American Journal of Plant Sciences, 2012, 3, 1328-1335
http://dx.doi.org/10.4236/ajps.2012.39160 Published Online September 2012 (http://www.SciRP.org/journal/ajps)
Changes in the Ovarian Polyamine Content and Seed Set
Efficiency of Cotton by the Plant Growth Regulator BM86
Androniki C. Bibi, Derrick M. Oosterhuis, Evangelos D. Gonias
Department of Crop, Soil, and Environmental Sciences, University of Arkansas, Fayetteville, USA.
Email: an-bibi@hotmail.com
Received June 7th, 2012; revised July 6th, 2012; accepted July 19th, 2012
ABSTRACT
Naturally occurring polyamines in plants have been implicated in floral initiation, and fruit development. The plant
growth regulator BM86 was formulated to stimulate seed production and fruit growth by enhancing polyamine synthe-
sis. The objectives of this study were to determine the effect of BM86 on free polyamine content in cotton (Gossypium
hirsutum L.) ovaries and on seed set efficiency, and also to investigate genotypic differences in ovarian polyamine con-
tent. BM86 was applied at the first flower stage and two weeks later at 2.34 mL/ha. This study showed that application
of BM86 had a significant positive effect on ovarian polyamine content of cotton. Putrescine and spermidine one week
after the 1st BM86 application and putrescine two week after the 1st BM86 application, were significantly increased
compared to the untreated control. Higher seed set efficiency with the BM86 application was observed when the total
number of seeds was used for the calculation. However, when the number of harvestable seeds was used to calculate
seed set efficiency BM86 had no significant effect. In addition, application of BM86 did not significantly alter seedcot-
ton yield of the crop. Only small differences in ovarian polyamine content were detected among the genotypes tested,
possibly due to the narrow genetic pool of the commercial cotton genotypes used. Application of BM86 can signifi-
cantly increase cotton seed number by enhancing polyamines biosynthesis, but further research is needed to determine
how to capitalize on the increased potential number of harvestable seeds.
Keywords: Putrescine; Spermidine; Spermine; BM86; Gossypium hirusutm L.; Ovaries
1. Introduction
Cotton (Gossypium hirsutum L.) is a major industrial
crop grown for fiber, fuel, and feed, but yield suffers
from various biotic and abiotic stresses during the repro-
ductive development. The number of seed per boll is an
important basic component of both cotton yield and fiber
quality. Seed number is a function of the number of lo-
cules (carpels) per boll and the number of ovules per
locule [1] (Stewart 1986). Several factors such as the
lack of seed fertilization, post-fertilization termination of
embryo growth, cultivar and environment can contri-
buted to the variation in the number of seed per boll [2]
(Turner et al. 1977).
Polyamines are organic polycations involved in DNA
replication and cell division [3] (Bais and Ravishankar
2002), and they are intimately involved in successful re-
productive development of the crop, cell multiplication
and cell differentiation during organogenesis [4] (Costa
et al. 1984). Polyamines can act as regulators of many
physiological processes including flower induction [5-9]
(Evans and Malmberg 1989; Faust and Wang 1992;
Bagni et al. 1993; Bouchereau et al. 1999; Kakkar and
Sawhney 2002). According to Costa et al. (1984) [10]
polyamines play an important role in flowering, pollina-
tion and early fruit development. Kloareg et al. (1986)
[11] indicated that polyamines are indispensable to plants
at the time of flowering and early fruit development.
The documented importance of polyamines in repro-
ductive development directed many researchers to exo-
genously applying polyamines in an effort to enhance
fruit development. Reports in mango (Mangifera indica
L.) [12] (Singh and Janes 2000) and apricot (Prunus ar-
meniaca L.) [13] (Alburquerque et al. 2006) showed im-
proved fruit retention and yield. However, exogenous
application of polyamines in row crops is not a practical
management practice due to the high cost of the material.
An affordable alternative is to apply synthetic plant
growth regulators.
The plant growth regulator BM86 (active ingredient
GA14; Goëmar Laboratories, Saint-Malo, France) is an
elicitor that contains oligosaccharides reported to re- gu-
late the synthesis of endogenic polyamines. Broquedis et
al. (1995) [14] investigated the effect of GA14 on the
composition of polyamines in grapes (Vitis vinifera L.),
Copyright © 2012 SciRes. AJPS
Changes in the Ovarian Polyamine Content and Seed Set Efficiency of Cotton by the Plant Growth Regulator BM86 1329
and showed an increase in the accumulation of poly-
amines particularly at the end of the first stage of fruit
development. Treatment with GA14 increased the num-
bers, weight and growth of the fruit, and this was related
to a significant increase in polyamine content in the
flowers and in the fruit. It appears, therefore, that the
BM86 contains elicitors which stimulate the metabolism
of polyamines of endogenous growth.
