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Detasseling increases kernel number in maize under shade stress

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 Agricultural and Forest Meteorology 280 (2020) 107811

Contents lists available at ScienceDirect

Agricultural and Forest Meteorology

journal homepage: www.elsevier.com/locate/agrformet

Detasseling increases kernel number in maize under shade stress T

a,b,1 a,1 c,1 a a a a
Zhen Gao , Lu Sun , Jian-Hong Ren , Xiao-Gui Liang , Si Shen , Shan Lin , Xue Zhao ,
⁎
Xian-Min Chena, Gong Wua, Shun-Li Zhoua,d,
a
College of Agronomy and Biotechnology, China Agricultural University, Beijing 100193, China
b
College of Agronomy, Hebei Agricultural University/ Key Laboratory of Crop Growth Regulation of Hebei Province, Baoding, Hebei 071001, China
c
College of Resources and Environmental Sciences, China Agricultural University, Beijing, 100193, China
d
Scientific Observing and Experimental Station of Wuqiao for Crop Water Use Efficiency, the Ministry of Agriculture and Rural Affairs, Wuqiao 061802, China

A R T I C LE I N FO A B S T R A C T

Keywords: Kernel number is one of the critical components of maize yield and is sensitive to environment variation around
Tassel removal tasseling when the tassel, stem and ear grow simultaneously. Similar to other areas in the world, the North China
Shading Plain faced reduced irradiance during the maize growing season, especially the critical window for determi-
Apical dominance nating actual kernel number. Shade stress that occurs between 15 days pre-silking and 15 days post-silking
Kernel number
significantly reduces kernel number and grain yield. In the present experiment, detasseling was conducted under
Assimilates
70% and 97% shade treatments to mitigate the environment stress at the 14-leaf (V14) and tasseling (VT) stages,
respectively. Shading treatments significantly reduced kernel number per plant, especially when shading oc-
curred at VT stage. Tassel removal did not relieve kernel losses under 97% shade, but detasseling did drama-
tically reverse the kernel losses under 70% shade. Removing tassel sink at V14 and VT promoted assimilates
being apportioned to ear sink under 70% shade stress, thus encouraging more carbohydrates to be made
available for ear growth. Consequently, detassseling increased soluble sugar concentration in the ear section,
shortened anthesis-tasseling interval, accelerated ear growth, offering a method to improve kernel number under
shade stress.

1. Introduction previous researches have demonstrated that abiotic stress-induced

Maize yield, as most crops, is mainly related to harvested kernel
number and, to a lesser extent, kernel weight, which is generally re-
cognized as a component not intensively influenced by agronomic and
environmental variables (Andrade et al., 1999; Borrás et al., 2004;
Otegui et al., 1995). Mechanisms involved in kernel number have been
broadly researched through approaches focused on individual plant and
crop levels (Andrade et al., 1999, 2002). It is currently accepted that
kernel number is largely determined by biomass accumulation during
the critical period bracketing silking (from 15 d presilking to 15 d
postsilking), and the proportion of crop growth that is partitioned to
reproductive organs during this period (Andrade et al., 1993;
Borrás and Vitantonio-Mazzini, 2018; Charles-Edwards, 1984;
Fischer, 1985).
Stress factors such as heat, drought and reduced irradiance reduce
kernel number and grain yield in maize (Cheikh and Jones, 1994;
Fedoroff et al., 2010; Schmidhuber and Tubiello, 2007). However,
kernel losses in maize can be reduced by feeding sucrose to the stem
(Boyle et al., 1991a, b; Hiyane et al., 2010; McLaughlin and
Boyer, 2004; Zinselmeier et al., 1995, 1999). The central role of sucrose
in this restoration raised the possibility that increasing assimilates
could improve kernel number in maize under stressful environments.
Solar radiation showed a significant decreasing trend over the last
several decades in the North China Plain (Fig. 1; Huang et al., 2018;
Wang et al., 2012). Moreover, local precipitation is mainly con-
centrated in July and August, when maize silking stage occurs
(Gao et al., 2017). Both factors limit photosynthesis and plant growth
rate during the critical period bracketing silking in maize, which would
lead to kernel number and yield decline. As demonstrated by foregoing
research results, sucrose feeding could reverse shade-induced kernel
abortion in maize. As a consequence, increasing photosynthates ap-
portioned to reproductive organs might play a dominant role in im-
proving kernel number under stress conditions. However, it is im-
possible to conduct assimilates feeding to stems at the scale of field

