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GENETIC RELATIONSHIPS BETWEEN ABOVE-GROUND
‘PLANT PARTS IN CABBAGE,

(Brassica oieracea var. capitata Linn.)

Dissertation for the Degree of Ph. D.
MICHIGAN STATE UNIVERSITY
WILLIAM LYNN SUMMERS
1977

:4" Y

n.”t x_‘

This is to certify that the

thesis entitled

GENETIC RELATIONSHIPS BETWEEN ABOVE-GROUND
PLANT PARTS IN CABBAGE,

(Brassica oleracea var. capitata Linn.)

presented by
WILLIAM LYNN SUMMERS

has been accepted towards fulfillment
of the requirements for

degree in Horticulture

“4%.“, 4,. h”\

(/ Major professor

Ph.D.

Dateh"? A? (27/7

0-7639

‘ABSTRACT

GENETIC RELATIONSHIPS BETWEEN ABOVE-GROUND PLANT PARTS

IN CABBAGE, (Biassica oleracea var. capitata Linn.)
By

William L. Summers

Thirty open-pollinated cabbage cultivars were screened for quan-
titative traits during 1973. Using an efficiency index, (total non-
wrapper leaf weight + stalk size (weight))/(head weight). 2 efficient
cultivars. 'Baby Head' and 'Badger Ballhead'. and 3 less efficient cul-
tivars. 'Red Danish', 'Chieftain Savoy' and P.I. 215514. were hybridized
to obtain F1 and F2 populations using a NC II design. For the selected
cultivars. data were collected on the effects of 1 generation of in-
breeding. Three plantings of the F1 population and 2 plantings of the
F2 population were made for this study. Narrow sense heritability,
genetic correlation and direction of dominance determinations were made.
F2 segregation ratios for 7 traits including maturity. head weight.
total non-wrapper leaf weight. stalk size. non-wrapper leaf number,
efficiency index and non-wrapper leaf size were compared to weighted
ratios produced by postulated models.

Crosses involving the cv. Red Danish generally exhibited dominance
patterns which were opposite to patterns observed in crosses involving

only green cabbage cultivars. A digenic model with 5 alleles per loci

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William Lynn Summers

is proposed for each trait. Higher heritability estimates were ob-‘
served for green cabbage crosses than for red x green cabbage crosses.
Heritability estimates suggest that progress could be made in selecting
for better plant efficiency, since most of the desirable traits, i.e.,
increased head weight, reduced days to maturity, and reduced total
non-wrapper leaf weight and number were dominant in this study. Gene-
tic correlations suggest progress can be made with 2 or 3 traits by
selecting for a trait that is highly correlated with these traits.
Added inbreeding of open pollinated cabbage cultivars may be desirable
for studies of head weight, total non-wrapper leaf weight, non-wrapper

leaf number and efficiency index. Studies of maturity, stalk size

and non-wrapper leaf size did not require inbreeding in this study.

GENETIC RELATIONSHIPS BETWEEN ABOVE-GROUND PLANT PARTS

IN CABBAGE. (Brassica oleracea var. capitata Linn.)

By
William Lynn Summers

A DISSERTATION

Submitted to
Michigan State University
in partial fulfillment of the requirements
.for the degree of

DOCTOR OF PHILOSOPHY
Department of Particulture

1977

To Kent L. Phillips
without whose help and motivation the data

Collection phase of this study would not have been possible.

ii

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ACKNOWLEDGEMENTS

I would like to thank Dr. Shigemi Honma for his
guidance and encouragement throughout this study. The
writer also wishes to express his appreciation to Drs.
L. R. Baker, J. L. Gill, J. N. Hanover, B. 8. Dean, and
H. C. Price for their interest and suggestions. Special
thanks is due to those people without whose help this
study could never have been completed including: Amos
Lockwood, for his field assistance and council, Kent
Phillips. for his help in harvest and data collection,
vegetable breeding graduate students who provided sug-
gestions and encouragement, and my wife. Sylvia for her

continued understanding and patience.

iii

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ACKNOWLEDGEMENTS

I would like to thank Dr. Shigemi Honma for his
guidance and encouragement throughout this study. The
writer also wishes to express his appreciation to Drs.
L. R. Baker, J. L. Gill, J. W. Hanover. B. B. Dean, and
H. C. Price for their interest and suggestions. Special
thanks is due to those people without whose help this
study could never have been completed including: Amos
Lockwood. for his field assistance and council, Kent
Phillips, for his help in harvest and data collection,
vegetable breeding graduate students who provided sug-
gestions and encouragement, and my wife, Sylvia for her

continued understanding and patience.

iii

TABLE OF CONTENTS

LISTOFTABLES........................ .......................... V
LIST OF FIGURES ................................................. xvii
Immumoi .............................. N ...................... 1
REVIEW or LITERATURE ............................................ 3
MATERIALS AND METHmS .................... . ......... ..... 6
RESULTS AND DISCUSSION ........................ 31
concwsmis . ............................ ..... 232
BIBLIOGRAPHY ................. ........................ 235

iv

Table

Table
Table
Table

Table

Table

Table

Table

Table

Table

Table
Table
Table
Table

"Table

10.

11.
12.
13.
14.

15.

LIST OF TABLES

Efficiency index for 30 ca bbage cultivars

grown in 1973 .................. . ....................... 14
Plant populations grown ................................ 15
Dependent variables used in this study ................. 18

Analysis of variance for cultivar, in-
breeding and planting effects .......................... 19

F1 data separation for variance component

analysis. Twelve (12) data groups each
composed of nine elements. three crosses
by three replications .................................. 22
Variance component analysis for F1 data ................ 24

Seventeen trait combinations generated
by adding data on one trait to data for
another trait from the same cabbage head ............... 26

Genetic covariance and correlation analy-

'siS from AOV #1 ........................................ 27

Analysis of variance table for cabbage
maturity in days from transplanting to

harvest ................................................ 32
Cultivar x planting interaction means

in days from transplanting to harvest .................. 33
1974L Cultivar and F1 performance means ................ 35
19755 Cultivar, F1 and F2 performance means ............ 36
1975L Cultivar, F1 and F2 performance means ............ 37

Parent.F , F2 and backcross maturity
distributionsz in percent for the 1975E
planting of the cross Baby Head x Badger .
Ballhead ............................................... 38

Parent.F % and F maturity distributions
in percen for t e 1975L planting of
the cross Baby Head x Badger Ballhead..

V

'I

 

Table

Table

Table

Table

Table

Table

Table

Table

Table

Table

Table

Table

Table

16.

17.

18.

19.

20.

21.

22.

23.

24.

25.

26.

27.

28.

Postulated cultivar genotypes for the . -
1975E planting ......................................... 41'

Postulated F1 genotypes for the 1975E
planting ................................. . ............. 42

Chi-square test for goodness of fit ,
to the postulated model for maturity in
the F2 generation (1975E planting) ..................... 43

Chi-square test for goodness of fit to
the postulated model for-maturity in the
F2 generation (1975L planting) ......................... 45

Parent, F1 and F maturity distributions
in percent for tEe 1975E planting of the

cross Baby Head x P.I. 215514 .......................... 47
Parent. F and F2 maturity distributions

for the 1575L planting of the cross Baby .
Head x P.I. 215514 ..................................... 48

Parent, F . F2 and backcross maturity

distributions in percent for the 1975E

planting of the cross Baby Head x Chief-

tain Savoy ............................................. 49

Parent. F1 and F maturity distributions
in percent for tGe 1975L planting of the
cross Baby Head x Chieftain Savoy ...................... . 51

Parent, F , F2 and backcross maturity

distributions in percent for the 1975E

planting of the cross Red Danish x

Badger Ballhead. ....................................... 53

Parent, F and F maturity distributions
in percen for t e 1975L planting of the
cross Red Danish x Badger Ballhead ..................... 54

Parent, F1 and F maturity distributions
in percent for tfie 1975E planting of the
cross Red Danish x P.I. 215514....' ..................... 55

Parent and F maturity distributions in
percent for She.1975L planting of the
cross Red Danish x P.I. 215514 ......................... 57

Parent. F . F2, and backcross maturity

distributions in percent for the 1975E

planting of the cross Red Danish x Chief-

tain Savoy..w ........................................... 58

vi

Table 29.
Table 30.

Table 31.
Table 32.
Table 33.
Table 34.
Table 35.
Table 36.
Table 37.
Table 38.
Table 39.

Table 40.

Table 41.

Parent, F and F maturity diStributionS
in percen for t e 1975L planting of
the cross Red Danish x Chieftain Savoy .................

Heritability estimates for seven traits
from green and red x green cabbage crosses
grown in three plantings ...............................

Genetic correlation between maturity and
six other traits in cabbage ............................

Analysis of variance table for head
weight in kilograms ....................................

Cultivar x planting interaction means
for head weight in kilograms.......... .................

Cultivar x inbreeding interaction means
for head weight in kilograms ...........................

Parent, F1, F2, and backcross head weight
distributions in percent for the 1975E

planting of the cross Baby Head x Badger

Ballhead ...............................................

Parent, F1 and F head weight distributions
in percent for tge 1975L planting of the
cross Baby Head x Badger Ballhead ......................

Chi-square test for goodness of fit to
the postulated model for head weight in
the F2 generation (1975E planting) ............ .........

Chi-square test for goodness of fit to the
postulated model for head weight in the
F2 generation (1975L planting) .........................

Parent, F1 and F2 head weight distribu-
tions in percent for the 1975E planting
of the cross Baby Head x P.I. 215514 ...................

Parent. F1 and F2 head weight distribu-
tions in percent for the 1975L planting
of the cross Baby Head x P.I. 215514 ...................

Parent, F . F2. and baCkcross head weight
distributions in percent for the 1975E

planting of the cross Baby Head x Chief-

tain Savoy .................................. ‘ ...........

vii

59

61

63'

66

67

68

70

-72

73

76

78

79

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Table 42.

Table 43.

Table 44.
Table 45.
Table 46.

Table 47.

Table 48..

Table 49.
Table 50.

Table 51.
Table 52.
Table 53.

Table 54.

Parent.F and F2 head weight distribu-
tions in Bercent2 for the 1975L planting
of the cross Baby Head x Chieftain Savoy ...............

Parent,F . F2 , and backcross head weight
distributions2 in percent for the 1975E

planting of the cross Red Danish x Badger

Ballhead ...............................................

Parent, F and F2 head weight distribu-
tions in Bercent2 for the 1975L planting
of the cross Red Danish x Badger Ballhead ..............

Parent, F and F2 head weight distribu-
tions in Bercent2 for the 1975L planting
of the cross Red Danish x P. I. 215514 ..................

Parent and F head weight diStributions
in percent fdr the 1975L planting of
the cross Red Danish x P.I. 215514 .....................

Parent,F , F2 , and backcross head weight
distribUtIons2 in percent for the 1975E

planting of the cross Red Danish x Chief-

tain Savoy .............................................

Parent.F and F2 head weight distribu-
tions in fiercent2 for the 1975L planting
of the cross Red Danish x Chieftain Savoy ..............

Genetic correlation between head weight
and four other traits in cabbage .......................

Analysis of variance table for total non-
wrapper leaf weight in kilograms ..... ~ ..................

Cultivar x inbreeding interaction means
in kilograms for total non-wrapper leaf
weight .................................................

Cultivar x planting interaction means in
kilograms for total non-wrapper leaf
weight.... ..................... . .......................

Inbreeding x plantings interaction means
in kilograms for total non-wrapper leaf -
weight................... ..............................

Cultivar x inbreeding x planting interaction

means in kilograms for total non-wrapper
leaf weight ............................................

viii

82

84

85

86

88

89

90

92

94

95

96

98

99

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Table 55.

Table 56.

Table 57.

Table 58.

Table 59.

Table 60.

Table 61.

Table 62.

Table 63.

Table 64.

Table 65.

Parent, F1, F2 and backcross total non-

wrapper leaf weight distributions in

percent far the 1975E planting of the

cross Baby Head x Badger Ballhead ...................... 101

Parent, F1 and F total non-wrapper leaf

weight distributIons in percent for the

1975L planting of the cross Baby Head x

Badger Ballhead ........................................ 102

Chi-square test for goodness of fit to

the postulated model for total non-wrap-

per leaf weight in the F2 generation

(1975E planting) ....................................... 103

Chi-square test for goodness of fit to
the postulated model for total non- wrap-
per leaf weight in the F2 generation
(1975L planting) ....................................... 106

Parent, F1 and F total non-wrapper leaf
weight distributions in percent for the
1975E planting of the cross Baby Head x
P.I. 215514 ............................................ 108

Parent. F1 and F? total non-wrapper leaf
weight distribut ons in percent for the
1975L planting of the cross Baby Head x
P.I. 215514 ............................................ 109

Parent, F1. F . and backcross total non-

wrapper leaf weight distributions in ~

percent for the 1975E planting of the cross

Baby Head x Chieftain Savoy ............................ 110

Parent, F1 and F total non-wrapper leaf
weight distributions in percent for the
1975L planting of the cross Baby Head x
Chieftain Savoy ........................................ 112

Parent, F1. F . and backcross total non-

wrapper leaf weight distributions in per-

cent for the 1975E planting of the cross

Red Danish x Badger Ballhead ........................... 113

Parent,F and F total non-wrapper leaf

weight diStributions in percent for the

1975L planting of the cross Red Danish

x Badger Ballhead ...................................... 115

Parent.F and F total non-wrapper leaf

weight di tribut ons in percent for the

1975E planting of the cross Red Danish x . '

P.I. 215514.. ............... . ..... . ............. . ...... 116

ix

 

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Table

Table

Tably

Table

Table

Table

Table

Table

Table

Table

Table

Table

103.

104.

105.

106.

107.
108.
109.
110.

111.

112.

113.

114.

115.

Parent and F non-wrapper leaf number

distribution; in percent for the 1975L

planting of the cross Red Danish x P.I.

215514 ................................................ 172

Parent, F1. F and backcross non-wrapper

leaf number distributions in percent for

the 1975E planting of the cross Red Danish

x Chieftain Savoy ..................................... 174

Parent,F and F2 non-wrapper leaf num-

ber distributions in percent for the

1975L planting of the cross Red Danish

x Chieftain Savoy ..................................... 175

Analysis of variance table for efficiency
index (non-wrapper leaf weight + stalk

size (weight)/head weight) ............................ 178
Cultivar x inbreeding interaction

means for efficiency index ............................ 179
Cultivar x planting interaction means for

efficiency index............................ .......... 181
Inbreeding x plantings interaction means

for efficiency index .................................. 182
Cultivar x inbreeding x planting interac-

tion means for efficiency index ....................... 183
Parent,F

F2 and backcross efficiency
index disiributions in percent for the
1975E planting of the cross Baby Head
x Badger Ballhead ..................................... 185

Parent,F and FE efficiency index dise
tribution; in pegc cent for the 1975L
planting of the cross Baby Head x .
Badger Ballhead ....................................... 186

Chi-square test for goodness of fit to
the postulated model for efficiency index
in the F2 generation (1975E planting) ................. 188

Chi-square test for goodness of fit to
the postulated model for efficiency in-

dex in the F2 generation (1975L planting) ............ 189
Parent,F and F2 efficiency index distri-
butions 1A percent for the 1975E planting

of the cross Baby Head x P. I. ,215514 .................. 192

xiii

 

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Table

Table

Table

Table‘

Table

Table

Table

Table

Table

Table

Table

Table

116.

117.

118.

119.

120.

121.

122.

123.

124.

125.

126.

127.

Parent.F and F2 efficiency index

distributions in2 percent for the 1975L

planting of the cross Baby Head x

P. I. 215514 ........................................... 194

Parent.F F2 and backcross efficiency
index disiributions in perCent for the
1975E planting of the cross Baby Head x

Chieftain Savoy ....................................... 195
Parent,F and F? efficiency index dis-

tribution; in pegc cent for the 1975L

planting of the cross Baby Head x Chief-

tain Savoy ..... - ....................................... 196
Parent.F

, and backcross efficiency
index disiribations in percent for the
1975E planting 0f the cross Red Danish x
Badger Ballhead.’ ...................................... .198

Parent. F1 and F2 efficiency index distri-
butions in percent for the 1975L plant-
ing of the cross Red Danish x Badger

Ballhead .............................................. 199
Parent,F and F efficiency index distri-

butions iA percefit for the 1975E plant-

ing of the cross Red Danish x P.I. 215514 ............. 201

Parent and F1 efficiency index distribu-
tions in percent for the 1975L planting
of the cross Red Danish x P.I. 215514 ................. 202

Parent,F F2 and backcross efficiency

index disiributions in percent for the

1975E planting of the cross Red Danishx

Chieftain Savoy ....................................... 203

Parent, F and F efficiency index distri-

butions 1A perceflt for the 1975L plant-

ing of the cross Red Danish x Chieftain

Savoy ................................................. 205

Analysis of variance table for non-n .
wrapper leaf size in kilograms ........................ 207

Cultivar x planting interaction means in
kilograms for non-wrapper leaf Size.. ................. 208

Cultivar x inbreeding x planting inter-

action means in kilograms for non-wrapper‘
leaf size ....................................... . ...... 209

xiv

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Table 92. Parent, F , F and backcross non-wrapper
leaf numbér distributions in percent
for the 1975E planting of the cross
Baby Head x Badger Ballhead ........................... 156

Table 93. Parent, and F2 non-wrapper leaf number
distributions in percent for the 1975L
planting of the cross Baby Headx Badger ‘ ,
* Ballhead .............................................. 157

Table 94. Chi- -square test for goodness of fit to
the postulated model for non-wrapper leaf
number in the F2 generation (1975E
planting) ............................................. 158

Table 95. Chi-square test for goodness of fit to the
postulated model for non-wrapper leaf num-
ber in the F2 generation (1975L planting) ............. 159

Table 96. Parent, F and F2 non-wrapper leaf num-
ber distributions in percent for the
1975E planting of the cross Baby Head -
x P.I. 215514 ......................................... 162

Table 97. Parent, F and F2 non-wrapper leaf num-
ber distributions in percent for the
1975L planting of the cross Baby Head x
P.I. 215514 ........................................... 163

Table 98. Parent, F1.F and backcross non-wrapper
‘ leaf number distributions in percent for
the 1975E planting of the cross Baby
Head x Chieftain Savoy ............................... ‘. 165

Table 99. Parent. F and F non-wrapper leaf num-
ber distributions in percent for the
1975L planting of the cross Baby Head x
Chieftain Savoy ....................................... 167

Table 100. Parent, F1, F and backcross non-wrapper
leaf number distributions in percent for
the 1975E planting of the cross Red Danish
x Badger Ballhead ..................................... 168

Table 101. Parent, F1 and F non-wrapper leaf num-
ber distributions in percent for the
1975L planting of the cross Red Danish x
Badger Ballhead ....................................... 169

Table 102. Parent. and F2 non-wrapper leaf num-
.ber distributions in percent for the 1975E
planting of the cross Red Danishx
P.I. 215514. .......................................... 171

xii

TabTe 140.

Table 141.

Parent, F1, F and backcross non-wrapper

leaf size disEributions in percent for

the 1975E planting of the cross Red

Danish x Chieftain Savoy .............................. 229

Parent, F1. and F2 non-wrapper leaf

size distributions in percent for the

1975L planting of the cross Red Danish

x Chieftain Savoy ............................ - ......... 231

xvi

Figure 1.

Figure 2.

Figure 3.

'LIST or FIGURES

The efficient cultivar Baby Head:
A - cabbage head; B - non-wrapper .
leaves; C - stalk ...................................... .8

The inefficient cultivar P. I. 215514:
A - cabbage head; 8 - non-wrapper
leaves; C - stalk ....................... ' ............... 10

The inefficient cultivar Chieftain

Savoy: A - cabbage head; 8 - non-wrapper
leaves; C - stalk ........ _ .............................. 12

xvii

INTRODUCTION

Energy and food shortages have placed increased emphasis on the
efficient use of existing resources to produce greater amounts of
food. Vegetable cUltivars grown during a time of abundant energy and
low cost fertilizer may no longer provide the yield per acre or pro-
fit needed to provide income. In an attempt to evaluate existing
vegetable cultivars and to breed new lines with greater efficiency
and less waste i.e.. (unsalable above ground portion - kg / salable
above ground portion - kg) open-pollinated cabbage was selected as
a crop system for such a study.

Presently. the breeding of cabbage for yield is based on infor-
mation derived from crosses made between green non-savqyed lines. De-
velopment of red and savoy cabbage cultivars is also dependent on the
same information. If previous studies represent an adequate sample
of genes and gene action for all cabbage cultivars. then. inbreeding of
cabbage would be ideally suited to create efficient lines for hybrids
since large head weight. few non-wrapper leaves and small non-wrapper
leaf size are transferred as dominant traits. Experiments to increase
cabbage efficiency by reducing plant parts such as stalk size and total
non-wrapper leaf weight and to maximize other plant parts such as head
weight have not been undertaken in cabbage.

The objectives of this study were:

1. Collect data on above-ground plant parts in cabbage to

provide information for future breeding programs.
I 1

2
Test whether inbreeding is necessary prior to using open-

pollinated cabbage varieties in an inheritance study.
Determine whether inheritance patterns of above-ground
plant parts are Similar for red and green cabbage with
either a smooth or savoy leaf.

Provide paternal half sib heritability estimates for
individual above-ground plant traits and genetic corre-
lation estimates for pairs of these traits.

Simulate a breeding program using the heritability and
genetic correlation estimates to provide estimates of

expected genetic gain.

REVIEW OF LITERATURE

Although cabbage is widely planted and consumed in many 'areas of
the world, genetic investigations have been limited to studies of
maturity. head weight. non-wrapper leaf number and various head dimen-
sions. These studies were further limited to smooth-leaf types of
green cabbage.

DetJen and McCue (5) found that open-pollinated lines possessed
variation for horticultural characters even when released for commer-
cial use and that selection for plant type and color could be prac-
ticed in these lines. These authors also noted large phenotypic dif-
ferences in quantitative characters between years and concluded that
environment played a large role in the phenotypic expression of cabbage.
They also found that non-wrapper leaf size was directly related to
head and stalk size and realized that selection for one of these traits
may predetermine the other two. Pearson (20) found environment played
a large role in head formation and non-wrapper leaf production. Hot
weather promoted greater non-wrapper leaf number at the expense of_early
head maturity. He also found a low number of non-wrapper leaves to be
dominant to a large number of non-wrapper leaves and noted a direct.
correlation between days to maturity and head wieght. In 1956 Yarnell
(37) found small numbers of non-wrapper leaves were completely dominant.
He also noted that head weight was only mildly affected by inbreeding ‘
depression. He also reportedthat earliness was dominant to later
maturing cultivars. In a study of growing media on transplant

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performance. Honma (9) reported that certain cultivars were more effi-
cient at producing heads than others. Using a half diallel with 6
open-pollinated commercial cultivars, inbred 2 additional generations.
Swarup et a1. (33) found early maturity and large head weight were
both dominant to late maturity and small head weight. Research in
India and Canada by Swarup at al. (32) and Chiang (3) showed close
agreement in spite of widely different envirbnments. Swarup et a1.
(32) found heritabilities for maturity. head weight and non-wrapper.
leaf number were 0.80. 0.37, and 0.52 respectively while Chiang (3)
using a full 5 parent diallel in Canada found heritabilities for these
traits to be 0.83. 0.04, and 0.35. Dickson and Stamer (7) found
heritability for percent dry matter in cabbage to be between 0.5 and
0.6. Their research also noted a direct correlation between high dry
matter content and resistance to frost damage.

At present. genetic information for red and savoy cabbage consists
of genetic models for red leaf color inheritance proposed by Sutton
(31). Kristofferson (13), and Pease (21) and for savoy leaf inheritance
by Swan (14). RasmusSon (24). and Tschermak (35). Currently, genetic
improvement in red and savoy cabbage lines is based on research done
only on green cabbage cultivars with smooth leaves.

- A better understanding of inheritance patterns for total basal leaf
weight and leaf number could possibly lead to solutions for other cab;
bage problems. Maynard (18) attributes tipburn in cabbage to an accum-
ulation of Ca. Mg and K in basal leaves. The absence of a rapid trans-
fer path for these ions from older basal leaves to younger head leaves
is thought to increase tipburn incidence. Wiebe et a1. (36) have shown

that the high transpiration of basal leaves depletes meristem water

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supply causing a reduction in both plant volume and plant growth during
the day. Night time reductions in transpiration allow a more favorable
plant water balance and resumption of plant growth. Ion mobility stu-
dies by Plazkill et al. (19) using 456a, lend Support for a link between
basal leaf transpiration rate and increased tipburn in cabbage. Cab-
bage trait correlation studies by Sharma et a1. (28). suggest that se-
lection for a reduced number of non-wrapper leaves may produce an ap-
preciable increase in head weight. With new knowledge of inheritance
patterns for leaves it may be possible to reduce both the number and

size of basal leaves and minimize tipburn in cabbage without sacrific-

ing total head production.

 

MATERIALS AND METHODS

Screening Studies
During the summer of 1973. 1 Plant Introduction (P.I.) and 29

open-pollinated cultivars of cabbage. a random sample of open-polli-
nated cultivars. were screened for parental material. The varieties
were seeded in peat pots filled with soil in a greenhouse. The seed-
lings were watered and fertilized as needed. When the stems reached
a diameter of 4-5 mm the plants were moved out-of-doors for hardening.
One month after seeding the plants were transplanted in the field in
a randomized complete block design with 5 replications and 5 plants
per plot. The plants were transplanted 0.46 m apart in the row and’
1.07 m between rows. As the plants approached maturity. the lines were
tested by thumb pressure to determine maturity. Head formation began
in late August and by mid-September the wrapper leaves of some lines
began to slip. The heads of these lines were firm and were harvested
by cutting the plant at groung level. Plants were harvested daily_after
mid-September. . .
Recorded data included harvest date. individual weights for the
head. stalk. and non-wrapper leaves and the number of non-wrapper leaves.
Parental selection was based on a cabbage plant efficiency index.
(tatal non-wrapper leaf weight + stalk size (weight))/(head weight).
This measurement provided the number of kg of non-wrapper leaves and
stalk required to produce a kg of head. Using this method the 30 culti-
vars were ranked from efficient (Figure 1) to least efficient (Figures

6

Figure l. The efficient cultivar Baby Head: A - cabbage head;

8 - non-wrapper leaves; C - stalk.

 

 

Figure 2. The inefficient cUltivar P.I. 2l5514: A - cabbage head;

B - non-wrapper leaves: C - stalk.

 

11

Figure 3. The inefficient cultivar Chieftain Savoy: A - cabbage head;

8 - non-wrapper leaves; C - stalk.

 

 

a
II.
II‘
ID

"‘1: ’

.~vu D

Ice-"a

...;:‘

‘n! l- n.

0.. C
a

ll}
I’-

.Q

‘6

r
5

N

.‘h'

13
2-3). The 2 most efficient cultivars and the 3 least efficient cultivars
were'selected as parents (Table 1). Single plant selections of each of

these cultivars were vernalized for 13 weeks at 4°C to induCe flowering.

Crossing_Scheme

The crossing scheme used in this study was the North Carolina "Fac-
.torial" Design 11 (NCII) proposed by Comstock and Robinson (4). TIhis
design was chosen because it produced the maximum amount of genetic in-
formation per family group evaluated (10. 22). The design required di-
vision of the parents into 2 groups: lines to be used as males and lines
to be used as females. In this study single plant selections of 'Badger
Ballhead'. P.I. 215515. and 'Chieftain Savoy' were used as males while
single plant selections of 'Baby Head'. and 'Red Danish' were used as
famales. anch female line was reciprocally crossed to each male line.
These 6 reciprocal combinations constituted the F1 population. Four F1
seedlings of each cross were randomly selected. grown. vernalized and
selfed to provide-the F2 population. Each of the cultivars were vegeta-
tively propagated to facilitate reciprocal backcrosses;

Incompatibility was encountered and resulted in a limited back-
Cross (BC1 a 8C2) population. Original seedlots of each of the culti-
vars were also grown in each test and were designated the P1 population.
Each parental plant was selfed once (51 population) and was planted in
EACH test (Table 2). The parental populations P1 and 51 were analyzed
to determine the effect of 1 generation of inbreeding. Estimates of ge-
"etic variance components. heritabilities within traits and estimates of
genetic correlation betweén traits for each of the 3 F1 plantings were
"Ede using the N011 analysis and covariance. correlation techniques de-

scribed by Kempthorne (11). Frequency distributions of the 12 F2

. 14

 

 

 

Table 1. Efficiency index for 30 cabbage cultivars grown
in 1973.
' Mean*'
Cultivar Efficiency
Index
Baby Head 0.3569 a **
Badger Ballhead IO.4795 ab **
Early Jersey Wakefield 0.4885 ab
Badger Market ‘ 0.5020 ab
Greenback 0.5216 abc
Early Marvel 0.5305 abc
Metor 0.5313 abc
Copenhagen Market Early 0.5376 abc
Stokes Wisconsin Ballhead 0.5496 ' _ abc
Emerald Acre 0.5664 abc
Early Round Dutch 0.5725 . abc
Bonanza 0.6043 abc
Early Greenball 0.6248 abc
Copenhagen Market Late 0.6570 abcd
Red Acre 0.6570 abcd
Savoy Cabbage 0.6635 abcd
Stokes Super Acre 0.6787 abcd
Stokes Viking Golden Acre 0.6817. abcd
Green Acre 0.6902 abcd
Jumbo 0.6922 - abcd
Viking Extra Early Strain 0.7268 abcd
Autumn Marvel 0.7657 abcd
Penn. State Ballhead 0.7869 abcd
Danish Ballhead 0.8600 bod
Golden Acre 84 0.9970 cde
Wisconsin Hallander No. 8 1.1080 de
Eastern Ballhead 1.1157 . de
Red Danish 1.3316 e **
Chieftain Savoy 1.3714 e **
P. I. 215514 2.0312 4 **

 

* Means followed by the same letter are not significantly different
at the 5% level by Duncans Multiple Range Test.

*? SeleCted as parents.

Table 2. .Plant populations grown.

15

 

 

Plant
Population

1974
Late

PlantingS’
1975
Early

T975
Late

 

 

16
populations were compared with the P1 and F1 distributions. to determine
inheritance patterns.

1974 Field Experiments-

A randomized complete block experiment with 3 replicates was planted.
Seed from the P1, 51. and F1 populations were planted in greenhouse flats
(an June 28. Seedlings were mechanically transplanted 4 weeks later.
thrvest began October 26 and ended November 20. A total of 1625 plants
were harvested. The thumb pressure method was used to determine maturity.
Other measures of maturity were used to obtain an objective measurement.
These included: 1) pressure testing each cabbage head 4 times with a Mag-
. ness-Taylor Penetrometer (16) and recording the pounds of force required
for a 3/16" probe to penetrate 1/2" into the head and. 2) use of water
displacement to determine head volume and density. Penetrometer head
measurements increased to a peak and remained constant prior to over-ma-
turity or bursting. .Density readings increased toward 1.0 as air spaceS‘
inside the head diminished with maturity. These 2 measures were corre-
lated to determine how well pressure readings predict density measures.
For the 1625 cabbage heads measured. the correlation between the mean of
4 pressure readings per head and head density was a highly significant

(P<O.01)Ir-0.32.

1975 Field Experiments

Two plantings were made. The first planting included the P1,
51, F1. F2. ac1 and ac2 populations. The second planting included all
populations except the BCI and 802 populations. Each planting was a ran-
-domized complete block with 3 replications. The first planting was
seeded in a cold frame with heating cable on May 5. The second planting

n
“i

In.‘

‘3.

I01
I
Ch

17 .
was seeded on May 29. All seedlings were mechanically transplanted in

the field. Harvesting began on August 9 for the early planting and on
October 20 for the late planting. Above-ground plant parts were har-
vested when the mean reading of 2 penetrometer punches was equal to or
exceeded the appropriate parent or midparent mean penetrometer reading
determined in 1974. In addition each plant was classified for leaf col?
~or: red or green. for leaf type: normal or savoy. and for head Shape:
drum. pointed or flat. There were 7 dependent variables included for

analysis (Table 3).