Numerous anecdotal reports with horticulture plants
have shown that BM86 acts to stimulate reproductive
development of the plant. According to these reports
(Anonymous 2008) [15] BM86 increased yield and oil
production of olives (Olea europeae L.), promoted uni-
form fertilization and fruit set in grapes (Vitis vinifera L.)
and resulted in increased fruit size and total yield in cit-
rus (Citrus sp. L). In cotton there is a single report that
associates BM86 application with increased yield [16]
(Rethwisch 2006). However, there are no reports that
examine the effect of BM86 on polyamines in cotton.
Therefore in this study it was hypothesized that poly-
amine concentration in cotton ovaries will increase with
the application of BM86 and this will result in increased
seed set efficiency of cotton. Also, it was hypothesized
that genotypic differences in cotton will exist. The objec-
tives were to determine the effect of the BM86 on free
polyamines in cotton ovaries, and seed set efficiency of
cotton, and to investigate genotypic differences in ovule
polyamine content
2. Material and Methods
2.1. Plant Establishment
A field study was conducted at the Lon Mann Cotton
Research Station in Marianna Arkansas (34˚5'N, 90˚5'W)
in 2005 and 2006. The cultivars used were DP444BR,
ST5599BR, and FM960BR. The soil was a Captina silt
loam (typical fragiudult). The experimental plot size was
four rows by 15m and the plant population 10 plants per
m2. The study was furrow-irrigated based on an irrigation
scheduler program [17] (University of Arkansas Coo-
perative Extension Database Service 2007). The fertiliza-
tion program was determined by preseason soil tests and
recommended values for cotton. Weed and insect control
were conducted according to Arkansas extension recom-
mendations. At first flower BM86 (Goëmar Laboratories,
Saint-Malo, France) was applied to the right 2 rows of
each plot at 2.34 mL/ha (recommended rate from Goë-
mar Laboratories) with a backpack CO2 sprayer cali-
brated to deliver 94 L/ha of spraying solution. The left
two rows were used as the control. The day before the 1st
application of BM86, the flowering node was determined
and 10 first-sympodial fruiting position white flowers
were collected from each plot. Sampling was performed
weekly using first-position flowers two nodes higher than
the previous position, for a total of three weeks. BM86
was reapplied two weeks after the first application.
2.2. Number of Ovules/Ovary
Half of the flowers that were collected were used to de-
termine the number of ovules per ovary. The procedure
involved separating the ovary from the petals and sepals
and dissecting the ovaries to determine the number of
ovules in each ovary.
2.3. Polyamines Analysis
Chemical: Putrsecine dihydrochloride, spermidine trihy-
drochloride, spermine tetrahydrochloride, hexamethyla-
nediamine, dansyl chloride, 99.9% acetone, and sodium
carbonate were purchased from Sigma Aldrich.
Extraction of Tissues: Polyamines were extracted
based on Smith and Davies [18] (1985) with modifica-
tions needed for cotton. Cotton ovaries were separated
from petals and sepals. Ovary tissue (0.1 g) was homoge-
nized in mortars with pestles in 0.2 N HClO4. For unfor-
tified samples, 100 μL 1 mM hexamethylanediamine in
0.2 N HClO4 was added to the tissue prior to the ho-
mogenization as an internal standard. The final volume
of 2 mL was obtained by adding 1900 μL 0.2 N HClO4.
For the fortified samples, 100 μL 1 mM hexanediamine
in 0.2 N HCLO4 was added plus the desired volume of
fortification solution in 0.2 N HClO4, which was 120 μL
1 mM of Putrescine in 0.2 N HClO4, 120 μL 1 mM Sper-
midine in 0.2 N HClO4, and 120 μL 1mM of Spermine in
0.2 N HClO4. The final volume of 2 mL was obtained by
adding 1540 μL 0.2 N HClO4. An aliquot of 1.5 mL of
the homogenate was transferred to 2 mL plastic micro
centrifuge tubes and the samples were centrifuged at 4˚C
for 20 minutes at 13,000 rpm.
