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ABSTRACT ince commercial cotton cultivars producing
insecticidal Bt proteins were planted in 1997, Bt cotton has
Changes in vegetative and reproductive growth have been
been planted widely in China and other areas of the world
widely observed in Bacillus thuringiensis (Bt) transgenic cotton
(Clive, 2012; Huang et al., 2010).The Bt technology improved
(Gossypium hirsutum L.). The objectives of this study were to
growers’ profitability, reduced environmental pollution from
determine the characteristics of vegetative and reproductive
synthetic insecticides, and increased workers’ safety (Huang et
growth after the Bt gene was introduced into cotton and to
al., 2001; Kranthi et al., 2005; Pray et al., 2001). However, unsta-
study the effect of N on vegetative and reproductive growth to
ble lint yields have been observed frequently among farms in
determine if N nourishment could provide a potential mechanism
cotton production regions; some farms have a higher lint harvest,
for changed growth characteristics in Bt cotton. Three Bt
while other farms growing the same Bt cultivars have a lower
cultivars (Sikang1, Ccsi41, and Ccsi45), with their current parents
lint harvest under similar ecological conditions (Dong et al.,
(Simian3, Ccsi23, and Jimian7, respectively), were chosen to
2011; Li et al., 2005). Lint reduction has been attributed to the
compare the characteristics of vegetative and reproductive growth
application of traditional cultural technology resulting in heavy
and N distribution, in 2011 to 2012; the same six cultivars with
abscission of reproductive organs in Bt cultivars. Because vegeta-
two N fertilizer application rate treatments (0 and 450 kg/ha)
tive and reproductive growth characteristics were changed by the
were further designed to investigate the N regulation impact on
introduction of the Bt gene, management practices also require
growth in 2013. The results indicated that the Bt cotton cultivars
change. Tian et al. (2000) reported that Bt cultivar CSC30 had
had higher growth rates of main stem leaf area, sympodium leaf
smaller bolls and a lower lint percentage than its parent. Higher
number, and plant height and a lower growth rate of sympodium
vegetative growth, including increased plant height and higher
leaf area than their parents. Higher numbers and development
relative growth rate and biomass, were also observed in Bt cot-
rates of fruiting nodes, fruiting branches, bolls, and retained boll
ton (Chen et al., 2002; Tian et al., 2000). In contrast to their
rates were also observed but lower fruiting branch positions than
parents, Bt cultivars showed earlier boll opening, more fruiting
their parents. The N-deficit and N-rich treatments affected both
nodes, and a boll-setting rate that was maintained during repro-
vegetative and reproductive growth of Bt cultivars. The Bt cultivars
ductive growth. However, the causes of the changed characteris-
had reduced numbers and rates of vegetative and reproductive
tics during growth and development were unclear. Chen and Gu
organs under N deficit compared with their parents. These
(2004) and Zhang et al. (2007) measured a changed N metabo-
results show that the introduction of the Bt gene changed growth
lism in Bt cotton. We, therefore, hypothesized that the changed
characteristics, including reduced vegetative and reproductive
vegetative and reproductive growth in Bt cotton was related to a
organ development under N deficit.
changed N distribution.
The objectives of this study were: (i) to determine the
characteristics of vegetative and reproductive growth after the
Core Ideas
introduction of the Bt gene into cotton; and (ii) to study the effects
• Bt cotton cultivars had higher sympodium leaf number and
plant height. of N on vegetative and reproductive growth to determine if N
• Bt cotton cultivars had higher growth rates of main stem leaf and nourishment could account for the changed growth characteristics
lower growth rate of sympodium leaf area. in Bt cotton.
• Bt cotton cultivars had higher numbers of fruiting branches,
fruiting nodes, bolls, and retained boll rates. MATERIALS AND METHODS
• N-deficit and N-rich treatments affected number and rates of Plant Material and Experimental Design
vegetative and reproductive organs.
