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V

FLOWER COLOR INHERITANCE IN SALVIA SPLENDENS
By
Elizabeth Ann Robertson
A THESIS
Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

MASTER OF SCIENCE

Department of Horticulture

1985

ABSTRACT

FLOWER COLOR INHERITANCE IN SALVIA SPLENDENS

By

Elizabeth Ann Robertson

Eight distinct flower color classes of Salvia splendens
were intercrossed.. The F1 plants were self-pollinated to
produce the F2 generation and backcrossed to both parents.
The F2 and backcross pOpulations were statistically analyzed
and compared. Seven genes were found to control flower color
inheritance. The R and L genes control the colors red, R_L_,
rose, rrL_, salmon, R_ll, and pink, rrll. The Int gene had
no effect on the R and L genes when dominant, but when reces-
sive with dominant R and L, it produces rose color. The P
gene controls colored, P_, versus white, pp, flowers. Purple
color is produced by the V gene and burgundy color is control-
led by the B gene. The E gene dilutes the colors produced by
the other loci. The R, L, Int, and P genes exhibit complete
dominance while the V, B, and E genes are incompletely dom-

inant. The R, B, and E genes were discovered in this study.

ACKNOWLEDGMENTS

Very special thanks are due to my generous mentor,
Dr. Lowell Ewart. Through his guidance I drew inspiration
and motivation for my work. I would also like to thank my
other committee members, Dr. A. Iezzoni and Dr. J. Kelly,

for their interest and counsel.

ii

List of Tables. . . .

List of Figures . . .

Introduction

Literature Review . .

Materials and Methods

Results . . .
Discussion .

Summary . .

Appendix. . .
Bibliography

TABLE OF

CONTENTS.

iii

Rage
iv

12
31+
37
38
79

Table
1.

2.
3.
4.
5.
6.
7.
8.
9.
10.
11.

LIST OF TABLES

Salvia Parents Used in This Study

Color of the F1 Plants

Description of the Salvia Flower Color Genes
Segregation Data for the R, L, and Int Genes
Segregation Data for the P Gene

Orange X Pink F2 Segregation

Segregation Data for the V Gene

Burgundy X Pink F2 Segregation

Parent Genotypes

Summary of Crosses

Status of Genes

iv

1.
2.
3.
1+.
5.

Interaction
Interaction
Interaction
Interaction

Interaction

of the
of the
of the
of the
of the

LIST OF FIGURES

<"U"U"U!U

w<wwt€llfl

and Int Genes

R, L, and Int Genes
R, L, and Int Genes
R, L, and Int Genes

and E Genes

P_a_g_e_
29
3o
3 1
32
33

INTRODUCTION

Salvia splendens F. Sellow ex Roem. & Shult. (1) is a
member of the Lamiaceae (Labiatae) or mint family. Original-
ly from the Brazilian trOpics, it is a perennial in warm
regions but grown as an annual in areas exPeriencing freezing
temperatures. S. splendens has racemose inflorescences and
deciduous bracts. The calyx is campanulate with the tubular
corolla extending 1 to 1% inches. The lower lip of the
corolla is much reduced (1). According to the most recent
study, the gametOphytic chromosome number is 22 (2). Common-
ly called Scarlet Sage, scarlet red is the original and most
pOpular color, but once in cultivation, purple and white
forms were quickly discovered. Plant breeders have expanded
the color range to rose, salmon, pink, bicolors, and various
shades of red.

Unlike its herbal relatives, S. splendens is a popular
ornamental Species used primarily as a bedding plant but also
as a cut flower. Salvia comprises 5 to 6 % of the bedding
plant crOp for most growers (9). In 198# the wholesale value
of bedding plants for the top 25 states in the country was
over $209 million (16). According to these figures over 310
million is Spent annually on salvia alone and its pOpularity

is growing. The objective of this study was to determine

the number of genes controlling flower color inheritance
in g. splendens. Such a determination would be quite
valuable to plant breeders wishing to expand the current
color range available to the gardening public.

LITERATURE REVIEW

In 1960 Paris, Haney, and Wilson (12) conducted a sur-
vey of flower color inheritance studies. They found that
Vavilov's (17) theory of homologous variation was quite

valid. The gene types most commonly encountered were:

W produces color; ww produces white
Iv produces non-ivory; iviv produces ivory
Y produces non-yellow; yy produces yellow
B produces purple or magenta; bb produces blue
P produces purple or magenta; pp produces pinks, roses,
or reds
Dil produces intense color; dil dil produces dilute color

The genes were named for their recessive phenotype and in
some species the dominance was reversed or the genes were
incompletely dominant in effect. No ivory (Iv), yellow (Y),
or blue (B) colors were encountered in this study, but the
presence of W, P, and Dil genes was probable.

In 1964 Hendrychova-Tomkova (7) investigated flower
color inheritance in S. splendens. She found that four
genes controlled the parental colors white (S. splendens
forma glgg), purple (f. violacea), red (f. 2223;), rose
(f. ragga), and salmon (f. carnea). The pigment, P, gene
controlled colored (P_) versus white (pp) flowers. Watts

(18) also made reference to this gene for white color in his

1980 text on plant breeding. This gene is analogous to the
W gene previously mentioned.

Another common gene is the purple versus nonpurple type,
and indeed Hendrychova-Tomkova (7) found a single gene for
purple color (V_) versus red color (vv). She noted its
incompletely dominant nature but did not quantify the hetero-
zygous (Vv) class separately from the VV purples. Both the
purple (V) and white (P) genes were epistatic to the other
genes and partly epistatic to each other. A cross of white
and purple segregated some ppV_ individuals that she called
white but counted separately rrem the ppvv whites, indicating
some color difference due to the V gene. In that cross (and
white backcross) disprOportionately low numbers of red and
ppV_ white recombinants indicated a strong linkage relation-
ship between the P and V genes.

Hendrychova-Tomkova (7) also did a pigment analysis of
the parental colors. White contained no anthocyanins, and
purple was due primarily to violet and purple derivatives of
delphinidin and cyanidin with small amounts of pelargonidin.
The red group (red, rose, and salmon) was composed largely
of salvianin, a pelargonidin derivative, with small amounts
of purple cyanidin. Red, rose, and salmon differed from
each other in the concentration and localization of the
pigment which was found to be governed by two genes working
in complementary fashion. She called these genes Int, inten-

sity, and L, limited localization. Red resulted when the

dominant alleles of both loci were present (Int_L_). A weaker
concentration of the intense red color produced rose (int int
L_). Salmon (Int_ll) was also a dilution of red color, but
in this case, the pigment was largely confined to the outer
corolla epidermis, the inner epidermis being nearly white.
Crosses of red with rose or salmon produced monogenic ratios
of 3:1. Rose and salmon crossed together produced red F1
plants which segregated in a dihybrid ratio of 9 red: 3 rose:
3 salmon: 1 pink with light rose or pink as the double reces-
sive (int int 11). Crosses of purple with rose and salmon
segregated two different shades of light violet (V_int int

L_ and V_Int_ll) with the expected colors of purple, red, and
rose or salmon.

In addition to the above mentioned colors, orange and
burgundy colors were also included in this study.

Orange color in many flower Species is the result of
yellow and red c0pigmentation (10,6), but the orange in this
study is actually closer to a very deep shade of salmon than
to the vivid orange color the word implies. This indicated
that the orange color may be the result of a diluter gene or
multiple allele system of the red color. A diluter gene
like the type mentioned by Paris et al. (12) would dilute
all colors, such as, in stocks where Schnack et al. (13,14)
found the gene P diluted purple to light purple, wine to
pale wine, and light red to rose. Similarly, Mehlquist and

Geissman (10) working on flower color in carnation found a

gene M that altered the sugar portion of the pigment molecule.
The result was diluted shades of the colors that carried the
dominant allele. Another possibility for orange color was

a multiple allele system, such as MHntzing (11) found in
Galeogsis where the locus R had four alleles that differenti-
ated red R, light red R1, faintly red R2, and white r.

The burgundy color in salvia is a relatively new devel-
opment. It could even be described as variegated Since the
body is burgundy colored while the upper lip is scarlet red.
Hendrychova-Tomkova (7) found that the red color contains
small amounts of cyanidin. Genes that regulate cyanidin to
pelargonidin are known. Lawrence, Scott-Moncrieff, and
Sturgess (8) found that cyanidin (magenta/rose) versus pelar-
gonidin (pink/salmon) pigment was monogenically controlled
in Streptocarpu . Very similar results were reported by
Mehlquist and Geissman (10) who found a single gene distin-
guished cyanidin (magenta/crimson) from pelargonidin (rose/
scarlet). A magenta colored mutant of Salvia coccinea was
found in a red population. D'Cruz and Jadhav (4) made
crosses with magenta and concluded it was monogenically

inherited. The pigments, however, were not analyzed.