Dansylation: The polyamines were derivatized by
adding 50 μL aliquots of the supernatant to 500 μL 21.2
mM aqueous Na2CO3, 200 μL 99.9% acetone, and 50 μL
of 12.5 mM dansyl chloride in acetone. The mixture was
incubated in a thermal reaction block at 60˚C for 1h in
the dark. After 1 h in the thermal block the samples were
removed and cooled to near room temperature, and 50 μL
of 1 N HClO4 was added to the mixture and mixed. The
samples were then centrifuged at 4˚C for 20 minutes at
13,000 rpm after which 700 μL of centrifugate was trans-
ferred into 2 mL sample vial and 700 μL of 0.02 N
HClO4 was added. The samples were capped and mixed
before injection into the High Performance Liquid Chro-
matography (HPLC). Derivatization needs to be in a ba-
sic solution, whereas the final solution for HPLC needs
to be acidic.
Standard Preparation: A total of six standards were
used for the preparation of the standard curves. The stan-
dards included putrescine, spermidine, spermine and the
Copyright © 2012 SciRes. AJPS
Changes in the Ovarian Polyamine Content and Seed Set Efficiency of Cotton by the Plant Growth Regulator BM86
1330
internal standard hexamethylanediamine. The concentra-
tion of putrsecine and spermidine in the six standards
ranged from 30 to 1 nmole/mL, while the concentration
of spermine ranged from 60 to 2 nmole/mL. A 500 μL
aliquot of 1mM hexamethylanediamine was added to the
first five standards, while an amount of 1.25 mL of 1mM
hexamethylanediamine was added to the last one. The
final volume was 10 mL for the first five standards and
25 mL for the final standard. To bring the standards to
the final volume 0.2 N HClO4 was used.
HPLC Analysis: HPLC analysis was performed in a
Hitachi HPLC (Hitachi High Technologies America, Inc.,
Canada) system that included a model L-7100 pump, an
L-7200 autosampler, a D-7000 interface, and an ERC-
3415a degasser, and an L-7480 fluorescence detector.
The column used in this analysis was a 25 cm × 2 mm id
0.5 micron Phenomenex Gemini C18. Injection volume
was 40 μL. Polyamines were eluted from the column at
0.3 mL/min with a methanol:water (v/v) gradient from
70% methanol to 95% methanol over 6 min and then
remaining at 95% methanol for 16.4 min. The system
was re-equilibrated with 70% methanol for 15 min before
the next injection. For dansyl polyamines, an excitation
wavelength of 365 nm was used with an emission wave-
length of 510 nm. Data collection and processing was
with Hitachi System Manager (HSM) software on the
internal standard concentration.
2.4. Seed Set Efficiency (SSE) and Seedcotton
Yield
At harvest five bolls were hand-picked from each plot
from the same node from which flowers had been previ-
ously collected, for both control and BM86 treated plants.
The final number of seed per boll was determined from
the hand-picked bolls, and seed set efficiency was calcu-
lated using the equation: [seed set efficiency = (# of
seeds/# of ovules) × 100]. In a cotton boll at harvest there
are undeveloped ovules and harvestable seeds. For our
experiments seed set efficiency was calculated using both
the total seed number (undeveloped ovules + harvestable
seeds) and the number of the harvestable seeds. Seedcot-
ton yields were determined by mechanically harvesting
each individual sub-plot.
2.5. Statistical Analysis
The experimental design was a randomized complete
block (RCBD) with five replications. The treatment de-
sign was split-plot in strips (split-block) with main factor
being “cultivars” that were randomly assigned to the ex-
perimental plots, and sub-factor being “BM86 applica-
tion” that was not randomly assigned to allow easier
harvest. The year effect was considered random and not
included in the model. Data from each year was analyzed
and presented separately. Statistical analysis was per-
formed with the JMP 6 Software (SAS Institute Inc.,
Cary, NC). Interactions and main effects were tested with
Analysis of Variance (ANOVA) at α 0.05, when sig-
nificant effects were detected means were separated with
Student’s t-test (α 0.05).
3. Results
The analysis of the data showed no significant “cultivar ×
BM86 × year” interaction therefore “cultivar × BM86
application” was analyzed separately for each years of
the study for each parameter measured. In addition there
was no significant “cultivar × BM86 application” inter-
action in either year (Table 1). The lack of interaction
allowed us to analyze the main effects of “nodal posi-
tion” and “BM86 application” on putrescine spermidine,
spermine, seed set efficiency (calculated with the total
seed number), seed set efficiency (calculated with the
harvestable seed number), and seedcotton yield.
