Inheritance of awnedness, kernel color, glume pubescence, and reaction to two physiologic races of
Tilletia in spring wheat
by Samuel C Litzenberger
A THESIS Submitted to the Graduate Committee in partial fulfillment of the requirements for the
Degree of Master of Science in Agronomy
Montana State University
© Copyright by Samuel C Litzenberger (1939)
Abstract:
This investigation Is primarily an Inheritance study of awnedneee, kernel color, and glume pubescence'
In spring wheat. In a cross of Reliance- Reward x Comet-(Hussar-Hard federation) the inheritence of
awnednees, kernel color, and glume pubescence was studied on 100 F3 hybrid progeny as well as their
reaction to two races, T-8 and L-7, of tillletia. Only awnedness one lnveetlgated In 100 F3 lines of a
second cross, Comet-(Hussar-Hard federation) x Ceres-(Hope-florence).
Two major factors were found to govern the awn development between the awnless and awned
varieties, since these were the two types represented In each cross, intermediate dominance prevailed
with the factors for the awnless condition being move potent than the awned, four true-breeding types
were obtained In the f3,. namely two intermediates, weekly awn-letted and strongly a waist ted, and the
other two like the parents, awnless and awned, A few epicalIy owned and half owned true-breeding
linen ware observed la the F4 of the first cross, a 1:2:2:4:1:1:2:2:1 gene-typic ratio whs observed In
both crosses, however, different phenotypic ratios were obtained, A 1:8:1:3:2:1 ratio of awnless,
apically awnletted, weakly awnletted, strongly awnletted, half awned, and awned, respectively, was
obtained in the Comet-(Hussarr-Hard federation) x Ceres-(Hope-Florence) cross sad a 1:4:5:3:2:1
phenotypic ratio mao found In the other cross, A single factor difference wee obtained for kernel color
and glume pubescence In the cross Reliance-Reward (dnrte-red seeded with pubescent gimmes) x
Cos*t-(Hussar-Hard federation) (Hatred seeded with glabrous glumes). The glabrous condition wan
found to be recessive to the pubescent.
The seed ef the F3 hybrids and parents In the first cross studied were Inoculated with n oosposlte of
races M and L-7 of Tilletia. The parents differed In their reaction to the Individual races, each being
resistant to one race and susceptible to the other. Nine lines mere obtained which mere move resistant
than the resistant parent, demonstrating it Is possible to combine the resistance ef each parent Into a
single variety ef wheat.
All characters studied were found independently assorted except seed color and per cent bunt Infection.
A weak linkage apparently exists between dark-red-seeded plants and susceptibility to bunt and
light-red-seeded plants and resistance to bunt. IlHKRITAJtCl Cf A MHOtms, KERMEL UCJLDFe GWfm Ptm^SCKSCXe AlD
RKACTlCR «J TTO PHTSICLCOIC RAClS CT TI UJETIA IR SPRING WHEAT
ty
LKymel Ce LItseenbeT^er
A TKISIS
Stttsdtted to the Orndtmte Committee
in p a r t i a l fu lfillm e n t o f the requirem ents
f o r the Degree o f Master o f Science
In Agronomy a t
Montana S ta te College
In Charge o f Major Hork
/ Chniimmm Omdnate Committee
Bosemn, Ibmtaaa
June, 1939
)V37£
L7 3^
coi»- ^
ACKNC « LKIXrKMSSTS
The w r ite r w ishes to r-rr-tof u l l y rvckTiowled/re h ie In­
debtedness to r>r.
He %mberp tm lo r T?hoae d ir e c t io n th e In-
v e e t l ^ t l o n e wore nede, to Or. $. J .
e l Itxmisen f o r h ie help­
f u l su p e r v isio n and e u fy e s tlo a e Anrlng to e f i r s t y e a r o f stu d y,
to Or. A. m. Schlehuber f o r h e lp fu l c r i t i c i s m nnd Bxir tia tio n e
during th e I n s t year o f stu d y , to P r o fe sso r
J1.
o a t f o r sug­
g e s t io n s and c r i t i c i s m in p rep aration o f th e m n n e c r lp t, to
Nr. A. a. Clarh fo r p rovid in g the n o to r ia l and g iv in g perm ission
fo r I t e u se In t h is stu d y , and a le e to to e M v ie lo n o f C ereal
Cro ie end M eo- s e e , U. s . Do ir tm n t o f A gricu lture fo r fln a n o ia l
? >
CO
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a e e lsta n c e throu^i th e Research fe llo w s h ip in Agronmy.
68498
-IIITABto cz c a e t m n
Zmm
UST CF t m - S .................................................................................................
LIST CF TABUS........................
IY
f
ABSTRACT ...........................................................
W T I C B .....................................................................................................
ITFJUTDH* BETIW ...............................................
YII
I
3
A -nodnee® ...................
3
Kernel Color
7
............................................................ . . . • •
Oltune Pubescenoe . ...............................
10
B m t Her c tlo n
10
...........................................
MA??JilAJLS AMD M TWDS.....................................................................................
%b
OO JUKWTAl RBSQiaS
If
. ................................................................................
SegpeeeUen o f C^tsumeters
Asnedaees . . . . .
................................................................
..................... . . . . . . . . . . . .
17
18
Cornel C o l o r ...............................................
Ug
Olnme Pubescence
50
.............................* ....................... ....
Bunt B m o t l m ....................
51
L lnkire S t u d i e s ................................
56
PfSCUS I ' ABD CCBCU . IC M S ........................................................................
$g
SONfUKT.................................................................................................................
67
LIfLRATlfPK CITKD ............................................................................................
Tl
-IV LIST CF FIGUmS
Pigttre I
Figure 2
Pedigree diagram o f the 100 P , lin e s o f R ellaneeSeward x Comet- (Hassa^Hard Fedem ti« a )
2%
Awn tgr aa found In progeny o f Reliance-Reward x
uonet- (Ifuasar-H' rd Federation) and Comefc-( Huesaritnrd Federation) x Ceree-( Hope-Flertmoe) t A and B,
v a ria tio n s in awnednesB from each o f two p la n ts c la s s ­
ed a s > awn tjr as from toe too c ro sse s, re sp e c tiv e ly ;
C and D, re p re se n ta tiv e spikes from toe two c ro sse s,
re s p e c tiv e ly , I , awnless; 2, n p ie illy sw nletted; > ,
weakly aw nletted; > , stro n g ly aw nletted; H, h a lf awaed, end 5» owned
29
Figure 3 F1 atm types in Reliance-Reward x Cow et-(Httss n r<v rd Fedem tion)* A, p a re n t I awn type x p aren t
5 *** type; B, progeny I awn type x progeny 5 msn
^rpe; C, progeny > awn type x progeny > atm tFpe;
P, progeny 3- x p e rM t 5 awn type; and X, progeny
> x p a ra a t 5 Q*n type
Figure 4
Frequency d is tr ib u tio n o f the average percentage
o f bunted p la n ts when inoculated w ith a composite
o f r ceo o f T-<? and L- 7 o f T i l l e t i a in paren ts
and progeny o f Reliance-Row rd x Comet-OhissarHerd Federation)
39
5*»
Uif
CF TAJlLtS
Pa#*
Table I
Table I I
Table I I I
Table IF
Table F
Table Fl
Table FII
The c la s s if ic a tio n o f p la n ts f o r awnednea:., p e r
cent b o a t, k ern el c o lo r, and glome pubescence in
F , lin e s and m re n t rows in the cro ss PellaneeSeeexft (*•*• 1366) x C oast-(Rsmearaffarft Federation)
( . . 1315) grown e t Josem n, Hoatana In I; >37
19
The c la e e l f l c a tio n o f p la n ts f o r mmednees In Fi
lin e s and p aren t rows l a the c ra ss Cons t - ( Hnesara
Jfard Federation) x C eres-( Fiope-Florenoe) grown a t
Boseaan, Montana in 1938
25
Honbor o f Fj p la n ts in each awnedness c la s s In the
nine F , breeding grasps o f Reliance-Reward x C om t(HttsearaRnrd Federation)
28
Breeding behavior In the F-. generation f o r the s ix
aimedness c la s s e s in the c ro ss Rellaitoe-Rerard x
Conet-OfasRaraHard Federation) grown a t Bosemaa9
Montana In 1937
30
C alculation o f Coodness o f F i t to a I* 2*2*4*I*I*2; 2*1
r a tio ( a) and a I*4*9*3*2:1 r a t i o ( ) on a tw o-factor
d iffe re n c e fo r awnedneae from a R dlaace-R erard x
Coas t - ( Rus aaraffard Federation) sp rin g wheat cross
grown a t Boseaan, Moatone In 1937
34
Munber o f F , p la n ts In each otmednees c la s s in the
nine F , breeding g re a ts o f the I(X) lin e s o f C om t(HuesaraHard Federation) x C eree-(Hoi>e-Florence)
4l
Breeding Xmkmrlov In the F j generation fo r the s ix
awtodness c la ss e s In the c ro ss Comet-( Hhmsaraffard
Federation) x C eres-(Hoito-Florence) grown a t Boaewm,
Montana In 1938
42
Table F i l l l-alo alatto n o f Ooedneee o f F it to a I t 2»2*4* I t I* 2»2*I
r a t i o (A) and a 1*8:1*3*2*1 r a tio (B) on # fio -fo e to r
d iffe re n c e fo r awnedneae from a tionet-(Sassnraffard
Federation) x Ceree-(H opraFlerm ee) spring wheat e rase
#rtnm a t Bosnma, Montana la 1936
47
Table IX
C alcalatlo n o f Ctoodnees o f f i t on a o n e-facto r
d iffe re n c e f o r k ernel c o lo r l a the I*, generation
o f f>ellan o e-Kerrnrrl * Gorot-Ohum- r-FTrd Federation)
grown a t Bosemn, Montana In 1937
Table X
G aleulatlon o f Geodnssa o f f i t fo r In heritance o f
Klome pubescence on a s in g le -fa c to r d ifferen c e In
100 Fi lin e s o f Hellaaee-Reward x Conet-(HueenrHard Foderatlon) grown a t Roxemn1 Moatona In 1937
Table XI
Rerceatogee o f bunted who t p la n ts In ro w o f
Reliance x Renrard and Ctonet x Hnesar-Hard Federation
p a re n ts and Fi hybrid stra in * o f the ernes grown a t
Rosoman. Montana l a 1937
Table XlI
A e o n ro rlson o f emit re a c tio n to composite o f T-S
and L-7 smut r%cee w ith seed co lo r In 100 Fj lln o e
o f RoUawcn Wemnri x Comet-CHusear-Hard Federation)
-VII
In h eritan ce o f Aroedneee, Kernel Color, Glwme Pubescence, and
Keactltm to two Physiologic Rneee o f T lllo tle in S pring Wheat
ABStRACT
Thie in v e s tig a tio n Ie p rim a rily an in h erita n c e study o f aroedneee,
k ern el c o lo r, and glume nb^ coace in et>ring whe t . In a c ro ss o f RelianceReenrd z Cotaet - (ilu s s-r-’Lard Kedor t l on) the Inh eritan ce o f eiraednees, ker­
n e l c o lo r, and glume pubescence ro e stu d ied on 100 F , hybrid progeny as
w ell as t h e i r r u c tio n to two ra c e s, T-R and L-7, Ofj T ll l o t i n . Cnly aroedness Be Inveotlgatod in 100 F , lin e s o f e mecond c ro ss. Comet-( its r dnrd Federation) % U erea-(K m ocH orm ce),
Two major fa c to rs were found to govern the awn development between
the aro leee and aimed v a r i e ti e s , sin c e these wore the two types represent­
ed in each c ro ss, interm ediate dominance p rev a ile d w ith the fa c to rs fo r
t i e -umleoB condition being o re o te n t than the rained, to u r tm o -b rre d Ing types were obtained in the F ,, namely* tvo In term ed iates, weakly awnle t t e d and stro n g ly a w aist ted , and the o th e r two lik e the p a re n ts, awnle s s and owned. A few a p ic a lIy owned and h a lf owned true-b reed in g lin e s
w re observed In th e F4 o f the f i r s t c ro ss. A I t 3 t2 t4*I*I* 2*Pt I geno­
ty p ic r a tio was observed In both c ro sse s, however, d if f e r e n t phonot y Io
r a tio s were o b t lned. a I t Rt1*3*2*I r tlo o f s ro le s s , a p le a lly aw nletted,
weakly a ro le tte d , stro n g ly aw nlettod, h a lf armed, and owned, re sp e c tiv e ly ,
was obtained in the Comst-(lfassaxfc»Bard Federation) % Cereo-(Keim-Florenee)
c ro ss and a I i *5*3*2*1 ;Aenotyplc r tlo was found In the o th e r cross,
A sin g le fa c to r d iffe re n c e was obtained fo r k ernel c o lo r and glume
tubeocenee In the c ro ss Reliance-Reward (dark-red seeded w ith pubescent
glumes) x Comet-(Bsssar-Kard fed eratio n ) (Ilfdbt-red seeded w ith glabrous
glum es). The glabrous condition was found to be recessiv e to the pubes­
c e n t.
The seed o f the F j hybrids and p a re n ts in the f i r s t cro ss studied
were inoculated w ith a cow ooito o f races T-K and L-7 o f Tl I I e t l a. The
p a ren ts d iffe re d in t h e i r re c tio n to the individual ra c e s, each being
r e s i s t a n t to one race sad enecontlble to the o th e r. Mine lin e s were ob­
tain ed which were more r e s i s t a n t than the r e s is ta n t p a re n t, dem onstrating
i t i s p o ssib le to combine the re s ista n c e o f each p a rro t In to a sin g le
v a rie ty o f wheat.
A ll ch arac ters stu d ied were found independently a sse rte d except
seed c o lo r and p e r cent bunt in fe c tio n . A weak linkage apparently e x is ts
between dark-red-seeded p la n ts and s u s c e p tib ility to bunt and lig h t-re d seeded p la n ts and re s is ta n c e to bunt.
In h ari t nce o f Awnednces, I ru e l Color, Olume Pttbeeconee, and
e ao tian to Tim Phyeiolo^ie Rooee o f T ll l e t i a in Spring wheat
Seroel G. L itzen b erro r
tw m m m tm
th e ro le played 6gr g enet!ee in e s trb lla h ln g toe underIy lap
p rin c ip le # o f la h e ritro e e eannot be over * t phmei#ed.
Bwrot inform #tiro
on the node o f In h eritan ce o f variou s character# ha# aided in ro ttin g
up mere e f f i c ie n t breeding progmraa,
Hrsqr e f tow charnotor# o f the
whmt p la n t have hero stu d ied and much has been accomplished, but l i t t l e
i« known about the mode o f in h e ritones o f many character# which should
be cnnbined in to the p e rfe c t v a rie ty ae conceived tgr t)» p la n t breeder.
Sony problems remain to be solved: genetic# can rod w ill play an lm e rto n t
ro le in th e ir so lu tio n .
Oregor
andal furnished ua with the plan fo r in h e rita n c e , deal­
in g w ith u n it c h a rac ter# , dominance, segregation rod recom bination.
It
wee not u n t il 1900 when Be f r i e s , Gorrena, and von Techesmk independent­
ly and s in u lt neously rediscovered Uendel*s c la s s ic findings jm bllshed
in 1-66 th a t wide i n te r e s t developed In the science o f g e n e tic s.
While
u n it c h aracter# have not proved to Ue as simple l a whe»t as was f i r s t
thought, and dominance i s incomplete In most case s, the sin g le c h aracter
I s s t i l l the foundation fo r studying wheat ln h e rl tnnce, rod doniaroce
tow parim ry b a sis f o r g en etic i n t e r p r e t felons.
The presence and imr'ort nce o f d isease re e l stance in wheat i s well
-3 knovn.
In rec en t ye^ore marked progress has been m da in breeding wheat
f o r re s is ta n c e to d ise a se .
lo g ic ra c e s.
(l.ierte.)
th ie typo.
Many Amgus d ise a se s contain nuraercms phyeio-
JSmt o r stin k in g erstt o f wheat, censed by T i l l e t i a t r l t l c l
l n t . and Tt Levis (Kuehn), Is one o f the most d e stru c tiv e o f
A v a rie ty may be r e s is ta n t to some physiologic races but may
Os v ery su sc e p tib le to o th e rs .
The g e n etic foundation f o r re s ista n c e to
bunt has been proved beyond a doubt, and i t s mode o f l n h a r i t nce has been
shown in some oases.
Blnce these conditions p re v a il, i t i s the ho e o f
the p la n t breeder through recom binations to combine In a sin g le v r l e t y
the near-immune re a c tio n to a l l r ees o f boat as w ell as to o th e r d ise ases.
The importance Cf g en etic s in a crop Improvement program has teen
c le a r ly defined by Salmon and Laude (19^?) who s ta te th a t "the random
c ro ssin g o f v a r ie tie s to induce v a ria tio n o r In the hope o f securing
favorable chance combinations i s too u n cart* in a method an which to base
a crop-Improvement r o g r -»*.
Time, in o rd e r to proceed most e f f ic ie n tly
in s breeding program, i t Ie Iwmrfemt to know the mode o f Inheritance
fo r most o f the im portant Char- C ters o f the p a re n ta l m a te ria l.
The In v e stig a tio n s reported in th is pa # r c o n sist o f Inheritance
stu d ie s in two sp rin g w h a t c ro sse s.
In one c ro s s, ) aIiance-Reward *
Cotnet-(Hua«ar-H"rd F ed eratio n ), stu d ie s were made «m awnedness, kernel
c o lo r, glume m beseence, and re s ista n c e to bunt; in the o th e r c ro ss.
Comet- ( Huswar-R'-rd Federation) * Ceres-(Hope-Flerfince), only the In­
h e rita n c e o f srmedneaa wee stu d ied .
LITSRATUKE R::vim
ATOedaess
Whest v a r ie tie s g en erally have been separated Into two major
groups on the b a sis o f awnednesa c h a ra c te r, namely*
aimed and nTOless.
I t I s now recognised th a t a l l wheats cannot be c lR sa lfie d Into these two
simple groups end th a t v rlo u s Interm ediate ty re s e 1 st.
The fa ilu r e to
recognize these Interm ediate types probably accounts fo r much o f the d ie agreement b e t ^ e n the r e s u lts o f the e a rly workers who sta«lled the Iah e rl t nce o f th is c h a ra c te r.
The f i r s t g enetic study on asmedness In w#ra% hybrids was re p o rt­
ed in 1905 ty B lffen, who concluded th a t the beardless condition la a
(loadnant, the bearded, a r e c e v lv e c h a ra c te r.
He made m ny cro sses be­
tween bearded and so -c a lle d beardless wheats.
The F« w s always "b*-rd-
Iese* , ^nd the Tg segregated Into 3*1 o f "Swnlesew to a-mad.
very c lo se f i t s to a *t I monohybrid r a t i o .
t e r m in i awns
c la s s ,
He obtained
P la n ts occurred w ith sh o rt
to h a lf eta Inch in length; these were put in to the sunless
'dome o f these
e re
Iso found In the
re n ts .
I thou t exception
the owned types bred tru e , while the "beardless* types e ith e r bred true o r
segregated in to bearded and "beardless* t y ee.
Cther e a rly workers obtained sim ila r r e s u lts in the f i r s t generation
and in the second generation a ls o , Wiem the "ownless" and awned p lan ts 00curred la a sim ple Hendellan r a tio o f 3*1»
Saw ders ( 1907) f i r s t queatloned
the id ea th a t the f i r s t g eneration between an awnless and an owned wheat a l­
ways i s su n less nnd aointolned th a t the awns la the
v a rie s w ith the wheats
used.
