Inheritance of awnedness, kernel smoothness, kernel length, and reaction to two physiologic races of
Tilletia in a spring wheat cross
by Orlie W Smith
A THESIS Submitted to the Graduate Committee in partial fulfillment of the requirements for the
Degree of Master of Science in Agronomy at Montana State College
Montana State University
© Copyright by Orlie W Smith (1941)
Abstract:
TMs investigation is an inheriatance study of awnedness, kernel smoothness, kernel length, and mode
of reaction to two physiologic races of Tilletia in the progeny of a cross between
Ceres-(Hope-Florence) and Renown. Ceres-Hope-Florence) is fully awned (awn type 5)? moderately
resistant to two physiologic hunt races (T-11 of Tilletia tritioi and L-3 of T. levis_), and has kernels
with the following characteristics: cheeks, mostly angular and knife-edged; crease, mid-wide to wide,
mid-deep to deep, and mostly pitted; and an average length of about 6.8 millimeters* Renown is
weakly anwletted (awn type 3-), more resistant to the two bunt races, T-11 and L-3, than
Ceres-(Hope*Florence), and-has kernels with the following characteristics: cheeks, mostly rounded to
semi-angular; crease, narrow to mid-wide, shallow to mid-deep, and usually closed but some may be
slightly pitted; and an average length of about 6.4 mm.
One major factor was found to govern the awn development between the parents of this cross with the
awnletted condition being incompletely dominant to the awned.
The F2 and F3 were classified into eight classes depending upon the degree of roundness or angularity
of cheeks, depth and width of crease, and pittedness of the kernels. A uniform breeding difference was
demonstrated for the parents and the prog ny segregated for a complete range of an between the parents
but no definite factorial analysis was established.
A total length of fifteen selected kernels from each plant were measured in millimeters to determine
kernel length differences. In the F2 a very regular segregation seemed to be indicated with a weak
dominance ox the smoother types suggested. The parents and F3, were so variable it was impossible to
check the F2 classification by the F3 breeding behavior.
Bunt infections in the parents, progeny, and susceptible check, Ulka, were so slight it was decided from
observational data that a differential reaction of the progeny could not be demonstrated. V
IWHERI TM CE OP AMEDWESS, KEEEEL SMOOTHERSS , KEEEEL LEEGTH, AED
REACTIOE TO TWO PHYSIOLOGIC RACES OP TILLETIA IE A SPRIEG WHEAT CROSS
Ly
.
O r lie W. S m ith
A THESIS
S u b m itte d t o th e G ra d u ate Committee
i n p a r t i a l f u l f i l l m e n t o f th e re q u ire m e n ts
f o r th e D egree o f M a s te r o f S c ie n ce
i n Agronomy a t
M ontana S t a t e C o lle g e
A pproved:
I n C harge o f M ajor Work
C hairm an E xam ining Committee
C hairm an G ra d u ate Committee
Bozemen, M ontana
Ju n e 1941
I
'-V// Y
' cii Hj
:»j/
'tl'
■<)
ACKN0V9LED03MFZTS
The w r i t e r w ish e s t o e x p r e s s h i s a p p r e c i a t i o n t o
D r s . E . H. Bamberg and A. M. S c h le h u b e r u n d e r whose d i r e c t i o n
th e i n v e s t i g a t i o n was m ade; t o S . C. L i t z e n b e r g e r , A s s i s t a n t in
Agronomy, f o r h e lp and s u g g e s tio n s w ith th e aw nedness s tu d y ; to
P r o f e s s o r A. H. P o s t f o r s u g g e s tio n s i n p r e p a r a t io n o f th e manu­
s c r i p t ; t o L . A. C la r k , D iv is io n o f F o rag e C rops and D is e a s e s ,
and D r. F. B. C o tn e r , P r o f e s s o r o f B otany and B a c te r io lo g y , f o r
h e lp and s u g g e s tio n s w ith th e p i c t u r e s ; and to th e D iv is io n o f
C e re a l C rops and D is e a s e s , U. S. D epartm ent o f A g r i c u l t u r e , f o r
f i n a n c i a l a s s i s t a n c e th ro u g h th e R e s e a rc h F e llo w s h ip i n Agronomy.
117292
' -III-
TABLE OP CONTENTS
LIST OP PIGU EES.....................................................................................................
Page
IV
LIST OF T A B L E S ............................................................ ■........................................
V
ABSTRACT..................................... "...........................................................................
VI
INTRODUCTION.............................................
I
■LITERATURE REVIEW.................................................
Awnedness
4
....................................
K e rn e l Sm oothness
4
...................................................................................
K e rn e l L en g th
.......................
. . . .
Bunt R e a c t i o n .............................................
8
8
9'
MATERIALS AND METHODS..........................................' . . ..................................
10
EXPERIMENTAL RESULTS
15
.............................................................................. . . .S-
.............................................
15
K e rn e l S m o o t h n e s s ...........................
17
aw nedness
K e rn e l L en g th
................... . .....................................
. . . . . .
31
Bunt R e a c tio n
.......................................... . . . . . . . . . . .
36
DISCUSSION AND CONCLUSIONS
. . . . . .
..............................................
37
SUMMARY.............................................................................. ■ ......................................
42
LITERATURE C I T E D ................................................................ ' ..............................
44
-IV -
LIST OF FIGUEES
Page
F ig u re I
Eange o f k e r n e l sm o o th n ess fo u n d i n th e Fg o f a
c r o s s betw een O e re s -(H d p e -F lp re n c e ) and Eenown
20
-V -
LIST OF TABLES
page
T a b le I
T a b le I I
T a b le I I I
T a b le IV
T a b le V
T a b le VI
The c l a s s i f i c a t i o n o f p a r e n t s and F? p l a n t s f o r
aw nedness c o r r e c t e d on th e b a s i s o i F , b r e e d in g
b e h a v io r o f q u e s t i o n a b l e ' ty p e s i n a c r o s s o f
C e re s-(H o p e -F o re n c e ) w ith Renown grown a t Boze­
man i n 1939 and a t Bozeman and M o ccasin , M ontana
i n 1940
16
C a l c u la tio n o f G oodness o f F i t t o
h y b r id r a t i o f o r aw nedness from a
F lo r e n e e ) x Renown c r o s s grow n a t
1939 and a t Bozeman and M o c c a sin ,
a 1 :2 :1 mono­
C e r e s - ( E o p eBozeman i n
M ontana i n 1940
17
C l a s s i f f i c a t i o n i n t o k e r n e l sm o o th n ess c l a s s e s o f
p a r e n t s ■and Fg p ro g en y from a c r o s s o f C e r e s (H o p e -F lo re n c e ) w ith Renown grown a t Bozeman,
M ontana i n 1939
18
The c l a s s i f i c a t i o n o f p a r e n t s and p ro g en y from a
c r o s s o f C e re s -(h o p e * F lo re n e e ,) w ith Renown f o r
aw nedness and k e r n e l sm o o th n ess i n Fg l i n e s grown
a t Boaeman i n 1939 and FL l i n e s grown a t Bozeman
and M o ccasin , M ontana in ^ l9 4 0
21
C l a s s i f i c a t i o n o f p a r e n t s , F and F^ f o r k e r n e l
sm o o th n e ss. ( F . p ro g en y p l a c i d in to ^ t h e g ro u p o f
g r e a t e s t fre q u e n c y )
29
T o ta l le n g th i n m i l l i m e t e r s Cf 15 k e r n e ls from
p a r e n ts and 560 F p l a n t s grown a t Bozeman,
•Montana i n 1939
32
T a b le V I I l T o ta l le n g th in m i l l i m e t e r s o f 15 k e r n e ls o f P a r e n ts
and p ro g e n y from a c r o s s o f C e re s -(E 6p e - F lo r e n c e )
w ith Renown grow nin th e Fg (1 9 3 9 ) and F , (1 9 4 0 )
a t Bozeman, M ontana
34
T a b le V I I I C l a s s i f i c a t i o n o f th e Fg p rd g e n y f o r k e r n e l l e n g t h .
( A ll C la s s e s b a se d on th e p ro b a b le e r r o r l i m i t s o f
th e p a r e n t s —-F. p ro g e n y p la c e d i n t o th e c l a s s o f
g r e a t e s t fre q u e n c y )
36
/
-V IIHHBRITMCE OF AWMEDHESS, EEMEL SMOOTHNESS, KEMEL LENGTH, AND
REACTION TO TWO PHYSIOLOGICE RACES OF TILLETIA IN A SPRING WHEAT CROSS
ABSTRACT
T h is in v e s tig a tio n i s an i n h e r i a t a n c e s tu d y o f avm edness, k e r n e l
s m o o th n e ss , k e r n e l l e n g t h , and mode o f r e a c t i o n t o two p h y s io lo g ic r a c e s
o f T i l l e t i a i n th e p ro g e n y o f a c r o s s betw een C e re s -(H o p e -F lo re n c e ) and
Renown. C e re s -H o p e -F lo re n c e ) i s f u l l y awned (awn ty p e 5 )? m o d e ra te ly
r e s i s t a n t t o tw o p h y s io lo g ic b u n t r a c e s ( T - l l o f T i l l e t i a t r i t i o i and L -3
o f Te l e v i s j , and h a s k e r n e l s w ith th e f o llo w in g c h a r a c t e r i s t i c s : c h e e k s ,
m o s tly a n g u la r and k n if e - e d g e d ; c r e a s e , rad-wide t o w id e , m id -d eep t o d e e p ,
and m o s tly p i t t e d ; and an a v erag e le n g th o f ab o u t 6 ,8 m i l l i m e t e r s . Re­
nown i s w eak ly a n w le tte d (awn ty p e 3 - ) , more r e s i s t a n t t o th e two b u n t
r a c e s , T - I l and L -3 , th a n .C e re s -(H o p e ^ F lo re n c e ), and- h a s k e r n e ls w ith th e
f o llo w in g c h a r a c t e r i s t i c s : c h e e k s , m o s tly ro u n d ed t o s e m i-a n g u la r; c r e a s e
n a rro w t o m id -w id e , s h a llo w t o m id -d e e p , and u s u a l l y c lo s e d b u t some may
be s l i g h t l y p i t t e d ; and an a v e ra g e lergbh o f ab o u t 6 ,4 mm.
One m a jo r f a c t o r was fo u n d t o g o v e rn th e awn d ev elo p m en t betw een
th e p a r e n ts o f t h i s c r o s s w ith th e a w n le tte d c o n d itio n b e in g in c o m p le te ly
dom inant t o th e awned.
The F_ and F- w ere c l a s s i f i e d i n t o e i g h t c l a s s e s d e p en d in g upon
th e d e g re e o f ro u n d n e ss o r a n g u l a r i t y o f c h e e k s , d e p th and w id th o f c r e a s e
and p i t t e d n e s s o f th e k e r n e l s . A u n ifo rm b re e d in g d i f f e r e n c e was demon­
s t r a t e d f o r th e p a r e n ts and th e p ro g ny s e g r e g a te d f o r a com plete ra n g e
o f an betw een th e p a r e n ts b u t no d e f i n i t e f a c t o r i a l a n a l y s i s was e s t a b l i s h
ed.
A t o t a l le n g th o f f i f t e e n s e l e c t e d k e r n e ls from e a c h p la n t w ere
m e asu re d i n m i l l i m e t e r s t o d e te rm in e k e r n e l le n g th d i f f e r e n c e s . I n th e
Fp a v e ry r e g u l a r s e g r e g a tio n seemed t o be i n d i c a t e d w ith a weak dom inance
ox th e sm o o th er ty p e s s u g g e s te d . The p a r e n ts and F- w ere so v a r i a b l e i t
was im p o s s ib le t o check th e F^ c l a s s i f i c a t i o n by th e F^ b re e d in g b e h a v io r .
Bunt i n f e c t i o n s i n th e p a r e n t s , p ro g e n y , and s u s c e p t i b l e c h e c k ,
U lk a , w ere so s l i g h t i t was d e c id e d from o b s e r v a tio n a l d a t a t h a t a
d i f f e r e n t i a l r e a c t i o n o f th e p ro g e n y c o u ld n o t be d e m o n s tra te d .
IHHEEITAHOB OF AiMEDNESS, KERNEL SMOOTHNESS1 , KERNEL LENGTH, MD
REACTION TO TWO PHYSIOLOGIC RACES OF TILLBTIA IN A SPRING WHEAT CROSS
O r lie W. S m ith
INTRODUCTION
From th e e a r l i e s t tim e s m en.have re c o g n iz e d th e f a c t t h a t
" l i k e L e g e ts l i k e " and t h a t th e o f f s p r i n g d i f f e r somewhat among them ­
s e l v e s and from t h e i r p a r e n t s .
They have lo n g u s e d t h i s k n o w led g e, more
o r l e s s u n c o n s c io u s ly , p e r h a p s , i n c h o o sin g f o r b r e e d in g p u rp o s e s th o s e
i n d i v i d u a l s among t h e i r d o m e s tic a te d a n im a ls and p l a n t s w hich b e s t s u i t ­
e d t h e i r r e q u ire m e n ts ..
O nly r a r e l y w ere d e l i b e r a t e b r e e d in g m ethods u s e d
w hich depended upon an e m p ir ic a l know ledge o f th e m ethods o f r e p r o d u c tio n , .
su ch a s th e a r t i f i c i a l p o l l i n a t i o n o f th e fem ale d a te p alm , w hich was
p r a c t i c e d i n Egypt and M esopotam ia many c e n t u r i e s b e fo re th e C h r i s t i a n
e ra .
The e a r l y husbandmen b re d t h e i r a n im a ls and p l a n t s w ith o u t any g e n e r a l
know ledge o f th e r e p r o d u c tiv e p r o c e s s e s , and th e le g a c y o f v a lu a b le c u l t i v
v a te d p l a n t s , w hich i s th e earm ark o f p erm anent c i v i l i z a t i o n s , r e s u l t e d
fro m ta m in g th e w ild s p e c ie s and s e l e c t i n g f o r t u i t o u s v a r i a t i o n s among th em .
A s c i e n t i f i c u n d e r s ta n d in g o f th e p ro b lem s o f h e r e d i t y and v a r i a t i o n , how—
e v e r , h a s b e fu n t o be re a c h e d o n ly r e c e n t l y .
In a l i t t l e
o v e r o n - t h i r d o f a c e n tu r y , g e n e t i c s h a s d e v elo p e d
i n t o an e x a c t s c ie n c e com parable t o su ch o ld e r s c ie n c e s a s p h y s ic s and
1K e rn e l sm oothness a s u s e d th ro u g h o u t t h i s p a p e r p e r t a i n s o n ly
t o th e c o n d itio n o f th e c h e e k s , c r e a s e , and p i t t e d n e s s o f th e k e r n e l s .
-2 -
c h e m is tr y .
The mode o f i n h e r i t a n c e o f many p la n t c h a r a c t e r s h as been
s t u d i e d and i n many c a s e s a f a c t o r i a l a n a l y s i s h a s b een e s t a b l i s h e d b u t
t h i s r e s e a r c h i s s u r e l y y e t i n i t s in f a n c y .
Many p ro b le m s rem ain t o be
s o lv e d b e fo re th e g e n e t i c i s t w i l l be a b le t o combine a l l d e s ir e d q u a l i t i e s
;
i n t o a p re c o n c e iv e d p e r f e c t v a r i e t y . . E x a c t in f o r m a tio n r e g a r d in g th e
^
u n d e r ly in g p r i n c i p l e s o f i n h e r i t a n c e o f v a r io u s c h a r a c t e r s h a s a id e d i n
,
e s t a b l i s h i n g more e f f i c i e n t b re e d in g o p e r a t i o n s .