3.1. Effect of BM86 Application on Ovarian
Polyamine Content
One week after the 1st BM86 application: Ovarian putre-
scine concentration was significantly increased compared
to the untreated control plant in 2005 (P = 0.0047; Fig-
ure 1(A)). However, application of BM86 did not sig-
nificantly affect spermidine and spermine concentration
with P = 0.3927 and P = 0.9337, respectively (Figure
1(I)). These data were further supported in 2006 where
again one week after the BM86 application the putre-
scine content was significantly higher compared to the
control (P = 0.0390; Figure 1(II)). In addition in 2006
the BM86 application significantly increased the sper-
midine content of cotton ovaries (P = 0.0094), but again
the effect on spermine was not significant (P = 0.3791)
(Figure 1(B)).
Two weeks after the 1st BM86 application: The putre-
scine content of cotton ovaries continued to be signifi-
Table 1. The statistical significant for the “cultivar × BM86
application” interaction of all the parameters measured in
2005 and 2006.
“Cultivar × BM86 application”
Variables measured 2005 2006
Putrescine 0.17 0.57
Spermidine 0.99 0.46
Spermine 0.19 0.24
Seed set efficiency (total seed
number) 0.54 0.26
Seed set efficiency (harvestable
seed number) 0.99 0.59
Seedcotton yield 0.64 0.46
Copyright © 2012 SciRes. AJPS
Changes in the Ovarian Polyamine Content and Seed Set Efficiency of Cotton by the Plant Growth Regulator BM86
Copyright © 2012 SciRes. AJPS
1331
A
A
A
A
A
B
Putrescine
Spermine
Spermidine
BM86
CONTROL
0 500 1000 1500 2000 2500
nmoles/g fresh wt
A
A
A
B
A
B
Putrescine
Spermine
Spermidine
BM86
CONTROL
0 500 1000 1500 2000 2500 3000
nmoles/g fresh wt
(I) (II)
Figure 1. The effect of the plant growth regulator BM86 polyamine concentration of cotton ovaries one week after the first
BM86 application in 2005 (I) and 2006 (II). Pair of columns for each polyamine with the same letter are not significantly dif-
ferent for α = 0.05 (± 1 std error bars are shown).
3.3. Effect of Cultivars on Ovarian Polyamine
Content
cantly increased compared to the untreated control plant
(P = 0.0492; Figure 2(I)). However the BM86 applica-
tion did not significantly affect either spermidine or sper-
mine concentration (P = 0.0643 and P = 0.2852, respec-
tively) (Figure 2(I)). Similar effects of BM86 were re-
corded in 2006 with the putrescine content significantly
higher compared to the control (P = 0.0192; Figure
2(II)), and spermidine and spermine concentration not
significantly affected (P = 0.0611 and P = 0.2539, re-
spectively) (Figure 2(II)).
One week after the 1st BM86 application: In 2005 the
cultivar effect was not significant for putrescine (P =
0.5119), spermidine (P = 0.1812), or spermine (P =
0.3451) (Table 2). In 2006 there were no cultivar diffe-
rences for putrescine and spermine (P = 0.1926 and P =
0.2767, respectively), but there were differences for sper-
midine (P = 0.0439) (Table 2). Among the genotypes
tested, FM960BR showed significantly higher ovarian
spermidine content compared to DP444BR and ST5599-
BR.
One week after the 2nd BM86 application: The BM86
application did not significantly affect any of the three
polyamines quantified in 2005 (Figure 3(I)). The P-values
for putrescine, spermidine, and spermine were P = 0.5529,
P = 0.0717, and P = 0.2273, respectively. In 2006, one
week after the BM86 application the ovarian putrescine
content was not significantly altered (P = 0.1780; Figure
3(II)), whereas the spermidine concentration of cotton
ovaries significantly increased compared to the untreated
control (P = 0.0124). Spermine was not detected in cot-
ton ovaries one week after the 2nd BM86 application.
Two weeks after the 1st BM86 application: In 2005, the
putrescine, spermidine and spermine concentrations of
cotton ovaries did not differ significantly among the cul-
tivar (Table 3). In 2006 there were cultivar differences
for putrescine and spermine (P = 0.0087 and P = 0.0369
respectively), but the spermidine concetration was not
significantly changed (P = 0.1499) (Table 3). Among the
genotypes tested FM960BR showed significantly higher
ovarian putrescine content compared to DP444BR and
ST5599BR, and DP444BR showed significantly higher
spermine concentration compared to ST55899BR.
3.2. Effect of BM86 Application on Seed Set
Efficiency and Seedcotton Yield
One week after the 2nd BM86 application: In 2005,
there were no cultivar differences for spermidine and
spermine concentration of cotton ovaries (Table 4).