MATERIALS AND METHODS

All the plants used in this investigation were grown at
the Plant Science Greenhouses at Michigan State University.
Dwarf, early flowering and primarily day-neutral cultivars
were used, therefore, no supplemental lighting or black cloth
was necessary. The plants were grown under normal conditions
recommended for salvia culture. The parent and F1 plants
were grown in 6 or 8 inch pots and fertilized at regular
intervals.

Salvia plants have long flower stalks with the individ-
ual tubular flowers whorled about the inflorescence. The
flowers eXpand from the bottom of the inflorescence upward,
with each flower lasting a single day. The stamens are curved
back within the flower bud. By inserting tweezers into the
side of the bud, the stamens become wrapped around the tweezer
points and can be pulled outward with minimal damage to the
style and stigma. Bud emasculations were done daily on
flowers to be pollinated the next morning. The pollinated
flower usually yielded 2 to 3 seeds with a maximum of A
possible. A whole spike would be labeled and pollinated
repeatedly, until the amount of seed needed was accumulated.

Once fertilization was achieved, the seed grew quickly

with seed harvest following 2 to 3 weeks later. It was

necessary for the seed to be completely dried before planting,
otherwise, germination was severely delayed. Optimally, seed
germination requires 12 to 15 days at 210 C. A fungicide
drench was applied at the time of planting and again after
transplanting. The F2 generations were grown in flats of #8
plants each, while the backcrosses were grown in either #8

or 32 insert flats. Flowering in the flat occurred approxi-
mately 8 weeks after transplanting.

Color evaluations were made using both volumes of the
Horticultural Colour Chart, H.C.C. (3). AS a British publi-
cation the H.C.C. color names do not correspond to the names
used in the United States. To avoid confusion, the color
class names are those currently used in the horticultural
trade and only the H.C.C. number is listed.

Colors were evaluated in the greenhouse on overcast
days when the colors were most intense, and light quality
was bright but diffuse. This was necessary Since some color
Shades are indistinguishable under bright direct light. All
evaluations were made on newly Opened flowers. The main
body of the flower differs in color intensity from the upper
lip, but, in most cases it differed by only one color grada-
tion. The white color class is a pure bright white and there-
fore does not have an H.C.C. number. The parent lines with
their respective color class, seed source, and H.C.C. numbers

are listed in Table 1.

 

 

 

Table 1. Salvia Parents Used in This Study
Color H.C.C. Number
Class Parent ' Source Vol.# Color #
White Carabiniere White Pan American - ----
White CISOpatra Mix Royal Sluis - ----
Pink F2 Selection --- 1 21/2
from Pixie
Rose Cle0patra Mix Royal Sluis 2 .20/1
Rose Carabiniere Cherry Pan American 2 .20/1
Rose F2 Selection --- 2 e20/1
from Pixie
Salmon CleOpatra Mix Royal Sluis 2 620/1
Salmon Champagne Harris 2 620/1
Salmon F2 Selection --- 2 620/1
from Pixie
Red Pixie Harris 1 18/0
Red Tally Ho Harris 1 18/0
Red CleOpatra Mix Royal Sluis 1 18/0
Orange Carabiniere Orange Pan American 2 619/0
Burgundy Burgundy Harris 2 82A/3
Purple CleOpatra Mix Royal Sluis 2 931/2
Purple Purple Blaze Sluis & Grut 2 931/2
* The two or three digit number represents the flower color
while the number after the slash indicates the degree of

intensity.

10

In order to cross each color class with every other
color, it was necessary to self-pollinate the parent lines
to determine if they were true-breeding for flower color.
All parents proved true except Pixie, which was known to be
an F1 hybrid from rose and salmon parents (5). Therefore,
the self seed would segregate as the F2 generation. A true-
breeding plant may carry inhibitor or epistatic genes that
hide the effects of other genes. Those other loci may or

may not be homozygous. Consequently, F plants were not

1
interpollinated and seed was collected separately from each
plant. .

In most cases 10 to 20 F1 plants from each cross were
used to evaluate color uniformity. The F1 plants were self-
pollinated for the F2 generation and backcrossed to each
parent. Backcross pOpulations numbered at least 20, while
the F2 generation contained 200 to 300 plants. F2 and
backcross segregations were quantitatively analyzed using
the Chi Square statistic. Sometimes it was necessary to
combine color classes of an incompletely dominant gene (i.e.
BbrrLL and BBrrLL where the B gene is incompletely dominant),
when the colors were not Significantly different. By compar-
ing segregations, it was possible to develOp genotypes for
the phenotypes. From there, line drawings were made to
demonstrate the interaction of all seven genes for flower
color. Calyx color was not evaluated quantitatively, but

there seems to be a strong linkage between calyx and flower

11

color genes. As new colors arose in the study, they were
matched with the Horticultural Colour Chart and their H.C.C.

numbers appear in the line drawings.

RESULTS

Parents of the same color class were crossed together
when parents from different sources were available. The
intercrossing of whites produced white F13 and all white F2
progeny. Likewise, crosses within the color classes salmon,
red, and purple also produced self-colored progeny. Only
the cross rose (Cleo) X rose (CA Cherry) exhibited complemen-
tary genes for the Single color phenotype. The F1 was red
and the F2 segregated 9 red: 7 rose/cherry (Table A), demon-
strating complementary epistasis. The two genotypes for
rose color are designated hereafter as rose (R gene) and
cherry (Int gene). The colors of the F1 plants are presented
in Table 2.

The genes discovered by Hendrychova-Tomkova (7) retain
their original designations. Only the R, E, and B genes
are new to this study. The R and E genes are named for the
recessive phenotype, while the B gene name represents the
dominant phenotype. The practice of naming genes for the
biosynthetic pathway they control or effect is a good one,
but this aspect was not studied, and therefore, it was neces-
sary to use phenotypic designations. The gene names with their
recessive and dominant phenotypes are presented in Table 3.

The gene action is also noted.

12

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The R and L genes control the colors red R_L_, rose
rrL_, salmon R_ll, and pink rrll (Table 4). Aside from these
colors, dominant R and L genes are found in all other parent
colors.

The Int gene was discovered from the parent cultivar
Carabiniere Cherry. Rose and cherry are the same color but
they are designated separately due to different genotypes.
The cherry genotype results when recessive int int is com-
bined with dominant R and L genes. A cross of cherry X
salmon segregated the color ratio 9 red: 3 cherry: 3 salmon:
1 pink (Table A). The pink color from cherry X salmon is
indistinguishable from the pink color by rose X salmon. Thus,
rose and pink colors can be formed in two ways. Rose color
is Int_rrL_ or int int R;L_ while pink color is Int_rrll or
int int R_ll. Recessive int int with recessive rr produces
a shriveled flower stalk. The color is either rose or pink
depending upon the status of the L gene. The shriveled
condition is characterized by reduced and distorted flower
buds which tend to fall off before Opening. Those that do
open have pollen but fail to set seed when pollinated.

The P gene controls colored (P_) versus white (pp)
flowers. ReceSSive pp is totally epistatic to the R, L,

Int, and E genes and partially epistatic to the V and B

genes (Table 5). Recessive epistasis with two genes produces
a 9:3:A ratio where A/16 would be white. White crossed with
purple separates the white portion into 1 pure white (VVpp)

16

Table 4. Segregation Data for the R, L, and Int Genes

 

 

Color Genotype Ratio Obs. Exp.
e se F2

Red 1 R_ Int_, 9 129 133.9

Rose/Cherry rr Int_ & R_ int int 6 94 89.2

Rose Shriveled rr int int ‘1 15 15,9
16 238 238.0

D2/e : .438 P > .80

Rose Salmon F2

Red R_ L_, 9 132 135.6

Rose rr L_ 3 39 45.2

Salmon R_ ll 3 54 45.2

Pink rr 11 l, 15 15,0
16 241 241.0

DZ/e = 2.724 p > .30

Chagry X Salmon F2

Red L_ Int_ 9 135 133.31

Cherry L_ int int 3 44 44.44

Salmon 11 Int_ 3 ,45 44.44

Pink 11 int int 1| 13 1S,§1
16 237 237.00

DZ/e = .233 P > .95

Pink X Cherry F2

Red R_ L_ Int_ 27 44 54

Rose/Cherry rr L_ Int_ & R_ L_ int int 18 35 36

Salmon R_ ll Int_, 9 27 18

Pink rr 11 Int_ & R_ ll int int 6 15 12

Rose Shriveled rr L_ int int 3 4 6

Pink Shriveled rr 11 int int 1 3 2

64 128 128
Da/e==8.294 P) .10

 

17

 

 

Table 5. Segregation Data for the P Gene

Color Genotype Ratio Obs. Exp.