The m echanism o f h e r e d i t y was f i r s t c o n c e iv e d and worked o u t
by G reg o r M endel whose e x p e rim e n ts i n p l a n t h y b r i d i z a t i o n l a i d th e fo u n d a ­
t i o n f o r m ost o f th e modern work i n g e n e t i c s .
M en d el’ s f o u r law s d e a l t
w ith u n i t c h a r a c t e r s , dom inance, s e g r e g a tio n and re c o m b in a tio n .
A lth o u g h
p u b lis h e d i n 1866, h i s f in d i n g s l a y i n o b s c u r ity u n t i l 1900 when th e y w ere
in d e p e n d e n tly and s im u lta n e o u s ly r e d is c o v e r e d by D e m e s , C o rre n s , and
Von T scherm ak.
U n it c h a r a c t e r s have n o t p ro v ed to . be a s sim p le i n w heat
a s was f i r s t th o u g h t and dom inance i s in c o m p le te i n m ost c a s e s , b u t i n
s p i t e o f t h i s th e s i n g l e c h a r a c t e r i s s t i l l th e fo u n d a tio n i n s tu d y in g
w heat i n h e r i t a n c e and dom inance th e p rim a ry b a s i s f o r g e n e t i c i n t e r p r e t a ­
tio n s .
D u rin g th e p a s t 15 o r 20 y e a r s c o n s id e r a b le in f o r m a tio n h a s
b e en a c c u m u la te d on th e mode o f i n h e r i t a n c e o f many c h a r a c t e r s o f w h e a t.
The i n h e r i t a n c e o f some o f th e s e c h a r a c t e r s h a s been fo u n d t o be r e l a t i v e ­
l y s im p le , w h ereas i n o t h e r c a s e s com plex r e a c t i o n s a r e fo u n d w hich i n ­
d i c a t e t h a t a la r g e number o f g e n e t i c f a c t o r s a re in v o lv e d .
The s o - c a l l e d
" q u a l i t a t i v e c h a r a c te r s " ,, such a s aw n ed n ess, k e r n e l c o l o r , c o lo r o f s tr a w ,
p u b e sc e n ce and c o lo r o f g lu m e s, and d w a rf and n orm al p l a n t s , a re u s u a l l y
-3 ~
more sim]3e t o s tu d y th a n '’q u a n t i t a t i v e c h a r a c te r s " w hich in c lu d e su ch
c h a r a c t e r s a s tim e o f m a t u r i t y , y i e l d , w i n t e r h a r d i n e s s , h e ig h t o f p l a n t ,
and strerg b h o f s tr a w .
T h is i s b e c a u se th e q u l a i t a t i v e c h a r a c te r s can
r e a d i l y be p la c e d i n t o d i s t i n c t c l a s s e s and g ro u p s , w h e rea s q u a n t i t a t i v e
c h a r a c t e r s a re c l a s s i f i e d i n n u m e ric a l o r p e rc e n ta g e f r e q u e n c i e s .
K e rn e l sm o o th n ess and k e r n e l le n g th a re two c h a r a c te r s o f w heat
on w hich t h e r e h a s been v e ry l i t t l e
mode o f i n h e r i t a n c e .
s y s te m a tic s tu d y t o d e te rm in e t h e i r
These q u a l i t i e s a re v e ry im p o r ta n t s in c e th e le n g t h
o f k e r n e l i s th e m ain d i f f e r e n t i a t i n g c h a r a c t e r betw een s p r in g and w in te r
w heat and sm oothness o f k e r n e l seems t o be c l o s e l y a s s o c i a t e d w ith t e s t
w e ig h t and c o n s e q u e n tly y i e l d o f f l o u r .
A sm ooth, plum p, and s h o r t k e r n e l
i s th e d e s i r e d ty p e i n h a rd re d s p r in g w heat and w ith th e s e c h a r a c t e r s i n
m ind th e w heat b r e e d e r i s a b le t o e lim in a te i n e a r l y g e n e r a tio n s tiaose
s e l e c t i o n s n o t h a v in g th e d e s ir e d k e r n e l c h a r a c t e r i s t i c s .
I t seems e v id e n t
t h a t k e r n e l sm oothness and k e r n e l le n g th have a d e f i n i t e mode o f i n h e r i t a n c e
b e c a u s e i t h a s been p o s s ib le to s e l e c t v a r i e t i e s t r u e b re e d in g f o r sm o o th ,
plum p, and s h o r t k e r n e l s from th e p ro g e n y o f p a r e n ts c o n tr a s te d i n these..!
re s p e c ts .
O b v io u s ly , th e n , t o p ro c e e d more e f f i c i e n t l y i n a w heat b re e d ­
i n g p ro g ram , i t i s d e s i r a b l e t o know th e e x a c t mode o f i n h e r i t a n c e o f th e s e
two c h a r a c t e r s .
The i n v e s t i g a t i o n s r e p o r te d i n t h i s p a p e r c o n s i s t o f i n h e r i t a n c e
s t u d i e s i n a s p r in g w heat c r o s s , C e re s -(H o p e -F lo re n c e ) x Renown, on aw nedn e s s , k e r n e l sm o o th n e ss, k e r n e l l e n g t h , and r e a c t i o n t o two p h y s io lo g ic
r a c e s o f b unt
.
-4 -
L ITEEATURE REVIEW
A1WnecLness
I n th e e a r l y r e p o r t s on s t u d i e s o f i n h e r i t a n c e o f aw nedness
i n w h e a t, o n ly two c l a s s e s , awned and a w n le s s , w ere d i f f e r e n t i a t e d .
More r e c e n t s t u d i e s have e s t a b l i s h e d th e f a c t t h a t t h e r e a re t r u e b re e d ­
i n g awn ty p e s in te r m e d ia te betw een c o m p le te ly a w n less and f u l l y awned
w h e a ts .
U n t i l th e s e in te r m e d ia te ty p e s were f u l l y r e c o g n iz e d , much con­
f u s io n r e s u l t e d among e a r l y w o rk e rs who s tu d ie d th e in h e r ita n c e o f t h i s
.c h a ra c te r.
A c c o rd in g t o Hayes and G a rb e r (1 9 2 7 )? B if f e n i n 1905 r e p o r te d
th e f i r s t s y s te m a tic g e n e t i c s tu d y o f aw n ed n ess.
He c o n clu d e d from th e
s tu d y o f s e v e r a l c r o s s e s o f b e a rd e d w ith s o - c a l l e d " b e a r d l e s s ” w h eats
t h a t th e " a w n le s s" c o n d itio n was dom inant and th e awned c o n d itio n r e ­
c e s s iv e .
P l a n t s h a v in g awns up t o o n e - h a lf in c h i n le n g t h were r e g a r d ­
e d a s " a w n le s s " , th u s th e
i n t o 3 :1 " a w n le ss" t o awned.
was alw ay s " b e a r d le s s " and th e F^ s e g r e g a te d
The awned ty p e s alw ays b re d t r u e , w h ile a b o u t
o n e - t h i r d o f th e " b e a r d le s s " ty p e s b re d t r u e and tw o - t h i r d s s e g r e g a te d i n ­
t o awned and " a w n le ss" t y p e s .
S e v e r a l o th e r w o rk d rs o b ta in e d s i m i l a r r a t i o s i n th e F^ and a
3 :1 b e a r d le s s t o b e a rd e d s e g r e g a tio n i n th e F^.
S a u n d e rs (1907) seem s t o
be th e f i r s t w o rk e r t o q u e s tio n th e id e a t h a t th e F1 o f a c ro s s b etw een a
b e a r d l e s s and a b e a rd e d w heat i s alw ay s b e a r d l e s s .
He m a in ta in e d t h a t th e
ty p e o f awns o b ta in e d i n th e F^ d ep en d s upon th e d i f f e r e n c e betw een th e
p a r e n t form s u s e d .
P e r c i v a l (1 9 2 1 ) l a t e r r e p o r te d t h a t i n num erous c r o s s e s
th e F^ p l a n t s have awns w hich a re e a s i l y d i s t i n g u i s h e d from .th e b e a r d l e s s
-5 -
p a r e n t and t h a t th e
s e g r e g a te d i n t o a 1 :2 :1 r a t i o o f h e a d le s s , Semi­
t e a r d e d , and b e a r d e d .
Many w o rk e rs have v e r i f i e d th e s e r e s u l t s .
Hayes
and Asmodt (1 9 2 3 ) found t h a t th e F^ o f a c r o s s b etw een and awned and a
sem i-aw ned v a r i e t y was s tr o n g l y a w n le tte d and th e F^ s e g r e g a te d a s
a sim p le m on o h y b rid .
C la rk and Q u is e n b e rry (1929) v e r i f i e d th e s e r e s u l t s
by a r e c i p r o c a l c r o s s o f th e same v a r i e t i e s .
S e v e ra l o t h e r i n v e s t i g a t i o n s
have r e v e a le d t h a t a o n e - f a c t o r d i f f e r e n c e would a c c o u n t f o r th e awn ty p e s
fo u n d i n th e p ro g e n y o f c r o s s e s b etw een sem i-aw ned and awned v a r i e t i e s .
W orkers who have r e p o r te d su ch r e s u l t s a r e :
G ain es and S in g le to n ( 1 9 2 6 ) ,
S te w a rt and Woodward ( l 9 3 l ) » S te w a rt and H a lle y (1 9 3 2 ) , Q u is e n b e rry and
C la rk ( 1 9 3 3 ) j Ausemus (1 9 3 4 )? and T in g e y and T o lm a n (l9 3 4 ).
S te w a rt and T in g ey (1 9 2 8 ) fo u n d t h a t when a t r u e b e a r d le s s
w heat was c r o s s e d w ith s e m i-b e a rd e d w heat t h a t a m onhybrid r a t i o was ob­
ta in e d .
A lso Q u is e n b e rry and C la rk (1 9 3 3 ) and Ausemus (1 9 3 4 ) o b ta in e d
s i m i l a r r e s u l t s from s i m i l a r c ro s s e s .,
The f i r s t work w ith t r u e a w n le ss and f u l l y awned w h eats was r e ­
p o r t e d by Howard and Howard (1912 ? 1915) and th e y c o n c lu d e d t h a t two f a c t o r s
w ere n e c e s s a r y t o e x p la i n th e d i f f e r e n c e i n i n h e r i t a n c e betw een th e s e two
ty p e s .
They g ro u p ed a l l awned and tip - a w n e d c l a s s e s t o g e t h e r a s aw ned,
w hich i n c o m p ariso n w ith th e a w n le ss gave a 1 5 :1 r a t i o .
They th u s con­
c lu d e d t h a t th e awned condition was d o m in a n t, w hich was an o p p o s ite con­
c lu s i o n t o t h a t o f o th e r w o rk e rs .
A number o f o th e r i n v e s t i g a t o r s have o b ta in e d a tw o -m ajo r—f a c t o r
d i f f e r e n c e f o r aw nedness when a w n le ss and awned ty p e s w ere c r o s s e d .
In
m ost c a s e s i t was assum ed t h a t th e a w n le ss and sem i-aw ned c o n d itio n was
-6 -
dom inant t o th e aim ed c o n d itio n *
A lso i n a number o f c a s e s th e w o rk e rs
have fo u n d i t n e c e s s a r y t o assume t h a t one o r more m in o r m o d ify in g f a c t o r s
w ere p r e s e n t t o f u l l y e x p la i n t h e i r r e s u l t s .
I n a K o ta ( fu lly -a w n e d ) x
H ard F e d e r a tio n ( t r u e a w n le s s ) c r o s s , C la rk (1924) c o n c lu d e d t h a t th e awn­
l e s s c o n d itio n was d o m in a n t, a s th e F^ ( a p i c a l I y a w n le tte d ) a p p ro a ch e d more
n e a r l y th e a w n le ss th a n th e awned v a r i e t y , and a l s o t h a t two g e n e ti c f a c t o r s
c o u ld n o t e n t i r e l y a cc o u n t f o r th e b r e e d in g b e h a v io r i n th e F^ and F^ g e n e r­
a tio n s ,
C la r k , F l o r e l l , and H ooker (1 9 2 8 ) s tu d ie d two c r o s s e s o f b e a r d ­
l e s s and b e a rd e d w h e ats and fo u n d a tw o -m a jo r f a c t o r d i f f e r e n c e i n one
c r o s s and a t h r e e - f a c t o r , two m a jo r and one m in o r, d i f f e r e n c e i n th e o t h e r .
A p p a re n tly th e o n ly e f f e c t o f th e m in o r f a c t o r , i n th e p re s e n c e o f one
m a jo r f a c t o r , i s t o u n ifo r m ly re d u c e th e e x te n t o f aw n ed n ess.
From t h i s
.
s tu d y th e y c o n c lu d e d t h a t a s many a s f o u r f a c t o r s may be in v o lv e d i n th e
i n h e r i t a n c e o f aw nedness i n w h e a t.
O th e r i n v e s t i g a t o r s who have fo u n d
t w o - f a c to r d i f f e r e n c e s i n w heat a r e :
S te w a rt and Heyward (1 9 2 9 ), S te w a rt
and Ju d d ( l 9 3 l ) , C la r k , Q u is e n b e rry , and Pow ers (1 9 3 3 ) , and Ausemus ( 1 9 3 4 ) ®
L i tz e n b e r g e r (19 3 9 ) r e p o r te d a tw o -m a jo r f a c t o r d i f f e r e n c e i n two c r o s s e s
o f b e a r d le s s and b e a rd e d w h e a ts ,
A p h e n o ty p ic r a t i o o f 1 : 4 : 5 : 3 : 2 : 1 f o r
•aw n less, a p i c a l l y a w n le tte d , w eak ly a w n le tte d , s t r o n g l y a w n le tte d , h a l f
aw ned, and awned r e s p e c t i v e l y was. fo u n d i n one c ro s s and a 4 : 8 : 1 : 3 : 2 : 1
p h e n o ty p ic r a t i o f o r th e s i x c l a s s e s i n th e o th e r c r o s s .
■
He a ls o made
s e v e r a l t e s t h y b r i d i z a t i o n s in v o lv in g th e p a r e n ts and p ro g en y ofone o f
th e o r i g i n a l c r o s s e s and from t h i s he c o n clu d e d t h a t th e aw n-group c l a s s i ­
f i c a t i o n a s o r i g i n a l l y made was c o r r e c t .
A number o f i n v e s t i g a t o r s have r e p o r te d t h a t t h e r e a re g e n e t i c a l l y
-7 -
d i f f e r e n t ty p e s o f sem i-avm ed w h e a ts .
Love and C ra ig (.1926) s t a t e t h a t
when S o n o ra i s c ro s s e d w ith a w n le ss ty p e s o f common w heat ( Tritipnm
v n l g a r e ) , th e
i s a w n le s s , w h ile i n th e
and p a r t l y awned ty p e s a p p e a r.
and l a t e r g e n e r a tio n awned
The r e s u l t s a p p ro a ch e d a 1 5 :1 r a t i o w hich
i n d i c a t e d t h i s v a r i e t y a lth o u g h sem -aw n le ss c a r r i e d a f a c t o r f o r th e
b e a rd e d c o n d i t i o n .
S te w a rt (1 9 3 2 ) c r o s s e d two t r u e b r e e d in g a w n le tte d
ty p e s w hich he o b ta in e d a s s e g r e g a te s from a c ro s s o f S e v ie r x F e d e r a tio n
and o b ta in e d a l l ty p e s a g a in . . F o u r t r u e b re e d in g ty p e s w ere r e c o v e r e d one f u l l y aw ned, one c o m p le te ly a w n le ss and two a w n le tte d ty p e s . C la rk
and Q u isen b er r y - ( 1933) c ro s s e d two a w n le tte d w h e a ts , S o n o ra and Q u a l i t y ,
and o b ta in e d s i m i l a r r e s u l t s .