P erctv n l (1921) I n t e r reported th a t Fg eepreraticm l a nunerews
cro sses ap in c h e d a 1: 2*1 r a tio when interm edia tea occurred, anfl when
the sent-be rded forms o f T rttlc n n v u l ^ r e wore crossed w ith ty p ic a lly
bearded f o r m , the F1 p la n ts have heads e a s ily d istin g u ish ed from those o f
the b eard less p a re n t.
Bie Fg generation from these *eefid-beardod* F1 hybrids
co n sisted o f ^beardleas*, sesd-bearded and bearded p la n ts In the r a tio o f
1*2*1.
Many workers here v e rifie d thee* r e s u lts .
Hrsyee and Aauaedt (1923)
found s o n e -fa c to r d iffe re n c e In n c ro ss between awned and seml-iwned
v a r ie tie s .
The
w s stro n g ly aw nletted and segregated a s a nwaoSybrid.
Vltii a re c ip ro c a l cross Clark and Quleenberry (1929) v e rifie d these re ­
s u lts .
Gaines and S ingleton ( 1926) , S tseaF t and Hoodwnrd (1931), S tsv n rt
BH* Dnllsy ( 1932) , ^ulsenberry and C lark (1933). Ausema (193^). m d
Tlngey and Tolaan (193*0 e l l reported s in g le - f e to r d ifferen c e s when armed
and eeml-rvmed paren ts were crossed.
In SKNit eases monohybrid r a t i o s have been obtained when th ese Mmimmed whm t s h ve been crossed w ith the tru e awslese tan>es.
orkors who
found s in g le -fa c to r d iffe re n c e s when awnless wheats were crossed with awnle tte d t T 1** are*
Stew art and Tingey (1928). Stew art (1932), fpieenbsnry
and Clark (1933) and Ausesns (193*0.
S everal r-orkers have reported minor fa c to rs p laying a p a r t In the
tgr 0 o f aim produced.
Clark m d Hooker (1926) In making a Rnrd fed e ratio n
* Mmrouie cross found an wmeunl imeber o f p la n ts w ith extrem ely sh o rt beaks
which v e rs not e x actly lik e the awnlees p a re n t, and v e rs n o t lik e the Mixt
h ig h er c la s s .
Then these were put In the nwnless e l s e they obtained a
5*1 r a t i o , but when m t w ith the a w le tte d c la s s a 15 to I r a t i o o f aw nletted
5te
CORtf
le t e l y Ivm lm n were obtained.
were the sans.
The r e s u lts o f fche re c ip ro c a l cross
They concluded th a t two fa c to rs (I mejer and I minor)
governed the awn development in th is c ro ss.
Two minor fa c to rs were needed to explain the r e s u lts obtained toy
C lark, T T orell, and Hooker (1928) when Bobs rnd Hard fe d e ra tio n were cross­
ed.
Both p aren ts are c la ssed *# ownless who tn .
The Bobs v a rie ty possesses
no beaks whatsoever, whereas JHhrd Fedor tlo n Has sh o rt beaks o f I to sevem l
na. In lengtii a t the t ip s .
An approximate 13*3 r a tio in the F^ was o btain­
ed.
P v a re n tly the awnless w h e t s re fe rre d to In most o f the e a rly
g en etic s tu d ie s were weakly o r stro n g ly aw nletted.
Howard nd Howard
(1912, 1915) in In d ia were the f i r s t to re p o rt r e s u lts o f c ro sses between
tru e awnless wheats and owned v a r ie tie s and they reported a tw o-factor
d iffe re n c e .
They grouped the mmed and tip-swned c la sse s to g eth er a#
mated and commred them to the awnless types which gave a 15*1 r tlo .
They thus concluded th a t the double m ceasiv es were
m le ss.
The F^ was
n e a rly awnless.
Various workers have obtained a two-major fa c to r d iffe re n c e fo r
wnednees wi«m awnless and mwued type# vmre crossed.
In a Kata r H-rd
F ederation cro ss C lark (192^) found a two-major fa c to r d iffe re n c e , but
sin c e no c o ^ T e te ly awnlfs * o r aimed types were obtained which bred true
he assumed th a t minor modifying fa c to rs were p resen t.
C lark, F lo r e ll, and
Hooker (1926) reported a tw o-factor d iffe re n c e la one c ro ss o f 0 bearded
w ith a b eard less wheat but e th re e -fa c to r <2 major and I minor) d ifferen c e
in another.
Vba only e f f e c t o f the minor fee to r in the presence o f one o f
th* «*Uor fa c to r s Ie to reduce the saount o f nenedneee a s ftom to e a tr o n ^
e r to the weaker I lm lte o f a c la s s .
From t h is stu d y they have concluded
th a t a s miyr a s fou r fa c to r s may be Involved In awsedaess lito e r lte n c e l a
eome wheat c r o e s e s .
Cther workers who found tw o -fa cto r d if fe r e n c e s w)iea
a im less wheats were cro ssed w ith owned typos ore*
(1 9 2 9 ), Etew ert and Judd (I Q J l)e C lark,
laan b erry, and Powers (1 9 3 3 ),
* u e e m s (1 9 3 4 ), and Tlngay and T olm a (1 9 3 4 ).
e m le t ted and the
Stewart and Reyward
AU found to e F1 s p i c i l y
- roz-enlee were c l a s s i f i e d Into n in e g en o ty p ic gretr s .
Kvldentiy th ere a re two g e n e tic a lly d if f e r e n t ty re s o f sesd-aroI m s wheats, since Love and C m lg (1926) found when Senom (semi—^wnleas)
was crossed w ith c e r ta in o th e r aesd-awnlees ty p es, the F1 was n early nwnIeee while the Fg end I n t e r gm iem tlons -wned and p a rtly awned ^rpes Ojipenm
ed.
The r e s u lts showed a 15*1 r a tio whl<to in d icated th is v a rie ty although
seml-awnlees c a rrie d a fa c to r fo r the aimed co n d lti n.
Sten r t (1932)
crossed two tru e breeding aw nletted types whlto he obtained os segregates
from e S ev ier * Federettcm cross and obtained a l l types s p a te .
breeding type* were recovered*
In term ed iates.
Kour true
one com pletely awnlee , one awned, and two
M lsenberry and C lark (1933) Obtained aI r d I a r r e s u lts from
n cro ss between two aw nletted vhe t* , Konom x M allty.
A co%)lete range
o f segregation from aimed to awnlees was obtained In to e Fg w ith four types
breeding tru e .
I t w e assumed th a t Samore contains the g e n e tic fa c to rs
seSB end Q uality contains fa c to rs AAbb, whereas the awnless segrem toa were
AiBB and the sm ed ones were abb.
?h iltlp le allelom orphs f o r half-owned nnd fullyavm ed plmnto a ris in g
by com,lex m t a t ion from the awnloas p la n ts has been used m a basis tor
la te r- r a tin g the awedaewa In whesta I y Mlleaoar- M e (1 9 2 0 ).
He reported
th a t he obtained by n a ta tio n tru e breeding fem e o f SWBlee*, h a lf-a m e d ,
aad
ened ch eat*.
mmed,
Awnloes fo rm ear# p o rtly dominant to half-owned mad to
nd K^lf--Iwned fe rn s to nimsd.
The b a sis f o r ln h o rit-nce o f sanedneas M e been found to be simple
In some cro sses mad very complex l a 0th e re .
A d iffe re n c e o f only a sin g le -
f a c to r p a i r eae found In some cro sses b a t me m ay a s four fa c to rs (2 major
sad 2 minor) have been found accessary to ex plain r e s u lts o b t Ined In com.
b ia rtio n o f crosses la r o lr la g Hard fe d e ra tio n , Kota, and Bob*.
Incomplete
dominance fo r awaleeeaeee h ie been demons t r i e d , the F1 more a a rly approachla g tiie awalees p a re n t than the owned type.
Aimless, awnod, and too ln te r -
medlnte ly res which may be recognised a re the four tru e breeding types m e t
o fte n found, although tru e breeding e p le o lly ow let,tod o r h a lf-a w e d ty re s
hare been obtained in some cro sses.
Kernel Color
Kernel c o lo r w e e n rly recognised am an im portant W inrncter In
rfcaet w r l e t l e s and consequently w e among th e f i r s t used fo r a study o f
Wendellan I n h e rit nce In wheat.
Most r n r l e t le e o f stie&t were observed to
h are e it h e r red o r white k e rn e ls, although some Alysslnlsua durum sh w te
and e m e rs are purple.
In the e a rly work Iy B lffea (1905), red was found dominant to s h ite
In the F1 and segregated in a simple 3*1 r a t i o in the Fg.
M llsaon-m is
(19U ) w s the f i r s t to re p o rt croeaes shlcdi In the F^ #mve 3*1, 15*1, and
• 3* I r a t i o s o f red-kem eled and whl te-kerrw led p la n ts , ln d le o tln g the presence
©f I , 2 and 3 gene t i e fac to r e .
Sewxzd and H e ^ rd (1912), Ckxlsea (1917),
Love and Craig (1918), Clark (19A ) . Etewart (Ig g ), Stew r t and Tlagfly
(1928), Stm m rt mad Soedw rd (1931)» Stew art and Bailey (1932), Clark,
onlsenberry, and Poeere (1933). Tlngay and Tolnsn (193%). and Schlehttber
(1935) have obtained el i d Ir r ra tio # depending ©n the v r i e t l e a need when
studying the progeny la red x white seeded wheats.
Thus, from F3 stu d ie s,
the presence o f I , 2, ©r 3 Independent g e n etic facto r# fo r kernel c o lo r
has been proved l a d if f e r e n t v a r ie tie s .
m eso a-D ile* # theory w-s th a t,
taken to g eth er, the th ree rad c o lo r fa c to r# beh w m in ulnlively, follow ing
the law o f a t r I hybrid, and lncreaee the in te n s ity o f the seed co lo r.
Some v a r ie tie s e f wheat have been reported to possess s tr a in s haw
ta g d if f e r e n t numbers ©f f r o to rs fo r g ra in c o lo r,
flbrrlagton ( I 1)62) re ­
p o rted llte h e n e r contains two independently ln h o rlted fa c to rs f o r red c o lo r,
and th a t some o f the s t r ln s e f Red Bobs and Rnrd Red C rla a tta contain two
independently In h erited fa c to rs fo r g r in c o lo r while o th ers co n tain only
e sin g le f c to r.
Clark
nd looker (I 26) found th a t in
•
nrpule x Rard
te d e r tlo n c ro sses two fa c to rs were most frequently p re se n t, b a t th a t in
sons cro sses only a s in g le -fa c to r d iffe re n c e wa» o b t lned.
th e f a c t th a t
two d if f e r e n t r a tio s were o b t Iaed in d ic a te s e ith e r th a t Ktrgui* does not
always haws two dominant fa c to rs fo r kernel c o lo r o r th a t tits white Hard
Foder tlo n parent# d iffe re d in t h e ir genotype.
The u su al r e s u lt o f crossing two w hite-grained wheats Is the pro­
duction o f w hite-grained d rs c e n d n ts In the
and a l l subsequent generation#,
however, P e rc iv a l (1921) has ret orted th a t V llro rln obtained some ro d -tra in ed
p la n ts In the F2
two white tdioats, C rltlw m r-olonlcnra -md JTgi tnrgidam.
-9 -
were orosned.
Perol v t l f a rth e r e ta te a th a t P lte tii recorded the appe rraice
o f a few yellow o r red dlah-^rslned p lant# In the aIx th generation from a
Squarehead x Challenge cream.
Both v n rle tle e a re white seeded.
Qalnea
(1917) rep o rted th a t In two eases when w hite-grained tn d lrld a a le were
crossed some rod-kom eled p la n ts occurred.
Se f a r ae known these Jresnlta
have never boon repeated ty o th e r workers, nor have sim ila r r e s u lts been
obtained by cro ssin g two w hlte-kerneled v r l e t l e s w ithin the anno »i»eciea.
The r e c n lts , i f a u th e n tic , would in d ic a te dominant gen etic fa c to rs fo r white
k e rn els and rocesslve complimentary fa c to rs fo r red.
P e rrtv a l ( 1921) suggested th a t when red-grained v a r ie tie s are
crossed the progeny Ie gener l l y red in a l l g en eratio n s.
However, he c ite s
Tsehenmk am having obtained In some lnsfcnces w hite-grained Individoals
^mong the descend ints o f lyrtaride between d if f e r e n t red-grained v a rie tie s .
P erciv al a lso re p o rts M llsaon-Shle as having recorded the occurrence o f
w hite-grained
la n ts In Tp o f the hybrid "More" wheat x " Kxtra Squarehead",
both red-grained p a re n ts.
Ikgree and Robertson (1924) noted the frequency o f the occurrence
o f the whl to -V erm led p la n ts In the ?g stu d ie s o f Minturki x M rqule and
Kanred x Marquis cro sses where the p a ren ts were both re*-! e m e le d .
From
the d ata o b t in ed . they concluded th a t the red c o lo r i n Marquis i s the
r e s u l t e f two independently in h e rite d f a c to rs , e ith e r one o f which leads
to the production o f seed c o lo r and th a t Minteurld and K n red have a sin g le
f a c to r f o r red d if f e r e n t from e ith e r o f these in M nrqde.
The crosses ap­
proximated th e r a tio o f 65 r ed-ke rn e led p la n ts to I w h lte-k sm eled p la n t.
Kanred Md Minteortri have th e follow ing g en etic c o n s titu tio n rrr* r* 8"R*
10*
w hile IPr-Pqnls te e *SR*1 * r* r" f o r I t s congtlfcutlon.
Olune I^tbegconc e
I1Uhesewit end ^Labrous glumes u su a lly c o n s titu te an allelom orphic
palp o f c h a ra c te rs.
Blffsei ( I 905) w e f i r s t to re p o rt f e lte d o r velvety
c h a ff dominant over glabrous c h a ff In * QInyJe 3*1 r a t i o .
hybrids stu d ied the progeny segregated In to a 3*1 r a tio .
dominance w e ahown In a few cmsee sin ce the
In e l l the
Incomplete
wee Interm ediate In imbee-
cence, but th e Fg segregation w e n o rm l as complete dominance o f 3*1
pubescent and glabrous glumes.
co n d itio n s and r e s u l ts .
P e r c lw l (1921) r leo reported these same
According to th is author Ruemkrr obtained both
glabrous and v elv et-ch affed p la n ts In th e W3 generation in the r a tio
3 velvet* I glabrous, In the hybrid between a ubeacent and beardless
f r ltlc u a v u lg ire nnd n pubescent and bearded T. port ^otu
( I t-iU l).
Perclvnl
rep o rted sim ila r r e s u lts from a hybrid between tee pubescent-chaffed wheats,
T. dicocciun * J 4 snhaerococcnm.
one o th e r than a 3 tibesoentt I glabrous
r a t i o te e been found In crosses between v a r ie tie s o f co wm whe t s .
Bunt BBectioa
Boat or stinking: smut o f w hert. T l l l e t l - t r l t l c l (FJerk)
ln t. or
T ll l e t l o lo v ls (Knehn), Hoe been end continues to be e d e stru c tiv e disease
o f steiat In the IInlted S t t tea and In o th e r co u n tries where wheat Is pro­
duced.
In these wheat producing a reas the pre ence o f Hetsgr physiologic
races o f bunt or Vw appearance o f new races increases tits d i f f ic u lt y for
the wheat brooder o f ob t in in g Soam erelel v a r ie tie s r e s i s t n t to bunt.
Reed (1924), Rodenhleer and StsJcaaa ( 1927) , Oainee (19285, Breesran (1931),
Holton Bad Heald (1936). HodenMaer and Holton (1937), %nd o th er s ivtre
stu d ie d and review ed th e
h y s lo lo g lc s p e c ia lis a t io n work w ith bunt.
The
eode o f lr iio r lta n c e o f r e e ls tsn ce to p h y sio lo g ic r oes o f bunt ha* bean
ex p lain ed on one, two, th r e e , and m u ltip le fa c to r d if fe r e n c e s .
S r iile im ^ r
(1938) has review ed the lit e r a t u r e on the nature e f seg r e g a tio n ra th er
e x h a u stiv e ly and hue grouped the authors accordin g to the number o f fa c to r s
in v olved In th e c r o s s w ith which they worked.
th e plan ner work In breeding wheat fo r r e s le tr n c e to bnnt was done
in A u e tr slla by Ih r re r
nd In th e U nited S k te e by Cnlnes o f Washington.
Irirrer in 1901 reported h ie f i r s t attem p ts to luring about bant r e s is ta n c e
in wheat by h y b rid lz t ie n .
Having observed th a t 10 A u stra lia n v a r ie t ie s
Cflirried in f e c t io n s varyin g from 12 to 9 5 .5 p er c e n t, he thought a s im ila r
degree o f v a r ia tio n might occur in th e hybrid gen era tio n o f h ie c r o s s e s .
H is p la n was to s u b je c t
th e b u n t-fre e p la n t s .
and
T1Opnlntiona to a hessvjr a t t ck and s e l e c t
Hs # As no attem p ts to observe the seg r e g a tio n Into
r e s is t a n t and stubc a j t I b le s tr r ln s in order to e s t a b lis h U arileltan r a t io s .
Ths v a r i e t i e s , F lorence and Rmnoa, which nre h ig h ly r e s i s t a n t , were pro­
duced.
C aines (1920) begsn h ie In veetig o tlr m s , in clu d in g the g e n e tic s o f
r e s is ta n c e in 1915, and In 1923 he d escrib ed h ie e x te n siv e r e s u l t s .
He
grouped v a r ie t ie s In to fou r cl- Sflea— s u e c o ^ tib le , in ter m e d ia te, r e s is t a n t ,
and lm sm e,
Wrvm r e s is t a n t v r i e t i e e were cro ssed w ith sm scen tib le o n es,
e u e c e p t ib lllt y woe dominant, w ith o n ly about 2 per cen t o f the progeny
s e r e s is t a n t a s th e r e s is t a n t p aren t.
When s u s c e p tib le v a r ie t ie s were
cro ssed w ith immune ones (M artin aril Hussar) th ere seemed to be e pre­
ponderance o f r e s is t a n t p la n ts .
More than h a lf o f the hybrids In the
-L -V
th ir d g en eration m duced Ieee than 5 p e r cent o f bunted bends, vittereas
about 20 p e r cent were b u n t-fre e .
A ltogether he studied more than 25
sep arate cro sses and explained the r e s u lts obtained in each on a netliIp le f a c to r b o ats,
"he Influence o f any sin g le fa c to r was not w ell defined,
Whwi a l l fa c to rs are p re s e n t, a s In immune v a r ie tie s , re s is ta n c e i s
apparently dominant, but a le s s o r number gives the recessiv e e f f e c t.
R rlgrs
( 1926) , however, found th a t the M r t i n v a rie ty had one dominant fa c to r fo r
n e ar Inwunity and th a t Hueear hod two.
Zn mn e x c e lle n t s e rie s o f papers,
Trlgge (1930. 1930». 1931. 193?. 1932a. 1933. 193%. and 1936) has shown the
In h eritan ce o f boat re a c tio n in d if f e r e n t cro sses to be q u ite s ta p le , due
to one o r two fa c to r d iffe re n c e s.
lo g ic races o f bunt,
He worked p rin c ip a lly w ith sin g le physio­
He (193*0 hos determ ined the genetic c o n s titu tio n o f
10 bunt r e s is ta n t v a r ie tie s o f wheat r-nd l i s t s them as follow s:
Y nristy
G snetlc C onstitu tio n
ite r tls
Fhhtt
White Odessa
MMWKtt
Banner Berkeley
MlfM itt
Odessa
MMMltt
Sherman
KfM ltt
Museaf
MMKHtt
!tesenff-Sft, Fed. * L i t t l e Club Bel. lHlS
XWfKtt
Hussauh-nd. Fad. x M t t l e Clnb S al, 1U03
PfffPHt
Turkey C .I. 1558
Tturfeay C .I. 3055
iwdihTT
Crc
SfMiTT
These v a r ie tie s contain the Martin (MtO, Htisesr (BH), o r Tmrtmy (TT)
fa c to rs fo r re s is ta n c e .
f o r r e s i s t ncG.