2i2§.$lfl2$£2 F2

Red P_ R_ L_ 27 159 144

White pp -- -- 16 77 85

Rose P_ rr L_ 9 43 48

Salmon P_ R_ 11 9 48 48

Pink P_ rr ll 3 _1_5 _1_§_
64 341 341

132/e : 3.084 P > .50

W F2

' Rose Purple P_ Vv 6 113 110.6

Purple P_ VV 3 62 55.3

Red P_ vv 3 67 55-3

Purple-White pp V_ 3 42 55.3

White pp vv 1. _ll 1§,§
16 293 295.0

DZ/e=9.576 P >.01

W F2

Red Burgundy P_ Bb 6 77 79.0

Red P_ bb 3 48 39.6

Burgundy P_ BB 3 37 39.6

Pink-White pp B_ 3 31 39.6

White pp bb _1_ fl 13,2
16 211 211.0

DZ/ez5.613 P) .20

 

18

to 3 purple-tinged whites (VvPP). Likewise, white X burgun-
dy segregated pure whites and pink-tinged whites (Table 5).

-The E gene dilutes the color produced by the other
genes. The only source of the E gene was the parent color
orange which was found to be a dilution of red. Since E
is incompletely dominant, there are three shades of every
color: intense color ee; light shades Be; and dilute shades
EE (Table 6). For example, rose X orange (Cross # 24) seg-
regated red, light red, and orange, plus rose, light rose,
and dilute rose. The E gene also dilutes the purple and
burgundy colors, but, Since they are incompletely dominant
as well, the ratios are greatly expanded. In the case of
orange X purple (Cross # 34), the diluted forms of VV and Vv
were identical in color so they were combined into a single
class called dilute purple. Similarly, diluted BB and Bb
were combined into one color class (Cross # 33).

The incompletely dominant V gene controls purple versus
red color. Homozygous dominant VV with dominant L and Int
genes produces purple color while VvL_Int_ is rose purple
color. Salmon X purple and purple X pink crosses segregated
a new light violet color with the genotype V_llInt_ indicat-
ing 11 dilutes V in some way (Table 7). The Int gene also
effects purple color. Cherry X purple segregated dusty
purple, VVR_L_int int, and dusty rose purple, VvR_L_int int
(Table 7). The color in both cases was greyed and the

flower stalk was somewhat reduced but not shriveled. The R

19

Table 6. Orange X Pink F2 Segregation

 

Color Genotype Ratio Obs. Exp.

 

WFZ

Light Red Ee R_ L_ 18 65 67.50
Red ee R_ L_, 9 40 33.75
Orange EE R_ L_ 9 33 33.75
Light Rose Ee rr L_ 6 22 22.50
Light Salmon Ee R_ 11 6 21 22.50
Rose as rr L_, 3 15 11.25
Salmon ee R_ ll 3 11 11.25
Dilute Rose EE rr L_ 3 9 11.25
Dilute Salmon EE R_ 11 3 12 11.25
Light Pink Ee rr ll 2 6 7.50
Pink ' ee rr ll 1 3 3.25
Dilute Pink. EE rr ll 1 __3 3,25

64 240 240.00

D2/e23.731 P>.975

 

20

Table 7. Segregation Data for the V Gene

 

Color Genotype Ratio Obs. Exp.

 

Eurp;e X Pink F2

Rose Purple Vv -- L_ 24 79 90.00
Purple VV -- L_ 12 53 45.00
Light Violet V_ -- ll 12 42 45.00
Red vv R_ L_, 9 41 33-75
Rose vv rr L_ 3 11 11.25
Salmon vv R_ 11 3 7 11.25
Pink vv rr ll 1 __2 3,23
64 240 240.00
DZ/e:8.9l+9 P>.10
W F2
' Rose Purple Vv Int_, 6 75 72
Purple VV Int_ 3 36 36
Red vv Int_ 3 34 36
Dusty Rose Purple Vv int int 2 22 24
Dusty Purple VV int int 1 14 12
Cherry vv int int 1 11 _lg
16 192 192

132/e: .819 P>.975

 

21

gene seems to have no effect on the V gene since the rose X
purple cross showed no new segregates. The B gene also had
no effect on the V gene, at least in the presence of dominant
R, L, and Int genes which both parents carried.

The burgundy color is rather unique because it is actu-.
ally two colors. The body is burgundy (or wine) colored
while the upper lip is intense red. Burgundy body coloring
is produced by the B gene with dominant R, L, and Int genes.
In the hybrid condition (Bb), the body color becomes more
reddish than burgundy. The R, L, and Int genes determine the
basic color of the flower while the B gene lends a purple
cast to the body, thus red becomes burgundy and rose becomes
fuchsia (Table 8). -The lip color is generally as it would
appear if the B gene were recessive._ The H.C.C. numbers
given to the burgundy related colors represent the body color
only.

By analyzing the F2 and backcross populations, it was
possible to determine the parent genotypes presented in Table
9; as they were all homozygous, only the haploid genotypes
are given. A summary of crosses is presented in Table 10.
For each cross, the F2 ratio tested, the Chi Square value,
and the probability is given. Table 11 gives the segregating
genes for each cross. The complete F2 and backcross segrega-

tions can be found in the appendix by using the cross number.

22

Table 8. Burgundy X Pink F2 Segregation

 

 

 

 

 

Color Genotype Ratio Obs. Exp.

Burgggdy x Eink F2

Red Burgundy Bb R_ L_, 18 127 112.50

Burgundy BB R_ L_. 9 51 56.25

Red , bb R_ L_, 9 53 56.25

Peach-Rose B_ R_ 11 9 41 56.25

Fuchsia Bb rr L_ 6 42 37.50

Maroon BB rr L_, 3 23 18.75

Rose-Pink B_ rr ll 3 21 18.75

Rose bb rr L_ 3 9 18.75

Salmon bb R_ 11 3 23 18.75

Pink bb rr ll 1’ 1O §,23
64 400 400.00

Da/e=16.735' P>.05

Table 9. Parent Genotypes

Parent Color Class Genotype

White p v b e R L Int

Purple P V b e R L Int

Burgundy P v B e R L Int

Orange P v b E R L Int

Red P v b e R L Int

Rose P v b e r L Int

Salmon P v b e R 1 Int

Pink P v b e r 1 Int

Cherry P v b e R L int

 

23

 

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mo. om:.m _"mumum seas x eem m—
mum. _ms.m _"_umumumumumumumumumuw. . Seam x emsuao m.
0.. :mm.w _"mumumum_num . Sheena x ream _—
om. Smo. _um seem x Read 0.
0.. mom._ _"m sonata x read m
_o. mum.m _"mumumuo . manned x means a
om. n_o.m _"munumuw . assemasm x eras; S
on. rem. _um sum x than; 6
om. sum. numuduw emeuao x mass; m
on. us... manna assess x seas; :
om. mms._ musum enom x than; n
on. .__._ mueum desaem x mass; m
on. Smo.n mnmumum_uum . even; x Adam _
nomwmwm maosdm H20 oanmm mm mmoao Amnssz
m mocha

 

mommoao mo hamsssm

.O— manna

24

 

N

 

axon as vopaomohm owam Sump m a
0m. mm..e .u..mum.m"0 seesmasm x seem 0m
0m. .wu. .un sem x emom mm
0.. ssm.s .u..mum.m.m emasao x seem em
00. one. ..0.0 . seem x saneno mm
0m. s0m.m ."mumumum manned x dosaem mm
on. mon.. .umumumnm sessmasm x aosaem .m
00. mmo. ."m sem x seaHsm 0m
0.. 000.S .....m.m.m”0 emeeao x sesaem 0.
mm. mmm. .“m.m.m . sesaem x aaaeno 0.
0m. nmu.m .unumum . eeaasm x enom S.
00. nan. .umunum damn maxed 0.
0.. 020.0 .umumuaum..m.usm . read x manned 0.
m0. mmu.0. ..m.m.m.mumumumumum. . Read x.»asemasm S.

nowwwww mamzwm Hno oaaom mm mmoao 909852
m . mmoao

 

A.0.neoev 0. eases

25

 

 

4x0» SH umpsmmoam omHn 0000 mm *

 

00. 000.. .4..m.0.0 044404 0 00000400 mm
00. 00.. ....m 000 x e40404 0m
00. 000. , .4..m 000 0 00000400 00
00. 00..m .u....m.m.m.m"0 044404 x 000040 00
00. 000.0 .4....mumum.m.0 00000400 0 000040 00
00. ..m. .4..m 000 x e00040 mm
000. 0.0. .4."m.m.m"0 . 040404 x 044000 .m
00. 000.0 .u..m.m.m.0 00000400 x 044000 00
00. 00.. ..m 000 x 444000 00
00. 000.0 .u..m.mum.0 044000 x 000040 00
.0. 000.0 ..m.:.0 044404 x 0000 mm
ammwwww oawsvm Hno 0Huum NH 00040 409832
m mmoho

 

A.u.u:oov 0— 0H968

26

 

 

 

 

Table 11. Status of Genes

Cross Status of Genes *

Number cross P v B E R L Int
1 Pink X White S r r r S S d
2 White X Salmon S r r r d S d
3 White X Rose S r r r S d d
4 White X Cherry S r r r d d S
5 White X Orange S r r S d d d
6 White X Red S r r r d d d
7 White X Burgundy S r S r d d d
8 White X Purple S S r r d d d
9 Pink X Salmon d r r r S .r d
10 Pink X Rose d r r r r S d
11 Pink X Cherry d r r r S S S
12 Orange X Pink d r r S S S d
13 Red X Pink d r r r S S d
14 Burgundy X Pink d r S r S S d
15 Purple X Pink d S r r S S d
16 Pixie self d r r r S S d
17 Rose X Salmon d r r r S S d
18 Cherry X Salmon d r r r d S S
19 Salmon X Orange d r r S d S d
20 Salmon X Red d r r r d S d

* S -segregating gene

1'

-recessive in both parents

d -dominant in both parents

Table 11 (cont'd.)

