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This is to certify that the

thesis entitled

PATTERNS OF STAMINATE FLOWER EXPRESSION
IN HERMAPHRODITIC PICKLING CUCUMBER

INDUCED BY SILVER NITRATE
presented by

Mary Holt Hunsperger

has been accepted towards fulfillment
of the requirements for

 

 

M.S. Jegreein Horticulture

Jam 4444

Major professor

Date Ml—

0-7539

 

HERARY
mewgan state
University

*—

 

OVERDUE FINES:
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RETURNING LIBRARY MATERIALS:

Place in book return rauove
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PATTERNS OF STAMINATE FLOWER EXPRESSION IN

HERMAPHRODITIC PICKLING CUCUMBER INDUCED BY SILVER NITRATE

By

Mary Holt Hunsperger

A THESIS

Submitted to
Michigan State University
in partial fulfillment of the requirements

for the degree of

MASTER OF SCIENCE

Department of Horticulture

1981

ABSTRACT

PATTERNS OF STAMINATE FLOWER EXPRESSION IN

HERMAPHRODITIC PICKLING CUCUMBER INDUCED BY SILVER NITRATE

BY

Mary Holt Hunsperger

A series of experiments was conducted to examine the effect of
silver nitrate (AgNOB) on conversion to staminate flowering in two
hermaphroditic pickling cucumber lines used in the production of

gynoecious x hermaphrodite F hybrids. The stage of plant development

1
at which treatments were initiated strongly influenced the effectiveness
of both silver concentration and application number upon the responses
investigated. Based upon days to conversion, duration of conversion,
and total number of staminate flowers, the optimal conversion for hybrid
seed production was achieved by three applications of 200 mg/l AgNO3 at

four day intervals beginning at the first true leaf stage. Evidence for

supernumerary bud development is presented.

ACKNOWLEDGMENTS

I would like to thank Dr. Larry Baker for suggesting the research
problem and for his guidance through the initial stages during his
time at Michigan State University.

My sincere appreciation is extended to Dr. Diana Helsel, not only
for her help and encouragement during the writing of this manuscript,
but also for her professional example.

I would like to thank Dr. Wayne Adams and Dr. Jon Fobes for
serving on my committee and for their review of this manuscript.

I also appreciate all those who helped me throughout the course
of my studies, especially Dr. John Gill for his excellent instruction
in statistics and his help in the design of Experiment III.

A special thank you is extended to my fellow students, Anand
Nandgoankar and Neil Cowen, for their assistance in the execution
of my research.

Above all I am indebted to my husband, John, for encouraging me
to pursue my education and seeing me through with counsel and

constructive criticism.

ii

To the Reader:

This thesis is prepared in journal-style format.

iii

TABLE OF CONTENTS

List of Tables . . . . . . . . . . . . . . .
List of Figures . . . . . . . . . . . . .
Introduction . . . . . . . . . . . . .
Materials and Methods . . . . . . . . . . .
Results and Discussion . . . . . . . . . . .

Experiments I and II . . . . . . . . .

Experiment III .

List of References

Appendix A .
Appendix B .

Appendix C .

iv

23
24

25

LIST OF TABLES

Summary of treatment factors and experimental designs
used to assess the efficiency of staminate flower

induction in hermaphroditic pickling cucumber . . . . .

Average number of staminate nodes per plant as
influenced by silver nitrate concentration and
application number in hermaphroditic pickling
cucumber line MSU 7152 H. Data from Experiments I
and II . . . . . . . . . . . . . . . . . . . . . . .

Means for proportion of staminate to total number of
flowers per plant ( averaged over concentration and
application number) resulting from AgNO application
to two hermaphroditic cucumber lines. ata from
Experiment III . . . . . . . . . . . . . . . . . . . .

Means for total number of flowers per plant (averaged
over stage and application number) resulting from

AgNO application to two hermaphroditic cucumber lines.
Data from Experiment III . . . . . . . . . . . . . . .

18

18

LIST OF FIGURES

Mean number of staminate flowers per plant resulting
from AgNO applications on hermaphroditic pickling
cucumber Tine MSU 7152K. Data from Experiment I . . .

Mean weekly per plant totals of staminate flowers
resulting from AgNO applications on hermaphroditic
pickling cucumber line MSU 7152B. Results from
Experiment II . . . . . . . . . . . . . . . . . . .

