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Transcript
Genetic characterizations of three male-steriles in wheat, Triticum aestivum L.
by Duane Lee Johnson
A thesis submitted in partial fulfillment of the requirements for the degree of DOCTOR OF
PHILOSOPHY in Crop and Soil Science
Montana State University
© Copyright by Duane Lee Johnson (1978)
Abstract:
Male-sterility provides a quick and easy way to formulate genetic recombination in wheat. The
inheritance and chromosome involvement of two spontaneous male-sterile mutants in 'Siete Cerros'
spring wheat and a single gene male-sterile in 'Chancellor' winter wheat were studied.
Chi square analyses of homogeneous F2, F3, F4, and F5 families were made for various expected ratios
for the three male-sterile sources using spring and winter grown populations. Selections made from the
41 original sibcrossed families of the Siete Cerros mutants were evaluated as spring grown F3, F4 and
F5 headrows. F2 and F3 plantrows of Chancellor male-sterile x various winter wheats were evaluated
as winter wheats.
An abnormal 7:1 segregation predominated in most Siete Cerros families with an unexpected high
number of nonsegregating headrows.
One family of the original 41 segregated for male-sterility as a single recessive allele.
The influence of background genotype on the Chancellor male sterility was attributed to a superior
gene(s) in the parental cultivars.
Monosomic analyses of the three male sterile sources suggested complex inheritance. The male-sterile
expression in Siete Cerros may be due to aneuploidy or gamete lethality. The major factor controlling
male-sterility in Chancellor confirmed the results of Driscoll (10) as being associated with chromosome
4A. Five additional chromosomes may also be involved. i
DEDICATION
This thesis is dedicated to the memory of C. A. Suneson,
pioneer of genetic male sterility in small grains.
GENETIC CHARACTERIZATIONS OF THREE MALE-STERILES
IN WHEAT, Triticum aestivum L.
by
DUANE LEE JOHNSON
A thesis submitted in partial fulfillment
of the requirements for the degree
of
DOCTOR OF PHILOSOPHY
in
Crop and Soil Science
Approved:
MONTANA STATE UNIVERSITY
Bozeman, Montana
June, 1978
iv
ACKNOWLEDGMENTS
The author wishes to extend his deepest gratitude to Dr. G. Allan
.Taylor for his advice, guidance and support in the course of this study
'and the preparation of this manuscript.
The writer also expresses his appreciation to Professor Robert E .
Eslick for his interest and support during the course of this study.
The author also wishes to thank the other members of his examining
committee, Drs. Raymond L. Ditterline, Harry McNeal, and Don E . Mathre,
x.
for their help and tolerance.
Thanks are expressed to Mr* Duane E . Falk.
His ideas and assis- ■
tance were invaluable in the pursuance of this investigation.
The author would also like to thank Mr. G. Hollis Spitler and the
wheat crew for their assistance in the field and greenhouse research,
the excellent typing of Mrs. Homer (Jean) Julian,, and the Montana Wheat
Research and Marketing Committee without whose assistantship the pur­
suit of this degree would have been impossible.
Special gratitude is extended to my wife, Janet, and to my parents
for their patience, encouragement and support in the completion of this
degree.
TABLE OF CONTENTS
Page
DEDICATION
.......................'................... .
V I T A .................................. . . . . ' . ..........
i
iii
ACKNOWLEDGMENT.................... '........... iv
TABLE OF CONTENTS .............'........................
LIST OF TABLES
. ...........................' ........ .. . .
v
vii
LIST OF FIGURES . .'.................... l
. .......... .
ABSTRACT
.........
INTRODUCTION
x
xi
. .............................................
I
LITERATURE REVIEW ..........................................
3
Male S t e r i l i t y ..........................................
History ..............................................
Cytoplasmic Male-Sterility................ .
Genetic Male-Sterility ................................
Genetic Male Sterility in Wheat
................ .. .
Types and Patterns of GMS in W h e a t ....................
3
3
4
5
6.
7
Monosomic A n a l y s i s ...................................... .
Aneuploidy - A.Genetic and Plant Breeding Tool . . . . .
Monosomies - Cytological Problems .....................
9
9
11
Study I.
The Inheritance of Genetic Male-Sterility
in Siete Cerros and Chancellor ........ . . . . .
MATERIALS AND METHODS . . .............^......... ' ........ ..
Genetic Materials..........................
Siete Cerros .....................* .............
.Chancellor............
;■ Cytologic and Genetic P r o c e d u r e s .........
13
14
14
14
14
15
Greenhouse Procedures............................ . . . .. .
17.
Field P r o c e d u r e s ....................
18.
vi
Page
RESULTS AND DISCUSSION.
. . . . . .
The Siete Cerros Male-Steriles
........................
................
21
Chancellor Male-Sterile ..................................
33
Allelism Tests Among Selected Families of the
■.Three Male-Steriles . ............................ .. . . .
38
Study II.
......
21
Monosomic Analyses..........................
42
MATERIALS AND M E T H O D S ..................
Genetic Stocks
43
..........................................
43
Field and Greenhouse Procedures..........................
43
RESULTS AND DISCUSSION
....................................
44
Siete C e r r o s ..............
44
C h a n c e l l o r ..........................
46
GENERAL DISCUSSION
........................................
48
The Inheritance of Genetic Male-Sterility in
Siete Cerros and Chancellor............ ............. .. .
Siete Cerros ................
Chancellor . . . . . . ....................
REFERENCES
..........................
APPENDIX
............................ ......... ' . . . . .
V
48
48
49
51
■
59
vii
■ LIST OF TABLES
Table
I-
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
.
Page
Allelism tests between meiotically normal
Siete Cerros A and B .......................... .
21
Siete Cerros A
family fertile and male-sterile
segregation and associated probability values for
genetic ratios ...............
23
Siete Cerros B F^- family:fertile and male-sterile
segregation and associated probability values for
three genetic ratios . . . ...........................
24
Siete Cerros A F^ family:fertile and male-sterile
segregation and associated probability values for
four genetic r a t i o s ............................
25
Siete Cerros B F^ family:fertile and male-sterile
segregation and associated probability values for
four genetic r a t i o s ................................
26
Number of Siete Cerros A F^ headrows best fitting
five genetic r a t i o s ..................
27
Number of Siete Cerros B F^ headrows best fitting
five genetic ratios . . . . ; ................
28
Siete Cerros A F^ family:fertile and male-sterile
segregation and associated probability values for
four genetic r a t i o s ................................
29
Siete Cerros B F^ family fertile and male-sterile
segregation and associated probability values for
four genetic r a t i o s ............................
30
Field grown F^ headrow distributions within families
of Siete Cerros A for four genetic ratios of fertile:
malersterile . .......................................
31
Field grown F^ headrow distributions within families
of Siete Cerros B for four genetic ratios of fertile:
male-sterile................................
32
viii
Table
.
Page
headrow distributions of greenhouse-grown Siete
Cerros A and B families for four genetic ratios of
fertile:male-sterile ..............................
34
F,_ headrow distribution of greenhouse-grown Siete
Cerros A and B families for various genetic ratios
of fertile:male-sterile . . ......................
35
Siete Cerros spring wheat ms A and ms B x HRWW
grown as F^ spring and winter wheats
. . ........
36
Probability of fit for ms C/Centurk F progeny
using chi square tests of heterogeneity and
goodness of f i t .................. ...............
37
Chi square test for heterogeneity for F populations
of ms C x HRWW and ms C x HRSW ....................
38
Probability, values for six genetic ratios in ms C/
winter wheat F families using chi square tests of
goodness of fit ................................
39
Probability values for six genetic ratios in ms C/
spring wheat F 's.using chi square tests of goodness
of fit . . . .......................................
40
Allelism tests between Siete Cerros male-sterile
A-1, A-3, B-20 and Chancellor male-sterile .' . .
41
20. Chromosome location of Siete Cerros ms A-I . . .
45
12 .
13.
14.
15.
16.
17.
18.
19.
21. Chromosome.location pf Siete Cerros male-sterile
A-3
.............................. ..
45
22. Chromosome location of Siete Cerros ms B-20
.46
23.
47
Chromosome location of ms C
ix
Appendix Tables
1.
2.
3.
4.
5.
Page
Allelism tests of Chancellor male-sterile and
other male-sterile sources......................
60
Fertile and male-sterile progeny of heterozygous
Siete Cerros ms A-I x monosomies ........ . . . . . .