However the cultivar effect was significant for putrescine
concentration (P = 0.0349). Among the genotypes tested
FM-960BR and ST5599BR showed significantly higher
putrescine compared to DP444BR. In 2006 neither putre-
scine nor spermidine concentration of cotton ovaries
were significantly affected by the cultivar effect with P =
0.1780 and P = 0.9024, respectively (Table 4).
In 2005 BM86 application significantly increased the
seed set efficiency of cotton when calculated using the
total number of seeds (P = 0.0423; Figure 4(I)), however
application of BM86 did not significantly affect seed set
efficiency when it was calculated with just the number of
harvestable seeds (P = 0.3287). Similar data were ob-
served in 2006, the seed set efficiency calculated by the
total seed number was significantly increased after BM86
application (P = 0.0046; Figure 4(II)), while when cal-
culated with just the number of harvestable seeds the
effect was not significant (P = 0.7434). The non-signifi-
cant effect of BM86 application on seed set efficiency
(i.e. of harvestable seeds) was reflected in the non-signi-
ficant effect on seedcotton yield in either year of the
study (Figure 5).
3.4. Effect of Cultivars on Seed Set Efficiency
and Seedcotton Yield
There were no cultivar differences in 2005 and 2006 for
seed set efficiency calculated by the total seed number
Changes in the Ovarian Polyamine Content and Seed Set Efficiency of Cotton by the Plant Growth Regulator BM86
1332
A
A
A
A
A
B
Putrescine
Spermine
Spermidine
BM86
CONTROL
0 500 1000 1500 2000 2500
nmoles/g fresh wt
A
A
A
A
A
B
Putresc ine
Spermin e
Spermi dine
BM86
CONTROL
0 500 1000 1500 2000 2500 3000
nmoles/g fresh wt
(I) (II)
Figure 2. The effect of the plant growth regulator BM86 on the polyamine concentration of cotton ovaries two weeks after the
first BM86 application in 2005 (I) and 2006 (II). Pair of columns for each polyamine with the same letter are not significantly
different for α = 0.05 (± 1 std error bars are shown).
A
A
A
A
A
A
Putrescine
Spermine
Spermidine
BM86
CONTROL
0 500 1000 1500 2000 2500
nmoles/g fresh wt
A
A
A
B
Putrescine
Spermine
Spermidi n e
BM86
CONTROL
0 500 1000 1500 2000 2500 3000
nmoles/g fresh wt
(I) (II)
Figure 3. The effect of the plant growth regulator BM86 polyamine concentration of cotton ovaries one week after the second
BM86 application in 2005 (I) and 2006 (II). Pair of columns for each polyamine with the same letter are not significantly dif-
ferent for α = 0.05 (± 1 std error bars are shown).
B
A
A
A
0
10
20
30
40
50
60
70
80
90
100
Total Seed NumberHarvestable seeds
Seed Set Efficien cy (%)
CONTROL
BM86
B
A
A
A
0
10
20
30
40
50
60
70
80
90
100
Total Seed NumberHarvestable seed s
Seed Set Efficiency (%)
CONTROL
BM86
(I) (II)
Figure 4. The effect of BM86 on seed set efficiency of cotton in 2005 (I) and 2006 (II). Pairs of columns with the same letter
are not significantly different for α = 0.05 (± 1 std error bars are shown).
and in seed set efficiency calculated by the harvestable
seed number (Table 5). In addition, the cultivar effect
was not significant for seedcotton yield in either year of
the study with P = 0.2209 (2005) and P = 0.6572 (2006)
(Table 6).
4. Discussion
This study showed that application of BM86 had a sig-
nificant positive effect on ovarian polyamine content of
cotton. The results were positive for putrescine and
spermidine one week after the 1st BM86 application and
Copyright © 2012 SciRes. AJPS
Changes in the Ovarian Polyamine Content and Seed Set Efficiency of Cotton by the Plant Growth Regulator BM86 1333
A
A
A
A
0
500
1000
1500
2000
2500
3000
3500
4000
4500
2005 2006
Year
Seedcotton (kg/ha)
CONTROL
BM86
Figure 5. The effect of BM86 on seedcotton yield in 2005
and 2006. Pair of columns within a year with the same letter
are not significantly different for α = 0.05 (± 1 std error
bars are shown).
and for putrescine two weeks after the 1st BM86 applica-
tion. Broquedis et al. (1995) [14] investigated the effect
of GA14, the active ingredient in BM86, and showed an
increase in the accumulation of polyamines in grapes
(Vitis vinifera L.). In addition, there are several anecdotal
reports in horticulture crops that associate the BM86 ap-
plication with increased polyamine levels; however this
is the first documented report of increased polyamines in
cotton ovaries. Among the genotypes tested only FM-
960BR showed increased polyamine content, however
this was not consistent from year to year. Polyamines
have been quantified in diverse rice cultivars [19] (Lepri
et al. 2002), while post-harvest changes in polyamines
have been measured in two diverse rose species [20]
(Sood and Nagar 2008). This is the first report of cotton
cultivar screening for polyamines in ovarian tissue after
Table 2. Cultivar effect on ovarian polyamine content one week after the 1st application of BM86 in 2005 and 2006.