T here was a com plete ra n g e from a w n le ss
t o awned i n th e F^ and i n th e F^ t h e r e w ere o b ta in e d tr u e , b re e d in g a w n le ss
and awned s t r a i n s a s w e ll a s s t r a i n s a w n le tte d l i k e b o th p a r e n t s .
I t was
assum ed t h a t S onora c o n ta in s th e g e n e t i c f a c t o r s aaBB and Q u a lity AAbb,
w h e rea s a w n le ss s e g r e g a te s a re AABB and awned ones a a b b .
The l i t e r a t u r e re v ie w e d i n d i c a t e s t h a t th e mode o f i n h e r i t a n c e
o f awns i n w heat i s r e l a t i v e l y sim p le i n some c r o s s e s and r a t h e r com plex in
o th e rs .
U s u a lly a s i n g l e - f a c t o r d i f f e r e n c e w i l l e x p la i n th e s e g r e g a tio n o f
sem i-aw ned x awned o r sem i-aw ned x t r u e aw n less c r o s s e s and two f a c t o r s ,
th e t r u e a w n le ssx f u l l y awned c r o s s e s .
However, a s many a s two m a jo r and
tw o m inor f a c t o r s w ere n e c e s s a r y t o f u l l y e x p la in th e s e g r e g a tio n o b ta in e d ,
i n one c r o s s .
The f o u r t r u e b r e e d in g ty p e s most o f te n fo u n d a re a w n le s s ,
aw ned, w eak ly a w n le tte d , and s t r o n g l y a w n le tte d .
T rue b re e d in g a p i c a l l y
awned and h a l f awned ty p e s have b een o b ta in e d from a few c r o s s e s .
Dominance
o f a w n le s s n e s s i s in c o m p le te — th e F^ more n e a r ly a p p ro a c h e s th e a w n le ss
•8-
p a r e n t th a n th e awned t y p e .
K e rn e l Sm oothness
The w r i t e r was u n a b le t o f i n d any l i t e r a t u r e d e a li n g w ith a
s y s te m a tic s tu d y o f the- i n h e r i t a n c e o f k e r n e l c h e e k , c r e a s e , and p i t t e d
c h a ra c te ris tic s .
K e rn e l L en g th
The le n g th o f k e r n e l i s u s e d a s a m ajo r
in g v a r i e t i e s o f w h e a t.
c h a ra c te r in d is tin g u is h ­
The d o r t e r ty p e s a re u s u a l l y c h a r a c t e r i s t i c o f
th e h a rd r e d s p r in g w h e a ts and th e lo n g e r o n e s , th e r e d w in te r w h e a ts .
T h is g e n e r a l i z a t i o n , how ever, i s becom ing more c o n fu s in g e v e ry y e a r b e ­
c au se o f th e d i s t r i b u t i o n o f more v a r i e t i e s o f h a rd r e d s p r in g w heat
w ith lo n g k e r n e l s and some h a rd r e d w i n t e r w h eats w ith s h o r t k e r n e l s .
Engledow (l$ 2 0 ) seems t o be th e f i r s t p e rs o n t o r e p o r t any i n ­
h e r i t a n c e s t u d i e s o f g r a i n le n g th o f w h e a t.
He c r o s s e d T ritio u m c o lo n !cu m .
w hich h ad a mean g r a i n le n g th o f a b o u t 1 0 .2 mm. and T . durum w ith a man
le n g th o f a b o u t 7»7 mm. and found- t h a t th e
was i n t e r m e d i a t e .
Fg t h r e e ty p e s w ere fo u n d w ith th e f o llo w in g mean g r a i n le n g th s :
I n th e
8 .8 4 ,
8 .6 7 , and 8 .3 3 mm. r e p r e s e n t i n g th e e x t r a c t e d p o lo n ic u m s , h e te ro z y g o u s
i n t e r m e d i a t e s , and e x t r a c t e d durums r e s p e c t i v e l y .
He c o n clu d e d th e
s e g r e g a tio n was su ch a s t o s u g g e s t a o n e - f a c t o r d i f f e r e n c e — a p p ro x im a te ­
l y a 1 :2 :1 r a t i o .
Ho g r a in s w ere fo u n d a s lo n g a s th e g ra n d p a r e n t a l
p o lonicum and no® a s s h o r t a s th e f u r u m .an c e s t o r , an in w ard " s h i f t " , t o ­
w ard th e mean o f th e two g r a n d p a r e n ts h a v in g o c c u r r e d .
I n a r e c e n t s tu d y o f k e r n e l l e n g t h , W eb ster (1 9 3 7 ) u s e d 10
“9-
p rim a ry k e r n e l s s e l e c t e d a t random from one s p ik e o f e a c h ,F2 and F^
p l a n t and r e c o r d e d th e t o t a l le n g th i n m i l l i m e t e r s .
The p a r e n ts u s e d
i n th e c r o s s s tu d ie d w ere B a a rt ( T r itic u m V u lg a re ) w hich h ad a mean
l e n t h o f 7 5 « 6 0 -.5 5 mm. and S hot w heat ( T. sphaerbococcum ) w ith a mean
le n g th o f 4 8 .2 7 -'.3 5 mm., a d i f f e r e n c e o f 27.33mm.
He fo u n d t h a t i n
th e F2 t h e r e was a c e d id e d dom inance o f th e lo n g e r ty p e k e r n e l s and i n
th e F ^ , 24 homozygous lo n g ty p e s a p p e a re d h u t o n ly one homozygous s h o r t
ty p e was r e c o v e r e d .
He c o n c lu d e d t h a t s e g r e g a tio n f o r k e r n e l le n g th
c o u ld n o t he s a t i s f a c t o r i l y e x p la in e d on a sim p le f a c t o r b a s i s .
Bunt R e a c tio n
The mode o f 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 p h y s io lo g ic ra c e s , o f
T i l l e t i a t r i t i c i ( B j e r k .) W in t. and T^. L e v is (KtIhn) h a s b een s tu d ie d by
a la r g e num ber o f w o rk e rs and e x p la in e d on o n e -, tw o -, t h r e e - , and
m u ltip le - fa c to r d iff e r e n c e s .
S c h le h u b e r (1938) h a s re v ie w e d th e l i t e r a t u r e
on th e n a tu r e o f s e g r e g a tio n r a t h e r e x h a u s tiv e ly and h a s t rouped th e
a u th o r s a c c o rd in g t o th e number o f f a c t o r s in v o lv e d i n th e c ro s s w ith
w hich th e y w orked.
S in c e th e w r i t e r was u n a b le t o f i n d any b u n t i n h e r i t a n c e
w ork more r e c e n t t h a n , t h a t sum m arized by S c h le h u b e r, no a d d i t i o n a l
l i t e r a t u r e re v ie w h a s been a tte m p te d .
-1 0 -
MATERIALS MD METHODS
The p u rp o se o f th e s tu d y was t o d e te rm in e th e mode o f i n ­
h e r i t a n c e o f aw n ed n ess, k e r n e l s m o o th n e ss, k e r n e l l e n g t h , and r e a c t i o n
t o two p h y s io lo g ic r a c e s o f T i l l e t i a i n a c r o s s betw een C e re s-(H o p e -I
F lo r e n c e ) (C . I . 11872)
and Renown (C . I . 11709)»
The C e re s-(H o p e -
F l o r e n c e p a r e n t i s a s e l e c t i o n o u t o f a c r o s s o f O eres w ith Hope x
F lo r e n c e , i s f u l l awned (awn ty p e %) and h a s k e r n e ls o f th e fo llo w in g
c h a ra c te ris tic s :
c h e e k s , m o s tly a n g u la r and k n if e - e d g e d ; c r e a s e , m id ^:
w ide t o w id e , m id -d eep t o d e e p , and m o s tly p i t t e d ; a v e ra g e le n g th a b o u t
6 ,8 m i l l i m e t e r s ; and m o d e ra te ly r e s i s t a n t t o h u n t r a c e s T - I l
2
2
and 1 -3 .
The o th e r p a r e n t , Renown, i s w eak ly a w n le tte d (awn ty p e 3 - ) and p o s s e s s e s
k e r n e l s o f th e s e c h a r a c t e r i s t i c s :
c h e e k s , m o s tly ro u n d e d t o s e m i- a n g u la r ;
c r e a s e , n a rro w t o m id -w id e, s h a llo w t o m id -d e e p , u s u a l l y c lo s e d b u t some
may be s l i g h t l y p i t t e d ; a v e ra g e le n g t h a b o u t 6 .4 m i l l i m e t e r s ; and more
r e s i s t a n t t o th e two b u n t r a c e s , T - I l and L -3 th a n C e re s = (H o p e _ F lo re n c e ). ■
O e re s -(H o p e -F lo re n c e ) was p ro d u ced a t a N o rth D akota A g r i c u l t u r a l Ex­
p e rim e n t S t a t i o n and i s r e s i s t a n t t o m ost o f th e known p h y s io lo g ic r a c e s
o f b u n t and Renown was p ro d u c e d a t th e D ominion R u st R e s e a rc h L a b o ra to ry
a t W in n ip eg , C anada, and i s a l s o r e s i s t a n t to m ost o f th e known p h y s io lo g ic
r a c e s o f b u n t.
^ 0 ,1 . r e f e r s t o a c c e s s io n num ber o f th e D iv is io n o f C e re a l C rops
and D is e a s e s .
^ T - I l and L-3 r e f e r t o known s p e c i f i c p h y s io lo g ic r a c e s o f
T i l l e t i a t r i t i c i and T . I e v i s r e s p e c t i v e l y .
-1 1 -
The p a r e n t v a r i e t i e s , C e re s -(H o p e -F lo re n e e ) and Renown, w ere
grown i n a b u n t p h y s io lo g ic ra c e n u r s e r y a t Bozeman, M ontana, and P u l l ­
man, W ash in g to n , i n 1937•
B ecause th e s e w ere two v a r i e i t i e s g iv in g low
p e r c e n ta g e s o f i n f e c t i o n t o a l l known r a c e s o f T i l l e t i a t r i t i c i and Ti*
l e v i s , a c r o s s was made i n an e f f o r t t o combine th e sm ut r e s i s t a n c e o f
th e two p a r e n ts w ith th e h ig h y i e l d o f C e re s -(H o p e -F lo re n c e ) and th e s h o r t
sm ooth k e r n e l ty p e and w eakly a w n le tte d c o n d itio n o f Renown'.
The c r o s s was
made a t Bozeman, M ontana, u n d e r th e s u p e r v is io n o f B r. R. H. Bamberg and
advanced th ro u g h th e F^ i n 1939 a s a p a r t o f th e w heat im provem ent p ro g ram .
The Fg p ro g e n y c o n s i s t i n g o f s i x f a m i l i e s o f a p p ro x im a te ly 100
p l a n t s e ac h and th e p a r e n ts w ere s t u d i e d i n th e l a b o r a t o r y f o r aw n ed n ess,
k e r n e l sm o o th n e ss , and k e r n e l l e n g t h .
F o r aw nedness i n h e r i t a n c e s t t i d i e s , th e Fg h y b r id p l a n t s w ere
c l a s s i f i e d a c c o rd in g t o th e f o llo w in g c l a s s e s : *234
Awn c l a s s ________ " ______________ D e s c r ip tio n o f awn ty p e ____________ ;_______ ___
L
A w nless, no d ev elo p m en t o f a w n le ts o v e r 2 mm. i n le n g th .2
A p ic a lly a w n le tte d , h a s awns from 2 t o 15 mm. lo n g a t th e
apex o f th e s p ik e .
3
W eakly a w n le tte d , h a s a w n le ts from 3 t o 20 mm. lo n g , th e
s h o r t e r o c c u r r in g a t th e base o f th e s p ik e and th e le n g t h
i n c r e a s i n g to w a rd th e a p ex ,
3+
S tr o n g ly a w n le tte d , h a s a w n le ts from 3 t o 40 mm, lo n g , th e
s h o r t e r ' o c c u r r in g a t th e b ase o f th e s p ik e and th e le n g t h
i n c r e a s i n g to w a rd th e a p e x . The lo w e r a w n le ts i n t h i s
c l a s s a re u s u a l l y in c u r v e d .
4
H a lf awned, h a s s h o r t awns v a r y in g th ro u g h o u t th e sp ik e
from 18 t o 50 mm. lo n g — a p p ro x im a te ly o n e - h a lf th e le n g th
o f a f u l l y awned t y p e .
5
Awned, th e awns v a r y from 30 t o 100 mm, i n le n g t h .
*12-
E or k e r n e l sm oothness i n h e r i t a n c e s t u d i e s , th e s e e d s o f th e
p l a n t s w ere c l a s s i f i e d a c c o rd in g t o th e fo llo w in g c l a s s e s ;
K e rn e l
sm o o th n ess c l a s s e s _______ D e s c r ip tio n o f k e r n e l sm o o th n ess c l a s s e s
2
C heeks, ro u n d ; c r e a s e , n a rro w , s h a llo w and c lo s e d
3
C heeks, s e m i- a n g u la r ; c r e a s e , n a rro w , s h a llo w and c lo s e d .
4
C heeks, m o s tly ro u n d e d ; c r e a s e , s h a llo w t o sem i-d ee d
and c lo s e d e x c e p t a few k e r n e ls may he s l i g h t l y p i t t e d
n e a r c e n te r o f c r e a s e .
5
C heeks, m o s tly s e m i- a n g u la r to. a n g u la r ; c re a s e
m o s tly n a rro w t o mid—w ide and m id -d e e p . May he
m o d e ra te ly p i t t e d .
6
C heeks, m o s tly a n g u la r ; c r e a s e , m o s tly m id-w ide to
w id e , m id -d eep t o d eep and p i t t e d .
7
C heeks, m o s tly a n g u la r , k n if e - e d g e d , and ro u g h o r
s c a l y ; c r e a s e , m o s tly w ide to s e m i-w id e , deep t o
m id -d e e p , and p i t t e d . V e n tr a l s u r f a c e u s u a l l y
s lo p in g from ch eek m a rg in s to b o tto m o f p i t and
w rin k le d aro u n d m a rg in s o f p i t .
8
C heeks, a n g u la r , k n if e - e d g e d , ro u g h and s c a ly ;
c r e a s e , m o s tly w id e , d eep and e x t e n s i v e l y p i t t e d .
V e n tra l" s u r f a c e u s u a l l y rough and s lo p in g from
m a rg in s o f ch eek i n t o th e deep p i t . D epth o f p i t
a b o u t o n e - h a lf o f d o r s o v e n tr a l th i c k n e s s o f k e r n e l . *2
F o r k e r n e l le n g th i n h e r i t a n c e s t u d i e s , h e a d s w ere s e l e c t e d from
th e th r e e m ain t i l l e r s o f e ac h F
p l a n t , F i f t e e n k e r n e l s were th e n ta k e n '
2
from th e two lo w e r f l o r e t s o f s p i k e l e t s s e l e c t e d a t random from th e c e n t e r
o f th e s e s p i k e s , l a i d end t o e n d , and m easured i n m i l l i m e t e r s , '
F o r F^ i n h e r i t a n c e s t u d i e s o f aw n ed n ess, k e r n e l sm o o th n e ss,
k e r n e l l e n g t h , and r e a c t i o n t o two p h y s io lo g ic r a c e s o f T i l l e t i a , two
s e p a r a te n u r s e r i e s o f th e same
l i n e s w ere grow n.
One n u r s e r y was
se e d e d f o r ,a w n e d n e s s , k e r n e l s m o o th n e ss , and k e r n e l le n g th s t u d i e s and
th e o t h e r was t r e a t e d w ith two r a c e s o f h u n t and se e d e d f o r h u n t i n ­
h e rita n c e s tu d ie s .
I n th e
’aw n ed n ess, k e r n e l sm o o th n e ss, and k e r n e l le n g th
n u r s e r y , 60 l i n e s o f 30 s e e d s e a c h w ere s e l e c t e d a t random from e a c h o f
f o u r Fg f a m i l i e s .