Mean number of staminate flowers per plant resulting
from AgNO applications on hermaphroditic pickling
cucumber Iines MSU 7152K and MSU 669R. Data from
Experiment III . . . . . . ... . . . . . . . . .

Duration of staminate conversion as influenced by
genotype and plant developmental stage at first
application. Data from Experiment III on staminate
flower induction by AgNO on hermaphroditic cucumber
lines MSU 7152H and MSU S69H . . . . . . . . . . .

Mean duration of conversion to staminate flowering
(averaged over genotype) resulting from AgNO
applications on hermaphroditic cucumber line;
MSU 7172B and MSU 669E. Data from Experiment III .

vi

10

. l3

l6

. l9

INTRODUCTION

Single-cross hybrids involving gynoecious (G) x hermaphroditic (H)
parents have been used to maximize pistillate expression in hybrids used
for pickling cucumber (Cucumis sativus L.) production (7,10). Alterna—
tively, an acceptable level of female expression occurs in three—way
crosses utilizing (G x H) F1 seed parents and monoecious (M) pollen
parents (9). Both hybrid schemes fall short of their potential seed
yield in the G x H cross as well as in H stock seed increases. Although
the exact basis has not been determined, we speculate that the floral
structure of the hermaphrodite inhibits bee visitation or the efficiency
of pollen transfer. Conversion of the H line to staminate flowering
in G x H seed production would equal the effectiveness of the M phenotype
as a pollen parent, yet retain the greater contribution of the H geno-
type to female flower expression in the resulting F1.

Silver nitrate (AgNOB) has been successfully used on G lines to
induce staminate flowering for stock seed increases (4,6,12). We con-
ducted several preliminary experiments to determine an effective, non-
toxic concentration range and application number for staminate flower
induction in the hermaphrodite based on developmental stages recommended
for G lines. A more comprehensive experiment was then performed to
evaluate the effect of developmental stage on concentration and appli-

cation number in two unrelated H lines.

MATERIALS AND METHODS

Three experiments were conducted in the greenhouse during the
winter and spring of 1980. Each experiment utilized the H line
MSU 7152H. A second H line, MSU 669R, was included in Experiment III

to evaluate the effect of AgNO on conversion in different genetic

3
backgrounds.' Plants were spaced 30 cm by 80 cm and trained to upright
supports in raised benches containing an artificial soil medium (VSP
Peat Lite Mix, Michigan Peat products, Co., Houston, TX). Temperatures
were maintained at 25° 112° C in the day and 15° :;2° C at night. Due to
the low winter light levels prevailing during Experiments I and II,
supplemental cool white fluorescent lighting was provided for 16 hours
each day.

Treatment factors and experimental designs are presented in Table
1. Plant developmental stages were defined in the following manner:
stage 0 was attained when the cotyledons were fully expanded, while
stages 1,2,3, and 4 were reached when the first through fourth true
leaves, respectively, had attained a 4 cm.diameter. Silver nitrate was
applied as a foliar spray to the entire plant until run-off. To avoid
the possibility of toxicity due to surfactants, none was included. First
applications were made when plants had grown to the desired developmental
stage. Subsequent applications were made when each succeeding leaf had

attained a 4 cm size; this correSponded to an interval of approximately

four days for the greenhouse conditions in these experiments. The

2

3

Table 1. Summary of treatment factors and experimental designs used
to assess the efficiency of staminate flower induction in
hermaphroditic pickling cucumber.

 

 

Treatment Experiment
Factors
I II III
1-14-80z 2-4-80 4-7-80
developmental 4 3 0
stage 1
at first ' 2
applicationy 3
AgNO3 0 O 0
concentration 75 75 100
(mg/l) 150 150 200
300 300 300
400
number of 1 l 1
applications 2 2 2
3 3 3
4
lines MSU 7152H MSU 7152H MSU 7152B
MSU 669H
experimental randomized randomized randomized
design complete complete incomplete
block block block
replications 5 6 2

 

z - date of sowing

y - stage 0 is defined as full cotyledonary expansion.
stages 1 through 4 are defined as the lst through 4th true
leaves at 4 cm in diameter;

plastochron was chosen as the interval between applications since it is
directly related to plant development and can be extrapolated to field
growing conditions.