27

 

 

 

Cross Status of Genes *

Number cmss P v B E R L Iii't'
21 Salmon X Burgundy d r S r d S d
22 Salmon X Purple d S r r d S d
23 Cherry X Rose d r r r S d S
24 Rose X Orange d r r S S d d
25 Rose X Red d r r r S d d
26 Rose X Burgundy d r S r S d d
27 Rose X Purple d S r r S d d
28 Orange X Cherry d r r S d d S
29 Cherry X Red d r r r d d S
30 Cherry X Burgundy d r S r d d S
31 Cherry X Purple d S r r d d S
32 Orange X Red d r r S d d d
33 Orange X Burgundy d r S S d d d
34 Orange X Purple d S r S d d d
35 Burgundy X Red d r S r d d d
36 Purple X Red d S r r d d d
37 Burgundy X Purple d S S r d d d

* S -segregating gene

r -recessive in both parents

d -dominant in both parents

 

28

Line charts were developed in order to clearly demon-
strate the gene interactions. Figure 1 shows the interaction
of the R, L, and Int genes. The genes being considered are
listed horizontally across the t0p of the chart with the
color phenotype and H.C.C. number at the far right. The
chart works like a botanical key for each gene. There are
either two choices if the gene is dominant, or three choices
if it is incompletely dominant. To find the genotype of a
color, start at the t0p left corner with the first gene
Shown. In the case of Figure 1, there is R, or moving down-
ward recessive r.- If the genotype for the pink shriveled
phenotype is desired, then r would be selected. By following
the lines downward and across the chart, the genotype for
pink shriveled is found to be rrllint int. Only the haploid
genotypes are shown on the charts, but homozygosity is as-
sumed unless the gene is incompletely dominant, in which
case, the intermediate class is also given.

Figures 2, 3, and 4 Show the interaction of the E, B,
and V genes with R, L, and Int. The interactions of V with
B, V with E, and B with E are Shown in Figure 5.

 

 

 

 

 

 

 

 

Figure 1.

 

 

 

Genes

R L Int
int

1 Int

int

r L Int
int

1 Int

int

Color

Red

Cherry

Salmon

Pink

Rose

Rose Shriveled

Pink

Pink Shriveled

Interaction of the R, L, and Int Genes

H.C.C.#

18/0

oZO/l

620/1

21/2

oZO/l

oEO/l

21/2

21/2

30

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Genes
P E R L Int
int
1
r L
I
1
Es R L Int
int
1
r L
l
e
P

 

 

Figure 2.

 

 

Color

Orange

Dilute Cherry

Dilute Salmon

Dilute Rose

Dilute Pink

Light Red

Light Cherry

Light Salmon

Light Rose

Light Pink

Figure 1

All White

H.C.C.#

619/0

o621/2

520/3

0621/2

623/3

17/1

20/2

621/2

20/2

623/2

Interaction of the P, E, R, L, and Int Genes

 

 

 

 

 

 

 

 

31

 

 

 

 

 

L

 

L

 

t“

H

 

 

Int

int

Int

 

int

 

 

Genes
P B R
r
Bb -- 1R
r
b
p B

 

Figure 3.

 

 

 

 

 

 

 

Interaction of the P,

Color

Burgundy/red lip

Maroon/rose lip

Peach-Rose

Maroon/rose lip

Rose-Pink

Red Burgundy

Fuchsia

Peach-Rose

Fuchsia

Rose-Pink

Figure 1

Pink-White
White

B, R, L, and Int Genes

H.C.C.#

82¢}

824/3

25/2

824/3

27/2

820/0

22/ 1

25/2

22/ 1

27/2

427/3

32

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Genes
P V R L Int
int
1
r L
I
1
VV R L ' Int
int
1
r L
l
v
R V.

<—

Figure 4.

 

 

 

 

 

Color

Purple

Dusty Purple

Light Violet

Purple

Light Violet

Rose Purple

H.C.CO#

931/2

834/3

30/2

931/2

30/2

928/2

Dusty Rose Purple 033/0

Light Violet

Rose Purple

Light Violet

Figure 1
Purple-White

White

30/2

928/2

30/2

433/2

Interaction of the P, V, R, L, and Int Genes

 

 

 

 

 

 

Genes
V B
I
Eh
I
b e
I
Be
I
E
Vv B
I
Bb
I
b e
I
Ee
I
E
v B e
I
Be
I
E
Bb e
I
Ee
I
E
b e
I
Be
I
E

Figure 5.

33

Color

Purple
Light Purple
Dilute Purple

Rose Purple
Light Rose Purple
Dilute Purple

Burgundy
Light Burgundy
Dilute Burgundy

Red Burgundy
Light Red Burgundy
Dilute Burgundy

Red
Light Red

Orange

Interaction of the V, B, and E Genes

H.C.C.#

931/2
830/1
..31/1

928/2
..31/0
..31/1

82MB
722/0
25/2

820/0
20/0
23/2

18/0
17/1
619/0

DISCUSSION

The L and Int genes were discovered by Hendrychova-
Tomkova (7). Salmon (L gene) crossed with white, red, rose,
and purple produced data confirming Hendrychova-Tomkova's
(7) results. She also found that rose X purple segregated a
light violet color. The R gene did not segregate any light
violet shades when combined with V, but the Int gene did,
therefore her rose color must have been due to the Int gene.
Since she had no other source of rose color, She would not
have discovered the R gene.

Hendrychova-Tomkova (7) also discovered the pigment, P,
and violet, V, genes. She found these genes to be closely
linked, but in this study, the cross white X purple showed
no indication of linkage. Possibly in the twenty years
since her study, the linkage between the P and V genes has
been broken, at least in cultivated p0pulations. Crosses
of salmon, rose, and red with white and purple produced
data substantiating her results.

The R, B, and E genes are new to this study. The R and
Int genes both produce rose color and segregate identical
colors when combined with the B and E genes. Only the V
gene is affected differently. Recessive int int with V pro-

duces dusty purple, while recessive rr has no effect on the

34

35

V gene. The R and Int genes are deleterious when their
recessive alleles are combined. The shriveled flower stalk
phenotype is not lethal but effectively unreproductive.

The B gene can be considered a variegation gene since
it primarily affects the body color of the flower. The lip
color is the same as if the B gene were recessive. The R,
L, and Int genes determine the basic color of the flower
while the B gene adds a purple cast to the body. Hendrychova-
Tomkova (7) did a pigment analysis and found small amounts
of purple cyanidin in the red color. Theoretically, the B
gene could increase the cyanidin content and cause the pelar-
gonidin colors to be purple-tinted. Obviously, a pigment
study is needed to verify this hypothesis.

The E gene dilutes the colors produced by the other
genes. Since E is incompletely dominant, there are three
Shades of every color: intense color ee, light Shades Be,
and dilute shades EE. The E gene had limited effect on the
B gene. The light burgundy and light red burgundy colors
were not greatly different from their intense forms. A side
by side comparison was necessary to differentiate them. If
the E gene acts primarily on the pelargonidin pigment and
burgundy color is due to cyanidin, then this could explain
the limited dilution of B by E. Another possibility is
modifier genes. All of the incompletely dominant genes
(V, B, and E) were somewhat variable in their coloring,

particularly within the intermediate class. The colors were

36

always within their color chart designations, but minor
differences were apparent in the intensity of the colors.