The f i r s t fiv e l i s t e d depend on the Martin f e to r
Besear coat in s W th the Mmrtirn rn& ftusssr fac to r* ,
Briggs
( 19J>b) concluded Tailcey C .I. 257^ CtBitalnod the Turkey f c to r fo r re s is ta n c e ,
Ttiressman (1933) foaad A lb it c en t ,Ino one m in fa c to r fo r resistan c e to c e r­
ta in i^ y s le le g ie races o f bunt and he suggested th a t th is fa c to r i s the sees
ae th a t c s r r ie d by i t s male p a re n t White Odessa.
The near-immune re a c tio n to some races o f bunt possessed by Hueear
faac been tra n s fe rre d to hard red sp rin g sh e e t in t Komr x Bnsssr cro ss
by Smith (1932).
According to Clark ( I 936) F lo r e ll mnd ^nylee have develop­
ed ,another n early lrmune hard red sp rin g sh e a t (S.So. IOlSt C .I. 11, ! 2)
from a c re ss o f Hneear, a hard red w in ter i * m i ?nd H rA Fedetw tion, a white
e?>rlag wheat.
Both o f these s tr a in s have b en crossed with su scep tib le
v a r i e ti e s , and the Fj p la n ts were found by Clark (1936) to be b u n t-fre e ,
roving dominance o f n e ar immunity.
According to Clark (1936) in the Fg
s tu d ie s made by C lark and Eodauhiser these c re sse s showed a s in g le -fa c to r
d iffe re n c e .
This i s sharply in c o n t r e t to th e in h eritan ce from such
v a r ie tie s as Hope, Knrquie, R ld it , f r o , Turkey and Florence crossed u on
stiseepU ble v a r i e ti e s in which the F1 p la n ts a re au sco p tlb le.
The Btitmg re s ista n c e o f Hope wheat to bunt was stu d ied by Clark,
n is e n b erry , and Powers (1933) l a cro sses involving the weaker ree l stance
o f Karquie end d if f e r e n t degrees e f s u s c e p tib ility in Cseres and BrrA
F ederation.
The re s is ta n c e e f these three cro sses was stu d ied follow ing
i SQCttlit i on w ith a c o lle c tio n o f bunt C T llle tla le v is ) fro* Koaksna.
The
r e s u lts in d ic ate d th a t sev era l fa c to rs were Involved but th a t the atre n g er
the r e s i s t ace in the ; re n ts the lee* complicated i s the in h erita n c e.
m r Z R l J & S AMD M S M C D S
th e p aren ts o f the cro ss In ^hich g en etic a W iles on "wnedness,
kernel c o lo r, glome pubescence, and roa c tio n to two physiologic r^ees
Cf T il l e t i * were Mursery Mtosbcr I.>88, » ne le c tio n from R eliance x Rewmrtl
end Wureery Number 1315» a s e le c tio n from Comet x Fhtasnr-Herd Federation,
fh e R eliance x Reward p a re n t i s com pletely mmed (awn type 5) , h&* dark
red k e rn e ls, pubescent g lm e a , i s highly re s is trm t to race tug o f g. t r i t i c .
^att i s Stw centiiae to rnee L-J o f X t le v i* .
The o th e r p a re n t, Uoset x
Fhtcsnr-Hard F ederation, i s awnleas (asm type I ) , has l i g h t red k e rn e ls,
glabrous glumes, r e s i s t n t to L-? but la su sce p tib le to m ce T-S.
In M d itlo n to the stu d ie s made on the cross described slew s, m
F j progeny o f another c ro s s , C om t-(Rusanr-H- rd FM errttlon) x Ceree-Oiepe■lorence), wi-s c la s s if ie d f o r atm type only,
Cne p r a n t, Gewet x fiussur-
Ssurd ^edem tlcm s W. M. 1383, i s ssmlesa (awn type I ) ,
Howewer, a few beaks,
one to se v e ra l mi U im oters in len g th , appear on a few o f the p la n ts .
The
Ceres x ( H o p e - I r e n e s ) . S electio n % nber 60.4,19, 1« on award v a rie ty
(awn type 5) ,
The p a re n t wheat v a r ie tie s (R eliance x Reward and Genet x HnesarHard F edem ti on) and F j hybrid progenies were grown a t the
c u ltu r a l tPcverinent s ta tio n , Soeewn, ltont na in 1937.
w * made by J ,
a.
C lark,
Monten
Agri­
The o rig in a l cross
!v isio n o f Cem al Crops and D iseases, Washington,
Se G., and advanced to the F j Igr him as p o rt o f the wheat Iw rovam nt pro­
gram in the s e le c tio n o f a bunt r e s i s t n t v a rie ty w ith d e sira b le agronomic
-1 5 -
d v im o te re fo r Witama condition#.
The Fg ^ ps grown by Mm xmd r banV-
fre e condition# In 1936,
Fomr hundred and twenty seeds o f e ch o f the parents and JO seeds
* ch o f 10 ' p le a ts taken a t random from the spaced Fg population were Inoim lated w ith a cost o e ito o f m coe L-7 aM T-8, f I l l o t l a l e r l a and T1
t r t t l c l re s p e c tiv e ly , hy shaking an excess q u an tity o f sjo ro s and the
seed wheat in a email paper envelope.
The F^ hybrid and p a re n t p la n ts
were grown on summer fallow land, spaced 3 inches apmrt in rod rows 12
inches apart*
the
Mine rows o f progeny were followed by one row o f p aren t,
re n ts o ccurring ml te rn te ly .
S h o rtly before rip e (atiff-d o u g h etnge) the Indivldm a p la n ts in
each row wore p u lled and c la s s if ie d In the f i e l d in to the fiv e groups;
0 , 25, 5°, 75»
100 p e r cent bunt, based on the p e r cen t o f Infected
heads on e ch p la n t.
The percentage o f stm t in a row was c a lcu late d according to the
follow ing I llu s tr a tio n :
Row 80 o f Reliance x Reward had 25 cmV*free
p la n ts , twelve 25 p e r cent smutted p la n ts , s ix 50 p e r cent smutted p la n ts ,
th ree 75 p e r cen t smutted p la n ts , and one 100 p e r cent smutted p la n t,
( 5x0) ♦ (12*25) ♦ (6*50) ♦ 6x75) » ( I x i p ) - 1^,7 r ^ r cant bunt
47 ( t o t a l number o f p la n ts)
This method n o t only gives the p e r cent o f bunt In the whole fam ily but
mainV ln a the Indivlihrql p la n t as a u n it.
The main spike from each p la n t in an F^ fend.Iy wns Wrwemted,
placed in envelopes, and brought to the lab o ra to ry fo r stmednee# c la s s !-
f l C atlcm.
The hybrid p la n ts o f t h is c ro ss were c la s s if ie d Into s ix c la sse s
Cl ss I , aim less, included those types w ith no development o f aw nlets as
w ell a s those types lik e Hard Federation and Comet % Roeonr-Rnrd Federation
H, Ho. 1383, which eometlmee nay have ownlete up to 2 m . Io n r a t the tip
o f th e eplke.
C lass 2, aj)lealIy aw nletted, has awnlete from 2 to 15 mm.
long a t the nrex o f toe sp ik e , but r a re ly occurring in the c e n tr a l o r b asal
p o rtio n o f the spike.
about 20
mb.
Claes > , weakly p tm letted , has awnlete from 3 to
long, th e s h o rte r occurring a t the base o f the eplke and the
len g th In creasin g toward the apex.
C lass > , stro n g ly aw nletted, has awn-
l e t s from 3 to 1TO no. long, the s h o rte r occurring a t the base o f toe spike
sad to e len g th in c re a sin g toward to e epee where they eeeesd to e length o f
the awnlete on Hnrqule (a ty p ic a l > ) by 10 to 20 mm.
In ad d ltlo n , toe
lower mnlete in c lo se > a re c h a m e te r ia tle a lly incurved.
ULsse U, h a lf
owned, has sh o rt none varying througlwut the spike from 18 to 50 mm. long,
b u t only about h a lf toe lengtin o f normal awns and freq u en tly Incurred o r
re c u rre d , much more than the stro n g ly swmletted c lo se .
toe s u e vary from 30 to MX) ms. in len g th ,
In c la s s 5, awned,
questionable trm ty ee w re
grown in in d iv id u al head rows in 193$ , o n e -h a lf the seed o f entii hood was
plan ted a t Moccasin and th e o th e r h a lf a t Bozeman.
Both p la n tin g s were
need to c o rre c t the F^ c la s s if ic a tio n .
The same spikes need fo r th e
airaedrosee study were c la s s if ie d
fo r c h a m e te r o f gIu see.
The in d iv id u a l heads were then threshed and th e g ra in placed in
se p a ra te envelopes.
vmrut
The th re e c la sse s f o r c o lo r used In g ro w in g th e seed
( I ) dark re d . lik e th e Reliance x Reward p a re n t, (2) medium re d .
»17*
«ad (3) l i g h t re d . lik e the Uoiaet x Hnseay-Hafd fe d e ra tio n
.re n t.
f o r a nor# d e ta ile d etndy o f the In h eritan ce o f awnedneae In the
Reliance-Regard x Comet-(Hnssar-K' rd fed e ratio n ) cro ss rarlem s crosses
In ro lrln g the p aren ts o r progeny were m de in the gresWwwao a t Boeenao,
Uoatasa during the w in ter o f I *37.
Th*
and fg o f these c ro sses wore
grown a t Boaenroi l a the greenhouse during 1938 and e a rly 1939.
notes were taken on both the I r1 and Fg p la n ts .
Awnwlnees
The follow ing crosses
were made;
(I
awn tfl>e)
x l rm it <5 awn type)
Progeny Cl
awn type)
x Progeny (5 owntype)
Progeny ( 3-
awn type)
x P aren t (5 awn type)
Progeny ( >
awn type)
x P arent (5 son type)
f re n t
Progray ( > awn type) x Progeny ( > awn type)
Cne hundred F^ p la n t rows from the Cone t - (Ruas^r-Hard Federotl on)
x G eres-(Hopo-Florence) cro ss were grown a t Bosemmn, Moatmna In I 938
under the same cond itio n s me the 100 s tr a in s from the o th e r c ro ss.
rip e th e p la n ts In each F j fam ily were pu lled and tie d In a bundle.
Whem
These
were taken to the latxirotory and c la s s if ie d In to the 6 awn e la ee ee , an
mweroge being token o f a l l he ds on a p la n t to determine toe c la s s Into
which i t ro e p a t.
The awn c la sse s need were the earns a s in the o th e r c ro ss.
IBOTmHEtiA FSSUMS
Segre- - tlo n o f Uhnroctors
The c h a ra c te rs stu d ied in the Bellanns Bewwrd x ComeV-(HuasrroHfird
Federo tlon) cro ss were a mmdnoes, kernel c o lo r,
rui glume nbesconce.
The
18tipeedtn^ behavior o f a l l Ihren chargeterg f o r the llo e e stu d ied are record­
ed la Table I .
A pedlfprme d l ^ r a a o f th is croes Ie riv en In N ru re I .
In
the Comeb-(Ham9 r-R#r& Fede ra tio n ) x Uereo-(Hope-Plorenee) only oenedneee
has been stu d ied
nd the breeding behavior In ree; e o t to title cl»um cter o f
the 10 Vy lime# Ie given In Table IX.
The r e s u lts o f the stu d ie s In the
Iy g en eratio n follow In the o rd er l i s t e d above.
Asnedneee
The F j end the la rg e r number o f Fg p la n ts o f Seltnnee-Ro ^ r d x
CSomet - ( Hueenr-Kard Federation) were weekly srwnletted (Claes >•) which
aprro®idi more c lo se ly th e awnleee than the owned p a re n t.
A t o ta l o f 100 Fg p lant# was se lec te d a t random fo r growing in
the F j g en eratio n .
The d a m n ific a tio n o f then# p la n ts i s shorn in Table I I I .
The m ateria l proved to be r a th e r v a ria b le , a s there were 36 apparently
se p ara te seg regating grouse, however, se v era l o f these from '* may not
d i f f e r in h ere n tly from the eth e r* .
The modal d&eeee e re Indicated by
UBilerllned number* and the various types a re arranged In the o rd er o f length
o f awns, and to to I s are drawn between groui>a not regarded a s in h ere n tly d if­
f e r e n t.
V arln tloa w ithin the groups m y be due e ith e r to minor genetic
f a c to r s , to environment, o r to the sin g le head e ln e s if lc tlo n method used.
N gttre SC shows photographs o f the 6 asm c la ss e s which were obtained in
th is cro ss and used fo r e la e s if Ic- tlo n .
Aimmg the F^ lin e s th ere was a wide range e f overlapr in g o f owned?ness c la esee .
*!lowing fo r a wide range o f overlapping o f c V s^ e e , the
m a te ria l wne separated Into nine groups (Tables I I I , IV, and V) as
Table I The a in s s l f ten t i r o o f plant# fo r ewnHaese, p e r cent bunt, tm m el c o lo r,
and glume ^ubeecenoe In Fi lin e s and p aren t roes In the cro ss Rellanoei aiswupd (S.S , I 388) * Comet-(Nueenr- ! rd Federation) ( . . 1315) green
a t Bozeman, Mont m l a 1937
1937
*•* o f p la n ts
row
,..having n m Y e
M i____r e r e n t o r cross I 2 > 3t
I S .' t. 13ns (P arent)
23
12 12 5
2 Cross 3513L62 A—1—1
- 3 27 6
I
do
4
do
-7 1 ** 8
5 3
-9 2 9 2 8
5
do
0
do
- U 7 13 5
3
7
do
-12 9 7 5 9
8
do
-13 2
3 10
10
9
do
7 U
-15 3 7
10 M.N. 1315 (P arent) 27
14 13
11 Croee 3511&2 a-1-17
-18
12
do
13 18
-29 4 8 5 I
8
13
do
14
do
6
-21
8
16
2
4
15
do
-2 2 1 0 8
9
lb
do
-23 2 13 10 fa 3 2
-24 6 11 5 7
17
do
18
do
-2 5 1 6 7
-26
2 13 I
2
19
do
20 !*,». 1388 (P arent)
33
21 Cross 351162 A-1-27 4 11 13 3 2
-28 I 2 14 8
22
do
^o, o f p la n ts Ho, p lan ts
having kernel
having
Mo. o f p la n ts
c o lo r
glnaos
h-vlng r s r cent bunt
Dark Med, !pht Glab- Pnbee59 Tt ;oo Ave. red red
red
1
12
3
5
?3
16
8
3 U I
.
5
29
4 2
I
10
>
19
23
29
13
7.6 11
8
lb
7
18
3
14
8
8
16
9.1 10
5
8
10.3
2
16
10
19
21
* 7
8
22
I
8 15
8,1
6
30
I
6
11
11.6
10
21
7
5
6
|
I
6.1
I
7 27
34
24 3
.8
27
19 9
8,0
18
7 13
7
9
0.8
12 16
I
31 I
25
16
S
S
7 2 2
8
16
V;. 7 U
23 6
9.4
6
5 27
26
3
4
lU 12
.8
7
8
23
23
9 .0
I
25 9 2
36
35
10.4
14
6
23 11 2
2
27
9
6
8
2 35.7 14
7 3 3
23
10
.
12
7 2
16
3
3
U .4
8 2
I
22
33
1 16.4 20
21 8 4
I
11
33
4 20
16 4 7 2
20.7
16
7
Table I
(continued)
1937
row
So.
P aren t o r o s o tl I
P i t i r m 351162 >^l-2«35
24
do
"30 5
1- 2-1 0
25
do
26
do
-2 3
27
do
-3
28
do
-4 5
-5
I
29
do
10 ' . s . 1115 (I re n t) 19
31 tiroes 351162A^ - 6 1
32
do
-7 3
-8
33
do
34
do
•11
-12
35
do
30
do
-13
1- 3- I
37
do
-2
3«
do
-3 2
39
do
40 K.H. 1388(Parent)
Ul tiros* 351162 A -> 4
42
do
- 5 15
-6 2
4i
do
44
do
-?
-8
2
45
do
46
do
-9 14
47
do
-11 3
So. o f p la n ts
2
10
7
5
11
314
9
14
10
13 7
U 13
9
5
7 12
>
I
4
9
8
24
15
7 12
6 10
I
4
14
4
2
6
7
3
Mo. o f p le a ts
7
7
6
3
30
5
2 6
4 2
21
27
7
I
10
I
27
10
I
I
6
1
16
I
2
g
7 3
11 11
16 6 4
8
4 2 I
17
23
17
26
20
22
25
21
17
26
19
12
18
15
25
17
12
18
12
21
29
15
26
W
11
26
25
13
4
7
11
S
6
6
2
6
19
7
8
9
5
9
12
4
18
8
4
11
4
15
13
4
50 75 100
2
I
2
I
2
I
I
I
2
I
I
2
2
I
2
5
I
2
2
we.