For Experiment I the number of staminate flowers occuring at each
node on the main stem.was recorded for the entire four week flowering
period. Additionally, the height of each plant was determined at the
end of the flowering period. An analysis of variance was computed for
the number of nodes bearing one or more staminate flowers, total number
of staminate flowers, and plant height (Appendix A). For Experiment II
the date of anthesis for each staminate flower was also recorded. An
analysis of varaince was calculated for the number of nodes bearing
staminate flowers, the total number of staminate flowers during each of
the four weeks of flowering, the cumulative staminate flower total, and
plant height(Appendix B).

In Experiment III the first three weeks of flowering were evalu-
ated. The date of anthesis, nodal position, and sex type were recorded
for each flower on the main stem as well as for each staminate flower on
the laterals. An analysis of variance was calculated for the following
responses: (1) days to first flower, (2) days to first staminate flower,
(3) duration of staminate flowering, (4) per cent nodes bearing staminate
flowers, (5) total number of staminate flowers, (6) hermaphroditic
flowers on the main stem, (7) total number of flowers on the main stem,

and (8) per cent staminate flowers on the main stem (Appendix C).

RESULTS AND DISCUSSION

Experiments I and II

 

The average number of nodes hearing at least one staminate
flower varied from 0.2 in the control to 13.5 when 300 mg/l of AgNO3
was applied (Table 2). There was no detectible interaction between
AgNO3 concentration and application number on staminate node frequency
in either experiment (Appendix A). The number of staminate nodes was
more strongly influenced by increasing the silver concentration than by
increasing the application number.

The total number of staminate flowers per plant was also examined
since our H lines bear a succession of flowers at each node. Signifi-
cant interaction for this character appeared in both experiments: hence,
comparisons were made between levels of one factor within each level of
the other factor. Analysis of Experiment I indicated that 75 mg/l did
not differ significantly from the control (Figure 1). Totals for the
control reflect the tendency of MSU 7152H to produce a few early
staminate flowers. No gain in staminate flower number was realized by
increasing concentration from 150 to 300 mg/l when a single application
was made. With two applications, however, the increase in staminate
flowers was substantial. Increasing the application number to three
failed to increase staminate flower production, a result possibly due

to the approaching termination of vine growth. Results of Experiment II,

Table 2. Average number of staminate nodes per plant as influenced
by silver nitrate concentration and application number in
hermaphroditic pickling cucumber line MSU 7153H. Data from
Experiments I and II.

Number of Staminate
Nodes per Plant

 

 

 

 

 

 

Treatment

Factors Level Experiment I Experiment II

AgNO3 o 0.9 0.2

(mg/1)y 75 5.5 6.8

150 10.3 9.3

300 13.5 11.1

MSDz 2.8 1.6

number of l 6.3 5.3

applicationsx 2 8.4 6.9

3 8.1 8.4

MSD2 2.1 2.1

 

 

z - Scheffe's MSD (52 level)
y - averaged over all application frequencies
x - averaged over all AgNO3 concentrations

Figure 1. Mean number of staminate flowers per plant resulting from AgNO3
applications on hermaphroditic pickling cucumber line MSU 7152H.

Data from Experiment I.

 

2 applications

 

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in which applications were made at stage 3, corresponded well with those
of Experiment I.

Examination of the weekly totals for Experiment II revealed that
conversion to staminate flowering did not begin until the second week of
flowering (Figure 2). Plants treated with a single application demon-
strated increasing numbers of staminate flowers with increasing concen-
tration. All other application frequencies were indistinguishable,
regardless of the AgNO3 concentrations used. The peak of flowering
occured during the third week at which time the greatest treatment
differences were seen. Maximum numbers of staminate flowers were induc-
ed by three treatment combinations: three 150 mg/l applications, two
300 mg/l applications, or three 300 mg/l applications. Plants treated
once or twice with 75 mg/l or once with 150 mg/l were reverting to the
hermaphroditic sex type by the third week. This evidence suggests that

higher AgNO concentrations remain active longer in the plant. By the

3
fourth week of flowering, conversion had ended for all treatment combin-
ations with the exception of plants treated with three 300 mg/l applica-
tions.