All parents except rose, salmon, and pink carried domin-
ant R and L genes, and the other five genes were each found
from different colors. With this situation, only three genes
could be segregating in any single cross, and consequently,
all gene combinations were not recovered. For example, both
purple and burgundy parents carried dominant R, L, and Int
genes, and the B gene seemed to have no effect on the V gene
in the cross purple X burgundy. If B, however, was combined
with V and recessive 11 or int int, then the effect of B
might be seen. The orange (E gene) parent also carried
dominant R, L, and Int genes, therefore the effect of B and/
or V on the diluted colors was not seen. The potential for

new colors is very high.

SUMMARY

This study was undertaken to determine the number of
genes for flower color in Salvia splendens. The colors
were divided into eight distinct classes and intercrossed.
The resulting F2 and backcross populations for each cross
were grown and analyzed genetically. Seven genes were
identified and line charts were deve10ped to Show their

interaction.

R produces red with L; rr produces rose with L

L produces red with R; 11 produces salmon with R
Int produces red with R and L; int int produces rose
with R and L
produces colored flowers; pp produces white
produces purple; vv produces red with L and Int
produces burgundy; bb produces red with R, L, and Int
produces dilute colors; ee produces intense colors

mw<p

The R, L, Int, and P genes exhibit complete dominance while
the V, B, and E genes are incompletely dominant. The R, B,

and E genes were discovered in this study.

37

APPENDIX

APPENDIX

The data to support the conclusions in this thesis are
presented here. The order of the crosses is the same as in
Table 10. The F2 and backcross segregations are included
for each cross, and the non-segregating genes are also given.
In some cases, it was necessary to combine classes when the
color was not able to be differentiated. An * indicates the

Chi Square value has been calculated using Yates' Correction

for Continuity (15).

38

Sepgtype

P_RSD_
pp----
P_rrL_
P_R_ll
P_rrll

P_RrLl
P_rrLl
P_erl
P_rrll

df=3

P_R_L_

pp----

df=1

39

Cross 1

Pink X White

non-segregating genes - v, b, e, Int

Ratio

27
16
9

\O

Ruler. 9122:.
Red ' 159
White 77
Rose 43
Salmon 48
Pink _lg
341

.70>P> .50

(Pink X White) X Pink

Red 21
Rose 17
Salmon 16
Pink 15
7O

.90) P) .80

(Pink X White) X White

Red 12
White _1_8_
30

.30>P> .30

pgp,
144
85
48.
48
.16
341

17.5
17.5
17.5

1205
70.0

15
12
30

p312

1.362
.732
.320
.000

3.084

.700
.014
.128
,lgg
.970

.416

.832 *

ESEQEXRE

P L

pp—-
P_ll

P_Ll
P_ll

df::1

P L

PP"

df::1

40

Cross 2

White X Salmon

non-segregating genes - v, b, e, R, Int

Ratio

9
4

.3
16

d

9242.1: 04s..
'Red 156
White 63
Salmon _55
263
.70 >P > .50

(White X Salmon) X Salmon

Red 37
Salmon ' SS
65

.50>P> .30

(White X Salmon) X White

Red 29
White g3
52

.50) P > .30

E_xp_. 2310
148.0 .432
65.7 .110
.4903. .552
263.0 1.111
32.5 .492
32.2 .422
65-0 .984 *
26 .240
25 .249
52 .480 *

P R

PP"

P_rr

df:2

P Rr

P_rr

df:=1

P R

PP"

df::1

41

Cross 3

White X Rose

non-segregating genes - v, b, e, L, Int

Ratio

9
4

.3
16

9.9.1.9.: 012.4...
'Red 211
White 81
Rose .19.
366

.50>P>.30

(White X Rose) X Rose

Red 43
Rose ' 33
78

.50)P> .30

(White X Rose) X White

Red 38
White SS
64

.20>P>.10

Egg.
205.9

91-5

§8.§
366.0

39

78

32
3g
64

pi;
.126
1.204
.0425
1.755

.314

.628 *

.945

1.890 *

.92221122
P_Int__

PP"

P_int int

df

II
N

P_Int int

P_int int

df=1

P_In t_

PP"-

dle

42

Cross 4

White X Cherry

non-segregating genes - v, b, e, R, L

Ratio
9
4

.3
16

-.

M 20s...

'Red 105
White 41

Cherry _32

185

.70>P> .50

(White X Cherry) X Cherry

Red 39
Cherry ;5
64

.20>P>.10

(White X Cherry) X White

Red 22
White S3
45

.90>P) .80

002.2%
104.0 .009
46.3 .606
.3402 0232
185.0 1.147
32 1.320
3; 1. 20
64 2.640
22.5 .011
2203 0911
45.0 .022

1511
(D

*0 0
I I
F1

F1

PP--
P_Ee

P_ee

df

11
N

White X Orange

43

Cross 5

non-segregating genes - v, b, R, L, Int

Ratio

00100

16

d

Color

White
Red

'Light Red

Orange

Light

.90)P> .80

Red

Orange

White
Light
Red

.95 >P ) .90

Red

.975 >P> .95

Obs,

58
35
30
.25
148

(White X Orange) X Orange

33

34
67

(White X Orange) X White

02010220
55.50 .112
37.00 .108
27.75 .182
22025 0222
148.00 .674
33.5 .007
3305.. 0992
67.0 .014
23 .043
11.5 .021
1105. 0921
46.0 .085

44

Cross 6
White X Red
non-segregating genes - v, b, e, R, L, Int
_XP..Gen0t e 3041.0 9010.1: 900.. 0:20.. Pie.
P_ 3 Red 100 104.25 .134
pp _1_ White _32 35.22 33.19
4 139 139.00 .544 *

df=1 .50>P).3O

(White X Red) X White

Pp 1 Red 15 I5 .000
pp 1 White _12 12 _._9_Q_Q

30 30 .000
df = 1 P) .99

(White X Red) X Red

P 1 Red 30 30 --

45

Cross 7
White X Burgundy

non-segregating genes - v, e, R, L, Int

___1LGen0t e 302.10 9.9.10.1: 900... .1202. 2200
P_Bb 6 Red Burgundy 77 79.0 .050
P_bb 3 Red 48 39.6 1.781
P_BB 3 Burgundy. 37 39.6 .170
ppB_ 3 Pink-White 31 39.6 1.745
ppbb ._1 White .19. .1302. 10992
16 211 211.0 5.613
df:4 .30>P>.20
(White X Burgundy) X White
PpBb 1 Red Burgundy 20 17.75 .285
Ppbb 1 Red 20 17.75 .285
ppBb 1 Pink-White 15 17.75 .426
ppbb 1 White ;§_ 12,23 0122
71 71.00 1.168
df=3 .80>P>.W)
(White X Burgundy) X Burgundy
P_Bb 1 ' Red Burgundy 31 31.5 .007
P_BB I Burgundy 3S 3133 .00

63 63.0 .014
(if-:1 095>P>090

#6

Cross 8
White X Purple

non-segregating genes - b, e, R, L, Int

mu 1: e Rail). .922: 9.1%.. 829.. 13312.
P_VV 6 Rose Purple 115 110.6 .052
P_VV 5 Purple 62 55.5 .811
P_vv 5 Red 67 55-3 2.475
ppV_ 5 Purple-White 42 55.5 5.198
ppvv _1_ White _1_1 M £0.49

16 295 295-0 9.576
(if-:4 .05)P>.01

(White X Purple) X White

Pva 1 Rose Purple ‘ 18 16 .250
vav 1 Red 14 16 .250
pva 1 Purple-White 20 16 1.000
ppvv 1 White _1_; _15 M90

6h 64 2.500

df=3 .50>P>.30
(White X Purple) X Purple
P_VV 1 Rose Purple 58 51 1.562
P_VV 1 Purple 23 1; 1,562
62 62 2.720 *

#7

Cross 9
Pink X Salmon

non-segregating genes - v, b, e, L, Int, P

8.113.213 01.1.2; size. 1222. file
5 Salmon 76 69.75 .070
.1. Pink 12 wine;
4 93 95.00 1.896 *
.20)P).10

(Pink X Salmon) X Pink

1 Salmon 15 15.5 .016
1 Pink 5 11,2 ,016
51 51.0 .052

.90 > P ) .80

(Pink X Salmon) X Salmon

‘

Salmon 21 21 --

df::1

Ll
ll

df::1

48

Cross 10

Pink X Rose

non-segregating genes - v, b, e, R, Int, P

Ratio
3
1

4

QQLQE QQR.
Rose 149
Pink _5;
201
.90) P ) .80

(Pink X Rose) X Pink

Rose 11
Pink _2
20

.90)P>.80

(Pink X Rose) X Rose

Rose 33

emf/.2
130.7 .009
.29.3 .Qéfl
201.0 .037 *
10 .023
1.0. .922
20 .050 *

33

22222122
R_L_Int_

rrL_Int_ &
R_L_int int

R_llInt_

rrllInt_ &
R_llint int

rrL_int int

rrllint int

df

H
W

R_L_Int int
R_L_int int

df::1

RrLlInt_

rrLlInt_

erlInt_

rrllInt_

df

H
\N

non-segregating genes - v, b, e, P

Ratio

27

18

64

1+9

Cross 11

Pink X Cherry

Color Obs,
Red 44
Rose/Cherry 35
Salmon 27
Pink 15
Rose Shriveled 4
Pink Shriveled 5

128

.20>P).10

(Pink X Cherry) X Cherry

Red 20
Cherry 12
39

.90 )P ) .80

(Pink X Cherry) X Pink

Red 3
Rose 5
Salmon 4
Pink ._1

11

.70>P> .50

£22.