Putrescine Spermidine Spermine
2005 2006 2005 2006 2005 2006
Cultivars
nmoles/g fresh wt
DP444BR 1300.3 a1 1403.5 a862.1 a 853.1 b187.1 a 517.3 a
FM960BR 1750.8 a 1461.9 a1105.3 a 1320.6 a196.7 a 682.9 a
ST5599BR 1442.2 a 1074.5 a883.6 a 979.9 b157.1 a 528.6 a
P-Value 0.5119 0.1926 0.1812 0.0439 0.3451 0.2767
1Cultivars with the same letter in a column are not significantly different for α = 0.05.
Table 3. Cultivar effect on ovarian polyamine content two weeks after the 1st application of BM86 in 2005 and 2006.
Putrescine Spermidine Spermine
2005 2006 2005 2006 2005 2006
Cultivars
----------nmoles/g fresh wt---------
DP444BR 590.1 a1 904.8 b431.9 a 273.2 a 128.3 a 125.5 a
FM960BR 609.4 a 1337.6 a472.7 a290.7 a 144.4 a 111.7 ab
ST5599BR 640.9 a 975.6 b405.5 a 253.8 a 120.7 a 103.6 a
P-Value 0.7463 0.0087 0.3135 0.1499 0.9632 0.0369
1Cultivars with the same letter in a column are not significantly different for α = 0.05.
Table 4. Cultivar effect on ovarian polyamine content one week after the 2nd application of BM86 in 2005 and 2006.
Putrescine Spermidine Spermine
2005 2006 2005 2006 2005 2006
Cultivars
----------nmoles/g fresh wt---------
DP444BR 106.8 b1 79.9 a 479.4 a34.1 a 21.4 a -
FM960BR 157.3 a 145.4 a 872.2 a30.5 a 25.9 a -
ST5599BR 163.8 a 137.2 a 822.9 a31.9 a 30.2 a -
P-Value 0.0349 0.1780 0.2813 0.9024 0.6550 -
1Cultivars with the same letter in a column are not significantly different for α = 0.05.
Copyright © 2012 SciRes. AJPS
Changes in the Ovarian Polyamine Content and Seed Set Efficiency of Cotton by the Plant Growth Regulator BM86
1334
Table 5. The effect of cultivars on seed set efficiency of cotton calculated by the total seed number and by the number of the
harvestable seeds in 2005 and 2006.
Cultivars Seed Set Efficiency
2005 2006
Seed Record
%
Total seed number DP444BR 70.3 a1 80.9 a
FM960BR 66.4 a 77.7 a
ST5599BR 76.3 a 80.7 a
P-Value 0.0584 0.4405
Harvestable seeds DP444BR 88.0 a 97.5 a
FM960BR 87.4 a 98.8 a
ST5599BR 91.3 a 96.3 a
P-Value 0.3008 0.4386
1Cultivars in a column for each parameter with the same letter are not significantly different for α = 0.05.
Table 6. The effect of cultivars on seedcotton yield of cotton
for 2005 and 2006.
Cultivars Seedcotton
2005 2006
kg/ha
DP444BR 3839.8 a1 3158.3 a
FM960BR 3692.6 a 3155.6 a
ST5599BR 4056.2 a 2989.9 a
P-Value 0.2209 0.6572
1Cultivars in a column with the same letter are not significantly different for
α = 0.05.
BM86 application.