The study was conducted on the farm of Yangzhou
University (32°30¢ N, 119°25¢ E) in 2011 to 2013. Three Bt
transgenic cotton cultivars, Sikang1, Ccsi41, and Ccsi45,
Published in Agron. J. 109:168–174 (2017) widely grown in China, and their parents were used in this
doi:10.2134/agronj2016.04.0209
Received 12 Apr. 2016 E. Abidallha, L.I.M. Tambel, L. Heng, X. Zhang, and D.H. Chen, Jiangsu
Accepted 22 Sept. 2016 Provincial Key Lab. of Crops Genetics and Physiology, Yangzhou Univ.,
Available freely online through the author-supported open access option Yangzhou 225009, P.R. China; E. Abidallha, Dep. of Forest Management,
Faculty of Forestry, Univ. of Khartoum, 13314, Khartoum, Sudan.
Copyright © 2017 American Society of Agronomy
*Corresponding author (zhangxiang@yzu.edu.cn, cdh@yzu.edu.cn).
5585 Guilford Road, Madison, WI 53711 USA
This is an open access article distributed under the CC BY-NC-ND Abbreviations: Bt, Bacillus thuringiensis FP, first fruiting branch
license (http://creativecommons.org/licenses/by-nc-nd/4.0/) position.
168 A g ro n o my J o u r n a l • Vo l u m e 10 9, I s s u e 1 • 2 017
study. Sikang1, Ccsi41, and Ccsi23 were the conventional Bt leaf and sympodium leaf, main stem nodes, plant height, and
cultivars developed by introducing Bt genes into Simian3, growth rate. Ten leaf samples for each experimental cultivar
Ccsi23, and Jimian7 (non-transgenic cultivars), respectively. from each plot were harvested. Leaf samples were the four
The use of the three Bt cultivars with their current parents was top leaves of the main stem plus 10 sympodial leaf samples
designed to compare the characteristics of vegetative growth, collected from the first nodes of the fruit branches at mid-
reproductive growth, and N distribution between the Bt cultivar plant. All the leaf samples were dried at 80°C to constant
and its parent during cotton growing seasons in 2011 and 2012. weight. The H2SO4 –H2O2 Kjeldahl digestion method
The plot dimensions were 5.4 by 8 m, containing six rows spaced was used for the measurement of total leaf N concentration
at 0.90 by 0.30 m. Cultivar was used as the experimental factor. (Richards, 1993). Nitrogen content was expressed as the
The experiments were arranged according to a randomized product of concentration and dry weight.
complete block design with three replications. Seeds were sown Growth indices of the reproductive organs were investigated
on 4 April in a warm room and covered with plastic film in every 15 d from peak boll setting to boll opening and included
both years. Seedlings were transplanted to the field on 17 May. the position of the first branch, the numbers of fruit branches
The field had a sandy loam soil (Typic Fluvaquents according to and fruit nodes, boll number, and their growth rates. At 10 d
the US taxonomy), which contained 24.5 g/kg organic matter, after flowering, 10 bolls were harvested from the first nodes of
and available N, P, and K at 108, 40.5, and 82.0.0 mg/kg, the fruit branches at mid-plant. All the boll samples were dried
respectively. Nitrogen (60 kg/ha as urea), P (300 kg/ha as single and used for measurement of N content.
superphosphate), and K (120 kg/ha as KCl) were applied and In 2013, the growth characteristics of vegetative organs
incorporated before transplanting. Nitrogen (54 kg/ha as urea), and reproductive organs were investigated under different N
P (300 kg/ha as single superphosphate), and K (120 kg/ha as treatments for Bt cultivars and their parents.
KCl) were applied again at the early flowering stage. Nitrogen
was also applied at early boll development (126 kg/ha as urea) Statistical Analysis
and peak boll (30 kg/ha as urea) stages. Mepaquat chloride A Student’s t-test was conducted for each characteristic of
(1,1-dimethyl piperidinium chloride, C7H16ClN) was applied at growth and N distribution between Bt cultivars and their
peak square (15 g/ha), early flowering (30g/ha), peak flowering parents, using proc ANOVA in SAS 6. The data are presented
(45g/ha), and peak boll (60g/ha) stages. separately by year.