Cultivar Inbreeding Analysis .
Because hybrids in genetic research should be both genotypical-

ly and phenotypically homogeneous. open-pollinated cultivars should be
tested to measure their degree of homozygosity. Since the degree of
homozygosity for each cultivar was unknown. analysis of variance (AOV)
tests were applied to plot means of the P1 and S1 populations within
each planting to determine whether added inbreeding significantly af-
fected the mean performance of plant traits in this study. Comparisons
were also made for cultivar variance across and within inbreeding levels
to provide a measure of parental homozygosity assuming some dominant
gene action was present. Cultivars with low variances: possess greater
homozygosity and phenotypic stability than do cultivars with large
variances. Homozygous parents provide greater precision in estimating
Segregation ratios and environmental variance from F1 and F2 data.
These tests provided a way of determining whether added inbreeding.

as previouslypracticed (33). was actually required for this study.
This on is shown in Table 4. I I

18

Table 3. Dependent variables used in this study.

A

 

 

* Units of Variable

Dependent

Variable . Measure Code

1. Days from transplanting Days Maturity
to harvest

2. Head weight Kilograms Head Wt

3. Total non-wrapper Kilograms Total Leaf Wt
leaf weight

4. Stalk size Kilograms Stalk Size

5. Non-wrapper leaf Leaves Leaf No.
number

6. Efficiency Index E. I.
(3 + 4) / (2)

7. Non-wrapper leaf size Kilograms ' Leaf Size

 

19

Table 4. Analysis of variance for cultivar. inbreeding and planting

effects. Yhijkl =‘u + 0i + 83 + (a8)ij + Yk + (“7)ik +

(37)jk + (oBTI1Jk + Rh T ehijkl : Zhijk will denote plot

means in the k-th planting with a common to all observations.

a1 is the effect of cultivar. 83 is the effect of inbreeding.

”k is the effect of planting and Rh is the effect of the h

replication. The term ehijk denotes error.

 

 

 

 

Source .Df SS MS .F
2
Total 60 2222 Z
hijk hijk
2
CF T Z. . ..
(3)(5)(2)(2)
z z2
Replication (R) 2 h h... - CF SSR/2
2 z2
Cultivar (C) 4 1 .i.. - CF SSC/4 MSC/MSE
(3)(2)(2)
z: 22
Inbreeding (I) I j .. . - CF 551/] MSI/MSE
3 5 2
z}: 22.
(C) x (I) 4 13 .1 -CF - SSC SSCI/4 MSCI/MSE
3 —SSI
I: z2
Planting (P) I .k - CF SSP/l MSP/HSE

nah
.

um,

Table 4. (cont'd)

20

 

 

Source Of SS MS F
2

22 Z.i.k - CF - ssc SSCP/4 MSCP/MSE
(c) x (P) 4 ik 137T?) - SSP
(I) x (P) l ‘2 22 'k - or - SSI. SSIP/l MSIP/MSE

3" (SITE) - SSP

222 z2
(c:) x (P) x (I) 4 i.k .1 k - cr - ssc SSCPI/4 MSCPI/MSE

, J - ssr - SSP
- ssc1 - SSCP - SSIP

Error 38 SST - cr - ssc - ssr SSE/38

- ssc1 - SSCP - SSIP

- SSP - SSCPI

 

‘4

.L.‘

21

 

ParentalI Midparent2 F1 and F., Mean Analysis

For each trait studied, reciprocal F1 and F2 population means were
compared to their appropriate parent and midparent means to determine
maternal inheritance. additive and dominant gene action. degree and di-
rection of dominance and the presence of abnormal gametic transmission.
Evidence of maternal inheritance was noted where reciprocal F1 population
means significantly differed. The presence of either additive or domi-
nant gene action was determined by comparing the reciprocal F1 means with
the appropriate midparent mean. The larger the difference between each
F1 and midparent mean. the greater the action of dominant genes. The
closer each F1 and midparent mean were. the greater the action of addi-
ti ve genes. The direction of dominance is toward the parent closest to
the F1 mean. Abnormal gametic transmission can be Shown if differences

exist between the F1 and F2 means.

Variance Component Analysis

Analysis was conducted on data from 3 plantings of the F1 popula-
tion. Although attempts were made to have each F1 plot mean represented
by an equal number of plants. poor seed production on some crosses and
field losses due to inclement weather in 1975 contributed non-orthogonal
data. ’Initial analysis of the means for F1 data showed large differences
between planting time. reciprocal crosses and female parents. Appro-
Prlfite AOV tests were made to find whether these differences were signi-
ficant. Based on these tests. the data were separated by planting time.
"Cipr‘ocal cross and female parent to provide 12 unique data groups each
"Presenting 3 F1 crosses and 3 replications (Table 5). Within each

data group. F1 plants were related as paternal half sibs. Variance

22

mcopaauwpqoc ms

 

usage mama

 

 

 

 

 

 

 

N. __ o_ m m A e m e m N F
m m m m m m m m m m m m Assam eeaeee_eo
m m m m m m m m m m m . m .epmm_~ .H.a
m m m m m m m m m m m .m eaeep.am eemeem
emaeeo new: emceeo new: emeeee one: emeeee ems: serene new: emeeee emu: e_ez
ea: seem ace seem ewe heme eta seem ,eee Anne ewe Anne _ee_m_ec
mpg—wu—
Peewmemm
waxy
pauoeawumm Pouoenwomm .euoeapumm mmocu
mung xpgmm one;
meme m~m_ seep ame_eeepe

 

 

 

 

 

.mzowumup_aoe ages» an mammoee emcee .aeeaee_c.aeEe as esneeeee
53 250.5 3.2... 35 0303—. .mwmA—oeo 95:09.8 35.—Lg g8 203233. 33. F. .m ~32.

 

23
component analysis was completed for each of the 12 data groups using
one AOV based on unbalanced plot means and a second AOV based on indivi-
dual data (1. 11). The variance components of this analysis are shown
in Table 6. The first analysis provided estimates of paternal and be-
tween-plot variance. The second analysis provided a combined estimate
of 3/4 additive genetic variance and environmental variance between
full sib related individuals in the same plot. .Use of the paternal var-
iance and the variance from the second analysis provided estimates of

herdtability based on paternal half sib data.

Covariance. Correlation Analysis

Covariance and genetic correlations were calculated by taking the
data for 7 traits and adding traits together in combinations of 2 with-
out replacement to provide 18 new "traits" (Table 7). These-new traits
were analyzed using variance component AOV #1. Results of the AOV #1
'tests for the original and combined traits (Table 8) were substituted
into appropriate formula (11) to provide estimates of genetic correla-
tion and covariance between traits. These formulas were derived from
the following assumptions: '

1) Expectations from Table 8 are:

E(Vu) + ogu + rag.u
£(vy) = azy+ rdgy
E(vz) ' 0:2 + "“32
as.) - oi.

 

24

 

 

 

 

. w see + No A_-zVAP-mv\aau am:-am¢-eu-ame A_-:VA_-¢V p eeeem
a
”be + use: + we F-z\aaz to - .WNea _-= ae_e=
~-¢\ame to - a. Na“ _-¢ eeeeee__as¢
N
,zu _ au
NN
2 .3
.wNww szagv peace
mzu m: mm mo ouczom
Pa >o<

uopn we mm: monococ nFN." gm + mm + _a + n u 8m?» .aumv Pm Low awn»

.m_m»~m=m m.:» cL memos
page ucozonsoo oueewge> .m m_amh

25

up: we

1 _ . i. n x: «Lug:
.uopa can mpoaup>wvcp we» mo cams baggage; on» we vauocn_uoa .A p NNVAzmxpv «

 

 

 

 

2..
. Hp xnp
.n_ . amp -....e mee_a eeee_:
we szamvu....:\mm3 ~>ww . ~>www AzVAav
Er
$25358 - flaw 2-25.: 32.. e858
mussom
mm m: 8 .3
«a >o<

~u.»eeev .m a_ee»

 

26

Table 7. Seventeen trait combinations generated by adding data
on one trait to data for another trait from the same
cabbage head.

 

 

 

Original Original New
Trait Trait Trait
Maturity Head Ht. Ul
Maturity Total Leaf Ht. ‘UZ
Maturity Stalk Size U3
Maturity Leaf Num. U4
Maturity E. I.* ' U5
Maturity Leaf Size U6
Head Ht. Total Leaf Ht. U7
Head Ht. Stalk Size U8
Head Mt. Leaf Num. - U9
Head Ht. * Leaf Size U10
Leaf Wt. Stalk Size U11
Leaf Wt. Leaf Num. U12
Leaf Ht. Leaf Size U13
Stalk Size Leaf Num. U14
Stalk Size Leaf Size U15
Leaf Num. E. I. U16
Leaf Mum. Leaf Size U17

 

* Efficiency index

27

 

 

 

ANN-»W-=MVN\_ 1 Na“ am Nu 5” A_-=VA_-¢V . Loaam
A~>-»>-=>v~\F u N», => ~> », _ A_-¢v nape:
~+xu= ~ a
“Page . «Page ovate
venues; coo: mwgmaam saw: . mu . mugaom

 

 

.F* >o< soc» mwmxpmcm coppapugsoo new mocupsm>oo opuoeow .m «pane

2)

3)

4)

28

g 2
E(Ey) oey

- 2
E(Ez) - oez

Therefore variance estimates were as follows:

A) 0 3y = 1/r (Vy-Ey)

a) o :2 = 1/4 (Vz-Ez)

C) a m = 1./r (Vyz-Eyz)
D) a g, = Ey

a a; = a:

F) Gleyz = Eyz

which produces an estimate of genOtypic correlation:

s ogyz C

2—T__ ..
pgyz (ass! ”92) V2 ' NAME)-

 

And an estimate of the error correlation of y and Z:

a 2 06 2 F
.pgyz hey aez) ”2 ' ND) (E)

 

29
F2 DATA AMALXSIS:

1 Individual F2 frequency classes were produced by grouping small
segments of F2 data. Since most data occurred in frequency groups,
i.e., 1.1.1; 0.0; 4.3.4; etc.. groups with similar numbers could be
pooled. i.e., 1. 0. 4. This pooling technique showed that a majority
of groups were pooled over a fixed number of quantitative units. F2
frequency distributions were redefined by dividing the range of data
into groups of fixed length. This system provided groupings based
on units of 0.49. l. 2. 4. etc. Estimates of the number of genes in-
volved and the location of phenotypic boundaries were provided using
techniques developed by Powers et a1. (23) and outlined by Leonard
et a1. (15). An estimate of the number of genes involved was provided
by sequentially summing and comparing frequency percentages from the re-
cessive parent and the F2 distribution. The formula (Fe/P1) (100) was
used. Estimates of 6.25 suggest a 2 gene model while a single gene
model might produce a result of 25.0. Sequential results usually fluc-
tuate around a given percentage. i.e., 25.0. or 6.25. A sudden increase
in the formula estimate suggests the boundary for the recessive phenotype
has been passed. Cultivar, F1, and backcross means were also used in
defining class boundaries. Theoretical ratios for F2 populations were
calculated normally if parental distributions did not overlap. Over-
lapping cultivar distributions were considered in the calculation of
theoretical F2 ratios by the use of a weighting factor. The weighting
factor was calculated by subtracting the absolute value of the net over-
lap,.found by subtracting the percentage of misclassified plants of 1
cultivar from the percentage misclassified from the other cultivar, from

100. This factor was divided by 100. multiplied by the expected frequency

30

of the deficient class and by the total number of F2 plants in the
sample to produce an expected ratio corrected for F2 misclassification
and cultivar overlap (17). Segregating F2 populations were tested for
heterogeneity prior to pooling the data for comparisons and inter-
pretation. Chi-square tests were used to compare observed and theoreti-
cal ratios. In all cases a chi-square significance level of (P = 0.05)
was used. The probability of rejecting a genetic model when it was cor-
rect was 5 per cent. Backcross ratios were also tested and used to

support the postulated models.

RESULTS AND DISCUSSION

Populations in this study have been divided into 2 groups which
include crosses between red and green cabbage and crosses between 2
green cabbage cultivars. This division was necessary since both
groups differed in dominance direction for certain traits and showed
further differences in other areas. The results and discussionwhich

follow will show how and in what areas these 2 groups differed.

Maturity
One generation of inbreeding did not affect the maturity of the

cultivars; however. significant differences were found between culti-
vars. plantings and the interaction cultivar x plantings (Table 9)..
Since no significant differences between inbreeding means were observed

‘ maturity may be genetically fixed or dominance may be absent.. The sig-
nificant cultivar x plantings interaction was caused, in hart. by differ-
ent responses to the environment in the 1975 early (E) planting. Two
cultivars. Baby Head and Badger Ballhead matured earlier in 19755 than

in the 1974 late (L) planting. The other 3 cultivars took longer to
mature in the 1975L planting than in 1974L. Sequential plantings of
Badger Ballhead should provide an orderly sequential harvest since its
maturity is fairly constant for various planting dates. (Baby Head ap-
pears to mature earlier when planted early in the season. If sequential-
ly planted this cultivar may produce a prolonged harvest period. The

other cultivars seem to mature earlier with later planting dates (Table 10).
' 31 i

Table 9. Analysis of variance table for cabbage maturity in days
from transplanting to harvest.

 

 

 

 

Source
of Mean
Variance DF Square F*

Replication (R) 2 ‘ 14.16 0.42
**

Cultivar (C) 4 3190.12 94.22

Inbreeding (I) l 75.01 2.22

c X I 4 65.01 1.92
**

Plantings (P) 2 1413.20 41.74

c x P 8 848.40 25.06"

I x P 2 6.36 0.19

C x I x P 8_ 11.34 0.33

Residual Error 58 33.86

* . * **

F test significant at the 5% . or 1% level.

33

 

 

 

 

Table 10. Cultivar x planting interaction means in days from
transplanting to harvest.

. Plantings Cultivar
Cultivar 1974L 1975E 1975L Means
Baby Head ,99.73 6* 72.60 a 114.33 c 95.56 v
Badger Ballhead 107.33 b 100.58 a 115.38 c 107.76 M
P.I. 215514 113.82 a 129.52 b 141.96 c 128.11 2
Red Danish 112.25 a 124.37 b 117.70 ab 118.11 x
Chieftain Savoy 116.67 a 132.42 c _124.13 b 124.41 Y
Planting Means 109.96 A 111.90 A 122.70 B

 

*Mean separation within rows by Duncan's Multiple Range Test at

the 5% level.

34 .
Early maturity has been reported by Pearson (20) and Rasmusson

(24) as dominant. Both the 1975E and 1975L Baby Head x Badger Ball-
head F1 means suggest early maturity is dominant (Tables 11-13).

Tables 14 and 15 show frequency distributions in percent for populations
involved in this cross. Powers et a1. (23) partitioning tests were
applied to these distributions to determine the number of genes control-
ling maturity; and. to define the boundary between early and late pheno-
typic classes. Powers et a1. (23) formula (F2/P2) was applied sequential-
ly from the recessive ends of both the F2 and P2 distributions. Using
the 19755 F2 planting of Baby Head x Badger Ballhead (Table 14) as an
example. the formula suggested recessive frequencies of 10.4/0 for the
144 day class. 16.4/0 for the 126 day class. 20.4/92.3 or 0.22 for the
104 day class and 53.7/92.3 or 0.58 for the 96 day class. This fre-
quency. 0.22. exhibited close agreement to an expected frequency of 0.25
from a 3:1 model. A monogenic model with a dividing point between the
96 and 104 day classes was proposed (Table 14). The 1975E reciprocal
F2 population produced a different recessive frequency estimate. As in
the previous example. the formula (F2/P2) suggested frequencies of
13.3/0 for the 144 day class. 14.9/0 for the 126 day class. 28.6192.3
or 0.31 for the 104 day class and 36.7/92.3 or 0.40 for the 96 day
class. The 88 day class produced an estimate of 86.7/92.3 or 0.94. A
dividing point between the 88 and 96 day classes seems apparent. The
ovserved frequency of 0.40 shows close agreement to the 0.44 frequency
expected from a 9:7 model (Table 14). Since Powers et a1. (23) tests
suggest a 2 locus model for other F2 populations in this study.a mul-
Itiple allelic 2 locus system was tested for each population. The pose
tulated loci. a and 0. each have 5 alleles which were inherited in the

following manner:- for the 1975E planting ‘5 was dominant to a and

35
Table 11. 1974L Cultivar and F1 performance means.

 

 

 

 

 

Cultivar Maturity Head Leaf Stalk. Leaf E.I:* Leaf
Ht. Wt. Size Mum. Size
Days kg. kg. kg. Leaves kg. kg.
Baby Head 99.08 1.14 0.23 0.07 8.25 0.33 0.03
2. Badger 106.96 2.25 1.79 0.28 18.28 1.12 0.10
Ballhead .
3. P.I. 215514 113.00 1.74 2.35 0.60 24.00 1.68 0.10
4. Red Danish 108.22 1.26 1.85 0.28 14.80 2.01' 0.12
5. Chieftain 116.32 1.55 2.59 0.34 20.30 2.20 0.13
Savoy *
F1
1 x 2 99.34 2.22 0.65 0.11 11.96 0.35 0.06
2 x 1 98.39 2.34 0.62 0.11 11.81 0.32 0.05
1 x 3 106.07 3.20 1.34 0.28 13.57 0.51 0.10
3 x 1 103.74 2.98 1.24 0.27 13.45 0.56 0.09
1 x 5 106.60 2.55 1.55 0.26 18.18 0.87 .0.08
5 x 1 104.60 2.38 1.39 0.21 18.21 0.76 0.08
4 x 2 106 60 1.24 2.26 0.42 20.72 2.90 0.11
2 x 4 107.74 1.26 2.16 0.43 20.28 2.62 0.11
4 x 3 119.55 0.99 2.70 0.62 20 72 4.34 0.13
3 x 4 114.03 1.51 3.07 0.70 21.16 2.99 0.14
4 x 5 117.14 1.14 2.73 0.49 20.46 3.56 0.13
5 x 4 112.68» 1.42 2.98 0.49 21.39 2.96 0.14

 

*Efficiency Index

36

Table 12. 1975E Cultivar. F1 and F2 performance means.

 

 

Maturity

Leaf E.I.*

 

 

 

 

Cultivar Head Leaf Stalk Leaf
Ht. Ht. Size Num. Size
Days kg. kg. kg. Leaves kg. kg.
1. Baby Head 69.36 0.70 0.38 0.09 14.19 0.71 0.03
2. Badger 99.38 3.09 1.23 0.26 15.85 0.59 0.08
Ballhead
3. P.I. 215514 129.67 3.29 2.21 0.61 23.78 0.96 _ 0.09
4. Red Danish 121.88 2.15 1.44 0.38 16.43 0.90 0.09
5. Chieftain. 135.94 2.95 1.17 0.32 16.94 0.56 0.07
Savoy
F1 .
1 x 2 74.93 1.33 0.76 0.14 16.57 0.74 0.05
2 x 1 ,74.92 1.41 0.71 0.12 16.08 0.65 0.04
1 x 3 95.62 2.99 1.03 0.23 14.12 0.46 0.07
3 x 1 97.33 3.28 1.12 0.23 13.88 0.43 0.08
1 x 5 115.48 2.73 0.78 , 0.24 12.52 0.48 0.07
5 x 1 120.25 3.37 0.86 0.28 11.52 0.36 0.07
4 x 2 99.21 2.89 1.68 0.34 17.20 0.79 0.10
2 x 4 99.00 2.38 1.68 0.42 18.29 1.00 0.09
4 x 3 112.57 2.91 2.21 0.62 19.65 1.12 0.11
3 x.4 104.89 2.75 2.48 0.68 21.06 1.27 .0.12
4 x 5 122.33 3.80 2.27 0.62 19.09 0.90 0.12
5 x 4 123.02 3.36 2.15 0.60 19.68 0.91 0.11
F2
1 X 2 96.04 2.05 0.73 0.18 13.73 0.51 0.05
2 X 1 94.68 1.44 0.55 0.13 12.84 0.56 0.04
1 X 3 105.02 2.12 0.78 0.22 13.38 0.49 0.06
3 X 1 106.96 2.76 1.00 0.27 13.26 0.47 0.07
1 X 5 119.48 2.51 0.71 0.22 11.39 0.38 0.06
5 X 1 117.39 2.38 0.73 0.23 14.57 0.45 0.05
4 X 2 123.84 2.34 1.45 0.45 19.24 0.91 0.08
2 X 4 119.21 2.33 1.46 0.41 19.03 0.96 0.08
4 X 3 - - - - - - -
3 X.4 122.92 2.65 2.12 0.69 21.49 1.34 0.10
4 X 5 126.11 2.74 1.97 0.58 20.13 1.08 -0.10
5 X 4 131.46 2.56 2.28 0.68 21.46 . 1.47 0.11

 

”Efficiency Index

1. £111
551’:
\

 

37
Table 13. 1975L Cultivar. F1 and F2 performance means.

 

 

 

 

 

 

Cultivar Maturity Head Leaf Stalk Leaf [.13' Leaf
Ht. Ht. Size Num. Size
Days kg. kg. kg. Leaves kg. kg.
Baby Head 113.23 0.84 0.06 0.07 3.08 0.17 0.02
2. Badger 114.26 2.12 0.43 0.21 10.14 0.35 0.04-
Ballhead
3. P.I. 215514 142.00 3.60 1.96 0.69 19.23 0.87 0.10
4. Red Danish 114.69 1.12 0.64 0.19 11.12 0.90 0.05
5. Chieftain 127.51 1.24 0.65 0.23 15.14 0.84 0.04‘
Savoy
F1
1 x 2 112.79 1.37 0.35 0.11 9.55 0.33 0.03
2 x 1 114.88 1.35 0.12. 0.15 3.87 0.35 0.03
1 x 3 114.70 1.79 0.29 0.21 7.88 0.30 0.04
3 x 1 114.49 2.06 0.37 0.25 8.48 0.32 0.04
1 x 5 138.25 2.24 0.37 0.18 9.80 0.32 0.04
5 x 1 121.74 2.03 0.25 0.16 7.24 0.24 0.03
4 x 2 112.48 1.42 0.52 0.26 12.74 0.84 0.04
2 x 4 115.84 1.00 0.53 0.23 13.42 1.09 0.04
4 x 3 121.00 1.62 0.83 0.47 15.00 1.34 0.05
3 x 4 125.14 0.94 1.13 0.37 20.25 1.98 0.06
4 x 5 112.46 1.67 0.92 0.26 9.32 0.84 0.10
5 x 4 112.30 1.79 0.89 0.26 8.57 0.70 0.10
F2
1 x 2 114.31 1.27 0.17 0.13 5.82 0.27 0.03
2 x 1 114.08 1.07 0.17 0.11 5.94 0.29 0.03
1 x 3 116.43 1.10 0.36 0.19 9.98 0.66 0.04
3 x 1 114.62 1.45 0.33 - 0.19 8.68 0.44 0.04
1 x 5 114.79 1.56 0.26 0.16 7.40 0.30 0.03
5 x 1 119.98 1.23 0.26 0.13 8.70 0.41 0.03
4 x 2 116.30 0.99 0.65 0.26 14.43 1.29 0.04
2 x 4 113.80 1.06 0.67 0.32 13.60 1.04 0.04
4 x 3 - - - - - - -
3 x 4 - - - - - - -.
4 x 5 125.71 1.11 0.86 0.33 17.54 1.54 0.05
5 x 4 121.28 1.00 0.94 0.33 18.19 1.73 0.05

 

*Efficiency Index

.Oav Ion-l larva-a. uaa\nv- 0.... zone. don-o i.eut-c as. stable. H‘s-c... hat-nu \ld--h!-tl\-I nu}n\.I I,‘ II\P‘ FiaII‘ Q . \allllll‘a‘ .1! !\§-\Ci,h.

38

 

 

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m. H m. m... 3 New NS 8 .me x ....
a. H a . Sm ... an 3: 685.8 .688
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e8: 3. a. g. 8 8 8 N. 85.... .
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39

 

H 5... ...... .... 5... 5.5.. 5.3 5:5. 5.5. ..N N. .... x N....
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... H 5.. 5.... ....N 5... 2.... an .N. x ....
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555: 5m. 55. 5.. 5.. 5.. 5.. 5.. ~.. ... 5.. “wemwn

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40

A was dominant to a3 and a4. In 19755. red x green cabbage crosses A

was dominant to A2 and A2 was dominant to a3 and 34. The D locus was

5

inherited in the Same manner as the a locus. Allelic inheritance pat-

terns for the 1975L planting suggest A was dominant to a and a5

3’ a4

for green cabbage crosses. In red x green cabbage crosses A was domi-

2
nant to a3. a4 and as. The b locus was inherited in a similar manner.
Postulated genotypes for each cultivar and Pi are shown in Tables 16
and 17. In cases where parental distributions overlap. a weighting fac-
tor was used to weight the expected segregation ratio. The 19755 plant-
ing of Baby Head x Badger Ballhead will again be used as an example
(Table 14). For a division point of 96 days. the parental distributions
suggest that Badger Ballhead produces less than 100% of its plants in
the same phenotypic class. This class. lateness will be deficient when
F2 segregation ratios are obtained. In this study. the net overlap.
7.7%. was subtracted from 100%. divided by 100 and multiplied by the ex-
pected probability of the deficient class and the total number of 52
plants sampled to obtain an expected segregation ratio. The number of
recessive plants was ((100-7.7)/100)(0.2s)(252) = 58.15. The number

of dominant plants was found by subtraction. (252-58.15) = 193.85. The
observed 52 segregation ratio of 201:51 for the 19755 planting of Baby
Head x Badger Ballhead fits (P=D.23) the expected 193.85 early maturity
: 58.14 late maturity expected from a weighted 3:1 segregation ratio
(Table 18). The 3:1 segregation ratio suggests that the presence of a.
single A allele in the genotype produces an early phenotype. The
dominant A allele was contributed by Baby Head (AABB) while the recessive
genotype a3a3b3b3 was contributed by Badger Ballhead. The reciprocal

52 populations were significantly different. The observed ratio of 157

41

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42

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43

Tab1e 18. Chi-square test for goodness of fit to the postulated
mode1 for maturity in the F2 generation (1975E p1anting).

 

 

 

 

 

 

 

 

Cross Observed Expected Hode1 X2 P
Baby Head x
Badger Ba11head

*
.F2 201: 51 193.85: 58.15* 3: 1.4400 0.23
RFZ 157: 91 147.85:100.15 9: 1.4034 0.24 .
Baby Head x
P.I. 215514
F2 197: 62 194.25: 64.75 3: 0.1557 0.69
RF2 331:120 338.25:112.75 3: 0.6216 0.43
Poo1ed 528:182 532.50:177.50 3: 0.1521 0.70
Baby Head X
Chieftain Savoy
F2 45:147 48.00:144.00 1: 0.2500 0.64
RF2 61:136 49.25:147.75 1: 3.7377 0.05
Red Danish x
Badger Ba11head
F2 51:158 52.25:156.75 1: 0.0399 0.84
RF2 51:103 38.60:115.50 1: 5.4112 0.02
Poo1ed 102:261 90.75:272.25 1: 1.8595 0.17
Red Danish x
P.I. 215514
F2 150:161 144.39:166.61 9: 0.4064 0.52
Red Danish x
Chieftain Savoy
F2 68: 39 60.19: 46.81 9: 2.3179 0.13
RF2 50: 52 57.38: 44.62 9: 2.1668 0.14
Poo1ed 118: 91 117.56: 91.44 9: 0.0037 0.25

 

*
= Weighted

44
early maturity: 91 late maturity fit (P=0.24) a weighted 9:7 expected
ratio of 148:100 (Table 18). A segregation ratio of 9:7 suggests that
the presence of l dominant allele at each locus. A or 8 conditions an
early phenotype. The dominant A and B alleles were contributed by Baby
Head while the recessive a3 and b3 alleles came from Badger Ballhead.
Backcrossing the reciprocal F1 to the recessive cultivar. Badger Ball-
head produced a 5:28 ratio. This ratio fit (P=0.21) a 1:3 model with
an expected ratio of 8.25:24.75 (Table 14). The Observed data fit the
postulated model. Following a chi-square test for homogeneity. both
1975L F2 populations of Baby Head x Badger Ballhead were pOoled. Using
a dividing point of 116 days, calculated as illustrated (23). an observed
ratio of 276 early to 62 late gave a good fit (P-O.79) to a 13:3 model
(Table 19). A segregation ratio of 13:3 results when a dominant geno-
type at l locus. A and the recessive genotype at the other locus, b3b3
produce the same phenotype. earliness. The dominant A allele was cdntri-
buted by Baby Head while the recessive b3 allele was contributed by
Badger Ballhead. The genotype a3a335 conditions lateness (Table 15).

F1 populations of Baby Head x P.I. 215514 show early maturity to
be dominant in both the 1975E and 1975L plantings. Since the 1974L mid-
parent and F2 means were similar, additive inheritance for maturity in
the planting seems apparent (Tables 11-13). Within each planting F2
populations of Baby Head x P.I. 215514 were pooled as suggested by chi-
square homogeneity tests. The dividing points for the 1975E and 1975L
plantings were 120 and 119 days respectively. These points were calcu-
lated as previously explained (23). An observed ratio of 528:182 for
the 1975E planting gave a good fit (P=0.70) to a 3:1 model of 532:178-
(Table 18). The 3:1 model suggests that the presence of 1 dominant

45

Table 19. Chi-square test for goodness of fit to the postulated
model for maturity in the F2 generation (1975L planting).

_A

 

 

 

 

 

 

Cross Observed Expected Model ‘ X P
Baby Head x
Badger Ba11head
F2 103: 27 105.62: 24.38 13: 3 ' 0.3479 0.56 -
RF2 176: 35 171.44: 39.56 13: 3 0.6476 0.42
Pooled _279: 62 277.06: 63.94 13: 3 0.0722 0.79
Baby Head X
P.I. 215514
F2 148: 20 157.50: 10.50 15: 1 9.1682 <:0.01
RF1 229: 15 228.75: 15.25 15: 1 0.0044 0.95
Pooled 377: 35 . 386.25: 25.75 15: 1 3.5442 0.06
Baby Head x
Chieftain Savoy

*
F2 143: 72 136.25: 78.75* 7: 9 0.9126 0.34
RF2 93: 85 91.07: 86.93 1: 3 0.0837 0.77
Red Danish x
Badger Ba11head

*
F2 105:186 92.52:19B.48* 1: 3 2.4702 0.12 -
RF2 14: 16 9.54: 20.46* 1: 3 3.0610 0.08
Pooled 119:202 102.05:218.95 l: 3 4.1262 * 0.04
Red Danish x
Chieftain Savoy

*
F2 34:165 39.12:159.88* 1: 3 0.8342 0.36
RF2 42:182 44.03:l79.96* 1: 3 0.1170 ' 0.73
Pooled ’76:347 83.16:339.84 1: 3 0.7663 ' 0.38

 

*
a Heighted

. 46
A allele in the genotype conditions an early phenotype. The dominant A

allele was contributed by Baby Head and the recessive a4 and b4 alleles
were contributed by P.I. 215514 (Table 20). The pooled F2 population
from the 1975L planting produced a 377:35 segregation ratio which fit
(Ps0.06) a 15:1 model (Table 19). A 15:1 segregation ratio suggests
that only 1 dominant allele. A or B . was required to produce an early
phenotype. Only the recessive genotype a4a4bhb2 from P.I. 215514 pro-
duced a late phenotype (Table 21).