These l i n e s w ere se e d e d i n s in g le row s 12 in c h e s a p a r t
w ith th e s e e d sp a c e d a b o u t t h r e e in c h e s a p a r t w ith in th e ro w s.
w ere grow n a t Bozeman and two a t M o c c a sin , M ontana.
was in c lu d e d a f t e r e v e ry 20 rows o f p ro g e n y .
Two f a m i l i e s
One row o f e a c h p a r e n t
A ls o , t h i r t y s e e d s e a c h o f
a l l q u e s tio n a b le awn ty p e s fou n d i n th e Pg were grown a t Bozeman and M o ccasin
The aw nedness c l a s s i f i c a t i o n was made i n th e f i e l d by d e te r m in in g
w h e th e r th e p l a n t s i n e a c h row w ere s e g r e g a tin g o r b r e e d in g t r u e f o r awn
ty p e s a s d e te rm in e d i n th e F^.
At Bozeman 20 p l a n t s w ere p u l l e d from e a c h o f th e 120 F^ l i n e s
and from 18 p a r e n t ro w s.
The s e e d s w ere th e n th r e s h e d and c l a s s i f i e d f o r
k e r n e l sm o o th n ess and m easured f o r k e r n e l le n g th i n th e same m anner a s
was done f o r th e Fg,
Twenty p l a n t s w ere p u lle d from e a c h l i n e a t M occasin
fo r s im ila r s tu d ie s .
F o r th e F^ b u n t s t u d i e s , 140 s e e d s o f th e same l i n e s a s w ere
s t u d i e d f o r aw n ed n ess, k e r n e l s m o o th n e ss , and k e r n e l le n g t h were p la n te d
i n d u p l i c a t e row s o f 70 s e e d s e a c h .
Seed f o r one o f th e d u p lic a te row s
o f e a c h l i n e was in o c u la te d w ith r a c e T - I l o f Ti. t r i t i c i and th e o t h e r
w ith r a c e 1 -3 o f T^. I e v i s by s h a k in g an e x c e s s o f th e s p o r e s and th e s e e d
-1 4 -
t o g e t h e r i n a s m a ll p a p e r e n v e lo p e .
One row o f e a c h p a r e n t and a
s u s c e p t i h e l c h e c k , U lk a , w ere in o c u l a t e d i n th e same m anner and in c lu d e d
a f t e r e v e r y 15 row s o f p ro g e n y .
A ls o ,
p o p u latio n s o f e a c h fa m ily w ere
in o c u l a t e d w ith e ac h o f th e p h y s io lo g ic r a c e s o f h u n t u s e d and grown ad­
j a c e n t t o th e
l i n e s i n o r d e r t o compare th e h u n t r e a c t i o n o f h o th th e
Fg and F^ g e n e r a tio n s i n th e same y e a r .
One h u n d red f o r t y s e e d s o f e a c h
fa m ily w ere in o c u la te d w ith e a c h h u n t r a c e .
-1 5 -
EXPERIMMTAL RESULTS
Awne dn.es B
The Eg g e n e r a tio n c o n s i s t i n g o f s i x f a m i l i e s o f from 94 t o
108 p l a n t s e a c h w ere s tu d ie d i n th e l a b o r a t o r y and c l a s s i f i e d f o r aw nedness.
The b r e e d in g b e h a v io r o f e a c h f a m ily and 68 p l a n t s o f C e re s -(H o p e -
F lo r e n c e ) and 98 p l a n t s o f Renown a re g iy e n i n T a b le I .
c l a s s i f i e d i n t o th r e e b r e e d in g g r o u p s , nam ely:
The p ro g en y w ere
( l ) t r u e ' b re e d in g w e ak ly
a w n le tte d t y p e s , (2 ) t r u e b r e e d in g f u l l y awned t y p e s , and ( 3 ) a p i c a l I y
a w n le tte d ty p e s w hich s e g r e g a te d f o r a l l th r e e c l a s s e s .
The p a r e n ts
w ere homozygous' and c l e a r . c u t f o r t h e i r r e s p e c t i v e aw n. ty p e s and no
q u e s tio n a b le ty p e s w ere fo u n d .
I n th e p ro g e n y , 24 q u e s tio n a b le ty p e s
w ere fo u n d w hich c o u ld n o t be d e f i n i t e l y p la c e d i n e i t h e r th e 3 - o r 3+
c la s s .
They a p p e a re d a s i n t e r g r a d e s b etw een th e s e two c l a s s e s .
These
w ere grown i n th e F^ and found t o be a l l b re e d in g t r u e and w eakly awnl e t t e d e x c e p t t h r e e w hich were s e g r e g a tin g and c l a s s i f i e d a s s t r o n g l y
a w n le tte d .
Of th e 240 F^ rows grown- a t Bozeman and M o cc a sin , o n ly two
l i n e s b re d d i f f e r e n t th a n was e x p e c te d from th e Fg c l a s s i f i c a t i o n .
Out o f 621 p l a n t s c l a s s i f i e d i n th e Fg and c o r r e c t e d a c c o rd in g
t o t h e i r F^ b r e e d in g b e h a v io r , a s g iv e n i n T ab le I , 156 were found t o be
w eakly a w n le tte d ( 3 - ) , 136 f u l l y awned ( 5 ) , and 329 a p i c a l l y .a w n le tte d
(3 + ).
T hese d a t a i n d i c a t e a c lo s e f i t t o a 1 :2 :1 r a t i o w hich may be
e x p la in e d by a s in g le m a j o r - f a c t o r d i f f e r e n c e w ith th e a w n le tte d c o n d itio n
b e in g in c o m p le te ly dom inant to th e f u l l y awned c o n d itio n .
I.
T a b le I
The c l a s s i f i c a t i o n o f p a r e n ts and F3 p l a n t s f o r aw nedness c o r r e c t e d on th e
b a s i s o f F^ b r e e d in g b e h a v io r o f q u e s tio n a b le ty p e s i n a c r o s s o f C e re s - '
(H o p e -F lo re n c e ) w ith Benown grown a t Bozeman i n 1939 and a t Bozeman and
M o cc a sin , M ontana i n 1940
C e re s -(H o p e -F lo re n c e ) x Benown
F am ily and p l a n t number
Humber o f
T o ta l num ber q u e n tie n a b le
o f p l a n t s i n 3 - o r '3+ awn
ty p e s
e a c h f a m ily
f3
c la s s ific a tio n
o f Fg q u e s ti o n ­
a b le ty p e s
3+
3=
Humber o f p l a n t s
a f t e r F., c o r r e c t io n
having I awn ty p e
33+
*
Bow-26 P l a n t - 4
98
8
7
I
25
52
21
Bow-26 P l a n t - 8
108
4
4
O
18
58
28
Bow-2 P la n t- 1 1
94
4
3
I
22
49
23
Bow-26 P l a n t —9
104
6
5
I
27
51
Bow-23 3 1 a n t -7
111
O
O
O
31
61
26
19
2
2
O
27
58
19
24
21
3
1$6
,329
136
Bow-2 P l a n t -2 .
T o ta l
106
621
Benown C . I . 11709'
98
C e re s -(H o p e -F lo re n c e ) C . I . 11872
68
98
68
»,
T
-1 7 -
The c a l c u l a t i o n o f c h i- s q u a r e f o r g o o d n ess o f f i t to a 1 :2 :1
r a t i o o f th e 621 Fg p l a n t s , a s g iv e n i n T a b le I I , shows th e o b se rv e d
num ber t o be v e r y c lo s e t o th e e x p e c te d .
The P v a lu e f o r g o o d n ess o f
f i t betw een th e o b s e rv e d and c a l c u l a t e d -d a ta i s b etw een 0 .9 0 and 0 . 8 0 .
T a b le I I
C a l c u la tio n o f G oodness o f F i t to
h y b r id r a t i o f o r aw nedness from a
F lo r e n c e ) x Eenown c r o s s grown a t
1939 and a t Bozeman and M o ccasin ,
Fg c l a s s e s b a se d on
b r e e d in g b e h a v io r o f F i
W eakly a w n le tte d ( 3 - )
O bserved
C a lc u la te d on
one f a c t o r
d iffe re n c e
a 1 :2 :1 mono­
Cer e s - ( Hope-'
Bozeman i n
M ontana i n 1940
( o -c )2
o -c
(o -c )2
C
156
155
I
I
0 .0 0 8
329
311
18
324
1 .0 4 2
F u ll y awned (5 )
136
155
-1 9
361
2 .3 2 9
T o ta l
621
621
S tr o n g ly a w r ile tte r (
)
.
X2 = 3 .4 7 7
D eg rees o f freedom ,
=
2
P = b etw een 0 .9 0 and 0 .8 0
K e rn e l Smoothmess
S in c e th e p a r e n t a l v a r i e t i e s o f th e C e re s -(H o p e -F lo re n c e ) x
Eenown c r o s s d i f f e r e d i n d e g re e o f ro u g h n e s s and p i t t e d n e s s o f s e e d , an
a tte m p t h a s been made t o d is c o v e r th e mode o f i n h e r i t a n c e o f t h i s c h a r a c t e r .
S ix Fg f a m i l i e s c o n s i s t i n g o f 635 p l a n t s w ere c l a s s i f i e d i n t o
e i g h t g ro u p s a c c o rd in g t o th e s e c h a r a c t e r i s t i c s :
d e g re e o f ro u n d n e ss o r
a n g u l a r i t y o f c h e e k s , d e p th and w id th o f c r e a s e , and degree o f p i t t e d n e s s .
—3.8—
The c l a s s i f i c a t i o n o f th e Fg and p a r e n t m a t e r i a l i s g iv e n i n T ab le I I I .
A ssum ing from an a n a l y s i s o f th e Fg and p a r e n t d a ta t h a t c l a s s 5 i n th e
b r e a k in g p o in t o r in te r m e d ia te c l a s s betw een th e ro u g h e r and sm o o th er
t y p e s , a p p ro x im a te ly o n e - h a lf o f th e p ro g e n y a p p ro a c h e s a l i k e n e s s o f
th e sm o o th e r Eenown p a r e n t and o n e - h a lf l i k e th e ro u g h e r C e re s-(H o p e F lo r e n ce) p a r e n t .
T h is i s a ls o t r u e o f f i v e o f th e Fg f a m i l i e s when
c o n s id e re d i n d i v i d u a l l y .
H ow ever, one o f th e Fg f a m i l i e s (.R2P2) had a
g r e a t e r p e rc e n ta g e o f p l a n t s re s e m b lin g th e ro u g h e r p a r e n t ty p e and more
n e a r l y a p p ro a c h e s a 3 :1 th a n a 1 :1 r a t i o .
T a b le I I I
C l a s s i f i c a t i o n i n t o k e r n e l sm o o th n ess c l a s s e s o f
p a r e n ts and Fg p ro g e n y from a c r o s s o f C e re s-(H o p e F lo r e n c e ) w ith Benown grown a t Boaeman-, M ontana i n
1939
K e rn e l
sm o o th n ess
P a r e n ts
c la s s e s
Eenown C .H .F .
E26P8
6
3
3
41
10
4
29
3
5
14
6
6
2
.
E26P9
Fp p ro g en y
E23P7 E23P14
E2P2
E2P11
TF?
o ta l
17
4
3
7
15
30
15
24
4
98
27
28
33
28
23
7
146
6
24
15
17
23
19
13
111
23
20
31
15
21
24
33
144
7
24
16
13
9
9
4
28
79
8
13
6
5
4
4
9
28
9
I
2
3
2
I
2
2
12
70
108
HO
114
104
103
96
635
T o ta l
96
■
-1 9 -
P h o to g ra p h s o f th e c l a s s ra n g e s o f k e r n e l sm o o th n ess fo u n d
i n th e Fg a re in c lu d e d i n F ig u re I .
The Eenovm p a r e n t re s e m b le s th o s e
ty p e s i n c l a s s e s , 2 , 3 , and 4 w ith th e g r e a t e r fre q u e n c y o f c l a s s 3 and
th e O e re s -(H o p e -F lo re n c e ) p a r e n t more n e a r l y re s e m b le s th o s e i n c l a s s e s
6 , 7 ? and 8 w ith a g r e a t e r fre q u e n c y o f c l a s s 7 ®
I n th e F3 , t h r e e o f th e Fg f a m i l i e s w ere s t u d i e d —B26P9 grown
a t boseman and E2P2 and E2P11 grown At M o cc a sin , M ontana.
A com plete
c l a s s i f i c a t i o n o f e a c h Fg l i n e and F3 p l a n t s o f th e s e l i n e s , and o f e a c h
p a r e n t i s g iv e n i n T a b le IV .
A co m p ariso n o f th e s ta n d a r d d e v ia t io n
o f th e p a r e n ts grown w ith th e Fg a t Bozeman i n 1939 and w ith th e F^
a t Bozeman and M occasin i n 1940 a r e a s f o llo w s :
Bozeman 1939; Eenown
3 . 7 - 1 . 0 ; O e re s-(H o p e -F lo re n c e ) 6 . 7 - 1 . I ; Bseman 1940, Eenown 3 . 8 - 1 . 2 5
C e r e s -(H o p e -F lo re n c e ) $ . 6 - 1 . 5 ; M o ccasin 1940, Eeno1Wn 3 . 2 - 1 . 0 ; C e re s (H o p e -F lo re n c e ) 6 .8 - 0 .9 7 .
The d e v i a t i o n s o f th e 1939 Bozeman p a r e n ts
and th e 1940 M occasin p a r e n ts a re v e ry s i m i l a r , w h e rea s th e 1940 Bozeman
p a r e n ts have a much l a r g e r s ta n d a r d d e v i a t i o n a n d , t h u s ,' much more v a r i a b l e
i n t h e i r ra n g e o f c l a s s i f i c a t i o n .
U n fo rs e e n c o m p lic a tio n s i n th e e n v ir o n ­
m e n ta l c o n d itio n s a re p ro b a b ly th e m ain c a u s e s o f t h i s g r e a t e r v a r i a b i l i t y .
G rouped c l a s s sum m aries o f th e b re e d in g b e h a v io r o f th e t h r e e
Fg f a m i l i e s s tu d ie d i n th e F^ a r e g iv e n i n T a b le V.
G rouped c l a s s e s w ere
a l s o d e te rm in e d by th e p ro b a b le e r r o r o f th e p a r e n ts b u t th e g ro u p m ethod
a s u s e d i n T a b le V seemed more s a t i s f a c t o r y s in c e f r a c t i o n a l num bers a s
c l a s s e s w ere a v o id e d and th e l i n e s e g r e g a tio n s i n th e F^ f e l l more e a s i l y
and d e f i n i t e l y i n t o th e 2 , 3 , and 4 ; 4 , 5 , and 6 ; and 6 , 7 , and 8 g r o u p s .
B oth g ro u p in g m ethods i l l u s t r a t e d a p p ro x im a te ly th e same s e g r e g a tio n a l
Cl*** 2
C la s s S
Figure I
C l* .. 4
Class 6
Range of kernel smoothness found in the F2 of a cross between
Ceres-(Hope-Florence) and Renown
T a b le I v
The o l a s s i f i c a ti o n o f p a r e n ts and p ro g e n y from a c r o s s o f C e re s -(H o p e -F lo re n c e )
w ith Eenown f o r aw nedness and k e r n e l sm o o th n ess i n F2 l i n e s grown a t Bozeman i n
1939 and F3 l i n e s grown a t Bozeman and M o c c a sin 5 M ontana i n 1940
P a r e n t o r p ro g en y
1940
row
No.
