULKED F ^ PROGENIES OF BARLEY

By
B alw ant Singh J o g i

A THESIS
S u b m itte d to t h e School o f G ra d u a te S t u d i e s o f M ichigan
S t a t e C o lle g e o f A g r i c u l t u r e and A p p lie d S c ie n c e
i n p a r t i a l f u l f i l l m e n t of th e requirem ents
f o r the degree of

DOCTOR OF PHILOSOPHY

D e p a r t m e n t o f Farm C r o p s
195*1-

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acknowledgments

T h i s s t u d y was c a r r i e d o u t u n d e r t h e g u i d a n c e o f
D r . K. T. P a y n e , Head a n d P r o f e s s o r o f Farm C r o p s , D r .
J . E. G r a f i u s , A s s o c i a t e P r o f e s s o r o f Farm C r o p s ,
D r . K. J . F r e y ,

and

a t p r e s e n t A s s o c i a t e P r o f e s s o r o f Farm

C r o p s a t I o w a S t a t e C o l l e g e , Ames, I o w a .

I take t h i s

o p p o r t u n i t y t o e x p r e s s ray s i n c e r e g r a t i t u d e t o t h e s e
t h r e e men.
A ppreciation i s

e x t e n d e d t o D r . W. D. B a t e n and

P r o f e s s o r H. M. Brown f o r h e l p w i t h t h e s t a t i s t i c a l
an aly sis.

G r a t e f u l a c k n o w l e d g m e n t i s g i v e n t o t h e Mid­

w e s t B a r l e y I m p r o v e m e n t A s s o c i a t i o n and M i c h i g a n Brew­
e rs A sso ciatio n f o r t h e i r f in a n c ia l a s s is ta n c e .
I a l s o e x p r e s s a s p e c i a l n o t e o f g r a t i t u d e t o my
b ro th er,

B a c h i t t e r Singh B a s s i ,

f o r h i s a s s i s t a n c e and

encouragem ent th ro u g h o u t th e e n t i r e program of s t u d i e s .
To a l l o t h e r s who c o n t r i b u t e d

to t h e s t u d y , b u t go u n ­

m e n t i o n e d , I am d e e p l y g r a t e f u l .

TABLE OF CONTENTS

Page
No.
INTRODUCTION,

1

REVIEW OF LITERATURE,

2

E s t i m a t i o n o f v a r i a n c e c o m p o n e n t s .......................

2

D ate o f h e a d i n g .

6

..............................................................

S p o t B l o t c h ( H e l m i n t h o 8 p o r lum s a t i v u m )
(Pam) K i n g .

e t c . . . ............... .......................... ..

g

P o w d e r y M ild ew ( E r y s i p h e g r a m l n l s h o r d e i )
L o d g i n g R e s i s t a n c e ...............................................................
EXPERIMENTAL PROCEDURE.
EXPERIMENTAL RESULTS,

©©©ecoeenpeove

The Com ponents o f V a r i a n c e

9
10

13
e © • e • » • ©• ■ > • ©« ©

21

...................... ..

21

E s t i m a t i o n o f G e n e t i c C o m p on en ts o f V a r i a n c e
f r o m F^ B u l k e d Y i e l d T r i a l s ..................................

27

E s t i m a t i o n o f G e n e t i c C o m p on en ts o f V a r i a n c e
f r o m O t h e r C h a r a c t e r s . ......................... ..................

30

D i s e a s e R e a c t i o n . .............................

31

A g r o n o m ic C h a r a c t e r s . .................

37

DISCU S S I 0N
CONCLUSIONS.. . . . . . . . . . . . . . . . . . . . . . . .

^7
. ...1

53

SUMMARY

55

LITERATURE CITED

56

INTRODUCTION
I n f o r m a t i o n i s now a v a i l a b l e on t h e u s e ©f e a r l y g e n e ­
ratio n

t e s t i n g i n s m a l l g r a i n s a s a means o f d e t e r m i n i n g t h o s e

c r o s s e s w h i c h w o u ld m o s t l i k e l y p r o d u c e t h e g r e a t e s t n u m b e r o f
high y ie ld in g seg reg ates.

From t h e s e s t u d i e s t h e c o n c e p t h a s

been d e veloped t h a t i t i s p o s s i b l e to o b ta in th e h i g h e s t
y i e l d i n g s e g r e g a t e s from th e h i g h e s t y i e l d i n g p a r e n t a l v a r i e ­
tie s.
As a r e s u l t o f g e n e t i c s t u d i e s w i t h s m a l l g r a i n s , v a r i o u s
m o d els have b e en c o n s t r u c t e d f o r s e p a r a t i n g th e com ponents o f
variance.

Such m o d e l s h a v e b e e n a p p l i e d t o t h e s e g r e g a t i n g

p o p u l a t i o n s a n d t o a l a r g e nu m b er o f b u l k e d F2 a n d F^ p r o ­
genies.
The p r e s e n t p a p e r i s a r e p o r t o f a s t u d y i n w h i c h an
a t t e m p t was made t o e s t i m a t e t h e g e n e t i c c o m p o n e n t s o f
v a r i a n c e i n v o l v i n g two c r o s s e s i n b u l k e d F^ p r o g e n i e s o f b a r ­
ley .

E f f o r t s h a v e a l s o b e e n made t o o b t a i n a d d i t i o n a l i n f o r ­

m a t i o n on t h e g e n e t i c c o m p o n e n t s o f v a r i a n c e f o r d i s e a s e r e ­
a c t i o n , d a te of h e a d in g , p l a n t h e i g h t , p e r c e n t of lodged
p lan ts,

lo d ging r e s is ta n c e ,

a n d awn t y p e .

REVIEW OF LITERATURE
E s t i m a t i o n o f V a r i a n c e C om ponents
F i s h e r ( 1 7 ) i n 1 925 i n t r o d u c e d t h e a n a l y s i s o f v a r i a n c e
a n d s u g g e s t e d i t s u s e i n t h e f o l l o w i n g ways;
1.

To o b t a i n t e s t s o f s i g n i f i c a n c e o f
treatm en ts.

2.

To e s t i m a t e v a r i a n c e c o m p o n e n t s .

The a n a l y s i s o f v a r i a n c e t o e s t i m a t e v a r i a n c e c o m p o n e n t s
h as been w id e ly used i n

th e f i e l d o f g e n e t i c s and i n s e l e c t i o n

f o r e f f i c i e n t sam pling d e s ig n s .

E x c e lle n t d is c u ssio n s of

t h e s e a p p l i c a t i o n s a r e f o u n d i n C o c h r a n ( 1 1 ) , Y a t e s and Z a z o p a n y (SO),

and Youden and M e h l i c h ( S i ) .

F i s h e r (16) f i r s t s e p a r a t e d g e n e t i c v a ria n c e i n t o
com ponents;

t h a t due t o a d d i t i v e e f f e c t s o f gen es,

three

t h a t due to

d om inance d e v i a t i o n s from t h e a d d i t i v e scheme, and t h a t du e to
d e v i a t i o n s fr o m t h e a d d i t i v e

scheme a t t r i b u t a b l e

to i n t e r -

a lle lic in te rac tio n s.
M ather (^7)

s u g g e s t e d t h a t t h e com ponents o f v a r i a n c e

can b e p a r t i t i o n e d i n t o
tab le,

two c o m p o n e n t s ; n a m e l y ,

and ( 2 ) h e r i t a b l e ,

(1) n o n - h e r i-

w h i c h i n t u r n may b e s u b d i v i d e d i n t o

f i x a b l e and u n f l x a b l e g e n e t i c v a r i a n c e .
C h a r le s and S m ith (S)

s e p a r a t e d g e n e t i c from t o t a l

v a r i a n c e by th e u s e o f e s t i m a t e s o f e n v ir o n m e n ta l v a r i a n c e
b a s e d on n o n - s e g r e g a t i n g p o p u l a t i o n s a n d p o s s i b l e r e l a t i o n s
b e t w e e n m ea n s a nd v a r i a n c e .

3

G r i f f i n g ( 2 6 ) i n a s t u d y on t o m a t o e s , c a l c u l a t e d t h e
c o m p o n e n t s i n a n a l y s i s o f v a r i a n c e a n d c o v a r i a n c e o f no n s e g r e g a t i n g p o p u l a t i o n s f o r t h e P ^ , P g , and F ^ .

T his a n a ly ­

s i s was f o u n d t o g i v e a s i m p l e , d i r e c t a n d p r o b a b l y t h e m o s t
a c c u r a t e m e t h o d o f o b t a i n i n g e s t i m a t e s o f p h e n o t y p i c and e n v i ­
r o n m e n t a l c o r r e l a t i o n s a nd r e l a t i o n s h i p s d e v e l o p i n g fr o m
these e stim a te s.

G e n o t y p i c c o r r e l a t i o n s w e r e c a l c u l a t e d mak­

i n g u s e o f g e n o t y p i c c o v a r i a n c e and t h e g e o m e t r i c means o f t h e
two p a r e n t a l g e n o t y p i c v a r i a n c e s .
are

the e r r o r ( w ith in v a r i e t i e s )

Of more i m m e d i a t e i n t e r e s t
v a r i a n c e and c o v a r i a n c e w h i c h

h e u s e d a s e n v i r o n m e n t a l v a r i a n c e and c o v a r i a n c e .
W r i g h t (79) d e v e l o p e d f o r m u l a e f o r the mean and v a r i a n c e
o f s q u a r e d d e v i a t i o n s f r o m a n optim u m f o r t h e c a s e s o f no
dom inance and o f c o m p le te d om inance,

f i r s t a s s u m i n g no e n v i ­

ro n m en tal c o m p l i c a ti o n s b u t l a t e r rem oving t h i s r e s t r i c t i o n .
The v a r i a n c e i n e a c h c a s e was s p l i t i n t o c o n t r i b u t i o n s d u e t o :
1.

A dditive g e n etic e f f e c t s .

2.

D o m inanc e d e v i a t i o n s .

3.

E p is ta tic d ev iatio n s.
E nvironm ental e f f e c t s

5.

N on-additive J o in t e f f e c t s of h e re d ity
and e n v ir o n m e n t.

He f u r t h e r d e v e l o p e d t h e f o r m u l a e w h i c h a p p l y t o e p i s ­
t a t i c r e l a t i o n s i n c a s e o f co m p lete dom inance.
posed form ulae f o r the

He a l s o p r o ­

c o r r e l a t i o n s betw een p a r e n t and o f f ­

s p r i n g a n d b e t w e e n two o f f s p r i n g .

He c o n c l u d e d t h a t i n a

.

p o p u l a t i o n i n w h i c h t h e mean o f some m e a s u r a b l e c h a r a c t e r i s
a t t h e o p t im u m ,

t h e p a r e n t - o f f s p r i n g and f r a t e r n a l c o r r e l a ­

t i o n s i n a d a p t i v e v a lu e a r e a p p ro x im a te ly the s q u a r e s of th e
c o r r e s p o n d i n g c o r r e l a t i o n s w i t h r e s p e c t to th e c h a r a c t e r i t ­
s e l f , w h a t e v e r e n v i r o n m e n t a l c o m p l i c a t i o n s t h e r e may b e .
Where t h e mean i s n o t a t t h e optim um t h e r e i s l e s s d i f f e r e n c e
b etw een th e c o r r e l a t i o n s i n a d a p t i v e v a lu e and c o r r e s p o n d i n g
o n e s w ith r e s p e c t to th e c h a r a c t e r i t s e l f .
The s t u d y o f h e r e d i t a r y and e n v i r o n m e n t a l c o m p o n e n t s o f
v a r ia tio n had i t s

b e g i n n i n g s i n t h e work o f J o h a n n s e n ( 3 7 ) >

who f i r s t d e m o n s t r a t e d t h a t b o t h h e r i t a b l e a n d n o n h e r i t a b l e
a g e n c ie s c o n tr ib u te d to som atic v a r i a t i o n in a s e g r e g a tin g
p o p u l a t i o n and t h a t v a r i a t i o n i n p u r e l i n e s w a s e n t i r e l y e n ­
vironm ental.

E a s t ( 1 5 ) f u r t h e r c o n f i r m e d t h e w o rk o f J o h a n n ­

s e n and d e m o n s t r a t e d how s u c h r e s u l t s c o n f o r m e d w i t h t h e c o n ­
c e p t o f M endelian g e n e t i c s .
F i s h e r , Xmmer, and T e d i n ( I S ) , M a t h e r ( ^ 7 ) »

L ush ( ^ 1 ) ,

P a n s e ( 5 $ ) a n d o t h e r s h a v e s t u d i e d h e r i t a b l e v a r i a t i o n and
have f u r t h e r su b d iv id ed i t

in to

th a t p o rtio n a ttrib u ta b le

to

a d d i t i v e g e n e t i c e f f e c t s a nd t h a t due t o d e v i a t i o n s f r o m t h e
ad d itiv e

scheme.

In a d is c u ss io n of h e r i t a b i l i t y

of c h a r a c te r s in a n i­

m a l s , L u s h ( ^ 2 ) d e f i n e s h e r i t a b i l i t y a nd d e s c r i b e s i t s u s a g e
in

th e broad and narrow a s p e c t s .

lity

In a broad sense, h e r i t a b i ­

r e f e r s t o t h e f u n c t i o n i n g o f t h e w hole g e n o t y p e a s a

u n i t and i s used i n c o n t r a s t w ith e n v ir o n m e n ta l e f f e c t s .

In

5.

th e narrow se n se h e r i t a b i l i t y i n c l u d e s o n ly the a v e ra g e e f f e c t s
o f g e n e s t r a n s m i t t e d a d d i t i v e l y from p a r e n t t o p r o g e n y .
L u sh and P a n s e ( ^ 3 ) p r o p o s e d t h e u s e o f t h e r a t i o o f t h e
a d d i t i v e g e n e t i c component o f v a r i a n c e t o t o t a l v a r i a n c e a s a
m easure o f th e d eg ree o f h e r i t a b i l i t y .
R o b i n s o n , C o m s t o c k , and H a r v e y ( 6 6 ) u s e d a m eth o d i n v o l v ­
i n g e s t i m a t e s o f com ponents o f v a r i a n c e t h r o u g h s t u d y o f b i p a r e n t a l p r o g e n i e s to m easure h e r i t a b i l i t y i n c o rn .