61
Fertile and male-sterile progeny of heterozygous
Siete CerrOs ms x Rescue monosomies . . . . . . . . .
61
Fertile and male-sterile progeny of heterozygous
Siete Cerros ms B-20 x m o n o s o m i e s ........ '.........
62
Fertile and male-sterile progeny of heterozygous
Chancellor male-sterile x monosomies ...................
63
X
LIST OF FIGURES
Figure
I.
Page
Generation procedures for Siete Cerros A and B
. . . .
16
xi
ABSTRACT
MaIe-sterility provides a quick and easy way to formulate genetic
recombination in wheat. The inheritance and chromosome involvement of
two spontaneous male-sterile mutants in 'Siete Cerros * spring wheat and
a single gene male-sterile in 'Chancellor' winter wheat were studied.
Chi square analyses of homogeneous F^, F3, F4 , and F families
were made for various expected ratios for the three male-sterile
sources using spring and winter grown populations. Selections made
from the 41 original sibcrossed families of the Siete Cerros mutants
were evaluated as spring grown F 3, F4 and F^ headrows. F3 and F 3
plantrows of Chancellor male-sterile x various winter wheats were
evaluated as winter wheats.
An abnormal 7:1 segregation predominated in most Siete Cerros
families with an unexpected high number of nonsegregating headrows.
One family of the original 41 segregated for male-sterility as a single
recessive allele.
The influence of background genotype on the Chancellor male ste­
rility was attributed to a superior gene (s) in the parental cultivars.
Monosomic analyses of the three male sterile sources suggested
complex inheritance. The male-sterile expression in Siete Cerros may
be due to aneuploidy or gamete lethality. The major factor controlling
male-sterility in Chancellor confirmed the results of Driscoll (10) as
being associated with chromosome 4A. Five additional chromosomes may
also be involved.
INTRODUCTION
The goal of plant breeding for yield has not changed over time but
knowledge and control of the environment have changed the methodology
used to attain that goal.
Many present day plant breeding programs
emphasize developing high yielding cultivars adapted to a specific en­
vironment with high quality standards and insect and disease resistance
that must be met.
As a result of these requirements, the technology
for determining these factors is improving but the breeder is still
manipulating the same pollination systems used by his prehistoric
ancestors.
Using genetics, modern man is learning to circumvent the limita­
tions of natural pollination mechanisms. Corn (Zea mays), normally a
cross-pollinated crop species, can now be treated as a self-pollinated
species.
Barley (Hordeum vulgare), a ■self-pollinated species, can
likewise be manipulated as a cross-pollinated crop.
Genetic male-sterility in a self-pollinating species provides the
plant breeder with a combination of factors which reduce the costs and
increases the number of potential crosses substantially.
The major
advantage of genetic male-sterility in a self-pollinated crop, however,
in the ability to shift from self-pollination to cross-pollination and
back to self-pollination at the desire of the researcher.
The occurrence of two spontaneous male sterile mutants in the
spring wheat (Tricitum aestivum L.) cultivar 'Siete Cerros1 led to the
initiation of this study.
A winter wheat cultivar, 'Chancellor', which
2
contained a simply-inherited genetic male-sterile was included as a
standard by which progress within the selection of Siete Cerros could
be measured.
The objectives of this research were to:
I.
Examine the inheritance of male-sterility within two spontan­
eous male sterile mutants of Siete Cerros spring wheat and a single
gene male-sterile isolate of Chancellor winter wheat.
2.
Locate as to chromosome the factor(s) causing male sterility.
LITERATURE REVIEW
Male Sterility
■History
Wheat (Triticum aestivum, L.), a self-pollinated species, has very
low levels of cross-pollination.
Natural outcrossing in .wheat varies
between 0.16 and 8% (19,22,24,74) and can be affected by environmental
conditions (3,26,27,66).
The low, natural levels of cross-pollination
are insufficient and too unreliable to be used effectively for genetic
recombination in wheat.
The importance of controlled crossing has been recognized by wheat
breeders since the latter part of the 18th century (46).
The popular- ■
ity of producing hybrid populations for selection in the production of
new cultivars increased and spread from Great Britain to Europe and the
United States.
By the end of the 19th century, wheat breeders through­
out the western world recognized the importance of obtaining new
genetic recombinants for varietal improvement (46).
Since, that time, breeders of self-pollinated crop species have
attempted to accelerate their breeding programs through controlled
genetic recombination.
Most of these efforts have been involved with
rendering the male gametes inviable in selected plants.
This phenom­
enon, termed male-sterility, can be attained by emasculation (common
in self-pollinated crops) or othrough genetically controlled manipu­
lation of factors affecting fertility.
4
Genetically controlled male-sterility is particularly useful in
wheat breeding.
This phenomenon relieves the wheat worker of the tedi­
ous task of.hand emasculation and allows opportunity for the manipu­
lation of wheat as a cross-pollinated species.
methods of creating male-sterility in wheat:
There are two common
(I) cytoplasmic male
sterility (CMS), and (2) genetic male-sterility (GMS).
Cytoplasmic Male-Sterility
•During the 1930's, CMS was discovered in corn (12).
The use of
CMS in the development of corn hybrids resulted in. improved corn yields
at a reduced cost and caused breeders to examine the potential of male
sterility in other crops (13).
CMS in rye (Secale montanum Guss.)(41), barley. (56) and wheat
(32,49,82,83) involve the use of alien cytoplasms.
In rye, only one of
the parent cytoplasms was effective in evoking male-sterility (40).
Currently, wheat hybrids derive their cytoplasmic male sterility
from Tricitum timopheevi, first reported by Wilson and Ross (84).
In
wheat, Aegilopis caudata cytoplasm commonly gave a high incidence of
pistilloidy and .consequently partial or complete female sterility (6.,
49).
Other cytoplasmic sources, such as Aegilops ovata, tended to pro­
duce late flowering female wheat lines with decreased.seed set in
crosses with hexaploid wheat pollen parents (49).
CMS has been developed in hexaploid wheat specifically for
hybrid wheat production.
CMS is therefore limited in its poential
5
uses.
The problems of uneconomical yield increases (21), incomplete
restoration and the necessity of multigenic restorers at northern
latitudes (83) limit the use of cytoplasmic male-sterility in Montana.
Genetic Male-Sterility
GMS offers a flexibility unattainable in the CMS system (29,30).
GMS-derived hybrids can potentially be used as commercial
hybrids,
in recurrent selection populations and for conventional varietal
improvement (16., 29,30,70,77) .
GMS also offers wheat breeders the
ability to shift easily from a program of conventional breeding to
recurrent selection to hybrids (30).
The first isolation of GMS in small grains was made by C. A.
Suneson (73).
In 1936, Suneson discovered a single male-sterile
barley plant which phenotypically resembled some male-sterile wheat he
had previously studied.
In the earlier study, freezing temperatures
had caused complete male-sterility in some wheat tillers (66).
How­
ever, the male-sterility observed in the barley plots proved to be a
simply-inherited single gene male-sterile (68)
In 1945, Suneson (67) suggested the use of a simply-inherited,
recessive, male-sterile in composite cross populations of barley to
allow continued recombination coincident with plant competition favor­
ing the most vigorous plants.
In 1951 (69)> he suggested the use of
GMS in a program of male-sterile facilitated synthetic hybrid barley.
Suneson later termed this male-sterile accelerated recurrent
6
recombination (72).
Eslick suggested a similar program termed male-
sterile facilitated recurrent selection (14).
With the discovery of GMS in barley, an interest grew in the
possibility of producing commercial F
hybrids.
Several systems have
been proposed involving chemically-induced lethals (80), aneuploidy
(51,53)., balanced genetic lethals and male-steriles (13,23), and
others.
Since the breeding systems of barley and wheat do not differ
greatly, the potential exists for similar developments in wheat.
Since the identification of the first genetic male-sterile in
barley in 1940 (67,68) twenty-seven additional genes for male sterility
have been reported (22).
Genetic Male Sterility in Wheat
In 1922, Sax (55) reported an increased proportion of sterility
in wheat in conjunction with an increased proportion of univalents
from interspecific crosses.
The observed male and female-sterility
was attributed to aneuploidy.
In 1938, partially sterile F^ plants were noted in the cross
'Pathology'4592' x 'Nebawa1 (65).
The partial male-sterility was
attributed to chromosomal aberrations.