C e re s -(H o p e -F lo re n c e )C . 141872 3003
do
3029
3065
do
do
3083
3101
do
do
. 3119
do
3137
do
3155
3158
do
Eenown C . I . 11709
3027
do
3045
3063
do
3081
do
do
3099
do
3117
do '
3135
do
3153
3156
do
C ro ss
3031
3048
do
do
3087
3106
do
do
3115
3132
do .
do
3143
3092
do
Awnedness
P a re n t; o r
F2
F3
F3
l i n e pheno­■ geno­
ty p e
ty p e
No.
Bozeman
3 -5 -2 4 5
5
3 -5 -2 1 5
5
3 -5 -2 2 5
5
3 -5 -2 3 5
5
3 -5 -2 2 5
5
3 -5 -2 1 5
5
3 -5 -2 4 5
5
9 -8 -2 8 5
5
988-7
5
5
37 -1 0 -3 3 7 -1 0 -2 3 331 -6 -2 4 3 31 -6 -2 4 3 7 -1 0 -1 3 37 -1 0 -1 3 37 -1 0 -2 3 37 -1 0 -5 3 337 -1 0 -7 3 2 6 -9 -1 7 3 326- 9-31 5
5
26—9—64 3 326- 9 -8 9 5
5
3+
26- 9-89 3+
26- 9- 10;I 3+
3+
3+
26- 9- 14:} 3+
26- 9-69 3 3-
P a re n t
and F2
c la s s i­
fic a tio n
.
'7
7 •
8
8
8
7
7
7
6
3
3
4
4
2
2
3
4 •
3
3
3
3
3
3
3
3
4
K e rn e l sm o o th en ess c la s s e s
P a r e n ts and F3
No. o f p l a n t s
2 3 4 5 6 7 8 9 Mean
6
7
4
I
>.
2
I
8
8
6
9
3
5
16
8
7
4
7
4
I
4
5
I
4
6
I
I
6
3
6
2
5
I
9
'5
6
4 3
2 5
I 3
4 7
3 9
8 4
5 6
7 4
4 10
2
2
4 5
4
2
5 5
9 10
7 9
8 5
7 3
3 6
7 7
7 3
4 6
I I
*1 I
3
I
4
2
7
5
7
I
5
3
I
I
4
5
3
4
5.8
3 .2
2 .9
4.8
3.5
3.3
2.3
4.2
4 .6
4.9
I
4 .4
3 .6
5 .1
4 .2
I
3
I
4
8
4
4
4 .2
5 .6
7 .4
6 .3
6 .3
4 ,7
5 .6
5 .2
6
2
I
6
I
4.3
5.4
5
7
7
7 .3
6 .7
Table IV
( co n tin u ed )
P a r e n t o r p ro g en y
C ro ss
-
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do.
do
do
do
do
Avmedness
P a re n t o r
1940 F3
F2
F3
row
l i n e p h en o - g e n o No.
No.
ty p e ty p e
269
-7
2
5
3095
5
3096 26- 9 -7 3 5
5
3102 26- 9 -7 6 5
. 5
3098 2 6 -9 -7 5 5
5
3113 26- 9 -8 7 5
5
31122 26- 9 -9 3 3+
3+
3130 26- 9- 1013+
3+
3131 26- 9- 1023+
3+
3142 26- 9- 1103+
3+
3144 26- 9- 1123+
3+
3145 26—9—1133+
3+
3146 26- 9- 1143+
3+
3147 26- 9- 1153+
3+
3086 26—9—63 3+
3—
3090 26-967 3 33106 26- 9 -8 0 5
5
3107 26- 9 -8 1 5
5
3127 26—9—98 3+
3+
3150 2 6 -9-1183+
3+
3139 26—9—107 3+
3+
3084 26- 9 -6 1 333085 26- 9-62 333088 26—9—65 3—
33091 26- 9 -6 8 333097 26- 9 -7 4 5
5
3104 2 6 -9 -7 8 5
5
3105 26- 9 -7 9 5
5
' 3109 26- 9 -8 3 5
5
3110 26- 9 -8 4 5
5
P a re n t
and F2
c la s s !flo a tio n
4
4
4 .
4
4
3
4
4
S
4
4
4
4
5
5
5
5
5
5
5
6
6
6
6
6
6
6
6
6
2
4
2
3
I
I
2
4
K e rn e l sm oothness c la s s e s
P a r e n ts and !F3
No. ojf p l a n t s
3 4 5 6 7 8 9 Mean
5 3 6 d
4 .7
2 5 6 6 I
4 .9
I 5 10 3 I
5 .9
4 7 6 3
5 .4
6 4 5 I
3 .7
I 6 9 2 I I
5 .0
I 3 6 7 3
5 .4
5 2 7 6
4 .7
4 6 6 2
3 .9
2 8 7 2 I 6 .6
6 9 5
6 .0
2 2 9 6 I
6 .1
I I 4 6 6 2 7 .1
3 6 8 2 I 6 .6
2 9 5
2 I I 4 .9
2 7 5 ■5 I
5 .8
7 7 3
3 .6
5 8 3 4
4 .3
2 4 6 7 I
6 .0
I I 2 6 4 6
6 .5
5 2 5 5 2 6 .6
I I 10 4 4
6 .5
I 3 6 4 6 7 .5
3 5 5 4 I I I 5 .1
' 5 4 6 4 I
5 .6
2 9 7 I
4 .3
10 6 3
3*6
8 7 3
3 .6
2 7 4 3
4 .0
T a b l e IV
( c o n tin u e d )
1940
ro w '
Ho,
P a re n t o r p ro g en y
C ro ss
do
do
do
do
do
do
do
•do
do
do
do
do
do
do
do
do
do
do
do
do
do
3111
3116
3121
3123
3124
3126
3133
3138
•3141
3149
3151
3089
. 3094
3103
3104
3112
3114
3148
3152
3093
3108
3125
Awnedness
- P a re n t1 o r
F3
F2
(ff3
l i n e p h e n o -g e n o H o. ty p e ty p e
Bozeman
2 6 -9 -8 5
5
5
2 6 -9 -9 0
3+
3+
2 6 -9 -9 2
3+
3+
3+
2 6 -9 -9 4
3+
26-9-95
3+
33+
3+
26-9-97
26- 9-1 0 4 3+
3+
26- 9-106 3+
3+
26- 9-109 3+
3+
26- 9-117 3+
3+
26- 9-1 1 9 3+
3+
26—^—66
3326- 9 -7 1
3326- 9-77
5
5
26- 9 -7 8
5
5
26*—
5
5
26-9-88 .5
5
26- 9-1 1 6 3+
3+
36- 9 -1 2 0 3+
3+
26- 9- 7©
3326- 9 -8 2
5
5
26- 9 -9 6
3+
3+
P a re n t 6
and F2
c la s s i­
fic a tio n
6
6
6
6
6
6
6
6
6
6
6
7
2
5
I
2
K e rn e l sm o o th n ess c la s s e s
P a r e n ts and F3
Ho. o f p l a n t s
3 4 5 6 7 8 9 Mean8
I
I
4
I
3
2
I
8
I
7
I
I
6
7
7
7
7
7
7
7
8
8
8
3
4
5
9
9
I
3
I
I
4
3
7
3
7
6
5
3
7
5
2
2
2
2
2
4
I
5
11
5
4
3
7
7
I
6 I
5 2
4 I
2
7 3
6 4
5 3
2 4
6 7
3 7
4 6
5 ■7
5
8
2
7
3
5
8
4
3.3
I
5-3
4.9
4 .7
4 .0
2
2
7
5
3
4
7
3
5.9
5.9
3
7
3
3
2
2
6
7
5
/
6
7
4
I
3
I
I
2
7 .2
7 .7
6 .4
7 .0
7 .5
6.8
3 .7
7 .5
6 .0
6 .7
I
6.3
6.7
5.3
5.3
4.9
2 7 1
3 2
1 3 5
1 6 3
6 .9
5 .5
7 .2
7 .2
I
M occasin
C e r e s - ( H o p e -P lo re n c e ) C»1 .1187 2
do
do
do
4003
4029
4047
4065
9- 8 -2 8
9 -8 -2 7
89- 8 -2 9
9 -8 -2 7
5
5
5
5
5
5
5
5
7
8
7
8
2
3
1
T a b le IV
( c o n tin e u e d )
P a r e n t o r p ro g en y
C e re s ^ H o p e -P lo re n c e ) C. I .11872
do
do
do
do
Eenown C . I .11709
do
do
do
do
:>
do
do
do
do
C ro ss
do
do
do
do
do
do
do
do
do
do
. do
do
do
do
. .
1940
EOW
Ho.
&P83
4101
4119
4137
4155
4001
4827
4045
4063
4081
4099
4117
4135
4153
4022
4057
4059
'*4060
4067
4012
4014
4015
4018
4020
4032
4035
4044
4049
4050
F3
lin e
Ho.
3 -5 -2 6
3- 5-26
3- 5-25
3- 5-25
9- 8 -2 9
1- 6 -2 5
1- 6 -2 6
1- 6 -2 5
1- 6 -2 6
1 -6 -2 7
1 -6 -2 7
1- 6 -2 8
1 -6 -2 8
7 -1 0 -4
2 -2 -1 1
2 -2 -4 2
2 -2 -4 4
2 -2 -4 5
2- 2 -4 9
2 -2 -1
2 -2 -3
2 -2 - 4
2 -2 -7
2 -2 -9
2 -2 -2 0
2 -2 -2 3
2 -2 -3 2
2 -2 -3 4
2 -2 -3 5
Awnedness
P a re n t o r P a re n t
and F2
F2
F3
phenogeno- - c l a s s i ­
ty p e ty p e f i c a t i o n
7
5
5
7
5
5
6
5
5
6
5
5
7
5
5
3
3333
32
333. 4
334
33
3— 3 3
334
33• 2
,
3—
32
332
3+
3+
2
3+
3+
2
3+
3+
2
3+
3+
333
3
3333
333
3333
3
333
3+
3+
3+
3+
3
3+
3
3+
3
3+
3+
K e rn e l sm o o th n ess c la s s e s
P a r e n ts arid F3
Ho . o f p l a n t s
2 3 4 5 6 7 6 9
Mean
6 3 1
6 .5
I 5 3
7 .2
2 5 3
7 .1
3 3 4
7 .1
2 4 3 I
6 .3
I 2 2 3 2
4 .3
3 .0
3 5 I I
2 .1
9 I
4 .0
5 I 3 I
3 4 I 21 I
3 .5
I 5 3 I
3 -4
2 I 3 4
4 .9
5 3 2
2 .7
6 4
2 .4
I 3 4 I
5 .2
I 3 2
4
4 .1
4 .1
9 I
3 5 2
6 .9
5 .0
4 3 2 I
I 4 4
I
5 .6
I 3 4 2
5 .7
3 4 2 I
5 .1
2 2 3 3
■ 6 .7
I I 3 5
2 .5
I 3 3
2 I
4 .2
I
3 6
2 .9
I 5 i 2 I
3 .8
I 2 2 4 I
5 .3
I 5 I 3
5 .8
&
f
‘
T a b le IV
( c o n tin u e d )
p a r e n t o r p ro g en y
C ro ss
do
do
do
d9
do
do
do
do
do
do
do
do
do
do
do
do
do
- do
do
do
do
do
do
do
do
do
do
do
1940
rov/
Ho.
4056
4061
4062
4066
4075
4077
4030
4031
4033
4034
4036
4039
4040
4041
4052
4058
4069
4073
4074
4070
4080
4102
. 4114
4120
4025
- 4038
4043
4054
4055
Awnedness
P a re n t o r
F2
F3
F3
p h e n o -g e n c
lin e
ty p e ty p e
Ho.
3+
3+
2 -2 -4 1
2- 2 -4 6
3+
3+
Sn
3+
2 -2 -4 7
2 -2 -4 8
3+
3+
2 -2 -5 7
5
5
5
. 2- 2 -5 9
5
2 -2 -1 8
33™
3— 32 -2 -1 9
2 -2 -2 1
3+
3+
2 -2 -2 2
3+
3+
3+
3+
2-2-24
3+
3+
2 -2 -2 7
2-2-28
3+
3+
3+
3+
2 -2 -2 9
3+
3*4"
2 -2 -3 7
3+
3+
2—2—43
3+
3+
2- 2 -5 1
5
5
2 -2 -5 5
2- 2 -5 6
5
5
2- 2 -5 8 . 5
5
332—2—13
2 -1 1 -1 6
5
5
2- 11-28
33332- 11-31
2 -2 -1 6
332- 2-26
3+
3+
3+
3+
2 -2 -3 1
3+
3+
2 -2 -3 9
2 -2 -4 0
3+
3+
P a re n t
and F2
c la s s iflo a tio n
3
3
3
3
3
3
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
5
5
5
5
5
K e rn e l sm o o th n ess
______
P a r e n ts and
Uo. o f p l a n t s
2 3 4 5 6 7 8
4 5
I I 4- 4
I 3 5 I
I I I 41 2 I
I
4 5
3 4 2 I
I 5 2 3
I
3 6
I 3 2 3 I
4 I 4 I
3 4 3
I 2 2
I
2
3 2 2 I I I
2 5 I 2
I 3 2 2 2
I 3 2 3
2 4 4
I 4 3 2
2 4 3 I
I 5 3 I
I 3 3 2 I
2 5 3
2 4 4
I 2 5 2
2 3
2' 3
I 5
3 I
3 2 4 I
2 5 3
I
7 2
c la s s e s
F3_____
9
Mean
5.5
5 .1
5.6
5.8
4 .3
5 .2
3 .7
2 .9
2
I
4.0
4.2
3.0
6.5
4.8
3.3
6.1
7.0
6 ,0
6 .6
3.3
6 .4
5.9
6 .1
7 .2
6.8
4 .1
5.2
4.3
6 .1
5.9
T a b le IV
( c o n tin e u d )
P a r e n t o r p ro g en y
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do .
do
do
do
do
do
1940
F3
lin e
row
Ho.
Ho.
. 4068 2- 2 -5 0
4071 2 -2 -5 3
4072 2 -2 -5 4
4086 2- 1 1 -3
4087 2 -1 1 -4
4090 .2 -1 1 -7
4093 2 -1 1 -1 0
4106 2 -1 1 -2 0
4116 2 -1 1 -3 0
4121 2 -1 1 -3 2
4131 2 -1 1 -4 2
4132 2 -1 1 -4 3
4141 2 -1 1 -4 9
4148 2 -1 1 -5 6
4152 2- 11-60
4016 2- 2 -5
4017 2 -2 -6
4019 2 -2 -8
4021 2—2—10
4023 2 -2 -1 4
4024 2 -2 -1 5
4026 2 -2 -1 7
4037 2 -2 -2 5
4042 2- 2 -3 0
4048 2 -2 -3 3
. 4053 2 -2 -3 8
4070 . 2- 2-52
4978 2- 2 -6 0
4079 2 -2 -1 2
Awnedness
P a re n t o r
F2
F3
p h e n o -g e n o ty p e ty p e
3+
3+
5
5
5
5
5
5
5
5
5
5
5
5 .
333■3333+
3+
3+
3+
3+
3+
3+
3+
3+
3+
333333333333333+
3+
3+
3+
3+
3+
3+
3+
5
5
5
5
33-
P a re n t
and. F2
c la s s !f i c a ti o n
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
6
6
6
6
6
.6
6
6
6
6
6
6
6
6
2
I
I
I
I
I
K e rn a l sm o o th n ess
P a r e n ts and
Ho. o f p l a n t s
3 4 5 6 7 8
9 I
2 6 I
I 5 4
I 6 3
I I 3 5
3 4 I 2
3 4 I 2
2 4 3
2 I 3 4
2
3 3 2
I 5 3 I
3 I 2 3
4 6
2 4 2 I
3 4' 3
3 6 I
2 4 4
3 5 I I
I 4 3 2
2 I I 3 3
2 I I 2 4
I 3 2 2
I I 2 3
2
3 3 3
5 I I I I
I 3 6
3 3 4
I 2 4 3
2 5 3
c la s s e s
F3
9
.1
I
Mean
6 .1
4 .3
5 .3
6 .2
5 .2
5 .2
6 .2
7 .3
6 .9
6 .3
6 .5
6 .8
5 ,6 .