Planting time influenced the F1 performance of Baby Head x Chief-
tain Savoy. The 1974L planting data show maturity to be inherited
additively with an F1 mean of 106 and a midparent mean of 107. How-
ever. both the 1975E and 1975L plantings show incomplete dominance for
lateness (Tables 11-13). The 1975E F2 division point was shown to be
116 days while the reciprocal F2 division point was 104 days (23). The
separation of classes for the 1975L F2 planting of Babthead x Chief-
tain Savoy was at 115 days. The 19755 F2 population of Baby Head x
Chieftain Savoy produced a segregation ratio of 45:147. while the re-
ciprocal F2 population produced a ratio of 61:136. Both populations
fit (P-0.64. 0.05) a 1:3 model (Table 18). In each population. the 1:3
segregation ratio suggests that the presence of the b5 allele in the
genotype conditions lateness regardless of the condition of the alocus.
Baby Head contributed AABB and Chieftain Savoy contributedA ”A:

The genotypes A5 Asbb and aabb condition earliness (Table 22). Back-
crosses to the dominant cultivar. Chieftain Savoy. fit the 0:1 model.
Backcrossing the reciprocal F1 to the recessive cultivar produced a ratio
of 9: 46. This ratio showed poor fit (P<0. 005) to the expected 1:1 ratio
(Table 22). Due to an overlap of the 1975L cultivar distributions.a

47

 

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e .H ea e._ . e.wm o.oe om ape x may
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48

 

 

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49

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50
weighting factor was used as previously described (23) to obtain a

weighted theoretical segregation ratio for the 1975L F2 populations.
The F2 population produced a 143:72 ratio which fit (P=0.34) a 7:9
weighted theoretical ratio. An F2 ratio.of 7:9 suggests that the pre-
sence of 1 or more homozygous recessive loci in the F2 genotype condi-
tions early maturity. The recessive alleles. AABB. were contributed by

Baby Head, and the dominant alleles. ASA were contributed by Chief-

55535 '

tain Savoy. The genotype A conditions a late phenotype (Table 23).

5—95—
The reciprocal F2 population produced a ratio of 93:85 which showed good
fit (P-0.77) to a weighted 1:3 ratio (Table 19). The 1:3 segregation
ratio suggests that the presence of the 35 allele in the genotype condi-
tions lateness regardless of the condition of the a locus. 'The geno-

types ASA bb and AABB condition earliness (Table 23).

5
The mean performance of 6 red x green cabbage crosses also show
dominance for earliness; however, the environment greatly affected the
performance estimates. Both the 1974L and 1975L F1 population means
of Red Danish x Badger Ballhead show maturity to be additively inherited;
however. the 1975E F1 population mean shows early maturity to be domi-
nant (Tables 11-13). The 1975E F2 populations of Red Danish x Badger
Ballhead were pooled. The division point of 124 days was calculated as
suggested by Powers et a1. (23). The pooled segregation ratio of
102:261 agreed (P-0.17) with the expected 1:3 ratio of 91:272 (Table 18).
The 1:3 segregation ratio suggests that the presence of a single.A2 allele
in the genotype was sufficient to condition a late phenotype irrespec-
tive of the genotypic make up of the b locus. Red Danish contributed
dominant alleles 52523232 for lateness while Badger Ballhead contributed

recessive genes a3a3b§b§ for earliness (Table 24). Due to a limited

51

 

 

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52
nunber of backcross plants. the backcross data suggest different models.

Backcrossing the F1 to the recessive parent. Badger Ballhead..produced
a 0:12 ratio while backcrossing the reciprocalfl to the dominant par-
ent, Red Danish. produced a 14:33 ratio. Although Badger Ballhead was
dominant in the F1 no F2 plants with the F1 phenotype were recovered.
Chi-square tests suggest the 1975L F2 populations should not be pooled.
Using a dividing point of 112 days (23) the 1975L F2 population produced
a segregation ratio of 105:186 which fit (P-0.12) a 1:3 weighted theo-
retical ratio of 93:198 (Table 19). The reciprocal F2 population also
fit (P80.08) a 1:3 weighted theoretical segregation ratio (Table 19).
In both plantings the 1:3 segregation ratio suggests that the A? allele
was completely cominant to the e3 allele and masks the action of the b
locus. The recessive genotype a3a3 conditions earliness (Table 25).
The F1 population means of Red Danish x P.I. 215514 were affected
by the environment. Both the 1975E and 1975L population means show i
early maturity to be completely to incompletely dominant. The 1974L
planting shows complete dominance for late maturity (Tables 11-13).
The Red Danish x P.I. 215514 F2 population was only grown in the 1975E
planting due to short seed supply. Using a dividing point of 124 days
(23) the F2 produced a segregation ratio of 150:161. This ratio fit
(P20.52) a 9:7 weighted segregation ratio (Table 18). The 9:7 segrega-
tion ratio suggests that homozygous recessive genotypes at either loci
condition lateness. Dominant alleles for earliness. A2A23282 were con-
tributed by Red Danish while recessive alleles for lateness. a4a4bgbz.
were contributed by P.I. 215514. The genotype A2_§2_ conditioned
earliness (Table 26).

53

 

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55

 

 

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56
Both the 1975E and 1975L F1 population means of Red Danish x

Chieftain Savoy show complete dominance for early maturity while the
1974L planting shows dominance for late maturity (Tables 11-13). The
1975E F2 populations of Red Danish x Chieftain Savoy were pooled.
A dividing point of 128 days (23) produced a pooled ratio of 118 early
maturity:91 late maturity (Table 28). The data showed a good fit
(P-0.95) to a 9:7 segregation ratio (Table 18). An F2 segregation ratio
of 9:7 suggests that homozygous recessive genotypes at either loci con-
dition lateness. Dominant alleles for earliness. Aznzazaz. were contri-
buted by Red Danish while recessive alleles for lateness. asasbsbs. were
contributed by Chieftain Savoy. The genotype.A?_p2__conditioned
earliness (Table 18). Backcrosses to the dominant Red Danish parent fit
the expected 1:0 ratio and support the postulated model. The A 2__a2_
genotype was dominant for lateness in the 1975L planting (Table 29).
The 1975L F2 data were pooled and a dividing pointof 114 days (23)
produced an obserVed segregation ratio of 76:347. This ratio fit
((P-0.38) a 1:3 weighted theoretical ratio of 83:340 (Table 29). The
1:3 segregation ratio suggests that the presence of 1 allele from Red
Danish masked the action of the b locus and conditioned a late pheno-
type. The recessive genotype a5a5b5b5 conditioned an early phenotype
(Table 19).

These F2 populations show that in all but one case. Baby Head pro-
vided dominant genes for earliness. In red‘x green cabbage crosses.
the 1975E planting data showed Red Danish phenotypes to be dominant:
however. in the 1975L planting Red Danish genes were dominant only if
their phenotype was later than the other parental phenotype.

 

57

 

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59

 

 

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60

Swarup (30) and Chiang (3) have previously estimated herit-
abilities for maturity to be 0.80 and 0.83 respectively. This study
shows heritability estimates of some of the traits to be greater than
1 or less than 0 (Table 30). Heritability estimates greater than 1
may be produced when 05‘302. Since residual error mean square for
ADV #1 (Table 6) is equal to a: + nk as. small values for as may
occur when “E: the environmental variance of the differences between
plots common to all individuals within a plot. is large. .Large herita-

E. the covariance of paternal

2 2 2
w or “c + nk w' Robinson et al.

(25) reported negative heritabilities in corn and suggested these es-

bility estimates may also occur when a

half sib data. is large relative to a

timates might arise by chance. Thompson and Moore (34) suggest nega-
tive estimates of variance components. used in estimating heritability.
may occur because of sampling error. Gill and Jensen (8) have calcu-
lated the probability of obtaining negative heritability estimates. In

this study 52 W111 denote the mean heritability estimate taken from all

2
x

bility estimate of all estimates. where estimates greater than 1 are

2
x

estimate to be included

estimates between 0 and 1. The term 5 will denote the mean herita-

set to 1 and negative estimates are set to 0. Both?2 and S estimates

were calculated in the following manner. Each h2

in a mean estimate was weighted by the factor 1/(S.E. estimate)2.

Weighted estimates were summed and divided by the sum of the various

weights. i.e., 2(h2/(S.E. estimate)2)//2(1/(S.E. estimate)2) - 52 or

2 2
x

Standard errors were calculated as l’l/lx(1/(S.E. estimate)2)1 The B

2
x

estimate was 0.66 i=0.36. Heritability estimates between 0 and 1 ranged

5 depending on the type of h estimates included in the calculation.

 

2

estimate for green cabbage F1 maturity was 0.62 j=0.38 while the S

61

Table 30. Heritability estimates for seven traits from green and
red x green cabbage crosses grown in three plantings.

 

 

w

 

 

 

 

 

 

 

 

 

F.l Cabbage Populations ,
l974L Green Green Red Red
Trait Reciprocal Reciprocal

MatUrity 0.61 0.62 A 8
Head Ht. A 0.89 . B 0.05
Total Leaf Ht. A A 0.17 A
Stalk Size A A 0.57 A
Leaf Num. A A B B
E. I.* 0.96 A 0.15 8
Leaf Size A A 0.40 A
l975E Green Green Red Red
Trait Reciprocal . Reciprocal
Maturity A A A A
Head Ht. A A 0.l7 A
Total Leaf Ht. 0.54 A A A
Stalk Size 0.96 A A A
Leaf Num. 0.67 A 0.42 0.57
E. I. A 0.18 f A 0.56
Leaf Size 0.52 A 0.50 0.70
1975L Green Green Red Red
Trait Reciprocal Reciprocal
Maturity A A A A
Head Ht. 0.86 A B 0.93
Total Leaf Ht. 8 A 0.88 A
Stalk Size 0.78 0.28 A A
Leaf Num. 8 A B A
E. I. B B B A
Leaf Size 8 0.84 A A

 

 

 

*Efficiency Index
A =Estimate greater than 1
B =Estimate less than 0

 

 

 

62

from 0.61 to 0.62. Red x green cabbage Fl's provided 5 estimates of hz

. greater than 1 and 1 negative estimate. The 5: estimate was 1.0 i 0.81.
It appears that red x green cabbage heritability for this trait is large.
The green cabbage cross 52 estimate suggests 62% of the total variance
in these crosses was due to additive genetic variance. This additive
portion may be genetically fixed using inbreeding and selection tech-
niques. The other 38% is dominance. to be used to an advantage in F1
hybrids, and an environmental effect (Table 30).

Genetic correlation (r9) estimates between some traits in this
study were greater than 1 and less than negative 1 (Table 31). Genet-
ic correlation values greater than 1 occur when the covariance be-
tween 2 traits is greater than the square root of the product of 2
trait variances. Genetic correlation values less than -1 can occUr
if the covariance is negative and larger than the square root of the
product of 2 trait variances. The term 59 will be used to denote a
mean genetic correlation of all estimates between +1 and -1 and the
term ng will denote a mean genetic correlation of all estimates
where estimates greater than 1 are set to 1 and estimates less than
negative 1 are set to negative 1. Both the E and the E x estimates

9 9

were calculated by weighting each r estimate by the inverse of its

variance and summing all estimates.9 The final answer was divided by

the sum of the inverse variances to produce a formula similar to that for
52. Genetic correlation variances were calculated using Scheinberg's
method (27). Standard errors for both 59 and ng were calculated

similar to those for 52. Genetic correlations between maturity and other
. above ground plant parts in cabbage have not been previously reported.

Far green cabbage crosses the mean genetic correlation between maturity

63

Table 31. Genetic correlation between maturity and six other
traits in cabbage.

 

 

 

 

 

 

 

 

 

Traits Plantings Green Crosses Red Crosses
R* R*

Maturity x 1974L 0.92 0.37 -0.10 -0.53
Head Height l975E 0.75 0.88 A A

1975L A 0.54 a B
Maturity x l974L A A A B
Total Non-wrapper 19755 -0.28 0.34 A 0.48
Leaf Height 1975L 0.03 0.01 -0.02 0.65
Maturity x 1974L A 0.92 A B
Stalk Size 19755 0.94 0.99 0.95 0.53

1975L 0.10 -0.54 A 0.91
Maturity x l974L 0.91 0.91 0.11 A
Non-wrapper l975E B B A 0.04
Leaf Number 1975L -0.14 0.31 0.45 A
Maturity x 1974L 0.88 A A A
Efficiency 19755 -0.85 a A -0.68
Index 1975L 0.29 0.98 0.23 A
Maturity x 1974L A 0.89 A -0.83
Non-wrapper l975£ 0.70 0.74 A 0.82
Leaf Size 1975L -0.08 -0.44 -0.13 -0.53

 

 

 

 

 

*Reciprocal cross

A 8 Correlation estimates greater than 1.0
B = Correlation estimates less than -1.0

 

64

and head weight was 39 = 0.84 a 0.53 while 5 x . 0.98 a 0.20. Genetic

9 .
correlation estimates ranged from 0.37 to 0.92 (Table 31). Mean genetic
correlation between maturity and total non-wrapper leaf weight was low.

29 = 0.05 t 0.99. however. numerous estimates were larger than 1 show-

ing 3? . 0.39 a 0.34. Estimates ranged from 50.28 to 0.34. The E

x

estimjte between maturity and Stalk size was 0.98 t 0.12 suggestingglate
maturity might be related to regulatory genes controlling stalk size.
Estimates ranged from -0.54 to 0.99. Genetic correlations between
maturity and non-wrapper leaf number. efficiency index and non-wrapper
leaf size were 0.63 i 0.73. 0.11 1 0.54 and 0.41 t 0.42 respectively.
However. ng values were -0.85 1 -0.21. -0.75 a 0.20 and 0.50 a 0.35 re-
spectively. Genetic correlation estimates ranged from -0.14 to 0.91.
-0.85 to 0.98 and -0.44 to 0.89 respectively. Environmental effects on
various :9 values were evident. Correlations between maturity and to-
tal non-wrapper leaf weight in the 1975E planting were -0.28 for crosses
and 0.34 for reciprocal crosses. It is also evident in genetic correla-
tions between maturity and efficiency index where the 1974L. 1975E and

1975L plantings gave r estimates of 0.88. -0.85 and 0.29 respectively.

Red x green cabbage crisses exhibit generally lower genetic correlations
compared to green cabbage crosses. The genetic correlation between
maturity and head weight was negative and low. .Eg--0.20 t 0.81 although
the ng is 0.76 t 0.73. Genetic correlation estimates range from -0.58

to -0.10. Mean genetic correlation estimates between maturity and total
non-wrapper leaf weight. stalk size and non-wrapper leaf number were

0.50 t 0.67. 0.93 t 0.34 and 0.16 t 0.89 while ng values were 0.98 t 0.22.
0.92 i 0.33 and 1.00 t 0.12 respectively. Genetic correlation estimates
ranged from -0.02 to 0.65. 0.53 to 0.95 and 0.04 to 0.45 respectively.

Mean genetic correlations between maturity and efficiency index and

65
non-wrapper leaf size were -0.28 t 0.76 and 0.13 i 0.59. Values from ng
were larger. 0.85 i 0.52 and 0.98 i 0.18 respectively. It appears that
both green and red x green cabbage crosses grown in the 1925L planting
showed negative correlations between maturity and non-wrapper leaf size
although prior plantings produced high positive correlation estimates.
The negative correlation between maturity and head weight for red x green

(cabbage crosses compared to a high positive correlation for green cabbage

crosses suggests different directions of dominance for head weight.

Head Height
Analysis for head weight showed significant differences between

treatment means for cultivar. inbreeding and planting time. Signifi-
cant interactions include cultivar x plantings and cultivar x in-
breeding (Table 32). The cultivar x plantings interaction was sig-
nificant since the head weight of Baby Head showed no increase or de-
crease for the 3 plantings while Badger Ballhead. Red Danish and
Chieftain Savoy all produced significantly heavier heads in the 1975E
planting when compared to the other 2 plantings. P.I. 215514 pro-
duced significantly heavier heads in the 1975L planting than in the
1974L planting. Head weights ranged from 0.78 kilograms for Baby
Head to 2.98 kilograms for P.I. 215514. An increase in head weight
of 1 kilogram or more may be achieved by planting cabbage earlier in'
the growing season (Table 33). The interaction. cultivar_x inbreeding
suggests that 1 generation of inbreeding did not have the same effect
on each cultivar. Inbreeding had no effect on the head weight of 4

cultivars; while. 1 generation of inbreeding a randomly selected plant
of Badger Ballhead produced a significant reduction in head weight

(Table 34). Although inbreeding produced significant differences. the

Table 32. Analysis of variance table for head weight in kilograms.

 

 

‘Source
of Mean

Variance DF Square F*

Replication (R) 2 0.58 '2.89
*‘k‘

Cultivar (C) 4 l2.02 59.89

Inbreeding (I) 1 1.19 5.91*

c x I 4 0.54 2.70'
**

Plantings (P) . 2 7.40 36.88

c x P 8 2.20 10.94”

I x P 2 0.35 1.77

c x I x P 8 0.15 0.73

Residual Error 58 0.20

 

* 'k H
F test significant at the 5% . or l% level.

67

Table 33. Cultivar x planting interaction means for head weight
in kilograms.

 

’-

A

Plantings Cultivar

 

 

Cultivar l974L f l975E l975L Means
Baby Head l.0l a*’ 0.66 a 0.68 a 0.78 H
Badger Ballhead 1.96 a 2.66 b l.63 a 2.08 Y
P.I. 2l5514 l.94 a 3.40 b 3.62 b 2.98 2
Red Danish l.2l a 2.27 b 0.76 a l.4l X
Chieftain Savoy l.27 a 2.96 b l.25 a 1.83 Y
Planting Means 1.48 A 2.39 B 1.59 A

 

*Mean separation. within rows. by Duncan's Multiple Range Test
at the 5% level.

Table 34. CultiVar x inbreeding interaction means for head weight

in kilograms.

68

 

 

 

Inbreeding Cultivar
Cultivar P1 S1 Means
Baby Head 0.89 a* 0.67 a 0.78 H
Badger Ballhead 2.49 b 1.68 a 2.08 Y
P.I. 215514 2.93 a 3.04 a 2.98 z
' Red Danish l.45 a l.37 a l.4l x
Chieftain Savoy 1.89 a l.76 a 1.83 Y
Inbreeding Means l.93 B l.70 A

 

*Mean separation. within rows. by Duncan's Multiple Range Test

at the 5% level.

69.
200 gram difference noted may not be great enough to Justify an extra

year of inbreeding for head weight within these cultivars prior to
using them in a breeding program.

. . It has been previously reported by Rasmusson (24) that large

head weights are dominant to small head weights. Data from green ,

‘ cabbage crosses in this study support Rasmusson's conclusions (Tables
11-13). During 1974. all of the F1 green cabbage crosses showed domi-
nance and overdominance for large head weight. Due to environmental
influences in the 1975L planting. dominance for this trait was reduced.
The F1 populations of Baby Head x Badger Ballhead grown in both the'
1975E and 1975L plantings suggest incomplete dominance for small head
wedght. The 1974L F1 planting of this cross showed overdominance for
large head weight (Tables 11-13). Tables 35 and 36 Show frequency dis-
tributions in percent for populations involved in the cross Baby Head

x Badger Ballhead. Powers et a1. (23) partitioning tests were applied
to these distributions to determine the number of genes controlling
head weight and to define the boundary between small and large head
weight classes. Powers et a1. (23) formula (FZIPI) was applied se-
quentially from the recessive ends of both the F2 and P1 distributions.
Using the 1975E planting of Baby Head x Badger Ballhead (Table 35) as
an example the formula suggests recessive frequencies of 0.4/19.0 or
0.02 for the 0.5 kg class. 7.5/90.5 or 0.08 for the 1.0 kg class and
28.9/100 or 0.29 for the 1.5 kg class. The large increase in frequency
between the 1.0 kg and 1.5 kg classes suggest a dividing point of 1 kg.
The mean of estimates for the 0.5 kg and 1.0 kg classes was 0.05. This
closely approximates the 0.06 estimate expected from a 1:15 model
(Table 35). The 1975E reciprocal F2 population of Baby Head x Badger

7O

 

 

 

 

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71
Ballhead suggests small head weights were dominant. Applying the same
formula(F2/P2) to the recessive end of the Badger Ballhead distribu-
tion. frequencies of 1.6/7.8 or 0.20 for the 9.5 kg class. 2.8/10.6 or
0.26 for the 5.0 kg class. 4.4/30.8 or 0.14 for the 4.5 kg class. i
4.8/43.6 or 0.11 for the 4.0 kg class. 6.0/53.8 or 0.11 for the 3.5 kg
class. 10.4/66.6 or 0.16 for the 3.0 kg class. 16.8/76.8 or 0.22 for
the 2.5 kg class and 34.9784.6 or 0.41 for the 2.0 kg class were observed
-(Table 35). The increase in frequency between the 2.5 kg and 2.0 kg
classes suggest a division point of 2.0 kg to separate the small and
large head weight phenotypes. The mean of frequency estimates from
classes 2.5 kg and larger was 0.17. This result suggests that a 3:1
model with a frequency equal to 0.25 might be appropriate. In both the
F2 and reciprocal F2 populations. overlap of the cultivar distributions
was accounted for by weighting the expected genetic ratios in the fol-
'lowing manner. For the 1975E F2 Baby Head x Badger Ballhead popula-
tion a net overlap of (12.8-9.5) - 3.3 was observed with the large
head weight class deficient (Table 35). The expected number of plants
in the deficient class was calculated to be ((100-3.3)/100)(0.9375)
(252) = 228.45. The expected number of recessive plants was 252-228.45
= 23.55 (Table 37). The expected ratio in the 19756 reciprocal F2
population was calculated in a similar fashion.. The net overlap
from a 2.0 kg dividing point was (12.8 + 2.6 + 7.8) a 23.2 (Table 35).
Again the large head weight class was deficient. The expected number
of plants in the deficient class was ((100-23.2)/100)(0.25)(248) -
47.62. The expected number of early plants was (248-47.62) - 200.38
(Table 37). Similar results for other crosses suggested head weight
might be controlled by 2 loci. The Simplest system available to insure

a digenic inheritance system and explain the cultivar differences

72

 

 

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m.c.u.m._ m.o m._ m._ _.m o.o_ m.~_ m..m m.m~ m.c_ cm. Na Awe x Fay
m.o.u e._ “.PF o.e~ e.em e.e~ m._ as ..a x Nay
eons. ma.m8_egeo;~ 8 arxpt
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73

Table 37. Chi-square test for goodness of fit to the postulated
model for head weight in the F2 generation (l975E planting).

 

 

 

 

 

 

 

 

Cross Observed Expected Model X2 P
Baby Head x
Badger Ballhead ,
*
F2 l9:233 23.55:228.46* l:l5 0.9653 0.32
RF2 206: 42 200.38: 47.62 3: 1 0.8208 0.40 -
Baby Head X
P.I. 2l55l4
*
F2 60:l99 64.75:l94.25* l: 3 0.4646 0.50
RF2 40:4ll 5l.68:399.32 l:l5 2.980l 0.08
Baby Head x
Chieftain Savoy
F2 25:167 33.18:158.82* 1:15 2.4360 0.12
*
RF2 33:164 34.04:]62.96 l:l5 - 0.0384 0.84
*
Pooled 58:33] 67.22:32l.78 l:l5 l.5278 0.22
Red Danish x f
Badger Ballhead
*
F2 l28: 8l l21.92: 87.08*_ 9: 7 0.7285 0.39
RF2 l02: 52 89.83: 64.l7* 9: 7 3.9547 0.05
Pooled 230:l33 21l.75:l51.25 9: 7 3.7750 0.05
Red Danish x
P.I. 2l55l4
*
F2 l58:153 l54.ll:l56.84 7: 9 0.l945 0.66
Red Danish x
Chieftain Savoy
*
F2 46: 6l 49.09: 57.9l*. 7: 9 0.3590- 0.55
RF 56: 46 59.06: 42.94 9: 7 0.3772 0.54

2

 

*
= Weighted

74
observed was a 2 locus multiple allelic system with 5 alleles per loci.
The loci were designated C and D since the small heads of the base
cultivar. Baby Head. were recessive in most crosses. Tables
16 and 17 show the cultivar and F1 genotypes under this system.
Allelic inheritance patterns for 19756 green cabbage crosses suggest.'
cs. c4
pattern. The inheritance pattern for both loci was reversed in the

and 03 were all dominant to? c. The D locus exhibited a similar

1975L planting with c , c4 and c3 recessive to c. Red x green cab-

bage crosses in the 1:75E planting showed an allelic dominance pat-
tern of (:5 and c4 being dominant to 02. and c2 being dominant to 03.
A similar pattern was observed for the Lilocus. Only the F2 populations
of Red Danish x Badger Ballhead were evaluated in the 1975L planting.

No seed of Red Danish x P.I. 215514 was available and no segregation

for head weight occurred from the cross Red Danish x Chieftain Savoy.
The Red Danish x Badger Ballhead cross suggests an allelic inheritance

pattern where c was dominant to c3 and 02 was dominant to 43. The

2‘
results of chi-square homogeneity tests within both the 1975E and 1975L
F2 plantings of Baby Head x Badger Ballhead suggest the F2 and recipro-
cal F2 populations within each planting should not be pooled. Using a
dividing point of 1 kg the 1975E F2 data produced an observed ratio of
19 small:233 large heads which compared favorably (P=0.32) with a 1:15
weighted theoretical ratio of 23.6:228.4 (Table 37). The 1:15 segre-
gation ratio suggests that dominant alleles. c3c50305. were produced by
Badger «Ballhead while recessive alleles. ccdd, were produced by Baby
Head. The genotype ccdd’produced a small head phenotype (Table 35).
The 1975E reciprocal F2 population. using a dividing point of 2.0 kg.
produced a segregation ratio of 206 small:42 large heads. This ratio.

fit (P=0.40) a 3:1 weighted theoretical ratio of 200:48 (Table 37). A

75
segregation ratio of 3:1 suggests that the presence of 1 dominant c
allele conditioned the production of a small cabbage head. The genotype
c3c3_ conditioned a large head phenotype. A backcross of the recessive
cultivar Badger Ballhead to the reciprocal F1 produced an observed ratio
‘ of 11:22. The ratio fit (P=0.05) an expected 1:1 model of 16.5:16.5.
Backcross data Support the postulated model. Since the F2 population
was dominant for large heads as its male parent Badger Ballhead and the
reciprocal F2 population was dominant for small heads as its male parent
Baby Head some evidence for a male inheritance factor exists (Table 35).
Both F2 populations grown in the 1975L planting showed dominance for
small heads. The observed ratio of 92 small:38 large heads from a 1.5
kg division point fit (P=0.48) a 9:7 weighted theoretical ratio of 88:42
(Table 38). Homozygous recessive genotypes at either loci produced a
large cabbage head. Small head weights were conditioned by the pre-
sence of 1 or more dominant alleles at each loci. Using a 1.5 kg divid-
ing point the reciprocal F2 population produced a ratio of 171:40 which
fit (P=0.82) a 3:1 weighted theoretical ratio of 172:39 (Table 38). The
presence of 1 dominant 0 allele conditioned small heads. In both cases
dominant alleles. ccpa. were contributed by Baby Head and recessive
alleles. c3c3djd3. were contributed by Badger Ballhead (Table 36).

Both the 1973E and 1974L plantings of Baby Head x P.I. 215514 F1
showed incomplete to overdominance for large head weights. The.1975L Fl
planting suggested incomplete dominance for small head weights
(Tables 11-13). Chi-square homogeneity tests suggest that both 1975E
F2 populations of Baby Head x P.I. 215514 were statistically different
and therefore were analyzed separately. Using a dividing point of 1.5
kg. calculated as shown previously (23). the F2 population produced a

76

 

 

 

 

 

 

Table 38. Chi-square test for goodness of fit to the postulated
model for head weight in the F2 generation (l975L planting).
I Cross Observed Expected Model * X2 P
Baby Head x
Badger Ballhead
F2 92: 38 88.21: 41.78* 9: 0.5055 0.48
R52 171: 40 172.24: 38.76* 3: 0.0490 0.82
Baby Head x
P.I. 215514
52 , 126: 42 135:69: 32.31* 3: 3.6000 0.06
*
RF2 202: 42 197.08: 46.92 3: 0.6395 0.42
Baby Head x
Chieftain Savoy
*
F2 204: 11 201.56: 13.44 15: 0.4717 0.49
*
RFZ 168: 10 162.99: 15.00 15: 1.8236 0.18
Red Danish X
Badger Ballhead
F2 241: 50 232.67: 58.33: 13: 1.4886 0.22
RFZ 25: 5 23.99: 6.01 13: 0.2136 0.64
Pooled 266: 55 256.65: 64.34* 13: 1.6977 0.19

 

*
= Weighted

77
segregation ratio of 60:199 (Table 39). Due to an overlap of par-
ental distributions the data were compared (P-0.50) to a 1:3 ratio of
65:194 (Table 37). The observed 1:3 segregation ratio suggests only 1
dominant allele ch‘was required to produce a large head. The absence
of c41alleles conditioned small cabbage heads. The dominant alleles.
c4c4b404. were contributed by P.I. 215514. The reciprocal F2 population
produced a segregation ratio of 40:411 using a dividing point of 1.5 kg.
The ratio fit (P=0.08) a 1:15 weighted theoretical ratio of 52:399 (Ta-
ble 37). The presence of 1 dominant allele at either loci was required
for a large head phenotype. The genotype ccdd conditioned small cab-
bage heads. Chi-square homogeneity tests suggest differences between
the 1975L F2 populations of Baby Head x P.I. 215514 (Table 40). The
populations were each analyzed separately. The observed F2 segregation.
ratio of 126:42 was produced using a dividing point of 1.5 kg. The
data fit (P=0.06) a 3:1 weighted theoretical ratio of 135.6:32.3. Using
a dividing point of 2.0 kg the reciprocal F2 population produced an ob-
served ratio of 202:42 which fit (P=0.42) a 3:1 weighted expected ratio
of 197:47 (Table 38). Large heads were produced by the genotype C4048 4(14
Only 1 dominant C'allele was required to produce small heads in this
planting (Tables 16-17). In both the 1975E and 1975L.plantings the
geneotype CCDD produced small head weights (Tables 15-16). In summary
F2 performance in 2 plantings during the same year. 1975. suggest that
alleles behaved differently under different growing conditions.

Three F1 plantings. 1974L. 1975E and 1975L. suggest large head
weights were overdominant in Baby Head x Chieftain Savoy F1 (Tables
11-13). 1975E F2 populations of Baby Head x Chieftain Savoy were pooled.
Using a dividing point (23) of 1.5 kg an observed ratio of 58:331 was

78

 

 

F...“ m.~ ~.e N.N a.e e.m m.m_ a.- o.m. «.mp P.~ a.~ ~.o _me Na “_a x may
a.o.u ..N N.. e.o m.~ _.m m.o. o.~_ e.m~ m.e~ N.N. ..m m.~ am~ Na Ana x pay
m.o.u m.m m.~ m.e_ e.m~ e.w~ a.» N.N ~.m m.~ e.P o“ A.a xAmav.
o.F.H O." N.. m.~_ m.~_ _.a~ ~.m~ ~.m m.m e.e m.~ m.~ am Ana x _av
a...“ m.m m.e_ o._P N.NN o.__ ~.e_ ~.e_ a.m a. Amav came—u .~.a
~.o.H “.0 m.m m.pa o.a_ Ne “Fav eao: seam

and: m.m o.m m.e c.e m.m o.m m.~ o.~ m._ o.P m.o meee_a

ea .62

mumpu we “we“; swan:

msncmo—Px c, agape: new:

 

.c—mmpw .~.m x new: anon mmogu mg» we
mcwucn—a mmump on» com acmucoq cw mcopuanmcummu agave: vow: Nu uce.pu .ucocua .mm w—nmh

79

:93

 

¢.o «.0 ~.p p.o o.a o.~

N.. A... x we

 

e.o.n e.P N m.m~ ..NN P.e eeN

e.o.u P.P e.o m._ a.m ~.m_ m.n~ a.am a... me, me fine x _av

m.o.H ..N m.~ m.~ e.m .o.e.. a.m~ ~.A~ o.ep ~.e Pm Ape x may

gum; N.N N.N 9: a: 98 9; 3.. me A”: r:

_.~.u e.m m.om N.N m.am c.m~ m. Amav epmm_~ .H.a

eoneo mammgmog_ 8 ieeazzs
eaoz m.a e.m m.e o.e m.m o.m m.~ o.~ m._ c.F m.o mace—a

mo .02

mmmpu mo aver; swan:
msmcmoppx cw agave: new:

 

 

mcwucmpa Amnm— ecu com ucmucma cm meow»:

.e_mm.~ .~.a x nee: seam emcee 6:» co
aacbaae o;m_oz neon Na eee pa .beocea .oe o_aae

80 ‘
produced. Due to parental overlap (Table 41) a 1:15 weighted theoreti-
cal ratio was used. The observed data fit (P-0.22) the weighted theo-
retical ratio (Table 37). Small head weights were recessive and condi-
tioned by the ccdd genotype. The presence of either a dominant c5
or 05 allele in the genotype conditioned large heads. The dominant C5
and 05 alleles were contributed by Chieftain Savoy. The model was fur-
ther supported by backcross data. Backcrossing the F1 and reciprocal
F1 to the dominant cultivar Chieftain Savoy produced ratios of 3:29 and
0:15 respectively (Table 41). 1975L F2 populations of Baby Head x Badger
Ballhead were not pooled. Both parents were small headed. However. in
both of the F2 populations segregants with head weights larger than
either parent were observed (Table 42). Separating the F2 data based
on a dividing point (23) of 3.0 kg produced a segregation ratio of
204:11. ,The reciprocal F2 population was separated using a 2.5 kg divide
ing point (23) to produce an observed segregation ratio of 168:10. Due
to parental overlap a 15:1 weighted theoretical ratio was compared to
both observed ratios. The data fit (P=0.49. 0.18) the weighted expected
ratios of 201.6:13.4 and 163:15 respectively (Table 38). Since both cul-
tivars were small headed large heads in the F2 suggest that C was domi-

nant to c' and D was dominant to d. The presence of complementary homo-

5 5
zygous loci. egos and dd. in a genotype produced a large headed phenotype.