Pugsley and Oram (50) found what appeared to be a genetic malesterile in an F _ family of 1Kenya Farmer1 x 1Javelin 48'.
3
The inher-
itance of this male-sterility was unclear (50,71). ■This male-sterile,
was used in■conjunction with induced male-sterile mutants in the first
7
wheat composite cross population (73,76).
Suneson's investigations of
the inheritance of this particular male-sterile were not conclusive.
Suneson hypothesized the male-sterility, in this instance, could be
due to a single recessive gene which was environmentally sensitive
(71).
Cytological studies by Wanige (79) and Zeven et al. (85) were
also inconclusive concerning this male-sterile mutant.
The chromosome
number varied irregularly (79,82,85) and no apparent relationship
between aneuploidy and the male-sterile character was noted.
From Pugsley and Oram's original male-sterile germplasm, Briggle
(5), through backcrossing, isolated and transferred a single gene for
male-sterility to the cultivar 1Chancellor'.
The gene behaved irradi-
cally when placed in different parental combinations.
Other genetic
backgrounds differentially influenced the expression of male-sterility.
•Briggle (5) hypothesized that modifiers from other cultivars or a
gamete competition might be responsible for this difference in reac­
tion.
Types and Patterns of GMS in Wheat
Two types of GMS appear to exist in wheat.
The first involves
multiple genes which act in a cumulative fashion to express malesterility (1,18,27).
The multigenic male-steriles have common characteristics.
All
are reported to involve three recessive genes having cumulative effects
and exhibit partial sterility (1,19,28).
An ability to set 5% selfed
8
seed has been advocated for the maintenance of male-sterile lines in
hybrid wheat production (1,19,28).
■ The second type of GMS involves single gene inheritance such as
the spontaneous male-sterile mutant found by Kruysnov (34) in
1Saratovskaya-29' in 1964.
Fourteen of twenty-two F^ families from
the original mutant, segregated for the male-sterile character.
Thirteen of the fourteen segregated for a single, recessive malesterile gene.
The fourteenth family segregated for a complimentary ■
two-gene recessive ratio (34).
Plants grown under field conditions
deviated more from the monofactorial ratio than did greenhouse grown
material.
Krupnov (34) proposed that decreased viability of the male-
sterile phenotypes affected the ratio of fertile to sterile plants.
Bozzini and Scarascia-Mugnossa (4) found a spontaneous mutant in
the tetraploid wheat cross 1Yuma1 x 1Capeits1.
The mutant allele
acted as a single recessive gene for male-sterility.
Bingham (2) reported a spontaneous male-sterile mutant in the
hexaploid cultivar 'Maris Widgeon'.
ically normal.
This single gene mutant was meiot-
Pollen transmission of the character was reduced while
female transmission of the gamete appeared to be normal.
This finding
corroborated■other studies involving monogenic male-steriles in wheat
(11,17,34).
Monogenic male-steriles have also been induced.
Fossanti and
Ingold (16) induced a single gene male-sterile in the cultivar 'Probus'
9
using x-ray.
Driscoll
(11 )
induced a similar male-sterile in the
cultivar 1Pitic-62' using gamma radiation.
The GMS isolated from the original material of Pugsley and Oram
and transferred to 1Chancellor1 (5) is allelic with the Probus mutant
and located on chromosome 4A (10).
The 'Cornerstone' mutant gene,
isolated from 'Pitic-62' by Driscoll (11), is also on chromosome 4A.
The Cornerstone source of male-sterility appears to involve a terminal
deletion of the alpha arm of that chromosome (11).
Maan (41) found the alien substitution line involving the trans­
fer of a single chromosome of Aegilops longissima (20" Th aestivum .
+ I" A. longissima) to be completely fertile.
Telosomic and isosomic
analyses showed the single A. longissima chromosome to be homologous
to that of chromosome 4A in wheat.
self-fertile.
The substitution line is completely
All male gametes carrying the A. longissima chromosome
are fertile and functional.
Thus, chromosome 4A in wheat.appears to ■
perform a vital function in fertility expression.
Monosomic Analysis
Aneuploidy - A Genetic and Plant Breeding Tool
Aneuploidy involves an imbalance of chromosome pairs or between
chromosomal homologues.
Initially, aneuploid types such as monosomies,
nullisomics and trisomics were used only for the chromosomal location
of specific characters.
Morris (43).
These characters have been summarized by
More recently, nullisomics and monosomies have provided
10
wheat researchers with the opportunity, to transfer specific chromo­
somes from cultivar to cultivar (54,64,78) and from species to species
(39,41).
Monosomies involve the loss of one member of a homologous pair of
chromosomes.
Instead of the expected 21 pairs of chromosomes (21"),
the monosomic has 20 chromosome bivalents and a single univalent (20".
+ I')(33).
.
Monosomic plants, when self-pollinated, produce disomic (21"),
monosomic (20" +1') and nullisomic (20") progeny.
The proportion of
these progeny depend upon the frequency with which the 20-chromosome
(n-1) gametes function in fertilization and upon the degree of via­
bility of the nullisomic progeny (35).
Approximately 75 percent of the functional female gametes are n1, regardless of the chromosome involved (35,61).
Functional male
gamates are generally of the 21 chromosome type since certation favors
the 21-chromosome pollen (35).
The percentage of functional 20-
chromosome pollen varies from approximately I to 10 percent depending.
upon the chromosome concerned (61).
The failure to transmit specific male gametes may be effective in
plant breeding research.
Recently, in conjunction with genetic male-
sterility, proposals have.been made to use aneuploids for hybrid wheat
and barley production (8,9,51,53).
11
Monosomies - Cytological Problems
Monosomies are meiotically unstable.
The absence of one dose of
chromosome, IB, 4B, 2D or 4D can promote misdivision of the centromere
(31) and other meiotic irregularities (47,57,58).
Chromosomes IB, 4B
and 6B appear to control regular bipolar segregation of homologous
chromosomes and, in the hemizygous state of the monosomic, can produce
elevated numbers of double monosomies (31).
In the monosomic series
of 'Cheyenne' and 'Wichita' wheat, plants which lacked bivalents for
6A, 6B or 6D showed some necrosis in the seedling stage (44).
Appar­
ently, there was no suppression of leaf necrosis when one of these
chromosomes was in the hemizygous state.
of n-1 gametes is common.
Reduced pollen transmission
The failure to effect fertilization by n-1
gametes is remarkably consistent across observed polyploid species
(20> 36,45,61).
The actual number of viable n-1 gametes varies between
monosomic lines within a species (39,61).
More information concerning
the theoretical aneuploid ratios is available from Kuspira and Unrau
(.35).
Univalent shift is another potential problem with the use of
monosomies.
Person (47) found univalent shift occurred frequently.
Univalent shift involves the mispairing of a chromosome with a uni­
valent from one of its homologues.
For example, a univalent of 4A
could occasionally mispair with a univalent of 4B leaving the 4B
homologue to become the new univalent.
Since all wheat chromosomes
12
are essentially metacentric, it is difficult to discriminate when ,
univalent shift has occurred (47).
asynapsis to occur.
Person (41) also found partial
New aneuploid types such as double monosomies
could result.
A final potential problem in using monosomies is non-expression
of a character in the hemizygous state (61).
As a result, evidence
gathered from monosomic analyses may be inconclusive.
Study I
THE INHERITANCE OF GENETIC MALE-STERILITY
IN Siete Cerros AND Chancellor
MATERIALS AND METHODS
Genetic Materials
Siete Cerros
Two spontaneous male-sterile mutants, found.in the spring wheat
cultivar 'Siete Cerros1 (C.I. 14493) by University of Arizona Profes­
sor R. K. Thompson, were studied.
Seeds from open-pollination with
Siete Cerros were assigned the population-designation A and B for
identification.
this study.
These identifying letters were maintained throughout
Each seed was assigned an Arabic number as a family desig­
nation (A—I through A-21 and B-I through B-20).
Chancellor
A single recessive gene for male-sterility was transferred from
the original male-sterile mutant population of Pugsley and Oram, com­
monly called Pugsley1s male-sterile, to the soft, red, winter wheat.
cultivar, 'Chancellor'
(C.I. 12333) by Dr. L. W. Briggle (5).
found no logical segregation pattern in the
Briggle
of the original cross.
Continued selection for a single, recessive male-sterile gene with four
backcrosses to Chancellor was, however, successful.
plants in the F
Partially sterile
and early backcross generations were discarded by
Briggle.