They

f u r t h e r compared t h i s method w i t h h e r i t a b i l i t y a s c a l c u l a t e d
from p a r e n t - o f f s p r i n g r e g r e s s i o n s .
G r a f i u s ( 2 2 ) d e v e l o p e d a model f o r t h e e s t i m a t i o n o f t h e
com ponents of g e n e t i c v a r i a n c e betw een s e l f “ f e r t i l i z e d ,
p r o g e n i e s from c r o s s e s o f i s o g e n i c l i n e s .
ta il

bulked

He p r e s e n t e d i n d e ­

t h e m a t h e m a t i c a l b a s i s f o r t h e m eth o d o f e s t i m a t i o n .
G - r a f i u s , N e l s o n and D i r k s

(2*0 s t u d i e d t h e h e r i t a b i l i t y

o f y i e l d i n b a r l e y a s m e a s u r e d by e a r l y g e n e r a t i o n b u l k e d p r o ­
genies.

They r e p o r t e d t h a t t h e n o n - h e r i t a b l e f r a c t i o n c o m p r i s ­

e d t h e m a j o r p o r t i o n o f t h e g e n e t i c v a r i a n c e i n t h e F g.
F^, w ith i n c r e a s e d hom ozygosity,

In the

the n o n - h e r i t a b l e f r a c t i o n d e ­

c r e a s e d i n com parison to th e a d d i t i v e

(h eritab le)

f r a c t io n of

the g e n e tic v a rian ce.
Mahmud and K r a m e r ( 5 0 ) made e s t i m a t e s o f h e r i t a b i l i t y
f o r y i e l d and p l a n t h e i g h t i n a s o y b e a n c r o s s .
the

They u t i l i z e d

s e g r e g a t e s a s t h e t o t a l v a r i a n c e and t h e s q u a r e r o o t o f

the p ro d u c t of th e p a r e n t a l v a ria n c e s a s the en v iro n m en tal
variance.

They f o u n d h e r i t a b l l i t i e s o f *J-3 and *1-1 p e r c e n t f o r

y i e l d and h e i g h t r e s p e c t i v e l y ,
W eber ( 7 5 )

u s i n g t h i s method.

a** i n t e r s p e c i f i c c r o s s o f s o y b e a n s , made

u se o f th e cube r o o t o f th e p r o d u c t o f t h e v a r i a n c e o f P^,
P g , and F^ a s t h e b e s t e s t i m a t i o n o f e n v i r o n m e n t a l v a r i a n c e .
C a l c u l a t i o n o f g e n e t i c and e n v i r o n m e n t a l c o r r e l a t i o n s i n v o l v e d
t h e u s e o f r e c i p r o c a l r e g r e s s i o n c o e f f i c i e n t s b e t w e e n F 2 and
t h e i r F^ p r o g e n i e s .

G e n e t i c c o r r e l a t i o n s were g e n e r a l l y h i g h ­

e r t h a n t h e o b s e r v e d c o r r e l a t i o n s i n Fg and F ^ .
Weber and M o o r t h y ( 7 6 ) i n t h r e e s o y b e a n c r o s s e s ,
seed y i e l d ,

studied

f l o w e r i n g t i m e , m a t u r i t y d a t e , p e r i o d fro m f l o w e r ­

i n g to m a t u r i t y , p l a n t h e ig h t ,
an i n d i v i d u a l p l a n t b a s i s .

s e e d w e i g h t and o i l c o n t e n t on

The e s t i m a t e s o f t h e g e n o t y p i c and

e n v i r o n m e n t a l v a r i a n c e s i n t h e s e s e v e n c h a r a c t e r s w e r e made.
They n o t e d t r a n s g r e s s i v e s e g r e g a t i o n f o r m o s t o f t h e c h a r a c t e r s
in a l l of these c ro sse s.

The h i g h e s t h e r i t a b i l i t y was n o t e d i n

f l o w e r i n g t i m e ( 75*7 p e r c e n t ) a n d t h e l o w e s t s e e d w e i g h t ( 5^*3
percent).
b ility

O i l c o n t e n t was 5^*7 p e r c e n t h e r i t a b l e .

The h e r i t a ­

o f s e e d y i e l d was e r r a t i c d u e t o a l a r g e e n v i r o n m e n t a l

v ariance.
D ate o f H eading
G riffee

(25) found an a s s o c i a t i o n betw een h e a d i n g d a t e

and tw o-row v e r s u s s i x - r o w c h a r a c t e r i s t i c .

He s u g g e s t e d t h a t

t h e i n h e r i t a n c e o f d a t e o f h e a d i n g i n b a r l e y was d u e t o one
facto r p air.
N eatby (5^)

r e p o r te d t h a t the i n h e r i ta n c e o f e a r l i n e s s

i n b a r l e y i s due to t h r e e f a c t o r p a i r s .

When t h e s e f a c t o r s

7

a r e I n t h e h o m o zy g ou s r e c e s s i v e

co n d itio n ,

velo p w i n t e r h a b i t s o f grow th.

He f u r t h e r s u g g e s t e d t h a t

.

the p l a n t s de­

t h e r e i s a l i n k a g e b e t w e e n t h e e a r l i n e s s and s h o r t v e r s u s
lo n g o u t e r glume.
Hehn ( 3 1 ) i n v e s t i g a t e d l i n k a g e i n b a r l e y i n c o n n e c t i o n
w i t h many c h a r a c t e r i s t i c s .

He c o n c l u d e d t h a t t h r e e f a c t o r

p a i r s d e te rm in e d the e a r l i n e s s o f h e ad in g in b a r le y .
W e x e l s e n ( 7 ^ ) r e p o r t e d two f a c t o r p a i r s d e t e r m i n i n g
d a te of heading In b a rle y .

He s u g g e s t e d t h a t h e a d i n g d a t e

was a s s o c i a t e d w i t h t h e two v e r s u s s i x - r o w c h a r a c t e r i s t i c ,
r o u g h v e r s u s s m o o t h awn and t h e n u m b e r o f b a r b s on t h e n e r ­
ves of

t h e lemma.
G -riffee ( 2 5 ) a ls o

reported th a t e a rlin e s s in b arle y is

l i n k e d w i t h t h e f a c t o r f o r s h o r t v e r s u s lo n g o u t e r glume.
Frey (20) i n v e s t i g a t e d

the h e a d in g d a t e In f o u r c r o s s ­

e s o f b a r l e y i n t h e F 2 g e n e r a t i o n and t h e p a r e n t s a n d p r o ­
g e n ie s o f F 2 p l a n t s o f seven o t h e r b a r l e y c r o s s e s .

He f o u n d

t h a t th e O g a l l t s u v a r i e t y o f b a r l e y d i f f e r s from H a r la n by
o ne f a c t o r f o r d a t e o f h e a d i n g .
H a r l a n by two g e n e s .

J e t and Anodium d i f f e r f r o m

B o t h J e t and Anodium c o n t a i n t h e same

gene p a i r s f o r d e te r m in in g h e a d in g d a te .

D a t e o f h e a d i n g was

f o u n d t o b e a s s o c i a t e d w i t h t h e r o u g h v e r s u s s m o o t h awn
c h a r a c t e r i s t i c i n a t h r e e way c r o s s , Moore x ( D o r s e t t x
M ^O O l),

a nd w i t h t h e b l a c k v e r s u s w h i t e h u l l c h a r a c t e r i n

a c ro s s o f H arlan x J e t .

S p o t B l o t c h ( H e l m l n t h o s p o r l u m s a t i v u m ) (Pam) K i n g
S p o t b l o t c h d i s e a s e o f b a r l e y was r e p o r t e d by Pammel
( 5 6 ) i n 1909 a n d was a t t r i b u t e d
Pammel, K i n g a n d Bakke ( 5 7 ) *

to H e l m i n t h o s p o r i u m s a t i v u m .

an<^ C h r i s t e n s e n ( 9 ) h a v e r e v i e w ­

e d p r e v i o u s s t u d i e s r e l a t i n g t o t h e p a t h o g e n e c i t y o f |L_ s a t i ­
vum and s u g g e s t e d t h a t t h e p a r a s i t e i s r e s p o n s i b l e f o r l e a f
spot,
ley*

root ro t,

f o o t r o t and s e e d l i n g b l i g h t o f w heat, b a r ­

r y e a n d many o t h e r g r a s s s p e c i e s .
Hayes e t a l (3 0 )

H.

sativum and i t s

s t u d i e d the i n h e r i t a n c e o f r e a c t i o n

a s s o c ia tio n w ith b o ta n ic a l c h a ra c te rs in a

M a n c h u r i a and L i o n c r o s s .
and Lion i s

to

su scep tib le

M anchuria i s a r e s i s t a n t v a r i e t y

t o t h e d i s e a s e . One h u n d r e d

tw enty-

f o u r F^ l i n e s w e r e grown i n rows o f 25 p l a n t s e a c h a n d s p r a y ­
ed w i t h s p o r e s u s p e n s io n s of

sativum .

Every t e n t h l i n e

was p l a n t e d a s a c h e c k , a l t e r n a t i n g t h e M a n c h u r i a and L i o n
paren ts.

From t h i s ,

t h e r e s i s t a n t and s u s c e p t i b l e l i n e s

w e r e s e p a r a t e d and t e s t e d i n

t h e F^ g e n e r a t i o n .

E ig h t lin e s

w e r e f o u n d t o b e a s s u s c e p t i b l e a s L i o n i n F^ and F^ and s i x
l i n e s were a s r e s i s t a n t a s M an ch u ria.

The c o r r e l a t i o n c o ­

e f f i c i e n t f o r d e g r e e o f i n f e c t i o n i n F-j and i n Fij. o f 36 h y ­
b r i d l i n e s o f L i o n x M a n c h u r i a was 0 . 2 5 6 + : . 1 0 5 n i f i c a n t a t the 5 p e rc e n t p o in t.

T his i s

sig ­

They c o n c l u d e d t h a t r e s i s ­

t a n c e t o H^ s a t i v u m i s a n i n h e r i t e d c h a r a c t e r a l t h o u g h a p p a ­
r e n t l y d e p e n d e n t on m ore t h a n a s i n g l e g e n e t i c f a c t o r .

9

G -riffee (25)
sativum i n r e l a t i o n

.

s t u d i e d , t h e r e a c t i o n o f Fj l i n e s t o H.
to o t h e r c h a r a c t e r s .

He c o n c l u d e d t h a t

a t l e a s t t h r e e f a c t o r s a r e c o n cern ed i n the r e s i s t a n c e of th e
t y p e p o s s e s s e d by t h e v a r i e t y S v a n h a l .

One f a c t o r was l i n k ­

ed w i t h t h e f a c t o r f o r tw o-row s, one w ith th e f a c t o r of
r o u g h awn a n d o n e w i t h t h e f a c t o r o f w h i t e g l u m e .
age o f

the f a c t o r s u s c e p t i b i l i t y

to r of

e a r l i n e s s i s much c l o s e r t h a n t h a t w i t h t h e f a c t o r

f o r six-row ed.

I t i s also

s e l f may p r e d i s p o s e

to

The l i n k ­

sativum w ith th e f a c ­

suggested th a t e a r l i n e s s in i t ­

t h e p l a n t to a t t a c k b y t h e p a t h o g e n .

P o w d e r y M ildew ( E r y s l p h e g r a m l n l s H o r d e l )
B r i g g s ( 6 ) s u g g e s t e d t h a t i n an i n h e r i t a n c e s t u d y o f r e ­
sistan ce

to any d i s e a s e i n p l a n t s th e f o l l o w i n g p o i n t s a r e o f

in terests
1.

The n u m b e r o f f a c t o r s f o r r e s i s t a n c e
p r e s e n t in each v a rie ty .

2.

The i d e n t i t y o f e a c h f a c t o r .

3.

The e f f e c t o f e a c h

The i n h e r i t a n c e o f r e s i s t a n c e t o

f a c t o r a c tin g alone.
b a r l e y m il d e w was s t u d i e d

h y b rid s in v o lv in g th ree r e s i s t a n t v a r ie ti e s ,

A rlington ,

C h i n e r m e , N i g r a t e and a s u s c e p t i b l e v a r i e t y ,

A tlas.

in

The

t h r e e r e s i s t a n t v a r i e t i e s d i f f e r from s u s c e p t i b l e A t l a s in
two i n d e p e n d e n t m a j o r f a c t o r s f o r r e s i s t a n c e t o t h i s d i s e a s e .
S u s c e p t i b i l i t y was r e c e s s i v e .
S t a n f o r d and B r i g g s ( 7 2 ) r e p o r t e d

t h a t t h e r e a r e seven

10.

d i f f e r e n t f a c t o r s f o r m il d e w r e s i s t a n c e ,
o ne r e c e s s i v e .

s i x d o m i n a n t and

The n u m b e r o f f a c t o r s i n a s i n g l e v a r i e t y

v a r i e s f r o m o ne t o t h r e e .
are d e f in ite ly lin k ed .

Of t h e s e v e n i d e n t i f i e d ,

two

The o t h e r f i v e a p p e a r t o be i n d e ­

pendent.
Ray e t a l

(62)

s tu d ie d e le v en wheat c r o s s e s in

t h e F2

g e n e r a t i o n f o r the i n h e r i t a n c e of m ature p l a n t r e s i s t a n c e
t o n a t u r a l i n f e c t i o n w i t h p o w d e r y m il d e w i n t h e f i e l d .
also

They

exam ined t e n c r o s s e s u n d e r g r e e n house c o n d i t i o n s f o r

seed lin g re sista n c e
fungus.

t o i n f e c t i o n by f o u r r a c e s o f t h e m il d e w

The s e e d l i n g r e a c t i o n a p p e a r s t o b e c o n d i t i o n e d by

a s i n g l e dom inant f a c t o r i n th r e e c r o s s e s .