Polyamines have been associated with fruit set and the
initial phase of fruit development in many horticulture
plants, such as apple (Malus domestica L.) [10,21] (Costa
and Bagni 1983; Biasi et al. 1991), pear (Pyrus commu-
nis L.) [22,23] (Crisosto et al. 1986; Crisosto et al. 1988),
pepper (Capsicum annuum L.) [24] (Serrano et al. 1995),
olive (Οlea europeae L.) [25] (Rugini and Mencuccini
1985), mango (Mangifera indica L.) [26] (Singh and
Singh 1995), and tomato (Solanum lycopersicum L.) [27]
(Antognoni et al. 2002). This can be attributed to a pos-
sible involvement of polyamines in cell growth [3] (Bais
and Ravishankar 2002). Therefore the increased levels of
polyamines reported in our study were expected to result
in improved seed set. Higher seed set efficiency with the
BM86 application was observed when the total number
of seeds was used for the calculation. However, when the
number of harvestable seeds was used to calculate seed
set efficiency BM86 did not have a significant effect. In
addition, BM86 application did not significantly alter
seedcotton yield of the crop. These data suggest that
BM86 application may set the potential for higher seed
set efficiency, but the cotton crop in Arkansas does not
have the resources to develop the set seeds to harvestable
seeds. In other locations such as California, higher cotton
yields after application of BM86 [16] (Rethwich 2006)
are possibly due to the higher productive environment
[28] (Gonias et al. 2008). The lack of significant diffe-
rences among the genotypes tested can be attributed to
the narrow genetic pool of the current commercial geno-
types.
5. Acknowledgements
The authors would like to acknowledge Dr. John Mattice
for the significant help with the HPLC analysis.
REFERENCES
[1] J. McD. Stewart, “Integrated Events in Flower and Fruit,”
In: J. R. Mauney and J. M. Stewart, Eds., Cotton physi-
ology, The Cotton Foundation, Memphis, 1986, pp. 261-
300.
[2] J. H. Turner, J. M. Stewart, P. E. Hoskinson and H. H.
Ramey, “Seed Setting Efficiency in Eight Cultivars of
Upland Cotton,” Crop Science, Vol. 17, No. 5, 1977, pp.
769-772.
doi:10.2135/cropsci1977.0011183X001700050023x
[3] H. P. Bais and G. A. Ravishankar, “Role of Polyamines in
the Ontogeny of Plants and Their Biotechnological Ap-
plications,” Plant Cell, Tissue and Organ Culture, Vol.
69, No. 1, 2002, pp. 1-34. doi:10.1023/A:1015064227278
[4] G. Costa, R. Baraldi and N. Bagni, “Influence of Putre-
scine on Fruit-Set of Apple,” Acta Horticulturae, Vol.
149, 1984, pp. 189-195.
[5] P. T. Evans and R. L. Malmberg, “Do Polyamines Have
Roles in Plant Development?” Annual Review of Plant
Physiology and Plant Molecular Biology, Vol. 40, 1989,
pp. 235-269. doi:10.1146/annurev.pp.40.060189.001315
[6] M. Faust and S. Y. Wang, “Polyamines in Horticulturally
Important Plants,” Horticultural Reviews, Vol. 14, 1992,
pp. 333-356.
[7] N. Bagni, M. M. Altamura, S. Biondi, M. Mengoli and P.
Copyright © 2012 SciRes. AJPS
Changes in the Ovarian Polyamine Content and Seed Set Efficiency of Cotton by the Plant Growth Regulator BM86 1335
Torrigiani, “Polyamines and Morphogenesis in Normal
and Transgenic Plant Cultures,” In: K. A. Roubelakis-
Angelakis and K. Tran Thanh Van, Eds, Morphogenesis
in Plants, Plenum Press, New York, 1993, pp. 89-111.
[8] A. Bouchereau, C. Duhaze, J. Martin-Tanguy, J. P.
Guegan and F. Larher, “Improved Analytical Methods for
Determination of Nitrogenous Stress Metabolites Occur-
ring in Limonium Species,” Journal of Chromatography
A, Vol. 836, 1999, pp. 209-221.
doi:10.1016/S0021-9673(99)00078-3
[9] R. K. Kakkar and V. P. Sawhney, “Polyamine Research
in Plants: A Changing Perspective,” Plant Physiology,
Vol. 116, No. 3, 2002, pp. 281-292.
doi:10.1034/j.1399-3054.2002.1160302.x
[10] G. Costa and N. Bagni, “Effect of Polyamines on Fruit
Set of Apple,” Horticultural Science, Vol. 18, 1983, pp.
59-61.
[11] B. Kloareg, M. Broquedis and J. M. Joubert, “Fruit De-
velopment: Elicitor Effects of Biostimulants,” Arboricul-
ture Fruitiere, Vol. 498, 1996, pp. 39-42.
[12] Z. Singh and J. Janes, “Regulation of the Fruit Set and
Retention in Mango with Exogenous Application of Poly-
amines and Their Biosynthesis Inhibitors,” Acta Hor-
ticulturae, Vol. 509, 2000, pp. 675-680.