The Chancellor male-sterile (ms C) is cytologically normal (21")
in both fertile and male-sterile phenotypes (5).
Driscoll (10)
15
reported the male-sterile allele from ms C was on chromosome 4A and
allelic to the 1Probus1 male-sterile.
Cytologic and Genetic Procedures
Figure I provides an outline of generation procedures for green­
house and field materials of Siete Cerros A and B populations.
mother cells (PMC) from twenty A and twenty-one B
Pollen
plants, hetero­
zygous for the male-sterile allele(s), were examined cytologically.
PMC analysis followed the techniques of Zeven et al.
(85).
Preliminary allelism tests were made between cytologically normal
F^ families of A and B during the summer of 1975.
Allelism test eval­
uations were based upon primary tiller phenotype.
Additional allelism
test crosses were made among elite F^ families of A, B and C furing the
summer of 1977.
Test cross offspring were evaluated for fertile to
male-sterile segregations in 1978.
Crosses were made between male-sterile individuals from F^ fami­
lies of Siete Cerros A and B with various hard red winter wheats
(HRWW) to examine the influence of genotypic background.
These F^
progeny were fall and spring planted for field observation in 1977.
Crosses were made between male-sterile individuals of Chancellor
(ms C) and the HRWW cultivar 1Centurk1 in the greenhouse during 19741975.
The F^ analysis of the cross.was made during the summer of
1976.
All field grown materials were evaluated for male-sterility at
16
Year
Field
1974-1975
1975
1. crosses ms x
HRWW
2. allelism
crosses (AxB)
3. 20 fertile
heads har­
vested for
headrows
within fami­
lies
1975-1976
1976
1. selection for
3:1 (fertile
heads har­
vested for F jl
4
headrows)
1976-1977
1977
I. selection
for 3:1
segregating
headrows
fertile
heads to be
grown as F 5 F
headrows
5
.F ' fa H
\ 2 planted
F \ spring
2 planted
2. selected
samples for
greenhouse
V
I. allelism
crosses
1977-1978
Figure I.
Generation procedures for Siete Cerros A and B
17
the Plant and Soil Science Field Research Laboratory, Bozeman, Montana.
1Individual F^ progeny (ms C x Centurk) were harvested and classified as
to awnless, awnletted or awned plant types.
F 3 C families from indi­
vidual F2 plants.were planted in the fall of 1976 and classified for
male-sterility in 1977.
Greenhouse Procedures
Materials derived from populations A and B and families of C were
greenhouse grown in steam-pasteurized benches of a sandy-loam soil (2
parts sand:2 parts Amsterdam silt loam:I part peat) and were irrigated
with a complete nutrient solution.
Greenhouse temperatures were main­
tained at 60-75°F for spring and 4O-SO0F for the winter wheat with a
gradual increase to 60-75°F.
The original open-pollinated F^ progeny of Siete Cerros popula­
tions A and B were greenhouse grown from December, 1974 to April, 1975.
Progeny resulting from field sib-crossing within Chancellor (ms C x C)
were also grown at this time.
F
4
and F
5
headrows were greenhouse grown during 1976 and 1977.
Greenhouse grown selected fertile F
4
plants, segregating for single
gene male-sterility, provided seed for F 3 headrows.
18
Field Procedures
The F
families obtained from selfing individual F
plants of
populations A and B were field planted in'April, 1975 to determine F^
segregation ratios for male-sterility.
available, from twenty fertile F
Twenty random spikes, when
plants from each A and B F
family
were harvested to provide seed for F^ families.
3
Greenhouse sub-crossed material from Chancellor (ms C x C) and
F^ progeny of ms C x Centurk were field grown during the summer of
1975.
Male-sterile plants from sib-crosses were crossed with various
hard red winter wheats.
Determination of fertile and male-sterile individuals within seg­
regating families of A, B and C was based upon the phenotypic expres­
sion of the primary tiller and 0% seed set from bagged heads of sus­
pected male-steriles.
F^ A and B family segregations were tested using chi square tests
of goodness of fit to three Mendelian ratios.
The tested ratios were:
1 . ' 3:1, a single gene, recessive male-sterile;
2.
15:1, a two gene, recessive male-sterile;
3.
63:1, a three gene, recessive male sterile.
■
Sample sizes for determining genetic ratios (p = .90) were based
on the formula of Mather (42).
Samples sizes exceeding 146 observa­
tions were considered definitive for all ratios tested at the family
level.
19:
Chi square analyses were used to determine goodness of fit within
Chancellor for the following ratios:
1.
3:1, a single gene, recessive male sterile with complete
gamete transmission of the male-sterile allele;
2.
4:1, a single gene recessive male-sterile resulting from an
assumed 10% reduction in pollen transmission of the malesterile containing gamete;
3.
5:1, a single gene recessive male-sterile resulting from an
assumed 15% reduction in pollen transmission of the malesterile containing gamete;
4.
7:1, a single gene recessive male-sterile resulting from
an assumed 18% reduction in pollen transmission of the malesterile containing gamete;
5.
15:1, a two gene recessive male-sterile ratio with complete
gamete transmission;
6.
13:3, an epistatic, two gene male-sterile ratio with com­
plete gamete transmission.
In 1976, segregation ratios for male-sterility of mx C x Centurk
F21S and spring planted F 3 headrows of A and B were determined.
F3 and
F^ A and B family analyses included the ratios:
I.
7:1, an observed segregation ratio of unknown origin.similar
to one found by Falk (15)
2.
1:0, a nonsegregating
Homogeneity of
(fertile:male-sterile) headrow ratio.
and F 3 families of C and F 3 and F4 families of
A and B were tested using heterogeneity chi square (42).
Headrows with
within each A and B F 3 family were analyzed using chi square tests of
goodness of fit to the expected ratios.
RESULTS AND DISCUSSION
The Siete Cerros Male-Steriles
. Allelic factors producing male-sterility were not evident in
crosses between heterozygous, cytologically normal (2n = 21") indivi­
duals within families of A and B (Table I).
Consequently, the two
populations were maintained separately and considered two mutational
events..
F2 family analyses of A and B were essential to the determination
of inheritance in the Siete Cerros mutants (Tables 2 and 3).
Proba­
bility values provided information concerning the actual segregation
ratio.
Since random pollen parents were used in crosses to the orig­
inal male-sterile mutants, individual families could segregate for
different numbers of genes.
Table I.
Allelism tests between meiotically normal Siete Cerros .A.
and B .
.Parentage
Frequency
Male-Sterile
Fertile
X2
(1:1 expected)
Probability
Value ■
<.001
ms A-l/B-8
• 15
O
15.0
ms A-5/B-8
3
O
3.0
ms A—8/B—8
7
O
7.0
ms A-ll/B-8
3
O
3.0
17
O
17.0
5
O
5.0
ms A-6/B-6
ms B-8/A-15.
.05-.10
<.01
.05-.10
<.001
<.01
22
PMC analysis showed chromosomal observations (telosomy and trans­
locations) and provided a determination of meiotically normal and
aberrant
percents of Siete Cerros A and B (Tables 2 and 3).
Family A-19.(Table 2) does not fit any of the expected ratios.
Family A-2 (Table 2) best fits a three gene model (p = .50 - .70).
Other families of A best fit 3:1 or 15:1 fertile to male-sterile seg­
regation.
Most F2 B families best fit 3:1 segregation ratios (Table 3).
Families B-7, B-17 and B-18 best fit a 15:1 ratio.
Families B-5, B-6>
B-16, B-19 and B-21 did not fit any of the tested ratios.
F
families (Tables 4 and 5) of Siete Cerros A and B did not fit
segregation patterns consistent with observed segregations within
families.
;
There wa.s an unusual, high number of nonsegregating F^ headrows
in A and B (Tables 6 and 7).
The reduced number of segregating classes
may be due to a gametic failure, aneuploidy. or a factor conditioning
gametic lethality such as pollen killer, Ki (62).
F4 families generally fit a 7:1 ratio of fertile:male-sterile•
(Tables 8 and 9).
Falk (15) reported an 8:1 ratio which may'be asso­
ciated with gametic lethality.
A-15 and B-20 were the only families
which fit a desired 3:1 ratio.
F^ and F4 data are similar for a number of non-segregating headrows (Tables 10 and 11).
Greenhouse F4 and F^ headfow
23
Table 2.