One c r o s s s e g r e ­

g a t e d f o r two f a c t o r d i f f e r e n c e s f o r s e e d l i n g r e s i s t a n c e .
I t was r e p o r t e d t h a t u n d e r f i e l d

c o n d i t i o n s t h e ge ne

co n tro l­

l i n g t h e e x p r e s s i o n o f glume p u b e s c e n c e a p p e a r e d t o b e c l o s e ­
l y l in k e d w ith the m ajor f a c t o r f o r s e e d l i n g r e s i s t a n c e

to

one r a c e o f pow dery m ildew .
M a i n s a n d M a r t i n i (*J-5) t e s t e d a n u m b er o f v a r i e t i e s
a g a i n s t m i l d e w and f o u n d t h a t v a r i e t i e s d i f f e r e d
ly In t h e i r r e a c tio n s .

considerab­

Some showed d i f f e r e n c e s i n t y p e and

s t a b i l i t y o f r e a c t i o n and o t h e r s e x h i b i t e d marked d i f f e ­
r e n c e s i n r e a c t i o n to th e t h r e e p h y s i o l o g i c form s of m il ­
dew.
Lodging R e s i s t a n c e
G a r b e r and O l s o n ( 2 1 ) r e p o r t e d t h a t l o d g i n g i n c e r e a l s
i s a s s o c i a t e d w i t h many c h a r a c t e r s b u t t h a t no o ne c h a r a c t e r

se em s t o b e c o r r e l a t e d c l o s e l y w i t h l o d g i n g .
A t k i n s ( 1 ) s u g g e s t e d t h a t no o n e l o d g i n g i n d e x c a n b e
considered r e l i a b l e

s i n c e l o d g i n g i s d e p e n d e n t on a n u m b er

o f f a c t o r s t h a t v a ry g r e a t l y from y e a r to y e a r .

He s u g g e s t ­

ed t h a t t h e c h a r a c t e r o f s t r a w s t r e n g t h i s an i n h e r i t e d o n e .
R a m i a h a n d D h a r m a l in g a r a ( 6 l )

found i n c r o s s e s betw een

two t y p i c a l l y l o d g i n g a n d n o n - l o d g i n g v a r i e t i e s o f r i c e
th e re i s only a sin g le f a c to r p a i r d iffe re n c e .

th at

T h e r e se em s

to b e g e n e t i c a s s o c i a t i o n betw een th e c h a r a c t e r o f th e s tr a w
on one h a n d a n d s u c h c h a r a c t e r s a s t i l l e r i n g a n d f l o w e r i n g
d u r a t i o n i n d a y s on t h e o t h e r h a n d .

The non-S>dging n a t u r e

o f t h e s t r a w i s l i n k e d w i t h p o o r e r t i l l e r i n g and l o n g e r du­
ratio n .

T h e r e a p p a r e n t l y i s no r e l a t i o n s h i p b e t w e e n p l a n t

h e i g h t a n d s t r a w c h a r a c t e r n o r b e t w e e n s t r a w c h a r a c t e r and
flo ret s te rility .
C lark e t a l (10)

stu d y in g th e lin k a g e r e l a t i o n In b a r ­

l e y fo u n d t h a t t h e f a c t o r p a i r s o f aw nedness and l o d g i n g a r e
lin k ed .
Bose e t a l ( 5 ) f o u n d t h a t a t l e a s t two g e n e s w e r e r e s ­
p o n s i b l e f o r t h e d e v e l o p m e n t a nd d i s t r i b u t i o n o f s c l e r e n chyma t i s s u e .

T h e s e c h a r a c t e r s w e r e r e p o r t e d t o be a s s o c i a t

ed w i t h l o d g i n g r e s i s t a n c e .

They a l s o f o u n d weak s t r a w t o

be dom inant.
In an i n h e r i t a n c e stu d y of w eight p e r u n i t le n g th of
cu lm and o t h e r c h a r a c t e r s i n a c r o s s o f K a n r e d and C o p p e i ,
A tkins (2)

re p o rte d the fo llo w in g s i g n i f i c a n t c o r r e l a t io n

c o e f f i c i e n t s o f Ijb p a r e n t p l a n t s a nd t h e i r F-^ p r o g e n y l i n e s ;
diam eter of

culm 0 . 5 S 2 , h e i g h t o f t h e p l a n t 0 . ^ 3 0 , l e n g t h o f

th e head O .S3 0 .

On t h e b a s i s o f t h e s e r e s u l t s ,

he s u g g e s t e d

t h a t s e l e c t i o n f o r t h e s e c h a r a c t e r s can be made i n e a r l y
g e n e r a t i o n s and t h a t t h e c o n t i n u e d s e l e c t i o n w i l l l e a d t o
d e s ir e d type.
G - r a f i u s a n d Brown ( 2 3 ) d e f i n e d l o d g i n g r e s i s t a n c e
as the r a t i o

(L r)

o f t h e a d d i t i o n a l t o r q u e ( o v e r and a bove t h e

t o r q u e c a u s e d b y t h e w e i g h t o f t h e c u lm , l e a v e s a n d h e a d ) a
c ulm i s c a p a b l e o f r e s i s t i n g
by e x t e r n a l f o r c e s ,

(T r)

to th e to r q u e (Ta) a p p l i e d

s u c h a s w in d and r a i n .

They p r o p o s e d t h e

l o d g i r g r e s i s t a n c e f a c t o r ( c L r ) and s u g g e s t e d t h e f o l l o w i n g
form ula f o r i t s

calcu latio n ;

c L r = X where F i s e q u a l to f o r c e a p p l i e d
b
and b i s t h e h e i g h t o f t h e p l a n t .

13.

EXPERIMENTAL PROCEDURE
T his problem i s a c o n tin u a tio n of a b a r l e y p r o j e c t
which had been i n p r o g r e s s a t th e M ichigan E x p erim en t S t a t i o n ,
E a s t L a n s i n g s i n c e 19^-9*
g eneration

I t c o n s i s t s of a stu d y o f advanced

s e l e c t i o n s f r o m two c r o s s e s w h i c h i n c l u d e t h r e e

v a rie tie s.

The m a t e r i a l u s e d i n t h e e x p e r i m e n t was f r o m t h e

F^ b u l k e d r o w s w h i c h h a d b e e n a l l o w e d t o s e l f f o r two a d d i ­
t i o n a l g e n e r a t i o n s i n o r d e r to o b t a i n s u f f i c i e n t seed to con­
duct a y ield

t r i a l i n t h e F^ g e n e r a t i o n .

I n t h e f a l l o f 19^9 two b a r l e y c r o s s e s ,

S t e w a r d x Bay

a n d K i n d r e d x Bay w e r e made i n t h e g r e e n h o u s e .
K i n d r e d v a r i e t i e s a r e r o u g h awned a n d Bay i s

S t e w a r t and

s m o o t h awned.

S e l e c t i o n s f r o m S t e w a r t x Bay w e re d e s i g n a t e d A272 a nd K in ­
d r e d x Bay c r o s s e s w e r e I d e n t i f i e d a s A2 6 7 .

D uring the

s p r i n g o f 1 9 5 0 , th e F2 s e e d s were s p a c e p l a n t e d i n th e f i e l d .
E a c h p l a n t was h a r v e s t e d a n d t h r e s h e d s e p a r a t e l y .

The Fg

s e e d s f r o m I n d i v i d u a l p l a n t s w e r e grown i n s e p a r a t e ro w s i n
1951 a n d a t h a r v e s t e a c h row was b u l k e d .
c o n d u c t e d w i t h F^ s e e d i n 1 9 5 2 .

Y i e ld t r i a l s were

I n the f a l l of 1952, o u t of

t h e F-j b u l k r o w s , F^ s e e d s w e re t a k e n a t random f r o m b o t h s e ­
lectio n s.

T w en ty f a m i l i e s f r o m e a c h c r o s s w e r e s e l e c t e d .

E i g h t e e n s e e d s w e r e t a k e n fro m e a c h f a m i l y , m a k i n g a t o t a l o f
720.
The F ^ s e e d s t a k e n a t random fr o m I n d i v i d u a l F^ ro w s
w e r e g rown f o r i n c r e a s e I n t h e g r e e n h o u s e i n t h e f a l l o f

1952.

I n t h e s p r i n g o f 1953#

in c re a s e in the f i e l d .

s e e d s were p l a n t e d f o r

E a c h F ^ row i n t h e f i e l d was n u m b e r ­

ed so a s t o b e a b l e t o t r a c e i t s p e d i g r e e t o a s i n g l e F^ row.
A g r o u p o f t h e F^ ro w s t r a c i n g t o a d e s i g n a t e d F^ row was
c a lle d a fam ily .
Of 720 o r i g i n a l F^ s e e d s ,

223 p r o d u c e d a t l e a s t lj-0

g r a m s o f F^ s e e d , w h i c h i s n e c e s s a r y f o r y i e l d t r i a l s p l a n t e d
a t the
as is

calcu lated

r a t e of a p p ro x im a te ly f i v e p e c k s p e r a c re ,

shown i n T a b l e 1.

TABLE I .
THE NUMBER OF FAMILIES AND SUB­
FAMILIES IN SELECTIONS FROM TWO BARLEY
CROSSES IN WHICH SUFFICIENT SEED WAS
AVAILABLE FOR YIELD TRIALS IN THE
F^ GENE-RATION

S electio n s

Cross

A272

Stew art

Fam ilie s

S u b -fa m ilies in
each fam ily

T otal

19

k

76

19

g

152'

X

Bay
A267

K indred
X

Bay
G rand T o t a l

223

On t h e b a s i s o f s e e d s u p p l y a s i n d i c a t e d i n T a b l e 1 ,
t h r e e e x p e r i m e n t s were a r r a n g e d .

The l o c a t i o n a n d t o t a l

n u m b e r o f p l o t s i n e a c h may be s e e n i n T a b l e 2.

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THE
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NAME AND NUMBER
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15

16

The b u l k e d F ^ s e e d f r o m e a c h F ^ row
s e e d a n y i e l d p l o t i n 195 ^*

was u s e d t o

l n a l l th r e e e x p e rim e n ts eaoh

i n d i v i d u a l p l o t was a s u b - f a m i l y .

I t was i m p o s s i b l e t o r e ­

p lic ate

the s u b - f a m ilie s u sin g t h i s p l o t s iz e because of

lim ited

seed.

The f o u r s u b - f a m i l i e s i n e a c h m a in f a m i l y

were t r e a t e d a s ^ r e p l i c a t e s i n a l l t h r e e e x p e r im e n ts .

In

o r d e r to d e te rm in e th e e n v iro n m en ta l e f f e c t s in each e x p e ri
ment t h e p a r e n t p l o t s were u se d a s c h e c k s.
The c h e c k p l o t s w e r e grown a t random i n a l l t h r e e e x ­
p e r i m e n t s and a n a l y s i s o f v a r i a n c e f o r t h e p a r e n t s was c a l ­
cu lated

sep arately .

The mean s q u a r e f o r e r r o r was u s e d a s

e n v iro n m e n ta l v a ria n c e to t e s t th e s t a t i s t i c a l l y

sig n ifi­

c a n t d i f f e r e n c e s betw een and w i t h i n th e f a m i l i e s i n a l l
th re e experim ents.
A s i m p l e l a t t i c e d e s i g n was u s e d a t b o t h l o c a t i o n s .
E a c h p l o t was f o u r r o w s w i d e ,
s p a c in g betw een rows.

IS f e e t l o n g w i t h a one f o o t

At m a t u r i t y t h e 12 f o o t c e n t r a l p o r ­

t i o n o f t h e two i n n e r r o w s was h a r v e s t e d f o r y i e l d .
2 - ^ - D ( 2- ^ - D i c h l o r o p h e n o x y a c e t i c a c i d ) was u s e d f o r
weed c o n t r o l when t h e b a r l e y p l a n t s w e r e i n t h e f o u r - l e a f
stage.
w eighed,

A f t e r t h r e s h i n g th e i n d i v i d u a l p l o t seed l o t s were
c le a n e d and t e s t- w e i g h e d .

D u r i n g t h e g r o w i n g s e a s o n o b s e r v a t i o n s on d i s e a s e r e ­
a c t i o n a nd o t h e r a g ro n o m ic c h a r a c t e r s were r e c o r d e d .

The

d i s e a s e s c o n c e rn e d were powdery m ildew ( E r y s i p h e g r a m i n is
h o r d e ! ) and s p o t b l o t c h ( H elm lnthosporlum s a t iv u m ) .

.

17

.

A g r o n o m i c c h a r a c t e r s s t u d i e d were:; d a t e o f h e a d i n g ,
h e ig h t o f p l a n t , p e r c e n t o f lodged p l a n t s ,

lo d g in g r e s i s ­

t a n c e a n d s m o o t h v e r s u s r o u g h awn.
DISEASE REACTIONS
P o w d e r y Mildew
The a t t a c k o f m i l d e w was s e v e r e and t h e f i r s t f o u r
l e a v e s o f t e n random p l a n t s p e r p l o t were exam ined f o r m i l ­
dew i n f e c t i o n and r e a d i n g s w e r e r e c o r d e d on a d i r e c t p e r c e n ­
tage b a s i s .

The a v e r a g e o f t h e t e n r e a d i n g s p r o v i d e d t h e

p lo t ratin g .
The s u b - f a m i l i e s w e r e g r o u p e d on t h e b a s i s o f amount o f
in fe c tio n in to four c a te g o rie s.
51-75$ and 76-100$.

They a r e 0 - 2 5 $ ,

26 - 5 0$ ,

In eac h ex p erim en t the av erag e y i e l d

o f seed from th e s u b - f a m i l i e s f a l l i n g w i t h in e ac n o f th e
a b o v e c a t e g o r i e s was c o m p a r e d w i t h a v e r a g e f r o m t h e o t h e r
c a t a g o r i e s t o d e t e r m i n e w h e t h e r s e e d y i e l d was d i r e c t l y r e ­
l a t e d to m ildew r e a c t i o n .
S p o t B l o t c h ( H. s a t i v u m )
Ten t o t w e n t y p l a n t s w e r e
row s

p u l l e d fro m t h e two o u t e r

J u s t p r i o r t o h a r v e s t and t h e r o o t s and s t e m s w e r e

exam ined

from th e f i r s t

to th e f i f t h b a s a l n o d e s.

a m o u n t o f i n f e c t i o n on e a c h p l a n t was n o t e d .

The

The r a t i n g

s c a l e u s e d was z e r o t o f i v e w i t h f i v e i n d i c a t i n g c o m p l e t e
in fectio n .
The

s u b - f a m i l i e s were grouped a c c o r d i n g to

sativum

IS.

reactio n in to
zero o r one,
fiv e.

four c a ta g o rie s .

T h e r e w e r e no n e i n c l a s s e s

so a l l f e l l i n c l a s s e s tw o , t h r e e ,

four o r

The a v e r a g e s e e d y i e l d o f e a c h c a t e g o r y was co m p ared

w ith th a t in

th e o t h e r c a t e g o r i e s to d e te rm in e th e e f f e c t s ,

i f a n y , o f H»_ s a t i v u m on y i e l d .
AGRONOMIC CHARACTERS
D ate o f Heading
N e a r l y a l l of
m o n th o f J u n e .