11,2
4.8
5.3
10.2
6.7
4.8
8.6
2.6
4.7
13.5
11.9
9.3
13.5
4.2
12.5
15.4
4.5
17.2
9.7
5.1
13.4
3.3
17.9
25.0
3.3
»0. o f p la n ts
Se. ria n t#
hawing In rn e l
hawing
o o le t------sluTnee
Dark Med. L ight Qlab- Pubeersd red
red roue
cent
6
20
12
39
4 10
8
7 13
22 10
11
21
22
?4
8
9
6
16
8
30
10
10
21
7 14
6
16
6
6
24
I
19
26
6
23
9
22
11
ll
25
11
21
9
16
12
8
19
U
13
7
17
18
9
27
6
21
15
7
7
18
I
7
29
6
21
5 10
27
6
21
30
21
10 .10
23
4
22
5 18
3
11
11
19
19
16 22
22
13
4 15
8
10
19
4 15
11
13
17
TsMe I
(continued)
Se. o f p l a n t s Se. p la n ts
1937
row
,
So. o f p i s a te
,.m , .sp a ____h
P-treat 0 T c r o ss I 2
Us Creee 351162 * -> 1 2
-14
do
Ug
50 S.S. 1315 <[Parent) Uc
51 Croat 3511*
do * a o U i , " i
52
do
-17 4 5
53
54
do
-IS I 4
do
-19
55
-20
2
do
56
2
1
H
9
do
57
-22 I
do
58
do
-23
59
bO M.W. 13 S I[Parent)
6 l Croee 3511*52 A->2413 18
62
do
-25
dO
-26 7 8
U
do
-27
I
-26 2 4
do
65
66
do
-29 719
-3011 24
do
67
66
do
-31
A-U-I 6 1?
do
69
70 s . s . 1315 1(Parent) **4
I
Tl C rete 35H*>2 *-4-2
do
-3
2
6
72
Ho,
>
'*
21
9
34
j:
6
32
10 10
4 20
10 22
14 I
90
9
4
5
7
I
24 15
72b
15
32
28
3 24
20
U 23
i s 27
11 37
32
24
27 26
52 30
Uo
2 38
39
2
10
b 4
16
It
IS
6 13
15 4
7 6 25
I
33
27
47 42
23
35
13 29
6 3 29
Net. e f plant#
h arin # kerne I
having
c o lo r
Munee
Dark Med. L ight Clab- PubeoATSf ted red
re d roue
c en t
25 % 75 100
9
).9
3
10
7 I
3 27.1
8
5*0
6
2
6 .9
16.9
9 10
16 2
I
I 16.9
2
17
12.5
12.5
9 6
G I
I
6.7
11
2
8.3
a.2
15 10
3
I
0.9
16 4
13.5
I
1.8
I
4
2.4
15 7
8
4 .0
10.4
15 I
4
2.7
14.9
19 6
2
1.1
I
13*4
15 2
9
SI
6
8 .8
3
8
2
7*7
4
8
7
7
14
13
8
30
U3
2b
17
2b
8
10
8
5
5
5
18
20
20
23
26
22
17
23
20
7
9
15
20
15
29
15
29
22
23
23
14
IJ
lb
30
8
7
10
10
6
28
10
37
40
35
17
12
15
10
8
20
9
41
21
38
8
9
12
42
4
16
9
13
30
17
11
23
Ul
29
27
45
32
43
18
52
32
m
51
37
34
34
Ui
44
9
21
34
12
33
26
1937
row
(continued)
Xe* e f p la n ts
kfivlng aim t/rx
-Of .... FfXSM, ,.p/xma...JL.. i , ,.r. I*
10 2 39
73 ones 351162 a^4-4 2 3 21
do
-5
11 19
71*
11 25
-6
8 30 , 17 51
do
75
do
7»
- 7 3 %17 4 U 3 35
do
-9 10 19 9 2 9
77
3f
do
-10
76
I 3 36 36
do
79
-U
3 13 8 5 6 30
SO
1386 (Parent)
61 Cross 3511b2 A-M? h 9 13 , 1 3
62
do
20
5 26
-13
do
-14
6
19
22
42
sI
64
do
-15 I 5 12 10 8 13 33
-Ib 17 15 6
14
do
S
Sb
do
47 >K>
-16 U 12 21 2 S I 26
do
67
SS
do
-19 U is 19
13
do
-20
31 S
89
27
90 #.*. 1315 (Parent)IS
32
91 OKKS 351162 A-Mi 2 11 17 I 6 I 33
do
-22
lb
92
3%
do
-23 6 39
28
93
-24
do
13 20 25
94
do
45
"25
t
19
-26 3 4 9 4 5 6, 10
do
96
do
97
-27
3 21 26 23
:e . o f p lan ts
.
haring kernel
haring
___ co lo r
X lH S L
Dftftc Ned* L ight Olob* Iktbeea*
o. o f p la n ts
8 2
to 4
4
9 3
6
4 3
4 I
12 6
10
3
7
14 2
12 5
9 I
15 I
11 10
11 I
6
7
16 I
12 5
8 2
20 4
12 5
13 7
75 top
3
4
I
2
I
I
I
I
5
I
11.5
1.8
10.1
3*6
5.8
C3
'U
2.6
3.6
9.2
3 30.3
5.9
11.1
4 33.1
8*3
3.9
4.4
15.4
12.2
11.1
17*6
22.3
21.9
x s i21
14
15
29 9
3 23
10 20
9 16
*5 2
10 16
17
37 U
11 23
32 4
13
14
25 11
10 21
11
10
6 IS
20
18 14
7 24
16 8
11 25
l6
27
40
9
10
10
7
13
13
i
55
35
17
2
1
34
S
27
29
9
25
15
S
7
47
13
16
10
H
*10
34
45
12
49
13
9
15
32
34
7
cen t
V
4 ^ 3 *
Tnble I
30
S
n
39
?3
29
26
S
Table I
(continued)
No. o f p la n ts
1937
he v in e awn. tv e
row
I 2 > > 4 5
Ms.___Emrent or <sroee
98 Croee 351162 a-5-1
5 9 11
-2
99
Ho
7 20 15
100S.W. 1388 (Parent)
, k)
5 4
101 Crese 351162 *-5-3 9 8 b
4
102
do
21
9
-10 6 9 18
101
do
104
do
10
-13
7 20
-14
10
16
7
105
do
IOb
do
2 34
-15
I
107
do
-1 7 U 18 5 3 3
108
do
-18 7 13 17
109
do
-19
46
1 1 0 * .;. 1315 (Parent) 39
IllC ro e e 351162 A-961 IS 26
112
do
-22 11 IS 15
So. o f plant#
having kernel
No. plants
feav Ing
No. o f p la n ts
havtme o er cen t bunt
Dark Ned. Light SU b- HhetH
0 5 50 75 100
. l td red___red rone
cent
5.4
2
I
20
16
25
9
4.3
24
I
18
16
26
37
3
10
12.2
I
S
33
5
22
11
2 19.7
3
3 3
19
32
8 19
21 17 I
12.2
12
39
10
27 4
23
33
6 .4
29 10
5 19
13
37
24 8
I
I
9.6
I
20
9 16
13
10.0
8
22 12 I
11
26
9 a
7.4
8
I
16
25
17
33
I
2
22 15
15 17
17.9
i
I
I
28 18
9.2
29 15
12
34
32 7
4.5
39
4
g 10
5.7
9 28
7
8 23
3.4
8
38 6
36
13
C R IM E A N
K M R E D ______________
H ARD RED C A L C U T T A -
M M O U IK
---------------------------
P R E L UEE
f f i t t i i t k o CAL C U m L M A P O U IK
J
f f l m
l*
—
H k P O F E D E R A T IO N H ARD R E D CALCU TTA.
-n s -
M A R Q U IS
H U SSA R
.FEDERATION
HARO F E D E R A T I O N S
Figure I
P edigree diagram o f the 100 F3 lin e s
o f Rellence-Heward x Com et-(HussarHard Federation)
2 5-
T a tts IX The e li ^ a l f lc - tl o n o f p la n ts fo r oanedneeo in J%
lin e s and p aren t rows in the cro ss Cornst-( Htise- r R ari Federation) x Ceree-(^lope-Floronce) ^rmm a t
BoaeeiBne Montano in 1938
1938
row Mo.
UT
118
U9
120
121
122
Si
127
128
129
130
IJl
132
133
134
135
136
137
138
139
1*10
IUl
l i »2
Ceres
Cross
Coast
Cross
1I?
IWt
1H7
1*48
li>9
150
151
Ceres
Cross
k k
Si
Cross 361239 A-l-U
do
-5
do
-6
do
-1 0
flo
—11
do
-lU
do
—16
Coast x Rnssar-Hr rd ^ d e r a tio n
Cross 361239 A-I- 17
do
-21
g-g1
m
114
Pr re n t o r cross
do
—28
to
-29
do
-32
do
-33
do
-3 4
x Rone-Flarenco
361239 a—1-35
do
-36
do
-39
do
—42
do
- 7
do
—9
do
-38
do
—12
do
-3 7
x Ruoenr-U rd Federation
361239 A-l-13
do
4—2—1
do
-3
do
—4
do
-5
do
-6
do
-8
do
-10
do
—11
x Uope-Florenee
361239 A-2-12
I’O.
\
2
3-
5
23
8
3
17
24
27
U
3*»
22
I
26
15
30
I
U
22
34
12
28
1H
U
8
29
U
61
17
8
" 'H T
11
15
52
19
13
50
I
11
28
IU
10
26
5
I
5
34
15
14
18
5
U
15
5
67
58
16
3
U
7
63
4
64
lU
2
6
Ug
2
U
22
26
it
22
21
32
U
16
10
28
19
6
13
16
22
8
18
32
12
2
30
7
lU
5
28
13
2
16
31
9
30
IU
5
16
7
31
5
7
13
6
5
16
23
11
58
16
*5
17
I
35
13
15
14
10
I
62
—26 "»
Taible I I
( coatim e d )
193*
row Mo.
152
153
15*
155
15b
157
15*
iS
161
162
S
167
15
a
U
2
18
2
4
50
13
10
22
26
24
29
22
26
7
29
34
6
7
lb
8
4
29
57
13
8
15
6
6
5
3
15
11
10
5
U
2
20
I
9
29
14
12
5
6
16
*3
%
2
52
I
4
8
28
31
9
32
19
7
a
31
20
25
2
6
11
14
16
17
50
24
24
7
5
I
2
2
14
14
3
9
S?
22
12
15
12
12
29
20
28
4
6
5
2
26
3
18
11
33
30
fi
14
15
11
12
5*
lU
14
55
I
4
7
4
29
10
I
I
rrvS xs
I
2
3
31
30
50
3 S'
Croee 361239 >-9-17
do
—19
do
-21
do
-23
do
-2*1
do
-26
do
-2 7
do
—28
Cchmt x HuflsoiMfcrd fe d e ra tio n
Croee 361239 *-3-29
do
-31
do
- 3*
do
-38
do
-39
do
-MO
do
-* *
do
-13
do
-3 0
Ceroe x Hope-Florenoe
Croea 361239 A-2-Ul
do
*-3-3
do
-5
do
-6
do
-9
do
-13
do
-14
do
-1 5
do
-17
Const x Muaepr-Hnrd Feder t i on
Cross 361239 A-3-15
do
-a
do
-23
do
do
-27
do
-3 0
do
-32
do
-33
do
-3 l1
Ceree x Hope-Floranoe
Cross 361239 a- > 3 5
-36
do
do
-37
trsS
16s
169
170
171
172
173
17**
175
176
177
17*
179
180
181
1*2
1*3
184
185
186
1*7
188
1*9
190
191
192
193
0. o f v
27-
Tnble XZ (oontlnuod)
no. Of plants hnvlnic sen fame
Parent or crooa
Gross 361239 A-3-3S
dO
do
3
-U l
do
-42
do
do
-46
Const x Ifussnn-H- rd Federation
Cross 361239 A-3-48
do
-5 2
-54
do
do
-56
do
-58
do
do
-61
do
do
-62
Ceres %Hope-Florenoe
Cress 361239 * -> 2 0
do
-16
do
-28
-Jig
do
do
A—4—2
do
-5
do
-9
do
-I
-10
do
Const x Hussnn-Hard Federation
cress 361239 a- 4-13
do
-8
do
-3
Ceres x Hope-Florence
4
10
25
31
16
13
8
29
8
5
19
5
I
53
8
I
26
2
17
18
16
2
53
50
12
3
52
22
99
18
12
7
2
6
3
26
22
3%
53
8
4
6
4
26
13
9
2
15
32
29
19
6
17
3
24
5
4l
10
53
9
15
5
2
29
13
5
31
28
15
5
13
12
15
5
10
5
12
55
28
12
10
31
10
5
27
5
Vl H Iv
S S B B S g S g S B S g m
g ii s s s a m
s s s s s s
1938
WHT MO,
57
8
13
4
31
I
15
10
9
12
I
6?,
»«?R*
Teble I I I
AieW r o f F j p la n ts l a eadh —mednesa c la a s In the
nlno r* breeding ^roi^e o f Rellaaoe-Rewnrd * Comet(RaeenT-Rnrd Tederstlo a )
Fj w eeding
- T "' Wtmber o f f t plants la mewdaoeo e lse * Iteu
g
^
3»
R
«5
Tot^l
™ J Z ^ m _______ %
I
9«
86
II
1*19
aj
298
I
2
III
«3
117
89
hg
*»3
IT
6H
183
332
l 6l
119
T
MU
97
10
ISh
T ill
I
2
I
IX
39^
612
97h
996
108
167
TII
629
380
108
TI
Total
h
4hg
899
180
13
2hg
890
199
202
**38
2
231
23h
33«
782
3.999
Figure 2
Awn types found in progeny o f Reliance-Reward x Comet-(Hussar-Hard Federation)
and Comet-(Hussar-Hard Federation) x C eres-(H ope-Florence): A and B, v a r ia tio n s
in awnedness from each o f two p la n ts c la ss e d as 3* awn types from the two cro sses
r e s p e c tiv e ly ; C and D, r e p r e se n ta tiv e sp ik es from the two c r o s s e s , r e s p e c tiv e ly ,
I , aw nless; 2 , a p ic a lIy aw nletted; > , weakly aw nletted; 3* , str o n g ly aw nletted;
4, h a lf awned; and 5» awned.
Table IT Breeding behavior in the T-. feaer^tlea for the e lx awnednese elaeeee Im
the oroee RelIanoe-Heward $ Comot-(tfaeenr-Hnrd iedenstlem) grown a t
Besenen, Montinn la 1937
I , awnleee
.'nn br»gdlnr. for I
JL ?ind 2
I anda
J" lin e Ter cent
of
Number total
Humber o f Fi p la n ts
on
^ar g^nednees olnesee
Cbmerw- 9 -fao to r
> 3b ^
5 T otal ed
baste
3
2
2
7»» 21
PH 65
3
I
2
I
6
I
2
3
I
2
1
6
I
2
HG
15
22
13
33
7
13
16
16
3
95
69
-o&-
Fg elaeeee and th eir
breeding behavior In the
F] generation ♦
T otal
■mlett -I
eegrem tlng for I , 2, and >
I . 2 and 3I . 2. > t H, and 5
I* 2 , sad 3 .
1. 2 . > . ^ >
I , 2 , and >
I . 2. > , H, and 5
I . 2 , > , «md 5
Subtotal
32
IH
Hs
16
69
s
32
106
26
2
31
9
110
39
39
I
Ikg 215 258
I
I
1
33
2?
212
1
1
2
2
U
G29
16
12.50
Table XV (oontinned)
F? lin e#
P er eent
Fg elaaeee and th e ir
Number
to ta l
2
I
3
I
2
I
2
2
I
3
I
2
I
2
Subtotal
12
12
Totnl
28
28
2
3
2
3
15
F j generation *
eefreRT-tin*? fo r I ,
I , 2, > , and >
I . 2, > , > , and
I . 2* 3-» Mid >
I.
> . nnd 4
I . £ , > , > . and
I , 2, > , and 4
I , 2 , > , > , and
r e r c e n t a of
.1^ ria n t#
N^lcnlp t pfl
on
Number o f F j p la n ts
I
2
>
>
*
5 Total
ed
basis
12
12.50
? , and >
4
4
4
19 15 9 18
8 6
9
5 H
15 33 12 18
7 0 6
5 13
14 23 14 3 17
6
5 14 0
7
20
15
27 4 9
6l
32
78
31
Tl
32
75
48 43
3«0
83 117
89
28
> » weakly ftm letted
aefye^ntlng fo r I , 2,
I . &, > . > . 4 , and
I . £ , > . > . nnd 4
I , 2, > . > . 4 , and
I , 2,
nnd 3»
I . 2 , > . > . and 4
2. > , > , end 4
2. > . > . 4 , rnd 5
I . > . > . 4 , nnd 5
2, > . > . 4 , and 5
I . 2. 5- . > , 4 , ..aid
S ubtotal
> , >♦, 4 A 5
5
5
5
15
I
I
I
I
I
I
I
I
I
I
I
I
2
2
28
28
3 22 15 15
14 4g
S
4 l 89 214 73
I
2 14 8
U 8
I
5
2 13
I
3 13 8
2
3 10
4 9 13
2 9 22 20
64 185 332 l 6l
5 S
11
73 44
3
2
5 6
5 9
2 6
13 24
119 97
68
102
534
25
21
18
35
29
g
90
956
?S
^ .0 0
Table Iv
(continued)
Fg elaeees and th e ir
breeding behavior In the
F1 fe n e ra tio n •
True breeding >
>
F^ lin e s
Per cent
of
Nuifiber
to ta l
I
Percentages o f
Fo olante
Calculated
Sumber o f f? p la n ts
on
bar avnednees c lasses
Cbeerr- 2 -factor
ed
basis
3- > ** 9 Total
3
3
108
106
3
3
108
108
31
31
U
I
4
I
1S
13
193
27
9
9
167
13
I 80
9
3
9
3
Subtotal
12
12
Total
I?
17
Subtotal
Total
3
6.29
31
31.?9
9
6.25
12
12.50
17
16.75
> , s tro n fly aw nletted
Tue breeding fo r >
5 .4
Subtotal
se g re m tin g fo r > , > , «nd 5
> » J * . end 5
2, >*• J f , %nd 5
65 178
27 46
94
30
337
log
5 92 224
1?4
4^15
5
Table IV (eontim ed)
Ts
c la ss e s -nd t h e i r
breeding behavior In the
T j generation *
4, h a lf awned
segregating for } , 4 , «nd 5
3*. 4. »ad 5
and 5
>. 4, and _
2. >. 4 and 5
> . > . 4. and ^
4, and ^
>2
2 > > 4 5
4
33
12
I
5
I
I
6
6
were)
6
8
26
2
74 159 202
I
I
2
2
52
32
fiU
43«
16
38
197 197
34 37
231
234
394 612 97*» S99 33« 782 3,999
9 I? 13 15 6 9
562#
$12,9
750.0
• U nderlined numbers In d ica te modal clan*
33 73
13 3b
Calculated
on
Cbserv- 2 -fa c to r
T otal
ed
b asis
134
4l
115
37
9
f
2
5
R atio
) Jbmber c a lc u la te d
64
24 15 2
I
3
I
I
2
T otal
I
I
3
I
I
2
I
^
Grand t o t a l observed
Krpeeted I t fg p la n ts
bulked and s e l f ad
t o ta l
4
12
ond
Pxmber o f Ft p la n ts
by a^medness^claeaes
of
, Ailljr owned
True lareedlng fo r 5
5 ,
>, 4 ,
ltn ee
Per cent
Number
Total
5
Pereeategee of
r9 IRnt=
375.0
937.5
!i,000
562.5
I?
l ?. c0
6
6 .2 9
100
100
C alculation ot Gocdneea o f M t to n l « 2 |3 l ^ i I l l l 2 i 2 t l r a tio (A) and a
I i M i 3»2i I r a ti* (B) on a two* fad to r d ifferen c e fo r awnedneoo from a
Reliance-Eeword x Comet-( Husear-Hard fed eratio n ) spring wheat eroee
grown a t Woaemnn, Montana in 1937
T&Me ?
#2 else### based on breeding
behavior o f the f j and some
o f the questionable F^*e
Ratio
A
B
I
I , breeding tru e
I
0
0
(e-c)
(n-o)^
(0—c)<
e
5
6. 5
1,25
1.5625
.^500
16
12.50
3.90
1 2 .?c00
.0800
12
28
12.50
25.OO
0.90
4
.500
M.0000
.0200
.3600
25.00
3.00
6.55
3 .2 5
5
28
3
31
31.29
0.25
'-.OT-OO
10.5625
.O625
. ' 600
1.6900
.0020
I
5
6. 5
1.25
1.5625
• 2500
2
12.50
18.75
0.50
1.75
.500
3
12
17
3.0625
.0200
.1633
2
2
12
12.50
0.50
.5 0 0
.0200
I
16
I
16
7
100
6.-5
10 . 0
0.75
.9625
.0900
2 , segregating fo r 1, 2, end > ;
2
r a tio 1*2*1
2 , eegregatlng fo r 1 ,2, and > |
r a tio 1*2*1
T otal
2
> , seg reg ating fo r 1, 2, > , > , U , end 5|
r a tio It b*5*312i I
> , breeding tru e
to ta l
4
I
> , breeding tru e
> . seg regating fo r > , > , and 5t
r a tio 1*2*1
T otal
seg reg ating fo r > ,4 , end 5|
r a tio I* 2*1
5 , breeding tru e
Orand t o ta l
Pf 5
Of I
*? o f
X2
Ot
.6853
3.6500
P P e
between 0.90 - I . OO
between 0.90 and 0.80
3.00
fellow s t
I
Il
m
s tr a in s o f awn c la s s I o r 2 th a t haws a eetie o f c la s s I
o r c la s s 2 p la n ts and breed tru e o r w ithin the U n i t s o f
c la s s I and 2 ;
s tr a in s o f c la s s 2 th a t here a mode o f c la s s I , c la s s 2,
o r c la s s > p le a ts and e e g rsg rte fo r c la ss e s I , 2, nnd >$
s tr a in s o f e ls e 2 U e t hare a mode o f c la s s I , c la s s 2,
and c la s s > p la n ts and so u re r to fo r c la ss e s I , 2 . > . >
and 4# o r a t l e a s t U o f them;
If
ntr a in s o f c la s s 3- th a t segregate f o r c ls s ee I . 2 , > , > ,
H and 5» o r a t le a s t fiv e o f Utea;
V s tr a in s o f c la s s >• th a t breed tru e ;
VI s tr a in s o f c la s s 3* o r have a mode o f c la s s > and do not
breed beyond the lim its o f c la s s 3» and 4$
VII
s tr a in s o f c la s s 3* th a t segregate fo r c la sse s 3», 3» and
5, o r have a mode o f c la s s 3* p la n ts with only a few e lfa s
2 p la n ts ;
VIII
s tr a in s o f c la s s 4 th a t segregate fo r c la sse s > , 4 and 5
and have a mode o f c la s s 3*. c la s s 4 , o r c la s s 5 p la n ts
w ith only a few c la s s 2 and c la s s > p la n ts; and
IX s tr a in s o f c la s s 5 th a t breed tru e o r w ithin th e lim its
o f c la ss e s 4 and 5.