Family
A-I§
Siete Cerros A
family fertile and male-sterile
segregation and associated probability values for three
genetic ratios
Frequency
Fertile
Male--Sterile
Probability for tested ratios
3:1
15:1
63:1
55
6
A-2
. 37
I
A-3
41
9
A-4
48
6
A-5§
28
4
A-6§
19
I
A-7
46
12
.30-.50
A-8§
16
2
.10-.20
.30-.50
—
A-9
14
2
.20-.30
.30-.50
—
A-Il
29
11
.70-.80
—
A-12
44
15
>. 99
—
A-13
9'
.20-.30
—
.20-.30
—
.10-.20
—
2
.50-.70
A-14
20
7
.90-.95
A-15§
37
14
.50-.70
A-16
51
2
A-17
60
4
A-18
36
6
A-19
63
10
—
A-20
28
3
—
—
.10-.20
.30-.50
—
—
.50-.70
—
.10-.20
—
.10-.20
—
.80-.90
—
.10-.20
.20-.30
—
—
-—
—
™—
.30-.50
—
.10-.20
>.99
--
—
—
—
.-
.30-.50
t Probability determined by chi square tests of goodness of fit
4= P < .05 are not given
§ Meiotically normal
plants
24
Table 3.
.Siete Cerros B
family:fertile and male-sterile segregation
and associated probability values for three genetic ratios
4*
.
Frequency
Fertile
Male-Sterile
Probability for tested ratios
3:1
15:1
63:1 .
—
B-I
38
10
.50-.70
B-2
57
11
.05-.10
—
—
B-3
23
9
.50-.70
—
—
B-4
50
21
.30-.50
——
—
B-5
71
13
—
--
B—6
47
7
—
—
B-7
36
5
—
W
I
CO
Family
59
14
B-9
57
19
B-IO
8
I
.30-.50
B-Il
3
2
.30-.50
—
—
B-13
44
9
.10-.20
—
--
—
—
.20-.30
>.99
.10-.20
—
.05-.10
'
26
5
.20-.30
B-15
.
47
14
.70-.80
B-16
53
9
—
B-17
25 '
2
—
.80-.90
B-18
44
2
——
.50-.70
B-19
. 58
9
—
B-20
• 44
10
B-21
59
31
—
—
—
B-14
.30-.50
—
——
—
—
—
—
.10-.20
--
—
—
—
—
—
+ Probabilities determined by chi square test of goodness of fit
+ P < .05 are not reported
§ Meidtically normal F plants
25
Table 4..
Family
+
Siete Cerros A F 3 family:fertile and male-sterile segregation
and associated probability values for four genetic ratios
Frequency
Fertile
Male-Sterile
Probability for ratios tested
7:1
3:1
15:1
63:1
—
■
A-I
156
17
A-2
■ 72
15
A-3
112
16
■"—
A-4
130
20
—
A-5
156
21
---
A—6
114
13
—
.30-.50
.05-.10
—
A-7
47
6
—
.80-.90
.10-.20
—
A-8 •
160
25
—
.50-.70
A-9
165
29
—
.10-.20
A-Il
10
2
A-12
. Ill
20
A-13
128
18
—
A-14
95
12
—
.50-.70
A-15
133
12
—
.05-.10
A-16.
81
11
—
.50-.70
A-17
161
18
—
..50-. 70
—
A-18
109
18
—
.50-.70
——
A-19
76
10
-.
.80-.90
—
A-20
173
26
—
.80-.90
.05-.10
.50-.70
—
t Missing families due to hail damage
4 P < .05 are not reported
.20-.30
.10-.20
.05-.10
—
—
—
—
—
.20-.30
—
—
.80-.90
—
—
>.99
.50-.70
.20-.30
>.99
—
—
—
.10-.20
——
—
—
—
—
—
—
.30-.50
——
—
—
.——
—™
26
Table' 5.
Family
•f*
Siete Cerros B
family:fertile and male-sterile segregation •
and associated probability values for four genetic ratios
*
Probability for ratios tested
3:1
7:1
15:1
63:1
Frequency
Fertile
Male-Sterile
.50-.70
—
—
.20-.30
—
—
—
.80-.90
—
—
31
—
.80-.90
—
—
• 137
23
—
.20-.30
—
—
B-6
101
.21
—
.05-.10
—
—
B-7
72
8
-.
.30-.50
B-8
9
6
B-9
50
7
B-IO
48
12
B-Il
118
17
B-13
47
8
B-14
145
25
——
B-15
178
29
—
B-16
36
5
B-17
158
23
—
.90-.98
B-18
66
6
—
.50-.70 ■ .30-.50
B—19
218 .
38
—
.30-.50
—
B-20
173
26
—
.80-.90
—
B-21
166
17
—
.10-.20
B-I
87
12
B-2
76
14
B-3
142
22
B-4
209
B-5
'
—
.10-.20 '
■.30-.50
—
.05-.10
.05-.10
t Missing families due to hail damage
t P < .05 are not reported
-.
.90-.98
.I0— .20
—
.05-.10
—
—
—
.05-.10 ■
—
—
.90-.98
—
—
■.50-.70
—
—
'.30-.50
—
--
.20-.30
—
—
.80-.90
.10-.20
—
.05-.10
"
——
27
Table.6.
Family
Number of Siete Cerros A
genetic ratios
3:1
headro'ws best fitting, five
Number of headrows best fitting ratios
7:1
15:1
63:1
1:0
A-I
2
3
I
0
14
A-2
3
3
I
0
13
A-3
5
2
5
0
8
A-4
3
7
3
0
7
A-5
5
4
2
0
9
A-6 .
2
7
0
0
10
A-7
0
6
I
0
12
A—8
2
0
0
0
3
A-9
I
2
I
0
6
A-Il
3
3
3
0
10
A-12
2
7
3
0
7
A-13
2
0
3
0
5
A-14
4
5
2
0
9
A-15
3
'5
I
0
10
A-16
I
2
0
0
17
A-17
3
4
2
0
9
A-18
2
I
2
0
15
A-19
5
3
5
0
7
A-20
0
6
5
0
8
t Using chi square tests of goodness of fit
•
28
Table 7.
Family
Number of Siete Cerros B
genetic ratios
3:1
headrows best fitting five
Number of headrows best fitting ratios
7:1
15:1
63:1
1:0 .
B-I
3
I
6
0
10
B-2
4
I
I
0
14
B-3
2
4 ■
3
0 ■
11
B-4
0
4
3
0
10
B-5
2
I
4
0
12
B-6
0
5
3
0
12
B-7
■3
0
2
0
15
B-8
4
2
5
0
6
B-9
4
3
4
0
9
B-IO
0
I
0
0
3
B-Il
I
I
0
0
I •
B-13
3
4
2
0
9
B-14
2
3
2
0
13
B-15
I
4
4
0
11
B—16
I
4
2
0
13
B-17
'I
5-
5
0
.9
B—18
3
4
2
.0'
11
B-19
'I
3
2
0
14
B-20
5
6
-3
.0
6
B-21
0
4
I
• 0
15
t Using chi square tests of goodness of fit
29
Table 8 .
Family
Siete Cerros A
family:fertile and male-sterile segregation
and associated probability values for four genetic ratios
Frequency
Fertile
Male-Sterile
A-I
251
30
A-2
249
34
A- 3 ,'
85
A-4 ''
■j*
Probability for ratios tested
3:1
7:1
15:1
63:1
>.99
---
—
—
>.99
—
—
11
—
>.99
—
—
114
17
——
>.99
—
—
. 86
12
—
>.99
—
—
A-6
139
14
—
.90-.95
A-Il
151
28
——
.95-.98
A-12
234
35
—
A-13
76
12
A-14
284
38
A-15
78
18
A-17
103
18
—
A-19
455
76
-.
A-5
•
.80-.90
—
—
—
>.99
—
—
—
>.99
—
—
—
>.99
—
—
.30-.50
—
—
.90-.99
.
-
—
—
—
.70-.80
>.99 .
t. Probability determined by chi square tests of goodness of fit
+ P < .05 are not given
30
Table 9.
Fnmil-^
B-2 ■
Siete Cerros B F 'family fertile and male-sterile segregation
and associated probability values for four genetic ratios
Frequency
Fertile
Male-Sterile
t
Probability for ratios tested
3:1
7:1
15:1
63:1
>.99
—— .
—
>.99
—
—
—
—
169
28
B-3
97
16
B-8
60
12
B-9
314
62
—
B-13
59
9
—
B-14
80
44.