B a s e d on d a t e o f h e a d i n g ,

were g ro u p e d i n t o
15;

the p l o t s a t b o th l o c a t i o n s headed in th e
the s u b - f a m ilie s

f o u r c l a s s e s a s follow s^

J u n e 16 t o J u n e 20; J u n e 21 t o J u n e 25;

after.

J u n e 10 t o J u n e
a n d J u n e 26 o r

The a v e r a g e s e e d y i e l d f o r e a c h o f t h e f o u r c l a s s e s

was c o m p a r e d w i t h t h e o t h e r s t o d e t e r m i n e w h e t h e r s t a t i s t i ­
c a l l y s i g n i f i c a n t d i f f e r e n c e s e x i s t e d b e t w e e n t h e i r mean
yields.
H eight o f P la n t
H e i g h t n o t e s w e r e t a k e n on 15 t o 20 p l a n t s i n e a c h p l o t
and a v e r a g e d .

On t h e b a s i s o f h e i g h t c l a s s i f i c a t i o n ,

s u b - f a m i l i e s were d i v i d e d i n t o

th re e groups.

the

The f i r s t

g r o u p i n c l u d e s t h o s e s u b - f a m i l i e s w h i c h w e r e 20 t o 25 i n c h ­
es t a l l ,

th e second group i n c lu d e d

es in h eig h t,

t h o s e f r o m 26 t o 30 i n c h ­

and t h e t h i r d g r o u p was made up o f t h o s e 31

inches o r t a l l e r .

The a v e r a g e s e e d y i e l d s o f t h e s e g r o u p s

w e r e c o m p a r e d t o d e t e r m i n e w h e t h e r y i e l d was a s s o c i a t e d w i t h
p la n t h eig h t.

19

.

Lodging
The a m o u n t o f l o d g i n g was e s t i m a t e d and r e c o r d e d i n p e r ­
c e n t a t h a r v e s t tim e.
four categ o ries;

The s u b - f a m i l i e s w e r e g r o u p e d I n t o

0 - 2 5 $ , 26- 50$ ,

5 W 5 # and 7 6 - 1 0 0 $ .

The

a v e r a g e y i e l d s o f se e d o f t h e s e c a t e g o r i e s were compared t o
d e t e r m i n e w h e t h e r o r n o t l o d g i n g was a s i g n i f i c a n t f a c t o r
in seed y i e l d s .
Lodging R e s i s t a n c e
Two w e e k s p r i o r t o m a t u r i t y ,
lod g in g r e s is ta n c e .

A s p e c i a l c h a i n o f 150 l i n k s was u s e d ,

e a c h l i n k w e i g h i n g o n e gram.
of the ch ain .

r e a d i n g s w e r e t a k e n on

A h o ok was a t t a c h e d t o o n e end

The h o o k was f a s t e n e d t o t h e r a c h i s a t t h e

base of the head,

and t h e culm was a l l o w e d to b e n d u n t i l a

p o r t i o n o f t h e l i n k s w e r e on t h e g r o u n d a nd t h e r e m a i n d e r
w e r e s u p p o r t e d by t h e c u lm .

The r e a d i n g s w e r e t a k e n by

c o u n t i n g t h e l i n k s f r o m t h e g r o u n d to t h e h o o k .

Five re a d ­

i n g s w e r e t a k e n i n e a c h p l o t and t h e a v e r a g e was u s e d a s an
i n d ic a tio n of lodging r e s is ta n c e .

T h i s a v e r a g e was d i v i d e d

by t h e p l a n t h e i g h t o f e a c h r e s p e c t i v e p l o t p r i o r t o s t a t i s ­
tic a l an ly sis.

The l o d g i n g r e s i s t a n c e ( c L r ) f a c t o r i s

equal

to Z where F i s
b
p la n t h eig h t.

e q u a l t o f o r c e s u p p l i e d a nd b i s e q u a l to

I n Ingham E x p e r im e n ts I and I I r e a d i n g s were r e c o r d e d
two w e e k s p r i o r t o m a t u r i t y and a g a i n s h o r t l y a f t e r m a t u r i t y .
The n u m b e r o f c u l m s t h a t b r o k e w i t h t h e w e i g h t o f t h e c h a i n
and t h e t y p e o f c u r v e c r e a t e d i n t h e culm b y t h e w e i g h t o f

t h e c h a in were a l s o r e c o r d e d b u t a r e n o t b e i n g p r e s e n t e d a s
a p a r t of t h i s

th esis.
Smooth V e r s u s Rough Awn

I n Ingham E x p e rim e n t I ,
p a r e n t l y the

no s e g r e g a t i o n was n o t e d .

Ap­

s m o o t h awned s e g r e g a t e i n t h e F 5 g e n e r a t i o n was

used as the S tew art p a r e n t.
I n In g h am E x p e r i m e n t I I and T u s c o l a I I I ,
l i e s were g ro u p ed i n t o

sm o o th a n d r o u g h awn.

the su b -fa m i­
The a v e r a g e

s e e d y i e l d o f e a c h g r o u p was c a l c u l a t e d and c o m p a r e d f o r
p o ssib le

sta tistic a lly

sig n ific an t d ifferen ces.
T est-W eight

T e s t - w e i g h t s o f e a c h s u b - f a m i l y and t h e p a r e n t a l p l o t s
were t a k e n .
Y ield
Y i e l d o f p l o t s was d e t e r m i n e d i n g r a m s a nd p l o t s w e r e
compared f o r

sta tistic a lly

sig n ific a n t d ifferen ces.

21.

EXPERIMENTAL RESULTS
The C om p o nents o f V a r i a n c e
Assume t h a t k s e e d s w e r e t a k e n a t random fr o m F^ b u l k
row s a n d s e l f e d

j g enerations.

The F^ l i n e s u s e d w e re r a n ­

dom s a m p l e s o f k s e e d s f r o m e a c h o f m F^ b u l k r o w s .
o f t h e k s e e d s was i n c r e a s e d f o r two g e n e r a t i o n s .

Each
H e n ce ,

e a c h p l o t w a s t h e r e s u l t o f a random sa m p le o f s e e d s f r o m
b u l k F j r o w s w h i c h w e r e a l l o w e d t o s e l f f o r two a d d i t i o n a l
gen eratio n s.

F o r e x a m p l e , on a s i n g l e g e n e b a s i s i f

s e e d c h o s e n w e r e AA o r a a ,

t h e r e s u l t i n g p l o t means w i l l r e ­

m ain u n c h a n g e d a f t e r J g e n e r a t i o n s o f s e l f i n g .
hand, i f

the

On t h e o t h e r

t h e s e e d c h o s e n w e r e Aa t h e n t h e p l o t mean w i l l

change w i t h each g e n e r a t i o n of s e l f i n g .

T h i s c an b e s t be

i l l u s t r a t e d w i t h an a l g e b r a i c m odel.
F o llo w in g M ather ( ^ 7 ) i
a a e q u a l d ^ , hj_ a n d - d ^ ,
T ables I I I

a n d IV, e t c .

l e t t h e mean e f f e c t s o f AA, Aa,

r e s p e c t i v e l y , a s may be s e e n i n
The m o d e l a s s u m e s o n l y t h e a d d i ­

t i v e a c t i o n o f g e n e s and no e p l s t a s i s .

The v a l u e h e q u a l s

t h e d o m i n a n c e d e v i a t i o n f r o m t h e a d d i t i v e scheme and may b e
p o s itiv e o r n eg ativ e.
TABLE I I I .
DISTRIBUTION OF GENOTYPES AND THEIR
MEAN EFFECTS IN AN F2 POPULATION

G-enotype o f F

p lan ts

Frequency

AA

i

Aa
aa

*

Mean e f f e c t

hi
“ dL

22

From T a b l e I I I ,

o n e may c a l c u l a t e

th e F2 v a r i a n c e

w h i c h w i l l be*.
£< *!+ £

a?

&

h? - ( £ h ^ ) 2 =
Aa

£ h|
Summing o v e r n s u c h l o c i g i v e s s
(* a?-t- i h |

)

n : l
=

*

D+ iH

The d i s t r i b u t i o n o f g e n o t y p e s a t a s i n g l e l o c u s i n
F^ i s g i v e n i n T a b l e IV .

I t is

the

to be n o t e d t h a t t h e c e n t e r

c l a s s r e p r e s e n t s a s i n g l e F_ row w i t h a mean o f h * / 2 .
In
j
t h i s e x p e r i m e n t s e e d s w e r e draw n a t random f r o m e a c h F_ row

3

g iv in g the d i s t r i b u t i o n

shown i n T a b l e V.

The t h r e e c l a s s e s

i n t h e l e f t h a n d co lu m n o f T a b l e V r e p r e s e n t i n d i v i d u a l F^
p la n ts in a l l p o ssib le

types of fa m ilie s .

TABLE I V . DISTRIBUTION OF GENOTYPES AND THEIR
MEAN EFFECTS IN AN F^ POPULATION

E xpected genotype of
F j row

Frequency

F-z row mean
effect

aa

£ aa
iA a

h /

( £ d^-p & h£ - £ d l )

aa

-d;

£

i

£ aa

TABLE V„

DISTRIBUTION OF GENOTYPES AND THEIR MEAN
EFFECTS IN AN F^ POPULATION

E xpected g e n o ty p ic f a m i l i e s
o f s e l e c t e d p l a n t s i n Fj^

Frequency

AA

£

^ A a^.^ aaj

\

{*

aa

TABLE VI.

£

-A l

DISTRIBUTION OF GENOTYPES AND THEIR MEAN
EFFECTS IN AN F^ POPULATION

E xpected g e n o ty p ic f a m i l i e s
o f F c row s f r o m F)± p l a n t s
in Table I I I
AA
j|

Fam ily
mean e f f e c t

^ + 1

Fam ily
mean e f f e c t

£
Aa«f £ a a ) _ ^ aaj

AA

Frequency

ajL

£

&

a

4

I t w i l l b e n o t e d i n T a b l e s V and VI t h a t t h e f a m i l y
mean e f f e c t f o r t h e c e n t e r c l a s s i s h a l v e d i n e a c h g e n e r a t i o n .
T h i s i s t h e e x p e c t e d r e s u l t u n d e r s e l f i n g a nd i f
t h e t o t a l f a m i l y mean e f f e c t a p p r o a c h e s z e r o .

continued,

By t h e t i m e

t h e s e l e c t i o n s h a d r e a c h e d F ^ t h e f a m i l y mean e f f e c t f o r t h e
h e t e r o z y g o u s f a m i l y h a d b e e n r e d u c e d t o h» a s shown i n T a b l e
16
VII.

TABLE V I I .
DISTRIBUTION OF GENOTYPES AND THEIR MEAN
EFFECTS IN AN F$ POPULATION

E xpected genotype o f y i e l d
p l o t s i n F^

Frequency

F a m i l y mean
effect

AA

^ 4 -!

d >L
^ * 1

h^ / 1 6

*

a a ) + ^ ■ aa]

aa

*

F i e ld Experim ents
As havs b e e n n o t e d

i n Ingham E x p e r i m e n t I ,

t h e r e were

f a m i l i e s e a c h w i t h f o u r s u b - f a m i l i e s w hich were t r e a t e d
p lic ate s.
dom,

19

as re­

Inasm uch a s t h e s u b - f a m i l i e s were sampled a t r a n ­

the r e p l i c a t e

t o t a l s h o u l d show no g e n e t i c b i a s .

Twelve

p l o t s o f e a c h o f two p a r e n t s we re p l a n t e d a t random t h r o u g h ­
o u t t h e e x p e r i m e n t and were a n a l y z e d s e p a r a t e l y i n o r d e r to
o b t a i n an e s t i m a t e o f t h e e n v i r o n m e n t a l v a r i a n c e .
In g h a m E x p e r i m e n t s I a n d I I w e r e composed a s f o l l o w s ;
T o t a l n u m b e r o f f a m i l i e s ................................................ 19
T o t a l number o f s u b - f a m i l i e s i n eac h
m a in f a m i l y . .............................

4

T o t a l number of p l o t s f o r

f a m i l i e s 19 x

T o ta l number o f p l o t s f o r

two p a r e n t s 12 x

Gra nd t o t a l

4 ...7 6

100

25.

I n T uscola Experim ent I I I

t e n p l o t s were e l i m i n a t e d

due to a l a c k of a d e q u a te space.

At t h i s l o c a t i o n t h e r e

w e r e s e v e n t y p l o t s f o r f a m i l i e s and t w e n t y f o r p a r e n t s ,
m aking a t o t a l o f 9 0 .
A n a l y s i s o f v a r i a n c e c o m p o n e n t s may b e s e e n i n T a b l e
V III.
TABLE V I I I .
THE ANALYSIS OF VARIANCE WITH MEAN
SQUARE EXPECTATIONS OF NUMBER OF m BULK ROWS
IN k REPLICATION USED IN A CROSS WITH 1
PARENTS
Sources o f v a r i a t i o n

Betv/een f a m i l i e s

D.F.

ra- 1

M. S.

Mean s q u a r e e x ­
pectation

M1

< r

k

<#•

k-1

Reps
F a m i l i e s x Reps

P arents

It

( m-1 ) ( k - 1 )

1

m2

M3

cp

+•

o f

c 8-

i s n e c e s s a r y t o d e t e r m i n e ,2. and 2 ( See T a b l e I X ) .
0?
tjr

26.

TABLE I X .

MODEL FOR THEiNALYSIS OF VARIANCE INVOLV­
ING- A SINGLE LOCUS

Frequency of
fam ilies

Genotype
of o r i g i ­
n a l BJj row

T otal p o s s ib le c a te g o rie s
of p lo ts
hX

*

AA

\

Aa

- a -L

Fam ily
mean
effect

di.
3 ^

i(.|

hJ
i f

ap

j

1

i4

i

1
1

1
i
i

-3 d -

aa

The t o t a l

“ di

sura o f s q u a r e s w i l l t h e n bes
3 /saf

>-hi(

j2

JU i
=:

(i

-

3 /8 )^ + . h ^

7 /^ i+

-

7/102^ h

T o t a l S. S . - 7/SD + 7/1024- H
2
The sum o f s q u a r e s d u e t o f a m i l i e s = U f ~

< /

( * d 2 -+- i d 2 + 1 / 5 1 2 h 2 - l / 1 0 2 4 h 2 ]

s 1
J D + 1/1024 H

27.