Serious AgNO toxicity on G lines has been noted by other research-

3
ers when applications were made before stages 3 or 4 (4,12). Under the
winter conditions of Experiment I moderate, but not excessive, necrosis
and crinkling of treated leaves was observed. The only treatment combin-
ation that resulted in reduced plant size was that of three 300 mg/l
applications. Stunting was also accompanied by a reduction of staminate
nodes; however, the total number of staminate flowers was not unduly

affected (Table 2 and Figure 1). No stunting and only mild necrosis

were noted under the more favorable environmental conditions of

10

Figure 2. Mean weekly per plant totals of staminate flowers resulting

from AgNO applications on hermaphroditic pickling cucumber

3
line MSU 7152H. Results from Experiment II.

[IIIDO mg/1-75 mg/l E300 mg/l

 

 

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Experiment II.

Experiment III

 

Since the primary goal of these experiments was to provide
information useful to the producer of G x H seed, the most definitive
criterion for assessing effective conversion was determined to be
maximum staminate flower production during the period of most concen-
trated fruit set. The plant responses most strongly influencing this
criterion are days to conversion, duration of staminate flowering, and
number of staminate flowers. Evaluation of each response is necessary
to determine the optimal treatment combination for effective conversion.

Significant interaction among stage, concentration, and appli-
cation number appeared for all responses examined in Experiment III
except days to first flower and days to first staminate flower. Toxicity
was inconsequential at any stage or concentration.

With regard to the total number of staminate flowers produced
along the main stem and laterals during the first three weeks of flower-
ing, single applications at any concentration as well as multiple
applications at 100 mg/l were in general indistinguishable from the
control (Figure 3). These results were similiar to those for Experi-
ments I and II. Treatments initiated at stage 0 resulted in fewer
number of staminate flowers than similiar treatments begun at later
stages. Stage 2 proved to be the most responsive since it required
but two applications of 200 to 400 mg/l to produce as many staminate
flowers as were produced by three applications at stage 1. Had flower-
ing been recorded over a longer period, plants treated at stage 3

would have produced more staminate flowers since treatments at higher

13

Figure 3. Mean number of staminate flowers per plant resulting from

AgNO applications on hermaphroditic pickling cucumber lines

3
MSU 7152 H and MSU 669H. Data from Experiment III.

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15
rates were continuing to produce staminate flowers at the end of the
experimental period. However, early conversion during the first three
weeks of flowering is more desirable for G x H seed production since
this is the period of most concentrated fruit set (3).

Treatments initiated at stage 0 resulted in staminate flowers
at the onset of flowering (Figure 4). Treatments initiated at stage I
began staminate flowering approximately one day later. The lag time
from treatment to staminate flowering depended strongly upon the H line
and varied up to eleven days. MSU 7152H first flowered approximately
one day earlier than MSU 669H. For treatments initiated at stages 2
and 3, MSU 7152B also entered the staminate flowering phase earlier.
Although MSU 669H required a longer period to express staminate
flowering, it produced more staminate flowers than did MSU 7152H.
Examination of the proportion of staminate flowers on the main stem
revealed a significantly higher conversion rate in MSU 669H compared to
MSU 7152B for treatments initiated at stages 0 and 1 (Table 3). In
addition, at silver nitrate levels of 200 to 400 mg/l, MSU 669H produc-
ed a greater total number of flowers on the main stem than did its
control (Table 4). MSU 7152H showed a similiar increase in total
flower number at 300 mg/l. It therefore appears that silver nitrate
can stimulate supernumerary flower bud development as was noted by others
working with gibberellic acid (1,5).

The maximum period of conversion occured for treatments initiated
at stages 0 and 1 (Figure 5). Additional applications contributed
progressively less as the time of initial treatment was delayed. Con-
currently, the period of staminate flowering diminished as initial

treatment times were delayed. Decreased sensitivity of the plants at

16

Figure 4. Duration of staminate conversion as influenced by genotype and
plant developmental stage at first application. Data from
Experiment III on staminate flower induction by AgNO3 on

hermaphroditic cucumber lines MSU 7152H and MSU 669H.

A Day of first treatment.

 

17

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Table 4.

18

Means for proportion of staminate to total number of flowers
per plant (averaged over concentration and application number)
resulting from AgNO application to two hermaphroditic
cucumber lines. Da a from experiment III.