6 .2
" 5 .0
6 ,8
7 .2
6 .0
6 .6
6 .4
5 .6
4 .3
5 .3
3 .8
3 .9
7 .5
7 .3
6 .9
7 .1
Table IV
( con tin u ed )
P a r n e t o r p ro g e n y
i
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
do
1940
row
No.
4084
4085
4091
4092
4097
4105
4107
4111
4124
4126
4127
4133
4142
4146
4U 7
4149
4151
4013
4051
4094
4096
4098
4103
4108
41094113
4115
4122
4123
F3
lin e
No.
2 -1 1 -1
2 -1 1 -2
2 -1 1 -8
2 -1 1 -9 '
2 -1 1 -1 4
2 -1 1 -1 9
2 -1 1 -2 1
2 -1 1 -2 5
2 -1 1 -3 5
2 -1 1 -3 7
2—11—38
2—11—38
2- 11-50
2 -1 1 -5 4
2 -1 1 -5 5
2 -1 1 -5 7
2 -1 1 -5 9
2 -2 -2
2—2—36
2 -1 1 -1 1
2 -1 1 -1 3
2 -1 1 -1 5
2 -1 1 -1 7
2 -1 1 -2 2
2 -1 1 -2 3
2 -1 1 -2 7
2 -1 1 -2 9
2 -1 1 -3 3
2—11—34
A im edness
P a rn e t o r
E2
F3
pheno geno
ty p e ty p e
■ 5
5
5
5
5
5
5
5
5
5
33' 33333+
3+
3+
3+
3+
3+
3+
3+
3+
3+
3+
3+
3+
3+
3+
3+
3+
3+
333+
3+
5
5
5
5
5
5
5
5
333333333+
3+
3+
3+
P a re n t
and F2
c la s s !fic a tio n
6
6
6
6
6
6
6
6
6
. 6
6
6
6
'6
6
6
6
7
7
7 7
7
,
7
7
7
7
7
7
7
2
K ernel- is m oothness
P a r e n ts and
No. 1o f p la n ts
3 4 5 6 •7 8
I I 5 3
I 4 4 I
I 4 3 2
3 3 2 3
I 3. 2 4
2 '5 3
6 3 I
I 2 4 2 I
2 5 3
6 4
3 5 2
I 5 3
5 4 I
I 7 I
2 2 6
2 2 3 2 I
I 2 5 2
I 2 4 2 I
2 2 I 2 I 2
3 3 4
4 4 2
I 6 2 I
5 5
I
6 I I
I
3 I 5
I I 6 2
3 6.
.2 3 3 2
2 2 3 2 I
c la s s e s
F3
9
Mean
5 .0
5.5
5 .6
5.3
6.9
I
I
6 .1
5 .0
6 .0
7 .1
7 .4
5 .9
7 .4
5 .6
7 .2
6.4
5.8
5.8
6 .0
5.2
6 .1
5 .8
6 .3
6 .5
6 .1
6.9
I
5 .9
7.8
6 .5
5.8
Table IV
( oon tin eu ed )
P a r e n t o r p ro g en y
1
do- us.
do
do
do
do
do
do
do
do
do
do
do
do
do
.
1940
row
No.
4125
2128
4129
4134
4140
4143
4145
4150
4088
4095
4104
4110
4112
.4139
4130
Awnedness
P a re n t o r
F3
F2
F3
l i n e pheno-geno©'*’
N o.
ty p e ty p e
2- 11-36
3+
3+
3+
3+
2 -1 1 -3 9
2- 11 -4 0
3+
3+
3+
3+
2 -1 1 -4 5
2- 11-48
3+
3+
3+
2 -1 1 -5 1
3+
2 -1 1 -5 3
3+
3+
2- 11-58
3+
3+
2 -1 1 -5
5
5
2- 11-12
5
5
2- 11-18
5
5
32 -1 1 -2 4
32- 11-26
333+
2 -1 1 -4 7
3+
2- 11-41
3+
3+
P a re n t
and F2
c la s s !flo a tio n
7
7
7
7
7
7
7
7
8
8
8
.
8
8
8
9
K e rn e l sm oothness
______P a r e n ts and F3
, No. o f p la n ts
2 3 4 5 6 7 8
3 5 2
3 I 2 4
2 3 5
2 84 5 I
2
3 5 I
I 3 5 I
I 2 I 4 2
I 4 5
2 I 5 I
5 5
4 2 3
2 5 3
2 6 2
I 2 2 2 I
c la s s e s
9 . Mean
5-9
6 .7
7 .3
5 .8
6 .7
6 .5
6 .6
6 .4
5 .3
I
5 .9
7 .5
5 .7
7 .1
6 .0
2
6 .6
03
I
-2 9 -
T a b le 7 C l a s s i f i c a t i o n o f p a r e n t s , 3T2, and F3 f o r k e r n e l
sm o o th n e ss, (F3 p ro g e n y p la c e d i n t o th e g ro u p o f
g r e a t e s t fre q u e n c y )
F am ily R26P9 - Bozeman
F3
_______ F2 c l a s s e s f o r k e r n e l sm o o th n ess
g ro u p s
2
6
3
4
5
I
I
2 -3 - 4
5
4 -5 - 6
8
4
3
9
6 -7 -8
I
3
4
7
G rand t o t a l
F3
g ro u p s
2
I .
2—3—4
4 -5 -6
3
6—7—8
I
Grand t o t a l
F3
g ro u p s
2—3—4
4 -5 - 6
6—7—8
2
7
I
2
5
F am ily E2P2 - M o ccasin
F2 c l a s s e s f o r k e r n e l sm o o th n ess
6
4
3
5
7
■6
. 2
2
• 2
13
3
5
5
I
4
3
9
8
3
9
T o ta l
------------ 8----29
20
8
9
F am ily E2P11 - M o ccasin
F2 c l a s s e s f o r k e r n e l sm oothness
3
4
5
6
.7
8
9
4
8
4
4
3
10
7
4
13
2
T o ta l
22
35
G rand t o t a l
.F 3
g ro u p s
T o ta l
11
31
18
60
57
3
P a r e n t s - Bozeman
K e rn e l sm o o th n ess c l a s s e s
4
5
T o ta l
Eenown
4
3
5
2
2 -3 - 4
3
4 -5 - 6
2
6 -7 -8
G rand t o t a l __________ _______.______ ._____
9
_____________ ___ ______ P a r e n ts - M occasin
F3
________ K e rn e l sm ooth n ess c la s h e s
g ro u p s
2
5 T o ta l 6
3
4
Eenown
• 2
2
2 -3 - 4
3
7
4—5—6
2
2
6 -7 -8
2
Grand t o t a l
9
6
7
8
9
0 . H. F.
T o ta l
6
.6
3
3
______ __________________ __________ 9
7
5
8
C .H .F .
I
I
9
T o ta l
I
8
9
—3 0 -
t r e n d s o f th e p ro g e n y .
The F3 l i n e s w ere p la c e d i n th e g ro u p s a c c o rd in g
to t h e i r g r e a te s t re s p e c tiv e fre q u e n c ie s .
S in c e t h e r e was more v a r i a t i o n i n th e p a r e n t a l m a te r ia l a t
Bozeman th a n a t M o cc a sin , i t was e x p e c te d t h a t t h e r e s h o u ld he a g r e a t e r
v a r i a t i o n i n th e p ro g e n y .
T h is a n a lo g y seem s t o he t r u e , b ecau se th e
F3 l i n e s a t Bozeman a p p a r e n tly s e g r e g a te d l e s s d e f i n i t e i n r e g a r d s t o
th e e x p e c te d from th e Fg c l a s s i f i c a t i o n th a n th e M o ccasin F3 m a t e r i a l .
F a m ily B26P9 w hich was grown a t Bozeman d o es n o t seem t o a p p ro a ch any
d e fin ite h e rita b le r a tio .
Some s e g r e g a tio n f o r e v e ry F^ g ro u p o c c u r re d
i n a l l Fg c l a s s e s w ith a dom inance o f th e ro u g h e r ty p e s se e m in g ly a p p a r e n t.
I n fa m ily B2P2 w hich was grown a t M o c c a sin , th e F2 was c l a s s i f i e d m o s tly
i n t o c l a s s e s 2 , 3 , 4 , 5? and 6 .
The g r e a t e r p e rc e n ta g e o f th e Fg sm o o th e r
ty p e s s e g r e g a te d i n t o th e sm o o th er c l a s s e s i n th e F3 and th e Fg ro u g h e r
ty p e s had th e g r e a t e s t p e rc e n ta g e s e g r e g a tin g i n t o th e ro u g h e r c l a s s e s .
A g a in , a dom inance o f th e ro u g h e r ty p e s seem s t o he e v i d e n t , s in c e 48 F^
l i n e s s e g r e g a te d i n t o g ro u p s 4 , 5 » and 6 , and 6 , 7 , and 8 , and o n ly 11
w ere p la c e d i n g ro u p 2 , 3 , and 4-
The sec o n d Fg f a m ily , R2P11, w hich was
grown a t M occasin i n th e F3 had a m o jo r ity o f i t s p l a n t s f a l l i n t o th e
r o u g h te r c l a s s i n th e Fg.
3 , and 4 sm ooth g ro u p .
i.
I n th e F3 no s e g r e g a te s w ere fo u n d i n " th e 2 ,
Of th e 57 l i n e s c l a s s i f i e d , 22 f e l l i n t o th e 4 ,
5; and 6 g ro u p and 35 i n t o th e 6 , 7 , and 8 g ro u p .
T h is a g a in i l l u s t r a t e s
a dom inance o f th e ro u g h e r t y p e s .
S e v e r a l m ethods o f g ro u p in g w ere t r i e d h u t no p l a u s i b l e f i t t o a
o n e - o r tw o - f a c t o r r a t i o was o b ta in e d .
I t seems e v id e n t t h a t i f one o r
two f a c t o r s w ould e x p la i n th e d i f f e r e n c e betw een sm o o th n ess and ro u g h n e s s ,
-3 1 -
th e y a re m asked by a c o m b in a tio n o f e n v iro m e n ta l c o n d itio n s and i f
t h e r e i s a t h r e e - o r more f a c t o r d i f f e r e n c e th e num bers u s e d i n t h i s
e x p e rim e n t w ere n o t la r g e enough t o o b ta in s e g r e g a tio n w hich would
e s t a b l i s h su ch c o m p lic a te d i n h e r i t a n c e .
H ow ever, s in c e th e r e s u l t s o f
t h i s e x p e rim e n t i n d i c a t e d a r a t h e r c o n s ta n t b re e d in g d i f f e r e n c e betw een
th e sm o o th er Renown p a r e n t and th e ro u g h e r C e re s -(H o p e -F lo re n c e ) p a r e n t
and th e p ro g e n y s e g r e g a te d f o r s co m p lete ra n g e betw een th e p a r e n t s , i t
seem s l o g i c a l t o assume t h a t kernel sm o o th n ess fo llo w s some d e f i n i t e
scheme o f i n h e r i t a n c e .
K e rn e l L en g th
An a v e ra g e d i f f e r e n c e i n t t h e k e r n e l e ln g th o f C e re s-(H o p e F lo r e n c e ) and Renown s u g g e s te d th e p o s s i b i l i t y o f s tu d y in g th e mode o f
i n h e r i t a n c e o f t h i s c h a r a c t e r i n th e c r o s s o f th e s e two v a r i e t i e s .
A ll
d i f f e r e n c e s i n th e p a r e n ts and p ro g e n y w ere d e te rm in e d by th e t o t a l le n g t h
i n m i l l i m e t e r s o f 15 k e r n e l s .
The k e r n e l m easurem ents from p l a n t s o f s i x Fg f a m i l i e s and
p a r e n t s a re re c o rd e d i n T a b le V i.
I n 1939 C e re s -(H o p e -F lo re n c e ) k e r n e l s
ra n g e d i n le n g th from 101 t o 108 mm. w ith a mean o f 1 0 4 .7 - a s ta n d a r d
d e v i a t i o n o f 2 .0 mm. and Renown from 91 t o 109 mm. w ith a mean o f 97»4—2 .2
mm.
The Fg l e n g th s ra n g e d from 92 t o 109 mm. w ith a mean o f 1 01.1 mm.
Out o f 560 Fg p l a n t s c l a s s i f i e d , a p p ro x im a te ly 77 p e r c e n t f e l l i n t e r m e d i a t e
betw een th e means o f th e p a r e n t s .
C la s s e s o f th e Fg w ere s e t up by a d d in g
and s u b t r a c t i n g th e p ro b a b ly e r r o r o f e a c h p a r e n t from i t s mean and u s in g
t h e s e ra n g e s a s p a r e n t a l c l a s s ty p e s and th e v a lu e d o u ts id e and betw een
—32—
T a b le VI
T o ta l
le n g th
i n mm.
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
109
Renown
I
0
0
4
12
13
7
10
10
8
2
2
2
T o t a l? I
T o ta l le n g th i n m i l l i m e t e r s o f 15 k e r n e l s from p a r e n ts
and 560 Pg p l a n t s grown a t Bozeman, M ontana i n 1939
C. nyp
P a r e n ts and Fp f a m i l i e s
R23P14 E2P2
R26P8
R2609 R23P7
2
6
5
7
9
6
5
5
4I
0
0
I
3
8
5
13
9
21
13
14
’ 5
6
3
I
I
0
0
I
7
5
7
6
6
15
14
15
7
4
4
4
I
4
I
8
8
8
11
17
9
21
6
7
3
2
0
0
I
45
103
97
106
I
I
0
12
17
10
9
18
9
5
3
0
3
I
2
2
I
7
8
13
13
18
6
7
2
I
0
I
91
79
M o f Renown = 9 7 .4
P .E .
= ± I •5
M o f C .H .F . =104.7
P .E .
= ± I .4
M o f C ro ss
=101.1
R2P11
I
2
I
9
9
8
18
12
8 ■
9
6
3
3
3
I
I
94
P2
to ta l
o f each
le n g t h
5
2
9
12
20
29
60
43
81
64
86
55
41
17
14
14
5
3
560
--3 3 -
th e s e ra n g e s a s t h r e e o t h e r c l a s s e s .
C l a s s i f i c a t i o n o f th e t o t a l num ber
o f Fg p l a n t s i n t o th e s e f i v e c l a s s e s i s a s f o llo w s :
9 0 .1 t o
9 6 .O 48 p l a n t s
Benown l i k e c l a s s
96•I to
9 9 -0 132 p la n ts
I n te r m e d ia te c l a s s
99*1 t o
1 0 3 .0 266 p l a n t s
. 1 0 3 .1 t o
1 0 6 .0 72 p l a n t s
1 0 8 .1 t o
1 0 9 .0 22 p l a n t s
C e re s -(H o p e -F lq re n c e ) l i k e c l a s s
I n th e F3 , one f a m ily , E26P9, w h ich was grown a t Bozeman was
s tu d ie d f o r k e rn e l le n g th .
The m easu rem en ts o f k e r n e l s from 20 F3 p l a n t s
from e a c h o f 57 Fg l i n e s o f t h i s f a m ily , t h e i r mean l e n g t h s , t h e i r Fg
l e n g t h s , and th e same d a t a f o r n in e l i n e s o f e a c h p a r e n t grown w ith th e
p ro g e n y a re g iv e n i n T a b le T H . - Much more v a r i a t i o n and o v e rla p p in g
o f th e p a r e n ts o c c u rre d i n 1940 th a n i n 1939«
I n a r e s p e c t i v e co m p ariso n
o f th e s ta n d a r d d e v i a t i o n s o f e a c h .p a r e n t grown i n 1939 and 1940.