The r e m a i n d e r ( T o t a l S . S . - S . S . d u e t o f a m i l i e s ) r ( j f - .
2
( p r - ( 7 / S D -4.7/102*1 H) - ( 1 / 2 D + 1 / 1 0 2 4 H)
3

3 /S D-t-6/1024 H

At t h i s p o i n t i t
H is

so s m a l l t h a t i t

c an be s e e n t h a t t h e c o e f f i c i e n t o f
can s a f e l y be I g n o r e d .

The c o e f f i c i e n t

o f D w i t h i n f a m i l i e s i s 3 / 4 o f t h a t betw een f a m i l i e s .
E s t i m a t i o n o f G-enetic Co m ponen ts o f
v a r i a n c e f r o m F^ B u l k e d Y i e l d T r i a l s
I n t h e Fg g e n e r a t i o n t h e g e n e t i c c o m p o n e n t s o f v a r i a n c e
may b e c a l c u l a t e d a s f o l l o w s ;
1 / 2 D Hh 1 / 1 0 2 4 H

2
( J T T ..........................................( a )

3/ s D S- 6 / 1 0 2 4 H

< j f c .......................................... ( b )

In these

two e q u a t i o n s t h e c o e f f i c i e n t o f H i s

i n t h e F^ g e n e r a t i o n t h a t i t a p p r o a c h e s z e r o .

so low

The a d d i t i v e

g e n e tic v a ria n c e w ith in f a m ilie s in eq u atio n (b) i s only
3 /4 th e s i z e o f t h a t betw een f a m i l i e s .
I n T able 10,

the a n a ly s is of v ariance f o r y ie ld of a l l

th ree experim ents i s

g i v e n a nd d i f f e r e n c e s

s i g n i f i c a n t a t the

one p e r c e n t l e v e l w e r e o b t a i n e d b e t w e e n f a m i l i e s .

S ig n ifi­

c a n t d i f f e r e n c e s were n o t o b t a i n e d w i t h i n f a m i l i e s .
The h e r i t a b i l i t y

v a l u e s g i v e n i n T a b l e 11 w e r e c a l c u ­

l a t e d from t h e f o l l o w i n g fo rm u la ;
2

H eritab ility

-

(T F
2
C T~

2
4- CTF

28.

The h e r i t a b i l i t y

v a l u e s a r e } 1 , 3 7 > 32 p e r c e n t f o r

Ingham E x p e r i m e n t s I and I I
resp ectiv ely .

and T u s c o l a E x p e r i m e n t I I I

These r e l a t i v e l y h ig h h e r i t a b i l i t y

figures

i n d i c a t e t h a t s e l e c t i o n f o r y i e l d a t t h i s s t a g e o f homo­
z y g o s i t y would b e s u c c e s s f u l .

?9

*

TABLE X.
THE ANALYSIS OF VARIANCE FOR YIELDS OF Fg BULK
PROGENIES IN TWO BARLEY CROSSES GROWN IN INGHAM
AND TUSCOLA COUNTIES

X
id
<D
G
*d
G
P

♦
*
O

fr«

oa

aj

*

aj
VO
„
rH

S'
W
cd
H
O
o
CO
3
Eh

e
o

G

cd

150

lta
.

MS

cd
CQ

M
M
M

o
150
CVJ
OA

XA
ISO
•
o
OA
rH
LTV

P

UA
OA

G

«5
X

VO
OA
rH
XA

Vi

o

CQ
P
CQ

to

w

CQ
CO

O
OS

o

>5
a)
m

M
M

x

VS
&
w
£4
3
.G

'd
a>
G
Kt

G
P
«

X>

*
o

&«

xa

•
XA
VO
XA
•
VO
•=3"
XA
VO

CO
a

G
M

*
♦
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Xs
cd
CQ
X
P
g

cd
S*
(D
p
CO

M
•
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X
w

— ————

CO
2

s

XOA
rH
VO

a

cd
.G

XA
OA
•
rH
ISO
rH
9

IXA
O
X -

XA

•
•

LfA

x-

150
rH

•=t
■=t

•

aj
OA
rH

vo
o

OA
150
«
XXXA
aj

-d-

LTA

•

CQ

rH
0}
p
o
Eh

CQ
CD
P
rH
P

sCD
O
P
<P
P
G
5D
P
CQ

.G

cd

a>

P

s

o
G

<P
<D
P

—~ • - ■ •
150
ISO
•
150
XA
cu
CQ

150
rH

c

CD

G
cd

X

G
O

s

cd

X
CD
H
rH
03
.G
p
CQ
«d
flA
0/
©
o

CQ

fe .

3

o
rH
03
o

cd

O,
«

cc

<d
rH

rH

CQ
P
ip
O <D
O
CQ G
CD 03
O P
G G
3 03
o !>
CQ

G

O

bD

G
HI

cd

•
CD

U
o

rH
aj
— ——— ---o
i—i

rH

G X^
W “

X
CD
CD
3
rH
cd
X
Cti

*
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150
^d"

-d“
H

•

•
&«
•

•

O
cd

rH
O

O

CQ

3

O
cd

rH
O

o

CQ

3

Eh

Eh

G

G

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G
O
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G
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•

x»

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G
O

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cd
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G

S
G
O
G

G
cd
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V

G
W

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30.

TABLE X I .
THE CALCULATED VALUES OF E , 1 / 2 D
AND HERITABILITY PERCENTAGES FOR YIELD
FROM Fg BULK PROGENIES IN TWO BARLEY
CROSSES GROWN IN INGHAM AND TUS­
COLA COUNTIES

Cross

E xperim ent

E \J

1/2 D

H eritab i­
lity %

Ingham I

S t e w a r t x Bay

2 2 3 S .S S

939.55

31

Ingham I I

K i n d r e d x Bay

1924.44

1105.4S

37

T uscola I I I

K i n d r e d x Bay

3196.95

1520.91

32

3/ Where E i s

th e e s tim a te of environm ental v arian ce

and 1 /2 D i s

the e stim a te of a d d itiv e g e n e tic

variance.
The E s t i m a t i o n o f G e n e t i c C om ponents o f V a r i a n c e
o f O t h e r C h a r a c t e r s O b s e r v e d From Fg
Bulked P r o g e n ie s
The r e s u l t s o f s t u d i e s o f o t h e r c h a r a c t e r i s t i c s o b s e r v e d
i n t h e f i e l d a r e p r e s e n t e d u n d e r t h e f o l l o w i n g two s u b - h e a d s ;
D isease re a c tio n
1.

P o w d e r y Mildew (EL g r a m l n l s h o r d e i )

2.

S p o t B l o t c h (Hj_ s a t i v u m )

Agronomic c h a r a c t e r s
1.

D ate of heading

2.

P e rc e n t of lodged p l a n t s

3.

H eight o f p la n t

31

4.

Straw s t r e n g t h

5.

Sm ooth v e r B u s r o u g h awn
D ise ase R eaction
P o w d e r y M ildew

There were s t a t i s t i c a l l y

s i g n i f i c a n t d if f e r e n c e s in

am o u n t o f m i l d e w p r e s e n t b e t w e e n t h e f a m i l i e s i n a l l t h r e e
experim ents.
T ab le 12.

The a n a l y s i s o f v a r i a n c e may b e s e e n i n

The r e s u l t s g i v e n i n T a b l e 13 i n d i c a t e no s i g ­

n ific a n t d iffe re n c e s in
d iffe re n t q u artile

the a v erag e seed y i e l d s o f th e

classes.

I t may b e i n t e r p r e t e d t h a t t h e

p a t h o g e n d i d n o t h a v e a d i f f e r e n t i a l e f f e c t on y i e l d o f
seed.

I n a f u r t h e r a t t e m p t t o d e t e r m i n e w h e t h e r m il d e w I n ­

f e c t i o n was a f f e c t i n g s e e d y i e l d ,
ed I n 10 p e r c e n t i l e

th e f a m i l i e s were g ro u p ­

c l a s s e s r a t h e r than q u a r t i l e s .

Analy­

s i s i n d i c a t e d no s i g n i f i c a n t d i f f e r e n c e b e t w e e n c l a s s
y ield averages,

which f u r t h e r s u b s t a n t i a t e s t h a t se v e re

mildew i n f e c t i o n was n o t a f f e c t i n g
m ild i n f e c t i o n .

seed

s e e d y i e l d a n y more t h a n

.

32.

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TABLE X I I I .
THE EFFECT OF MILDEW ON THE AVERAGE YIELD OF
FAMILIES CLASSIFIED
IN PERCENT FROM F£ BULK PROGENIES
IN TWO CROSSES GROWN IN INGHAM AND TUSCOLA COUNTIES

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S p o t B l o t c h ( H. s a t i v u m )
The s p o t b l o t c h r e a c t i o n i n T a b l e 1^ i n d i c a t e s no
s i g n i f i c a n t d i f f e r e n c e betw een f a m i l i e s .
sta tistic a lly

In a d d itio n ,

s i g n i f i c a n t d i f f e r e n c e s were n o t n o te d i n

th e a v e r a g e y i e l d s betw een th e f o u r d i f f e r e n t c l a s s e s of
H.

sativum r e a c t i o n .

T h is i n d i c a t e s t h a t th e pathogen

had n o t produced a d i f f e r e n t i a l e f f e c t in r e g a rd to a v e r ­
age seed y i e l d s o f d i f f e r e n t c l a s s e s o f f a m i l i e s .

35*

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TABLE XIV.
ANALYSIS OF VARIANCE FOR H. SATIVUM
DATA OBSERVED FROM F6 BULK PROGENIE S~TN“ TWo
BARLEY CROSSES GROWN IN INGHAM AND TUSCO­
LA COUNTIES

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TABLE XV.
THE EFFECT OF H. SATIVUM ON THE AVERAGE SEED YIELD
OF FAMILIES IN THE F6 BU lK “ PROGENIES IN TV/O BARLEY CROSS­
ES GROWN IN INGHAM AND TUSCOLA COUNTIES

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A g r o n o m ic C h a r a c t e r s
H ig h ly s i g n i f i c a n t d i f f e r e n c e s betw een th e f a m i l i e s
i n d a t e o f h e a d i n g i n a l l t h r e e e x p e r i m e n t s may be s e e n i n
Table 16.

T h e r e w e r e no s i g n i f i c a n t d i f f e r e n c e s i n t h e

a v e ra g e y i e l d s of t h e f a m i l i e s a t d i f f e r e n t h e a d in g d a t e s ,
a s shown i n T a b l e 1 7 .

I t may b e c o n s i d e r e d t h a t t h e d a t e

o f h e a d i n g h a d no s i g n i f i c a n t e f f e c t on t h e a v e r a g e y i e l d s
o f the f a m i l i e s .
S i g n i f i c a n t d i f f e r e n c e s a t t h e one p e r c e n t l e v e l were
o b t a i n e d b e t w e e n t h e f a m i l i e s f o r l o d g i n g ( T a b l e XX) i n I n g ­
ham E x p e r i m e n t I I ,

w h e r e a s t h e r e w a s no s i g n i f i c a n t d i f f e ­

r e n c e i n Ingham E x p e rim e n t I b e tw e e n t h e f a m i l i e s .

The a v e r ­

age y i e l d s i n b o th Ingham e x p e r i m e n t s c l a s s i f i e d u n d e r l o d g ­
i n g i n T a b l e 19 show no s i g n i f i c a n t d i f f e r e n c e b e t w e e n t h e
catag o ries.
H ig h ly s i g n i f i c a n t d i f f e r e n c e s i n h e i g h t e x i s t e d be­
tw e e n f a m i l i e s ,

a s may be s e e n i n T a b l e s 20 a n d 2 1 .

may b e i n t e r p r e t e d

T his

to mean t h a t t h e r e w e r e d i f f e r e n c e s i n

the a v erag e y i e l d s of

the f a m i l i e s due to h e i g h t .

The a n l y s i s o f v a r i a n c e f o r l o d g i n g r e s i s t a n c e ( c L r )
p r e s e n t e d i n T a b l e 22 i n d i c a t e s s i g n i f i c a n t d i f f e r e n c e s b e ­
t w e e n f a m i l i e s a t t h e \% l e v e l i n In gh am E x p e r i m e n t I I .
The a v e r a g e y i e l d s o f s m o o t h and r o u g h awned f a m i l i e s
a r e g iv e n i n T able 23.

T his i n d i c a t e s t h a t

t h e r e a r e no

s i g n i f i c a n t d i f f e r e n c e s betw een f a m i l i e s i n Ingham E x p e r i ­
ment I I and T u s c o la E x p e rim e n t I I I .

As h a s b e e n s t a t e d ,

fam i­

l i e s I n I n g h a m E x p e r i m e n t I d i d n o t s e g r e g a t e f o r awn c h a r a c ­
ter.

3 *.

TABLE XVI „ ANALYSIS FOR DATE OF HEADING OB­
SERVED IN F £ BULKED PROGENIES IN TWO BAR­
LEY CROSSES GROWN IN INGHAM AND TUSCOLA
COUNTIES

CRO SSES
S t e w a r t x Bay
In g h am Exp. I
Sources of
variance

D . F.