Small head weights were produced by genotypes having 1 or more dominant
alleles at either loci (Tables 16-17).

When Red Danish x Badger Ballhead F1 population means were com-
pared with their midparent means small head weight was incompletely
dominant in both the 1974L and 1975L plantings and large heads were com-
pletely dominant in the 19755 planting (Tables 11-13). Fé populations
of Red Danish x Badger Ballhead grown in the 1975E planting were

81

 

 

a.o.u a.~ 8.”. a.“ ~.m_ m.a m.m~ o.o~ N.N. e.m mm Pa x A_a x may
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~.o.u.h.o m.a m._~ o.a_ Ne apav ewe: seam

cad: m.a o.m m.e o.a m.m o.m m.~ o.~ m._ 9., m.o abeapa

ea .62

mmm_u co «wee; saga:

msmcmopwx cw agave: new:

 

 

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mcwpcopa mmnmp ecu so» ucmucoa cw meowuanmcummu agave: new; mmocuxumn use . a

need on» we

..a .eeocaa ..e o_aae

82

 

 

9o“: ... 95 9c 92 99. 9mm 9... a: Na} xmav

9on9. 9o 9. 9~ 9a 9: 9a 9% 92 9e as £3. .13

9on9~ 92 9" 92 9mm 93 N..: 9.: 9. mm taxes:

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9o...H 9o 98 9mm 9: ea. 1.: .86: .33
can: m.a o.m m.a o.e m.m o.m m.~ c.~ m.c o.c m.o ”wewwn

mmmpu co «PEP; cman:
msmcmopwg cw usmwm: tam:

 

 

I .»o>mm :pmucowgu x wee: anmm mmocu as» we
mcmuceFQ AmumP 6;» coc acoucma cw m:o_u=awcumcu “cope: and; we can —u .pcmcua .Ne open»

83
pooled to produce a segregation' ratio of 230:l33. The dividing point
(23) was 2.5 kg. This segregation ratio fit (P=0.05) a 9:7 weighted
theoretical ratio of 212:151 (Table 37). A single dominant allele at
each locus was.required to produce a small cabbage head phenotype. Thus
2_p2__conditioned a small head. Dominant alleles. c2
and 02. were contributed by Red Danish. Backcrossing to the recessive

the genotype c

cultivar Badger Ballhead produced a ratio of 1:11. The ratio fit (P=0.20)
a 1:3 model of 3:9. Backcrossing to the dominant cultivar. Red Danish
produced a 26:22 ratio (Table 43). F2 populations for the 1975L plant-
ing were also pooled. The observed segregation ratio. 266:55. fit
(P-O.19) a 13:3 weighted theoretical ratio of 257:64 (Table 38). A
dividing point (23) of 1.5 kg was used (Table 44). Either a dominant
genotype at the c locus or a recessive genotype at the a locus produced
a small cabbage head. Large heads were produced by the genotype cijszz.
Dominant alleles c2 and 02 were contributed by Red Danish. Recessive
alleles. c3 and d3,
1974L Red Danish x P.I. 215514 F1 population means suggest head

were contributed by Badger Ballhead.

weight was additively inherited. The 1975E and 1975L F1 means suggest.
large heads were incompletely dominant and small heads were incompletely
dominant respectively (Tables 11-13). Due to limited seed. the Red
Danish x P.I. 215514 F2 population was grown only in the 1975E planting.
Using a dividing point (23) of 2.5 kg an observed segregation ratio of
158:153 was produced (Table 45). This ratio was compared (P'0.66)

to a 7:9 weighted theoretical ratio of 154:157 (Table 37). The presence
of 1 dominant allele at each locus provided a basis for the ratio.
Dominant c4 and D4 alleles were contributed by P.I. 215514 while Red

Danish contributed the recessive alleles c and d2. The genotype c4_p4_

2
conditioned a large head weight (Table 45).

 

 

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msacmoFVx cw agave: com:

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«awed ea a8 Ndcmaumacaécudceé Na 8 c3x~$
F...“ o.N c.a ~.a N.m c.a. a.ec m.~c m.m_ a.ec ~.m _.m as cNa x aav
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a.o.u a.~ c.m m.~ c.m m.ec c.mN o.o~ c.cc c.m mm ceav emceao com
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ca .62

 

 

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m:_ucmpa mmmmp on» coc acmucoa cw mcomuznpcummv azure: vow; mmocuxuma use .Nu .pd .ucocma

.me mpnmh

85

 

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co .0:

mum—u co peep; cane:
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.86

 

 

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cad: m.m o.m m.e o.e m.m o.m m.~ o.~ m._ o.2 abea2a
ca .62

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msmcmopmx cw agave: you:

 

 

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.e2mm_~ ._.a x emceao eox amoco oeb co
anccpm_c cameo: com; me use —c .ucmcma .me «peep

87

Red Danish x Chieftain Savoy F1 population means for the 1974L
planting show an additive inheritance pattern for head weight. However.
both the 1975E and 1975L plantings suggest overdominance for large head
weights (Tables 11-13). Differences in the dominance direction of head
weight between plantings coupled with the relatively stable dominance
direction of maturity may have contributed to problems in estimating.
the genetic correlation of these 2 traits. Both Red Danish and Chief-
tain Savoy distributions in the 1975L planting were nearly identical.
Red Danish x Chieftain Savoy F2 segregation patterns did not deviate
from the parental patterns observed (Table 48). Based on these distri-
butions no segregation could be detected in this planting (Table 48).
1975E Red Danish x Chieftain Savoy F2 populations produced a segrega-
tion ratio of 46:61 using a division point (23) of 2.5 kg (Table 47).
This ratio fit (P=0.55) a 7:9 weighted theoretical ratio of 49:58 (Table
37). Large heads were dominant and conditioned by a single dominant
allele at each locus. The genotype c5_p5__produced a large head weight

phenotype. Recessive alleles. c and d were contributed by Red Danish.

2 2’
The reciprocal F2 population of this cross produced an observed segre-
gation ratio of 56:46 which fit (P=0.54) a 9:7 weighted expected ratio
of 59:43 (Table 37). The dividing point (23) was 2.5 kg. The presence
of a single dominant 02 or 02 allele from Red Danish at each locus con-

ditioned a small head phenotype. Recessive alleles. c and as. were

5
contributed by Chieftain Savoy. F1 and reciprocal F1 backcrosses to Red
Danish produced observed ratios of 5:15 and 13:27 respectively. The
data show poor fit (P<0.005) to expected ratios of 13.5:4.5 and 40:0.
The backcross data suggest Red Danish was recessive and produced large
head weights. however cultivar distributions suggest Red Danish produced

small heads.

88

 

 

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c.o.u m.e m.~ m.~ m.~ c.- e.oe e.e~ ea Ana x eav

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mmepu co wes_e cwae:
msecmopex cw pzm_w3 new:

 

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.eceecm .~.a x eaweee ewe anaco age co
ea_b=accen2e “emcee eewe 2a eee peacea .ee a2aec

89

 

p.w ~.w ~.w

ea x cea x eav

 

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ca .62

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mocu we» co
.Pd .ucwcea

.me wpneh

90

 

a.e a.o e.e a.e2 e.e~ e.mm o.2~ e- we 2ea x ea2

 

e.e.u o.2

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9392 9e 9e 92 92 2.2 9” 92. on 222:9:

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ca .62

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.mv wEfl

91

Green cabbage F1 heritability estimates for head weight have been
previously Shown by Swarup (30) to be 0.4 and by Chiang (3) to be 0.04.
This study shows mean head weight heritability for green cabbage crosses
to be large. EZ=O.87 1 0.49 while $2=0.06 1 0.12 for head weight in red
x green cabbage crosses. Heritability estimates between 0 and 1 ranged

from 0.86 to 0.89 for green cabbage crosses and from 0.05 to 0.93 for

2
x

and 0.08 t 0.12 respectively. Reciprocal effects can be noted in the

negative and positive hz estimates Shown for head weight in the 1975L

red x green cabbage crosses (Table 30). The E values were 0.93 a 0.36

red x green cabbage F1 planting. Environmental effects can be sh0wn by

2=0.05

=0.93. These differences point out the

comparing the 1974L red x green cabbage heritability estimate of h

with the 1975L estimate of a2

need for numerous experiments prior to accepting a reliable heritability
parameter for a given location (Table 30).

Genetic correlation estimates between head weight and total non-
wrapper leaf weight for green cabbage crosses ranged from 0.35 to 0.92
(Table 49). The mean genetic correlation was 0.78 t 0.48 and the.£gx=
0.78 i 0.48. The mean genetic correlation between head weight and
stalk size was 0.96 i 0.18 with an :ng value of 0.97 t 0.17. Genetic
correlation estimates ranged from 0.33 to 0.98. A large number of nega-
tive correlation estimates were observed between head weight and non-
wrapper leaf number. It appears that larger heads tend to have fewer
non-wrapper leaves than smaller heads. Thel? estimate for head weight

9

and non-wrapper leaf number was -0.73 i 0.63 and thee-gx value was 0.39

i 0.37. The mean genetic correlation between head weight and non-wrapper
leaf size was 0.83 i 0.41. The ng value was 0.93 i 0.29. These genetic
correlations suggest that head weight was positively correlated with

total non-wrapper leaf weight and a few large leaves rather than many

92

 

 

 

 

 

 

 

Table 49. Genetic correlation between head weight and four other
traits in cabbage.
Traits Plantings Green Crosses . Red Crosses
R‘ R*
Head Weight x 1974L 0.62 0.40 A A
Total Non-wrapper 19755 0.56 0.77 A 0.46
Leaf Weight l975L 0.35 0.92 B -0.07
Head Height x 1974L 0.83 0.86 A A
Stalk Size l975E A 0.98 0.98 0.62
1975L 0.33 ‘O.73 B -O.85
Head Height x 1974L 0.08 -O.l9 -0.94 B
Non-wrapper Leaf 1975E -O.84 -l.00 A 0.21
Number 1975L 0.12 A A 8
Head Weight x 1974L 0.95 0.83 A A
Non-wrapper Leaf 19755 A A A i 0.65
Size l975L 0.12 0.60 -O.ll A

 

 

 

 

 

*Reciprocal cross

A = Correlation estimates greater than 1.0

B = Correlation estimates less than -1.0

 

93

smaller leaves. Red x green cabbage crosses produced a greater number
of negative genetic correlation estimates than did green cabbage crosses
(Table 49). The relationship between head weight and total non-wrapper
leaf weight for red x green cabbage crosses was 0.34 2 0.87 with an

ng estimate of 0.36 t 0.84. Genetic correlation estimates ranged from
-0.07 to 0.46. Genetic correlations between head weight and stalk size
ranged from -0.85 to 0.98. The mean genetic correlation between these

2 traits was 0.36 i 0.97 with a 29x estimate of 0.58 i 0.75. Mean gene-
tic correlations between head weight and non-wrapper leaf number and non-
wrapper leaf size were -0.23 i 0.93 and 0.54 i 0.90 respectively. The
ng estimates were -0.76 i 0.67 and 0.76 i 0.65 with rg estimates rang-
ing from -0.94 to 0.21 and -0.11 to 0.65 respectively (Table 49).

Total Non-Wrapper Leaf Weight_

Cultivar analysis showed that differences exist within cultivars.
inbreeding and plantings. Interactions between each of these effects
were significant (Table 50). One generation of inbreeding significantly
increased total non-wrapper leaf weight by 0.15 kg. This trait may not
be genetically fixed in the cultivars since negative heterosis was
still modifying the plant phenotype (Table 51). There was no differ-
ence between Red Danish and Chieftain Savoy for total non-wrapper leaf
weight. however. the remaining cultivars were each significantly differ-
ent from this group. Total non-wrapper leaf weight ranged from 0.19 kg
for Baby Head to 2.43 kg for P.I. 215514 (Table 51). Differences be-
tween plantings for total non-wrapper leaf weight suggest different
environmental conditions can produce leaf weights ranging from 0.78 to

1.97 kg (Table 52). The cultivar x inbreeding interaction was significant

94

Table 50. Analysis of variance table for total nonéwrapper leaf
weight in kilograms.

 

 

 

Source
of Mean
Variance . DF Square F*
Replication (R) 2 0.08 1.24
Cultivar (0) _ 4 11.70 169.81**
' ' **
Inbreeding (I) l 0.49 7.10
*‘k
C x-I 4 0.38 5.58
Plantings (P) 2 10.82 156.96**
0 x.P 8 0.92 13.33**
**
I x P _ 2 0.45 6.53
, **
C x.P x I 8 0.30 4.41
Residual Error ‘ 58 0.07

 

**
*F test significant at the 5%*. or 1% level.

95

Table 51. Cultivar x inbreeding interaction means in kilograms

for total non-wrapper leaf weight.

 

 

 

 

Inbreeding Cultivar

Cultivar P1 S1 Means
Baby Head 0.23 a* 0.15 a 0.19 W
Badger Ba11head 1.42 b 1.04 a 1.09 X
P.I. 215514 2.18 a 2.67 b 2.43 2
Red Danish 1.26 a 1.70 b 1.48 Y
Chieftain Savoy . 1.47 a 1.46 a 1.47 Y
Inbreeding Means 1.26 A 1.40 B

 

*Mean separation. within rows. by Duncan's Multiple Range Test

at the 5% level.

96

Table 52. Cultivar x planting interaction means in kilograms for
total non-wrapper leaf weight.

 

 

 

 

, Plantings ' Cultivar
Cultivar 1974L l975E 1975L Means
Baby Head 0.17 a* 0.36 a 0.04 a 0.19 W
Badger Ba11head 1.69 c 1.19 b 0.39 a 1.09 X
P.I. 215514 3.02 b 2.17 a 2.09 a 2.43 2
Red Danish ‘ a 2.45 c 1.27 b 0.73 a 1.48 Y
Chieftain Savoy 2.52 c 1.25 b 0.63 a 1.47 Y
Planting Means 1.97 C 1.25 B 0.78 A

 

*Mean separation. within rows. by Duncan's Multiple Range Test
at the 5% level. ‘

.97
due to an increase in total non-wrapper leaf weight after 1 additional
generation of inbreeding Red Danish. The other 3 cultivars were not
effected by inbreeding (Table 51). The cultivar x planting interaction
was significant due to differences in total non-wrapper leaf weight for
both Baby Head and P.I. 215514. ‘Baby Head showed no difference in'
total nonewrapper leaf weight for the 1974L and 1975E plantings; however.
P.I. 215514 and the remaining cultivars showed a decrease in total non-
wrapper leaf weight between these 2 planting times. Significant reduc-
tions in total non-wrapper leaf weight were noted for all cultivars ex-
cept Baby Head and P.I. 215514 between the 1975E and 1975L plantings
(Table 52). The significance of the inbreeding x planting interaction
was due to an increase in total non-wrapper leaf weight after a gener4
ation of inbreeding in the 1974L planting. Inbreeding did not effect
total non-wrapper leaf weight for the other plantings (Table 53). The
significance of the second order interaction cultivar x inbreeding x
planting can be attributed to differences due to inbreeding in.the 1974L
planting for Red Danish and P.I. 215514. The data show a significant
increase in total non-wrapper leaf weight after 1 generation of in-
breeding for both of these lines while no differences were shown for
the other 3 cultivars. Total non-wrapper leaf weight was not affected
by 1 generation of inbreeding in either the 1975E or 1975L plantings
(Table 54).

Means of F1 green cabbage crosses lie near the midparent mean and
show incomplete dominance for non-wrapper leaf weight. Although the
magnitude of the cultivar and F1 population means seem to be affected
by the environment. the F1 means lie near the midparent mean for each

planting. All 3 F1 plantings of Baby Head x Badger Ballhead suggest

98

Table 53. Inbreeding x plantings interaction means in kilograms
for total non-wrapper leaf weight.

 

 

Plantings Inbreeding

 

 

Inbreeding 1974L l975E l975L Means
P] 1.76 a*' 1.27 a 0.75 a 1.26 Y
S1 2.18 b 1.23 a 0.80 a 1.40 2
Planting Means 1.97 C 1.25 B 0.78 A

 

*Mean separation. within columns. by Duncan's Multiple Range Test
at the 5% level. j

99

Table 54. Cultivar x inbreeding x planting interaction means
in kilograms for total non-wrapper leaf weight._

 

 

 

 

 

 

 

Plantings

Cultivar Inbreeding . 1974L 1975E . ' 1975L
Baby Head P] 0.23 a* 0.39 a 0.06 a

S] 0.12 a ' 0.33 a 0.02 a
Badger P1 1.79 a' 1.24 a 0.42 a
Ballhead

S1 1.59 a 1.15 a 0.37 a
P.I. 215514 P.I 2.35 a 2.17 a 2.03 8

S1 3.68 b 2.17 a 2.15 a
Red Danish P] . 1.83 a 1.36 a 0.60 a

51 3.07 b 1.18 a 0.86 a
Chieftain P1 2.59 a 1.18 a 0.64 a
Savoy

S1 2.45 a 1.32 a 0.62 a

 

*Mean separation. within columns and cultivars. by Duncan's
Multiple Range Test at the 5% level.

100

incomplete dominance for total non-wrapper leaf weight. Reciprocal dif-
ferences between F1 population means were noted in the 1975L planting
(Tables 11-13). Powers et a1. (23) partitioning tests were applied to
these distributions to determine the number of genes controlling total
non-wrapper leaf weight and to define the boundary between Small and
large total non-wrapper leaf weight classes. Powers et a1. (23) for—
mula (Fa/P1) was applied sequentially from the recessive ends of both
the F2 and P1 distributions. Using the 1975E F2 planting of Baby Head

x Badger Ballhead (Table 55) as an example recessive frequencies of
6.0/4.8 or 1.25 for-the 0.2 kg class. 12.0/23.8 or 0.50 for the 0.3 kg
class. 21.5/61.9 or 0.35 for the 0.4 kg class. 31.8/88.1 or 0.36 for the
0.5 kg class. 46.1/95.2 or 0.48 for the 0.6 kg class and 67.3/100 or
0.67 for the 0.8 kg class were produced (Table 55). The increase in
frequency between the 0.6 kg and 0.8 kg classes suggested a dividing
point of 0.6 kg. Further. the mean of all values less than 1 for classes
0.2 to 0.6 kg was equal to 0.42. which closely resembles a 0.44 fre-
quency expected from a 7:9 model. Due to overlap of the parental dis-
tributions a weighting procedure was used to adjust the expected gene-
tic ratio to account for misclassified plants. The 1975E Baby Head x
Badger Ballhead cross required a weighting procedure which was calcu-
lated in the following manner. The net overlap around the 0.6 kg
dividing point was 2.5+2.5-4.3=o,4. The expected number of plants in the
deficient class was ((100-0.4)/100)(0.5625)(252)) = 141.23. By sub-
traction a weighted expected ratio of 110.77:141.23 was produced (Table
57). The weighted expected ratio for the 1975E reciprocal planting was '
calculated in the following manner from a 0.5 kg dividing point (23).

The expected number of plants in the deficient class was ((100-9.3)/100)

101

 

 

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102

 

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ca .62

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103

Table 57. Chi-square test for goodness of fit to the postulated

model for total non-wrapper leaf weight in the F

generation (l975E planting).

 

 

 

 

 

 

 

 

Cross Observed Expected Model X2 P
Baby Head X
Badger Ballhead

*
F2 109:143 110.77:141.23 7: 9 0.0504 0.82
RF2 130:118 126.47:121.53* 9: 7 0.2010 0.65
Baby Head X
P.I. 215514 1
52 201: 58 194.25: 64.75 3: 1 0.9383 0.33
RF2 331:120 338.25:112.75 3: 1 0.6216 . 0.43
Baby Head X
Chieftain Savoy

*
F2 110: 82 103.10: 93.90* 7: 9 0.7073 0.40
RF2 109: 88 100.49: 91.51* 7: 9 1.8893 0.17
Pooled '219:170 203.59:185.41 7: 9 2.4466 0.12
Red Danish x
Badger Ballhead

*.
F2 70:139 80.85:128.15* 7: 9 2.3761 0.12
RF2 59: 95 59.58: 94.42* 7: 9 0.0091 0.92
Pooled 129:234 l40.43:222.57 7: 9 1.5173 0.22
Red Danish x
P.I. 215514

*
F2 110:201 115.14:l95.86 l: 3 2.3649 0.54
Red Danish x
Chieftain Savoy
52 10: 97 8.59: 98.41: 1:15 0.2516 0.64
RF2 10: 92 8.19: 93.81* 1:15 0.4349 0.51
Pooled 20:189 16.78:192.21 1:15 0.6718 0.44

 

*
= Weighted

104
(0.5625)(248) = 126.47. The expected ratio was 126.47:121.53. The 1975E

reciprocal F2 population produced a different model and division point.
The distribution was bimodal with inflection points at 0.3 kg and 0.6
kg. The 0.5 kg class. with the fewest members between 2 inflection
points was selected as a division point. As in the previous example
(23) the formula (FZ/Pz) suggested frequencies 3.2/32.0 or 0.08 for the
3.8 kg class. 4.8/48.8 or 0.10 for the 1.4 kg class. 9.2/59.0 or

0.16 for 1.2 kg class. 18.5/76.9 or 0.24 for the 1.0 kg class. 31.9/94.8
or 0.34 for the 0.8 kg class. 47.6/97.4 or 0.49 for the 0.6 kg class.
59.7/97.4 or 0.61 for the 0.5 kg class and 73.0/100 or 0.73 for the 0.4
kg class (Table 55). Since the mean of classes 3.8 and 1.4 was 0.089
which compares favorably with an expected 0.06 frequency expected from
a digenic model a 2 gene model was postulated. The data also suggest
division points (23) between classes 1.4 and 1.2 kg and also between
classes 0.6 and 0.5 kg. An evaluation of the data suggested a 9:6:1
model might be appropriate. A multiple allelic 2 locus model was pos-
tulated as the simplest model available to insure digenic control of
total non-wrapper leaf weight. To account for_cu1tivar differences each
locus in the model was given 5 alleles. The loci. E and G. produce the
cultivar and F1 genotypes noted in Tables 16 and 17. For the 1975E

and a were dominant to E, and E

3 5

was dominant to 94. Alleles at the G locus were inherited in a similar

manner. Red x green cabbage crosses suggest an allelic dominance pat-

planting of green cabbage crosses. a

tern where E3 and E4 were dominant to £2. and E2 was dominant to as.

A similar pattern was noted for the G locus. In the late planting E was

dominant to e3, 64 and as; 32 was dominant to 63 and as. Similar patterns

were noted for the G locus. Chi-square homogeneity tests suggest both

105
1975E F2 populations of Baby Head x Badger Ballhead Should not be
pooled (Table 55). Using a dividing point of 0.6 kg an F2 segregation
ratio of 109 small non-wrapper leaf weight leaves to 142 large non—
wrapper leaf weight leaves showed good fit (P=0.82) to a 7:9 weighted
theoretical ratio of 11:141 (Table 57). A single dominant allele at
each locus was required to produce a large total non-wrapper leaf weight.

Dominant alleles. E and G3. were contributed by Badger Ba11head. while

3
recessive alleles. eland g. were produced by Baby Head. The reciprocal
F2 population produced a segregation ratio of 130:106:12 which fit
(P=0.43) a 9:6:1 weighted expected ratio of 126.47:104.17:17.36. Data
from a backcross between the reciprocal F1 and Badger Ba11head produced
an observed ratio of 6:27 which fit (P=0.92) an expected 1:3 ratio of
8.25:24.75. A single dominant allele at each locus conditioned small
total non-wrapper leaf weight. Medium non-wrapper leaf weight was pro-
duced by a single dominant locus genotype. Large total non-wrapper
leaf weight was produced by-a homozygous recessive genotype. Dominant
alleles. a and G were produced by Baby Head and recessive alleles. e3
and 93. were contributed by Badger Ballhead. The reciprocal F2 data
support a 9:7 weighted theoretical ratio for total non-wrapper leaf
weight inheritance (Table 57). The 96:34 F2 ratio observed from the
Baby Head x Badger Ballhead 1975L planting fit (P=0.07) a 3:1 weighted
expected ratio of 104:26. A 0.2 kg division point was used. The re-
ciprocal F2 population produced a segregation ratio of 169:42 using a
0.2 kg division point. Theratio fit (P=0.17) an expected ratio of
169:42 produced by a 3:1 weighted theoretical ratio (Table 58). The

Presence of E conditioned a small leaf weight while the recessive geno-

tYpe e363 conditioned a large leaf weight. Dominant alleles 5 and c

106

Table 58. Chi-square test for goodness of fit to the postulated
model for total non-wrapper leaf weight in the F2
generation (1975L planting).

 

 

 

 

 

 

 

Cross Observed Expected Model X2 P
Baby Head x
.Badger Ballhead
52 96: 34 104.13: 25.87* 3: 1 3.1921 0.07
RF2 161: 50 169.01: 41.98* 3: 1 1.9103 0.17
*
Pooled 257: 87 273.15: 67.85 3: 1 4.7976 0.03
Baby Head x
P.I. 215514
F2 128: 40 126.00: 42.00 3: 1 0.1269 0.72
RF2 195: 49 183.00: 61.00 3: 1 3.1478 0.08
Pooled 323: 89 309.00:103.00 3: 1 2.5372 0.11
Baby Head x
Chieftain Savoy
*

F2 165: 50 168.75: 46.25* 3: 1 0.3873 0.53
RF2 139: 39 .139.71: 38.29 3: 1 0.0167 0.90
*

Pooled 304: 89 308.46: 84.54 3: 1 0.2997 0.58

Red Danish x
Badger Ballhead
F2 55:236 49.37:24l.63* 1: 3 0.7744 0.38
*
RF2 14: 16 5.09: 24.91 1: 3 18.7889 '<0.01
Red Danish x
Chieftain Savoy
*
F2 51:148 51.48:l47.51* l: 3 0.0062 0.94
RF2 49:175 57.95:l66.05* 1: 3 1.8661 0.17
Pooled 100:323 109.44:313.56 1: 3 3.5405 0.06

 

*
= Weighted

107

were contributed by Baby Head while the recessive alleles e and g were

3 3
produced by Badger Ballhead (Table 56).

F1 population means for Baby Head x P.I. 215514 suggest total non-
wrapper leaf weight inheritance ranged from incomplete dominance for
small total weight in the l975E and 1975L plantings to additive inheri-
tance in the 1974L planting (Tables 11-13). Both 1975E F2 populations
of Baby Head x P.I. 215514 fit 3:1 models. The genotype 8_g__produced
a dominant small non-wrapper leaf weight phenotype (Table 59). Using
a division point of 1.1 kg an observed ratio of 201:58 was produced‘
which fit (P=O.33) a 3:1 model of 194.25:64.75. The reciprocal F2
population. using a 1.3 kg dividing point. produced an observed ratio
of 331:120 which fit (P=0.43) a 3:1 model of 338.25:112.75 (Table 57).
Baby Head x P.I. 215514 F2 populations grown in the 1975L planting were
pooled. Using a divison point of 0.5 kg an observed ratio of 323:89 was
produced (Table 60). The ratio fit (P=0.53) a 3:1 model of 309:103

(Table 58). The genotype e4e49 94‘conditioned a recessive large total

4
non-wrapper leaf weight phenotype. Dominant alleles a and 6 were con-
tributed by Baby Head.

Baby Head x Chieftain Savoy F1 populations suggest total non-
wrapper leaf weight was inherited in an additive manner. Both the 1974L
and 1975E Fl population means were larger than the midparent mean. .Fl
means for the 1975L planting suggeSt incomplete dominance for small to-
tal non-wrapper leaf weight (Tables 11-13). Baby Head x Chieftain
Savoy F2 distributions within the 1975E and 1975L plantings were pooled.
The 1975E F2 segregation ratio of 219:170 was compared (P=0.12) to a
7:9 weighted theoretical ratio of 204:185 (Table 57). The division

point was 0.7 kg (Table 61). One dominant allele at each loci conditioned

108

 

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110

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111

large leaf weight. A homozygous recessive locus in the genotype condi-
tioned small total non-wrapper leaf weight. Dominant alleles. 35 and
as. were contributed by Chieftain Savoy. Recessive alleles e and g were
contributed by Baby Head. Backcrossing the reciprocal F1 to the reces-
sive cultivar. Baby Head. produced a segregation ratio of 42:13 (Table
61). This data fit (P=0.84) a 3:1 model expected on backcrossing to
a 7:9 model. Both backcrosses to Chieftain Savoy suggest the absence
of complete dominance. Backcrossing to the F1 produced a 5:27 ratio
while backcrossing to the reciprocal F1 produced a 5:10 ratio. Ratios
of 0:32 and 0:15 were expected to support the 7:9 model. Data from the
pooled 1975L F2 population suggest the genotype a___ was dominant for
small non-wrapper leaf weight under these growing conditions (Table.62).
Using a dividing point of 0.4 kg the pooled 1975L F2 population. produced
a segregation ratio of 304:89 which fit (P=0.53) a 3:1 weighted theoreti-
cal ratio of 308:85 (Table 58). A single dominant gene controlled small
total non-wrapper leaf weight in this cross. The recessive genotype
e5959595 conditioned large total non-wrapper leaf weight in this planting.

Red x green cabbage crosses exhibit a different direction of domi-
nance and inheritance pattern for total nonéwrapper leaf weight. Al-
though 1975L F1 plantings of Red Danish x Badger Ballhead and Red Danish
x P.I. 215514 produce population means approximating the midparent mean.
suggesting additive gene action. the remaining crosses in the 1974L.
1975E and 1975L plantings exhibited dominance or overdominance for in-
creased total non-wrapper leaf weight (Tables 11-13). The pooled
1975E F2 population of Red Danish x Badger Ballhead produced a segrega-
tion ratio of 129:234 using a division point of 1.2 kg (Table 63). This

ratio fit (P=0.22) a 7:9 weighted theoretical ratio of 140:223 (Table 57).