C eres-
(H o p e -F lo re n c e ) was 1 0 4 .7 - 2 .0 and 1 0 0 .9 - 4 .0 and Benown was 9 7 * 4 -2 .2 and '
9 4 .9 * 3 * 4 .
A lso th e mean o f e ac h p a r e n t w.as a p p ro x im a te ly th r e e m i l l i ­
m e te rs l e s s i n 1940 th a n i n 1939.
I n T ab le T i l l a re g iv e n th e Fg c l a s s i f i c a t i o n and P3 b r e e d in g
b e h a v io r o f th e 57 l i n e s a s d e te rm in e d on th e b a s i s o f th e p ro b a b le e r r o r
l i m i t s o f th e p a r e n ts w hich w ere grown a lo n g w ith th e Fg and F3 .
The Fg
o f t h i s f a m ily s e g r e g a te d s i m i l a r t o th e t o t a l Fg p l a n t s o f a l l s i x
f a m i l i e s , t h a t i s , 32 p e r c e n t l i k e th e Benown and s h o r t e r c l a s s , 51 p e r
c e n t i n th e in te r m e d ia te c l a s s and 17 p e r c e n t l i k e th e G e r e s - ( HopeF lo r e n c e ) and lo n g e r c l a s s .
seem s t o be s u g g e s te d .
A weak dom inance o f th e s h o r t e r k e r n e l ty p e
H ow ever, th e F3 s e g r e g a tio n o f t h i s fa m ily seem s
I
'NI
-3 4
T a b le V II T o ta l le n g t h i n m i l l i m e t e r s o f 15 k e r n e l s o f p a r e n ts and p ro g jgny from a c r o s s o f C eres-(H ope-FliD B ence) wijth Eenown
grown i n th e Pg (1939) and
(1940) a t Bozemanj M ontana
I
}
1940
row
Ho.
P a re n t o r p ro g e n y
C e re s -(H o p e -F lo re n o e ) CyI , 11709 3003
3029
do.
do o
3065
3083
do.
3101
do.
d®.
3119
dO«
3137
do.
3155
3158
d o.
Eenown C. I . 11872
302?
do.
3045
do.
3063
3081
do.
do.
3099
do.
3117
do.
3135
do.
3153
3156
do.
C ro ss
30&4
do.
3085
3086
do.
do.
3087
3088
do.
3089
do.
3090
do.
do.
3091
do.
3092
d o.
3094
do.
3095
3096
do.
d o.
3097
3098
do.
3102
do.
do.
3103
do.
3104
do.
3105
3106
do.
do.
3107
3108
do.
• I inF3
d
Ho.
3 -5 -2 4
3 -5 -2 1
. 3 -5 -2 2
3- 5 -2 3
3-5-22
3 -5 -2 1
3 -5 -2 4
9- 8-28
9- 8 -7
7 -1 0 -3
7- 10-2
I - 6-24
1-63:24
7- 1 0 -1
7- 1 0 -1
7- 1 0 -2
7 -1 0 -5
7-IO -7
26-9-61
L ength
i n mm.
o f p a r —e n ts &
___ E e_____ 88
I 07
103
106
105
106
103
107
101
105
96
99
95
95
96
101
26- 9-65
26- 9 -6 6
26—
26-9~6S
26- 9 -6 9
26- 9-71
26- 9-72
2 6 -9 -7 3
26—9—74
2 6 -9 -7 5
26- 9 -7 6
2 6 -9 -7 7
96
2 6 -9 -7 9
26-9-80
26- 9-81
26- 9-82
92
93
I ■ 2
2
2
I
2
I
.1
2
I
5
3
3
I
4
3
I
I
I
2
2
I
2
3
2
I
2
2
2
I
I
I
3
2
I
96
98
26-9-64
91
'
100
100
90
I
I
96
99
26- 9-62
2 6 -9 -6 3
26-9-78
89
2
99
I
2
I
I
3
3
I
I
I
I
I
97
103
102
101
101
102
96
107
103
103
102
101
101
97
101
99
I
I
I
I
I
2
I
2
I
2
2
I
2
I
2
I
3
I
I
I
3
I
T o ta l le n g th o f 15 k e r n e l s in m i l l i m e t e r s
Humber o f F ^ p la n ts w ith e ac h le n g th !
94 95 96 97 98 99 100 101 102 103 104 105 106 107
I
I
2
2
2
I 11
4
5I
I
I
2
2
2
3
3
5
2
I
I
2
2
I
I
I
I
2
2
I
2
I
3
3
4
2
I
2
2
2
2
2
3
I
5
2
I
2
2
3
3
4
3
I
3
I
3
3
3
3
3
2
I
I
I
I
3
4
3
I
2
2
2
I
4
4
4
1
2
2
I
2
I
3
2
1
2
2
2
I
I
3
5
1
I ■ 3
I
2
1
2
I
2
I
3
3
4 ■2
3
2
I
2
2
I
2
3
3
3
I
I
I
2
I
3
I
2
I
I
I
3
3
I
2
I
4
3
5
I
I
I.
3
4
3
4
I
I
2
2
I
I
5
4
4
I
2
I
I
2
4
4
2
2
I
3
2
3
I
3 ' 2
2
I
2
2
3.
I
I
3
4
2
I
2 .4
I
3
3
3
I
I
I
2 .- 3
2
2
2
I
I
3
2
2
I
I
2
2
I
2
I
2
3
I
I
2
I
I
I
6
3
I
3
2
I
2
2
3
I
3
3
3
(I
2
2
2
5
4
3
I
I
I
I
2
3
4
3
4
2
I
2
2
2
2
I
I
4
2
I
I
2
2
I
3
I
2
I •4
I
3
I
4
3
2
I
2
I
2
2
I
5
4
2
3
2
I
3
I
2
4
I
I
I
2
I
2
I
4
I
I
I
2
2
I
2 I
I
I
I
I
I
2
I
2
3 • I . 2
I
I
4
I
2
2
I
2
I
2
4
5
I
I
L
-Mean
le n g th
102.5
1 0 0 .4
95.3
1 0 0 ,9
1 0 2 .1
1 0 2 .0
1 0 3 .4
9 7 .2
1 0 2 .5
9 4 .4
9 7 ,4
9 3 .1
9 4 .9
9 2 .5
9 7 ,9
9 3 ,9
94.I
95.5
9 7 ,0
9 9 .O
9 0 .7
1 0 2 .9
9 8 .1
9 9 .O
1 0 0 .8
9 7 .7
99.3
9 8 .1
98.2
9 9 .4
1 0 0 .5
■ 9 3 .8
95.3
1 0 0 .2
9 6 .5
9 1 .8
1 0 0 .9
1 0 0 .1
102.5
I
(
-3 5 -
i
T ab le Y II (c o n tin u e d )
i
do.
do.
do.
do.
dO.
do.
do.
do.
do.
do.
do.
do.
do.
do.
do.
do.
do.
do.
do.
do,
do.
do.
do.
do.
do,
do.
do.
do.
do.
do.
do.
3149
3150
3151
3152
2 6 -9 -8 3
26- 9 -8 4
26- 9-85
2 6 -9 -8 6
26-9687
2 6 -9 -8 8
26- 9-89
2 6 -9 -9 0
2 6 -9 -9 2
2 6 -9 -9 3
2 6 -9 -9 4
2 6 -9 -9 5
2 6 -9 -9 7
26- 9-98
2 6 -9 -9 9
26- 9-101
26- 9-102
2 6 -9 -1 0 3
26—9—10426- 9-105
26- 9—106
26- 9-107
26- 9-108
26- 9-109
2 6 -9 -1 1 0
2 6 -9 -1 1 1
26- 9-112
2 6 -9 -1 1 3
26—9—114
26- 9-115
26- 9-1 1 6
2 6 -9 -1 1 7
26- 9-118
2 6 -9 -1 1 9
26- 9-120
03
CO
PO
105
102
97
101
102
98
101
105
99
107
105
102
102
101
99
101
100
104
98
99
98
104
102
96
102
100
103
96
97
104
102
102
100
101
99
89
90
91
~2
92
"2
93
-I
94
4
I
3
I
I
I
I
95
3
3
3
I
2
2
I
I
I
I
2
I
I
I
I
I
I
I
2
2
I
I
2
I
2
2
1— I C O 1— I CM
do.
do.
do.
'
3169
3116
3111
3112
3113
3114
3115
3116
3121
3122
3123
3124
3125
3126
3127
3130
3131
3132
3133
3134
3138
3139
3140
3141
3143
3143
3144
3145
3146
3147
3148
T o ta l le n g th o f 15 k e r n e l s i n m i l l i m e t e r s
Number o f
p l a n t s w ith e a c h le n g th !______ _______________________ __
2
I
I
2
I
3
2
2
2
3
I
2
2
I
I
4
4
I
I
2
I
96 97
2
3
6
2
I
3
.2 . 2
I
I
I
I
I
5
I
3
2
2
3
2
2
I
3
2.
I
I
2
4
I
2
2
5
I
3
I
2
3
4
I
3
2
4
2
3
2
5
I
3
5
4
I
4
I
3
I
I
4
2
3
3
4
2
3
98
2
I
3
3
I
I
I
5
3
2
I
3
I
1I
2
I
I
2
4
3
I
I
2
2
2
I
•4
3
I
2
3
I
99
I
2
2
I
4
3
2
3
I
5
I
4
%
I
3
I
2
3
3
3
2
I
4
2
2
I
2
2
I
3
5
3
I
2
3
2
2
4
2
2
3
3
4
4
-2
2
3
3
101 102 103 104
2
2
2
2
3
3
2
4
2
5
i
.3
I
4
2
2
2 • 2
2
2
I
I
2
I
3
I
I
2
I
I
2
I
2
I
I
2
4
2
3
2
I
2
2
I
3
3
3
2
I
2
4
3
2
2
2
I
2
I
2
3
I
2
3
3
3- 3
-6 i '-8
I
4
3
3
I
I
2
I
I
3
2
I
I
I
2
3
4
3
I
3
%
S ro s s
A e
No.
0
0
1—
i
P a re n t o r p ro g en y
1940
row
Ho,
L ength
i n mm,
of p a re n ts &.
I
2
2
I
I
3
2
:-2
I
3
4
106 107
I
I
I
I
I
I
I
IT
I
2
2
I
3
I
2
I
4
I
3
I
I
I
2
I
j.
I
2
I
2
I
I
I
2
i
!3
,
Mean
l e n g th s
9 4 .8
9 7 .5
9 7 .4
9 9 ,7
1 0 0 .8
1 0 0 ,6
9 8 .1
9 8 .7
■ 1 0 0 ,3
9 8 .4
1 0 0 ,5
9 9 ,8
1 0 0 .8
1 0 1 .7
9 9 .6
1 0 1 .4
1 0 1 ,5
9 6 .1
9 8 ,1
9 9 .8
9 8 .1
9 7 ,2
9 6 .9
9 4 .4
9 6 .1
9 4 .1
9 6 .4
9 7 .0
9 6 .5
1 0 1 ,8
9 9 ,5
9 8 .6
9 9 ,0
1 0 0 .6
1 0 0 .7
—36—
t o s h i f t th e dom inance t o th e lo n g e r ty p e k e r n e ls w ith a p p ro x im a te ly
57 p e r c e n t o f th e p l a n t s f a l l i n g i n th e C e re s -(H o p e -F lo re n c e ) c l a s s ,
14 p e r c e n t i n th e in te r m e d ia te c l a s s and 29 p e r c e n t i n Eenown and
s h o rte r c la s s .
T a b le V I I I
F , c la s s e s
by P .B . o f
p a r e n ts
8 9 .0 - 9 2 .5
92 .6 - 9 7 .2
9 7 .3 - 9 8 .1
9 8 .2 - 9 8 .1
1 0 3 .7 -1 0 7 .0
T o ta l
C l a s s i f i c a t i o n o f th e F2 and F^ p ro g e n y f o r
k e r n e l l e n g t h . ( A ll c l a s s e s b a se d on th e
p ro b a b ly e r r o r IinL ts o f th e p a r e n t s — F^
p ro g en y p la c e d i n t o th e c l a s s o f g r e a t e s t
fre q u e n c y )
F2 c l a s s i f i c a t i o n b y P . E . p a r e n ts
” 96 e O 9 6 .1 - 9 9 .0 9 9 .1 - 1 0 3 .0 1 0 3 .1 -1 0 6 .0
I
I
■
2
2
7
3
I
4
3
18
3
9
3
15
30
.
6
1 0 6 .1 -
2
T o ta l
2
14
8
32
2
• 56
Bunt E e a c tio n
A lth o u g h a p a r t o f th e s e i n v e s t i g a t i o n s w ere d e s ig n e d to make
a g e n e t i c s tu d y o f th e r e a c t i o n o f th e p ro g e n y o f th e c r o s s t o two
p h y s io lo g ic r a c e s o f b u n t, i n f e c t i o n s w ere so s l i g h t a t b o th Bozeman and
M occasin i t was d e c id e d from o b s e r v a tio n a l d a ta t h a t a d i f f e r e n t i a l r e ­
a c t i o n c o u ld n o t be d e m o n s tra te d .
w ere su ch t h a t v e ry l i t t l e
A p p a re n tly th e e n v iro m e n ta l c o n d itio n s
i n f e c t i o n was o b ta in e d , b e c a u se th e s u s c e p t i b l e
c h e c k , U lk a 9 had l e s s th a n one p e r c e n t o f sm u tte d h e a d s .
-3 7 DISOUSSIOH MD COWCLUSIOITS
I n th e c r o s s o f C e r e s - ( H o p e ^ F lo re n c e ) (awn ty p e 5) w ith Eenown
(awn ty p e 3 - ) s tu d ie d i n th e F2 and F3 , a m on o h y h rid , 1 : 2 : 1 , r a t i o and
in c o m p le te dom inance more c l o s e l y re s e m b lin g th e a w n le ss p a r e n t was o h - .
ta in e d .
These r e s u l t s a re i n a c c o rd w ith w hat th e m a j o r i t y o f o th e r
w o rk e rs have fo u n d t o be t u r e when a sem i-aw ned w heat i s c ro s s e d w ith a
f u l l y awned o n e .
A p r e lim in a r y a n a l y s i s i n d i c a t e d t h a t th e r e was a d if f e r e n c e
i n k e r n e l sm o o th n ess betw een th e p a r e n ts o f th e C e re s -(H o p e -F lo re n c e ) x
Eenown c r o s s .
At f i r s t th e s u p e r f i c i a l c h a r a c t e r i s t i c s o f th e whole
k e r n e l w ere c o n s id e re d b u t a s th e s tu d y p r o g r e s s e d i t became a p p a re n t
t h e t check s h a p e , d e p th and w id th o f erea® , and p i t t e d n e s s o f th e k e r n e l s
w ere th e more c o n s ta n t c h a r a c t e r s .
Most w heat b r e e d e r s a g re e t h a t th e r e
i s a m arked d i f f e r e n c e f o r th e s e c h a r a c t e r s among v a r i e t i e s grown u n d e r
th e same e n v iro m e n ta l c o n d itio n s and r i g i d s e l e c t i o n i n e a r l y h y b rid
g e n e r a tio n s f o r d e s i r a b l e k e r n e l ty p e s i s p r a c t i c e d by m ost o f them .
S in c e s t r a i n s o f w heat have been s e l e c t e d w hich a re u n ifo rm f o r a c e r t a i n
d e g re e o f k e r n e l sm oothness u n d e r th e same e n v iro n m e n ta l c o n d itio n s , i t
seem s l o g i c a l t o assume t h a t th e s e c h a r a c t e r s must conform t o some d e f i n i t e
shceme o f MendeEan i n h e r i t a n c e .