Table I I I shows the number o f p la n ts in each awnedneee c la s s in each
o f the 9 groups.
In group I there were S6 p la n ts In 154 Uiat were c la s s if ie d as
o f c la s s
In gros^) IV, which should segregate as In the V2 generation,
th ere were more c la s s > than c la s s 2 p la n ts .
In e l l the o th e r groups
th ere was considerable overlap Ing but the range In tiw se groups was
mere o r le s s d e f in ite .
Probably the g re a te s t v a ria tio n w ithin a group
was in I I I ; however, the segregation u su a lly w*$» nover beyond c la s s 4,
The p la n ts which were placed in c la s s 4 c lo se ly resembled the >
types.
-3 6 fh » r e s u lts o f these stu d ie s In d ica te the presence o f two m jo r
fa c to rs f o r ewn development.
I t I s asexmed th n t the aimed R eliance *
Reward p aren t Ie aabb, and the aim less Coast * fhiesnr-Hnrd fed e ra tio n
p a re n t Ie A’RB.
Ca th is b a s is the phenotypic c la s s , g m e ra l breeding
beh v lo r. and geneIyplc grosze may be In d icated as follows*
Fhenotyyic c la s s , breeding behavior, and genotypic groups
Groups
Glass I , nwalese
I MB® - Breeding tru e
I
C lass 2 , a p ic a lIy o v alet ted
2 AABb - Segregating f o r c la s s I , 2 , and >»
£ M IS - Ssgregptlng f o r c la s s I , 2 , and >
IZ
III
Glass 3“ » weekly arm let tod
4 AaBb - S egrerntlng a s In f_ f o r c la sse s I , 2» > , > ,
4 and 5
2, AAbb - Breeding tru e
5
IT
V
Class 3o, stro n g ly aw nletted
I aaSB - Bnwdlng tru e
£ Anbb - Segregating fo r > , 3» and 5
TI
TI!
C lass 4 , h a lf awned
2 aaBb - Segregating fo r > , 4 , and 5
711%
Claes 5 , f u lly owned
I sabb - Breeding tru e
ZZ
On t h is g e n etic In te rp re ta tio n the gcmdnes® o f f i t f o r both the
6 phenotypic c la ss e s and 9 genotypic grewps, which Is based on the c a l­
cu lated mwibere fo r tvjo major fo n o tic f c to r s , I s shown In T b le 7.
The expected 1*4*5*3*2*1 phenotypic r a t i o f o r the s ix ownednese c la sse s
was eery c lo s e ly cp reached in the observed d a ta ,
th e F T rlae f o r goodness
Jh
o f f i t betaoea the observed and c a lc u la te d d a ta Ie between O.95 rmd 0. 90.
She F value between 0.90 and 0 . # fo r the 1*2*2;%; 1*1;2x2:1 ^eaaetypSc
r a t i o In d lcatee th a t e worse f i t , due to chance alone n lp h t be expected
80 to 90 times l a a hundred t r i a l s .
I t would e en, th e re fo re , th a t two
major g e n etic fa c to rs s a t is f a c to r i l y explains the In h eritan ce o f awnedness l a tills c ro ss.
Although I t i s recognized th a t some o f thn v a r ia b ility
observed m y be cau ed t y minor fa c to rs , i t has been lm a s a ib le to prove
the In te ra c tio n o f an a d d itio n a l run t i c fa c to r o r fa c to rs because o f the
e f f e c t o f u n co n tro lled v a ria tio n caused by environment.
Tronendees v a ria tio n s In ty r* » f awn* on d if f e r e n t head* were
found w ith in p la n ts o f various genotypes Arom th is c ro ss.
In Figure 2A
i s shown the v a ria tio n l a types observed l a p la n ts which in d ic a te s a
range o f awn expression from about >
found In lo se nwned types.
to 5.
S im ilar v a ria tio n s were
Frequently heads from * sin g le p la n t were
found having awns o f more than m e c la s s .
The presence o f those v a ria tio n s
may account f o r some o f the se g re m tln g type# which were n o t in h ere n tly
d if f e r e n t from e ch o th e r.
Te fu r th e r the Im vestlgstiim o f atmodnees In the Reliance-Reward
x Comet-(Husrnr-Rard fed e ratio n ) cro ss v ario u s cro sses were made.
These
were made to determ ine whether the awa - groap c la s s if ic a tio n ns made was
c o rre c t.
The follow ing c ro sse s, giv in g the
phenotypic awn expression
m d the Fg breeding behavior f o r those combinations with the modal c la sse s
u n d erlin ed , were made a s follows*
*2 tre a d in g behavior
P j pheno- fo r aeneAnaoe el ^ees
Croa*___________________
type
I 2 > j»
U 5
F a w a t I «wb tyrie z F eren t 5 ww type
>
♦ ♦
±
♦
♦ ♦
Fregeay I aen type % Progeny 5 man type
>
♦ ♦ jt
♦
♦ *
Progeny > asa ^nie % P aren t 5 smn type
>
♦
S
♦
> - 4
♦
±
♦
>
♦
♦
Progeny j e aan type % P aren t 5 awn type
Progeny > awn type z Progeny > sen type
Photographs o f th e various F ^ 'e obtained in th ese crosses a re given In
Figures 5*» &* C, 0 , and R.
Figure JA shows a fla v o r e f heads which a re
re p re se n ta tiv e e f the F1** obtained in th e p aren t I esn type % parent 5 awn
type; JB, the F1 o f the progeny I awn type % progeny 2 awn type; JC, the
Fx o f tiae progeny > awn type z progeny Je awn type; JD, th e F1 e f the
progeny > own type z p a re n t 5 awn type; and, Ut1 the F1 o f the progeny
> awn type x p aren t 5 awn ty re .
The fo u r types l i s t e d , namely* awn types,
I , J - , 3», and 5, were the only types need in cro ssin g since th ese were the
tru e-b reed in g types observed In tits I y
A few tru e-b reed in g a p lc a lly awn-
le tte d (awn type 2) and h a lf owned (aim type 4) were observed in the
but
no w o e Mes -e re mod# u sin g these a# p a re n ts.
Althowh com paratively email pepnlatione were grown in ootih case,
an in d ic a tio n e f the segregation o f each gron$) ums obtained.
These segregations
were e f prime l a a r t a n e e , since th is would in d ic a te hew the v a r i o u phesotypee would segregate i n l a t e r g en eratio n s.
-3 9 -
otftifiMt
DFffffIl!/^
JfIfHMIli
Figure 3
F, awn types in Reliance-Reward x Comet(Sussar-Hard F ed eration ): A, parent I awn
type x parent 5 awn type; B, progeny I awn
type x progeny 5 awn type; C, progeny 3awn type x progeny 3* awn type; Df progeny
3 - x parent 5 awn type; and E, progeny 3*
x parent 5 awn type
Xb the Comt»( Hussfir-ifard Federa tion) * C eres-( Hor>e-TfI o ronoe)
cro ss only the lnhorl t nos o f awnednees was studied*
The experim ental
deelfia and p lan o f study was the mum es th a t fo r the previous cross*
In t h is c re ss one o f the p aren ts was t a l l y awrv$4e and the o th e r p aren t
had e few In d lv ld aale p re se n t which
In le n g th .
Table I I .
oseessed splfeil beaks o f about 2 am.
The d a ta obtained on the v rio o e F.-, lin e s are presented in
a
photograph o f e re p re se n ta tiv e specimen o f each o f toe s ix
mwned c la sse s from th is cro ss I s glvan in Flgnre 2D.
Thle n o to ria l proved to be q u ite v a ria b le , bu t not me much *»o toe
RelIance-Rewmrd x Oomt - ( Htesemr-Hard Federation) which shewed JS d iffe re n t
types o f se g rsm tlo n .
This cross showed only 21 d iff e r e n t segregating
^TfHJS w ith 9 onto tending major breeding gro% s predom inating, which are
l i s t e d In ta b le # VI end V II.
The modal clnseyma In anch case a re in d ic ted
by u nderlined numbers In ta b le V II.
They a re arranged in the o rd er e f
len g th o f owns, and to ta ls are drawn a f t e r groups no t regarded as in h eren t­
ly d if f e r e n t.
Severn! o f these types n n m w n tly do n o t d i f f e r Inherently
from toe o th e rs .
V ariation w ithin the groups may be due e it h e r to the
e f f e c t o f a minor g e n etic f a c to r o r to environment.
I t seems reasonable
to n esuMB th a t a minor f a c to r may cause some o f th is v r i.a b ility since
some o f the a wales s p a re n t p la n ts had aw aists a t the tip e f the spikes and
one o f the awnless segregates had no awn types beyond the lim its o f awn
type I .
Allowing f o r overlapping o f the c la s s e s , the m aterial was separated
In to nine groups (Tables VI and VII) a# follows*
Table VI Haaber o f *'3 p la n ts In each amednees elnao In the
nine Vi breeding groups o f the 100 lin e s o f Conet( Nussnr-Rrrd fed eratio n ) x Ceres-(Hope-iflersnee)
V^ b reed las
___
I
I
301
27
2
330
II
291
162
58*
III
133
1U3
IV
101
777
,Aa.^?yianaa3,<?^qq
>
288
162
5
150
3
372
189
IA5
V
VI
%
19%
130
IX
I
67»
1.3*7
965
10
3«b
VIII
T otal
58b
10b
1.^70
276
l« 3
1.705
UbS
I
b33
VII
Total
2
Ubg
179
761
153
561
351
352
7*9
5.772
TabU Tl*
BreedUfi behavior In the I*, penerntlon fo r the e l* awnedaeee cU eeee in
the arose Oenet-(Hneeer-Hsfd F e d e ra tio n )* C^ree-(Hore-Tlorence)
at
Bosemml Montana in 1936
FereenUfiee o f
Tg oleseee and t h e i r
breedtofi behavior in the
Tm fioaermtion •
I . awnleee
Tmo breed Infi fo r I
I
I and 2
I , 2 . and >
T o tal
2. a p ic a lIy aw aletted
eegrefiatU fi fo r I , g , and >
I,
end >
I . I , > , and 4
S ubtotal
seg reg ating fo r I , £ , and >
I , g , and 3*
I . £ . > and 4
Subtotal
F er cent
of
HueAer
t o ta l
I
Number o f T, p la n ts
on
bar Armedneos ;classes
Cbserr- 2 -fa c to r
2
> > 4
5 t o ta l
ed
b asis
I
4
I
I
4
I
52
197 19
92 8
2
52
216
62
6
6
301
2
330
9
1
9
I
124 259 143
9 32 19
2
526
62
10
10
133 291 162
2
588
7
3
7
3
102 202
4 i SG
103
47
3
boy
177
10
10
143 288
190
3
584
27
6.00
6.25
10.00
12.50
10.00
12.50
(continued)
?2 c lo se o i and th e ir
breeding behavior In the
I 3 generation *
seg reg ating f o r I , I , 3» . > , H A 5
I, £. > . > ,
and 5
I,
> . >,
and
5
I.
> , >.
end
4
I , > , > . U, and 5
S ubtotal
T otal
_n '""('Iy awnlotted
Jr
J r . and 4
T otal
> , stro n g ly a a n le tte d
True breeding fo r >
>
J e . end 4
> . 4 , and 5
Subtotal
f? Iln ee
Ter cent
Of
Pereeatfiigee o f
---- f g r l o n t e
Colcnlotod
Number o f I* p la n ts
on
-y -•-n "d n o ''r''c .l---^ o o
_ Cbeerv- 3- fa c to r
2
> > h
5 ! fob'll
ed
b asis
Humber
te
IOto
toll
27
I
I
I
27
I
I
I
97 705 147 325 175 96 1.545
3 24
4 19
5
55
I 2« 5 14 5
53
20 b 14 9
3
52
30
30
101 777 162 372 159 104 1,705
7
1
7
1
I
3«7
58
I
44$
I ________4U6
30.00
25.00
50.00
50.00
g.QQ
6.25
8.00
.
387
59
8
8
2
I
I
I
167
213
53
9
1
2
167
222
56
8
8
433
10
2
445
-C tre
Table fIX
Table TH
( continued)
I s o laete* ^md th e ir
breeding behavior in the
F1 generation *
e e g re frttn g fo r > s > , and 5
> . j* » and 5
2. > , > . end 5
S ubtotal
TM lina*
P er cent
of
Stuober
to ta l
I
Percentages o f
. Fjjii^ la n te
c alcu lated
Sumber o f F- p la n ts
on
by stmednees^elseaee
Cbeerv2 3- 3b 4 5 T otsl ed
b asis
12
I
12
I
t 175 353
4 19 31
172
7
700
61
13
13
4 194 384
179
761
10
10
130 27* 153
10
10
1)0 27% 153
5
I
5
I
6
100
6
loo
11.00
T otal
Us h a lf earned
eeg resatln g fo r > , 4, and 5
> » 4, and 5
T otal
5, f u lly aimed
True breeding f o r 5
5
> and.5
T otal
Grand t o t a l observed
I
284
67
I
j55l
678
965
483
1.387
1,470
7*9
Expected i f Fg p la n ts were) Ratio
9 Io
9 15 6 9
bulked and. se lfe d
) Sunber c a lcu late d 611. g
811.8
941.3
• Oaderliaed numbers in d ic a te modal e la te
384
Sg
152 6.00
5,772 i w .60
5.772.6
13.*0
X o tra in s o f o laes I , 2, o r > th a t hr«ve a node o f c la s s I
p ir a te and bread tru e o r w ithin the lim its o f c la ss e s I and
2 w ith only a few c la s s > p la n ts;
IX s tr a in s o f c la s s 2 th a t have a mode o f c la ss 2 and se ^ rsm t#
fo r c la ss e s I , 2, and J - ;
XXX s tr a in s o f c la s s 2 awn type th a t have » mode o f c la s s 2 and
eeffpefiate fo r c la ss e s I , 2 , and > ;
XV s tr r in e o f c la s s 2 a m typo th a t have a mode o f c la s s 2 and
!WTesnte f o r c la sse s I , 2, J - , > , U, and 5, o r a t le a s t
fiv e o f them;
V s tr a in s o f c la s s J - th a t breed tru e ;
VX s tr a in s o f c la s s Je th a t breed tru e , o r have a mode o f c la ss
Je p la n ts w ith only a few o f c la s s 4 and c la s s 5 p la n ts ;
VTX s tr a in s o f c la s s Je th a t sSfTSfr to fo r c la sse s J - , J e , and 5
w ith a mode o f c la s s Je p la n ts ;
VXIX s tr a in s o f Clas 4 th a t have a mode o f c la s s k p la n ts and
segregate f o r c la ss e s > , 4 , and 5; and
IX s tr a in s o f c la s s 5 th a t breed tru e .
The number o f p la n ts in or oh awnedness c la s s l a each o f the nine breeding
groups i s shown in Table VI.
In Group I 2? p la n ts in JJO were c la s s if ie d in to c la s s 2 end two
in to c la s s J - .
Cne lin e o f 52 p la n ts wrs observed which showed no over­
lap Ing in to the c la s s 2 type.
In no o th e r /group w ith a p o ssib le ex­
c ep tio n o f Group VI (tru e breeding »
c la s s e s .
was th ere very much overlapping o f
There seems to be an overlapping o f Uie Je type in to type 4.
This might be expected sin c e some p la n ts had heads with types varying
from Je to 5#
This i s c le a rly shorn in Figure 2B.
In Table VIX i s given the breeding behavior by g roups based upon
the s ix owned c la sse s o f th e 100 F^ lin e s .
The Fg genotypes were determined
Igr the breed In#; behavior o f the T y
The re s n lte o f th ese stu d ie s In d lcote th a t two major fa c to rs are
$re s e n t which Infloence the development o f asms.
I t I s assumed th a t the
awnlees Comet x !W sar-R ard Feder tlo n p aren t Ie M M , and the armed
Ceres x Ffon'e - Florence p aren t la eabb.
On th is b a sis the m n o m l breeding
behavior and flonolgrplc croups megr be asmmed to bo the same ns In the pre­
vious c ro ss stu d ie d .
The goodness o f f i t , breed on the calcu late d numbers f o r two
g e n etic fa c to rs on both 9 genotypic and 6 phenotypic c la s s e s , i s shown In
frb ls f i l l .
Thsgrraament Ie good throughout.
The P value f o r the e r e c te d
genotypic r a tio 1*2*2*%;I i I*2:2*1 cornered w ith the observed wn> between
0.70 and 0.80.
The P value f o r the c a lc u la te d phenotypic r e t i e I t 8*1* 3*2* I
compared: to the observed dr to w-s m ch h ig h er, being between 0.80 nnd 0. 90.
In e ith e r ease the P value i s h i # and a worse f i t , due to elw ioe alone,
might be ear acted to occur on *a -v er ge o f 80 times la I O t r i a l s .
It
would seen, th e re fo re , th a t two g e n etic fa c to rs s a t is f a c to r i l y explain the
I n h e r it nee o f asnedness in th is c ro ss.
In th is cross as l a the former
minor fa c to rs any have played a ro le In awn development, b u t because o f the
e f f e c ts o f u n controlled v a ria tio n ceased by environment I t was im possible
to prove the presence o f such fa c to rs ,
In a few lin e s th ere were p la n ts which were widely d if f e r e n t In
pvm type from what might have been expected in th a t Inh r i t m c e group.
These
few widely d iv ergent tri e s may have been due to cro ss f e r t i l i s a t i o n In the
p r e v io u s generation a# i t la known th a t some c ro ss f e r t i l i s a t i o n does occur
In whes,*.
F3 p la n ts re s u ltin g from c r o s s - f e r t i l i s e tto n could and would be
ta b le f i l l
CalculaUcm o f Oeedneoe o f F it to e 1*2; *4;I : I * 2;2*1 r a tio ( a) and a
Is Si I i ) i 2i I r a tio (S) on # tw o-footer d iffe re n c e fo r awnedneos from a
Cemet - ( Hub as r-Hn rd Federation) ar Ceree-(Ikipe-Florenee) spring wheat
crone erotm a t Bosmmaa, .’Montana la 1938
Fg c la sse s based on brooding
behavior o f th e F^ generation
I , breeding tru e
Calcul- ted
on
Ratio
2 -fa e to r
A
B Cbeerved d ifferen c e
I
I
6
6.25
10
12.50
10
0-0
. 2$
(o -e )2
Ia=SlI
e
.0625
0.01
2.50
6.2500
.50
12.50
2.50
6.2500
.50
25.00
go.oo
5.00
0.00
25.0000
0.0000
1.00
0.00
2, segregating fo r l,2,am d > *
r a tio 1*2*1
2
2 . e eg rem tln g fo r 1 ,2 ,and > t
r a tio 1*2*1
2
2 , segregating fo r I
,
4, and 5*
r a tio It 611*3*211
4
Total
6
30
50
> , breeding tru e
I
I
6
6.25
1.75
3.0625
.**9
3», breeding tru e
> , e e g rem tln g fo r > , > , and %
r a tio 1*2*1
Total
I
6
6.25
1.75
3.0625
.Iw
3
13
21
12.50
M .75
.50
2.25
.2500
5.0625
.02
.27
2
2
10
12.50
2.50
6.2500
.50
I
16
I
16
6.25
1.25
1.5625
100.00
F s very high fo r r a tio doalpnatod
P a w ry high fo r r a tio dee l/matod
.25
4, segregating fo r > , 4 , and 5*
r a tio 1*2*1
5» breeding true
Ctund to ta l
%
2
Of 7 ^ f o r Ratio A - 3.76
Ot 5 a2 f o r Ratio % a 1.52
5
100
n eet cone icttmia l a Orotme I . XXI, TI, T H , snd Xx, sad I t Se l a theee
groups th a t the d e rla tlo M a re aeen.