Based on the results of 2011 and 2012, the same six cultivars
with two N fertilizer application rate treatments were further Results
tested on Yangzhou University Farm in 2013. The 2 N fertilizer Effect of the Introduction of the Bt Gene
treatments consisted of 0 (N deficit) and 450 kg/ha (N rich) into Cotton on Vegetative Organ Growth
The experiment was arranged in a randomized complete block The number of main stem leaves per plant was similar for all
design, with three replications for each treatment. The sowing the studied cultivars during the cotton growing seasons of 2011
date was 5 April, and the transplanting date was 18 May. The and 2012 (Table 1)—approximately 25 leaves. No significant
seedlings were planted at a spacing of 0.90 by 0.30 m. The plot differences in main stem leaf number were detected between
dimensions were also 5.4 by 8 m. With the exception of the N Bt cultivars and their corresponding parents. However, there
treatments, the same management practices were used in both were differences in main stem leaf area (Table 1). Cultivars
years according to local recommendations. containing the Bt gene (Sikang1, Ccsi41, and Ccsi45) had
higher leaf area than their corresponding parents (Simian3,
Measurements Ccsi23, and Jimian7), and the differences were significant
In 2011 and 2012, the growth characteristics of vegetative between Bt cultivars and their parents in both years (2.4 and
organs and reproductive organs were investigated every 20 d 3.3% increase between Sikang1 and Simian3, 3.0 and 2.4%
from square appearance to boll opening. The indices for the between Ccsi41 and Ccsi23, and 4.1 and 2.6% between Ccsi45
vegetative organs included number and area of the main stem and Jimian7 for 2011 and 2012, respectively).
Table 1. The number and leaf area of main stem leaves and sympodium leaves and plant height for Bt cotton cultivars and their parents in
the 2011 and 2012 cotton growing seasons.
Main stem leaf Sympodium leaf Plant height
Cultivar 2011 2012 2011 2012 2011 2012 2011 2012 2011 2012
––– leaves/plant––– ––– cm2/leaf ––– ––– leaves/plant ––– ––– cm2/leaf ––– ––––– cm –––––
Sikang1 25.9 26.2 4520 4573 103.2 105.5 4216 4167 108.4 112.3
Simian3 25.6 26.0 4415 4426 92.5 95.3 4394 4275 102.3 105.2
Significance ns† ns 0.014 0.036 0.043 <0.01 <0.01 <0.01 <0.01 <0.01
Ccsi41 25.1 25.4 4640 4710 99.6 102.6 4298 4212 112.3 116.7
Ccsi23 24.7 25.1 4505 4602 90.4 94.7 4452 4398 104.6 107.1
Significance ns ns <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01
Ccsi45 25.4 25.6 4705 4736 95.2 97.4 4315 4265 115.1 120.4
Jimian7 25.2 25.5 4520 4615 87.3 91.9 4521 4472 105.6 109.1
Significance ns ns <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01
† ns, not significant at the 0.05 level.
Table 2. The growth rate of vegetative organs for Bt cotton cultivars and their corresponding parents in the 2011 and 2012 cotton grow-
ing seasons.
Growth rate Growth rate Nodes of main Plant
of main stem leaf of sympodium leaf stem height
Cultivar 2011 2012 2011 2012 2011 2012 2011 2012 2011 2012 2011 2012
––– leaves/d ––– ––– cm2/d ––– ––– leaves/d ––– ––– cm2/d ––– ––– nodes/d ––– ––– cm/d –––
Sikang1 0.273 0.281 256.3 260.1 1.81 1.87 246.4 243.2 0.27 0.3 1.2 1.31
Simian3 0.268 0.28 247.5 251.1 1.58 1.6 250.3 247.8 0.27 0.29 1.09 1.15
Significance ns† ns 0.024 0.035 0.047 0.023 0.028 0.051 ns ns 0.052 0.026
Ccsi41 0.266 0.275 270.1 278.4 1.71 1.76 250.2 248.2 0.26 0.29 1.25 1.38
Ccsi23 0.258 0.273 251.6 256.3 1.54 1.55 255.6 254.1 0.26 0.28 1.12 1.19
Significance ns ns 0.011 <0.01 0.022 0.037 0.194 <0.01 ns ns 0.043 0.019
Ccsi45 0.264 0.277 278.6 281.3 1.62 1.64 250.9 249.6 0.27 0.28 1.29 1.45
Jimian7 0.261 0.275 264.1 266.5 1.52 1.53 259.2 256.7 0.27 0.28 1.15 1.21
Significance ns ns <0.01 <0.01 0.039 0.052 <0.01 <0.01 Ns ns 0.024 <0.01
† ns, not significant at the 0.05 level.