—
—
1 “—
—
B-16
■ 95
■ 11
—
.98
.30-.50
—
. 14
—
.98-.99 . .80-.90
—
.90-.95
—
B-18
131 .
'
■
B-19
15
2
B-20
102
23
—
.30-.50
.20-.30
>.99
.70-.80
—
>.99
—
-
—
,
—
.30-.50
—
t Probability determined by chi square tests of goodness of fit
4 P < .05 are not reported
—
-.
31
Table 10.
Family
Field grown F headrow distributions within families of
Siete Cerros A for four genetic ratios of fertile!malesterile
Parental headrow__________
Male-Sterile
Fertile
Number of headrows best.
_______ fitting ratios^
3:1
7:1
15:1
1:0
A-1—1
11
4
0
3
2
4
A-l-2
11
2
I
3
2
5
A- 2—1
.5
4
I
I
2
5
5
2
0
2
3
A-2-2
5 '
A-2-3
9
2
I
2
2
2
A-3-1
10
3
0
2
2
4
A—4—I
7
2
0
4
2
2
A-6-1
11
4
0
4
2
6
A—11—I
11
4
I
4
4
3
A—12-1
12
3
I
3
4
5
A-12-2
10
3
I
I
3
2
8
4
0
2
2
4
I
3
.2
7
A-13-1 .
.
A-14-1
■ 14'
4
I
3
A-15-1
■
16
4
3
. o
A-I7-1
9
3
2
A-19-1
7
7
2 .
A-19-2
11
4
2
'
•
3
'■3 '
. 5
5
6
6
3'
3
I.
t Classification based on highest probability values in chi square
tests of goodness of fit
32
Table 11.
Family
Field grown F 4 headrow distributions within families of
Siete Cerros B for four genetic ratios of fertile:malesterile
Parental headrow
Fertile
Male-Sterlie
Number of headrows best
fitting ratios^
3:1
7:1
15:1
1:0
B-2-1
6
2
0
2
I
B-2-2
11
3
I
'4
2
I
B-3-1
7
2
.0
2
.I
5
B-3-2
10
4
I
2
I
I ■
B-8-1
9
3
0
3
I
0
B—9—I
18
7
4
6
4
4
B-9-2
7
3
0
I
2
8
B—11—I
13
I
0
0
0
3
B-13-1
5
2
0 .
2 ■
I
B—20—I
4
2
I
0
I
2
B-20-2
3
I
I
0
I
I ’
B-20-3
7
3
0
I
3
2
.
3 .
■
.I
t Classification based on highest probability value in chi square
tests of goodness of fit
33
segregation (Tables 12 and 13) are similar to field grown F4 headrtiws.
Heterogeneity chi square failed to differentiate A- and B x HRWW■
F3S for male-sterility grown in spring and fall plantings (Table 14).
Results suggest an environmental! and genotypically insensitive genetic
male-sterile.
With the exception of Siete- Cerros B-8, B-9 and B-20, all fami­
lies segregate in a 7:1 ratio in crosses with other genotypes.
Reduced transmission of the male-sterile factor(s) in other
studies (2,5,11,34,71) were not associated with high numbers of non­
segregating progeny.
The genetic system causing male-sterility in
Siete Cerros may differ from other reported male-sterile sources.
The
factor affecting the expression of. male-sterility in Siete Cerros may
have been altered in advanced generations.
Initially, B-20 segregated
7:1 and in later generations segregated 3:1 (Tables 5 and 9).
In
crosses with Cheyenne winter wheat, 3:1 progeny ratios were obtained.
Testcross progeny derived from an F4 male-sterile of B-20 and a fertile,
heterzygous sibling fit a single gene male-sterile segregation with a
chi square value of 0.50 and a probability of fit of .30 to .50.
Chancellor Male-Sterile
The inheritance of the Chancellor male-sterile allele was deter­
mined in crosses with HRWW using F
Z
and F _ progeny.
6
Tests of the
34
Table 1 2 .
headrow distributions of greenhouse-grown Siete Gerros A
and B.families for four genetic ratios of fertile:malesterile
Parental headrow
Fertile
Male-Sterile
Family
Number of headrows best
fitting ratios’^
3:1
7:1
15:1
1:0
A—1-1
11
4
0
3
2
I
A-2-3
9
2
0
3
0
2
A-3-1
10
3
I
0
0
I
A-12-1
12
3
I
I
I
3
A-I2-2
10
3
0
3
0
2
A-14-1
14
4
0
4
3
I
A-15-1
16
4
0
I
0
4
B-2-2
11
3
4
0
0
I
10.
4
I
4
. 2 .
'0
B-8-1
9
3
2
I.
0
2
B-9-1
18
7
I
I
0
3
B-ll-1
3
I
0
•0
0
3
B-20-3
7
3
2
0
0
3
B-3-2
.
'
t Classification based bn higest probability values from chi square
• tests of goodness of fit
35
Table 13.
Family
headrow distributions of greenhouse-grown Siete Cerros
A and B families for various genetic ratios of fertile:
male-sterile
Parental headrow
Fertile
Male-Sterile
Number of headrows best
fitting ratios^
3:1
7:1
15:1
1:0
"NP
3
2
2
I
0
A—2—3—2"
20
3
2
2
0
3
A-3-1-1
16
5
I
2
2
6
B— 2— 2— 3
14
3
4
I
I
3
B—8—1—I
19
3
I
3
2
3
B-20-3-1
15
5
I
I
2
6
CN
I
CO
I
O
CN
I
m
CO
I
CN
20
17
6
I
3.
I
5
t Classification based on highest probability values in chi square
tests of goodness of fit
36
Table 14. ' Siete Cerros spring wheat ms A and ms B x HRWW grown as F
spring and winter wheats
2
Parents ■ ^
Environments
S
Frequency
Fertile
MaleSterile
Probability values for ratios
____________ .tested^
_______
3:1
7:1
15:1
63:1
SW
708
93
—
.30-.50
—
—
SW
462
56
T—
.20-.30
—
—
ms A-3/Sheepers
SW
174
26
——
.80-.90
—
—
913
107
—
.05-.10
—
—
SW
496
65
—
.30-.50
--------
SW
306
36
—
.20-.30
—
—
MS A-14/Trappen
SW
109
15
——
.80-.90
—
—
ms A-17/ID 7457
SW
189
18
—— .
.05-.10
ms B- 3/Cheyenne
■
SW
177
27
ms B-8/MT-6827
SW
323
31
1943
205
—
ms B-18/Winalta
W
114
13
—
ms B-20/Cheyenne
SW-
202
63
ms A-l/HRWW
ms A-2/HRWW
•ms A-4/HRWW
§
S
ms A-ll/HRWW
MS A-12/HRWW
SW
S
§
■S
ms B-9/HRWW SW
.70-.80
.-
.50-.70
—
—
.30-.50
—
-
.05-.10
—
—
—. —
.05— ;10
—
.05-.10 -
—
—
——
—
t Spring planted (S); fall planted (W); both plantings homogeneous (SW)
f p < .05 are not given (using chi square tests of goodness of fit)
§. More than one pollen parent used with no indicated difference deter­
mined by heterogeneity chi square
37
mx C x Centurk F^s showed two types of progeny or subfamilies (Table
15).
Subfamily I segregated for two independent complimentary genes
while subpopulation 2 best fit a single gene ratio.
These results
differ from those of Briggle (5) and Driscoll (10,11) in that they
assumed the reduced number of male-steriles was due to pollen trans­
mission problems.
A gene(s) in Centurk could have influenced the ex­
pression of the Chancellor male-sterile allele.
Probability values for ratios
tested^
3:1 4:1 5:1
7:1 15:1
13:3
—
>.99
—
O
—
I
I
294
—
I
I
1428
§
CD
2
—
A
HO
I
I
1889
I
I
I
1
Frequency
Fertile Male-Sterilet
H
Subfamily
Probability of fit for ms C/Centurk F progeny using chi
square tests of heterogeneity and goodness of fit.
0
Ul
Table 15.
t Homogeneous populations derived from heterogeneity chi square test
(P > .05)
+ Probability values determined by chi square tests of goodness of fit
§ P < .05 are not given
To further test this hypothesis, crosses were made between ms C
and various HRWW and HRSW cultivars.
The F s of ms C x HRWW and ms C
*■
x HRSW were fall and spring planted, respectively.