A few dev iatio n s m y he due to
f ix tu r e s , h a t aone dew latlona (Groups X and TI) occur toe freq u en tly to
be cloesed %# adxtures o r the r e s u l t o f c ro e e -fe r t l l l e tlo n .
I h ie ron^e
ie p robibly caused by a minor modifying fa c to r sad I t s a c tio n c m be noticed
o n ly la those gronpe.
Kernel Color
Since the parent* In the TfellanetWtmmrd x Oorot-(Ihiesa)Vfferl
Feder-itlo n ) c ro ss d iffe re d l a In te n s ity o f redness o f seed c o lo r, the
seed o f the In d iv id u al heeds from the p la n ts in the 100 Fg lin e s were
c la s s if ie d f o r seed c o lo r in to the classes*
red .
dark, in term ed iate, rod I l h t
The seed o f the Reliance z Reward p a re n t was much darker red than the
o th e r p a re n t.
In the Fg lin e s th ree breeding types appeared among the
progeny, namely*
( I ) lin e s In which only l ig h t and interm ediate red-seed­
ed p la n ts were found, and ( 3 ) lin e s in which th ere were found dark, in te r ­
mediate, and l ig h t red-seeded p la n t# .
Since th e l ig h t red-oeoded p arent
hod both l l # t and interm ediate types and the dafk red p aren t had some
Interm ediate ty r e s , i t was thought J a s t l f ls h le to clo se an Fg I l r o so
breeding lik e the l l # i t :« r e n t. I f i t
osseeeed only l ig h t - rnd medium-
red Iosroeled p la n ts , and lik e the dark re d p a rro t. I f only dark and In te r­
mediate type# were found.
The o rig in a l d a ta on d a s ! f i c tio n f o r kernel c o lo r Ie given In
T b le I .
The Iareedtng behavior o f t wonty-elg h t Fg lin e s warn lik e the l ig h t
red p a re n t f e r k ern el c o lo r; tw enty-six bred lik e the d ark red p a re n t.
-1*9v h il# tho re m ln d o r, o r **6 lin o s , m epom teA f o r a l l throe types.
I i a ^ the p la n ts f o r the
rod*
w g re m tln e M eea th ere no re obtained 357 dsrfe
5?«6 Interm ediate radi
•05,5 dark red*
T otnl-
Ul5 l i# h t red a* coapared to an expected o f
809 interm ediate red*
U0U.5 l i g h t rad .
GalcnlaM ng Chi-squiure fo r goednees o f f i t fo r kernel c o lo r cm a
sin g le -fa n to r d iffe re n c e , 1*2*1 r a t i o , on the 100 F-j lin e s showa the ob■erved number to be very clo se to the e ^iected vhloh i s given In fn b le IX.
P value f o r goodness o f f i t between the observed and c alc u la te d d ata
i s between
OmJ O sad CU 50 .
Table IX CalSBilation o f Goodness e f M t on a ona-fnotor
d iffe re n c e f o r k ernel c o lo r In tho IU generation
o f B e lianos- Havard x Comet- (Tbiss -r-m rd Federati<m)
groan a t Bozeaan, Montana in 1937
Cbaerred
C lass
C alcu lated
on 1*2*1
r a tio
O-C
( c - c )2
Co-c)2
c
Sag. fo r I i S i l o f
dark* lnt* l i g h t
U6
50
-U
16
.32
True breeding fo r
llg jh t c o lo r
28
25
♦3
9
.36
Tm e breeding fo r
dark c o lo r
26
25
♦1
I
.OU
100
100
T otal
Df 2
X2 o f 0.72
P
s
between O.70 and O.JO
50Clune Poboaomee
Slaee the p aren te d iffe re d l a ^ lone pobeecm ce, the IOO
lin e s
o f Rellaaeee-Seeaid * Gome#-(Heasar-Hard FederziUon) provided oxcollont
m ateria l f o r a study o f the In heritance o f th is c h a ra c te r,
the breeding behmrlor o f each I la e cure #lvam l a Table I .
grsaps eppeened l a A e progeny, nanolyi
th e d ata on
Three bm edlag
(I) tru e breeding fo r glabrous
gtoaoe. ( 2) tru e breeding pubescent tamee, and (3) those e eg reg rtln g fo r
glabrous and pubescent g lu aes.
I a a few o f the F , li n e s abnormal r a t io s r
eared.
For In sta n ce,
In son s fa m ilie s I o r 2 p la n ts were pubescent sa d th e r e m in d e r gl-brem a.
Karwle^s (1932) T ables o f P r o b a b ilitie s were used a s a b a s is fo r c l a s s i ­
fy in g th e se l i n e s la te e it h e r th e s e c r e t? tin g o r tru e-b reed in g group.
The d ata o b trln e d Aow a c lo se f i t to a 1 :2 :1 r a tio which m jr be
explained by e sin g le g en etic fa c to r d iffe re n c e w ith the pubescent con­
d itio n dominant to glabrous.
Twenty-four F-j lin e s bred tru e f o r m besew it
glumes; twenty-one bred tru e fo r glabrous g lu m e; A l l s f i f t y - f i v e seg­
regated fo r ubeecent and g la rmis glume* l a a 3:1 r tlo ,
In the f i f t y -
fiv e eeg rag atlag lla e e th ere were obt Iaed 1319 pubescent and UUg .glab­
rous p la n ts a s compared to the ext tee ted number o f 1320.75 pubescent and
UUo.? 5 glabrous types.
I t was thought advisable to t e s t A s observed number o f breeding
t y a s w ith the c a lc u la te d number.
o f f i t i s shown In Table X.
*50 and
The c a lc u la te d F value f o r goodness
The P value was determined to be bo twoon
.7 0 A le h I s a clo se f i t to a 1 :2 :1 r a t i o .
The discrepancies
between the observed and c a lc u la te d r a tio s coa be assasmd to be due to
51
chance and no% to
bcbki
o th e r gone t i c cans©.
fa b le X C alcn latio n o f OooAness o f F i t Ib r in h eritan ce o f
rltiiio ^rabescence on a s i ^ le - f r a c to r d iffe re n c e in
100 F j lin e s o f Roliance-BeiFard x Coneb-(HnosnrRard FeAmrrtioa) groan a t Bosersan, MmWm in 19J7
Breeding behavior
(o - o )2
C
S eg regation fo r pubeacm t nod
glabrous glumes; r a tio 3*1
55
50
5
25
•30
True breed ing f o r pu bescent
glumes
2*1
25
I
I
.(A
True breeding fo r glabrous
glumes
21
25
u
16
.64
100
100
Ib ta l
Df 2
X2 o f I . IR
F =
between 0.7<5 end O.50
Msy etion
tlthongh t h i s experlm m t m s d eal^rtrd to bo an in h e r it' nc© study,
the r e s ta te obt Ined ecn not be l n t e r reto d in th a t lig h t.
In stead o f
determ ining the re o tlo n o f the 100 F-, lin o s to e ch o f the two physio­
lo g ic r- ces o f bunt, as i s naosenary in % gene t i e study, a composite o f
the inoculum w-e used.
Various worlrere hrve concluded thmt i t i s e s s e n tia l
to work w lto sin g le p h y slo lo rlc rscee r a th e r th-.n w ith s Oeepeelte o f
se v e ra l to study the ln h o rltrn c e o f re s is ta n c e to bunt.
se v era l r-oot; composited i s
the h o st in d iv id u a lly .
This
The re o tto n o f
d if f e r e n t fchrn when r-tees a re noting m
rob bly accounts fo r the c n la x ity o f the
•53IalMVfttMMM o f reaetSoB to boat In e a r l / InooatM attoino.
H m i re o e e t work
has shown the Inhaxltaneo o f ro eletan ee to troat to be le e s complex than on#
In d icated by aome o f the e a rly to re # tlgR tlone.
Fercenta^ee o f in fe c tio n by boat fo r the paren ts and 100
lin e s o f
the Rollanee-Reward % Cone!-(Tluoer r-R ard Feder t i n) cro ss a re Iv e n In
Table I .
The range o f the Reliance x Reward p a re n t fo r bunt In fe c tio n wee
from 12. ? p er cen t to 23.9 p e r cent w ith an average o f 16.2 p e r c en t.
The
ottior p aren t Comet x Bueear-Fbrd FoAor tlo n shewed more re s is ta n c e and e
much narrower r age o f snece??t l b i l l ty , the range being from 2 .6 to 5*1 p e r
cen t In fe c tio n w ith an average o f 4 .0 p er c e n t.
Ihe range o f the F3
regeiy
extended beyond the lim its o f e it h e r the r e s i s t a t o r su sc e p tib le p a re n t.
Six o f the FjJ lin e s averaged lower In fe c tio n than the lower H a l t o f the
r e s is ta n t
re n t.
The meet r e s is ta n t F^ ' roPeiW lin e averaged 0 .8 f o r
cent bunt, while th e most su sc e p tib le lin e showed 3 5 .7 p er cent bunt.
Five
lin e s were more su sc e p tib le to the com o e ite than the snsce t lb l e p aren t.
Forty-two lin e s averaged bunt In fe c tio n between the lower l im i t o f the more
sn se e o tib le p a re n t and upper lim it o f the more r e s is ta n t p a re n t.
Ttto standard d e v ia tio n fo r percentage o f bant fo r OOdh o f the
I £ &
p a re n ts wee c a lc u la te d by th e fo rmula y H - I . Fren Ittle th e
probable e rr o r woe obtained by m ultiplying the etondnrd d e v ia tio n by
0.6745.
S r the use o f th is method the
.* . o f Reliance x Reward was
c a lc u la te d to be * 3 .4 g iv in g n c l^ a s r?mge o f 1 2 .8 p er cen t to 1 9 .6 per
c en t.
The P.K. o f Comet x «uesax^»fsrd Federation was c a lc u la te d to be
* 0 .7 , g iv in g a c la s s range o f 3*3 P»r cent to 4 . 7 p er c e n t.
From these
c a lc u la tio n s the c la ss e s wore determ ined, l . e , , those F j lin o s which would
-5 >
f a U wrlthla the a m t FMflMatAgM o f 12.8 p e r c en t and 11.6 j-er c e n t ml/^tt
be e j e c t e d to breed the sane ae the R ellaae % Heward pmr e n t, and those
f a l l l e e w ithiii the range o f 3«3 p e r cent and 4 .7 p e r cen t n iffit be RTpected to r e a c t lik e the note r e s is ta n t p a re n t.
Consequently, the lin e s giving
percentage* o f boot below 3.3 p e r c en t were c la ssed a s being mere r e s is ta n t
Vtan the r e s i s t n t perm it.
lBhoae lin e s f a l l in g between the n p e r I i M t
( .7 p e r cent) o f th e group brooding lik e the r e s i s t a n t p a re n t and below
the lower I l M t (12.8 p e r cent) o f those breeding lik e th e su sce p tib le p aren t
a re classed a* in term ed iates and comprise the M ass 4 .8 p e r emit to 12.7
IMwr e n t bant In fe c tio n .
Since 19.6 p e r em it i s the u-p^er l im i t o f the
c la s s f o r those breeding lik e the su sc e p tib le p aren t a l l the lin e s which
are showing Ixmt percentages o f 19.7 and above a re classed a s being ro re
su sc e p tib le thnn the su sc e p tib le p a re n t.
Thus the 5 bunt c la s s e s were as
follows*
O
-
3.2 p e r em it
Cl ®s Midpoint a t
1 .6
p e r cent
3*3 -
4.7 p « r cen t
Glnss M dp elat a t
4 .0
per sent
4 .8 - 12.7 p e r cent
uloee Midpoint a t
£.75 p e r cent
I .8 - 19.6 p a r cen t
Clnss Midpoint a t 16.2
p e r emit
19*7 - 35*7 p a r cen t
Glass Midpoint n t 27.7
p e r cent
Using these bunt c la s s e s , the p a re n t re a c tio n In percentage o f bunt­
ed p la n ts , th e number o f rows o f each p a re n t, mid the eegreg tio n in the
XMlsno#■Reward * G m et-(HnsoaMfcrd Pedern tlon) cro ss as shown tgr the
average o f the Pj hybrid s t r a in s , a re shown In fa b le XI, and g rsp h ie a lly
In fig u re 4.
511-
X REWARD
16.2 i 3 4
r e l ia n c e
_
COMET
PE
HUSSARHARD FEDER­
ATION
PE. 4.0± 0.7
PERCENTAGE OF BUNT
Figure 4
Frequency d is tr ib u tio n o f the average
percentage of bunted p la n ts when in­
oculated w ith a composite of races of
T-8 and L-7 o f T l l l e t i a in p a ren ts and
progeny o f Reliance-Reward x Comet(HusBar-Hard Federation)
55fn b le Xl J1Orcentc^es o f banted ',rheai p i'ia ts In rows o f
Helisaoe % Reward sad Coaet x Ruaaer-iI- rd le d e re tlo a
a re n te and
hybrid s tr a in s o f the c r o s s ^rrewn s i
Hosefem, Moatmio l a 1937
Comet % Kuaear-
C laeew l a
n e r cen t bunt
Munber to r cent
0 -3 .2
Per cent
Btimbcr Per csj
9
9
2
33.3
3 . y - * .7
12
12
2
33.3
b.ft-12.7
5H
*
3
50.0
2
33.3
12.8-19.6
15
15
I
16.6
1 9 7 - 35.7
10
10
2
33.3
100
100
6
99.9
6
99.9
T otal
The hybrids ee& rem ted beyond the U n ite o f th e p a re n ts.
Twelve
o f the IDO etr a in s had a boat m o tio n sim ila r to th a t o f the Comet x FhiaearBhrd F edem tloa p a re n t.
At the o th e r end o f the curve th ere
w ith in the lim its o f the Rellanee x Reward p a re n t.
ere 15 s tra in s
Considering the fiv e
c la s s e s f o r bunt which were formed on the b asis o f the breeding behavior
o f the p a re n ts toward the composite lnoeulnn o f the two m o w the number o f
fa m ilie s in e oh c la w would be a s follows!
9
12
more r e s is ta n t than the r e s is ta n t p aren t
lik e th e r e u le te n t p aren t
5& Interm ediate between the two parents
15
lik e the su scep tib le p a re n t
10
more su sc e p tib le than su sce p tib le
a rs n t
Lintenfra Stqdlca
Linkage e Indies have beat mmde in c e rta in phases e f g e n e tic s, and
many linteige grrmm have been determined in mxmroua crop p la n ts ,
Cmt^
p e ra tiv e ly fe e cases o f linkage have been rejjorted In Jhe-I croasee,
A fee
Im re F itlg a to re studying linkage have been c ite d by Rayes oM Or rb e r CI93H),
Some e f them mentioned a re
i f fen , XbgLedov, Cninoa and Care te a s .
I t Ie
always o f value and e f i n te r e s t to the p la n t breeder to know whether a
e h a r a c te r Ie linked w ith another, in th a t good ngrononic u s l i t i e s may be
lin k ed w ith o th e r good o r poor c h a ra c te rs,
Conso.piently, c e r t in good
agronomic q u a litie s may be tra n s fe rre d w ith e-\se o r d if f ic u lty depending
on the s tre n g th o f the linkage o f the o th e r fa c to rs .
In th is study th e d a ta on the three c h a ra c te rs, ownedneas, kernel
c o lo r and glume pubescence, are compared with the data on se n t r e s i s t no©
in o rd e r to determine whether o r not th ere i s linkage between the fa c to rs
c o n tro llin g these various c h a ra c te rs,
Centingency ta b le s were prepared
to coHTiare the various c h arac ters to determine whether o r n o t toe foe to rs
determ ining the c h a ra c te rs a re lin k e d .
Chi-Gpmre snd P values obtained
In each co^aarleo* o f c h a ra c te rs a re as follow s:
Ghi- Degrees
square
of
P
G M B s m a __ ___________ ..............................
13.#*%
Ib
0. 50- 0.%0 InAet . Ass1Id
Aimedaees v s . glum# pubescence
Aamedneee v s. k ernel c o lo r
IO.976
16
0. 80- 0.90
do
Mmedness v s. p e r cen t boat
3^.090
32
0. 3*6
do
0. 80- 0.90
do
Slums pubescence v s. kernel c o lo r
1.272
Slumc pubescence v s. p e r cen t boat 5*829
8
0. 50-0.70
do
Kernel c o lo r v s. p e r c e n t bunt
8
0. 05-0,02
e e s. linkage
16.S 17
'7-
C olculatlon o f ladepeeidence Iqr Jriaherfa ( 1936) method eae employ­
ed to o b tain th e Chl-equ-^re value when eoa^arlng tite obs rved w ith the
C--Iosilatod number.
The I' value was obtained a f t e r the Chl-equare value
nd degnioe o f freedom
ore d tcrn ln ed .
"a" has n o t been c alc u la te d beyond 30.
In F ish e r’s ( 193b) x2 fab le,
Since the decrees o f freedom
between awnedneee n i p e r c e n t boat was >2, F is h e r's method o f e b t ln ln g
a "mod ' . ro x lm tlo n o f i*
ondoyod.
He assumed th».t (2 .V2-S V it-J)
I e norm ally d is trib u te d about zero w ith u n it d e v ia tio n .
I f th e re s u ltin g
q u a n tity obtained I s g re a te r than two, th e w in e o f X2 I s n e t In aocofdanee
w ith the expected.
In th is oos%mrl#ou x2 equals 3U,09| s u b s titu tin g In the
formula (2 ViTx2 - V r n - I ) «# h.=tve (✓
. ,1:-
- V (jM-1) o r ", ;Ua fo r the
value o f F which I s w ith in the expectation f o r lndep«id«ace.
Very l i t t l e . I f any, re la tio n s h ip e x le te between amaedness
gluna
ibesconce,
nd
wnedneee and !cornel c o lo r, a«maednees and bunt per cant,
glume pubescence and bunt, and glume pubescence and kernel c o lo r, since the
obtained r e s u lts c^me w ith in W gaotatlon f o r lzv!e$indent assortm ent.
How­
e v er, acme re la tio n s h ip e x is ts between p e r c e n t bunt and k ernel c o lo r,
sin ce the progenies a re mere o r le e s confined to c e rt- in bunt elaeeee
dei ending on tho c o lo r o f the k e rn e l.
A e o rap r i eon o f smut re s ista n c e and seed c o lo r In th e 100 Fj
progenies I s shown In Table XII.
In th is ta b le I t may be observed th a t
the g re a te r number o f r e s is ta n t p la n ts tend to have lig h t red seed and the
more su sce p tib le ones to have d a rk e r k e rn e ls.
The n ature o f th le d i s t r i ­
bution In d ic a te s a p o ssib le llmkngs between #mn$ re s is ta n c e end seed c o lo r.