There were significant differences in the increase of number of Effect of Nitrogen Regulation on Vegetative and
fruiting nodes and retained boll rates between the Bt cultivars and Reproductive Organ Growth and Development
their corresponding parents (Table 4). Sikang1 had 0.15 and 0.20 Higher total N contents were detected for the Bt cultivars
fruiting nodes per day more than Simian3, Ccsi41 had 0.13 and in the main stem leaves during square and boll development
0.08 more fruiting nodes per day than Ccsi23, and Ccsi45 had periods. The differences were significant between Bt cultivars
0.11 and 0.09 more fruiting nodes per day than Jimian7 in 2011 and their corresponding parents, with 0.25 to 0.39% difference
and 2012, respectively. Correspondingly, Sikang1 had 0.12 and between Sikang1 and Simian3, 0.27 to 0.55% difference between
0.14 more bolls per day than Simian3, Ccsi41 had 0.08 and 0.09 Ccsi41 and Ccsi23, and 0.26 to 0.55% difference between
more bolls per day than Ccsi23, and Ccsi45 had 0.09 and 0.08 Ccsi45 and Jimian7 in 2011 and 2012, respectively. However,
more bolls per day than Jimian7 in 2011 and 2012, respectively. the total N content in sympodium leaves showed a different
pattern than that in main stem leaves, with Bt cultivars having
Table 4. The development rate of reproductive organs in Bt cot- significantly lower N contents (Table 5).
ton cultivars and their corresponding parents in the 2011 and Nitrogen application data showed that vegetative organ growth
2012 cotton growing seasons. was different under N deficit (0 kg/ha) and under N-rich conditions
Retained fruiting Retained (450 kg/ha) between the Bt cultivars and their corresponding
node rate boll rate parents. Under N-rich conditions (Table 6), no significant
Cultivar 2011 2012 2011 2012 differences were detected for main stem leaves and leaf area between
—— nodes/d —— —— bolls/d —— Bt cultivars and their corresponding parents. However, under an
Sikang1 1.18 1.25 0.53 0.57 N deficit, lower numbers of main stem leaves and leaf areas were
Simian3 1.03 1.05 0.41 0.43 detected in Bt cultivars than in their corresponding parents. The leaf
Significance <0.01 <0.01 <0.01 <0.01 number was reduced by 2.2, 2.6, and 2.3 leaves for Sikang1, Ccsi41,
Ccsi41 1.15 1.17 0.47 0.50 and Ccsi45, respectively, and the leaf area was decreased by 84, 108,
Ccsi23 1.02 1.09 0.39 0.41 and 141 cm2 for Sikang1, Ccsi41, and Ccsi45, respectively.
Significance 0.022 0.035 0.023 0.044 There were no significant differences in number of sympodium
Ccsi45 1.09 1.11 0.46 0.48 leaves and sympodium leaf area for Bt cultivars under the
Jimian7 0.98 1.02 0.37 0.40 450 kg/ha N treatment (Table 6). However, lower number of
Significance 0.039 0.043 0.052 0.047 sympodium leaves and leaf area were detected in Bt cultivars than
Table 5. Total N content in main stem and sympodium leaves in Bt cotton cultivars and their corresponding parents during the 2011 and
2012 cotton growing seasons.