Winter and Spring
habitat did not influence the expression of male-sterility (Table 16).
Within fall and spring planted F^s differences were observed.
cross ms C x MT 6827 resulted in 104 fertile F
The
plants (Table 17).
r
MT 6827 may possess a suppressor gene(s) for male-sterility.
The F^s
38
Table 16.
Chi square test for heterogeneity for F^ populations of
ms C x HRWW and ms C x HRSW
Frequency
Fertile
Male-Sterile
Subclass
.HRWW
1672
316
HRSW
225
55
1897
371
Total
t
y2
A
P
10-.20
2. 52
t Homogeneous populations derived from heterogeneity chi square test
(P > .05) .
of ms C x Cheyenne and ms C x PI 094436 (Table 17) were the only
winter wheat crosses with
segregation ratios which had a greater
than 99% probability of fitting a single gene male-sterile.
The F^s
of ms C x Bluebird, ms C x Tobari and ms C x Tobari 66 were the only
winter x spring wheat crosses with F^ segregation ratios which had a
greater than 99% probability of fitting a single gene male-sterile
(Table 18).
Allelism Tests Among Selected Families
of the Three Male-Steriles
Two testcrosses, ms A-I x B-20 and ms B-20 x ,ms C/Centurk fit a
two gene testcross ratio.
A third testcross, ms A-I x A-3, was of
insufficient sample size to.be definitive for a two gene testcross
ratio.
ratio.
None of the three allelism tests fit a single gene testcross
Table 17.
Probability values for six genetic ratios in ms C/winter wheat
using chi square tests of goodness of fit
Cross
designation .
ms
ms
ms
ms
ms
ms
ms
ms
ms
ms
ms
ms
C/Brinkle
C/Centurk
C/Cheyenne
C/Crest
C/Froid
C/Lancer
C/MT 6827
C/MT 7243
C/PI 094436
C/Sundance
C/Winalta
C/Winoka
Ratios tested
Ratio fertile/
male sterile^
3:1
4:1
81:17
233:60
95:30
108:9
434:78
118:14
104:0
86:10
83:25
179:33
46:16
105:24
.50-.70
.80-.90
>.99
—
—
—T
—
——
.95-.98
—
—
.30-.50
.90-.95
>.99
.70-.80
.■—
—
——
.80-.90
.90-.95
—
.98-.99
5:1 ■
>.99
.80-.90
—
—
>.99
.30-.50
■—
.30-.50
.30-.50
>.99
.20-.30
.95
families
*
7:1
.70-.80
—
..30-.50
>.99
.90-.95
—
.90-.95
—
.95-.98
.70-.80
—
15:1
**—
—
—
.90-.95
—
—
—
.30-.50
—
.50-.70
—
t Ratios from homogeneous populations using heterogeneity chi square (P > .05).
+ Probability values where P < .05 are not given.
13:3
.70-.80
—
.10-.20
—
——
—
—
——
.20-.30
.20-.30
->.99
Probability values for six genetic ratios in ms C/spring wheat F^'s using
chi square .tests of goodness of fit
Ratios tested
4:1
39:14
—
.05-.10
——
.
I
ms C/Tobari 66
CO
O
27:9
O
ms C/Tobari
.80-.90
.20-.30 '
0
r-
.10-,20
—
—
1
39:7 .
.10-.20
13:3
O
ms C/Siete Cerros
.80-.90
15:1
in
.90-.95
O
CO
43:13
7:1
.30-.50
O
LO
ms C/Bluebird 4
—
m
77:12
O
ms C/Anza
5:1
NJ
3:1
O
Ratio fertile/
male sterile^
?
Cross
designation
H
Table 18.
W
O
.05-.10
——
—
t Ratios from homogeneous populations using heterogeneity chi square (P > .05).
+ Probability values where P < .05 are not given.
.30-.50
1
—
0
CM
—
O
.10-.20
I— I
.80-.90
.30-.50
to
?
>.99
41
The phenotypic expression of male-steriles from ms B-20 x ms C/
.Centurk was different from either parent and may have been environ­
mentally influenced.
Table 19.
Allelism tests between Siete Cerros male-sterile A-l, A-3,
B-20 and Chancellor male-sterile
Female/male
Frequency
Fertile Sterile
2
2
X
(1:1
expected)
X
value
(3:1
expected)
••P ^
8
0
8.00
—
ms A-I/
B-20 (Msms)
20
8
5.14
—
ms A-I/
/ms C/Centurk
22
0
22.00
—
7.33
ms A-3/
B-20 (Msms)
78
.0
78.00
—
•26.00
ms A-3/
/ms C/Centurk
30
0
30.00
—
10. 00
——
ms B-20/
/ms C/Centurk
60
15
27.00
—
1.00
.20-.30
ms A-I/ .
A-3 (Msms)
O
.19 ,
.05-.10
£
LO
t P < .05 are not given
2.67
■-
—
Study II
MONOSOMIC ANALYSES
MATERIALS AND METHODS
Genetic Stocks
Siete Cerros A-I, A-3 and B-20 and Chancellor were the malesterile sources used in the monosomic analyses.
Three monosomic series were used in this study:
I) 'Chinese
Spring', from Dr. E. R. Sears (USDA-SEA, University of Missouri);
2) 'Rescue', from Dr. R. I. Larson (Canada Department of Agriculture,
Lethbridge); and 3) 'Kharkof', from Dr. G. A. Taylor (Montana State
University).
Field and Greenhouse Procedures
The Chinese Spring and Rescue monosomic series were field planted
in May, 1975.
Cytologically determined monosomies.(85) were emascu­
lated at spike emergence and pollinated by random plants from within
families of A-I, A-3 and C.
The use of fertile
pollen parents re­
quired progeny testing using the techniques described by Driscoll (7).
Primary tillers of greenhouse grown F s were examined for malesterility.
Monosomic analyses were similar to that of other workers
(10,36,37,38,48,62,63,81).
Remaining crosses of Rescue and Kharkof monosomies with A-3, B-20
and C were made in 1976 and 1977.
Examination for male-sterility was .
made in the greenhouse for the crosses made in the respective years.
RESULTS AND DISCUSSION
.Siete Cerros
Three Siete Cerros families, A-I, A-3 and B-20, were crossed with
monosomies to determine chromosome location of male-sterile factors.
Siete Cerros A-I appeared to involve chromosomes 1A, ID and 7A (Table
20).
No partial sterility was observed.
Telosomics and isosomics of IA increase fertility but do not pro­
vide complete fertility (61).
The male-sterile observation in mono-
somic IA may be due to univalent shift.
In other studies the main effect of chromosome ID has been to
increase fertility over nullisomic ID plants (61).
Homeologous group 7 also affects fertility in wheat (61).
Mono-
isosomic studies of chromosome 7A increased fertility over nullisomic
counterparts primarily by the suppression of pistilloidy and not fer­
tility restoration.
The partial monosomic analysis of A-3 produced
only for monosomic ID (Table 21).
male-steriles
No partial, sterility was observed in
the monosomic analysis of ms A-3.
Male-sterile expression in B-20 involves chromosome 3B and 7A
(Table 22).
Chromosome 3B is shown to produce irratic response to
fertility although the cause has not been determined (39).
45
Table 20.
Chromosome location of Siete Cerros ms A-I
Monosomic
Designation
Frequency
Sterile
Fertile
Semisterile
.IA
' 12
I
• 0
IB
10
O
0
ID
■ 8
2
0
2B
3
O
0
2D
5
O
3D
3
O
0
4D
7
O
0
SB
I
0.
0
SD
5
0
0
7A
6
I
0
Chromosome location of Siete Cerros male-sterile A-3
'Frequency •
2A
4
2D
8 .
' »
9
4A
9
4B
2
4D ■
4
5A
6
8
O
6
7
O
3B '
to .O
7
o
ID.
o
4.
o
IA
Sterile
O
Fertile
O -O O
Monosomic
■ 50
SD
0'
O
Table 21.
.
46
Table 22.
Chromosome location of Siete Cerros ms B-20
Frequency
Monosomic
Fertile .
Sterile
IA
18
O
IB
2
O
2B
3
O
2D
5
O
3A
8
O
3B
16
3D
3
0
4A
3
• O
4D
7
O
SB
2
O
6A
7
O
6B
13
.
4 ,
O ,
6D
5 '
O
7A
6
I
Chancellor
Monosomic analyses of Chancellor male-sterile involved crosses of
Chinese Spring, Rescue and Kharkof monosomies x F
heterzygotes of C.