56reelet-m ee to a m t ? e tn r ItakaA w ith l ig h t c o lo r ;snd e n s c e p tib lltty with
dark « l » r e Ihe P ra lu a w a caleulateA to W between OeOg aad 0,02 which
I e WfnrAeA a s n e t being InAexandent assortm ent, M t Ia d le tee a t Ie a e t a
weak lin k a g e ,
In In te rp re tin g re s u lt* i t Ie eafe to sBenm th s t i f the
value o f P fb r any d is tr ib u tio n Ie hig h er than 0.0$ there Ie no evidence
o f s lg n lf lc r n t c o rre la tio n between the c h a r c t - r s being considered,
Ibible XII
A cow: <ris e n o f a m t re e tio n to CFtsonite o f
&»8 and L-7 a m t roeem w ith weed c o lo r in 100
lln e a o f Rellanoe-Rewrrd ar Come!-(W unmrmrA fed eratio n )
Tme
breeding
dark red
SeereFn tin g dark,
in term ed iate, aad
0 - 3.2 p e r cent
I
3.3 - U.7 p e r cent
4.8 - 12.7 p a r cent
True
breeding
l l s h t rod
Total
2
6
9
2
6
4
12
12
26
16
5*
o r cent
6
9
0
15
19.7 - 35.7 p er cent
5
3
2
10
26
46
28
100
12.8 - 19.7
T otal
'-IwCV.,.I. M AMD Ct^CLUBK HS
Two cro sses Reliance-Reward % C onst-(Hnesrr-Rrrd FeAerntlo n ) rn d
Uomt-CIMastir-K rd Federation) x Ceree-(Kepe-Florenee) were studied fo r
the i n h e r i t nee o f asmadneas.
the 100 F j lin e s stu d ie d ,
5H a tin c t breeding groups
were
obtained in
In the R elisnce-Reward x Uoaet-(ftas8'>s^Hsr4
F edrr t i on) cro ss r a th e r wide d ev iatio n s from the exj sotod ^ore obtained.
59A muAer o f lin e s were observed w ith p la n ts d iffe rin g froei the m ajority
e f p la n ts In th a t lin e .
Comparatively fe* lin e s e f th is ts- e were ob­
served In the second c ro ss stu d ied .
Ih e method e f u sin g a sin g le head from a p la n t to e s ta b lis h the
asm type f o r th a t p la n t m j account fo r maeh o f the v a ria tio n which w e
observed in the Reliance-Pew rd x Comet-(Hnss-- r-P ard FOder tlo n ) erase,
wide v a ria tio n s were found to api^enr w ithin a sin g le p la n t.
V ariation#
on sin g le p la n ts were found to range Arom stro n g ly aw nletted tyue to
eem nletely owned sp ik e s.
Cn p la n ts possessing fewer awns the r nge was
from com pletely awmloss to weakly a ra le tte d ty p es.
Thus, considerable
d e v ia tio n s may appear depending neon the sin g le head se le c te d f o r
c la s s if ic a tio n .
-■r awns.
F ie ld e b s e rr tlo n s In d ica te the l a t e r culms to have Iong-
The f i r s t spikes which appee® o rd in a rily have much sh o rte r asms
tlian do the l a t e r heads.
Why these l a t e r culms a sse ss Spikes with
lo n g er sens then the e a r l i e r spikes has not been determined.
Further
research on th is phase needs to be conducted.
Miner modifying fa c to rs may have played en Im rcrteRt p a rt in
c using th ese v a ria tio n # w ithin some lin e s .
At a re n tly th e only e ff e c t
o f the Hlivtr fa c to rs In the preeance o f the major fa c to rs I s to reduce
the amount o f awnednees, a# f r e e the stro n g to tee weaker lim its o f a
c la s s .
I t has been I n o a slb le to dem onstrate the l n t e r c tia n o f an
a d d itio n a l gone t i e f a c to r bee use o f the e f f e c t e f uncontrolled v r i tirm
caused by environment.
The jeten o e o f th e minor fa c to r In the B elieace-
Reward % Comet-(itueear-Rard Federation) was ep- e r e n t, w hile In the Comet{Bnssnr-H rd Federation) x C eres-(Mope-Florsnos) tee e f f e c t o f minor fa c to rs
*6o*
w«* le s s pronmmeed.
^rldnnee p o in ts toward a Pdm r fa c to r W ing p resen t
l a the l a t t e r a re a s , sin c e a tru e br eding completely iw nless lin e v.m
obtained in the Wy even though the Coast x Haeaar-Rard F ederation was not
o f th is ty ro .
P a ren t p la n ts were o ccasio n ally found -rhlch possessed awn-
I e to rxp to sev eral m illin e to rs la len g th and consequently v e rs placed in
the ar ic a lly m m letted c la s s .
Cross f e r t i l i s a t i o n I s known to occur In »he t .
Xt has been shown
by the work o f L elg b tf RBd Taylor (1926) th a t as h i # a s 3*» p e r cent nay
h i c c acted l a c e r t In
Her t s and en v iro n m n ts.
A few o f tineas widely
d iv erg en t types m y have been due to cro ss f e r t i l i s a t i o n In the previous
g en eratio n .
In c e r ta in grotCHi n a tu ra l em eelng can no t W observed but
In c e r ta in breeding types i t I s ebvlons.
Evidently fo u r tru e W eeding awn types can be Is o la te d .
These
types a re apparently due to two major g e n etic fa c to rs (AA and SB) which
c o n trib u te d if f e r e n tly toward the expression o f the awns.
The f in a l
awn expression i s dependent upon which fa c to r i s o r fa c to rs a re p ro ren t.
I t l%s been shown th a t there i s an in e r f mot dominance o f awnleesness.
The awnod type i s the phenotypic expression o f the double rec essiv e o f
these two fa c to rs while the double dominant I s the avmleee expression.
F acto r M
when in combination w ith a double recessive o f th e o th e r fa c to r
fo r awnedneer produces a tru e breeding weakly aw nletted type w hile the
oi'poelte co n d itio n , a s t r a in o f wheat having the BB fa c to r w ith the double
rec essiv e o f the o th e r fa c to r , w ill produce a stro n g ly
a n le tte d spike.
True breeding apic l l y aw nletted and half-owned s tr a in s o f wheat have been
known to occur.
Sons o f those true
reeding ty a s were obs erved in the
fit In the Reliance-Reward % Gonet - (Husear-Hard f e d e r tio n ) c ro ss which
a^aln In d ic a te s th e presence o f a minor f a c to r In IM s c ro ss.
Since the o rig in a l cro ss has been m de and the F1 produced. I t
can be seen In Figure 3* Uiat In the Relianoe-Roeard x Gomb-( JftiaearHard Federation) c ro ss the
w s weakly a e n le tte d m th e r than -Iplc-Ily
a e n le tte d a s m e ap p aren tly found In the o th e r c ro ss stu d ied .
workers when studying awnless x nwned crosses have found the
apical I y aw nletted.
M l o th er
to be
Since the F jl was s p lc a lly a e n le tte d a d if f e r e n t
phenotypic r a t i o o f I t 4s 513*211 o f awnlese, apic l l y a e n le tte d , weakly
a e n le tte d , stro n g ly a e n le tte d , h a lf aimed to aimed, was obtained,
Horn
ev er. In the second c ro ss a phenoty- lo r e t i e o f I*SiI i J i a i l was obtained
f o r the s ix c la s s e s , re sp e c tiv e ly ,
th e r e s u lts cm the various cro sses involving the p a re n ts and progeny
o f ttie RelianeWWward x Comb-(Husear-Hard Federation) c ro ss confirm the
awn-group c la s s lf lc r tlo n with th e excej^tlon o f Uie cro sses involving tha
tru e-b reed in g segregate belonging to Uio asm group > .
Since the > and >
to’pee were thoa##it to re p re se n t d if f e r e n t interm ediate types, each g e n e tic a lly
o n e -fa c to r removed from e it h e r p a re n t, the R eliance x Reward lacking both
dominant fa c to rs and Coast x Rhseai^-Rard Fedem tion peseeselag both dominant
f a c to r s , i t wus thought by c ro ssin g these two interm ediate types the fa c to rs
would be re-assem bled and re s to re the o rig in a l p a re n ts.
In U iis c re ss Ute
grand p a re n ta l types wiH
pe a c t re s to re d , bu t only interm ediate progenies were
ob tain ed .
Alee when the J s true-breeding segrepnte was bnokcrossod with the
R eliance x Rmvrrd p a re n t (as* type 5) , Uie F1 was n e ith e r s > nor a 4 asm
type but ap reached mere n e arly th e J e earn type; n 4 mm type was expected.
f*o» these ZTititilts I t appears th a t the > progeny p la n t used I a e re e sla g
d id n e t possess the f a c to r ia l composition aaRB as sms expected I f the
3- eegreentee wgr he in d ic ate d tgr th e f a c to r ia l fo rs e la AAhh. since
tid e > awn type x 5 awn typ* eroee set-related very O ia ila r to the 3»
nsa type x 5 ana typo, an* the grand p a re n ta l types w r e n e t re sto re d from
the 3" * > awn types p la n ts , i t I s believed th a t the d iffe re n c e Ia mm
expression between these Interm ediates used I a th is cro ss i s due to a
sin g le n in o r f a c to r o r fa c to rs .
Although the model c la ss e s were o f the
3» typo l a both o f the backcroeees involving the 3»
nd 30 to the 5 earn
type p a re n t, th ere was a g re a te r e m b e r o f p la n ts o f th e 5 awn type in
the > c re ss then i s normally expected in a 1: 2*1 m tl o .
Since the p aren ts In th e Reliance-Reward x ( Rneacr-CarA Federation)
d iffe re d in I n te n s ity o f redness o f seed c o lo r. I t woe p o ssib le to study
the In h eritan ce o f th is c h a ra c te r In the 100
lin e s .
The seed o f the
Reliance x Reward p a re n t was m eh d ark er than those o f th e o th e r p aren t.
Three breeding groups aproarsd among the progeny, namely:
(I) Iln se
breeding I llm the dark red p a re n t, (2 ) U jw s breeding lik e the l ig h t red
p a re n t, and ( 3 ) lin e s seg reg atin g f o r l i # » t , lntorm odlate, end dtork-reAkem oled p lan t# In a 1:2 :1 m tl o .
Of the one hundred l in e s , tw enty-six
bred lik e the dastc p a re n t, tw enty-eight bred tru e f o r ligh t-reA -k em eled
p la n ts , and the rem aining, o r f o r ty - s ix , lin e s segregated f o r kernel c o lo r.
Cf the se g reg a tin g Iin e e th ere were 357 dark red: SU6 Interm ediate red*
415 Iip h t-rwd-l'om,tiled p la n ts obtained ae com afed to the expected o f
tKlUe dark red: 609 ln te m o d ia te red* U0U.5 H ^it-red-kem eX od p la n ts .
« 6>
The d a ta o b t lnod on the
Ila e e Ia d lc a te a o n e -fa c to r d ifferen ce
f o r la te n e lty o f red c o lo r boteeen R ellaaee z Re* rd ead Comet z JSaaaa**
Jtatd Federation*
From these d ata along w ith the v e rifie r tle a o f seed c o lo r
o f the Fji p le a t i t has been concluded th a t th ere Ie ire^orfeet dominance o f
redness*
Both p a re n ts possess fa c to rs fo r red kernel c o lo r, the lig h t
colored p a re n t possessing a t l e a s t one f a c to r and the drurte-red-ketneled
n ren t o eseseing a t Ie s t Wo.
Frtw th e re su lW obtained In t h is study i t
can be concluded th a t f a c to r ia l d iffe re n c e s l a In te n s ity o f c o lo r con be
observed.
This work furnishes; fu rth e r p ro o f o r evidence f o r th e explanation
o f kesiMd c o lo r Igr Sileeon-JShlet S theory.
The r e s u lts on glume pubescence su b s ta n tia te the fa c ts Jmawn w ^ ard Ing th e In h eritan ce o f
abeeceat and, ^Labrous glumes.
Titege Wo char c te rs
c o n s titu te an a lle lo n o rp h tc p a ir o f c h a ra c te rs , the pubescent condition be­
in g dominant over the plabroua ,fdtuae and in the Fg OOgrer' a te in to a 3*1
ra tio .
This Fg breeding behavior wan e sta b lish e d by the F^ s tu d ie s .
The
Wj d ata rev e al the preeenee o f about equal tru e breeding and se g re ra tln g
et r a i n s .
Qa th e b a s is o f a s in g le -fa c to r d iffe re n c e f o r glume -axbesconce,
25 a re eapocted to Weed tru e fo r glume pubescence, 50 to segregate 3*1,
nd
5 to brr-ed tru e f o r glabrous glumes.
Ca th is c a lc u la te d b a sis the P
value WHS determ ined to be beWesn 9.90 s a l 0, f ’o which In d ic a te s s good f i t
to the o n e -fa c to r 1*2*1 r a t i o .
Bute a re presented f o r 100 F j progenies o f Rellsmee-Remrd % Comet—
( Jbeurser-Kord Federation) th a t have demonstrated various degrees o f sus­
c e p t i b i l i t y to bunt when Inoculated w ith & composite o f physiologic races
T-8 and L-?.
-61*Slnoe th is e p e r lm n t was n e t e e t up ae an In h e rit'n e e study, the
r e s u lts e t t Ined Can not be In te rp re te d In th a t I W it.
A oowioalte o f the
two races Wfte used la s te d o f determ ining the re a c tio n o f each race In d l r l d u a lly to the progeny.
I t has been shown th a t the use o f o o n m sltes
o f a number o f rtc e e o f T ll l e t l a t r i t l c l and J
il
le v la 1« n o t a re lia b le
Riethod o f determ ining the ab so lu te bunt re s is ta n c e o f v a r ie tie s .
Tarlcms
fa c to rs e n te r which probably accounts f o r the e o e tile rlty o f the in h eritan ce
o f bunt by e a rly In v e s tig a to rs .
Here recen t wo** has shown the In heritance
o f burnt to be q u ite sim ple, probably due to the use o f sin g le races fo r the
inoculum r a th e r t h n a com os It©.
The range o f bunt in fe c tio n to the con o s l t e o f the two r coo f o r
the R eliance x Reward p a re n t wee from I . 2 p e r cent to 2 3 .9 p e r cent with
sn average o f 16.2 p e r c e n t, although i t w e highly r e s i s t n t to T-g and
su sce p tib le to race W
uhea te s te d In d iv id u a lly .
The o th e r p arent show
od mere re s is ta n c e to the composite and a much narrower range o f sus­
c e p t i b i l i t y , the range being fro* 2.6 to 5.1 p e r coat in fe c tio n w ith an
average o f UO p e r c e n t.
Thie p a re n t. Comet x fleeear-Rard F ederation, i s
r e s is ta n t to I#-7 but su sc e p tib le to T-8,
The range o f the Fj progeny e*-
tonded beyond the l l s d t s o f e ith e r the r e e ie t^ n t o r eumce t l b l e
re n ts.
Fine o f the progeny aweraged lower In fe c tio n than the lower l im i t o f the
r e s i s t a n t p a re n t, the most r e s is ta n t hybrid lin e averaging 0 .8 p e r cent
bunt.
There were ten lin e s more su sce p tib le than the su sc e p tib le p a re n t.
F ifty -fo u r o f the hybrid s tr a in s produced an Interm ediate re a c tio n to the
composite bunt, w hile 12 bred Ilk* the r e s i s t a n t parent and 15 bred lib *
the su sc e p tib le p a re n t.
Xt hne been denonatra te d ty various workers th a t r e s i s t nee o r
snscat tlb llltg r to se n t o f wheats eon be tra n s fe rre d to o th e r v a r ie tie s .
Good agronomic q u a litie s and smut r e s lsttuuso can be combined In to a
sin g le v a rie ty tgr means o f h y b rid !ac tio n .
In th is c ro ss both ] irren te
reacted d if f e r e n tly to the too races o f smut, each v a rie ty being r e s is ta n t
to d if f e r e n t ra c e s.
Since lin e s wore obtained In the F-j which segregated
beyond the lim its o f the r e s is ta n t p a re n t, i t was concluded these s tra in s
possessed the fa c to rs fo r r e s i s t ace to the two races o f smut.
Klas o f
these lin e s averaged le e s bunt in fe c tio n than the r e s is ta n t p a re n t.
A
number o f head se le c tio n s were node from those r e s is ta n t lin e s and grown
In th e Ffci to determ ine whether the r e s i s t a n t lin e s were tru e breeding o r
whether they were escapes,
being b u n t-fre e,
th ese lin e s demonstrated t h e ir Immunity by
thus the near-immune re c tlo n o f each p a re n t to the two
races In d iv id u a lly h ss been tra n sfe rre d and combined Into n sin g le lin e
which wne the primary o b je c t o f making the c ro ss.
In o rd e r to determine whether a re la tio n s h ip e x is ts between the
v ario u s morphological c h arac ters and re s is ta n c e to stin k in g smut, s s e rie s
o f contingency ta b le s was prepared shewing a comparison o f th e r e c t i on to
smut in fe c tio n and morphological c h a ra c te rs.
A measure o f the re la tio n sh ip
between the d is tr ib u tio n o f the v arious ch arac ters and bunt re a c tio n taken
two a t » time woe obtained by c a lc u la tin g x2 and determ ining the value o f
F from F is h e r 's (19)6) ta b le s .
In each c ase.
Table XII gives the
robm b ill tie # obtained
In I n t e r r a tin g the r e s u lts i t was considered sa fe to assume
th a t I f the value o f F f o r any ^lven d is tr ib u tio n i s hig h er than
.0 5 , th ere
i s no evtdm ce o f s ig n ific a n t c o rre la tio n between the c h a ra c te rs being con-
eld ered ; hamsver. I f the Talno obtained i s below to la p o in t, re la tio n sh ip
between the c h arac ters compared l a e x h ib ited .
< W one P TBine in Table X ll i s below 0, 05.
n e l c o lo r and se n t s u s c e p tib ility .
R d s i s between ker­
The remaining comparisons showed no
el@ alflean t c o rre la tio n s , slM e the lowest P value was above 0. 3.
R esults
obtained between asnedneee and bunt coffljsare favorably w ith Ausemss ( 193U)
who found awnednesa and bunt independently a sso rte d .
Kam el c o lo r and
awnednees were lndepetoleatly ae-sorted both in these esperiaiM te and those
conducted by C larh and Hooker (1926) .
A OtNamrleen o f d a ta by Schlehaber ( 1935) in d icated a p ossible
weak linkage between smut re a ls t-nce and eeed c o lo r in am A lb it x Mlnhordi
c ro ss.
A lb it1 w hite seeded, i s the r e s i s t a n t p aren t and y e t, o f the 20
pure red~seoded
progenies, 10 were found to be smut fre e .
Xe s n a c ^ tib le
pure white progenies were discovered, and only fo u r pure w hites were o b ta in ­
ed.
Be concluded th is number made I t im passible to draw any s a tis f a c to r y
conclusions concerning t h is p a r tic u la r group.
The nature o f the d i s t r i ­
bution o f the fo u r breeding groups f o r c o lo r in r e la tio n to the three smut
groups in d icated a p o ssib le linkage between smut re sista n c e and seed co lo r.
The r e s u l ts o f i:chlehuber a re o f in te rn e t in v ie
o f the f a c t th a t the r e ­
s u l t s obtained in th is study v e rify M e supposition th a t the dark colored
lin e s o r v a r ie tie s a re mors su sce p tib le to s tin k in g smut than a re the
H r h t colored v a r i e ti e s .
Apparently a re la tio n sh ip e x is ts betw*M# these
two e h em ctere from the standpoint o f in h erita n c e ra th e r th m a m atter
o f cH-vmn,
sotwtior
1.