112

 

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113

 

 

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e.e.u.2.2 m.e m.w m.e m.e e.mm s.e2 2.e2 N2 me x 2~w x ea2
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m.o H e.2 0.2 w.~ m.e ~.2 e.22 e.22 a.e2 m.m2 o.o2 e.e2 e.2 mow we 2~a x eav
e.o.u 2.2 N.N e.m 2.e m.m e.e2 N.Nw e.e~ 2.22 .2.e e.e 2.2 cm 2ea x ~22
e.o.H 2.2 e.2 2.m w.e e.w2 e.e2 e.e2 e.m2 e.e2 e.o2 e.2 0.2 em 2~w x eav
90H 92 9m 9w. 9” 92 92 92 92 922 2.2 2 2w: eew22ee cweeee
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ca .62

mwe2w ca 22522 cwaae
maecmo22x s2 usmmwz cews cwsseczisoz,peuo»

 

 

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coc uswocws s2 mso2wsnecum2u psm2w3,cew2 cwsseczisos peace mmocwxwen use .Ns .pd .uswces

.mm wpaeh

114
Data from the backcross F1 x Badger Ballhead suggest a poor fit (P<0.005)
to the expected 3:1 model. Data from the backcross F1 x Red Danish pro-
duced an observed ratio of 1:46. A 0.47 ratio was expected (Table 63).
The data support the postulated 7:9 model. Genes contributed by Red

Danish. E and G2. were dominant for increased total non-wrapper leaf

2
weight (Table 16). The presence of a homozygous recessive e

3 or 93 '
locus conditioned small total non-wrapper leaf weight. The same genes .
appear to be dominant in the 1975L planting. *The Red Danish x Badger
Ballhead F2 population produced 55 small total non-wrapper leaf weight
phenotypes and 236 large total non-wrapper leaf weight phenotypes. A
1:3 weighted expected ratio of 49:242 was compared (P=0.38) to the ob-
served segregation ratio (Table 58). A division point of 0.4 kg was
used to separate the F2 data into classes (Table 64). The genotype
52___ was dominant.for 1arge total non-wrapper leaf weight (Table 64). h
The recessive genotype e3e3g3g3 conditioned small total non-wrapper leaf
weight. The 93 and g3 alleles were contributed by ‘Badger Ballhead.
A1though data from the 1975L F1 planting of Red. Danish x P.I. 215514
suggest incomplete dominance for small total non-wrapper leaf weight.
data from both the 1974L and 1975E F1 plantings suggest over-dominance
for large total non-wrapper leaf weight (Tables 11-13).. The dominant
E4 allele donated by P.I. 215514 produced a large total non-wrapper leaf
weight phenotype when compared to the e2 allele (Tables 16. 65). Using
a dividing point of 1.8 kg the 1975E planting of RedDanish x P.I. 215514
produced an F2 segregation ratio of 110:201. The data fit (P=0.54) a
1:3 weighted expected ratio 0f 115:196 suggesting the E4 allele was
dominant (Table 57). No E2 populations of this cross were planted in

the 1975L planting (Table 66).

115

 

 

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9eu9e 92 92 2 92 92 92 92 2.2 92 9e 2e 22. we 32.:
2.22....922 92... 99a. 92e 95 2 22.22:“:
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a .2.

mme2u co peeve cwaa:
msecmopex sw usmewz mewe cwsneczusoz Peach

 

 

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coc uswucws se msoeusnecpmeu usmewz cew2 cwsseczisos peas» we use 2; .uswcee .em w—eeh

116‘

 

F.NF

Ne 2me x eev

 

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117

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.6 .az

mme2u co “~52; cwan:
maecmopex s2 usa2w= cewe cwsaeczisoz 2epo»

 

 

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coe uswucwa se msompenwcumwu usmewz wewp cwaneczisos Peace 2; use uswces .mo.wpneh

118

F1 data from the 3 plantings. 1974L. 19755 and 1975L. of Red Danish
x Chieftain Savoy suggest overdominance for large total non-wrapper leaf
weight (Tables 11-13). F2 populations within each planting were pooled
based on chi-square homogeneity test results (Tables 67. 68). Using
a dividing point of 1.2'kg the pooled 1975E F2 populations produced a
segregation ratio of 20:189 which fit (P=0.44) a 1:15 weighted expected
ratio of 17:192 (Table 57). Both the F1 and reciprocal F1 plants were
backcrossed to the dominant cultivar Red Danish. Ratios of 0:18 and
18:29 were observed. Ratios of 0:18 and 0:37 were expected on back-
crossing to a 1:15 ratio (Table 67). Results from crossing the recipro-
cal Fi to Red Danish suggest dominance may be incomplete. A single
dominant gene at either or both loci conditioned 1arge total non-wrapper
leaf weight. The recessive genotype esesgsgsconditioned small total
non-wrapper leaf weight. Recessive alleles 85 and 95 were contributed
by Chieftain Savoy. Using a 0.6 kg dividing point the pooled 1975L 52
population produced a segregation ratio of 100:323. The data fit
(P=0.06) a 1:3 weighted expected ratio of 109:314 (Table 58). Red
Danish wasthe source of dominant genes for large total n0n-wrapper leaf
weight in this population (Table 68). The presence of a dominant 32
allele in the genotype conditioned a large total non-wrapper leaf weight
phenotype regardless of the G locus genotype. F2 segregation ratios
support dominance patterns noted in the F1 generation. i.e., dominance
for small total non-wrapper leaf weight in green cabbage crosses and
dominance for large total non-wrapper leaf weight in red x green cab-
bage crosses.

Heritability estimates of total non-wrapper leaf weight for green

cabbage crosses were as follows: 4 of the 6 heritability estimates were

119

 

 

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mme2u co 22524 cwss:

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mo .02

 

 

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coe uswucws sw msowusnmcumwu unmewz cewn cwasecznsos pence mmocueuea use .Ne .2; .2swcee .Nu wpeeh

120

 

ea 2ee x eev

 

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e.e.u e.o m.e e.e e.e e.e2 e.e2 N.NN e.e o.e e.e e.e e.e ee2 es 2ee x eav
e.o.u e.o m.m 2.e m.m2 e.e2 e.ee e.e2 o.e2 m.e 2.e .o.e2 om 2ee x ea2
e.e.u e.o e.m c.e2 2.2 .e.2e e.22 m.e2 2.e2 e.m e.m 2.2 ee 2ee x eev
m.o.H e.o e.e m.e e.e e.e2 e.me. 2.ee m.e e.e m.e me 2me2 easwm e2wecw2ew
m.e.u e.e 2.e e.e 2.m e.e2 e.2e e.e2 e.e2 2.m e.2e em ‘ 2ee2 ea2eee ewe

sew: e.m e.2 e.2 e.2 e.2 e.o e.e m.o e.o e.e e.o meew2e

ca .az

ewe—u co peeps cwan:
msecmo2e¥ se unmewz cwsseczisoz peace

 

 

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coc uswucws se msoepsnecumeu psmewz cewp cwaneczisos 2e2ou we use pm .uswcee .wc wpaeh

121
greater than 1. One of the 6 estimates was negative (Table 30). The
1975E planting provided a heritability estimate of 0.54 t 0.54. suggest-
ing that at least half of the variation in the F1 population was addie
tive. The 5: estimate was 0.75 a 0.39. 'F1 red x green cabbage popula-
tions produced 4 heritability estimates greater than 1 and 2 estimates
’ within the range of 0 toll. The estimates were 0.17 for the 1974L
plantingand 0.88 for the 1975L planting. The 71% difference between
the 2 estimates suggests that a large number of estimates are needed to
obtain a reliable heritability estimate. The mean heritability eStimate
» for red x green cabbage crosses was 0.36 s 0.37 and Ex=0.60 i 0.29
(Table 30).

Green cabbage cross genetic correlation estimates between total
non-wrapper leaf weight and stalk size range from 0.16 to 0.96 (Table
69). The 59 value for this pair of variables was 0.93 i 0.26. The mean.
4 genetic correlation between total non-wrapper leaf weight and non-wrapper
leaf number was 0.33 s 0.11 and ranged from -0.96 to 0.98. The green
cabbage mean genetic correlation between total non-wrapper leaf weight
and non-wrapper leaf size was 0.90 t 0.37 with r estimates ranging from

g
0.51 to 0.95. The I value was 0.63 i 0.29. Red x green cabbage crosses

gx

exhibit a mean genetic correlation of 1.00 s 0.04 for total non-wrapper
leaf weight and stalk size. Selection for reduced total non-wrapper
leaf weight might also reduce stalk size. Estimates for this correla-
tion ranged from 0.22 to 1.00. For red x green cabbage crosses the mean
genetic correlation between total non-wrapper leaf weight and non-wrap-
per leaf number was 0.22 i 0.85 with a range of -0.39 to 0.53 (Table 69).

The mean genetic correlation between total non-wrapper leaf weight and

non-wrapper leaf size was 0.34 s 0.93. Only 1 r9 estimate between -1

122

Table 69. Genetic correlation between total non-wrapper leaf weight
and three other traits in cabbage.

 

J

 

 

 

 

 

Traits Plantings Green Crosses Red Crosses
8* 8*
Total Non-wrapper Leaf 1974L 0.96 0.84 0.57 0.69
Weight x Stalk Size l975E . 0.16 0.52 1.00 A
l975L 0.74 A . 0.22 0.96
Total Non-wrapper Leaf 1974L 0.85 0.82 A -0.39
Height x Non-wrapper 1975E 0.12 -0.34 A A
Leaf Number 1975L -0.96 0.98 A 0.53
Total Non-wrapper Leaf 1974L 0.88 0.85 A A
Weight x Non-wrapper l975E 0.51 0.95 A A
Leaf Size 1975L B A A 0.34

 

 

 

 

 

 

*Reciprocal cross
A - Correlation estimates greater than 1.0
8‘ Correlation estimates less than -1.0

123
and 1 was observed for this correlation. The rgx values for these rela-
tionships were 0.97 a 0.38 and 1.00 e 0.18 respectively (Table 69).

Stalk Size .

Significant differences for stalk size exist between cultivar means
and between interaction means for cultivar x inbreeding and cultivar x
plantings (Table 70). Since there were no differences due to inbreeding
or plantings. stalk size appears to be genetically fixed in the cultivars
selected and uneffected by the environment. Similar results could be
obtained if dominant genetic systems were absent. Baby Head. Badger
Ballhead. P.I. 215514 and Red Danish each were significantly different
for stalk size. Chieftain Savoy was not different from Badger Ballhead
or Red Danish but was different from Baby Head and P.I. 215514 (Table 71).
Due to an increase in the stalk size of Red Danish after 1 generation of
inbreeding the cultivar x inbreeding interaction was significant. The
other cultivars showed no difference in stalk size due to inbreeding
(Table 71). The significance of the cultivar x plantings interaction
was due to a decrease in stalk size between the 1974L and 1975L plant-
ings for Red Danish. During this period the other 4 cultivars showed
no difference in stalk size (Table 72).

The inheritance of stalk size has not been previously reported.
Small stalk size appears to be incompletely dominant in green cabbage
crosses. F1 population means of Baby Head x Badger Ballhead also suggest
incomplete dominance for small stalk size (Tables 11-13). Tables 73 and
74 show frequency distributions in percent for populations involved in h
the cross Baby Head x Badger Ballhead. Powers et a1. (23) partitioning
tests were applied to these distributions to determine the number of

genes controlling stalk size and to define the boundary between small and

124

Table 70. Analysis of variance table for stalk size in kilograms.

 

 

 

Source
of Mean

Variance DF Square F*
Replication (R) 2 0.01 1.15

. h **
Cultivar (C) 4 0.76 84.49
Inbreeding (I) 1 _0.01 0.24
c x 1 4 0.02 ‘ 2.56*
Plantings (P) 2 .0.02 (1.94
c x P ‘ 8 0.02 , 2.22*
I x P 2 0.01 0.86
C X I X P 8 0.01_ 0.88
Residual Error 58 0.01

 

a: shit ' .
*F test significant at the 5% . or 1% level.

125

Table 71. Cultivar x inbreeding interaction means for stalk size

in kilograms.

 

 

 

 

Inbreeding 'Cultivar
Cultivar - P1 S1 Means
Baby Head 0.08 a* 0.06 a 0.07 W
Badger Ballhead , 0.25 a 0.22 a ‘0.24 X
P.I. 215514 0.65 a 0.61 a 0.63 2
Red Danish 0.27 a 0.40 b 0.34 Y
Chieftain Savoy 0.29 a 0.30 a 0.30 XY
Inbreeding Means 0.31 A 0.32 A '

 

*Mean separation. within rows. by Duncan's Multiple Range Test at

the 5% level.

126

Table 72. Cultivar x planting interaction means for stalk size
in kilograms.

 

 

Plantings Cultivar

 

 

Cultivar 1974L l975E 1975L Means
Baby Head '0.06 a* 0.09 a 0.06 a 0.07 W
Badger Ballhead 0.27 a 0.24 a 0.19 a . 0.24 x '
P.I. 215514 0.59 a 0.60 a 0.70 a 0.63 2
Red Danish 0.41 b 0.34 ab’ 0.26 a 0.34 Y
Chieftain Savoy 0.34 a 0.32 a 0.23 a 0.30 xv
Planting Means ‘ 0.34 A 0.32 A 0.29 A

 

*Mean separation. within rows. by Duncan' 5 Multiplellange'Test at
the 5% level.

127

 

 

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meecw s2 w~2m.e2e2m

 

 

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m .—m .usmcum

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128

 

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maeco se w~2m e2e2m

 

 

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mseesepa ememp wse cow uswocwa sm msoeuse2cumeu w~2m epeum.~e use.2m .uswcee .em w2eee

129
large stalk size classes. Powers et a1. (23) formula. (FZ/PZ)’ was ap-
plied sequentially from the recessive ends of both the F2 and P2 distri-
butions. Using the 1975E planting of Baby Head x Badger Ballhead as an
example recessive frequencies of 4.8/7.8 or 0.62 forthe 900+ gm class.
5.6/12.8 or 0.44 for the 440 gm class. 7.6/20.5 or 0.37 for the 400 gm
class. 0.2/23.1 or 0.40 for the 360 gm class. 12.4/33.3 or 0.37 for the
320 gm class. 16.0/48.7 or 0.33 for the 280 gm class. 26.2/61.5 or 0.43
for the 240 gm class. 38.1/82.0 or 0.46 for the 200 gm class and 70.6/97.4
or 0.72 for the 160 gm class were produced. The large increase in fre-
quency between the 200 gm class and the 160 gm class suggests a dividing
point of 160 gm might be used to separate F2 data into phenotypic classes.
The mean of frequency estimates between the 200 gm and 900+ gm classes
was 0.43. This frequency compared favorably toha 7/16 frequency of 0.44.
A 2 gene 9:7 model with a dividing point of 160 gm seems appropriate. The
reciprocal F2 population of this cross produced the same estimate of 160
gm as a dividing point but produced a lower frequency estimate for the
recessive class. The mean of recessive class frequency estimates was 0.21
which suggests a monogenic 3:1 model with a 160 gm division point. Since
the distribution of both parents. Baby Head and Badger Ballhead. over-
lapped. a weighting procedure was used to account for misclassifications .
in F2 segregation ratios. For the 1975E planting of Baby Head x Badger
Ballhead the net overlap around the 160 gm class was 18.0% (Table 73).
The expected number of recessive plants in the F2 population can be cal-
culated as ((100-18)/100)(0.4375)(252) = 90.4. The expected ratio was
161.6:90.4. The expected number of recessive plants in the reciprocal
F2 population was ((100-18)/100)(0.25)(248) = 50.84. The expected ratio
was 197.16:50.84 (Table 75). Similar results for other crosses suggested
stalk size might be controlled by 2 loci. The simplest system available

130
to insure digenic inheritance and explain differences between the culti-
vars was a 2 locus system with 5 alleles per loci. The loci were desig-
nated H and I since the Small stalk size of the base cultivar. Baby Head.
was dominant in most cases (Tables 11-13). The genotypes of each culti-
var and F1 in this study are Shown in Tables 16 and 17.. Data from 1975E
green cabbage crosses suggest H5 was dominant to H and a was dominant

to h3 and h4. Inheritance of the I locus was similar. Red x green cab-

bage data suggest H5 and H4 were dominant to H and 82 was dominant to

2

h3 .. The I locus exhibited a similar inheritance pattern. Allelic in-

heritance patterns for both 1975L green and red x green cabbage crosses

suggest that H was dominant to h3. h and.h5? while H was dominant to H

4 3
The I locus was inherited in a similar man-

2
and H2 was dominant to as.
ner for both sets of crosses. Chi-square homogeneity tests suggest

that 1975E reciprocal F2 distributions of Baby Head x Badger Ballhead
were Significantly different for stalk size (Table 73). ‘The popula-
tions were not pooled. The F2 segregation ratio of 186’ small stalk
size:66 1arge stalk size produced using a 160 gm dividing point. fit
(P=0.47) a 9:7 weighted expected ratio of 161.6:90.4.. Homozygous reces-
-sive genotypes at either loci conditioned the same phenotype. large stalk.
size. while 1 dominant allele. H or I . at each locus conditioned small

wstalk size. Recessive alleles. h and 13, were contributed by Badger

3
Ballhead. The reciprocal F2 segregation ratio of 192:56 fit (P=0.44)

a 3:1 weighted expected ratio of 197.2:50.8 (Table 75). Small stalk
size was conditioned by 1 dominant H allele regardless of the I locus
genotype. The recesSive genotype. h3h3i313. from Badger Ballhead con-
ditioned a large stalk size phenotype (Tables 16-17). Backcrossing the

reciprocal F1 to Badger Ballhead produced a segregation ratio of 14:19.

131

Table 75. Chi-square test for goodness of fit to the postulated
model for stalk size in the F2 generation (l975E planting).

 

 

2

 

 

 

 

 

 

Cross Observed , Expected M0del X P

Baby Head x -

Badger Ballhead

F2 156: 96 161.60: 90.40* 9: 7 0.5400 0.47
* .

RF2 192: 56 197.16: 50.87 13: 1 0.6587 0.44

Baby Head x

P.I. 215514

F2 152:107 145.69:113.3l 9: 0.6252 0.43

RF2 254:197 253.69:197.31 ' 9: 0.0009 ' 0.98

Baby Head x

Chieftain Savoy a

52 59 133 61.71:130.28: 1: 3 0.1759 0.67

RF2 63:134 63.32:l33.68 l: 3 0.0024 0.96
*

Pooled 122:267 125.03:263.96 l: 3 0.1086 0.74

Red Danish x

Badger Ballhead

52 45:164 53.74:155.26* 1: 3 1.9146 0.17
* .

RF2 36:118 39.60:ll4.40 l: 3 0.4406 0.51
* ,

Pooled 81:282 93.32:269.66 l: 3 2.1971 0.14

Red Danish x

P.I. 215514

F2 126:185 l36.06:l74.94* 7: 9 1.3230 0.25

Red Danish x

Chieftain Savoy
*

F2 6:101 14.31: 92.69* 1:15 0.0754 0.79

RF2 6: 96 13.64: 88.36* 1:15 0.0235 0.88

Pooled 12:197 13:06:195.94 1:15 0.0922 0.77

 

* .
-’Weighted

132
Backcrossing to a 3:1 model should produce an expected ratio of 16.5:16.5.
The data fit (P=0.41) the expected 1:1 model. The 9:7 segregation ratio
for the 1975L F2 population suggests that 1 dominant allele at each 10-
cus is required to condition a small stalk size phenotype. The ratio
also requires that the homozygohs recessive condition at either loci
produce the same. large stalk weight. phenotype. Using a division
point of 140 gm. the 1975L F2 population. produced a segregation ratio
of 89:41 (Table 74). The data fit (P=0.16) a weighted segregation ratio
of 81.3:48.7 (Table 76). The 1975L reciprocal thpopulation produced
a 170:41 segregation ratio. The data were compared (P=0.49) to a 3:1
.weighted expected ratio of 165.9:45.1 (Table 76). The action of 1 domi-
nant gene. H. conditioned a small stalk size phenotype in this planting.
The recessive genotype h3h3i313 produced a large stalk size phenotype
(Tables 16-17). Dominant alleles. H and I. were contributed by Baby '
Head. 1

Baby Head x P.I. 215514 F data from 3 p1antings. 1974L. 1975E. and

1
'1975L. suggest incomplete dominance for small stalk size (Tables 11-13).
F2 data from both the 1975E and 1975L plantings also suggest small stalk
size was dominant (Tables 77-78). The 1975E Baby Head x P.I. 215514 F2
population with a division point of 190 gm produced a segregation ratio
of 192:56. A 9:7 weighted expected ratio of 181:67 was compared (P=0.12)
to the observed data (Table 75). Homozygous recessive alleles at eith-

er loci conditioned a large stalk weight phenotype. One dominant allele
at each loci. HeI_: was required for a small stalk weight phenotype.
Chi-square homogenity tests suggest that the 1975L F2 populations could

be pooled. The pooled F2 segregation ratio was 327:85. A division point

of 260 gm was used. The data fit (P=0.33) a 13:3 m0del of 335:77

133

Table 76. Chi-square test for goodness of fit to the postulated
model for stalk size in the F2 generation (1975L planting).

 

 

 

 

 

 

 

Cross _Observed Expected Model X2 P
Baby Head X‘
'Badger Ballhead 1
1 *
52 89: 41 81.34: 48.66* 9: 7 1.9283 0.16
RF2 170: 41 165.87: 45.13 3: 1 0.4815 0.49
Baby Head x
P.I. 215514
52 130: 38 136.50: 31.50 13: 3 1.6508 0.20
852 197: 47 198.25: 45,75 13: 3. 0.0421 0.34
Pooled 327: 85 334.75: 77.25 13: 3 0.9569 0.33
Baby Head x
Chieftain Savoy
52 120: 95 127.91: 87.09* 9: 7 1.2085 0.27
~ * .
RFZ 116: 62 105.90: 72.10 9: 7 2.3782 0.12
*
Pooled 236:157 233.81:159.19 9: 7 0.0506 0.82
'Red Danish x
Badger Ballhead
52 92 199 104.62:186.38: 1: 3 2.3784 0.12
RF2 . g 11: 19 10.79: 19.21* 1: 3 0.0066 0.94
Pooled 103:218 115.41:205.59 1: 3 2.0838 _ 0.15
Red Danish x
Chieftain Savoy
52 ', 39:160 35.08:l63.92: 1:15 0.5318 0.46
852 42:182 39.49:184.51*' :1:15 0.1941 0.66
Pooled 81:342 74.57:348.43 1:15 :. 0.6738 0.41

 

= Weighted

134

No. 222. .2 en:

 

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e.e2 .H 2.mme 2.e e.e .e.ee e.2 e.ee e.2 e.22 e.e e.e e.2 O2 222 x eev

m.ae .H e.eme e.2 m.e e.e. o.e2 e.e 2.2M e.e2 e.e2 e.e e.m m.e .ee 2ma x 2e2

2.e2e.H e.eee Ne.e2 2.22 2.22 e.e e2 2ee2 e2me2e .2.e

e.ee .H m.ee, e.e m.ee e.mm ea 2222 eewe cewe
sew: +eee oee oem ewe e2e oee ewe oe2 ee2 ,ee2 ee meew2e

mme2u co 222?; cwssz
maecu s2 w~2m eeeum

mo .02

 

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.e2ee2e .2.2 e ewwe enee anoco web co

ee2cww2e w~2n e2eew we eee 2e .wewcwa .22 w2ee2

135

 

 

2.222.“ c.2e2 e.2 e.e e.e e.e e.2 e.22 ~.e e.m2 e.22 esem e.e eee 22a x nae

2.ee2.H e.ee2 e.e e.e e.e e.e e.2 . m.e m.e 2.e2 2.e2. 2.ee 2.2 ee2 ea 2ea x 2e2

e.me2.fl e.eee e.a 2.m e.e e.e2 e.e N.NN e.e2 8.8 e.e e.e 2e 22a x nee

e.ee2.u m.e2~ 2.e e.m 2.e e.e 2.e e.22 2.m2 m.e2 m.e2 e.oe m2 2me x 222

2.eee.u 2.2ee e.ee 2.me 2.2 m2 2e22 e2ee2e .2.a

e2muee2 e2 eemeee 8 itesess
sews +eee eee oee oem e28 eee . ewe ee2 ee2 wee2 ee Hwewwn

mme2u mo H2524 cwsa:
msecw s2 w~2m e2eum

 

 

.epmmpm .2. e x uew: anem mmocu wsu co
ms22se2a em2m2 wsu com uswucws s2 mso22=s2c2m2u w~2m e2e2m we use 2; .uswces .m2 w2seh

136
(Table 76). A 13:3 ratio was produced when a dominant genotype at 1
locus and the recessive genotype at the other each produce the same.
small stalk weight, phenotype. The recessive genotype h4h4ii conditioned
a large stalk weight phenotype. Recessive alleles h4 and 14 were con-
tributed by P.I. 215514.

All 3 plantings of the F1 population. Baby Head x Chieftain Savoy,
suggest incomplete dominance for large stalk size. Both F2 populations
of Baby Head x Chieftain Savoy grown in each planting period. 1975E
and 1975L, were pooled as suggested by chi-square homogeneity tests.

A division point of 160 gm was used to separate the F2 data into class-
es. Pooled F2 data from the early planting suggest a 1:3 weighted ex-
pected ratio of 125:264 should be compared (P=0.74) to an observed ra-
tio of 122:267 (Table 75). Large stalk size was controlled by a single
dominant gene, H5. ‘The genotype ”5+-~ conditioned large stalk size
while recessive small stalk size was produced by the genotype hhii.
Recessive alleles were contributed by Baby Head while alleles from Chief-
tain Savoy were dominant. A segregation ratio of 26:29 was produced

by backcrossing the reciprocal F1 to the recessive cultivar Baby Head.
The data showed good fit (P=0.70) to the expected ratio of 27.5:27.5.
Backcrossing the F1 and reciprocal F1 to Chieftain Savoy produced
segregation ratios of 4:28 and 2:13 (Table 79). Ratios of 0:33 and 0:15
were expected. F2 data from the late planting show dominance can be
effected by planting time. The observed ratio of 236:157 fit (P=0.82)

a weighted expected ratio of 234:159 derived from a 9:7 model (Table
76). Phenotypic Classes were divided using a 140 gm division point.
Small stalk size was controlled by the presence of 2 dominant alleles

at different loci. Large stalk size was cantrolled by the presence of

137

 

.H m.mm2 e.m N.m m.u N.NP m.n m.o2 ¢.m ¢.om e.m

3 2a 12.. 2.:

 

2.o~2
N.m~2.u c.2om 2.2 o.o~ 2.2 N.mm 2.2 2.22 2.2 2.2 m2 ma x 22; x may
2.222.“ o.mm~ _ ~.m wm.w2 ‘~.o 2.22 ~.m 2.2 N.o ~.o N.m 2.2 mm ‘ ma 2 Ame x 2av
2.222.“ 2.~m~ 2.2 2.2 o.2 2.2 . 2.2 «.22 ~.o2 m.o2 2.2 m.- m.o 222 Nd 22a x.mav
~.2m .H 2.m2~ 2t~ 2.2 2.2 2.2 m.~ m.- 2.2 0.22 2.22 2.22 2.2 ~22 Na Ame x 2av
3232.23 92 ca «.2 32 3 2:8. 92 2.22 ma 2.2 . 8 22:22:
o.oo2.fl 2.22m _~.~ 2.22 2.2. ~.m2 2.2 2.22 2.22 2.2 2.2 ~.a2 mm 22a x.2av
o.mm2.w 2t-m 2.22 e.o~ a.~ 2.22 2.22 2.e2 w.m m.~ m.m em Away sasmm c2222a2gu,
2.m~ .H m.oa 2.N 2.22 2.22 No. . 22av new: 2aam
cam: +oom owe com cum O22 com o- om2 om2 922 cm wwcwwn _

mmm2u ca 22524 gang:
macaw =2 m~2m x2aum

 

 

.ao>mm =2m22m2su x new: An
mc22=m2a mm2m2 ms» co2 “smegma =2 mco2uzn2c2m2u m~2m x2m2m mmoguxuma ecu

mm mmocu mg» 20
m .2; .ucmcmm

.mn open»

138
a single homozygous recessive genotype at either loci. Dominant alleles.

a and I, were contributed by Baby Head. The difference in dominance
direction noted between these 2 plantings suggest a 30 day difference
in planting cabbage might expose the F2 population to conditions that
change the phenotypic expression of certain genotypes (Table 80).

Red Danish x Badger Ballhead Fl data from the 1974L. 1975E and
1975L plantings suggest overdominance for large stalk size (Tables
11-13). Chi-square homogeneity tests suggest that F2 populations of
this cross can be pooled within each planting. The pooled 1975E F2
population produced a segregation ratio of 81:282 using a dividing
point of 290 gm (Table 31). The data fit (P=0.14) a weighted expected
ratio of 93:270 from a 1:3 model (Table 75). Large stalk size was con-

trolled by the presence of a single dominant gene. H The recessive

.2'
genotype h3h3i3i3 conditioned small stalk size. The dominant allele
H was contributed bvaed Danish (Table 81). BackcrOssing the F1 to

2
the recessive cultivar Badger Ballhead produced a segregation ratio of
3:9. An expected ratio of 6:6 was compared (P=0.09) to the observed
data. Backcrossing the reciprocal F1 to Red Danish produced a segre- ,

gation ratio of 0:47. The data support an F2 model of 1:3. An ob-
served segregation ratio of 103:218 produced by the pooled 1975L F2
population fit (P=0.15) a 153 weighted expected ratio of 115:206
(Table 76). A division point of 200 gm was used. Large stalk genes
from Badger Ballhead were dominant. Large stalk size was controlled
by the presence of 1 dominant H3 allele. The recessive genotype
h2h21212 conditioned small stalk size. Recessive alleles were con-
tributed by Red Danish (Table 82).

139

 

 

m.om .H 2.222 0.2 2.0 N.N QLN 2.2 N.e e.m 0.2 e.me m.2~ w22 Nd 22¢ x may

2.22 .H 2.oe2 2.2 ~.2 m.~ 2.2 N.e .~.o2 2.2 2.2 2.2e 2.o2 m2N Ne Ame x 2ev

n.2c2.u 2.222 e.m 2.2 2.2 2.2 e.e o.- m.@ 2.22 e.e mm 22a x may

m.em .H c.2m2 0.22 o.o~ o.o~ o.o~ o.m~ om. “me x 2ev

m.~m .u 2.2NN. . m.m e.e e.22 m.~ 0.22 m.o~ 0.2 m.e2 2.2 me Amev sesem.e2eeee2eu

e.2m .u a.O2 . 2.2 2.22 m.em mm . A2ev new: 2522
eeez +oom owe com 0N2 O2N Dem ONN 022 cc. oe2 on ”wemwn

mmm2u 2o 22524 swan:
mange =2 m~2m x2mpm

 

 

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mc22=m2q 4m2m2 mc2 co2 pcmucma =2 mco2pan2eum2u mN2m x2m2m we use 2; ucmcea .om m2am2

140

 

 

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2.82.222: 92 2.2 we 2.2 2.2 2.2m 3 ed 2.: 23 0.2 $2 mien. xmt
m.§u2.eme 2.m .3. 3: 92 2.3 2.2.2 we #2 2.2 ea 92 SN mime xeev
twig: 2.2 N.N 2.2 :2 2.22 2.8 3 2.2 Te. N.N 8 3x»:
Ndfiueéem 2.N o.2 . we 2.: 92 SN 2.2 2.2 o.2 em 3 :5
22.82.2258 3 3 3 N52 3: 32 32 2.8 2.22 8 3:22:23 .583
932.“.ng 2.2 2.2. 3 ea 98 :2 3w 2.2. 2.m 2.2. 98 mm 3: 5.23 82.
5e: .68 Se 82 2.8 OS 2; 82 SN

mmm2u 2c 22524 Loan:
msmcw :2 m~2m x2mum

oem com .oop mace—a
_ 2o soz

 

 

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.ummz22em Leanna x gm2cmo emu mmogu «:2 20
2c «~2m x2e2m mmoeuxoea wen we .2; .2cwcee .2w m2ne2

141

 

 

.mmm2o 2c 2252; swan:
mseew :2 m~2m x2eum

,2.emN 2.02 2.2 2.e2 2.2 o.o2 2.2m om Nd Aee x Nev
2.mo2.w o.~2 e.m o.o~ e.m2 2.22 ~.e2 2mm . Ne Ame x eev
m.me .H m.o2 m.o2 m.o2 2.2m .e.2m 2.2 m2 flea x may
.m.em .H e.2~ 2.m m.em c.22 2.22 mm Ame x eev
2.22 .H 2.0 2.e e.m2 e.e~ m.e~ m.em me Away eeee22em eeeeem
t$.Ht82 <2 22 92 #8 9% gm Afvfizaeg

ooe cum cam oeN oo~ oe2 meee2a

2e.ez

 

 

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m=2uce2a AmNmF asp co2 peduema :2 mco2pzn2e2m2e m~2m x2e2m we use 2; .ucmeem .Nm w2QN2

142

1974L and 1975E plantings of Red Danish x P.I. 215514 F1 suggest
overdominance for large stalk size (Tables 11-13). Data from the
1975L planting show stalk size to be inherited in an additive manner
(Table 84). Only the F2 population of Red Danish x P.I. 215514 was
grown in the 1975E planting. Although both cultivar distributions were
nearly identical the F2 data showed transgressive segregation for large
stalk size. The data were compared to a 7:9 model which assumed that
H was dominant to h

2 4
of 600 gm an F2 segregation ratio of 126:185 was produced (Table 83).

and :4 was dominant to 12. Using a division point

This ratio was compared (p=o.23) to an expected ratio of 136:175 (Ta-
ble 75). Large stalk size was controlled by the presence of H2 and I4.
Homozygous recessive genotypes at either loci conditioned a small
stalk size phenotype.