54

36
18

12

6
._2
128

19.5
12.5
39.0

2.75
2.75
2.75
g,z§

11.00

Dale

1.851
.027

4.500

.750

.666

8.294

.012
.012
.024

.022
.022
.568
1.1.11
1.725

Genotype

EeR_L_
eeR_L_
EER_L_
EerrL_
EeR_ll
eerrL_
eeR_ll
EErrL_
EER_ll
Eerrll
eerrll

EErrll

df==11

EeR_L_

EER_L_

df::1

50

Cross 12

Orange X Pink

non-segregating genes - v, b, Int, P

Egp. D219

Ratio

18

NW'WWKJJmmOO

d

64

1

9212: 922.
Light Red 65
Red 40
Orange 33
Light Rose 22
Light Salmon 21
Rose 15
Salmon ‘ 11
Dilute Rose 9
Dilute Salmon 12
Light Pink 6
Pink 5
Dilute Pink _1

240

.99 ) P > .975

(Orange X Pink) X Orange

Light Red 17
Orange 14
31

.80>P> .70

67.50
33.75
33.75
22.50
22.50
11.25
11.25
11.25
11.25

7.50

3.25

.092

1.157

.016
.011

1.250

.005
.450
.050
.500
.150

5,25 2120
240.00 5.751

1505 0061+
12.2. .2é4
51.0 .128 *

Genotype
EeRrLl

EerrLl
Eeerl
Eerrll
eeRrLl
eerrLl
eeerl

eerrll

df

I1
Q

R‘L_
rrL
RLll

rrll

df

II
\N

R L

51

Cross 12 (cont'd.)
(Orange X Pink) X Pink

82112 99.12.12 9.1.11. 13112.
1 Light Red 6 5
1 Light Rose 6 5
1 Light Salmon 3 5
1 Light Pink 4 5
1 Red 6 5
1 Rose 6 5
1 Salmon 5 5
.1. Pink .11 .2
8 110 110

p '> .995
Cross 15
Red X Pink

non-segregating genes - v, b, e, Int, P
9 Red 142 155
5 Rose 55 45
5 Salmon 33 45

_1_ Pink _12. .12
16 240 240
.10 > P > .05
(Red x Pink) x Red
1 Red 72 72

.05
.05
.00

[9

.75 *

.562
2.222
3.200
1.666
7.450

52

Cross 15 (cont'd.)
(Red X Pink) X Pink

__xn_Genot e m acne}: 9.11s... £151.. 113/.12
RrLl 1 Red 15 15 .000
rrLl 1 Rose 11 15 .507
erl 1 Salmon 12 15 .076
rrll 1 Pink 16 15 152;
52 52 1.075

df::5 .80>P).70

Cross 14
Burgundy X Pink
non-segregating genes - v, e, Int, P

BbR_L_ 18 Red Burgundy. ' 127 112.50 1.868
BBR_L_ 9 Burgundy 51 56.25 .490
bbR_L_ 9 Red 53 56.25 .187
B_R_ll 9 Peach-Rose 41 56.25 4.154
BbrrL_ 6 Fuchsia 42 57.50 .540
BBrrL_ 5 Maroon 25 18.75 .965
B_rrll 5 Rose-Pink 21 18.75 .270
bbrrL_ 5 Rose 9 18.75 5.070
bbR_ll 3’ Salmon 23 18.75 .963
bbrrll _1 Pink __]_0_ __§:_2_5 2.2 0
64 400 400.00 16.755

df=9 JO>P>JB

9.9221129.
BbRrLl

BbrrLl
Bberl
Bbrrll
bbRrLl
bbrrLl
bberl
bbrrll

df

11
fl

BbR_L_
BBR_L_

df::1

Ratio

1

1
1

53

Cross 14 (cont'd.)
(Burgundy X Pink) X Pink

Color . Obs.

 

Red Burgundy
Fuchsia
Peach-Rose
Rose-Pink
Red

Rose

Salmon

Pink

5L.»
O \J'IO\\]\N\‘1-P'\n

.90>P> .80

(Burgundy X Pink) X Burgundy

Red Burgundy 25
Burgundy g3
48

.90)P >.80

$~ Ln Pd
<3 U1 \n \m U1 \n \n \fi x
O

g3
48

o o o o o o 0 UN
0 N (I) (I) (I) N O (0

IE»

3.6

.0104

,0104
.0208 *

921.121.29.12

Vv--L_
VV--L_
V_r-ll
vvR_L_
vvrrL_
vvR_ll

vvrrll

11
ON

df

Vv--L_
Vv--ll
verLl
vvrrLl
verll

vvrrll

df

11
U1

54

Cross 15

Purple X Pink

non-segregating genes - b, e, Int, P

Ratio

24
12
12

9
5
5

_1_

64

92122 .222.
Rose Purple 79
Purple 53
Light Violet 42
Red 41
Rose 11
Salmon 7
Pink .__2
240
.20>P>.10

(Purple X Pink) X Pink

Rose Purple 11
Light Violet 8
Red 7
Rose 8
Salmon 5
Pink ._5

42

.70>P> .50

21.112.
90
45
45

33.75
11.25

11.25

240.00

10.50
10.50
5.25
5.25
5.25

42.00

93,4;
1.344
1.422
.200
1.557
.005
1.605

8.949

.025
.595
.583
1.440
.964

,012

3.617

.QEBQEIEE
VvR_L_
VVR_L_

df::1

55

Cross 15 (cont'd.)
(Purple X Pink) X Purple

2111i. 2.1.2:. 9.1.1...
1 Rose Purple 25
1 Purple 25
' 48

.90>P> .80

£591.
24
214
48

Genotype

R_L_
rrL_
R_ll

rrll

RrL

rrL_

df

d

R_Ll
R_ll

df:1

56

Cross 16

Pixie self (Rose X Salmon)

non-segregating genes - v, b, 6, Int, P

Ratio

9

.2212; .222.
Red 154
Rose 40
Salmon 40
Pink _1§
250
.90>P > .80

(Rose X Salmon) X Rose

Red 17
Rose 15
52

.90 >P > .80

(Rose X Salmon) X Salmon

Red 12
Salmon 11_
23

.90 >P >.80

2122.11.21.
129.4 .165
45.1 .222
43.1 .222
..12.4 ..122
230.0 .784
16 .015
12 .215
32 .030
11.5 .021
11.5. .221
23.0 .042

57

Cross 17
Rose X Salmon

non-segregating genes - v, b, e, Int, P

me e 22.1.1.0. 921.02 9.1.12. 222.. 2.21.
R_L_ 9 Red 132 135. 6 .095
rrL_ 5 Rose 59 45.2 .850
R_ll 5 Salmon 54 45.2 1.715
rrll _1_ Pink _1_§ __1_5_,_o ..226

16 241 241.0 2.724
df::3 .50>P>.3O

(Rose X Salmon) X Rose
RrL_ 1 Red 25 ' 23 .097
rrL_ 1 Rose _2_1_ 25 1991
46 46 .194 *
df::1 .70>P>.50

(Rose X Salmon) X Salmon

R_Ll 1 Red 50 25 .810
R_ll 1 Salmon 20 25 .810
50 50 1.620 *

df:1 .50>P>.20

58

Cross 18
Cherry X Salmon

non-segregating genes - v, b, e, R, P

Manet e 22.1.1.9 29.1.02 91.12. .422. 22.42
L_Int_ 9 Red 155 155.51 .021
L_int int 5 Cherry 44 44.44 .004
llInt_ 5 Salmon . 45 44.44 .007
llint int ._1_ Pink __15 14,51 1.521
16 237 237-00 .253
df=3 .975>P>.95
(Cherry X Salmon) X Salmon
LlInt_ 1 Red 1 1 12 .020
llInt_ 1 Salmon 15 1g, ,ogo
24 24 .040
dfz1 . .90>P>.80
(Cherry X Salmon) X Cherry
L_Int int 1 Red 52 52.5 .007
L_int int 1 Cherry 55 5§15, 1992

65 65.0 .014
df::1 .95>P>.9O

Ge 0
EeL_
eeL_
EEL

Eell
eell
EEll

df=5

EeLl

eeLl

Eell

eell

df:5

EeL

EEL

df::1

e

59

Cross 19

Salmon X Orange

non-segregating genes - v, b, Int, R, P

Ratio

6

NKNKN

d

291% 222..
Light Red 111
Red 61
Orange 58
Light Salmon 35
Salmon 15
Dilute Salmon __5
288

.20>P>.10

(Salmon X Orange) X Salmon

Light Red 14
Red 10
Light Salmon 8
Salmon 1Q

42

.70>P>.50

(Salmon X Orange) X Orange

Light Red 20
Orange 12
39

.90 >P>.80

222.