 

 

Plant Line

Stage MSU 7152B MSU 669H
0 0.442 0.52*
l 0.56 0.63*
2 0.69 0.68
3 0.69 0.75

 

z - significance within rows at 5% level on arc sin
transformations of data presented in table

Means for total number of flowers per plant (averaged over
stage and application number) resulting from AgNO application
to two hermaphroditic cucumber lines. Data from Experiment
III.

AgNO

 

 

 

Concentragion Line 2
(mg/l) MSU 7152B MSU 669H
0 53.2 58.4
100 58.2 66.0
200 58.3 81.6
300 65.8 72.6
400 59.4 78.2

 

z- Scneffe's MSD within columns is 7.1 (5% level)

19

Figure 5. Mean duration of conversion to staminate flowering (averaged
over genotype) resulting from AgNO3 applications on hermaphro-
ditic cucumber lines MSU 7152H and MSU 669H. Data from

Experiment III .

Number of applications one 1' L“2 00000 3 III-Ill:

Days oi staminate flowering

20

.o
01

10

Days of stam’nate flowering

Figure 5.

 

20

100 200 300 400

Concentration AoNOaimg/l)

 

100 200 300 400

 

100 200 300 400

Concentration AgN03(mg/I)

21

later stages is supported by the increase in lag time between treatment
and conversion noted when treatments were initiated at progressively
later stages (Figure 4).

Other researchers have used the number of staminate nodes as an
indication of the effectiveness of treatments on sex expression (1,4,6,
8,11,12). An analysis of variance was calculated for per cent staminate
nodes to account for differences in growth rates between MSU 7152H and
MSU 669H. The conclusions from.this analysis were similiar to those
for duration of staminate flowering.

The maximum number of staminate flowers was produced when treat-
ments were initiated at stages 1 and 2. However, since the maximal
period of early conversion occured for treatments initiated at stages
0 and 1, stage 1 was determined to be the optimal stage at which to
begin treatment. The three responses selected for assessing effective
conversion to staminate flowering for G x H seed production were
optimized by three to four applications of 200 to 400 mg/l of AgNO3
initiated at stage 1.

A promising treatment combination suggested by these experiments

was examined in a preliminary field plot experiment. Excellent

conversion through node 20 was observed.

LI ST OF REFERENCES

10.

ll.

12.

22

LIST OF REFERENCES

Atsmon, D. and C. Tabbak. 1979. Comparative effects of gibberellin,
silver nitrate, and aminoethoxyvinylglycine on sexual tendency and
ethylene evolution in the cucumber plant (Cucumis sativus L.).
Plant and Cell Physiol. 20:1547-1555.

 

 

Beyer, E. 1976. Silver ion: a potent anti-ethylene agent in cucum-
ber and tomato. HortScience 11:195-196.

 

Bos, J.H.L. 1979. Manager, Worldwide Vegetable Seed Production,
Asgrow Seed, Co. Kalamazoo, Michigan. personal communication.

Den Nijs, A.P.M. and D.L.Visser. 1980. Induction of male flowering
in gynoecious cucumbers (Cucumis sativus L.) by silver ions.
Euphztica 29:273-280.

 

Fuchs, E., D. Atsmon, and A.H.Halevy. 1977. Adventitious staminate
flower formation in gibberellin treated gynoecious cucumber plants.
Plant and Cell Physiol. 18:1193-1201.

 

Kalloo, R.K.J. and S. Franken. 1978. Chemical induction of stami-
nate flowers in four determinate gynoecious lines of pickling
cucumber. Gartenbauwissenschaft 43:280-282.

 

Kubicki, B. 1965. New possibilities of applying different sex
types in cucumber breeding. Genetica Polonica 6:241-249.

Owens, K.W., C.E. Tolla, and C.E. Peterson 1980. Induction of
staminate flowers on gynoecious cucumber by aminoethoxyvinylgly-
cine. HortScience 15:256-257.

 

Pike, L.M. 1974. Tamu Triple Cross pickling cucumber. HortScience
9:83.

and W.A.Mulkey. 1971. Use of hermaphrodite lines in the
development of gynoecious hybrids. HortScience 6:339-340.

and C.E. Peterson. 1969. Gibberellin A /A7 for induction of
staminate flowers on the gynoecious cucumber Cucumis sativus L.).
Euphytica 18: 106-109.