All plantings of Red Danish x Chieftain Savoy F1 suggested over-
dominance for large stalk size (Tables 11-13). Chi-square homogeneity
tests suggestedpooling F2 populations of Red Danish x Chieftain Savoy
within each planting. Transgressive segregation for large stalk size
was noted in the 1975E planting (Table 85). Using a 320 gm division
point the F2 distribution produced an observed segregation natio of
12:197. The data show good fit (P=D.77) to a 1:15 model (Table 75).
The presence of either dominant allele conditioned large stalk size
while the recessive genotype hshsisis conditioned small stalk size.
Backcross segregation ratios suggest differences between reciprocal
Fl's. The dominant cultivar, Red Danish, was crossed to reciprocal
Flplants and produced segregation ratios of 0:18 (F1) and 28:9 (RF1)'
Ratios of 0:18 and 0:37 were expected. The difference between the

1 observed RF1 backcross ratio and its expected ratio suggest differences

143

 

we “as x eat

 

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e.mm2.H 2.m~e 2.22 N.22 N.o~ 2.m~ 2.22 2.2 2.2 N.N 2.2 . .am “as x may

m.m-.fl 2.eme N.e~ 2.22 m.e2 m.22 o.o~ e.m ~.2 22 22a x eav

2.e2~.u e.eoe N.NN 2.2 m.2~ 2.2m c.22 e.m m2 Anew e2mm2~ .2.a

o.m-.H 2.222 2.2 2.2 m.~ e.m o.o~ e.e2 e.w 2.2 2.2. e.~ o.o~ mm Aeav ea2eeo com
eaoz +oom com OO2 ace com ooe own oaw cam com. oe2 moea2a

mmm2u 2o 22524 Loan:
manta 2.2 m~2m x2o2m

2o .oz

 

 

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.e2mm2m .2.a x ea2eaa cog aaoco ego 2o
2=a2coa2a o~2a x2a2a Na eee 2a .2eocaa .mw o2ee2

144

 

e.2m 2.2 2.2m 2.2 2.22 , 2.22 e2 fies x may

 

2.-~.H 22~2e 2.2

2.222.“ m.2em m.~ m.~ 2.2 ..a.m2 m.a~ 2.2 2.22 e.e 2.m m.~ ea 22a x ea.

_.maN.u_2.2me m.wm 2.2 .2.2 2.om ,2.2 2.2 m2 Amav e2mm2m .2.a

2.Ne .H e.em2 2.2 m.m2 m.~2 e.m~ 2.22 mm Aaav ea2eac eox
eeoz +oom com OO2 com com ooe own cam cam com oe2 _apee2a

2o .02

mmm2u 20 22524 Loan:
manta =2 m~2m x2epm

 

 

.e2mm2m .2.a x cm2cma cum mmogu use 2o
mc2uce2a 4m2m2.m;2 s02 ucwusma :2 m=o2uan2cpm2u o~2m x2mum.2m use geese; .ew m2nm2

145

 

982H982 98 . 2.8 92 98 98 9: 22 ea 15 xmn:
9:82.982 98 98 9.8 2.: 9m . . 82 ea 12.. 2.:
9.28“ 2.28 9: 2.2 92 98 98 92 92 92 9: NS 8:: x25
953982. 92 9e 98 98 9: 9e 9e 9e 98 2.2 2 22 mime in:
983982. 92 92 9: 98 92 9e 98 .92 9: 8 3.2.:
98:23.80 2.: 92 98 98 2.: 2a 92 8 22:5:
982.2982 9m 9m 9: 9m 9: 9: 9: 92.2 9: 8 223.233 2.2.3235
9882.92... 2.2. 2.2.) 99 98 98 :2 9m 5 2.2 98 98 mm 22.: 8.252. as.

 

cam: .+oom com com com com cow omm omw oem cow oo2 mpcm2a
. . , mo .02

mmo2u 2c 2252; song:
manta =2 m~2m x2m2m

 

 

_ .xo>em c2e22w2zu x :m2ceo no mmocu on» 2o .
mc22=e2a 4m2m2 we» c02 uemucoa :2 mco2usn2s2m2u m~2m x2e2m mmOtbxumn new we .2; .2cmcea .mm m2ae2

146 .
between F1 and RF1 gametic transmission. Pooled F2 data for the 1975L

planting suggest Chieftain Savoy contributed dominant genes for large
stalk size (Table 86). A 1:15 weighted theoretical ratio of 75:348 was
compared (P=0.43) to an observed segregation ratio of 81:342. A divi-
sion point of 200 gm was used to separate F2 data into phenotypic
classes. The model suggests that the genotype hzhziziz was recessive
for small stalk weight while all other gene combinations produced.large
stalk size (Table 16).

Green cabbage cross stalk size heritability estimates between 0 and
one ranged from 0.28 to 0.96. The mean heritability estimate was 0.36
i 0.24. The data show a 50% difference in reciprocal cross heritability
estimates for the 1975L planting. This difference may be due to mater-
nal influences rather than to environmental causes. The 5: estimate
was 0.40 t 0.23. Only 1 red x green cabbage heritability estimate fell
within the D to 1 heritability range. That heritability estimate was 0.57
s 0.60. The Biestimate for red x green cabbage crosses was 0.81 1
0.40' (Table 30). ‘

Genetic correlations between stalk size and non-wrapper leaf number
and non-wrapper leaf size exhibited a wide range of estimates. Green.
cabbage Fl estimates of genetic correlation between stalk size and non-
wrapper leaf number ranged from 0.37 to 0.67 with a mean of 0.60 1 0.55
(Table 87). The ng estimate for the relationship was -0.58 a 0.42.

The mean genetic correlation between green cabbage F1 stalk size and
~non-wrapper leaf size was 0.93 1.0.32. Estimates for this genetic corre-

lation ranged from 0.21 to 0.98 and produced an-E x estimate of 1.00

9
0.04 (Table 87). Red x green cabbage F1 mean genetic correlations be-

tween stalk size and non-wrapper leaf number and non-wrapper leaf size

147

 

 

92.23988 9: 9: 90 9e 92, 98 .92 2.: 92 98 2.2 88 88 :22. em:

.82..“ 2.82 9o 9m 9m 93 98. 9e 2.: 9m 92. 9o 8: 812.. x 2.:

9.8 .2928 92 2.8 98 98 9a 2.2 2.2 8 A: x ma:

923928 922 2.2 9: 98 92 9mm 8 Amaxeav

98 H.988 2.2 9m 9m 98 98 98 98 me 22.: 882 £38.25

9% H98— 28 92 98: 98 98 82 22.: 5.28 88
eaoz +oom com OO2 _ com com Doe omm 028 028 com oe2 ”Newwm

mmm2u 2c 2252; song:
mango :2 m~2m x2mum

 

 

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mc22cm2n 4m2m2 8:2 co2 pauucma :2 mco2uan2cpm2u m~2m x2e2m Nu new 2a pause; .mw m2nm2

148

Table 87. Genetic correlations between stalksize, non-wrapper
leaf number, efficiency index and non-wrapper leaf

size in cabbage.

 

 

r

 

 

 

 

 

Traits Plantings Green Crosses Red Crosses
R* ' R*
Stalk Size x l974L 0.63 0.37 -0 l2 0.08
Non-wrapper Leaf Number l974E B B A 0.96
l975L 0.67 A -D.74 0.89
Stalk Size x 1974L A A 0.33 0.84
Non-wrapper Leaf Size “l975E 0.98 0.88 0.99 A
l975L 0.2l A -0.27 -0.09
Non-wrapper Leaf Number x l974L A 0.96 . B B
Efficiency Index l975E 0.93 A 0.96 0.80
, 1975L B -0.02 B' A
Non-wrapper Leaf Number x l974L 0.49 0.40 A -0.80
Non-wrapper Leaf Size l975E -0.84 -0.65 A A
l975L 0.98. 0.78 -0.65‘

 

-0.96

 

 

 

 

*Reciprocal cross
A

= Correlation estimate greater than 1.0
B = Correlation estimate less than -1.0

 

149 , _
were 0.88 s 0.44 and 0.91 s 0.43 respectively. The ranges associated

with these rg values were -0.74 to 0.96 and -0.27 to 0.99. fig

mates for these relationships were 0.89 s 0.42 and 0.94 s 0.37 respec-

x esti-

tively. Both green and red x green cabbage crosses produced a wide
range of genetic correlation estimates. Differences between estimates
may be a function of maternal inheritance as shown in red x green cab-
bage F1 correlations between stalk size and non-wrapper leaf number
estimated during the 1975L planting or a function of environment

as shown in red x green cabbage F1 correlation estimates between stalk

size and non-wrapper leaf size for the 3 plantings (Table 87).

Non-Wrapper Leaf Number

Analysis of the results for 1 generation of inbreeding for non-
wrapperleaf number suggests significant differences exist between
cultivars, inbreeding and plantings. Significant interactions also
exist for cultivar x inbreeding, cultivar x plantings and inbreed-
ing x plantings (Table 88). Four cultivars. Baby Head, Badger Ballhead;
P.I. 215514 and Chieftain Savoy were each significantly different for
non-wrapper leaf number. Red Danish was not different from Badger
Ballhead or Chieftain Savoy but was significantly different from the
other 2 cultivars (Table 89). The S1 plants produced an increase of
1 non-wrapper leaf (Table 89). Although this increase was significant
and suggests that non-wrapper leaf number may not be genetically fixed
in these cultivars the small increase in non-wrapper leaf number may
not justify time spent in added inbreeding for this trait prior to use
in a genetic study. A comparison of both the 1974L and 1975E plantings
shows no difference in non-wrapper leaf number, however, the 1975L

planting showed a significant decrease in non-wrapper leaf number when

150
Table 88. Analysis of variance table for non-wrapper leaf number.

 

 

Source

 

of Mean
Variance DF . Square F*
Replication (R) 2 0.74 0.22
Cultivar (C) 4 512.02 :32.lo**
Inbreeding (I) l l9.l9 5.70*
c x I 4 28.07 8.34**
Plantings (P) 2 230.52 68.48**
c x p 8 30.13 - 8.95**
' ' **
I x P. 2 16.36, . 4.86
c x 1 x p 8 6.33 l.88
Residual Error 58 3.37

 

*F test significant at the 5%*, or l%** level.

151

Table 89. Cultivar x inbreeding interaction means for non-wrapper
leaf number.

 

 

Inbreeding . cultivar

 

 

Cultivar P1 S1 Means
Baby Head ' 8.54 b* 6.47 a . 7.50 v
Badger Ballhead . 14.71 a 16.67 b _ 15.69 x
P.I. 215514 21.90 a 22.76 a 22.33 2
Red Danish 14.08 a 18.56 b 16.32 xv
Chieftain Savoy 17.51 a 16.9l a ‘ l7.2l Y
Inbreeding Means 15.35 A l6.27 B

 

*Mean separation, within rows, by Duncan's Multiple Range Test at
the 5% level.

152
compared to the previous plantings (Table 90). Cultivar response to
inbreeding in this study can be divided into 3 groups:.
(1) Baby Head, showing a reduction in nonewrapper leaf
number due to 1 generation of inbreeding.
(2) ~Badger Ballhead and Red Danish. showing an increase
in non-wrapper leaf number due to 1 generation of
inbreeding.
‘(3) P.I. 215514 and Chieftain Savoy uneffected by 1

generation of inbreeding (Table 89).
The significant cultivar x plantings interaction suggests that these
cultivars responded differently during the 3 planting periods. Baby
Head's non-wrapper leaf number increased between the 1974L and 1975E
plantings and decreased between the 1975E and 1975L plantings. Both
Badger Ballhead and P.I. 215514 showed no difference in non-wrapper
leaf number between the 1974L and 1975E plantings but showed a reduce
tion in non-wrapper leaf number between the 1975E and 1975L plantings.
Red Danish and Chieftain Savoy both produced significantly less non-
wrapper leaves between the 1974L and 19755 plantings and showed no
change between the 1975E and 1975L plantings (Table 90). The sig-
nificance of the inbreeding x plantings interaction was due to a sigé
nificant increase in non-wrapper leaf number due to inbreeding in the
1975L planting. No differences were shown for other plantings
(Table 91).

In 1934 Pearson (20) reported that in cabbage few non-wrapper

leaves were dominant to many leaves. He also reported on several .
modifying factors. The 1974L F1 planting of Baby Head x Badger Ball-

head supports Pearson's (20) conclusions. The data show an incomplete

153

Table 90. Cultivar x planting interaction means for non-wrapper

leaf number. '

 

 

 

 

Plantings Cultivar
Cultivar l974L l975E l975L Means
Baby Head 7.16 6* 12.86 c 2.49 a 7.50 w
Badger-Ba11head 18.57 b 17.10 b 11.40 a ,l5.69 x
P.I. 215514 23.72 b 23.48 b 19.80 a 22.33 2
Red Danish { l8.26 b l5.90 a l4.Bl a 16.32 XY
Chieftain Savoy 20.57 b l6.44 a l4.62 a l7.2l Y
Planting Means 17.66 B '17.16 B 12.62 A

 

*Mean Separation, within rows, by Duncan's Multiple Range Test at

the 5% level.

 

 

,[. Illill... J

154

Table 9l. Inbreeding x plantings interaction means for
non-wrapper leaf number.

 

 

 

 

Plantings Inbreeding
Inbreeding ' l974L l975£ l975L Means
111 17.12 a* 17.47 a 11.46 .a I 15.35 Y
51 l8.l9 a 16.84 a 13.78 b. 16.2.7 2 '4
Planting Means l7.66 B 1 l7.l6 B l2.62 A

 

*Mean separation, within columns, by Duncan' 5 Multiple Range Test
at the 5% level level.

155

. dominance for non-wrapper leaves. Data for the 1975E and 1975L F1
plantings of this cross do not agree with Pearson's (20) conclusions.
The data suggest that for the 1975E planting many non-wrapper leaves
were overdominant. However. in the 1975L planting the F1 population
showed incomplete dominance for many leaves and the reciprocal F1
population showed incomplete dominance for few non-wrapper leaves
(Tables 11-13). In contrast. all of the F2 populations of green cab-
bage crosses showed dominance for few non-wrapper leaves (Tables 94-95).

Tables 92 and 93 show frequency distributions in percent for

 

populations involved in this cross. Powers et a1. (23) partitioning
tests were applied to these distributions to determine the number of
genes controlling non-wrapper leaf number and to define the boundary
between few and many non-wrapper leaves. Powers et a1. (23) formula
(Fz/Pz) was applied sequentially from the recessive ends of both the

F2 and P2 distributions. Using the 1975E Baby Head x Badger Ballhead

F2 population (Table 92) as an example recessive frequencies of 5.2/30.8
or 0.49 for the 21 leaf class, 23.5/43.6 or 0.54 for the 19 leaf class,'
40.9/69.2 or 0.59 for the 17 leaf class and S6.7/79.5 or 0.71 for the

15 leaf class were produced. The rapid increase in frequency between
the 17 and 15 leaf classes suggested a dividing point of 15 leaves

would be appropriate to separate the F2 data into 2 classes. few (0-15)
and many (16+) leaves. A mean estimate of 0.54 was produced by

classes 17-21. The 1975E reciprOcal F2 population produced the same
dividing point. 15 leaves, and produced a mean frequency estimate of
0.53.. The data from both populations suggest a 9:7 model could fit

the data. however. due to a large amount of cultivar overlap a

weighted 3:1 model was used to explain the segregation patterns ob-
served. A 3:1 model was chosen since the bimodal distribution pattern

156

 

 

 

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158

Table 94. ‘Chi-square test for goodness of fit to the postulated model
for non-wrapper leaf number in the F2 generation
(l975E planting). . .

 

 

 

 

 

Cross Observed Expected Model X2 P

Baby Head x

Badger Ballhead *

F2 149:103 161.78: 90.22 3: 1 2.8199 0.10
*

RFZ . 169: 79 159.22: 88.78 3: l . l.678l 0.l9.

Baby Head x

P.I. 215514

' *

F2 181: 78 180.38: 78.62 13: 3 0.0072 0.93
* .

RFZ 328:123 314.09:l36.91* l3: 3 2.0295 . 0.15

Pooled 509:201 494.46:215.54 13: 3 ‘ 1.4076 . 0.24

Baby Head x

Chieftain Savoy , *
146: 46 150.66: 41.34

 

 

 

F2 * 3: 1 0.6681: 0.41
RF2 116: 81 122.76: 74.24 9: 7 0.9887 0.32
Red Danish x

Badger Ballhead

F2 59 150 69.02:l39.98: 1: 3 2.1702 0.14
RF2 43:111 50.85:103.15* 1: 3 1.8109 0.18
Pooled‘ 102:261 119.87:243.13 1: 3 3.9774 0.05
Red Danish x

P.I. 215514

F2 62:249 73.86:237.14* 7: 9 2.4985 0.11
Red Danish x

Chieftain Savoy .

F2 26: 81 26.32: 80.68 1:.3 0.0053 0.94
RFZ- 17: 85 25.09: 76.91 1: 3 3.4616 0.06

Pooled 43:166 51.42:]57.58 1: 3 1.8269 0.18

 

* ‘ .
= Heighted

Table 95.

159

Chi-square test for goodness of fit to the postulated
model for non-wrapper leaf number in the F
(l975L planting). .

2 generation

 

 

 

 

 

 

 

Cross (Observed Expected Model X2 P ,
Baby Head x
Badger Ballhead _
F2 99: 31 107.50: 22.50: 3: 1 3.8821 0.05
RF2 170: 41 174.48: 36.52* 3: 1 0.6644 0.42
. Pooled 269: 72' 281.98: 59.02 3: 1 3.4511 0.06
Baby Head x
P.I. 215514
F2 123: 45 129.23: 38.77: 3: 1 l.3018 ' 0.25
sz 189: 55 186.54: 57.46 3: 1 0.1376 0.71
Baby Head x .
Chieftain Savoy
F2 126: 89 126.14: 88.86: 9: 7 0.0004 0.98 '
R52 92: 86 104.43 "'57: 9: 7 3.5799 0.06
Pooled 218 175 230.57:162.43 9: 7 1.6576 0.20
Red Danish x
Badger Ballhead
F2 37 254 44.30 246.70 3:13 1.4179 0.23
R52 11: 19 10.66 19.34 7: 9 0.0172 0.90
Red Danish x
, Chieftain Savoy
.52 31:168 38.2o:160.80* : 1.6787 0.20
sz 33 191 43.00:181.00* : 2.8766 0.09

 

* .
= Heighted

160
expected for a 2 gene model was absent in the F2 data (Table 92).
The expected weighted ratio was calculated as follows. The net over-
lap around the 15 leaf claSs was (30.9 + 7.1 + 4.8 + 2.4 - 5.1 - 5.1 -
10.3 - 10.3) -- 14.4. The numberof dominant plants expected was
((100-14.4)/100)(0.75)(252) . 161.78. The expected ratio was 161.78:
90.22. Substituting the number 248 for 252 in the previous equation
produced the expected ratio for the 1975E reciprocal F2 population.
5 The ratio was 159.22:88.78 (Table 94). The results of similar tests
on other F2 populations suggest digenic control of non-wrapper leaf
number. The simplest gene system available to insure digenic control
and account for phenotypic differences between cultivars was a 2 locus
system with 5 alleles per.loci. .The postulated loci J and K produced
cultivar genotypes as noted in Table 16. F2 data from each of the
crosses suggest a was dominant to 13. 14 and 15. The x alleles were
inherited in a similar manner. Red x green cabbage crosses in the

1975E planting suggest J5 and J4 were dominant to J and 32 was domi-

2
nant to 13. The 1975L planting of red x green cabbage crosses sug-

gest 25 was dominant to 32

the dominance relationships of the Klocus were similar to those of .

and .72 was dominant. to 13. In both plantings

the J locus (Tables 16-17). For both the 1975E and 1975L F2 plantings
of Baby Head x Badger Ballhead few non-wrapper leaves were dominant.
-1975E F2 populations of this cross were not pooled. The F2 popu-
lation. using a 15 leaf dividing point. produced a segregation ratio
of 149:103 which fit (P=0.10) a weighted theoretical ratio of 161.78:
90.22 (Table 94). The reciprocal F2 planting. using the same dividing
point. produced an observed ratio of 169:79. A 3:1 weighted expected
ratio of 159.22:88.78 was compared (P=0.19) to the observed ratio

(Table 94). Crossing the reciprocal F1 to the receSsive cultivar.

161

Badger Ballhead. produced a segregatibn ratio of 19:14. This ratio
fit (P=0.41) an expected 1:1 model of 16.5:16.5. The pooled F2 data
for the 1975L planting of this cross showed a segregation ratio of
269:72. The data fit (P=0.06) a 3:1 weighted expected ratio of
282:59 (Table 95). A dividing point of 9 leaves was-used to separate
the phenotypic classes (Table 93). Few non-wrapper leaves were pro-
. duCed when a single dominant gene. J. was present in the genotype.
Many non-wrapper leaves were produced by the receSsive genotype
j3j3k3k3.. Dominant alleles were contributed by Baby Head (Table 16).

-All 3 plantings. 1974L. 1975E.and 19751. of Baby Head x P.I. 215514
F1 suggest incomplete to complete dominance for fewnon-wrapper leaves
(Tables 11-13). The results of a chi-square homogeneity test sug-
gest that both the 1975E F2 and 19755 reciprocal F2 populations of
Baby Head x P.I. 215514 could be pooled (Table 96). Using a dividing
- point 0f 17 leaves the pooled F2 population produced a ratio of 509:201
which fit (P=0.24) a 13:3 weighted expected ratio of 494.46:215.54
(Table 94). Few non-wrapper leaves were produced when the dominant
genotype at 1 locus. J_, or the receSsive genotype at the other locus.
k4k4, were present in the total plant genotype. Many non-wrapper
leaves were produced by the genotype 1414xx. Dominant alleles. J and
K. were. provided by Baby Head while recessive alleles 14 and k4 were
produced by P.I. 215514. F2 populations from the 1975L planting were
different statistically and were not pooled. Using a dividing point
of 14_leaves. the 1975L Fé population produced a segregation ratio of
123:45 while the reciprocal F2 population. using a dividing point of
13 leaves. produced a ratio of 189:55 (Table 97). Both populations fit
(P-O.25 and 0.71) 3:1 weighted expected ratios of 129:38 and 187:57

162

 

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163

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164

respectively (Table 95). Few non-wrapper leaves were controlled by
the presence of 1 dominant gene, J. provided by Baby Head. The reces-
Sive genotype j4j4k4k4 conditioned many non-leaves (Tables 96-97).
Both the 1974L and 1975L plantings of Baby Head x Chieftain
SavoyF1 suggest incomplete dominance for many non-wrapper leaves.
however. the 19758 planting of this F1 population showed incdmplete
dominance for few non-wrapper leaves (Tables 11-13). The data sug-
gest non-wrapper leaf number may be affected by planting time. The
1975E planting of Baby Head x Chieftain Savoy F2 produced a segrega-
tion .ratio of 146:46. The data fit (980.41) a 3:1 weighted expected
ratio. The presence of a single gene. J, from Baby Head was dominant
for few non-wrapper leaves. The recessive genotype jéjsksks from Chief-
tain Savoy produced many non-wrapper leaves. The reciprocal F2 popu-
lation of this cross produced 116 plants with few leaves and 81 plants
with many leaves (Table 94).‘ A dividing point of 16 leaves was used
for both populations (Table 98). The reciprocal F2 data fit (P¢0.32)
a 9:7 weighted theoretical ratio (Table 94). .A single dominant allele
was required at each locus, JԤ_. to produce few non-wrapper leaves.
Homozygous recessive genotypes at each locus produced many non-wrapper
leaves. Baby Head contributed dominant alleles for few leaves while
Chieftain Savoy contributed recessive alleles for many leaves. An
8:7 ratio was produced by crossing the reciprocal F1 of this cross
to the recessive cultivar Chieftain Savoy. The data produced a poor
fit (P=0.01) to the 3.75:11.25 ratio expected. In contrast. the same
cross using the F1 rather than reciprocal F1 produced better agreement
(P=0.08) to a 16:16 expected ratio. 'Backcrossing the reciprocal F1
to the dominant.cultivar Baby Head produced a ratio of 50:5. A ratio

 

165

 

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166
of 55:0 was expected. The data support the 3:1 and 9:7 models postu-

-lated for non-wrapper leaf number inheritance in the cross Baby Head

x Chieftain Savoy. F2 segregation data were pooled for the late plant-
ing (Table 99). Using a dividing point of 8 leaves a pooled segrega-
tion ratio of 218:175 was observed. The data fit (P=0.20) a 9:7
weighted expected ratio of 231:162 (Table 95). Chieftain Savoy contri-
buted recessive genes for many leaves while Baby Head contributed .
genes for few non-wrapper leaves. A homozygous recessive genotype

at either locus conditioned many non-wrapper leaves (Table 99).

, F1 crosses between red and green cabbage show dominance for many
leaves (Tables 11-13). All 3 plantings. 1974L. 19755 and 1975L. of
Red Danish x Badger Ballhead F1 suggest overdominance for many non-
wrapper leaves. A comparison of cultivar distributions suggest Red
Danish produced a greater number of non-wrapper leaves than did Badger
Ballhead (Tables 100-101). Pooled Red Danish x Badger Ballhead F2
segregation data from the early planting also show many non-wrapper
leaves to be dominant (102:261) (Table 94). The pooled data fit
(920.05) a weighted-expected ratio of 120 243 (Table 94). A divide
ing point of 17 leaves was used. Red Danish contributed a single .
dominant gene. J2, for many non-wrapper leaves. The recessive geno-
type. j3j3k3k3, from Badger Ballhead produced few non-wrapper leaves.
Crossing the F1 to the dominant cultivar Red Danish did not produce a
close fit to the expected 0:47 ratio. A ratio 0f 14:33 was observed
(Table 100).. Many non-wrapper leaves were also dominant in the late
planting. The observed 37:254 segregation ratio for the 1975L F2
population of Red Danish x Badger Ballhead fit (P-o.23) a 3:13 weighted
expected ratio of 44:247 (Table 95). Many non-wrapper leaveswere

167

 

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168

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169

 

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170

produced when the dominant genotype at 1 locus, . or the recessive

J2.
genotype at the other locus. k3k3, were present. -The genotype j3j3k2k2
conditioned few non-wrapper leaves. The reciprocal F2 population was
small. .The observed segregation ratio of 11:19 fit (P=0.90) a 7:9
weighted expected ratio (Table 95). The dominant genotype J2;K2-
conditioned many non-wrapper leaves. Few non-wrapper leaves were pro-
duced when homozygous recessive genotypes at either loci were present.
Red Danish contributed dominant genes for many leaves while Badger
Ballhead produced recessive alleles for few leaves (Tables 16. 100).
In both the F2 and reciprocal F2 populations a dividing point bf 11'
leaves was used (Table 101).

Data from the 1974L and 1975L F1 populations of Red Danish x
P.I. 215514 suggest incomplete dominance for few non-wrapper leaves.
Data from the 1975E planting of this population suggest an additive
inheritance for non-wrapper leafnumber (Tables 11-13). The trend
toward dominance for many non-wrapper leaves in red x green cabbage,
F2 populations was further noted in Red Danish x P.I. 215514 F2
Ipopulations. Phenotypic classes for the 1975L F2 population were
produced using a 17 leaf dividing point. An observed ratio of 62
plants with few leaves to 249 plants with many leaves was produced
(Table 102). The data fit (P=0.11) a 7:9 weighted theoretical ratio
of 74:237 (Table 94). A single dominant allele at each locus. J4_K4_,
produced many non-wrapper leaves.’ P.I. 215514 contributed dominant
alleles for many non-wrapper leaves. Few non-wrapper leaves were pro-
duced when homozygous recessive genotypes at either locus were pre-

sent. Red Danish produced recessive genes for few non-wrapper leaVes

(Tables 95.103).

171

 

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172

 

 

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.epmmpm .H. a x nm-ema veg «mote men to mcpucupn Jammp
ecu Low ucmugma :« «copuanpcum«u Logan: temp cwaamgz-:oc _m new names; .mop wrap»

173

Both the 1974L and 1975E F1 plantings of Red Danish x Chieftain
Savoy suggest overdominance for many non-wrapper leaves. However. the
1975L F1 planting of this cross suggests an overdominance for few non-
wrapper leaves (Tables 11-13). Non-wrapper leaf number may be affeCted .
by environmental conditions. 1975E F2 plantings of Red Danish x Chief-
tain Savoy suggest Red DaniSh contributed recessive alleles for few
non-wrapper leaves (Tables 104-105). The pooled 1975E F2 population
of Red Danish x Chieftain Savoy fit (P=0.18) a 1:3 model with an ex-
pected ratio of 52:157 (Table 94). A dividing point of 17 leaves was
used to separate the F2 data into phenOtypic classes. Many non-
wrapper leaves were controlled by the presence of 1 dominant gene.

J5, contributed by Chieftain Savoy. The recessive genotype, jzjzkzkz,
produced few non-wrapper leaves. Both the F1 and reciprocal F1 were
backerossed to the recessive cultivar Red Danish. The segregation
ratios. 8:10 and 14:23 fit (P=0.67.0.16) an expected 1:1.model. The
data support the 1:3 model for non-wrapper leaf weight inheritance in
the cross Red Danish x Chieftain Savoy (Table 94). Both 1975L F2 popu-
lationsof this cross were compared to 1:15 weighted theoretical ratios
since the parental cultivars overlapped. Powers et a1. (23) test sug-
gested a division point of 13 leaves should be used. The 1975L Fé,
population produced a segregation ratio of 31:168 while the reciprocal
population was composed of 33 plants with few non-wrapper leaves and-
191 plants with many leaves (Table 105). Both populations fit
(P=0.20.0.09) 1:15 weighted expected ratios (Table 95). The recessive
genotype. jzjzkzkz contributed by Red Danish. produced few non-wrapper
leaves.” Many nonewrapper leaves were produced by gentoypes with 1 or

more dominant alleles per loci. Chieftain Savoy. JSJSKSKS. contributed

174

 

mam—u co “_e_4 toga:

conga: emu; cmqqmczscoz

e.u_m_ N.N N.N m.m_ «.mp m.¢~ o.m. ..m ..w N.N hm an x flea x may
e.u.m. o.m _.P_ P.__ ”.NN «.mm _.__ m_ . ea x Ame x «av
m.H_F~ m.~. e.m_ w.m ~.o_ .e.¢P m.o_ w.m ¢.m o.F o.~ o._ mop. .Nd Ace x may.
m.“ owl _~.e m.m N.N, n.5F m.m. m.m_ m.mp ~.m m.~ o.o a.o Nap Nd Ame x any
e.“ ON .o._ m.~ m.mp o.o_ m.-. w.m~ w.m m.m w.m om Ace x mac
m.“ a_ o.m o.m ~.m_ m.om e.mm P.m. o.m mm Ame x eav
N.“ up o.~ m.m. m.m m.m ~.¢_ e.up m.m m._~ o.m m.~ N.e_ em Away xasam ccmucm_;u
M.“ e. m.~ ”.N. N.m e.m “.mm ~.m~ ,m.ep ¢.P_ . mm heav gmwemo emu
cam: mm on mm mm _N .G_ .N. m_ m_ PP m mucmpa
to .02

 

 

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on» com ucmugua cw meowpsnwcumwu consac mam, cmanmu3-:o= mmocoxomn new Nu .pu .ucmcma .eop wpnmh

175

 

m.NN. m.- N.pp o.mp

Nu Ac; x mav

 

o + m, m.NP ~.o_ e.m _.m o.m m.o m._ emw
on». 92 9: fig :2 9: c? 99 9m 9« to em m2 fififlxet
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~.H.m P.“ e..~ m.mm e.a~ P.“ am Ame x any
m.“ mp m.e o.~ m.m m.N~ m.a m.m m.o. e.¢ m.~ m.e o.e me Amav aosmm cwaucaagu
N.“ __ N.o m.Np ..mm e.a m.a_ «m Acav smegma wag
cam: mm mm , _N m. N_ m_ mp _P m N m mucmpa
to .oz

mumpu $0 umsmg Lwan:

.Lmnsaz mum; connect-co:

 

 

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ms» to» ucmugmg cw mcowusnwcummu Longs: mam. cmaamczucoc Nd ucm Fm .ucmcmm .mo— open»

176

dominant alleles for many non-wrapper leaves. A comparison between
F2 data from green cabbage crosses and red x green cabbage crosses
shows a striking difference in dominance direction for non-wrapper
leaf number. Further experiments are needed to determine whether this
difference was due to the use of red cabbage in these crosses or wheth-
er an interaction of the green and red germplasm produced this
difference. .