108
54
54
36
18

.12

288

1005
10.5
10.5

10,5
42.0

19.5
12.2
39.0

1.166
.023
.595

,025
1.807

.012

,012

.024

60

Cross 20
Salmon X Red

non-segregating genes - v, b, e, R, Int, P

_meenot e 2212.19. 29.1.2; 212. 2.1.1.. .1231.
L. 3 Red 168 169.5 .005
ll _1_ Salmon _55 _5§_,5 £11

4 226 226.0 .022 1
df::1 .90>P>.80

(Salmon X Red) X Salmon

Ll 1 Red 37 35.5 .028
ll 1 Salmon 54 5515, .02

71 71.0 .056 *
df=1 .90>P>.80

(Salmon X Red) X Red

L 1 Red 47 47 --

61

Cross 21
Salmon X Burgundy

non-segregating genes - v, e, R, Int, P

_meenot e 2211.112 92.1.9.1; 2122. .1211. Die
BbL_ 6 Red Burgundy 68 75 .653
BBL_ 5 Burgundy 41 57.5 .526
bbL_ 5 Red 56 57.5 .060
B_ll 5 Peach-Rose 40 57.5 .166
bbll _1_ Salmon _1_5 __1_2_,_5 1E
16 200 200.0 1.705
df:_4 .80>P>.7O I
(Salmon X Burgundy) X Salmon
BbLl 1 Red Burgundy 12 11 .090
bbLl 1 Red 1 1 1 1 .000
Bbll 1 Peach-Rose 10 11 ,.090
bbll 1 Salmon 1_1_ 1_1_ ,999
44 44 .180
df=5 .99>P>.975
(Salmon X Burgundy) X Burgundy
BbL_ 1 Red Burgundy 15 14 .017
BED_ 1 Burgundy 15 14 .4132

28 28 .054 *

Genotype

VVL_
VVL_
V_ll
VVL_

vvll

df::4

Vle

Vvll

val
vvll

VVL_
VVL_

df::1

Ratio

6
3
3
3
.l
16

d

62

Cross 22

Salmon X Purple

2329.1: 922..
Rose Purple 104
Purple 59
Light Violet 60
Red 53
Salmon _1;
288
.70>P>.50

(Salmon X Purple) X Salmon

Rose Purple 10
Light Violet 9
Red 10
Salmon _2
38

.99 >P >.95

(Salmon X Purple) X Purple

Rose Purple 25
Purple go
45

.70>P >.50

. non-segregating genes - b, e, R, Int, P

222.. .2342
108 .148
54 .462
54 .666
54 .018
._18 2.929
288 3.294
9.5 .026
9.5 .026
9.5 .026
3.5.2.22
38.0 .104
22.5 .177
22.5. .122
45.0 .354 1

22222122
R_Int_

rrInt_ &
R_int int

rrint int

II
N

df

ernt_

rrlnt_

df::1

R_Int int

R_int int

df::1

63

Cross 25

Cherry X Rose

non-segregating genes - v, b, e, L, P

Ratio

9

Color Obs.
Red 129
Rose/Cherry 94
Rose Shriveled 15

238

(Cherry X Rose) X Rose

Red ~ - 19
Rose 15
52

.50 > P > .30

(Cherry X Rose) X Cherry

Red 36
Cherry 5;
68

.80)P >.70

Eyp, Daze
133.9 .179
89.2 .258
14,9 .001
258.0 .438
16 .390
.12 . 390
52 .780 *
34 .066
54 -066
68 .152 *

212211.122

EeR_
eeR_
EER_
Eerr
eerr

EErr

df”

H
U1

EeRr
eeRr
Eerr

eerr

df

H
\N

EeR
BER

df=1

64

Cross 24

Rose X Orange

non-segregating genes - v, b, L, Int, P

Ratio

6

3
3
2

ad

16

dd

Color

Light
Red
Orang
Light
Rose

Dilut

Red

e

Rose

e Rose

.20>P>.10

Q11_s_,
92
64
49
32

(Rose X Orange) X Rose

Light
Red
Light

Rose

Light

Orang

Red

Rose

.90>P >.8O

Red

.70>P>.50

C) (D \O '0

(Rose X Orange) X Orange

21
12
38

.Egpo
98.2
49.1
49.1
32.8
16.4
15,4
262.0

8.5
8.5
8.5

34.0

19
12.
38

.264
.029
.029
,_2_§_4
.586

.118

.256 *

65

Cross 25
Rose X Red
non-segregating genes - v, b, e, L, Int, P
W 22.1.1.0 2.2.12.1: .122... 2.12. 2542
R_ 5 Red 225 216 .195
rr _1_ Rose £5 _2; 15%
4 288 288 .781 *

df =1 .50>P >.30

(Rose X Red) X Rose

Rr 1 Red 18 18 .000

rr 1 Rose 18 15 ,000
' 36 36 .000

df::1 P>099

(Rose X Red) X Red

R 1 Red 58 58 --

222012112

BbR_
BBR_
bbR_
BBrr
Bbrr

bbrr

df:5

BbRr

bbRr

Bbrr

bbrr

df:5

BbR

BBR

df::1

66

Cross 26
Rose X Burgundy

non-segregating genes - v, e, L, Int, P
mfia
112.50 1.175
56.25 1.521

32222
6
3
3

29.121: 222.
Red Burgundy 124
Burgundy 47
Red 51
Maroon 16
Fuchsia 41
Rose _2_1_

500

.70 >P >.50

(Rose X Burgundy) X Rose

Red Burgundy 6
Red 6
Fuchsia 10
Rose _5
28

.70>P>.5O

(Rose X Burgundy) X Burgundy

Red Burgundy 16
Burgundy 1g
52

P>.99

56.25
18.75
37.50

.490
.405
.526

1§,25 ,220
500.00 4.185

gkflflfl

12
32

011-42
.142
1.285

1.711

.000
,000
.000

Genotype

Vv--
vv--
vvR_

vvrr

df

H
\N

Vv--
ver

vvrr

df

H
h)

Vv—-

VV--

df::1

67

Cross 27
Rose X Purple

non-segregating genes - b, e, L, Int, P

Ratio

Ia 01 :- c»

16

QQLQE. QEE;
Rose Purple 139
Purple 96
Red 50
Rose _;g
309
.05>P>.01

(Rose X Purple) X Rose

Rose Purple 20
Red
Rose 19
35
.50>P> .30

(Rose X Purple) X Purple

Rose Purple 58
Purple 5;
80

.80>P> .70

EEE;
154.5
77.3
57.9
.1223
309.0

17.50
8.75

55.00

40

A9
80

gig

1.555
4.523
1.077
12134
8.299

.357
1.607

2.142

.056
.926

.112 *

W _iLRa 0

EeInt_

eeInt_

EEInt_

Eeint
eeint

EEint
df::5
EeInt
eeInt

Eeint

eeint

df

11
\N

int
int

int

int
int
int
int

EeInt_

EEInt_

df::1

68

Cross 28
Orange X Cherry

non-segregating genes - v, b, R, L, P

6

NKNKN

16

992-9.; £02..
Light Red 113
Red 1 54
Orange 48
Light Cherry 28
Cherry 17
Dilute Cherry _1_l+_
274

(Orange X Cherry) X Cherry

Light Red 12
Red 12
Light Cherry 9
Cherry _1_;
44

. 95 > P > . 90

(Orange X Cherry) X Orange

Light Red 15
Orange 15
28

.90 >P> .80

51.58
51.38

E22. .2342
102.75 1.022

.133
.222

54.25 1.140

17.12

m
1.4
28

.568

12,12 ,001
274.00 3.086

.090
.090
.363
,999
.543

.017

,012
.034 *

69

Cross 29
Cherry X Red

non-segregating genes - v, b, e, R, L, P

men t e 32.22 22.142 gm. 2m 2342
Int_ 3 Red 185 188.25 .040
int int 1_ Cherry _§§, _62‘25, 4129