 

Tolla, C.E. and C.E. Peterson. 1979. Comparison of gibberellin
A4/A7 and silver nitrate for induction of staminate flowers in a
gynoecious cucumber line. HortScience 14:542-544.

 

APPENDICES

APPENDIX A

23

APPENDIX A

Analysis of variance for Experiment I on staminate flower induction by

 

AgNO3 in hermaphroditic pickling cucumber line MSU 7152H
Total
Staminate Staminate Height
Nedes Flowers (cm)
Sources of variation df MS MS MS
concentration 3 458.3** 4127.5** 2222.1**
application number 2 26.3* 901.0** 4585.1**
concentration X 6 4.1 215.9** 1339.6*
application

blocks 4 22.8* 296.8** 1160.5**
error 44_ 6.9 70.0 495.0
total 59

 

*significance of F test at 5% level
** significance of F test at 1% level

APPENDIX B

24

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APPENDIX C

25

APPENDIX C

Analysis of variance for Experiment III on staminate flower induction
by AgNO3 on two unrelated hermaphroditic cucumber lines, MSU 7152H and
MSU 669H. (1) days to first flower, (2) days to first staminate flower,
(3) duration of staminate flowering, (4) per cent nodes bearing one or
more staminate flowers (are sin transformation), (5) total number of
staminate flowers, (6) per cent staminate flowers on the main stem

(arc sin transformation), (7) perfect flowers on the main stem,

(8) total number of flowers on the main stem.

26

Ho>oH NH can an ummu b no cosmonchHm «« “Ho>oH N0 ecu um ummu m we moccoHMchHm s

 

 

 

0.00N 0.00 No.0 0.NOH m0.0 5.0 5.0 m.N 00H nouns
«em.mHm m.mn No.0 «sm.mmm no.0 0.0 0.0H N.NH 0 mxooHn
«s0.0NNH *0.w0m «coN.0 «s0.0wNN x0H.0 H.m H.0 0.0 H chHumoHHaou
.oc :OHumoHHaam
x =OHumuucoosoo
0.50H 0.00 No.0 0.0mH m0.0 0.0 0.0 0.H 0m x ocHH x mwmum
.o: aOHumUHHaem
x ocHH
N.mmN 0.05 No.0 0.00N no.0 0.0 0.5 n.H NH x :OHumuucoocoo
.oc :0HumUHHaem
N.m0N 0.00 No.0 N.NON No.0 «0.HH 0.0 0.0 m x mcHH x ownum
cOHunuusmocoo
0.mn 0.0m No.0 m.NHH m0.0 N.N 0.N 0.H NH x oaHH x swoon
mcHH
«s0.000H H.N0 No.0 sa0.MMHH H0.0 0.H H.H «m.0 0 x :OHumuucoocoo
N.NNH «0.05H ss00.0 0.00 no.0 0.HH «s0.0N m.N m ocHH x wwnum
«sw.00MNH k«0.000 «wNN.0 ss0.0wm~ 0m.0 0.NH «s0.mwN «xH.0m H osHH
.o: :OHumUHHaam
_ x coHumuucoocoo
sN.Nmm sx0.HmH «x00.0 s«~.00m se00.0 «su.0H 0.0 m.N 0m x ammum
.o: GOHumoHHmam
asH.000 «em.0Hm «s0~.0 «sa.Hn0H «smm.0 s«0.00 0.0 0.0 NH x cOHumuucoucoo
.o: :OHumoHHaom
ss0.w0m «x0.mm0 ssm0.0 ssm.m0NN ss00.0 «s0.00 0.0 m.m m x mmcum
«sw.H~Hm «so.m00~ «sm0.m «sH.mmmmH ss00.m ssm.0cm 0.0 0.m m .0: :OHumoHHaan
. SOHumuucoocoo
H.HHm n.0NH ssm0.0 «£0.000 «cHH.0 «s0.hH 0.m N.0 NH x owcum
«sN.000N «em.Hm0HH «sN0.N ssm.n¢NMN sz~.0H «sm.mwNH H.N 0.0 0 :oHumuucouoou
«sm.~mm0 «e0.000 samH.0 ssN.NmmN «st.0 «sm.00N «sN.HONH «sm.wN m owmum
m: m: m: m: m: m: m: m: .0.0 cOHucHum>
0 m 0 0 0 m N H «o mmousom

o XHQ2mmm<

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