[13] N. Alburquerque, J. Egea, L. Burgos, D. Martinez-
Romero, D. Valero and M. Serrano, “The Influence of
Polyamines on Apricot Ovary Development and Fruit
Set,” Annals of Applied Biology, Vol. 149, No. 1, 2006,
pp. 27-33. doi:10.1111/j.1744-7348.2006.00067.x
[14] M. Broquedis, P. Lespy-Labayette and J. Bouard, “Com-
position en Polyamines des Grappes au Moment de la
floraison. Effect de Pulverizations de Crème d’Algue
Cryobroyée,” Phytoma-La Défense des Végétaux No. 474,
1995.
[15] Anonymous Ag & Garden Imports, “GOËMAR BM86
Improve Health, Increase Production Naturally,” 2008.
www.agandgarden.com
[16] M. D. Rethwitch, T. Cox, D. M. Ramos, M. Luna and J.
Wellman, “Effect of Goёmar BM86 and Mepiquat Chlo-
ride on DPL449BR and DPL 484R Cotton,” Arizona
Cotton Report (P-145), University of Arizona, 2006, pp.
179-193.
[17] University of Arkansas Cooperative Extension Database
Service, “Irrigation Scheduler,” 2007.
http://www.aragriculture.org/computer_programs/irrigatio
n_scheduling/default.asp
[18] M. A. Smith and P. J. Davies, “Separation and Quanti-
fication of Polyamines in Plant Tissue by High Perform-
ance Liquid Chromatography of Their Dansyl Deriva-
tives,” Plant Physiology, Vol. 78, No. 1, 1985, pp. 89-91.
doi:10.1104/pp.78.1.89
[19] O. Lepri, L. Bassie, P. Thu-Hang, P. Christou and T.
Cappell, “Endogenous Enzyme Activities and Polyamine
Levels in Diverse Rice Cultivars Depend on the Genetic
Background and Are Not Affected by the Presence of the
Hygromycin Phosphotransferase Selectable Marker,”
Theoretical Applied Genetics, Vol. 105, No. 4, 2002, pp.
594-603. doi:10.1007/s00122-002-0922-4
[20] S. Sood and P. K. Nagar, “Post-Harvest Alternationsin
Polyamines and Ethylene in Two Diverse Rose Species,”
Acta Physiologiae Plantarum, Vol. 30, No. 2, 2008, pp.
243-248. doi:10.1007/s11738-007-0113-7
[21] R. Biasi, G. Costa and N. Bagni, “Polyamine Metabolism
as Related to Fruit Set and Growth,” Plant Physiology
and Biochemistry, Vol. 29, 1991, pp. 497-506.
[22] C. H. Crisosto, P. B. Lombard, D. G. Richardson, R.
Tetley and M. D. Vasilakis, “Effect of Ethylene Inhibitors
on Fruit Set, Ovule Longevity and Polyamnies Levels in
‘Comice’ Pear,” Acta Horticulturae, Vol. 179, 1986, pp.
229-236.
[23] C. H. Crisosto, P. B. Lombard, D. Sugar and V. S. Polito,
“Putrescine Infuences Ovule Senecence, Fertilization
Time, and Fruit Set in ‘Comice’ Pear,” Journal of the
American Society for Horticultural Science, Vol. 113,
1988, pp. 708-712.
[24] M. Serrano, M. C. Marinez-Madrid, F. Riquelme and F.
Romojaro, “Endogenous Levels of Polyamines and Ab-
scisic Acid in Pepper Fruits during Growth and Ripen-
ing,” Annals of Applied Biology, Vol. 149, 1995, pp. 27-
33.
[25] E. Rugini and M. Mencuccini, “Increased Yield in the
Olive with Putrescine Treatment,” Horticultural Science,
Vol. 20, 1985, pp. 102-103.
[26] Z. Singh and L. Singh, “Increased Fruit Set and Retention
in Mango with Exogenous Application of Polyamines,”
Journal of Horticultural Science and Biotechnology, Vol.
70, 1995, pp. 271-277.
[27] F. Antognoni, F. Ghetti, A. Mazzucato, M. Franceschetti
and N. Bagni, “Polyamine Pattern during Flower Devel-
opment in the Parthenocarpic Fruit (par) Mutant of To-
mato,” Physiologia Plantarum, Vol. 116, No. 4, 2002, pp.
539-547. doi:10.1034/j.1399-3054.2002.1160413.x
[28] E. D. Gonias, D. M. Oosterhuis and A. C. Bibi, “Cotton
Radiation Use Efficiency Response to Plant Growth
Regulators,” Journal of Agricultural Science, Cambridge.
Published online 24 October 2011.
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