4 251 251.00 .160 *
df::1 .70>P).50

(Cherry X Red) X Cherry

Int int 1 Red 55 51.5 .285

int int 1 Cherry 28 51,5 ,285
63 6300 0570 *
df=1 050>P>030

(Cherry X Red) X Red

Int 1 Red 54 54 --

70

Cross 50
Cherry X Burgundy

non-segregating genes - v, e, R, L, P

m 32.2.2 20.1.2; 948.. 21.6.. 2242
BbInt_ 6 Red Burgundy 85 79.12 .456
BBInt_ 5 Burgundy 35 39.56 .525
bent_ 5 Red 58 59.56 .061
Bbint int 2 Fuchsia 55 26.58 .771
BBint int 1 Maroon 10 15.19 1.661
bbint int _1 Cherry . , _1_9 _1_1,_1_9 .4211

' 16 211 211.00 4.225
df:5 .70>P>.50

(Cherry X Burgundy) X Cherry

BbInt int 1 Red Burgundy 11 9 .444

Bbint int 1 Fuchsia 7 9 .444

bent int 1 Red 12 9 1.000

bbint int 1 Cherry _9 ‘_9’ 19999
36 56 2.888

df::5 .50>P>.50

(Cherry x Burgundy) x Burgundy-
BbInt_ 1 Red Burgundy 15 15 .175
BBInt_ 1 Burgundy ll _11 4.1.2}

26 26 .546 *
df=1 .70>P>050

gengtzpe
VvInt_

VVInt_
vant_
Vvint int
VVint int
vvint int

df

11
U1

VvInt int
Vvint int
vant int
vvint int

df

11
\N

VvInt_
VVInt_

df::1

Cherry X Purple

71

Cross 51

non-segregating genes - b, e, R, L, P

gatig legr

Rose Purple

Nkflkflm

16

d

Purple
Red

Dusty Rose Purple

Dusty Purple

Cherry

.99 >P > .975

QEE;
75
36
34
22

(Cherry X Purple) X Cherry

Rose Purple

Dusty Rose Purple

Red
Cherry

.95 >P ) ~90

6
8

29

, (Cherry X Purple) X Purple

Rose Purple

Purple

.80)P>.7O

Egg;
72
36
36
24
12

_12,

192

7.25.

7.25
7.25
.4222:
29.00

23.5
2125
47.0

9213
.125
.000
.111
.167
.333
rQflfi
.819

.215
.077
.008
,gzz
.377

.042

.042
.084 *

72

Cross 52
Orange X Red

non-segregating genes - v, b, R, L, Int, P

9.929.002 32212 2242: are. Em 13.212
Be 2 Light Red 137 134.50 .008
ee 1 Red 68 67.25 .046
BE ‘1 Orange _94 _52995, 9152

4 269 269.00 .211
df=2 .90>P>.80

(Orange X Red) X Orange

Be 1 Light Red 28 29 .008
BE 1 Orange ‘ 59 £2 9999

58 58 .016 *
df=1 095>P>090

(Orange X Red) X Red
Be 1 Light Red 16 14.5 .068
ee 1 Red 15 1_4_,_5 9969
29 29.0 .156 *
df=1 .80>P>.70

73

Cross 55
Orange X Burgundy

non-segregating genes - v, R, L, Int, P

Ge e 222212. 211.02 93.6....
BbEe 4 Light Red Burgundy 75
BBEe 2 Light Burgundy 25
Bbee 2 Red Burgundy 56
bbEe 2 Light Red 56
B_EE 5 Dilute Burgundy 59
BBee 1 Burgundy 16
bbee 1 Red 24
bbEE _l .Orange _19
16 283
df::7 .50>P>.30
(Orange X Burgundy) X Orange
BbEe 1 Light Red Burgundy 8
BbEE 1 Dilute Burgundy 8
bbEe 1 Light Red 14
bbEE 1 Orange l9
4O
df=5 .50>P>.50

Egg,
71.0
35.3
33.3
33.3
33.0
17.7
17.7

41.2

285.0

9312
.056

3.005
.013
.013
.679
.165

2.242

6.944

.400
.400
1.600

.000

2.400

74

Cross 55 (cont'd.)
(Orange X Burgundy) X Burgundy

Metemm 932.31.231.12
BbEe 1 Light Red Burgundy 14 15 .507
BBEe 1 Light Burgundy 11 15 .076
Bbee 1 Red Burgundy 15 15 .076
BBee 1 Burgundy 1;} 15’ 9192

52 52 .766
df::5 .90 >P>.80

Cross 54
Orange X Purple

non-segregating genes - b, R, L, Int, P

We t e 221.12 92.24.: 9.48.. 3.16.. 221.9
VvEe 4 Light Rose Purple 60 55.0 .454
VVEe 2 Light Purple 50 27.5 .227
Vvee 2 Rose Purple 25 27.5 .227
vae 2 Light Red 22 27.5 1.100
[_EE 3 Dilute Purple 37 41.25 .437
VVee 1 Purple 16 15.75 .568
vvee 1 Red 14 15.75 .004
vaE _1 Orange _1_6 .1112: _‘569

16 220 220.00 5.185
df:7 .90>P>.80

geggtype

VvEe
VvEE
vae

vaE

df

VvEe

VVEe

Vvee

VVee

df

11
\N

11
\N

Ratio

1
1

1

75

Cross 54 (cont'd.)

(Orange X Purple) X Orange

92l2£

Light Rose Purple
Dilute Purple
Light Red

Orange

.80>P > .70

Obs,
14
14

(Orange X Purple) X Purple

Light Rose Purple
Light Purple
Rose Purple

Purple

.99 >P > .975

12
11

324.222
12.25 .250
12.25 .250
12.25 .127
2.25.413
49.00 1.040
11 .090
11 .000
11 .000
.ll .929
44 .180

Qeggtxpe
Bb
BB
bb
df=2
Bb
bb
df::1
Bb

BB
df::1

76

Cross 55

Burgundy X Red

non-segregating genes - v, e, R, L, Int, P

38.12.19.
2
1
_1_
4

2949.2 .046...
Red Burgundy 112
Burgundy 56
Red _99
228
.90 >P > .80

(Burgundy X Red) X Red

Red Burgundy 17
Red I lg
52

.90>P > .80

(Burgundy X Red) X Burgundy

Red Burgundy 12
Burgundy 11
25

.90 >P >.80

E_xa. 9.242
114 .055
37 .017
.22 .122
228 .209
16 .015
la .21.:
52 .050 *
12.5 .020
.12.: .29
25.0 .040

Gengtype
Vv

VV

VV

df=2

Vv

VV

df=1

Vv

df::1

77

Cross 56
Purple X Red

non-segregating genes - b, e, R, L, Int, P

Ratio

2
1
1
4

5291—02 9.48..
Rose Purple 84
Purple 45
Red _9;
171
.95 >P > .90

(Purple X Red) X Red

Rose Purple 42
Red 59
8O

.80 >P >.7O

(Purple X Red) X Purple

Rose Purple 21
Purple 19
4O

.90) P > .80

32.221.
85.50 .026
42.75 .118

32.25.94:
171.00 .157
40 .056
49 .926
80 .112 *
20 .012
29. .942
40

.024 *

Genotype

Vv--
vv.-
vab
vaB
vvbb

df

1|
:-

VvB_
vab
vaB

df

II
N

Vv--

df::1

78

Cross 57
Burgundy X Purple

non-segregating genes - e, R, L, Int, P

9221.9 99.1.9.1; 0_b.§..
8 Rose Purple 162
4 Purple 92
2 Red Burgundy 45
1 Burgundy 23
_1_ Red ._92
16 337
.80>P >.70

d

(Burgundy X Purple) X Burgundy

Rose Purple 19
Red Burgundy 9
Burgundy 19
38

.99>P >.95

(Burgundy X Purple) X Purple

Rose Purple 18
Purple 19
36

P>.99

Egg;

168.5
84.2
42.1
21.1

21,1

337.0

19.0
9.5

.2.2
58.0

18
1g,
36

5342
.250
.722
.019
.171

..229

1.958

, .000

.026

3029
.052

.000

,000

.000

BIBLIOGRAPHY

1.

2.

5.

4.

5.
6.

7.

8.

9.

10.

11.

BIBLIOGRAPHY

Bailey, Liberty Hyde, and Ethyl Zoe Bailey. Hortus
Iggrd. Macmillan Publ. Co., New York, 1976, p. 1000.

Bhattacharya, S. 1978. Indian members of the genus
Salvia. Cytol. 45:517-524.

British Colour Council. Horticultural Colour Chg; .
Volumes I and II. H. Stone and Son, Banbury, 1958 and
1941.

D'Cruz, Rui and A.S. Jadhav. 1965. Flower colour in
Salvia. Sci. and Cult. 51:201.

Ewart, L.C. 1985. Personal communication.

Gairdner, A.E. 1956. The inheritance of factors in
Cheiranthus cheiri. Jour. Gen. 52:479-486.

Hendrychova-Tomkova, Jarmila. 1964. Genetic analysis
of colour mutants in Salvia splendens. Preslia (Praha)
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