Main stem leaf Sympodium leaf
25 June 15 July 5 Aug. 25 June 25 July 25 Aug.
Cultivar 2011 2012 2011 2012 2011 2012 2011 2012 2011 2012 2011 2012
––––––––––––––––––––––––––––––––––––––––––––––––––––––––– m/d –––––––––––––––––––––––––––––––––––––––––––––––––––––––––
Sikang1 3.75 3.82 3.66 3.78 3.43 3.61 3.43 3.51 3.56 3.64 2.73 2.83
Simian3 3.43 3.5 3.41 3.49 3.05 3.22 3.66 3.72 3.74 3.88 2.85 2.97
Significance 0.049 0.035 0.052 0.048 0.041 0.029 0.047 0.041 0.043 0.041 0.047 0.051
Ccsi41 3.83 3.89 3.72 3.76 3.51 3.65 3.58 3.67 3.68 3.79 2.87 2.91
Ccsi23 3.52 3.61 3.45 3.48 2.98 3.1 3.71 3.85 3.85 3.98 2.98 3.12
Significance 0.034 0.042 0.413 0.039 <0.01 0.016 0.051 0.052 <0.01 0.023 0.058 0.053
Ccsi45 3.96 4.02 3.84 3.91 3.76 3.84 3.67 3.75 3.75 3.86 2.93 2.98
Jimian7 3.61 3.67 3.58 3.63 3.15 3.29 3.83 3.98 3.89 4.01 3.15 3.27
Significance 0.019 0.021 0.022 0.024 0.023 0.018 0.064 0.052 0.055 0.013 0.032 0.024
in the corresponding parents under N deficit: the fruiting branch Nitrogen rates also had different effects on the reproductive
leaf number reduced by 4.1, 3.1, and 3.2 leaves for Sikang1, Ccsi41, organ development of the Bt cultivars and their corresponding
and Ccsi45, respectively, and the leaf area decreased by 47, 83, and parents under N-deficit (0 kg/ha) and -rich (450 kg/ha)
73 cm2 for Sikang1, Ccsi41, and Ccsi45, respectively. conditions. Under the N-rich condition, the FP of Bt cultivars
Main stem nodes of the Bt cultivars were similar to those of the and their corresponding parents were not significantly
corresponding parents under the N-rich treatment (450 kg/ha) different. However, under N deficit, the Bt cultivars had a
(Table 6), but lower main stem nodes were detected in Bt cultivars higher FP than their corresponding parents.
under N deficit. Similarly, plant heights were not significantly The fruiting nodes, fruiting branches, bolls, and retained
different between Bt cultivars and their corresponding parents boll rates of Bt cultivars were not significantly different than
under the N-rich treatment, but contrary results were observed for those of the corresponding parents under the N-rich treatment
Bt cultivars under N deficit. (Table 8). However, different results were measured for these
Larger total N contents were detected in bolls in Bt cultivars reproductive organs when Bt cultivars and their corresponding
during 2011 and 2012. The differences were significant between parents were compared under a N deficit. The fruiting nodes
Bt cultivars and their corresponding parents from peak flowering per plant were reduced by 3.4, 5.8, and 5.1 nodes , the fruiting
to boll opening, with a 0.26 to 0.43% increase between Sikang1 branches were decreased by 0.8, 0.9, and 1.7 layers per plant, the
and Simian3, a 0.36 to 0.51% increase between Ccsi41 and retained bolls were reduced by 1.5, 1.2, and 1.1 bolls per plant,
Ccsi23, and a 0.44 to 0.65% increase between Ccsi45 and and the retained boll rate was decreased by 2.4, 2.1, and 1.9%
Jimian7 in 2011 and 2012, respectively (Table 7). for Sikang1, Ccsi41, and Ccsi45, respectively.
Table 7. Total N in the boll after >10 d past flowering in Bt cotton cultivars and their corresponding parents during the 2011 and 2012 cot-
ton growing seasons.
30 July 15 Aug. 30 Aug.
Cultivar 2011 2012 2011 2012 2011 2012
––––––––––––––––––––––––––––––––––––––––––––––––––– m/d –––––––––––––––––––––––––––––––––––––––––––––––––––
Sikang1 3.56 3.62 3.41 3.5 3.29 3.35
Simian3 3.27 3.3 3.15 3.19 2.89 2.92
Significance 0.024 0.015 0.015 <0.01 0.037 0.023
Ccsi41 3.63 3.7 3.59 3.62 3.35 3.38
Ccsi23 3.21 3.25 3.08 3.11 2.93 3.02
Significance 0.016 0.022 <0.01 0.012 0.026 0.036
Ccsi45 3.75 3.79 3.64 3.69 3.42 3.5
Jimian7 3.19 3.25 2.98 3.12 2.98 3.05
Significance <0.01 <0.01 <0.01 <0.01 0.021 0.027