Heritability estimates for non-wrapper leaf number have been
shown by Chiang (3) and Swarup (30) to be 0.35 and 0.52 respectively.
Heritability estimates for green cabbage crosses included a negative

estimate and 4 estimates greater than 1. The only estimate within

2
x

(estimate was 0.90 1 0.48. It appears that 67% of the variation in the

the established 0 to 1 heritability range was 0.67 t 0.88.‘ The B

Fl-population may be additive. Heritability estimates from red x
green cabbage F1 populations included 3 negative estimatesand 1
estimate greater_than 1. Two estimates fell within the established
th0 1 heritability range. They were 0.42 and 0.37. The mean heri-
tability estimate was 0.50 1 0.33 with an 5: estimate of 0.51': 0.33.
Additive genetic control for non-wrapper leaf number may be greater in
green cabbage Fl's than in red x green cabbage Fl populations
(Table 30).

It appears from the data that there is a better genetic correla-
.tion between non6wrapper leaf number and efficiency index than between
non-wrapper leaf number and non-wrapper leaf size. ‘The F1 mean genetic
correlations between non-wrapper leaf number and efficiency index or
non-wrapper leaf size for crosses involving green cabbage cultivars

were 0.94 i 0.27 and -0.18 i 0.59. while the same E9 values for red x

177
green cabbage Fl's were 0.85 i 0.90 and -O.56 1 0.93. Correlation

estimates between non-wrapper leaf number and efficiency index ranged
from -0.02 to 0.96, while genetic correlations between non-wrapper

leaf number and non-wrapper leaf size ranged from -0.96 to 0.98 for

crosses between green cabbage cultivars. “g

0.10 and -0.18 i 0.59 respectively. Red x green cabbage F1 genetic

x estimates were 0.92 1

correlations between non-wrapper leaf number and efficiency index
ranged from 0.79 to 0.96 and for correlations between non-wrapper
leaf number and non-wrapper leaf size ranged between -0.80 and 0.78.

figx estimates were 1.00 t 0.10 and 0.34 t 0.92 respectively (Table 87).

Efficiency Index

Efficiency index is a ratio of total non-wrapper leaf weight and
stalk size (weight) to head weight. Efficient plants. those that pro-
duce a head weight equal to or greater than the sum of total non-
wrapper leaf weight and stalk size, produce efficiency index readings
that are equal to or less than 1., Analysis of the inbreeding data
showed significant differences exist between cultivars. inbreeding
and plantings. The interactions, cultivar x inbreeding. cultivar x
plantings, inbreeding x plantings and cultivar x inbreeding x plant-
ings were also significant (Table 106). Although Chieftain Savoy did
not differ from P.I. 215514, all of the other cultivars were signi-
ficantly different from each other. Between cultivars, efficiency‘
index estimatesranged from 0.36 units of unsalable plant parts per
unit of_cabbage head to 2.02 units of unsalable plant parts per unit
of cabbage head (Table 107). The difference of 0.5 units due to in-
breeding was significant. It appears that 1 generation of inbreeding
a randomly selected parent plant reduced plant efficiency to 50%

178

Table l06. Analysis.of variance table for efficiency index ‘
(non-wrapper leaf weight +_stalk size (weight)/head

 

 

 

. >weight).
Source A
of ‘ Mean ~
Variance DF Square F*
Replication (R) 2 0.07 0.38
. ‘ Hr
Cultivar (C) 4 7.07. 38.99 '
. a . i *1:
Inbreeding (I) l 5.4l 29.85
**
C x P 4 l.7l 9.43 I
**
Plantings (P) 2 l0.45 57.62 -
c x P ' 8 2.45. 13.50"
. ** .
I X P 2 1.52 8.40
*1}
c x I x P a 0.90 4.99

Residual Error 58 0.18 ‘

 

' 'k
*F test significant at the 5% , or l%f* level.

179

Table l07. Cultivar x inbreeding interaction means for efficiency
index. . '

 

 

 

Inbreeding Cultivar
Cultivar P1 7 51 Means
Baby Head . 0.4l a* 0.32 a 0.36 N
Badger Ballhead 0.68 a 0.99 a' 0.83 x
P.I. 215514 l.l6 a 1.47 a 1.31 v
Red Danish l.26 a 2.80 b 2.03 z
Chieftain Savoy 1.21 a l.60 a [1.40 v
Inbreeding Means 0.94 A

1.43 B-

 

*Mean separation, within rows, by Duncan's Multiple Range Test at

the 5% level.

180
(Table 107). The selection of only 1 plant from an open-pollinated

line may also have contributed to reduced plant efficiency. The data
show environmental effects modified efficiency index. Each planting

- was significantly different. A comparison between the 1974L and l975L
plantings show that efficiency estimates can double due to environmental
effects (Table 108). The significance of the cultivar x inbreeding
interaction was due to a difference in plant efficiency between in-
breeding levels for Red Danish (Table 107). Plant efficiency for Baby
Head was not effected by different plantings, however, Red Danish
showed a significant increase in efficiency from the 1974L planting

to the 1975E planting and a significant decrease in efficiency from

the 1957: to 1975L planting. Badger Ballhead, P.I. 215514 and Chieftain
Savoy all showed a significant increase in plant efficienCy between

the 1974L and 1975E plantings and remained unchanged between the 1975E
and 1975L plantings (Table 108). While plant efficiency decreased

due to 1 generation of inbreeding in both the 1974L and 1975L plantings,
the 1975E planting showed no change in plant efficiency due to inbreed-
ing (Table 109). The second order interaction. Cultivar x inbreeding

x plantings was significant due to a decrease in plant efficiency be-
tween inbreeding levels of the 1975L planting of Red Danish. Other
cultivars showed no change in efficiency due to inbreeding for either
the 1975E or 1975L plantings. Differences in cultivar, inbreeding re-
sponse during the 1974L planting contributed to a significant inter-
action. For the 1974L planting both Baby Head and Badger Ballhead
showed no difference in plant efficiency due to inbreeding while the
other 3 cultivars each showed a significant decrease in plant effi-

ciency after 1 generation of inbreeding (Table 110).

181

Table l08. Cultivar x planting interaction means for efficiency

 

 

 

 

Planting Means

0.73 A

index.

. Plantings - ' Cultivar '
Cultivar l974L l975E 1975L Means
Baby Head 0.25 ab* 0.69 o 0.15 a 0.36 w w

. Badger Ballhead l.25 b 0.66 a 0.59 a 0.83 X
P.I. 215514 2.12 b 0.95 a 0.87 a 1.31 v
Red Danish 2.87 b 0.77 a 2.45 b 2.03 2
Chieftain Savoy 2.78 b 0.59a 0.34 a 1.40 v

l.85 C 0.98 B

 

*Mean separation, within rows, by Duncan's Multiple Range Test at

the 5% level.

182

Table l09. Inbreeding x plantings interaction means for
efficiency index.

 

 

 

 

Plantings Inbreeding
Inbreeding ' l974L 1975E l975L Means
P1 - 'l.46 a* 0.74 a 0.62 a ' 0.94 Y
51 . ‘2.24 o 1 0.72 a 1.34 b 1.43 z
Planting Means l.85 C 0.73 A 0.98 B

 

*Mean separation, within columns, by Duncan's Multiple Range Test I
at the 5% level.

Table 110.

183

Cultivar x inbreeding x planting interaction means
for efficiency index.

 

 

 

 

 

 

_ Plantings .

Cultivar Inbreeding l974L l975E ' l975L.
Baby Head P1 0.32 a* . 0.73 a 0.17 a

a 0wa 0Ma qua
Badger P1 1.11 a 0.60 a 0.34 a
Ballhead

S1 l.40 a ' 0.73 a 0.83 a
9.1. 215514 P1“ 1.68 a 0.96 a 0.84 a

S1 2.56 b 0.93 a 0.90 a
Red Danish' P] 2.00 a. 0.86 a 0.90 a'

51 3.73 b 0.67 a' 4.00 o
Chieftain P1 2.21 a 0.58 a 0.84 a
Savoy . . .

S 3.34 b 0.60 a 0.84 a

 

*Mean separation, within columns and cu ltivars, by Duncan' 5 Multiple

Range Test at the 5% level.

184

No study has previously reported on the inheritance of a cabbage
efficiency index. Flmeans of green cabbage crosses Suggest incomplete
dominance to overdominance for increased plant efficiency. on the
Other hand all red x green cabbage crosses. with the exception of
the 1975L F1 mean of Red Danish x Chieftain Savoy, Show reduced plant
efficiency to be dominant. The 1974 F1 planting of Baby Head x
Badger Ballhead suggests that increased plant efficiency was completely
dominant. however, both the 1975E and 1975L F1 plantings of this cross
suggest complete dominance to overdominance for reduced plant efficiency
(Tables 11-13). Tables 111 and 112 show frequency distributions in per—
Cent for populations involved in the cross Baby Head x Badger Ballhead.
Powers et a1. (23) partitioning tests were applied to these distributions
to determine the number of genes controlling plant efficiency and to de-
fine the boundary between efficient and inefficient classes. Powers et
a1. (23) formula (F2/P1) was applied sequentially from the recessive ends
of both the F2 and P1 diStributions. Using the 1975E planting of Baby'
Head x Badger Ballhead as an example recessive frequencies of 3.6/9.6 or
0.38 for the 3.50+.class. 5.6/12.0 or 0.47 for the 1.09 class. 6.8/21.5
or 0.32 for the 0.99 class. 10.0/33.4 Or 0.30 for the 0.89 class. 13.6/
. 45.3 or 0.30 for the 0.79 class, 20.3/61.9 or 0.33 for the 0.69 class
and 31.4/80.9 or 0.39 for the 0.59 class were produced (Table 111). The
change in frequency between the 0.69 and 0.59 classes suggest a dividing
point of 0.59 could separate the phenotypic classes. Frequency estimates
were close to the 0.25 frequency expected from a 3:1 model. The 1975E
reciprocal population also suggests dominance for increased plant effi-
ciency. Applying the same formula (FZ/Pl) to the recessive end of the
Baby Head distribution. frequencies of 6.8/9.6 or 0.71 for the 3.50+ class.

185

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cop x xoncH aucmpu_$$m

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Panaam 9e 9” em ta 9m 92 92 t2 5: e: an «a Nataxwv
m.F~.H m.om .e.m o.~ ~._ ~.m e.m ~.a ..PP m.m~ m.mN a.m. e,m Nam Na ANa x _av
~.p~.u.a.ae e.m a.m _._ p.op. _.m_ a.~p ~.m~ m.ep “.0 ... mm , AFa x Nay
o.e~.H m.m~ .m.op o.a e.m w.mP o.N. e.a_ N.m_ o.~p N.N mm ANa x pav
“.mm.n e.mm m.~ e.~ N.N Fuop P.m~ m.o~. m.o~ N.N am away eaoeppam toaoam
aaflufifi ea a& ma eé_a;_ea_oa_mé.a& a8 afi tseazzs

eaoz +omm me, me am a“ me ‘am . me an mN m_ “wewwu

 

 

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mmmmp as» com acmucma cw mcopusnpgumwu xmvcp aucowupemo_Mmosoxuun can u

ea mcpucm—a
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186

 

 

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.oop x xmucfi xucowomewm .

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e.op .H a.~_ m.a~ ~.F~ mm . A_av eao=_»aem
eaoz aomm mo. mm mm a“ ma mm me an aw a, wwewwu

 

 

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e meoaosaaeoaae xoeer aoeoeo_eeo we ago .1 .oeoeaa .N.. opaae

187

8.8/12.0 or 0.73 for the 1.09 class. 11.6/21.5 or 0.54 for the 0.99
class. 16.0/33.4 or 0.48 for the 0.89 class. 21.6/45.3 or 0.48 for the
I 0.79 class, 34.1/61.9 or 0.55 for the 0.69 class and 48.0/80.9 or
0.59 for the 0.59 class were produced (Table 112). The frequencies
suggest a dividing point between the 0.69 and 0.59 Classes. The ob-
served mean frequency of 0.58 was higher than the 0.43 expected from
a 9:7 model. however. the data fit (P=0.10) the model (Table 114). In
both the F2 and reciprocal F2 populations weighted theoretical ratios
were Used to explain the data since the distributions of the parental
cultivars overlapped. The weighted ratio was obtained by the following
(methOd: first the net overlap around the dividing point was sub-
tracted from 100.and divided by 100. This number was multiplied by
the total number of plants in the sample and the expected frequency
of the deficient class. For example the 1975E Baby Head x Badger Ball-
head F2 3:1 weighted expected ratio was calculated in the following
manner (Table 113). The net overlap (38.1-28.2) = 9.9 subtracted from
100 and divided by 100 = (0.901). was multiplied by 252 and by the
. frequency 0f the deficient class. 0.25, to provide the numberof reces-
sive plants (56.76). The expected ratio was found by subtracting this
number from 252 to produce a ratio of 195.24:56.76 (Table 113). The
weighted expected ratio of the 1975E reciprocal F2 population was cal-
culated in a similar manner. For example ((100-9.9)/100)(0.4375)(248)
=97.76. The expected ratio was 150.24:97.76 (Table 114). Powers et a1.
(23) tests on other F2 populations suggest a maximum of 2 genes con-
trol this trait. Genes fOr components of this trait may be linked .
since such a complex trait Should be controlled by a large number of
genes. In order to explain the differences between cultivars and pro-

vide a simplified gene syStem a 2 locus 5 allele per locus model was

188

Table 113. Chi-square test for goodness of fit to the postulated model
for efficiency index in the F2 generation (l975E planting).

 

 

 

 

 

 

 

 

Cross Observed Expected Model X2 - P
Baby Head x

Badger Ballhead

F2 1 201: 51 195.24: 55.75: 3: 1 0.7552 0.41
RFZ 163: 85 150.24: 97.76 9: 7 2.7485 0.10
Baby Head x

P.I. 215514

F2 245: 14 242.81: 15.19 15: 1 0.3153 0.57
RFZ 424: 27 422.81: 28.19 15: 1 '0.0533 0.82
Pooled 669: 41 655:62: 44.38 15: 1 0.2738 0.50
Baby Head x

Chieftain Savoy

F2 181: 11 180.00: 12.00 15: 1 0.0889 0.75
Rsz 187: 10 184.59: 12.31 15: 1 0.4533' 0.30
Red Danish x

.Badger Ballhead

F2 35:174 43.4a:155.52: 1: 3 2.0903 .0.15
RFZ 27:127 32.04:121.96* 1: 3 1.0015 0.32
Pooled 52:301 75.52:287.47 1: 3 3.0585 0.08.
Red Danish x

P.I. 215514 _ ~

F2 18:293 19.44:291.56 1:15 . 0.1134 . 0.74
Red Danish x

Chieftain Savoy '

F2 8: 99 5.54:l0l.46: 1:15 l.l508 , 0.2a
RFZ 7: 97 5.28: 95.72 1:15 . 0.5892 0.44

Pooled 15:194 _10.Bz:19B.1a* 1:15 1.5999 0.19

 

*
= weighted

189

Table ll4. Chi-square test for goodness of fit to the postulated

model for efficiency index in the F

(l975L planting).

2 generation

 

 

 

 

 

 

 

Cross Observed Expected Model X2 P
Baby Head x
Badger Ballhead

*
F2 83: 47 91.87: 38.l3* 9: 7 2.9209 0.09
RFZ 142: 59 149.11: 51.88; 9: 7 1.1574 0.28
Pooled 225:115 240.95:100.01 9: 7 _ 3.5158 -0.05
Baby Head x
P.I. 215514
F2 130: 38 138.92: 29.08* 3: 1 3.3114 0.07
RFZ 183: 51 185.39: 58.51* 3: 1 0.1283 0.72
Baby Head X
Chieftain Savoy
F2 171: 44 157.97: 47.03* 3: 1 0.2495 0.52
RFZ 128: 50 , 139.05: 38.94* 3: 1 4.0249 0.04
Pooled 299: 94 307.04: 85.95* 3: 1 0.9517 0.33
Red Danish x
Badger Ballhead
F2 15 275 11.70:279.30* 1:15 0.9573 0.32
Red Danish X
Chieftain Savoy
F2 15 184 l2.44:l86.56 1:15 0.5532 0.45
are. 12:212 14.00:210.00 1:15 0.3523 0.55
Pooled 27:395 25.44:395.55 1:15 0.91

0.0128

 

* = Weighted

190 ,
postulated. The loci were designated L and M.’ Tables 16 and 17
show the cultivar and F1 genotypes under this system. A comparison of
cultivar distribution patterns and postulated models for segregation
data suggest that in both 15:1 and 1:15 models transgressive segre-
gation occurred. These observations suggest that parental homozygous
loci were in repulsion for crosses producing these ratios. Green cab-

bage crosses in the early planting suggest_L was dominant to 13, 14

and 15 and m was recessive to ME, M4 and M5. Red x green cabbage

crosses in the early planting suggest L2 was dominant to 13, i4
5 and m3 were recessive to ”2, and M: was recessive to M2.

1975L F2 distributions suggest L was dominant to 13, 1

and.15 while m
4 and 15. Simi-
lar inheritance patterns were noted for the M locus. Red x green
cabbage crosses grown in the late planting suggestzé _was dominant to
13 and 15 while m3 was recessive to M: and M? was recessive to M5
(Tables 16-17). 1975E Baby Head x Badger Ballhead F2 data, using the
0.59 class as a dividing point. produced a segregation ratio of 201:51.
The data fit (P=0.41) a 3:1 weighted expected ratio of 195.24:56.76
(Table 113).. Efficient plants were dominant and controlled by the pre- '

sence of a Single gene L The recessive genotype 11mm produced in-

3.
efficient plants. Baby Head was the sourCe of recessive alleles in this
cross. Reciprocal F2 data for the same planting, showed a segregation
ratio of 163 efficient plants to 22 inefficient plants. Again a divid-
ing point of 0.59 was used to separate the phenotypic classes. This
data fit (P=0.10) a 9:7 weighted expected ratio of 150.24:97.76 (Table
113). Inefficient plants were produced when 1 or more homozygous ref

cessive loci occurred in the plant genotype. The genotype L3‘M3_ con-

ditioned efficient plants. Dominant alleles for efficiency were

 

 

. 191
provided by Badger Ballhead (Table 111). The reciprocal F1 was crossed
to the dominant cultivar Badger Ballhead. This cross produced a segre-
gation ratio of 32:1. A ratio of 33:0 was expected. The data support
the 9:7 model with Badger Ballhead contributing dominant alleles for
increased cabbage plant efficiency. F2 data from the 1975L planting,
of Baby Head x Badger Ballhead was pooled to produce a segregation ratio .
of 225:116. A division point of 0.29 was used to separate the data
into classes. The data fit (P=0.06) a 9:7 weighted expected ratio of

 

241:100 (Table 114). Homozygous recessive genotypes at either locus
produced inefficient plants. Recessive alleles 13 and m3 were contri-
buted by Badger Ballhead.:

Both the 1974L and 1975L plantings of Baby Head x P.I. 215514 F1
suggest incomplete dominance for efficiency while the.1975E planting
suggests overdominance for efficiency (Tables 11-13). Both the 1975E
F2 and reciprocal F2 populations of Baby Head x P.I. 215514 were pooled.
The pooled data suggest efficiency was dominant (Table 115). Of the
710F2 plants sampled 669 were classed as efficient and 41 were
classed as less efficient. A dividing point of 0.89 was used. The
data showed good fit (P=0.60) to the 15:1 model (Table 113). Efficient
plants were produced when either L or M; were present in the genotype.~
Inefficient plants were produced by the genotype 14lgmm. Baby
Head contributed L and m alleles while P.I. 215514 contributed14and
u) alleles. Each cultivar contributed 1 dominant and 1 recessive-
allele to the F1 plant. erpopulations in the 19751 planting were not.
pooled. Fhe F2 population. using a diViding point of 0.69. produced.
a segregation ratio of 130:38 (Table 114). The data fit (P=0.07) a3:1
weighted expected ratio of 139:29 (Table 114).. The 1975Lreciprocal
F2 population of this_cross. with an observed ratio of 183:61. also fit

192

 

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193
(P=0.72) a 3:1 weighted theoretical ratio (Table 114). The presence
of a single dominant gene, L. from Baby Head conditioned efficient
plant production. The recessive genotype,1;14mam4, produced ineffi-
cient plants (Table 116).

F1 populations of Baby Head x Chieftain Savoy produced the same
results ashF1 populations of Baby Head x P.I. 215514. 1974L and l975L
F1 plantings showed incomplete dominance for efficiency while the 1975E
planting showed overdominance (Tables 11-13). Baby Head x Chieftain
Savoy F2 data from the early planting were not pooled. Using a divid-
ing point of 0.69 the F2 population produced a segregation ratio of
181:11 and fit (P=0.76) a 15:1 model (Table 113). ~The recessive geno-
type 1515mm conditioned poor efficiency (Table 117). The reciprocal
F2 population produced a ratio of 187:10 which fit (P=0.30) a 15:1 model
(Table 113). A dividing point of 0.89 was used to separate phenotypic
classes. Dominant efficient plants were produced when a dominant'
allele at either loci, L or M5, was present in the plant genotype. The
recessive genotype 1515mm conditioned inefficient plants. Chieftain
SaVoy contributed 1 dominant allele L5“and'1'receSSive‘allele ms;to the
gene pool while Baby Head produced both a dominant allele M and a re-
cessive allele 1. Crossing both the F1 and reciprocal F1 plants to
Chieftain-Savoy produced ratios of 26:6 and 15:0.‘ Ratios of 32:0 and
15:0 were expected. The reciprocal F1 was also backcrossed to Baby
Head. A ratio of 55:0 was produced. A 55:0 ratio was expected. The
backcross data support the 15:1 F2 model for Baby Head x Chieftain
Savoy efficiency index inheritance. Fz data from the late planting.
were pooled.) The segregation ratio of 299:94 suggests that efficiency
was dominant (Table 118). A dividing point of 0.39 was used to separate

194

 

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196

 

 

 

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197
the F2 classes. bThe data fit (P=0.33) a 3:1 weighted expected ratio
of 307:86 (Table 114). Efficient plants were produced by the presence
of a dominant L allele from Baby Head. Inefficient plants were pro-
duced by the 1515m5m5 genotype of Chieftain_Savoy. .

All 3 plantings, 1974L. 19755 and 1975L, of Red Danish x Badger
BallheadF1 suggest poor efficiency was dOminant to overdominant when
compared to efficiency (Tables 11-13). F2 data from the 19755 plant-
ing of Red Danish x Badger Ballhead were pooled as-suggested by
chi-sguare homogeneity tests.- A population of 363 plants was sampled
for efficiency index (Table 119). Using a dividing point of.0.59..
data from the pooled F2 population produced a ratio of 62:301. This
ratio fit (P=0.08) a 1:3 weighted theoretical ratio of 76:287 (Table
113). Inefficient plants were produced when a single dominant allele.
L . was present in the plant genotype. Efficient plants were con-

2
ditioned by the recessive genotype 1 I'm m Badger Ballhead contri-

3 3 3 3’
buted genes for efficiency while Red Danish contributed genes for in- '
efficiency. A ratio of 7:5 was obtained fromthe backcross between-l
the F1 and the recessive cultivar Badger Ballhead. The data fit (Pi0.59)
an expected 6:6 ratio. Data from a cross between_the reciprocal F1 and
Red Danish suggest a segregation ratio of 2:45. A ratio of 0:47 was
expected. Data from the 1975L Fé planting of this cross produced a
ratio of 15:276 (Table 120). A dividing point of 0.39 was used to
separate the F2 data into classes. The observed ratio fit (P-D.32) a
1:15 weighted expected ratio (Table 114). Inefficient plants were con-
trolled by 2 dominant genes. The presence of a dominant allele, Lz'or
M2 , at either loCus conditioned an inefficient plant. The homozygous
recessive genotype 13138383 from Badger Ballhead produced an efficient
plant phenotype. '

 

198

 

 

 

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199

 

 

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200

F1 populations of Red Danish x P.I. 215514 grown in the 1974L.
r9755 and 1975L plantings suggest poor cabbage plant efficiency was
overdominant (Tables 11-13). Due to poor seed production, F2 popula-
‘tions of Red Danish x P.I. 215514 were grown only in the early plant-
ing (Table 122). Of the 311 plants sampled. 18 were classified as
efficient While 293 were lessefficient (Table 121). The data pro:
duced a good fit (P=0.74) to a 1:15 model (Table 113). .A dividing
point of 0.49 was used to separate the F2 data. Inefficient plants
were produced when either of the dominant L2 or M2 alleles were pre-
sent in the plant genotype. Efficient plants were produced by the
1414m2m2 genotype. Red Danish produced a dominant L2 allele while
P.I. 215514 produced a dominant M4 allele. .

‘ 1975L and 19755 F1 plantings of Red Danish x Chieftain Savoy
suggest overdominance and complete dominance for reduced plant effi-
ciency. The 1975L F1 planting suggests incomplete dominance for in-
creased plant efficiency (Tables 11-13). Environmental conditions
affect changes in plant efficiency. Red Danish x Chieftain Savoy
F2 data were pooled within each planting based on the results of
chi-square homogeneity tests. Pooled F2 data from each planting were
compared to 1:15 weighted theoretical ratios. F2 data from the early
planting produced a ratio of 15:194 (Table 123).' Thisratio fit (P=0.10)-
an expected 11:198 ratio (Table 113). A division point of 0.59 was
used to divide F2 data into classes. Inefficient plants were produced
by genotypes with either a dominant L2 or M2 allele. The recessive
genotype 1515m5m5 from Chieftain Savoy conditioned an efficient plant
phenotype.' F1 plants backcrossed to Red Danish produced a ratio of
'1:17. A 0:18 ratio was'expected.‘ However. the 20:17 ratio produced

 

201

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204 y
from the reciprOcal F1 x Red Danish backcross did not agree with the .

0:37 expected ratio. Differences in F1 and RF1 gametic transmission
are suggested. Data from the 1975L planting of Red Danish x Chieftain
Savoy F2 show an observed segregation ratio of 27:396 (Table 124). The
data show good fit (P=0.91) to a 1:15 madel (Table 114). A division
point of 0.39 was used. Since Red Danish and Chieftain Savoy were both
inefficient in the 1975L planting (Table 124) the presence of efficient
plants suggest transgressive segregation. Transgressive segregation
could occur when 2 recessive loci recombine to produce a homozygous
recessive phenotype. The recessive genotype 1515m5m5-c0nditioned an
efficient phenotype. Dominant inefficient plants were produced when -
either of the dominant L2 or MS alleles were present in the plant
' phenotype. Red Danish contributed L2 and m2 alleles while Chieftain
Savoy contributed 15 and MS alleles. I

F1 green cabbage cross heritability estimates for efficiency in-
dex range from 0.18 to 0.96. The mean heritability estimate was 0.75

z 0.62. Two heritability eStimates were negative. 2 were greater than

2
x

estimate was 0.86 1 0.45. Red x green cabbage F1 heritability estimates

- l and 2 estimates were within the 0 to 1 heritability range. The B

for efficiency index ranged from 0.15 to 0.56. The mean heritability
was 0.35 i 0.34 with an 5: estimate of 0.58 1 0.27. Again 2 estimates
were negative, 2 were greater than 1 and 2 were within the D to 1 heri-

tability range (Table 30).

Eon-Wrapperjgaf Size
Non-wrapper leaf size is the ratio between total non-wrapper leaf
weight and non-wrapper leaf number. Small numbers suggest leaf size

(weight) was small while larger numbers suggest larger leaf size.

205

 

 

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206

Analysis of inbreeding data for non-wrapper leaf size showed signifi-
cant differences exist between cultivar and planting means. No dif-
ference due to inbreeding was noted (Table 125). However, this resultj
may also occur when dominance is absent. Non-wrapper leaf size ranged
from 0.02 kg for Baby Head to 0.11 kg for P.I. 215514. No difference
was observed between Red Danish and Chieftain Savoy, however, these
cultiVars differed from each of the other Cultivars (Table 106). Each
planting was significantly different for non-wrapper leaf size (Table
126). The significance of the cultivar x plantings interaCtion was
due to different cultivar responses to the 3 plantings. Baby Head '
showed no difference in non-wrapper leaf size for the 3 plantings,
however, Badger Ballhead, Red Danish and Chieftain Savoy showed signi-
ficant reductions in non-wrapper leaf size for the 3 plantings. P.I.
215514 showed a significant reduction in non-wrapper leaf size between
the 1974L and 1975E plantings and then showed no change between the
1975E and 1975L plantings (Table 126). The second order interaction,
cultivar x inbreeding x plantings was significant due to cultivar x
plantings responses and also due to the effect of P.I. 215514 and its
inbred grown in the 1974 planting. P.I. 215514 showed a significant
increase in non-wrapper leaf size after 1 generation of inbreeding
in the 1974L planting. Cultivars grown in the 1975E and 1975L plant-
ings produced no difference in non-wrapper leaf size after an added
generation of inbreeding (Table 127).

Baby Head x Badger Ballhead F1 plantings in 1974L and 1975L sug-
gest non-wrapper leaf size was inherited in an additive manner. The
1975EF1 planting of this cross suggested incomplete dominance for

small non-wrapper_leaf size (Tables 11-13). Tables 128 and 129 show

207

Table 125. Analysis of variance table for non-wrapper leaf size
. in kilograms.

 

 

 

' Source .

of - Mean i
Variance _ DF Square F*
Replication (R) 2 0.0003 1.48 ‘
Cultivar (C) 4 ' 0.0193 , 89.94**
Inbreeding (I) . 1 0.00001 . 0.03
c x I 4 0.0005 ' 2.43*
Plantings (P) 2 0.0l78 _ ' 83.0l**4
'c x P 3 0.0021 ' 9.7a**
I x P 2 0.0007 3.12*

**

c X 1 x P 3 0.0006 2.75
Residual Error 58 ‘ 0.0002

 

' 'k **
*F test significant at the 5% . or 1%. level.

208

Table.l26. Cultivar x planting interaction means in kilograms for
non-wrapper leaf size.

 

 

 

. Plantings Cultivar
Cultivar 1974L l975E l975L Means.
Baby Head 0.02 a* 0.03 a 0.02 a 0.02 H
Badger Ballhead 0.09 c 0.07 b 0.04 a 0.07 X
P.I. 215514 0.13 b 0.10 a 0.11 a _ 0.112
Red Danish 0.13 c 0.08 b 0.05 a 0.09 v
Chieftain Savoy 0.12 c 0.08 b 0.04 a 0.08 Y
Planting Means- 0.10 C 0.07 B 0.05 A

 

*Mean separation, within rows, by Duncan's Multiple Range.Test at

the 5% level.

 

209

Table 127.. Cultivar x inbreeding x planting interaction means in
. kilograms for nonewrapper leaf size.

 

 

Plantings

 

 

 

 

 

Cultivar Inbreeding l974L 19755 l975L
Baby Head P' 0.03 a* 0.03 a 0.02 a
0.02 a 0.03 a 0.01 a
Badger : 0.l0 a 0.08 a. 0.04 a
Ballhead ‘
. 0.03 a 0.07-a 0.03 a
0L2MM4~' 0J0a oma OJZa
owb oma ,owa
Red Danish 0.12 a 0.03 a 0.05 a
-0Ma 0%a owe
Chieftain 0.13 a 0.07 a 0.04 a
.Savoy
0.l2 a 0.08 a 0.04 a

 

*Mean separation. within columns and cultivars. by Duncan's'Multiple

Range Test at the 5% level.