MS

F

K i n d r e d x Bay
I n g h a m Exp. I I
MS

T otal

75

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16

21.20

13.59**

30.66

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5^

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4-. 3 6 **

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(P arents)

IS

1.56

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l e v e l of s i g n i f i c a n c e

F v a l u e s e x c e e d s 1% l e v e l o f s i g n i f i c a n c e
1J E n v i r o n m e n t a l e r r o r

F

2 0 . £ 6*#
2.06

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TABLE XVII.
THE EFFECT OF DATE OF HEADING- ON THE AVERAGE
SEED YIELD OF FAMILIES IN F 5 BULKED PROGENIES IN TWO
BARLEY CROSSES GROWN IN INGHAM AND TUSCOLA COUNTIES

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TABLE XV III,
ANALYSIS OF VARIANCE FOR HEIGHT OF PLANT
Fc BULK PROGENIES IN TWO BARLEY CROSSES GROWN IN
INGHAM AND TUSCOLA COUNTIES

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TABLE XIX.
THE EFFECT OF HEIGHT OF PLANTS ON THE AVERAGE
YIELDS OF FAMILIES IN F^ BULKED PROGENIES IN TWO BAR­
LEY CROSSES GROWN IN INGHAM AND TUSCOLA COUNTIES

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TABLE XX. ANALYSIS OF VARIANCE OF LODGING AT
MATURITY IN F 5 BULK PROGENIES IN TWO BARLEY
CROSSES GROWN IN INGHAM AND TUSCOLA COUN­
TIES

CROSSES
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I n g h a m E3sp. I
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MS

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Ingham E x p . I I
MS

F

T o tal

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2 . 23*

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F e x c e e d s X% l e v e l o f s i g n i f i c a n c e

*

F e x c e e d s 5$ l e v e l o f s i g n i f i c a n c e

1/ E nvironm ental e r r o r

TABLE XXI.
THE EFFECT OF LODGING ON THE AVERAGE
YIELD OF FAMILIES IN F£ BULKED PROGENIES IN TWO
BARLEY CROSSES GROWN IN INGHAM AND TUSCOLA
COUNTIES

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TABLE XXII,
ANALYSIS OF VARIANCE FOR LODGING- RESISTANCE
RECORDED TWO WEEKS BEFORE MATURITY IN Fg BULK PROGEN­
IES IN TWO BARLEY CROSSES GROWN IN INGHAM AND
TUSCOLA COUNTIES

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TABLE X X I I I .
THE EFFECT OF AWN TYPE ON THE
AVERAGE YIELDS OF FAMILIES IN F£ BULKED
PROGENIES IN TWO BARLEY CROSSES GROWN
IN INGHAM AND TUSCOLA COUNTIES

Awn t y p e s
Smooth

|

Rough

Mean y i e l d p e r p l o t
i n gram s

E xperim ent

F values

In g h am Exp. I I

K i n d r e d x Bay

412.^6

H-2 0 . 8 J

O.At-O

T u s c o la Exp.

K i n d r e d x Bay

^7 3 • 59

1J4S.69

1.76

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TABLE XXIV.
CALCULATED VALUES OF E, D/2 AND HERITABILITY PERCENTAGES
OF DISEASE REACTION AND AGRONOMIC CHARACTERS OBSERVED IN F£
BULKED POPULATION IN TWO BARLEY CROSSES GROWN IN INGHAM AND
TUSCOLA COUNTIES

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*7.

DISCUSSION
It

i s e v i d e n t t h a t th e g e n e t i c components o f v a r i a n c e

a r e c o m p o s e d o f a d d i t i v e and n o n - a d d i t i v e gene e f f e c t s .

In

o r d e r to s u p p o r t t h i s c o n c e p t th e f o l l o w i n g a p p ro a c h e s to
th e problem w i l l be d isc u sse d ^
1.

The g e n e t i c c o m p o n e n t s o f v a r i a n c e
a s d e te r m i n e d from th e a l g e b r i a c
model.

2.

The g e n e t i c c o m p o n e n t s o f v a r i a n c e
d e t e r m i n e d f r o m t h e F^ b u l k y i e l d
test.

3.

The g e n e t i c c o m p o n e n t s o f v a r i a n c e
a s d e t e r m i n e d fr o m d i s e a s e r e a c t i o n
and o t h e r a g r o n o m i c c h a r a c t e r s .

In the

a l g e b r i a c m odel,

the c a p i t a l l e t t e r s D fo r a d d i­

t i v e e f f e c t s and H f o r n o n - a d d i t i v e
d icate

su m m a t i o n o v e r a l l l o c i .

e f f e c t s were u se d to i n ­

The e x p e c t e d g e n o t y p e s w i t h

t h e i r f r e q u e n c i e s and mean e f f e c t s w e r e shown f r o m t h e F 2
t o t h e F^ g e n e r a t i o n ,

th is

e x p e r im e n t b e in g c o n d u c te d i n Fg.

From t h i s m o d e l t h e f o l l o w i n g two e q u a t i o n s w e r e d e v e l o p e d ,
s h o w i n g t h e f r e q u e n c y o f D an d H i n t h e F^ g e n e r a t i o n s
1 / 2 D -f- 1/102*1-

H = [ £ - * ...................................

3/S D +

H s. ( y - p ................................... ( b )

6/102^

E q u a t i o n ( a ) r e p r e s e n t s t h e mean s q u a r e e x p e c t a t i o n

*1-8,

f o r g e n e t i c e f f e c t s b e t w e e n t h e f a m i l i e s and e q u a t i o n ( b )
is

t h e mean s q u a r e e x p e c t a t i o n f o r g e n e t i c e f f e c t s w i t h i n

the f a m ilie s .

We h a v e s e e n t h a t t h e c o e f f i c i e n t o f D b e ­

tween f a m i l i e s r e m a in s c o n s t a n t from g e n e r a t i o n to g e n e r a ­
tio n .

The c o e f f i c i e n t o f D w i t h i n t h e f a m i l i e s i s

3/ g .

T h is i s e q u a l to 3 / ^ of th e v a l u e f o r betw een f a m i l i e s .
the h e te ro zy g o u s f a m ilie s ,

In

t h e c o e f f i c i e n t o f H g o e s down

o n e - h a l f i n e a c h s u b s e q u e n t g e n e r a t i o n and i t becom es so
low t h a t i t

c a n b e i g n o r e d a t t h e e nd o f F^ g e n e r a t i o n .

c o e f f i c i e n t of H w ith in fa m ilie s i s
t h a t betw een f a m i l i e s .

s ix tim es l a r g e r than

The r e d u c t i o n i n t h e n o n - a d d i t i v e

e f f e c t s i s an i n d i c a t i o n

t h a t t h e f a m i l i e s a r e a l m o s t homo­

z y g o u s a t t h e e n d o f t h e F^ g e n e r a t i o n .
in hom ozygosity,

it

Due t o t h e i n c r e a s e

i s p o s s i b l e t o s e l e c t e f f e c t i v e l y fro m

the p ro g e n ie s a t t h i s s ta g e .
H also f a c i l i t a t e

The

M oreover,

t h e v a l u e s o f D and

the c a l c u l a t i o n of the h e r i t a b i l i t y p e rc e n ­

tag es.
W ith th e h e l p o f th e a l g e b r a i c m odel,

t h e c o m p o n e n ts

c a n b e d i v i d e d i n t o a d d i t i v e a nd n o n - a d d i t i v e e f f e c t s i n any
selected generation.
used.

In th is

r e p o r t t h e F^ y i e l d d a t a was

The e n v i r o n m e n t a l v a r i a n c e was s u b t r a c t e d f i r s t fr o m

b e t w e e n f a m i l y mean s q u a r e s ( T a b l e 1 0 ) .

The v a l u e s o f k

w e r e d e t e r m i n e d i n I n g h a m E x p e r i m e n t s I and I I and T u s c o l a
E xperim ent I I I .

The v a l u e o f k i s ^ . 0 0 i n t h e f i r s t

e x p e r i m e n t s , w h e reas the v a lu e o f k i n
ment I I I

is

3 .68.

The v a l u e o f k i n

two

the T u sc o la E x p e ri­

the l a t t e r e x p e rim e n t

^9.

is

s m a l l e r s i n c e s i x f e w e r f a m i l i e s were c o n t a i n e d i n t h i s

test.

The h e r i t a b i l i t y

v a l u e s w e re d e t e r m i n e d f r o m t h e

fo llo w in g form ula:
2
H e r i t a b i l i t y ;=•

________
t r 2"

The h e r i t a b i l i t y

+

tff-

f o r y i e l d s a r e 3 1 , 37 and 32 p e r c e n t

f o r I n g h a m E x p e r i m e n t s I a n d I I and T u s c o l a E x p e r i m e n t I I I
resp ectiv ely ,
b ility

w i t h a mean v a l u e o f 33 p e r c e n t .

The h e r i t a ­

v a l u e s i n F^ g e n e r a t i o n s a r e r e l a t i v e l y h i g h , w h i c h

c o n f i r m s t h e v i e w p o i n t t h a t s e l e c t i o n i n F^ g e n e r a t i o n w i l l
be v e r y e f f e c t i v e f o r y i e l d .
From t h e a n l y s i s o f y i e l d d a t a ( T a b l e 1 0 ) ,

i t may be

s e en t h a t t h e F v a lu e e x c e e d s th e one p e r c e n t l e v e l of s i g ­
n i f i c a n c e f o r d i f f e r e n c e s betw een f a m i l i e s i n b o t h c r o s s e s
in a l l

th re e experim ents.
The e s t i m a t e o f g e n e t i c c o m p o n e n t s o f v a r i a n c e d u e t o

d i s e a s e and o t h e r agronom ic c h a r a c t e r s p r o v id e a d d i t i o n a l
h e ritab ility

estim ates.

The a m o u n t o f m i l d e w r e c o r d e d i n d i c a t e s t a t i s t i c a l l y
s i g n i f i c a n t d i f f e r e n c e s betw een f a m i l i e s
three

experim ents,

b u t s i g n i f ic a n t d if f e r e n c e s did not e x is t

betw een th e d i f f e r e n t

c l a s s e s ( T a b l e 13) when t h e c l a s s e s ,

b a s e d on d e g r e e o f i n f e c t i o n ,
y ie ld s of seed.

( T a b l e 12) i n a l l

were compared f o r a v e r a g e

T h i s a p p a r e n t l y was d u e to t h e f a c t t h a t

50.

t h e p a t h o g e n , e t c . , was n o t a f f e c t i n g s e e d y i e l d a p p r e c i a b l y .
The a n l y s i s o f v a r i a n c e ( T a b l e 1*0 f o r
in fectio n

shows no s t a t i s t i c a l l y

sativum

sig n ific a n t d ifference

betw een th e f a m i l i e s i n any o f th e t h r e e e x p e rim e n ts ;

a nd

when t h e s e f a m i l i e s w e r e g r o u p e d i n t o c l a s s e s o f i n f e c t i o n
a n d c o m p a r e d f o r a v e r a g e s e e d y i e l d s , no s i g n i f i c a n t d i f f e ­
r e n c e s were o b t a i n e d .

T h is i n d i c a t e s t h a t th e p a th o g e n had

no d i f f e r e n t i a l e f f e c t o n y i e l d .
F iv e agronom ic c h a r a c t e r s , d a t e o f h e a d in g , h e i g h t o f
p l a n t , p e r c e n t o f lodged p l a n t s ,

lodging r e s is ta n c e (cL r),

a n d s m o o t h v e r s u s r o u g h awn, w e re i n c l u d e d w h i c h p r o v i d e
estim a te s of h e r i t a b i l i t y .
D a t e o f h e a d i n g ( T a b l e 16) and h e i g h t d a t a ( T a b l e 15)
in d ic a te h ig h ly
the f a m i l i e s .

s i g n i f i c a n t d i f f e r e n c e s e x i s t i n g betw een
The f a m i l i e s w e r e c l a s s i f i e d i n t o f o u r g r o u p s

on t h e b a s i s o f d a t e o f h e a d i n g and a v e r a g e s e e d y i e l d s o f
each group were com pared.

N© s t a t i s t i c a l l y

sig n ifican t

d i f f e r e n c e b e t w e e n t h e s e g r o u p s was f o u n d ( T a b l e 1 7 ) .

The

h e i g h t , h o w e v e r , was a s s o c i a t e d w i t h s i g n i f i c a n t d i f f e ­
re n c e s in th e av erag e y i e l d s of the f a m i li e s (T able 19).
The t a l l e r f a m i l i e s t e n d e d t o p r o d u c e g r e a t e r y i e l d s o f
seed.
S i g n i f i c a n t d i f f e r e n c e s in lodging p e rc e n t e x is te d
b e t w e e n f a m i l i e s ( T a b l e 20) a n d w i t h i n f a m i l i e s i n
d r e d x Bay c r o s s ,

but s t a t i s t i c a l l y

were n o t d e t e c t e d i n

th e Kin­

sig n ific a n t differences

t h e S t e w a r t x Bay c r o s s .

The f a m i l i e s

51.

were g ro u p ed I n t o

f o u r c l a s s e s on t h e b a s i s o f p e r c e n t a g e

o f l o d g e d p l a n t s and t h e s e

c l a s s e s were compared f o r th e

a v erag e y i e l d s o f seed (T able 21).

No s i g n i f i c a n t d i f f e ­

r e n c e s were n o t e d .
It is
y ield .

concluded t h a t lo d g in g d id n o t e f f e c t

the seed

T h i s may b e d u e to t h e f a c t t h a t t h e l o d g i n g o c c u r ­

r e d o n l y s h o r t l y b e f o r e t h e p l a n t s m a t u r e d and t h e h a r v e s t ­
i n g m ethods u se d p e r m i t t e d

the re c o v e ry o f th e lodged as

w e ll as the e re c t p la n ts .
D i f f e r e n c e s betw een f a m i l i e s were h i g h l y s i g n i f i c a n t
f o r lod ging r e s is ta n c e

( c L r ) f a c t o r i n Ingh am E x p e r i m e n t I

and T u s c o la E x p e rim e n t I I I .

I n In gh am E x p e r i m e n t I I ,

the

d i f f e r e n c e s were s i g n i f i c a n t a t t h e 5 p e r c e n t l e v e l .
The a v e r a g e y i e l d s o f s m o o t h and r o u g h awn f a m i l i e s
w e r e c o m p a r e d and no s i g n i f i c a n t d i f f e r e n c e s b e t w e e n f a m i ­
l i e s e x iste d in

t h e K i n d r e d x Bay c r o s s .

The e s t i m a t e s o f h e r i t a b i l i t y

I n a l l t h e d i s e a s e and

a g r o n o m i c c h a r a c t e r s a r e g i v e n i n T a b l e 2^ a l o n g w i t h t h e
v a l u e s o f e n v i r o n m e n t a l v a r i a n c e s and a d d i t i v e
effects.

g en etic

The h e r i t a b i l i t y o f h e a d i n g d a t e b e t w e e n f a m i ­

l i e s r a n g e s f r o m 76# t o 2 3 ^ , w i t h an a v e r a g e o f
l i n e w i t h F r e y 8s r e s u l t s ( 2 0 ) ,

In

th is high h e r i t a b i l i t y e s t i ­

m a t e was n o t u n e x p e c t e d .
The e s t i m a t e s o f h e r i t a b i l i t y
ra n g e from bo$ to
experim ents.

f o r m il d e w i n d i c a t e a

w i t h a mean o f 6 2 ^ f o r t h e t h r e e

L odging h e r i t a b i l i t y
r a n g e from 16# t o 35#.

e s t i m a t e s f o r t h e two c r o s s e s

The c o n t r a s t i n g e s t i m a t e s a r e u n ­

d o u b t e d l y d u e to d i f f e r e n c e s i n
S t e w a r t x B ay c r o s s ,
straw ed.

t h e two c r o s s e s .