S e a e tlc s te a l re ©a the Ia h e rtta a o e e f avnedawie, kernel c o lo r,
»ad gimme mbeeeenee, aad re a ctio n to two physiologic rscee o f T ll l n ti r
a re re v e rte d on th e 100 F-j l i a r s o f Rellance-Reeard * Com t» ( itassai^-Hard
F ed eratio n ).
2.
Aenedneee se e stu d ied mad W jitF tei on 100 F j hybrid lin e s o f
n second sp rin g whent c ro s s . Const-(Russrr-Hnn$ Federation) x Cores-{RopeFloremoe) .
3« In both cro sses the in h e rita n c e o f neaedneee wae ste d le d ty
g ro ssin g the m ate ria l In to ttm s ix follow ing elms no*
( I ) aim less,
( ) a p ic a lIy aw aletted , (3) weekly aw nletted. (%) e tr o a ^ y aw aletted,
( 5) h a lf owned, and (6) amsed.
H.
TJm expression o f amedneea was shown to be r- th e r v a ria b le
in th e W llnnoo- Reeard * Cnmst - (Htteesr-Rard Federation) caress, but by
grwti^lng d if f e r e n t segregating types a good f i t was shown f o r s ix pheno­
ty p ic c la ss e s aad fo r nine genotypic gwmps.
Two major fa c to rs a re c le a r ­
ly shown, w ith a I t 1H S t j i 1?* I phenotypic r a t i o o f aim less, anlc a lly aw nletted,
weakly aw nletted, stro n g ly aw nletted, h a lf owned, re s p e c tiv e ly , and a gma»tyi ic to tlo o f I: :2A* I: Ii ; : I baaed u on th e segrega tin g grow #.
The
*3 I ' nedness c la s s if ic a tio n was baaed to on a sin g le head from 0 eh p la n t.
5.
Tartoos cro sses were made In v o lrtn g the paren ts and progeny
o f the cro ss Reliance-Reward x Const-(Hnsnor-Rard F ederation).
The I 1 and
*2 F e ro lts from these c ro sses in d ic ate d the awn-grown c la s s if ic a tio n so made
OTa c o rre c t.
6.
In the second sp rin g wheat cro ss the ev reoslnn o f mmodness
w^e baaed u on an e n tir e p la n t and m s found n o t os v a ria b le as the f i r s t
craea.
%r eroapin<g the nine aegrer^tin#; Igrjes a /mod f i t m s shorn fo r
the s ix phenetsr io c la ss e s and fo r the nine genoty Ic rrm n s .
fe e majer
fa c to rs a re c le a rly shown, w ith a I i S t l t 3*3*1 phenotypic r a tio fo r the
s ix Stmedneas c la ss e s sod a penoty i c r a t i o o f I l 2t ?t h, It It 2t 2t I fo r the
segrega tin g g rm ^e.
7«
V ariation occurs w ith in heads on the sane p la n t r e la tiv e to
am ednass ezi rsa sio n ,
I t i s considered advisable to use the whole p la n t
f o r c la s s if ic a tio n r a th e r than a sin g le head to determine th e expression
o f sens upon a p la n t.
S.
A sin g le major fa c to r d iffe re n c e f o r kernel c o lo r m e o btain­
ed between the R elianee-Sem rd x Com6- (Itusr,- r-H- rd fed e ratio n ) cro ss.
Both a re red wheats, the Reliaaee-R em rd being somewhat d ark er than the
e th e r v-orent.
The
progenies segregated in to 26 pure lig h t- r e d seeded, 26
d ark-red seeded, and W segregating I t P t l dark, interm ediate to l ig h t red.
9#
Rnbescwit M d glabrous c h arac ters a re expressions o f an
lle lo n o r h ie p a ir o f fa c to rs .
Md in th e
Pubemeent glumes ore dominant to glabrous,
g eneration the d ev ia tio n s in numbers were n e t s ig n ific a n tly
d if f e r e n t from the s i m l e 1* *1 r a tio in any o f the IO i
fam ilie s sttwlied.
fw enty-foer lin e s bred tru e fo r pubescent glumes; twenty-one bred tru e f o r
glabrous glumes; Md f i f t y - f i v e segregated f o r g la m tubosconce in a 3*1
r a t i o f o r rube scen t to glabrous.
10.
The Fj g en etic stu d ie s o f awns, kernel c o lo r and glume pubescence
in d ic a te te e g e n etic c o n s titu tio n o f the p arent m aterial to be a s fellows*
P ren t#
Reliance-Reward, %.!*. 1388
Airata K#m#l c o lo r Qlmm r%beoc#nc#
nabb
-R R R'R*
PbPh
Comet x HwmtiMtoxd Fedem tlont t,f». 1315 AABB
-R R r ’r '
rbpb
Comet x Sterneax-iiiaxd Pedomtlon, I.H. 1)83 AAHB
Ceree x Ho e-Plnrenoe b e l. 60.4.1$
11.
aabh
The probable e rr o r f o r p e r cen t o f bunt fo r each o f th e
: a ren te was c a lc u la te d , end from these c a lc u la tio n s the fiv e d if f e r e n t
bunt clo se a were determ ined, namely*
0 -3 .2 p e r c e n t, 3»> •7 p e r cen t,
4.8-1 .7 -pmr c e n t, 12.8-19*6 p e r c e n t, and 19.7-35*7 p e r c en t.
3 * 7
The
P«r c e n t c la s s was determined to be lik e the Comet x Rnsear-
Hord P edem tion p a re n t and the 12. 8- 19.6 p e r cent c la s s was I ik s the
Relisnoe x Reward oarent.
12.
Since th is experiment was n o t designed as a p e n e tle study,
i t was im possible to determ ine the muraber o f fa c to rs resp o n sib le fo r
r e s is t- nee to bunt In tills cro ss.
13*
The seg reg atio n o f the hybrids was beyond the lim its o f
e ith e r the re s is ta n c e o r s u s c e p tib ility o f the p a re n ts, showing th a t the
re s is ta n c e to both races T-S and b»7 Ims boon combined Into a sin g le s tr a in
o f wheat.
l4 .
There was no ovi onoe in the stu d ie s erde o f azyr r e la tio n be­
tween swnednesa and glume ^mbeocince, awnednees and kernel c o lo r, awnedneea and boat p e r c e n t, glume pubescence and bunt, and glume pubescence
and k ern el c o lo r, since the r e s u lts obtained ease w ithin expectation fo r
70-
i Bdeperulent assortm ent.
15» * ooi^xtrlson e f date on bunt ro- e tto n and seed c o lo r la d le tee
e p o ssib le weak H a t e r s between re a c tio n to toe coapoelte o f r cea o f f -8
nnd L- 7 and seed c o lo r.
% e P vslna fo r Intlo; endence wae c a lcu la ted to
be betwnen 0.05 %ad 0.02 which Is rep rtled a s n o t being lrtle ondent
assortm ent.
h v m w tim - e n tm
1.
K am m at
V j y i,
R*
Crtrrelt ted lriiierl tsn oe o f r e a c tio n to d leen e e
cor to I a hot? ile a l ch aractors In tr ia n g u la r wh
t cro 8B"8.
r e . bst 31- 57 ,
Jour, Agr.
2.
td o f
B lffer1, R, H.
I 905 ,
Hendelk lame o f ln h e r tte n e e and wheat breeding.
Jour.
AfT, Sol, I*
3.
Breeeema, R.
1931,
.
V e r le te l m e ls tn n c e , p h y sio lo g ic s p e c ia lis a t io n and Inheri­
tance stu d le a in bent o f w heat.
Crop. Apr.
- . Sto ,
"all.
281, Wi n p.
4.
________________ and H a rris, L. g.
1933,
In h eritan ce in d i b i t wheot o f r e s is ta n c e to b o a t, T l l l e t l a
tr ltic i.
5.
Jour. Agr. Res. 46* 351- 365.
arlng., r.
1926.
I n in r ltone# o f r e e ls to ce to hunt, T llto t tn t r l t i c i (BJork.)
In t o r , in wh" - t . Jou r, Agr. >'-«e. JPl975“<) 1 •
6.
_____________
193 .
Inherltnn oe o f th e seoo 4 fa c to r fo r r e s is ta n c e to bu t ,
T l l l o t l f t r l t i c i . in -hiesnr ??! rat.
Jour* Agr. i o s . -Kli
225-232.
7.
_____________
193 0 . In h eritan ce o f r o e ie tf oe to
M e s s ’ '.-eKf-1.
Jou . A T .
u:..t, .T illctin t r l t i c i . in V -Ite
e . ■ *31:
7215.
arif/w,
1931.
. }f.
In h eritan ce
o t
r e a le ta oe In bn t , T l l I e t l t t r l t i c ? . In
Iqrbrlds o f White F ederation and Fmawrr nrkeleqr ndieata.
Jou r. Kf T, Res. ^>*3 7-313*
9* ______________
1932.
In h erlten o e o f re s is ta n c e to bent..
crosses i f
t ie t r l t l c ! in
r i t e F- d e r a tio n and Cd^esa
-four.
Afcr. Pes•
. ________
10
1932s. In h e r ito ee o f r e s is ta n c e fce be; t , T l l l e t t r t r l t i c i . in
c r o s s e s o f d h ite F ederation w ith ^irkey w heats.
J-mr. Kpr,
F a s . UU11 2 1 -1 2 7 .
11.
_____________
1933.
A third genetic factor for resistance to
trltici. Iii wheat hybrids.
12.
burst,
Tllletia
Jmur. C--- -otlcs L1?: -3'.-'-'I.
_____________
193>i.
In b e r ttM ce o f r e s is ta n c e to b a n t, T lU e U fi t r l t i c i . In
Sherm a m 4 tiro wiieat b b r ld s .
O onetlcs I *73-82.
13* _____________
1936 .
In h eritan ce o f r e s is ta n c e to bunt, T lllo t lo t r l t i c . In
hybrids o f hur ay w heats, C. I . I r-LFi and C. I . 257E.
Rllfcfirdla 10*19-25.
7>
14.
C lark. J .
1924.
a.
SagregBtlon and c o rre la te d in h e rita n c e in eroasee betreen
Kota and Hard fe d e ra tio n wheat* fo r r u s t and drought
reaiataR co.
Joxur. A#cr. Roe. 29*1-47.
15. ____________
1936.
Xi^ rorement in wheat.
K r in tW O ffice,
16.
..........
1925.
TearWok o f v r ie u ltu r e .
V.
. Oort.
,--ahW to n t D. C. t > 7-302.
. H o r e l l e Te Hee nnd l ooker, J . Re
Xnherttanee o f aemednese. y ie ld , nnd q u a lity in croeeee
between Robe, Hord Fodemtlo n , and Prepe wheats a t Dmrise
C a lifo rn ia .
17»
U. Se De a . Tech. R oll. 39, W pp.
. and Hooker, J e Re
1926.
Sefdregntion and c o rre la te d in h eritan ce In Mnrqnie and
srd
Fedem tion c re s s e s , witii fa c to rs fo r y ie ld and q u a lity o f
« rin g wheat in Monten .
18.
0. S.
De
a.
R ail. 1 3 , J l pp.
. nnd gaieenberry, K. Se
I 929.
In h eritan ce o f y ie ld and
ro te in content In oroaeee o f
Morouie rod Koto s- rin g whe->te r-ro-m in c a t nae Jo u r. Agr.
Re*. 38*205-217.
19*
.
<t ilsen tierry t
1933*
K,
s ..
nd Po m re , LeHoy.
In h eri t-n ce o f hunt re c tlo n and o th er e h a m e te re in Hope
wheat c ro sse s.
Jo u r. Agr. Roe. '6* -13-5-; 5.
-J k ~
20.
Varrart W.
!!■XU.
R em ilte o f the ’ omhrigg Iwmt expeyii *>nte o f l$o .
0,^3.
21,
. S. Walee, 1 2 i U l ^ 3 0 .
Z lehor, R. A.
193>.
S t o t l e t t c a l method# fo r reeenrtih workers.
c i l w r r Kt
22,
ttalnoe,
1917*
-',yd.
dinbur/dx:
-d. 6 , 339 pp.
P.
Xrtfwrltfinee In whf t , h e r le y ,
E* . Rtf .
a ll. I] , 6l
oofc h y b rid s,
hash. <vr.
.
23. _____________
1920.
th e In Iw rlten ce o f r e s ie tm ie e to bent o r e tln k l g em it of
whent.
Jour. * '.or. S e e , A#gmn. 12t IS--IJ;-?.
2U. ___________
IfI-1";.
25.
V1U c e o f mnt r e e ie te i.c e In crhent, dour. Afr.
- a.
_____________
192d.
Sew jd y r e lo lo flc fo r m o f f i l l e t l o lo v lg end T ll mtle t r l t l c l .
P h y ty n th .
26 .
18*579- 588 .
. end SlnrhIotO:1-, R.
!,
Oermtlee o f Msrqale % Turkey Uhewt la w e p e c t to bant w e let.or.c e , w in ter h a b it, and aw nleeeneeo.
Jour. Arr. Wee.
32:16 -1 8 1 .
27.
H arrington, J . P.
1922.
Ih e aed# o f In h eritan ce o f c e r ta in choree to r s In whor .
s o l . Agr. 2 1 3 1 -3 2 5 .
7>
28#
''e y n a , I. X. and Aann-H, < . 5 .
192 .5.
A sttadr o f n e t r esla ta n o e In a eroe#
Kota ^ e e n tc .
2 '.
NiirtjQie end
-Io-UP Af--T. R##. .flH 97* 1012.
aart 6 river, ' . J .
192k.
U35
50*
.
swd Botortao?!* ’ .
1925 .
See Torlc 1 MeCSraw-Hill Haeflt Co. KU 2 ,
IiMWAtas am p f la n t s .
.
'Wio Inhorii--TiCO o f r ro ln
lo r In vhoat.
4anr. A o r . Soe.
Arrr-'n . lGi 7^7*790.
31 .
. « . ? . iwx" JToald* 1 .
193b.
S tu d io s OB th e e o n tr o l and e t h e r r s -met* o f \mn% o f wh#r?t ,
■s'oeiw att.
32.
Howard* A. and Hooaad*
1912.
z .
t&*
u l l . 33 -, 93 p .
-. >. C.
Ca the ln h er ita n e e o f Bonn chiaraetere in W w at.
Af -". '>•'.*
33«
.
-H. S e r ie s 5*1“ 7»
_____ ___________________________
I Vlr .
Ca lhm W am rltoaee o f eo-w d w im eto rs in #h# t .
A rr. UtfT. R o t. S -r ie s Jt 27
u.
L e lfh ly , 0 . F. end ' Y v; r ,
19? .
Rfcmtlea la
Uor ,
.
U and CkbI t ,
-. - .
Hur -I h y t r ia is a t le n o f
■• % -Iot .. Arr* hes*
. JU
-■>» The geaw tlea o f Sezaorst wheal.
-201307.
I I ia Sent.
.
35*865.
33 .
Indtn Bent,
-Iear. Aeer.
—76 ™
36.
-Ileenn-Kble1
1911.
KwommgmmIertMhwni,■» am h a fe r «ad weleee, I I .
: ;i l v . A resiHP. a. >". AtiU 2,
37-
UJ» So. 6 , S2
Igatdrn
•; .
_______________
1920.
‘Hilti la a llo lo nor he and tow lexmutet i 0 nea to la weieen.
Hereditas 1*277-311.
38.
rcie% l, John.
I 21.
39.
Heed, C,.
I v. .
The sto a t
Iant$ A eenorra h .
Lemlen, %) p - .
.
V arietal eusce tl I I lty o f wb t to T ill U a lewis Kuehn.
Phytorath. Ik* 437-2^0.
40.
___________
1928.
Phyeinleglc races o f ;. m t in sto a t.
An r. Jour. Bn
15*157-17 .
41.
i Odej^iIenr, H. 4 . and inlto =,, C. S.
1937.
h rgiolorlc races o f V llIetln t r l t l c l nnd T. I ? is . Jour.
Agr. Rne. 55*
42.
,
1927.
and Btsbman1 $. C.
Physiologic sn ec la lisa tio a in T illo tla le v le
tr ltlc l.
43.
Qule nberry,
I9 3 3 .
r.
B.
Phyt-v nth.
T illn tla
%
and UlarP1 J. A.
Inheritance o f awn develo merit in Sm ore shoot crosses.
Jour. Amr*
..
Af i T - - .
*483-492#
Th
^ •»
Salmn ,
19)2.
. Jtrni
-<i##
,
Tweftiy y ea rs o f te s tin g v e r le ile e and strm lne e f w inter whewfe
’:>t the Ka -sss J r r l e a l t a r e
‘a, erlmimt S t a t l- -u
x . S t s . Tech. lM li. j . 7)
**5.
Kar,". a#gr.
.
Saadwee ti. Ke
19 7,
The In heritance o f awns In whe-t .
<Fe, t . ) In te rr IU Co if.
Senetlce ( 19(6 ) 3*371^372.
46.
BehIehnh er. A, Me
1935«
Whmt ln h e rl t-neei
Reaction t i frmr bant bloty* <?#, SnIt#
d e n sity , M d Med c o lo r .
Wash. Agr. R^re S ts . Tech. Bel .
323. 32 PP«
47« ________________
1938«
The In h eritan ce o f re a c t to* to physiologic reces o f Tl I lo tl^
t r l t l c l ( Bje r k .) SShte in a w inter sh e e t c ro s s,
Resmrch
s tu d ie s . S ta te Coils## o f Sash. 6}75- 96.
48.
Sm ith, R. M.
193 .
T ran sferrin g s «6 Inmn I ty to h^rd rsd # r l n f wheat,
^enr.
Amor. So.;. Ajegron. 2lH*%3«
49.
S tew art, Oeorpe
1928.
C orrelated Inh eritan ce In Kanred * S e v l'r v a r ie tie s of H iest.
Jo u r. Apr. Bee, 36*873*896.
50. _______________
1932.
Fe-aesem Iin p the fa c to rs fo r asm# and fo r s p ite d nel ty In
S e v ie r x IV d em tlon who t cross**# end hoc c r o s s e s .
S rc. Apro . 24*543-861.
Jour. Aner.
51*
St
rt,
I
V fend
'11 / ,
A ccxim latlw tr? rreeaVf' a g r g allon Ia wheat.
See. Afren.
52.
.
24:7 »3-76b.
_________________ and Itoywrrode D.
19^).
.
C orrelated Ia h e ritanae In a Wltet?.t eroae between Federation
and a hybrid o f E e rie r x Blc' low.
53.
J e a r. Ah-tt.
Jox$r.
a^r .
Res. l i t 367- '92.
___ ___________ and Jxadd. % I.
I 931 .
In lw ritan ee o f eene In Keta x Rard Tad e ra tton c ro ss.
Ar- r . See. Afltoae
23*
Jonr.
V 1^H.
and Ttngey, I). C.
19 ?? .
TmnfTftSeivw
id
rs -1 R--^r-Ratien In a c ra ss o f
Federation whee ts .
5 .
________________ and ' -iodward,
1931.
Tlnfey, I. U. and Tmlnm .
1934.
Soae
A g ro
u
20; 6 ? t-bj*1*
. %.
Jtror. Agr. %**. ^2; Sr'!"*'? •
im .
In h eritan ce o f re s ista n c e to loose Snut in c e rta in wheat
cro sse s.
57 .
A rm r.
In h eritan ea in a wheat c ro ss between Hybrid 128 x White
Cdeesa and Ka; red.
n6.
Jour.
arq-ii# x
Jo u r. Agr. Kea. - : 631- 6 '/ .
hnnrick. B. L.
1932 .
T m h s b ility ta b le s f o r
endelian r a tio s w ith s n a il numbers.
Texsui Agr. -T •. S t >. d u ll. 463, 28 pp.
MONTANA STATE UNIVERSITY LIBRARIES
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