E nvironm ent i s a m ojo r f a c t o r i n c a u s in g v a r i a t i o n in th e d e g re e
o f k e r n e l sm o o th n e ss.
Wheat grown u n d e r a r i d c o n d itio n s w i l l be more
s h r i v e l e d and more a n g u la r th a n when i t i s grown u n d e r a humid c lim a te .
C la rk and B a y le s (1935) s a y t h a t th e d e p th o f th e c r e a s e i s o f tax o n o m ic
v a lu e o n ly when th e k e r n e l s a re n o rm a lly d e v e lo p e d and i s s u f f i c i e n t l y
—3 8 -
c o n s ta n t t o be u s e d i n f d e s c r i b i n g v a r i e t i e s grown u n d e r i d e n t i c a l and
u n ifo rm e n v iro m e n ta l c o n d itio n s .
T h e r e f o r e , i n a tte m p tin g t o make a
s y s te m a tic g e n e t i c s tu d y o f ..k e r n e l sm o o th n ess i t i s n e c e s s a r y t o s e l e c t
e n v iro n m e n ta l c o n d itio n s w hich a re a s i d e n t i c a l a s p o s s i b l e w ith l i t t l e
v a r i a t i o n from y e a r t o y e a r .
P a r e n t l i n e s and a p p ro x im a te ly 100 p l a n t s e a c h o f s i x Fg
f a m i l i e s w hich h ad been grown sp ac e p la n te d i n an i r r i g a t e d n u r s e r y a t
Bozeman w ere s tu d ie d f o r k e r n e l sm o o th n ess i n th e F g .
T h is p a r e n t and
p ro g e n y m a t e r i a l was s tu d ie d f o r d i f f e r e n t d e g re e s o f k e r n e l sm oothness
and f i n a l l y a ra n g e o f c l a s s e s a s i l l u s t r a t e d i n F ig u re I and d e s c r ib e d
u n d e r M a t e r i a l s and M ethods, page 1 2 , was e s t a b l i s h e d .
Of th e 635 Fg
p l a n t s c l a s s i f i e d , 261 f e l l i n t o c l a s s e s 2 , 3 , and 4 , 111 i n t o c l a s s 5 ,
and 263 i n t o c l a s s e s 6 , 7 , and 8 .
T h ere seems t o be no e v id e n c e o f
dom inance o f e i t h e r p a r e n t a l ty p e o r o f an in te r m e d ia te ty p e .
The same
ty p e o f b re e d in g b e h a v io r o c c u rre d i n a l l th e Fg f a m i l i e s when a n a ly z e d
i n d i v i d u a l l y e x c e p t one w hich had t h r e e - f o u r t h s o f i t s s e g r e g a te s f a l l
i n t o th e 6 , 7 , and 8 c l a s s e s . . I n t h i s f a m ily , B2P11, a dom inance o f th e
ro u g h e r c l a s s e s i s i n d i c a t e d .
P a r e n t a l l i n e s and t h r e e Fg f a m i l i e s w ere s t u d i e d f o r k e r n e l
sm o o th n ess i n th e F3 .
Two o f th e s e f a m i l i e s , one h a v in g 60 Fg l i n e s and
th e o th e r 57, w ere grown u n d e r th e s e m i - a r id c o n d itio n s a t M o ccasin , M ontana
and one f a m ily o f 57 Fg l i n e s i n a n o n - i r r i g a t e d n u r s e r y a t Bozeman.
The
means and s ta n d a r d d e f l a t i o n s o f th e p a r e n t a l l i n e s grown a t Bozeman i n
1939 and a t M occasin i n 1940 were v e r y s i m i l a r b u t th e 1940 Bozeman p a r e n t a l
l i n e s had a much l a r g e r s ta n d a r d d e v i a t i o n th a n e i t h e r th e 1939 Bozeman o r
-3 9 -
1940 M occasin p a r e n t s .
As w ould be e x p e c te d from th e b e h a v io r o f th e
p a r e n t s , th e F3 p o p u la tio n grown a t Bozeman d id n o t seem t o s e g r e g a te
i n any d e f i n i t e m anner, a lth o u g h a dom inance o f th e ro u g h e r ty p e s was
in d ic a te d .
H ow ever, th e M occasin m a t e r i a l i n w hich th e p a r e n ts were
more u n ifo rm seemed t o f o llo w th e t r e n d s s u g g e s te d i n th e Fg c l a s s i ­
fic a tio n .
I t was im p o s s ib le t o v e r i f y th e Fg c l a s s i f i c a t i o n - on th e b a s i s
o f th e F3 b r e e d in g b e h a v io r b e ca u se o f th e d if f e r e n c e i n e n v iro m e n ta l
c o n d itio n s u n d e r w hich th e m a t e r i a l was grow n.
The n u r s e r y a t Bozeman
i n w hich th e F3 p l a n t s w ere grow n, a lth o u g h p la n n e d t o be w ith o u t i r ­
r i g a t i o n , was so lo c a t e d t h a t see p a g e from th e m ain i r r i g a t i o n p r o j e c t s and d i t c h e s s u b i r r i g a t e d a p o r t i o n o f i t .
T h is f a c t p r o b a b ly a c c o u n ts
f o r th e g r e a t e r p a r t o f th e v a r i a t i o n o c c u r r in g w ith in th e . p a r e n ts and
p ro g e n y .
At M occasin th e p r e c i p i t a t i o n was u n ifo rm , b u t th e s o i l i s so
s h a llo w and v a r i e s so much i n d e p th w i t h i n a s h o r t d ita n c e t h a t v a r i a t i o n s
i n p l a n t v i t a l i t y , h e i g h t , and d a te o f r ip e n in g o c c u r r e d .
I t seem s l o g i c a l f o c o n clu d e from th e d a ta p r e s e n te d i n t h i s e x ­
p e rim e n t t h a t i n o r d e r t o s y s t e m a t i c a l l y s tu d y th e mode o f i n h e r i t a n c e o f
k e r n e l sm o o th n ess th e e n v iro m e n ta l c o n d itio n s m ust be a s n e a r ly i d e n t i c a l
w ith in th e n u r s e r y and from y e a r t o y e a r a s p o s s ib le and s in c e th e p a r e n ts
i n d i c a t e d a r a t h e r d e f i n i t e b re e d in g d i f f e r e n c e and th e p ro g en y s e g r e g a te d
i n t o a co m p lete s e r i e s , r a n g in g from th e ro u g h e s t c l a s s o f C e re s-(H o p e F lo r e n c e ) t o th e sm o o th e st c l a s s o f Renown, k e r n e l sm o o th n ess m ust f o llo w
some d e f i n i t e t r e n d o f i n h e r i t a n c e .
I n o r d e r t o s y s t e m a t i c a l l y s tu d y th e , i n h e r i t a n c e o f k e r n e l
-4 0 -
le n g th j i t was n e c e s s a r y t o d e v is e some m ethod w hich c o u ld he u s e d t o
o b ta in a s c o n s ta n t and a s r e p r e s e n t a t i v e m easurem ents o f k e r n e l s a s
p o s s ib le .
S e v e r a l d i f f e r e n t m ethods o f s e l e c t i n g k e r n e l s from th e
p l a n t s and d i f f e r e n e t num bers o f k e r n e l s w ere m easu red t o d e te rm in e w hich
m ethod w ould g iv e th e m ost c o n s ta n t r e s u l t s .
A s t a t i s t i c a l and g r a p h ic a l
co m p ariso n o f th e d i f f e r e n t m ethods t r i e d i n d i c a t e d t h a t th e t o t a l le n g th
o f 15 k e r n e l s ta k e n from th e lo w e r f l o r e t s o f s p i k e l e t s s e l e c t e d n e a r th e
c e n t e r o f t h r e e m ain s p ik e s o f e a c h p l a n t gave m easurem ents s u f f i c i e n t l y
r e l i a b l e f o r th e p u rp o s e .
The m easurem ent o f 20 k e r n e l s showed a r e ­
d u c tio n i n e e r r o r b u t t h i s r e d u c tio n was so s n a il i t d i n o t seem f e a s i b l e
t o m easure more th a n 15 k e r n e l s .
Out o f 560 Fg p l a n t s m easu red and p la c e d i n c l a s s e s a s d e te rm in e d
by th e p ro b a b le e r r o r o f th e p a r e n t s , 94 f e l l i n th e C eres-(H ope-F lorence§>
l i k e and lo n g e r c l a s s , 286 i n th e in te r m e d ia te c l a s s , and 180 i n th e Re­
nown l i k e and s h o r t e r c l a s s .
T h is v e ry r e g u l a r s e g r e g a tio n seems t o su g ­
g e s t t h a t some d e f i n i t e f a c t o r s a re p ro b a b ly o p e r a tin g i n th e d e te r m in a tio n
o f k e rn e l le n g th .
A ls o , a weak dom inance o f th e s h o r t e r ty p e s seems t o be
in d ic a te d .
Tw enty p la n ts , from e ac h o f 57 Fg I i n e s o f one fa m ily grown a t
Bozeman w ere s tu d ie d f o r , k e r n e l le n g th i n th e F3 .
The s ta n d a r d d e v i a t i o n s
o f th e p a r e n ts grown i n 1940 were ab o u t tw ic e a s la r g e and t h e i r means
a b o u t t h r e e m i l l i m e t e r s l e s s th a n f o r th e p a r e n ts grown i n 1939«
A ls o ,
t h e r e was a s h i f t o f th e dom inance i n th e p ro g en y t o th e lo n g e r ty p e
k e rn e ls .
T h is g r e a t e r v a r i a b i l i t y fo u n d i n th e F3 made i t im p o s s ib le t o
v e r i f y th e Fg c l a s s i f i c a t i o n on th e b a s i s o f th e F3 b r e e d in g b e h a v io r .
-4 1 -
T hese d a t a do n o t seem, t o a p p ro a c h th e e x p e c te d from th e Fg c l a s s i ­
f i c a t i o n , "because i f th e Fg- c l a s s i f i c a t i o n i s c o r r e c t and th e F3 was
n o t in f lu e n c e d by e n v iro n m e n t, th en , more p a r e n t a l ty p e s s h o u ld a p p e a r
i n th e F3 th a n i n th e F2 and a dom inoance o f th e s h o r t e r k e r n e l ty p e s
s h o u ld a l s o have b een e v id e n t i n th e F3 .
T h e r e f o r e , s in c e th e s ta n d a r d
d e v i a t i o n o f th e p a r e n ts grown i n 1940 u n d e r s e m i - s u b i r r i g a t ed c o n d itio n s
i s a b o u t tw ic e a s la r g e a s f o r th e p a r e n ts grown i n 1939 u n d e r i r r i g a t i o n ,
envirom m ent p ro b a b ly in f lu e n c e d th e e x p r e s s io n o f k e r n e l le n g th i n th e F^
more t h a n .t h e in n a te f a c t o r s and th e Fg c l a s s i f i c a t i o n p ro b a b ly more n e a r ­
l y a p p ro a c h e s th e t r u e mode o f i n h e r i t a n c e o f t h i s c r o s s .
Bunt i n f e c t i o n s were so s l i g h t a t b o th Bozeman and M occasin
i t was d e c id e d from o b s e r v a tio n a l d a ta t h a t a d i f f e r e n t i a l r e a c t i o n o f th e
p ro g e n y t o th e tw o •p h y s io lo g ic r a c e s u s e d c o u ld n o t be d e m o n s tra te d .
O nly
a t r a c e o f smut was fo u n d i n th e s u s c e p t i b l e check and a l s o o n ly s l i g h t
i n f e c t i o n s w ere o b ta in e d a t e i t h e r s t a t i o n i n any o f th e s p r in g w heat b u n t
n u rs e rie s .
Hot d ry w e a th e r and h ig h s o i l te m p e r a tu r e s f o r s e v e r a l d ay s
a f t e r s e e d in g th e bunt n u r s e r y p ro b a b ly e x p la in s why th e i n f e c t i o n s w ere
s l i g h t a t Bozeman.
H ow ever, c o o l m o ist c o n d itio n s c o n tin u e d a t M o ccasin
f o r s e v e r a l d ay s a f t e r s e e d in g b u t h e re a l s o only s l i g h t i n f e c t i o n s were
o b ta in e d .
The w r i t e r h a s no e x p la n a tio n why h ig h e r i n f e c t i o n s were n o t
o b ta in e d a t M oCcasin
-4 2 -
G
SUMffiAET
Fg
F3 g e n e t i c s t u d i e s w ere made on th e i n h e r i t a n c e o f
aw n ed n ess, k e r n e l sm o o th n e ss, k e r n e l l e n g t h , and r e a c t i o n t o two p h y s io ­
l o g i c r a c e s o f h u n t i n a C e re s -(H o p e -F lo re n c e ) x Eenown s p r in g w heat c r o s s .
F o r a m e d n e s s ,th e h F 2 p ro g en y s e g r e g a te d i n t o th e fo llo w in g c l a s s e s :
( l ) w eakly a w n le tte d , ( 2 ) s t r o n g l y a w n le tte d , and ( 3 ) f u l l y awned.
A c lo s e
f i t was o b ta in e d t o a m o nohybrid , 1 : 2 : 1 , r a t i o w ith in c o m p le te dom inance
o f th e a w n le tte d c o n d itio n .
F o r th e k e r n e l sm o o th n ess s t u d i e s , th e p a r e n ts and p ro g en y
w ere c l a s s i f i e d i n t o e i g h t c l a s s e s d e p e n d in g upon th e d e g re e o f ro u n d ­
n e s s o r a n g u l a r i t y o f c h e e k s , d e p th and w id th o f c r e a s e , and p i t t e d n e s s
o f th e k e r n e l s .
A u n ifo rm b re e d in g d i f f e r e n c e betw een th e p a r e n ts was
o b ta in e d and th e p ro g en y s e g r e g a te d f o r th e whole ra n g e o f and betw een
th e p a r e n t s .
So much v a r i a h l i t y o c c u re d i n th e F^ b e c a u se o f n o n -
u n ifo rm e n v iro n m e n ta l c o n d itio n s t h a t i t was im p o s s ib le t o v e r i f y th e
Fg c l a s s i f i c a t i o n .
K e rn e l le n g th was s tu d ie d by m e a su rin g th e t o t a l le n g th i n
m i l l i m e t e r s o f I 5 s e l e c t e d k e r n e ls from e a c h p l a n t .
I n th e Fg a f a i r l y
u n ifo rm s e g r e g a tio n was o b ta in e d w ith a weak dom inance o f th e s h o r t e r
k e r n e l s b e in g s u g g e s te d .
N o n-u n ifo rm e n v iro n m e n ta l c o n d itio n s c au sed
so much v a r i a t i o n w ith in th e p a r e n ts and F3 m a t e r i a l t h a t i t was im p o s s ib le
t o check th e Fg c l a s s i f i c a t i o n by th e F3 b re e d in g b e h a v io r .
Bunt i n f e c t i o n s were so s l i g h t i n th e p a r e n t s ^ p ro g e n y , and
s u s c e p t i b l e check t h t i t was d e c id e d from o b s e r v a tio n a l d a t a t h a t a d i f ­
f e r e n t i a l r e a c t i o n o f th e p ro g en y c o u ld n o t be d e m o n s tra te d .
A p p a re n tly
—4 3 -
e n v iro n m e n t a l co n ditions w ere su ch t h a t o n ly t r a c e s o f i n f e c t i o n were
o b ta in e d
-4 4 -
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ir f .'-.I"),
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'1
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MONTANA STATE UNIVERSITY LIBRARIES
CO
III I 111I l III Il
7132 100 1551 6 5
-
N378
Sm63i
1*4=1., c. 2
117292
Sm ith, O.W.
In h e r ita n c e o f aw nedness. . .
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