 

210

 

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211

 

 

 

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212
frequency distributions in percent for populations involved in this
study.. Powers et a1. (23) partitioning tests were applied to these
'distributions to determine the number of genes controlling non-wrapper
leaf size and to define the boundary between the small and large non-
wrapper leaf weight phenotypes. 'Powersiet a1. formula (F2/P2) was
applied sequentially from the recessive ends of both the f2 and P2 dis-
tributions. Using the 1975E F2 planting of Baby Head x Badger Ballhead
(Table 128) as an example recessive frequencies of 5.6/20.5 or 0.27 for

the 169 gm class, 6.4/30.7 or 0.21 for the 109 gm class, 7.6/35.8 or

 

0.21 for the 99 gm class, 11.6/40.9 or 0.28 for the 89 gm class. 15 4/
53.7 or 0.30 for the 79 gm class and 30.2/69.3 or 0.44 for the 69 gm
class were produced. The large difference in frequency between the

79 gm class and 69 gm class suggests a dividing point of 69 gm. The

mean frequency of 0.25 agrees closely with a 0.25 frequency expected

from a 3:1 model. A monogenic model with a dividing point between the

69 gm and 79 gm classes was postulated. The reciprocal F2 population

of this cross produced similar results. Powers et a1. (23) formula
(Fz/Pz’ was again applied to produce the following redessive fre-
quencies, 2.0/20.5 or 0.10 for the 169 gm class. 2.8/30.7 or 0.09 for

the 109 gm class, 3.6/35.8 or 0.10 for the 99 gm class. 6.4/49.9 or 0.16
for the 89 gm class, 10.00/53.7 or 0.19 for the 79 gm class and 16.1/ ' i
69.3 or 0.23 for the 69 gm class. The increase in frequency noted be-
tween the 69 gm and 79 gm classes suggest a dividing point of 69 gm..

The mean frequency of 0.128 when doubled (23) shows close agreement to
the 0.25 frequency expected from a 3:1 model. The 1975E reciprocal F2
data of this cross were screened using 69 gm as a separation. Since '

the results of Powers et a1, (23) tests suggest some F2 populations

213

used a digenic system to control non-wrapper leaf size a 2 gene system
was postulated. These loci were designated N and 0. Each of the 5
cultivars had a different set of alleles at these loci to explain the
observed differences between cultivars. The 1975E gene system suggests
that N5 and N4 were dominant to N and N was dominant to n3 for green
cabbage crosses. The 0 locus was inherited in a similar fashion.

1975L data suggests that N was dominant to n3, nd and us while 05 was

-dominant to o and 0 was dominant to o and 04. The early planting of

3
red x green cabbage crosses Suggest that N2 was dominant to n3, H4 and

n5 whileoz

that N3 was dominant to N

was recessive to 03. 04 and 05. The late planting suggests

and N2 was dominant to n Inheritance pat-

2 5'
terns at the Other-locus suggest that 05 and 03 were dominant to 02.
Dominance for large non-wrapper leaf size was noted for all populations
grown in the early planting while dominance for small non-wrapper leaf
size was noted in the late planting. Parental distributions of Baby .
Head and Badger Ballhead overlap in the 1975E planting. A weighting
factor was applied to the expected F2 ratio to account for the mis-
classification of phenotypes. The 1975E.F2 planting of Baby Head x
Badger Ballhead will again be used as an example (Table 128). Using

a division point of 69 gm the data suggest Badger Ballhead produces
only 53.7% of its plants with a large non-wrapper leaf size pheno-
type. Because of this overlap of 46.3% the small non-wrapper leaf size
F2 class will be larger than normally expected and the large

leaf size class will be smaller than expected. In this study the net

overlap, 46.3. was subtracted from 100, then divided by 100 and multi-
plied by both the expected probability of the deficient class and the

total number of F2 plants sampled to obtain the expected segregation

. RF

} 214

ratio. In this case the expected number of recessive large leaf size
phenotypes was ((100-46.3)/100)(252)(0.25) = 33.83.. The number of
dominant small leaf size phenotypes was found by subtraction (252-33.83)
= 218.17. The observed F2 segregation ratio of 211 42 for the 1975E
planting of Baby Head x Badger Ballhead fits (P=0.19) a 3:1 weighted
theoretical ratio of 218.17:33.83 (Table 130). The reciprocal F2 popu-
lation produced a ratio of 223:25 which fit (P=0.13) a 3:1 weighted
expected ratio of 215:33 (Table 130). The presence of a single domi-
nant allele. N, from Baby Head produced small non-wrapper leaf size.
The cultivar Badger Ballhead provided the recessive genotype n3n3o3o3
for large nOn-wrapper leaf size. The backcross ratio of 16:17 for '
1 x Badger Ballhead fit (P=0.88) an expected 1:1 model. Both the F2
and backcross data support weighted 3:1 F2 ratios for non-wrapper leaf
size inheritance. A chi-square homogeneity test suggested pooling the
1975L F2 data (Table 129). A dividing point of 34 gm was used to
divide the F2 data into phenotypic classes.‘ The pooled F2 data produced
a ratio of 274:67 which fit (P=0.20) a 3:1 weighted expected ratio
(Table 131). Small non-wrapper leaf size was controlled by a single
dominant gene, N, contributed by Baby Head. The recessive genotype
n3n3o3o3 produced a large size non-wrapper leaf. 7

Both the 1974L and 1975E plantings of Baby Head x P.I. 216514
suggest dominance for large non-wrapper leaf size. However, the 1975L
planting of the same cross suggests incomplete dominance for small non- '

wrapper leaf size (Tables 11-13). The F1 data suggests that environ-

mental conditions can greatly affect non-wrapper leaf size phenotypes.

F2 populations of Baby Head x P.I. 215514 grown in the 1975L planting
were pooled. Using a dividing point of 39 gm to separate the small and

 

215

Table 130. Chi-square test-for goodness of fit to the postulated model
for non-wrapper leaf size in the F2 generation
(l975E planting).

 

 

 

 

 

 

 

 

Cross Observed _Expected ‘Model X P.
Baby Head x
'Badger Ballhead . * ,
F2 211: 42 218.17: 33.83*1 3: l 1.7548 0.19
RF2 223: 25 214.71: 33.29 3: l 2.3897 . 0.13
Baby Head x
P.I. 215514 *
F2 59:200 61.67:197.33* 1: 3 0.1513 0.70
RF2 33:418 26.84:424.15 1:15 1.5004 0.22
Baby Head x
'Chieftain Savoy '
F2 68:124 68.57:123.43* 1: 0.0074 0.93
sz 119: 78 123.12; 73.88* . 9:.7 0.3685 0.54
Red Danish x .
Badger Ballhead
F2 . 15:194 13.06:l95.94 1:15 0.3066 0.58
RF2 6:148 '9.62:l44.36 1:15 1.4562 0.23
Pooled 21:342 22.69:340.31 1:15 0.1339 0.71
Red Danish x
P.I. 215514 . .
F2 19:292 19.44:291.56 1:15 0.0107 0.92
Red Danish x
Chieftain Savoy. .
F2 . 9: 98 6.69:100.31 1:15 0.8529 0.36
RF2 9: 93 6.38: 95.62 1:15 ' 1.1529 ‘ 0.28
Pooled 18:191 l3.06:l95.94 1:15 1.3476 0.24

'k
= Weighted

Table 131.

216

Chi-square test for goodness of fit to the postulated
model for non-wrapper leaf size in the F

(1975L planting).

2 generation

 

 

 

 

 

 

Cross Observed Expected Model X2 P

Baby Head x

Badger Ballhead

F2 100: 30 100.65: 29.35* 3: 1 0.0188 0.89
* ,

4 RFZ 174: 37 163.37: 47.63* 3: 1 3.0647 0.08
Pooled 274: 67 264.02: 76.98- 3: 1 1.6737 0.20
Baby Head x
P.I. 215514
52' 154: 14 157.91: 10.09: 15: 1 1.6135 0.20
RF2 233: 11 229.35: 14.65 15: 1 0.9686 0.32

*
Pooled 387: 25 387.26: 24.74 15: 1 0.0029 0.96
Baby Head x
Chieftain Savoy
F2 204: 11 201.56: 13.44 15: 1 0.4717 0.49
RFz 171: 7 166.88: 11.12 15: 1 1.6315 0.20
Pooled 375: 18 » 368.44: 24.56 , 15: 1 1.8702 0.17
Red Danish x
Badger Ballhead
52 244: 47 248.69: 42.31* 15: 1 0.6073 0.44
RF2 27: 3 25.64: 4.36* 15: 1 0.4978 0.48
. :1:
Pooled 271: 50 274.32: 46.68 15: 1 0.2770 0.60
Red Danish x
Chieftain Savoy
F2 ' 159: 40' 161.69: 37.31 13: 3 0.2383 0.62
RFZ 177: 47 182.00: 42.00 13: 3 0.7326 0.39
A Pooled '336: 87 343.69: 79.31 13: 3 lo.9171 0.34

 

*
- Heighted

217

large leaf size classes the 1975E F2 planting showed increased non-
wrapper leaf size to be dominant with a segregation ratio of 59:200
(Table 132). This ratio showed good fit (P=0.70) to a 1:3 weighted
expected ratio of 61.67:197.33 (Table 130). Increased non-wrapper leaf
size was controlled by a single dominant gene, N4, contributed by
P.I. 215514. The recessive genotype. nnoo contributed by Baby Head.
produced small size non-wrapper leaves. The reciprocal F2 population.
using a 39 gm dividing point. produced a ratio of 33 plants with small
leaf size to 418 plants with large non-wrapper leaf size. This ratio
fit (P=0.22) a 1:15 weighted theoretical ratio of 27:424 (Table 130).
Large non-wrapper leaf size was controlled by 2 dominant genes N4 and 04.
The presence of either or both of these genes in the genotype condi-
tioned large leaf size. The genotype, nnoo, of Baby Head produced
small leaf size. Opposite segregation patterns were observed in the
1975L planting of this cross (Table 133). A dividing point of 59 gm
was used to separate the F2 data into phenotypic classes. Pooled'F2
data from the 1975L planting showed a good fit (P=0.96) to a 15:1
weighted expected ratio of 387:25 (Table 131).‘ The genotype of P.I.
215514, n4n4o4o4, was recessive for large non-wrapper leaf size. Small
non-wrapper leaf size was produced when either or both of 2 dominant
alleles, N or 0, were present in the plant genotype. _Baby Head contri-
buted dominant genes for small leaf size in this planting (Tables
132-133). '

The 1975L planting of Baby Head x Chieftain Savoy F1 suggeSts
non-wrapper leaf size-was inherited in an additive manner. However,

the 1974L and 19756 F plantings suggest incomplete to complete domi-

1 .
nance for large non-wrapper leaf size (Tables 11-13). Baby Head x

218

 

N.mN.H ¢.~n N.N n.o— m.op m.P~ ~.op m.¢~ P.n o.o~ m.m m.— «.0 pme Nu A—m x may

 

e.e~.u e.em e.o a.e N.N e.m m.m o.ep m.m m.e~ o.~. 0.8 m.e emu Ne Ame x Pav

~.e~.w ..em N.e m.PN o.o_ P.~_ ”m.o~ e._F m.e e.m e._ e._ oe A_a x may

e.m~.u m.- e.~ o.~ m.~_ m.- m.~P m.Fm m.m ..m m.~ m.m N.. em , Ame x _av

m.e~.u ~.em e.m. N.NN o.PF m.mm N.e. e.m mp Amav epmmPN .H.a

~.e .H m.e~ . V m.e w.m~ m.ee F.“ we AFaV eao: seam
:88: me, mm_ mm mm as , me . am am _ mm mm m_ ”wemwn

mmm—u we “Paw; song:
macaw cw wNwm mam; cmqqmszucoz

 

 

.epmmpw .H.a x.eeo= seam aaeeo ago to neweea_a mm~m_
on» com “smegma :_ mcowu:n_cumwv m~_m 4mm— cmqamczuco: we use p; .pcmcma .Nmp mpamh

219

 

 

e..e.u m.wm e.o 4.0 .e.o m.m e.m a.~m m.oe e.~ m.o eam Ne APa x may

a NN.“ o.mm e.o N._ N.F m.m. m.~ m.m . w.m~ m.me _.m_ we, «a Ame x .av

m.ep.u o.me m._ m.P N.e e.e . m.ap m.ee m.m. m.P pm ape x may

N.__.u 8.5m N.N N.N m.e m.om a.~m F.e me Ame x Fav

N.Ne.u e.eo_ e.mm e.mp e.mp N.N e.m_ A.“ m. Amev e_mme~ .H.a

e.e_.w m.e~ m._ m.~ m.. _.m_ m.m~ N.Ne e.m mm Apav eao: seem
eaoz +mm_ m_F mm mm me am am me am .me a ”wewwn

mmmpu mo peep; swag:
manta cw mNPm mam; coaqmszucoz

 

 

. .epmm_~ .H.a x ewe: seam mmeao oeo co meweeapa 4m~m_.
mg“ so» “smegma cw mcowpsnwcumwu mNWm temp smaqmuzuco: we can pm .pcmsam .mmp apne»

220 .

' Chieftain Savoy'F2 populations grown in the early planting were not
pooled because of differences in segregation patterns (Table 134).

Both the F2 and reciprocal F2 data were phenotypically divided into
classes using a 54 gm dividing point. The 1975E F2 population pro-
duced a segregation ratio of 68:124. This ratio fit (P=0.93) a 1:3
weighted expected ratio of 68.57:123.43 (Table 130). Large non-wrapper
leaf weight was controlled by the presence of 1 dominant gene, N5, do-
nated by Chieftain Savoy. The absence of this gene in thegenotype
conditioned small non-wrapper leaf size. The 19756 reciprocal F2 popu-
lation showed small leaf size to be dominant with an observed segrega-
tion ratio of 119:78. This ratio fit (P=0.54) a 9:7 weighted theoreti-
cal ratio of 123:74 (Table 130). Small size non-wrapper leaves were
produced when both the dominant N and 0 genes were present in the
genotype. Large size non-wrapper leaves were produced by genotypes
that were homozygous recessive at either loci. Backcrossing the F1 x
Chieftain Savoy gave a ratio of 5:27. A ratio of 0:32 was expected.
The reciprocal F1 was crossed to both Chieftain Savoy and Baby Head. .
These crosses produced observed ratios of 3:12 and 32:23 respectively.-
These ratios were compared (P=0.68) to 1:3 and 1:0 models If the re-
ciprocal F2 had produced segregation ratios similar to those observed
for the F2. then backcross data would support a 1:3 model. however. it
appears that reciprocal differences exist between F2 and RF2 populations
(Table 134). 1975L F2 plantings of Baby Head x Chieftain Savoy were
pooled as suggested by a chi-square homogeneity test. A dividing

point of 59 gm was used to divide the F2 data into phenotypic classes.
The data suggest small leaf size was dominant (Table 135). An observed

ratio of 375:18 fit (P=0.17) an expected ratio of 368:25 from a 15:1

221

 

 

P.mm.u ~.mm e.m 0.5 o.m e.m m.m e.m e.m e.e_ m.e_ N.N? m.e_ mm .a x A_a x.mav
e._m.u_m.ee “.0 e.mp “.8 o.o~ ~.mp m.mp ~.e «.mp ,e.e my me x Ape x may
N.NM.H ..mm F.m anew m.NP m.~p ~.o m.m~ e.m m.- _.m mm me x Ame x Fav
o._~.H m.om o.~ o.~ ..e .~._F e.o. “.3 m.- n.5, N.NP m.~ Amp Na A_a x may.
e.m~.fl e.mo o.~ ..m e.e ..N_ e.e_ F.ep ..m m.m~ 5.8 e.m o.. Nap Na Ame x pay
m.m~.u ~.m~ ~., ~.m~ N.mp e.m o.-. N.m_ o.~_ .m.o~ ~._ ~.F mm A_a x may
4.8m.“ N.No m.m e.F_ m.m a.” m.m ..mm 3., N.Nm N.N w.m mm Ame x _av
~.-.H _.o~ m.~ e.m 0.4, m.__ N.o~ m.~ m.m e.o~ m.~ m.m em Away aesam e_eoco_eu
_.e .H m.e~ m.e w.mN m.ee P.~ Ne A_av.eao: seem
eaoz map amp mm mm as mm am am , mm mm a, Hwewwn

mmmpu we uws_4 swan:
manta cw mNPm mam; soaamc31=oz

 

 

.xo>am :Peuwowco x new: xnmm mmosu on» do mcwucmpa umsmp
on» so» ucmosmq cw mcowuzapspmwu wNPm mama smaamczuco: mmocuxomn wen we .Fm .ucwsma .emp mpaoh

222

 

 

w.em

 

mmmpu we peep; swan:
manta cw «New emu; smaqmsz::oz

N.ee.u m.~m _.F e.o e.ow e.o F.P a.“ m.mF F.mm N.N wee Na A_a x mav
.e.m_.fi m.mm m.o m.~ m.~ e.e m.mm N.Nm a.e, m.~ . mpw Na Ame x _av
m.o_.H m.mm m.e m.om m.om ~.op ~.F mm ape x may
m.oF.H arem o.m o.mm o.oe ON Ame x Fay
N.N_.H o.me m.~ m.~ m.~ ~.mp N.Fm e.mp e.e me “may sesam e_aocomeu
e.ep.u m.e~ 8.. 8., m._ _.m_ m.m~ N.Ne e.m mm Apav wee: seem

ewe: .mme m__ mm mm me am am me am me a moea_a

. to .62

 

 

.»o>mm :wwpmmwsu x vow: xamm.mmocu mg» mo mcwucopa Amump
as» so» ucmocma cw mcompznwsumwu m~_m temp swaamszuco: Nu wan Pm apnoea; .mmp m—nmh

223
model (Table 131). Small non-wrapper leaf size was produced when.1 or

both of 2 dominant genes, N or o . were present in the plant genotype.

5
The genotype nsnsoo produced large non-wrapper leaf size. This system
suggests that each cultivar contributed 1 dominant allele to the F1
plant. Each cultivar produced small nonrwrapper leaves.

Both the 1974L and 1975L plantings of Red Danish x Badger Ballhead
F1 suggest non-wrapper leaf size was additively inherited. The 1975E
F1 planting of this cross showed overdominance for large leaf size
(Tables 11-13). F2 populations of Red Danish x Badger Ballhead were
pooled within each planting. 'Data from the 1975E planting was separated
into phenotypic classes using a 44 gm division point. The data showed
small leaf size to be recessive (Table 136) while data from the 1975L
planting, using a 59 gm dividing point showed-large leaf size to be re-
cessive (Table 137). A pooled ratio of 21:342 was observed for the early
planting while a pooled ratio of 271:50 was observed for the late plant-
ing. The 19755 data showed good fit (2:0.71) to a 1:15 model (Table 130)
. while the 1975L data fit (P=0.60) a 15:1 weighted expected ratio (Table
131). Both backcross populations showed perfect agreement to their ex-
pected ratios. :The'backcross data support a 1:15 1975E F2 model with
both cultivars contributing 1 dominant allele to the F1 genotype. Since
both cultivars produced large leaf size, the 1975E data suggest that re-

cessive small leaf siZe was conditioned by the n3n genotype. The

392°2

presence of either N2 or 03 in the genotype conditioned the production of

large non-wrapper leaf size. 1975L data suggest that the Red Danish geno-

type n conditioned the recessive trait for large leaf size. The

2"2°2°2

presence of either N or o in the genotype produced small leaf size.

3 3

224

 

a

 

m.~N.u e.o__ m.e w.e_ m.m e.m~ m.NP. e.mp m.m _.~ N.e ea 8 x_Aea x av
tamedm 0m We 92 93 93 Gm 0m ~_ Na*%axfiv
e.eN.u N.Ne m.P N.m m.e e.~ m.~ m.m N.., m.o~ ~.m_ e.o~ m.m em_ Ne Rea x Nev
m.e~.u e.oe o.F a.~ m.m w.m “.8 e.m o.~. m.ep m.m_ m.P~ M.“ aow Na Awe x eav
e.a~.u e.ea _._ e.mp e.m a.m o.o_ o.__ m.m_ a.m m.m e.e N.e om Aea x may
o.e~.u N.mm o.m m.m _m.- e.__ ~.e_ .~.m e.ep e._P N.e ~.m o._ em Ame x eav
a.om.u ~.m~ A.“ N.N . _.m N.o_ _.m ..m m.N_ e.mp m.o~ N.op am away eeoepeam someem
e.em.H o.mm m.~ m.m m.m m.__ ~.m m.e_ e.m m.e_ m.__ m.~ m.e_ mm Aeav emeeaa wax
eaoz mew me_ am. app mo_ mm mm as me am ea wwewwn

mmmpu mo p_sw4 swan:
msmsu cw «New comb smagmszucoz

 

 

.eeo=.Fem tomeem x eaweea eom maeeo ago 46 oe_peapa “mew,
mgu com ucmucma cw mcowuzawsumwu mNWm camp cmaqmsznco: mmosuxumn ucm me ._u .ucmema .om_ mpamh

225

 

~.o~

on we Aee x Nev

 

m.m~.u 0.08 m.m e.e~ e.em 5.0 m.m

N.eP.H m.ee e.o 0.? m._ m.m m.m_ e.~e o.m~ e.~ _am we Ame x eav
e.m .H e.mm o._~. e.ee e._m. m, Ree x Nev
m.NP.H m.~e m.e m.om F.0N m.em mm Ame x eav
~.mP.H e.Pe o.~ ~.a m.e_ .o.me e.- _.e me Away eaoe_Pam Loosen
e.m~.H m.em N.e _.m m.~F e.m e.m e.mp o.m~ mm . Aeav emweao no“

see: +mm_ mp_ mm am am am me am a, a aoee_a .
ca .62

mmmpu do “wee; swan:
mamsw cw mNPm wow; smaaws31=oz

 

 

.eeoe~_am tomeam x smegma ode amoeo one to mewoeaee 4m~m_

one so» “causmq cw m:o_u:nwspmew mNPm wmmF smqamszlco: mu tam Pm .pcmcmm .mm— mpnm»

226

Both the 1974L and 1975E F1 populations of Red Danish x P.I.
215514 suggest overdominance for large non-wrapper leaf size. How-
ever, the 1975L F1 planting suggests non-wrapper leaf size was inherited _
in an additive manner (Tables 11-13). Environmental conditions can
affect non-wrapper leaf size.” Both Red Danish and P.I. 215514 pro-
duced large leaf size phenotypes in the 1975E planting (Table 138).

The 1975E F2 population of this cross pr0duced a 19:292 segregation

ratio using a dividing point of 59 gm. The observed data fit (P=0.92)

a 1:15 model (Table 130). A genotype of n4n4020 Zwas suggested for

small non-wrapper leaf size. The presence of either N2 ,from Red Danish,
or 04 ,from P. I. 215514, in the plant genotype produced large non-
wrapper leaves. .

All 3 plantings,_1974L. 19755 and 1975L, of Red Danish x Chief-
tain Savoy F1 suggest overdominance for large non-wrapper leaf size ‘
(Tables 11-13). Chi-square homogeneity tests suggest F2 populations
of Red Danish x Chieftain Savoy could be pooled. 1975E F2 data suggest
small non-wrapper leaf size was recessive (Table 140). Using a 59 gm
dividing point to separate phenotypic classes an obserVed ratio of
18:191 was produced. The data fit (P=0.24) a 1:15 model (Table 130).
Backcrossing the F1 to Red Danish produced a ratio of 0:18 while cross-
ing the reciprocal F1 to Red Danish produced a 4:33 ratio. Ratios
of 0:18 and 0:37 were expected. Since both parental distributions pro-
duced large non-wrapper leaf size the 1:15 model suggests that the re-
combinant genotype 1251250202 conditioned small non-wrapper leaf size.)
The presence of either N2 or 05 alleles in the plant genotype condi-
tioned the production of large non-wrapper leaVes. Using a 64 gm di-

viding point to separate the data into phenotypic classes the pooled

227

 

Nu.ama x «my

 

mpm

we .02

mmmpu 4o u_sw4 swag:
mango cw «New woo; cmaqaszucoz

e.mN.H m.mm m._ m.m_ 8.5 ,a.~_ N.N, m.~. e.__. m._P F.~ m.m e.N __m
. m.m~.fi ~.~__ N.N e.m~ _.m_ o.mp e.- m.P_ N.N e.m e.m F._ p._ mm fies x mev
m.mN.H e.mpp ..e_ w.m_ m.ep m.~_ _.mp N.op e.~ m.m N.P ~._ . m.~ mm Ame x adv
m.m~.H N.ea e.e_ we.m o.m o.F_ m.mm 4.8? e.m .mp “may epmm_~ .H.e.
a.am.H o.mw m.~ e.m .e.m m.PP ~.m m.ep e.» m.ep m.P_ m.~ m.eP .mm Aeav ;m_eao eom
eaoz me_ amp opp mo, mm am me mm mm ea woeePa

 

 

. .cpmm_m .~.a x ;m_:ao com mmosu we» mo acmacmpa mmmmp
we“ so; ucmucma cw meow»:a_sumwu «New vamp smaamczscoc mm new pm .“cmcwa .wmp w—nm»

228

 

..e _.e m.ep N.e~ e.P~ weep ..e e_ Rea x nee

 

m.e~.u N.em
e.-.H N.mm m.~ m.~ m.e m.~ e.mp e.e N.e_ e._m mlm_ _ ea Ame x eav
N.Ne.u e ee_ e.em e.mp e.mp N.N e.m_ N.N MP Amav e_mm,~ .H.a
“.mN.H e.em ~.e ,.m m.~_ e.e_ e.e e.e e.m_ e.m~ mm 3 Aeav eaeeee eoe
eaoz .mmp e_P me me me ee mm me an m_ . meea_a
. ....O . oz

mme_u we peeve gene:
maeso cw eNWm meme ceeeecz:cez

 

 

..epmm—N .H.e x smpceo emu mmecu one we mcwucepe emsmp
ecu Lem Hemesea =_ mcewu:e_gum_e m~_m weep Leaeesz:ce: Fm e:e.ucwgmq .mm— m—eeh

229

 

smut; N.N Te 3: ea. 3: 92“.: we: fie e..m 3 B 5:: xmn:
m.mm....e.§ 8% .m.- 7: e5. ea c: 2 Jim... xet
NRHmeS e; E: e.: 3: we 3: ea 3 3 3 ea NS Nieaxms
93.3.8 2 3w 3: ..2 N: :2 3: .2 me e.m e.~ BF Nimexeav
Nauumezee one; «.2 3: we 3. e.m mg e.m ....N Z 8 taxes;
eemnmefi 7.2 mam :2 ..e 3 ..2 Fe ea fie ea . mm . 31.:
INNS: , .3 at e.: 5.8 2 .....8 a: em 3: seem £3.55
e.em.+.e,.ee. e.~ est. e.e 9: 3 3: ed a: a: Z 3: mm 3: 5.28 E.
new: .

mpw may .mNF m—_ mop mm mm mm mm am we mucepm
. we .ez

mme_u so peeve seen:
mseeo cw eNpm meme ceeeeczncez

 

 

.>e>em eweueemsu x smegma eem mmece we» we memuce—e mmum~
one Lew aceegee cw meemueewsumwe erm temp seeeeuz::e= mmeeexeee ecu Nu ._m .ueesee .oeF open»

230
F2 data from the 1975L planting produced a segregation ratio of 336:87

(Table 141). This data fit (P=0.34) a 13:3 model (Table 131). The

presence of dominant o alleles or the action of homozygous recessive

5
alleles. ”sns’ conditioned the production of small non-wrapper leaves.
The Red Danish genotype N2N20202 conditioned the production of large
nonéwrapper leaves.

Non-wrapper leaf size heritability estimates for green cabbage
F1 populations range from 0.52 to 0.84 with an 52 value of 0.68 i 0.53
and an E: value of 0.79 i 0.43. Three h2 estimates were greater than
1 and 1 estimate was negative. Two estimates were within the 0 to 1
heritability range. Red x green cabbage F1 heritability estimates
range from 0.40 to 0.70 with a mean heritability estimate of 0.53 i
0.33 and an E: estimate of 0.61 t 0.30. Three estimates were within
the 0 to 1 heritability range and 3 estimates were greater than 1.
Red x green cabbage heritability estimates suggest that greater than
50% of the F1 population variance was due to additive genetic effects

that may be fixed in cabbage by inbreeding (Table 18).

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CONCLUSIONS

This study has shown that components of cabbage plant efficiency
are heritable traits. Seven traits in cabbage: maturity, head weight,
total non-wrapper leaf weight. stalk size, non-wrapper leaf number,
efficiency index and non-wrapper leaf size. are each controlled by
2 loci. This type of inheritance is not unique in cabbage. The length
of the longitudinal and transverse diameters of cabbage heads was
found by Krauze (12) to be controlled by 2 dominant genes. Studies
by Dickson (6) found cabbage core length to be controlled by 2 genes
each incompletely dominant. The dominance direction was affected by
the half sib related F1 and F2 populations using either Baby Head or
Red Danish as parents. The presence of gross changes in dominance
direction in cabbage may be triggered by environmental sensors that
modify genotypic dominance to produce an optimum phenotype. Reversals
in dominance patterns have been noted by Shifriss (28) in acorn squash.
Modifications in dominance have also been shown by Russell and Srb
(26) in Neurospora. They observed that while the bulk of their modi-
fiers "increased" the dominance of the wild type allele, 2 modifiers
"decreased" the dominance of the wild type allele. Both Russell (26)
and Shifriss (24) showed modification to be dosage dependent. stronger
in the heterozygous condition than in the homozygous condition. Fur-
ther studies by Srb and Basil (30) show that the genotypic background
of Neurospora can trigger a completely dominant or completely recessive
phenotype when dominant Pk-4 mutants are introduced into 3 species of

this organism. Dominance patterns suggest that more efficient cabbage

232

233 1
cultivars could be produced without sacrificing head quality or yield.
Heritability estimates were greatly influenced by the environment. The
wide range of estimates suggests that selection of traits with high)
heritability_estimates, days to maturity, head weight, and leaf number,
. for green cabbage crosses and stalk size for red x green cabbage crosses,
.may increase the rate of progress especially if they are favorably'
correlated with low heritability traits. Heritability estimates for
all.traits ranged from 0.96 to 0.05. Heritability estimates were larger
in green cabbage crosses than in red x green cabbagecrosses. The pre-
sence of very large and very smallheritability estimates between p1ant-
ings provide an area for further study. Genetic correlation data sug-
gest that components of plant efficiency are correlated and selection
for a given traitmay simultaneously affect other traits in cabbage.
Data from this study produced genetic correlations greater than 1 and
less than negative 1. These abberant estimates may be due to sampling
error or a violation of assumptions necessary for genetic correlation
estimates. Estimates of this type have also been found by Sharma (27)
in cabbage and Burton (2) in pearl millet.

A generation of added inbreeding produced significant changes in
the following traits: head weight, total non-wrapper leaf weight, non-
wrapper leaf number and efficiency index. Additional inbreeding should
be applied to open-pollinated cabbage cultivars prior to undertaking
genetic studies of these traits. For this sample of cultivars. matu-
rity stalk size and non-wrapper leaf size appeared to be genetically
homozygous. Genetic studies of these traits could be undertaken with-

out added inbreeding.

234

More study is needed to determine whether the change in dominance
pattern observed between green cabbage crosses and red x green cabbage
crosses is a result of the cultivars chosen or whether all red cab-
bage cultivars could be expected to provide the same response.) If this
response is conditioned by all red cabbage. it may be possible to use
red x green cabbage crosses to provide more desirable phenotypes in
greater abundance than could be done breeding red and green cabbage
separately.

-Efficient green cabbage cultivars can be developed by crossing
large headed. large leaf cultivars to small headed. small leaf culti-
vars. The resulting hybrid should have the large head of one parent
and the smaller leaVes of the other parent. Selection for efficient
plants could be practiced in the F2 generation of this cross. Large .
amounts of additive genetic control should increase the nUmber of plants
selected in future generations. The development of more efficient
compact plants should stimulate the development of new cultural prac-
tices including closer spacing. Information from this study may be
of value not only as an aid in cabbage breeding. but also as a stimulus

for future research in other Brassica species.

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10.

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12.

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