In the

both p a re n ts a re r e l a t i v e l y s t i f f

I n K i n d r e d x Bay c r o s s , K i n d r e d i s

a w eak-etraw -

e d p a r e n t a nd Bay a s t r o n g - s t r a w e d p a r e n t , r e s u l t i n g I n a
h igher g e n etic variance.
H eight d a ta i n d i c a t e s r e l a t i v e l y uniform h e r i t a b i ­
l i t y p e r c e n t a g e s i n b o t h th e c r o s s e s and t h e r a n g e i s from

3 5# to

bsfl.
The h e r i t a b i l i t y

r a n g e f o r l o d g i n g r e s i s t a n c e i s 1 7#

t o 6 2 # , w i t h a mean o f ^ 2# i n b o t h t h e c r o s s e s ;
t e s t w eight,

th e range i s

7^ p e r c e n t .

The h e r i t a b i l i t y e s t i m a t e s f o r

whereas,

in

55 t o 27 p e r c e n t , w i t h a mean o f
sativum a re

12 t o 32 p e r c e n t .
These e s t i m a t e s of h e r i t a b i l i t y a r e r e l a t i v e l y h ig h ,
and i t

i s p o s s i b l e t o make a n e f f e c t i v e

s e le c tio n in the

g e n e r a t i o n b e c a u s e h o m o z y g o s i t y I s a p p r o a c h i n g a maximum.
D e l a y l a t e r t h a n t h i s u n d o u b t e d l y would b e a w a s t e o f t i m e
on t h e p a r t o f t h e p l a n t b r e e d e r .

53.

CONCLUSIONS
In a l l

s e g re g a tin g g e n e ra tio n s the a d d itiv e g e n e tic

f r a c t i o n s re m a in o n e - h a l f and th e n o n - a d d i t i v e g e n e t i c
f r a c t i o n betw een f a m i l i e s i s h a lv e d in each su cceed in g
generation.

The n o n - a d d i t i v e g e n e t i c f r a c t i o n s i n t h e

g e n e ra tio n approach zero .

From t h i s i t i s

concluded

th a t w ith th e re d u c tio n of n on -additive gen etic fr a c tio n
t h e h o m o z y g o s i t y i n F^ h a s r e a c h e d a p r a c t i c a l maximum.
The h e r i t a b i l i t y p e r c e n t a g e s f o r y i e l d d a t a a r e 3 1 ,
37 and. 3 2 f o r t h e t h r e e r e p o r t e d e x p e r i m e n t s .

T hese

v a l u e s a r e r e l a t i v e l y h i g h and s u g g e s t t h a t s e l e c t i o n f o r
y i e l d i s m o s t e f f e c t i v e i n F^ g e n e r a t i o n .
From p o w d e r y m i l d e w d a t a i t

c an b e c o n c l u d e d t h a t t h e

p a t h o g e n was a f f e c t i n g t h e s e e d y i e l d s i n a l l f a m i l i e s t h e
same.

T h e r e was n o t a h i g h t y p e o f r e s i s t a n c e a s a l a r g e

p r o p o r t i o n o f t h e f a m i l i e s h a d more t h a n 25 p e r c e n t i n f e c ­
tio n .
H.

s a t ivum i n f e c t i o n d i d n o t p r o d u c e d i f f e r e n t i a l

e f f e c t s on t h e a v e r a g e s e e d y i e l d s o f t h e f a m i l i e s .
T h e r e was no e f f e c t o f d a t e o f h e a d i n g on t h e a v e r ­
age y i e l d o f th e f a m i l i e s .
be a s s o c i a t e d w i t h y i e l d ,

How ev er,

t h e h e i g h t se ems t o

as t a l l e r f a m i l i e s produced

g rea ter y ield s.
Seed y i e l d s o f d i f f e r e n t f a m i l i e s w e r e n o t a f f e c t e d
by t h e l o d g i n g w h ic h o c c u r r e d i n t h i s e x p e r im e n t.

5^.

No d i f f e r e n c e s i n y i e l d w e r e n o t e d w h i c h m i g h t h a v e
b e e n a s s o c i a t e d w i t h r o u g h o r sm o o th awn c h a r a c t e r .
The h e r i t a b i l i t y v a l u e s o f t h e s e d i s e a s e r e a c t i o n
and a g ro n o m ic c h a r a c t e r s c o n firm th e v ie w p o in t re a c h e d
f o r h e r i t a b i l i t y of y i e l d ; nam ely,
tio n ,

t h a t b y t h e F^ g e n e r a ­

g e n e t i c g a in from s e l e c t i o n w i l l be a p p ro a c h in g a

maximum.

55.

SUMMARY
To c o n d u c t y i e l d t r i a l s

i n t h e F^ g e n e r a t i o n ,

b u l k e d f a m i l i e s f r o m two b a r l e y c r o s s e s ,

tw enty

S t e w a r t x Bay and

K i n d r e d x B a y , w e r e s e l e c t e d i n t h e F^ g e n e r a t i o n .

E igh­

t e e n F ^ s e e d s w e r e t a k e n a t random fr o m e a c h o f t h e ^t-0
F ^ b u l k e d r o w s i n c r e a s e d f o r two g e n e r a t i o n s i n t h e g r e e n
h o u s e i n 1952 and th e f i e l d

in 1953*

The p e d i g r e e s o f

f a m i l i e s a n d s u b - f a m i l i e s w e re m a i n t a i n e d .
Three e ^ e r i m e n t s ,

two i n Ing h am C o un ty and one i n

T u s c o la C ounty, were c o n d u c te d .
sin g le,

The t e s t s w e r e on a

four-row p l o t b a s is in a l a t t i c e design.

E n v i r o n m e n t a l e r r o r was c a l c u l a t e d fro m t h e p a r e n t a l
checks.
tio n

The f o l l o w i n g c h a r a c t e r s w e r e s t u d i e d i n a d d i ­

to yield:,

powdery m ildew ,

spot b lo tc h , d a te of head­

i n g , h e i g h t o f p l a n t , p e r c e n t o f lodged p l a n t s ,
resistan ce,

a n d awn t y p e .

lodging

The e f f e c t o f e a c h o f t h e s e

c h a r a c t e r s on s e e d y i e l d o f f a m i l i e s was s t u d i e d .
An a l g e b r a i c m o d e l f o r t h e e s t i m a t i o n o f
was i n c l u d e d .

and

The h e r i t a b i l i t y v a l u e s f r o m s e e d y i e l d and

o t h e r c h a r a c t e r s w e r e c a l c u l a t e d f r o m t h e f o l l o w i n g formu=
la.
x 100.

H e rita b ility

These h e r i t a b i l i t y p e r c e n t a g e s f o r y i e l d and o t h e r c h a r a c ­
t e r s w e re r e l a t i v e l y h i g h a n d i n d i c a t e d
l e c t i o n can be p r a c t i c e d i n

th at e ffe ctiv e

t h e F^ g e n e r a t i o n .

se­

56.

LITERATURE CITED
1.

A tkins, I .
M.
R elatio n of c e rta in p la n t c h arac ters
to s t r e n g t h o f s tr a w and lo d g in g i n w i n t e r wheat.
J o u r . A g r . R e s. 56 s 9 9 - 1 2 0 .
1 9 3$.

2.

A tkins, I .
M.
I n h e r ita n c e of straw s tr e n g th p e r u n it
l e n g t h of culm and o t h e r c h a r a c t e r s i n Kanred x
Coppei w h e a t.
J o u r . Agr. Res. 7 6 s 53-72.
194$.

3.

A t k i n s , R. E. and H. C. Murphy.
E valuation of y ield
p o t e n t i a l i t i e s of o a t c r o s s e s from b u lk h y b r id t e s t s .
A g r o n . J o u r . 4 l ; ^-1—^ 5 .
19^9 .

4.

B a k k e , A. L.
The l a t e b l i g h t o f b a r l e y {H e l m i n t h o s p o rium t e r e s s a c c ) .
P r o c . Io w a Acad. S c i . 1 9 2 9 3 - 1 0 2 .
1912.

5.

B o s e , R. D . , M. A. A z i z ,
a n d M. P . B h a t n a g a r .
The i n ­
h e r i t a n c e o f some a n a t o m i c a l c h a r a c t e r s r e s p o n s i b l e
f o r l o d g i n g and some e a r h e a d c h a r a c t e r s i n an i n t e r ­
s p e c i f i c c r o s s b e t w e e n two p u s a b a r l e y s .
Ind. Jour.
A g r . S c i . 7s 4 S - S S .
1937.

6.

B r i g g s , N. F.
The i n h e r i t a n c e o f r e s i s t a n c e o f
Am. N a t . 72s 3 4 - 4 l .
1938.

m il d e w .

7.

Brown, A. R. and J . H. M i l l e r .
A com parative study of
H e l m i n t h o s p 0 r i u m s a t i v u m Pam, K in g a n d Bakke a n d H.
v i c t o r i a e Meehan a n d Murphy on o a t s .
Agron. J o u r .
4'6V '6 3- M .
1954.

$.

C h a r l e s , D. R. and H. H. S m i t h .
D i s t i n g u i s h i n g between
two t y p e s o f gene a c t i o n i n q u a n t i t a t i v e i n h e r i t a n c e .
G e n e t i c s 24; 3 4 - 3 ^ .
1939-

9.

C h risten sen , J. J.
S t u d i e s i n t h e p a r a s i t i s m o f H^
sativum .
Minn. A g r . E x p t . S t a . T e c h . B u i . 11s 4 2 - p p .
1922.

10.

C l a r k , R. E. a n d H. K. W i l s o n .
Lodging i n a l l g r a i n s .
J o u r . Amer. S o c . A g r o n . 25; 56.1-572.
1933•

11.

C o c h r a n , W. G.
Long t e r m a g r i c u l t u r a l e x p e r i m e n t s .
J o u r . Roy. S t a t . S o c . Supp. 6 : 1 0 4 - l 4 $ .
1949.

12.

C o m s t o c k , R. E. a n d H. E. R o b i n s o n .
The c o m p o n e n t s o f
g e n e tic v a ria n c e in p o p u la tio n of b i - p a r e n ta l progen­
i e s and t h e i r u s e i n e s t i m a t i n g the average d eg ree of
dom inance.
B i o m e t r i c s 4; 2 5 4 - 2 6 6 .
194$.

57

13.

C o m s t o c k , R. E . , H. E. R o b i n s o n and P . H. H a r v e y .
A b r e e d i n g p r o c e d u r e d e s i g n e d t o make maximum u s e
o f b o th g e n e r a l and s p e c i f i c combining a b i l i t y .
A g r o n . J o u r , *4-1*. 360- 3 6 7 .
19*4-9.

1*4-.

Crump, S, L.
The e s t i m a t i o n o f v a r i a n c e c o m p o n e n ts
in a n ly s is of v ariance.
B io m e tr ic s B ui. 2 ; 7-11
19^6.

15.

E a s t , E. M.
S t u d i e s on s i z e i n h e r i t a n c e i n N i c o tia n a .
G e n e t i c s 1; 16*1—1 7 6 .
1916.

16.

F i s h e r , R. A.
The c o r r e l a t i o n among r e l a t i v e s on
th e s u p p o s i t i o n s of M endelian i n h e r i t a n c e .
Trans.
R o y a l S oc . E d i n b u r g h . L I I : 39 9—*+33.
1918.

17.

F i s h e r , R. A.
S t a t i s t i c a l m e t h o d s f o r r e s e a r c h work­
ers.
O l i v e r a n d Boyd, E d i n b u r g h ( 1 9 2 5 “ 19*4-**).

IS.

F i s h e r , R. A . , F . R. Immer, and 0. T e d i n .
The g e n e t i c a l I n t e r p r e t a t i o n o f s t a t i s t i c s of the t h i r d degree
i n th e study of q u a n t i t a t i v e i n h e r i t a n c e .
G enetics
17; 107-12*4-.
1932.

19.

F l o r e l l , V. H.
n e ss i n w heat.

20.

Frey,
d ate

2 1.

S t u d i e s on t h e i n h e r i t a n c e o f e a r l i ­
J o u r . A g r . R e s . 29: 333—3^7192*4-.

K. J .
I n h e r i t a n c e a n d h e r i t a b i l i t y of h e a d i n g
in b arley.
A g r o n . J o u r . *4-6: 2 2 6 - 2 2 8 .
195^*

G a r b e r , R. J . a nd P . J . O l s o n .
A stu d y of the r e l a ­
t i o n s h i p o f some m o r p h o l o g i c a l c h a r a c t e r s t o l o d g i n g
in c e re a ls.
J o u r . Araer. Soc. A g r o n . 11; 1 7 3 - 1 8 6 .
1919.

22.

G r a f i u s , J . E.
A s t a t i s t i c a l m odel f o r e s t i m a t i n g
com ponents o f g e n e t i c v a r i a n c e in b u lk y i e l d t e s t s
of s e l f p o l l i n a t e d sm all g r a i n s .
S o u t h Dak. Agr.
Exp. S t , Tech.
B u i . No 9 . 1952.

23.

G r a f i u s , J . E . , and H. M. Brown.
L o d g in g r e s i s t a n c e
in o a ts .
A g r o n . J o u r . *4-6: *4-1*4—*1-18.
1951*-*

2*1-.

G r a f i u s , J . E. , W. L. N e l s o n a n d V. A. D i r k s .
The
h e r i t a b i l i t y o f y i e l d i n b a r l e y a s m easured by e a r l y
g e n e r a tio n bulked p ro g e n ie s .
A g r o n . J o u r . *4-*4: 253 257.
1952.

25.

G r i f f e e , F.
C o rrelated in h e rita n c e of b o ta n ic a l
c h a r a c t e r s i n b a r l e y a n d m a n n e r o f r e a c t i o n t o H.^
s a t i v u m . J o u r . A g r . R e s . 15: 9 1 5 —935*
3-925*

.

G r i f f i n g , J . B.
S t a t i s t i c a l a n a l y s i s o f g e ne a c t i o n
i n com ponents of tom ato y i e l d .
U n p u b l i s h e d PH.D. t h e s i s .
Ames, I o w a .
194-2.