Monosomic chromosomes 4A, 4B and 5A segregated for maler-sterility
(Table 23).
The largest proportion of male-sterile progeny involved
chromosome 4A.
Similar results fdr the location of ms C on 4A were
found by Driscoll (10).
Chromosome 4B may be involved,as a modifier
or may be a univalent shift.
Chromosome 5A is known to affect
47
fertility (61) or may. be a univalent shift to SB.
Partial sterility
(semisterile) was observed in monosomic crosses ID, SB, 6A, and 6Bi
All of these monosomies affect fertility (61) or may be suppressions
from other genotypes (Study I).
Partial sterility was not observed in
the Siete Cerros monosomic analyses.
The ms C factor may be environ­
mentally sensitive.
Table 23.
Chromosome location of ms C
Monosomic
IA
IB
ID
2A
2B
■ 2D
3A
3B
3D
4A
4B
4D
SA
SB
SD
. 6A
6D. '
7B
' 7D
Fertile
33
13
21
6
15
28
24
8
2
45
36
25
36
22
11
37
63
24
24
Frequency
Sterile
■
O
O
3
O
O
O
O
O
O
11
•4
O
I
O
O
O
O
.0
0
Semisterile
0
0
3
0
0
0
0
0
0
•0
0
.0
0
2
0
2
2
'0
0
GENERAL DISCUSSION
The Inheritance of. Genetic Male-Sterility
in Siete Cerros and Chancellor
Siete Cerros
Within families of Siete Cerros A and B, potential aneuploidrelated male-sterlies were observed.
The unselected E
3
families of A
and B generally fit 7:1 fertile to male-sterile segregation with an
unexpectedly high number of non-segregating headrows.
Large samples
sizes (from headrows to plantrows) may have allowed a simplified
Mendelian explanation.
Because of small sample sizes, data analyses
were insufficient to test fertile to male-sterile segregation ratios
greater than a two gene model.
Since pollen parents involved in the original crossed, seed may
have provided different numbers of factors, segregation ratios for
families were expected to differ.
The observed 7:1 (fertile:male-sterile) in the F , F , F , and F
j
4
(A- and B x HRWW) generations is not currently explainable.
3
Z
A similar
ratio observed by Falk (15) was attributed to gametic lethality.
Fac­
tors effecting gametic lethality in wheat are known (62).
One family, B-20, was selected for further evaluation as a single
gene male-sterile.
F^ family data and F^ data of crosses to Cheyenne
support a single gene model.
7:1 segregation,
F^ greenhouse data show a reversion to a
ms B-20 may have a very mutable locus (or loci) or is
currently in an unstable state.
49
Other F2 progeny of male-steriles of A and B families in crosses to
HRWW segregated 7:1 similar to the parent families of Siete Cerros.
The
factor(s) invoking male-sterility in Siete Cerros A and B therefore
appear stable in a variety of background genotypes.
Monosomic analyses implicate the involvement of several chromo­
somes in most.Siete Cerros families studied, suggesting multifactorial
control of male-sterility in Siete Cerros A and B.
The occurrence of
male-sterile progeny in a multitude of monosomic testers could be due
to:
I) a dominant-recessive factor relationship; 2) a translocation or
series of translocations; 3) a deletion or series of deletions which
affect gametic transmission rates; 4) a gametic lethal..
Allelism test results fitting a two gene model lend further, support
•to a multigenic cause for male-sterility in Siete Cerros.
Further
research is required to clarify the causal^response relationship of
male-sterility in Siete Cerros.
Chancellor
The presence of a suppressor gene(s) was apparent in F2 families
of ms C x HRWW.
Frequency of male-sterility in C was influenced by the
'
background genotype,
ms C x Cheyenne F 2 fit a single gene recessive .
model while ms C x MT 6827
did not segregate for the male-sterile
character.
MT 6827 has a parent, PI 178383, which is used extensively in the
Pacific northwest wheat improvement programs.
A test of ms C x
50 '
PI 178383 is necessary before widespread use of ms C is made in combi­
nation with other cultivars having PI 178383 in their, background.
The stability of the Chancellor male-sterile is exemplified by
test results involving crosses of heterozygous C plants crossed to a
random male-sterile individual in Composite Cross I (76).
Composite
Cross I (C.C.I) has been grown as a spring wheat bulk for 10 years at
Bozeman, Montana.
and the five
A test cross of ms C.C. I x C
(heterozygote) was made
progeny fit the expected 1:1 ratio (Appendix Table I).,
implying that the same gene conditioning male-sterility in Chancellor
is also functioning in the non-selected C.C. I population as a stable,
male-sterile allele.
Monosomic analyses of Chancellor implicated three and possibly
.6 chromosomes in the segregation of ms C.
The involvement of multiple
chromosomes is different from other studies (5,10).
The multifactorial
(multichromosomal) effects may be related to expression of suppressor
factors hypothesized in Study I.
REFERENCES
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Genetic male
2.
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'
_____ . 1973. A chromosomal male-sterility system- of producing
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■
_____. 1977. Registration of Cornerstone male sterile wheat
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*
. /,«
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•
54
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APPENDIX
60
Appendix Table I.
Allelism tests of Chancellor male-sterile and other
male- sterile sources
Frequency
Cross designation■
2
X
Fertile
Male-sterile
P +
F
(1:1 expected)
C .C.I ms/ms C
3
2
ms B-6/ms C
2
0
2.0
.10-.20
ms A-2/ms C
7
0
7.0
—
ms A-14/ms C
22
0
22.0
—
ms A-20/ms C
6
0
6.0
—
ms C/C.C. I.://ms B-14/Cnn
. 15
0
15.0
—
ms C/ms D/3/ms A-6/
MT 00906//MT 7015
. 3
2
.20
t Probability values less than '0.05 are not reported.
.20
.50-.70
.50-.70
61
Appendix Table 2.
Fertile and male-sterile, progeny of heterozygous
Siete Cerros ms A-I x monosomies
Female monosomies
Male ■
sterile
Fertile
Chinese Spring IA
IB .
6B
6D
7A
8
3
I
5 ■
6
Kharkof
ID
4D
7
I
O
O
Rescue
IA
IB
ID
2B
2D
3D
,40
4
7
3
3
5
3
6
I.
O
2
O
■O
O .
O
Appendix Table 3.
IA
. ID.
2A
2D
3A
3B
4A
4B
4D
5A
5D
6D
O
O
O
.O
I
Fertile and male-sterile progeny of heterozygous
Siete Cerros ms x Rescue monosomies
Fertile
Rescue
'
';
'
4
.
7
. 4
8
9
6
9
2
4
6
8 .
7
Male
sterile
O
I
O
O
O
'O
O
O
O
O
O
O
.
62
Appendix Table 4. Fertile and male-sterile progeny of heterozygous
Siete Cerros ms B-20 x monosomies
Female monosomies
■ Rescue
Kharkof
Fertile
Male-sterile
IA
18
O
IB
2
Q
2B
3
O
3A
8
O
3B
16
4
4A
3
O
SB
2
O
6A
7
O
6B
13
O
6D
5
0
7A
6
1
63
Appendix Table 5.
Fertile and male-sterile progeny of heterozygous
Chancellor male-sterile x monosomies
Female monosomies
Fertile
Chinese Spring IA
IB
ID
2D
3D
4A
4B
5A
SB
6A
6B
GD
30
7
. I
21
2
7
12
20
22
25
8
30
Rescue
Karkof
IA
2A
2B
.3B
4A '
. 4B
6B
GD
7D
IB
ID
2B
2D
3A
4B
4D
SA
SD . •
GA
GB
GD
7B
TD
Male
sterile
0
0 .
3
0
0
0
0
I
0
0
0
0
3
6
5
8
38
14
8
I
4
0
0
0
0
11
4
0
0
0
6
20
10
8
24
10
25
16
11
12
21
32
24
20
0
0
.0
0
0
0
0
0
0
. 0
0
0
0
0
'A
Semisterile
0
0
I .
0
0
0
0
0
2 .
2
I
2
.
■0
0
0.
I
0
0
I
0
0
0
2
0
0
0
0
0
0
6
0
I
0
0
0
D378
J&31
coo. 2
DATE
Johnson, Duane L
Genetic characteriza­
tions of three p a l e ­
st eri Ies in wheat . . .
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