Iowa S t a te C o lleg e L ib r a r y ,

H a r l a n , H. V. , M. L. M a r t i n i a n d S. A. H a r l a n d .
A
s t u d y o f method i n b a r l e y b r e e d i n g .
U . S . D . A . Tech.
B u i . No 7 2 0 .
1940.
H a r r i n g t o n , J . B. P r e d i c t i n g t h e v a l u e o f a c r o s s
f r o m an F 2 a n a l y s i s .
Can. J o u r . R e s . 6: 2 1 - 3 7 .
1932.
H a r r i n g t o n , J . B.
Y ie ld in g c a p a c ity o f wheat c ro s s e s
a s i n d i c a t e d by b u l k h y b r i d t e s t s .
Can. J o u r . R e s. 12:
572-524.
1 94 0 .
H a y e s , H. K . , E. C. S t a k e m a n , F . G r i f f e e a n d J . J .
C hristensen.
R e a c ti o n of b a r l e y v a r i e t i e s to Helmin. Minn. A g r . E x p t . S t a . TecE". B u i .
pp.
1923.
Hehn, E.
The i n h e r i t a n c e o f a g r o n o m i c c h a r a c t e r s i n
b arley .
U n p u b l i s h e d PH. D. t h e s i s .
1952.
Iowa S t a t e C o lle g e L ib r a r y .
Ames, Io w a .
H o r - K v e n , S.
I n t e r a c t i o n of g e n e t i c f a c t o r s in b a r l e y .
G e n e t i c s 9; 1 5 1 - 1 2 0 .
1924.
H u l l , H.
R e c u r r e n t s e l e c t i o n f o r s p e c i f i c combining
a b i l i t y in corn.
J o u r . Am. S oc . A g r o n . 3 7 : 1 3 4 - 1 4 5 .
19^5Im mer, F . R.
R e l a t i o n b e t w e e n y i e l d i n g a b i l i t y and
horaozygosis i n b a r l e y c r o s s e s .
J o u r . Amer. So c. Agron.
33: 2 0 0- 2 0 6 . 1 9 4 1 .
Im m er, F. R.
R e l a t i o n b e t w e e n y i e l d i n g a b i l i t y and
hom ozygosis i n b a r l e y c r o s s e s .
J o u r . Amer. Soc.
A g r o n . 3^» 2 4 4 - 2 5 0 .
1942.
J e n k i n s , M. T.
The s e g r e g a t i o n o f g e n e s a f f e c t i n g
y i e l d o f g r a i n i n m aize.
J o u r . Amer. S o c . A g r o n .
32: 55-63.
1940.
J o h a n n s o n , W. E l e m e n t e d e r e x a k t e n E r b b e h k e s t s l e h r e
gene.
G ustav.
F ish er.
1909.
K a l t o n , R. R.
B reeding b eh av io r a t su c ce ssiv e genera­
tio n s fo llo w in g h y b r id iz a t io n in soybeans.
Iow a
E x p t . S t a . R e s. B u i .
3 52 .
1942.

59.

39*

L erner,
ment.

J.
P o p u l a t i o n g e n e t i c s and a n i m a l i m p r o v e —
Cambridge a t t h e U ni. P r e s s ,
1950.

40.

L o v e , H. H. A p r o g r a m f o r s e l e c t i n g and t e s t i n g
sm all g r a i n s in su c c e ssiv e g e n e ra tio n s fo llo w in g
h y b rid izatio n .
J o u r . Amer. S o c . A gro n . 19; 7 0 5 712.
1927.

41.

L u s h , J . L. A n i m a l b r e e d i n g p l a n s .
The
P r e s s I n c . Ames, I o w a .
2nd E d i t i o n pp

42.

L u s h , J . L. H e r i t a b i l i t y o f q u a n t i t a t i v e c h a r a c t e r s
in f a r a anim als.
P r o c . E i g h t h . I n t . Cong. Gene.
194S.
H e r e d i t a s S u p p l . V o l. 356- 3 7 5 .
1949.

C o lleg iate
76- 7 9 . 1943.

43.

L u s h , J . L. a nd V. G-. P a n s e .
The i n h e r i t a n c e o f
q u a n t i t a t i v e c h a r a c t e r s and p l a n t b r e e d i n g .
Jour.
G enetics.
40; 2 & 3 -3 0 2.
1940.

44.

L u t t r e l l , E. S.
A c o m p a r i s o n o f H. s a t i v u m and H.
v i c t o r i a e ( A b s ) P h y t o p a t h . 4-2: 4 £ § - 4 7 5 .
1 9 52 .

^5.

M a i n s , E. B. and
M. L. M a r t i n i .
S u s c e p t i b i l i t y of
b a r l e y to l e a f r u s t ( P u c c i n i a a n o m a l ) and t o pow­
d e r y m i l d e w ( E r i s i p h e gram i n i s Hor'de'l ?

46.

M a r t i n , G. W., C. R. Weber a n d R. R. K a l t o n .
E arly
g e n e ra tio n t e s t i n g in soybeans.
Amer. S oc . A g ron .
39; 7 9 1 - 2 1 2 . 1 9 4 7 .

47.

M a t h e r , K.
B iom etrical g e n e tic s.
& Co. 162 p p .
19^9*

43.

M eehan, P . a n d H. C. Murphy.
A new H e l m i n t h o s p o r i u m
b lig h t of o a ts.
S c i e n c e , 104; 4 l 3 - 4 l 4 .
1946.

49.

M urp h y, H. C. a n d F . Meehan.
R e a c tio n of o a t v a r i e ­
t i e s t o a new s p e c i e s o f H e l m i n t h o s p o r i u m .
Phyto­
p a t h . 361 407 PP.
1946.

5 0.

Muhmud, Imam and H. H. Kramer.
S egregation f o r
y i e l d , h e i g h t and m a t u r i t y f o l l o w i n g a s o y b e a n
cross.
A g r o n . J o u r . 4 3 ; 605- 6 0 9 . 1 9 5 1 .

51 .

M y l e r , J . L. Awn i n h e r i t a n c e i n b a r l e y .
Res. 6 5 ; 4 0 5 - 5 1 2 .
19^2.

5 2.

London.

Methuen

Jour.

Agr.

N e a t b y , K. W. I n h e r i t a n c e of q u a n t i t a t i v e a n d o t h e r
ch aracters in b arley crosses.
S c i . Agr. f i 7 7 - S 4 .
1926.

6 o.

53.

N e a t b y , K. W. An a n l y s i s o f q u a n t i t a t i v e c h a r a c ­
t e r s and lin k a g e in b a rle y .
S c i . A gr. 6 s 7 0 1 - 7 1 g .
1926.

5^.

N e a t b y , K. W. An a n a l y s i s o f t h e i n h e r i t a n c e o f
q u a n t i t a t i v e c h a r a c t e r s and l in k a g e i n b a r l e y .
S c i. Agr. 9 s 701- 7 1 7 . 1929.

55.

56.

N e w to n , M a r g a r e t a n d W. j . C h e r e w i c k .
Er y s i p h e
g r a m i n i s i n C a n a d a . Can. J o u r . R e s. ( CT 25s 73 - 9 3 .
W T "
Paramel, L. H. W heat b l i g h t a n d b r o w n i n g
Breeders* G a z e tte .
56- 1 5 5 . 1 9 0 9 .

of

b arley.

57* Paramel, L. H . ,

C. M. K i n g and A. L. B a k k e .
Two b a r ­
l e y b l i g h t w i t h a co m p ariso n of s p e c i e s o f Helminth o sp o riu ra upon c e r e a l s .
I o w a Agr. Exp. S t a . B u i .
116. 1910.

53.

Panse,
ing.

59-

Pow ers, Leroy.
The n a t u r e o f t h e s e r i e s o f e n v i r o n ­
m e n t a l v a r i a n c e s a nd t h e e s t i m a t i o n o f t h e g e r t e t i c
v a r i a n c e a n d t h e g e o m e t r i c m ea n s o f c r o s s e s i n ­
v o lv in g sp e c ie s of L ycopersicon.
G e n e t i c s 2 7 ‘» 561-

575-

V. G.
Jour.

A p p l i c a ti o n o f g e n e t i c s to p l a n t b re e d ­
G e n e t i c s *K)s 2^ 3” 3 0 2 .
19*K).

19^2.

60.

Pow ers,
L e r o y , L„ F. Locke and J . C. G a r r e t t .
P arti­
t i o n i n g m eth o d o f g e n e t i c a n a l y s i s a p p l i e d t o quan­
t i t a t i v e c h a r a c t e r s of tom ato c r o s s e s .
U.S.D.A.
T e c h . B u i . 9 9 $ . 56 p p .
1950.

6 1.

R araiah, K. a n d S. D h a r m a l i n g a m . L o d g i n g o f s t r a w
an d i t s i n h e r i t a n c e i n r i c e .
I n d i a n J o u r . Agr.
S c i e n c e ki S S O - ^ .
193^.

62.

R a y , D.
A . , T. T. H e r b e r t and G. K. M i d d l e t o n .
In­
h e r i t a n c e o f r e s i s t a n c e t o p o w d e r y m ild e w i n w h e a t .
Agron. J o u r .
37 9 ~ 3 ^3 195^*

63.

R o b e r t s o n , D. W.
t i c s 1^ ; 1- 3 6 .

6^ .

R o b e r t s o n , D. W ., G. A. Wiebe a n d F . R. Immer.
A
summary o f o f l i n k a g e s t u d i e s i n b a r l e y .
Jour.
Amer. S o c . A g r o n . 33? *+7-6*1.
19*+1.

Linkage s t u d i e s
1929•

in b a rle y .

Gene­

61

65.

R o b e r t s o n , D. W. , 0 , A. Wiebe a n d F . R. Immer.
A
summary o f l i n k a g e s t u d i e s i n b a r l e y .
Supp. I
1940-1946.
J o u r . Amer. S oc . A g r o n . 3 9 s 4 6 4 - 4 7 3 .
19^7.

66.

R o b i n s o n , H. F . , R. E. Comstock and P . M. H a r v e y .
E s t i m a t e s o f h e r i t a b i l i t y a n d t h e d e g r e e o f d o m in ­
ance in corn.
A g r o n . J o u r . 4-1; 3 5 3 - 3 5 9 .
19*1-9.

67.

S a t t e r t h w a i t e , F . E.
An a p p r o x i m a t e d i s t r i b u t i o n
of v a r i a n c e com ponents.
B i o m e t r i c s B u i . 2s 1 1 0 114.
1946.

66.

S p r a g u e , G. F. a n d A. A. B r y a n .
The s e g r e g a t i o n o f
g e n e s a f f e c t i n g y i e l d , p r e p o t e n c y , l o d g i n g and
d i s e a s e r e s i s t a n c e i n F-, l i n e s o f c o r n .
J o u r . Amer.
S o c . A g r o n . 335 2 0 7 - 2 1 4 '
1941.

69.

S p r a g u e , G-. F. , L. A. Tatum.
G-eneral V/S s p e c i f i c
com bining a b i l i t y i n s i n g l e c r o s s e s o f c o rn .
Jour.
Amer. S o c . A g r o n .
3^ 9 2 3 “ 9 3 2 .
1 94 2 .

70.

S p r a g u e , 0 . F.
E arly te s tin g of
corn.
J o u r . Amer. S o c . A g r o n .

71.

S n e d e c o r , 0 . W.
S t a t i s t i c a l methods,
Ames, Iowa.
The C o l l e g i a t e P r e s s I n c .
Ed. 4.
1 946.

7 2.

S t a n f o r d , H. E. a nd N. F. B r i g g s .
Two a d d i t i o n a l
f a c t o r s f o r r e s i s t a n c e t o m il d e w i n b a r l e y .
Jour.
A g r . R e s . 6l : 2 3 1- 2 3 6 . 1 94 0.

73.

S u n e s o n , C. A. and G. A. W ie b e.
S u r v i v a l of b a r l e y
and wheat v a r i e t i e s i n m ix t u r e s .
J o u r . Amer. Soc.
A g r o n . 34s 1 0 5 2 - 1 0 5 6 . 1 9 ^ 2 .

inbred lin e s of
3 ^ IO6- I I 7 .
1946.

74.

W e a v e r , J . C.
American b a r l e y p r o d u c t i o n .
P u b l i s h i n g Co. M i n n e a p o l i s .
1950*

Burgess

75.

W ebe r, C. R.
I n h e r i t a n c e and i n t e r a c t i o n of some
a g r o n o m i c a n d c h e m i c a l c h a r a c t e r s i n an i n t e r ­
s p e c i f i c c ro ss in soybeans.
G l y c i n e max X G.
u ssu rien sis.
I o w a A g r. E x p t . S t a . R e s. B u i . 374s
765 - 6 1 6 . 1 9 5 0 .

76.

Weber, C. R. and B. R. M o o r t h y .
H e r i t a b l e and n o n h e r i t a b l e r e l a t i o n s h i p s and v a r i a b i l i t y o f o i l con­
t e n t a nd a g r o n o m i c c h a r a c t e r s i n t h e F2 g e n e r a t i o n
o f soybean c r o s s e s .
A g r o n . J o u r . 44s
202-209.
1952.

.

62.

77.

W e i s s , M. G. , C. R. Weber and R. R. K a l t o n .
E arly
g e n e r a tio n t e s t i n g in soybeans.
J o u r . Amer. Soc.
A g r o n . 39; 7 9 1 - 8 1 1 .
1947.

78.

W e x e l s e n , H.
Q u a n t i t a t i v e i n h e r i t a n c e and l i n k a g e
in b arley .
H e r e d i t a s 18; 3 ° 7 ” 3 i|-8.
1933*

79.

Wright, Sewall.

The an alysis o f variance and corre­

l a t i o n s betw een r e l a t i v e s w i t h r e s p e c t to d e v i a ­
t i o n s f r o m an o ptim um .
Jour. G enetics.
243-

256.

1935.

80.

Y a t e s , F , and I . Zacopanay.
The e s t i m a t i o n o f t h e
e f f i c i e n c y o f sam pling f o r y i e l d in c e r e a l e x p e r i ­
ment. J o u r . A g r . S c i . 2 5 : 5^5-577* 1935*

8 1.

Y ouden, W. J .

a nd A. M e h l i c h .

S election of e f f ic ie n t

methods for s o i l sampling. Contributions from Boyce
Thompson I n s t it u t e 9» 59”70. 1937*