⁎
Corresponding author. College of Agronomy and Biotechnology, China Agricultural University, No.2, Yuanmingyuan West Road, Haidian District, Beijing,
100193, China.
E-mail address: zhoushl@cau.edu.cn (S.-L. Zhou).
1
These authors contributed equally to this work.

https://doi.org/10.1016/j.agrformet.2019.107811
Received 20 February 2019; Received in revised form 7 October 2019; Accepted 13 October 2019
Available online 23 October 2019
0168-1923/ © 2019 Elsevier B.V. All rights reserved.
Z. Gao, et al. Agricultural and Forest Meteorology 280 (2020) 107811

Table 2
Yield components affected by shading and detasseling treatments.
Year Treatment Treatment Ear Number of Kernels Kernels
stage length kernel per row per ear
(cm) rows

2011 V14 CK 17.9a 14.0a 35.0a 490.0a

S1 11.7b 14.7a 18.5b 271.0b
S2 14.1b 12.7b 24.0b 304.7b
VT CK 17.9a 14.0a 35.0a 490.0a
S1 14.2b – – 246.3b
S2 13.1b – – 132.0c
2012 V14 CK 15.5a 14.9a 32.6a 484.7a
S1 12.9b 14.4a 22.9b 333.8b
S1Det 15.0a 14.5a 31.3a 454.3a
S2 11.8c 14.2a 16.3c 237.0c
S2Det 11.0c 14.1a 15.3c 220.2c
VT CK 15.5a 14.9a 32.6a 484.7a
S1 11.3b 14.3a 21.3c 273.0c
S1Det 13.0a 14.9a 26.5b 371.9b
S2 11.4b 13.4b 16.1d 154.8d
Fig. 1. Mean solar radiation variation from 1956 to 2016 at the experimental
S2Det 10.6b 12.5b 14.3d 168.3d
station. 2017 V14 CK 16.9a 15.4b 31.4a 483.6a
Det 16.5a 15.5a 31.9a 494.6a
S1 14.1c 14.3c 24.1b 344.5b
maize cultivation. How can we stimulate assimilates to be apportioned
S1Det 15.1b 16.3a 31.6a 513.9a
to ear in maize under stressful environment? VT CK 16.9b 15.4a 31.4b 483.6b
Maize is a monoecious plant with separate male and female flowers Det 18.3a 15.5a 34.0a 527.0a
on the identical plant. During the early stages of generative growth in S1 10.0d 14.2b 16.9d 239.2d
maize, tassel growth competes with ear growth for assimilates and S1Det 13.9c 14.4b 26.8c 385.9c

nutrients, especially under stressful environments when resources are

restricted (Fox et al., 2017). Notably, detasseling displayed a positive
trend in grain yield, especially under stressful conditions (Cabrera-
Bosquet et al., 2017; Moreira et al., 2010; Mostert and Marais, 1982).
However, the physical removal of tassel in the female inbred parent to
produce pure hybrid seed would decrease hybrid seeds yield because
the top leaves are destroyed during emasculation (Skibbeand
Schnable, 2005). In our previous result, detasseling was shown to re-
verse kernel losses caused by shading pre-anthesis and post-anthesis in
maize (Sun, 2013). We hypothesize that, under shade conditions, de-
tasseling removes tassel apical dominance, thereby promoting more
assimilates to be partitioned to ear for kernel formation.
2. Materials and methods
2.1. Experimental site
Field experiment was conducted in 2011, 2012 and 2017 at the
Wuqiao Experimental Station of China Agricultural University (Hebei
Province, China, 116.3°E, 37.4°N). The soil at this experimental field is
a light loam with 12.2 g kg−1 organic matter, 0.90 g kg−1 total N,
84.0 mg kg−1 available potassium and 46.2 mg kg−1 available phos-
phorus. Soil chemical data were measured in the upper 0.2 m of soil at
the beginning of the field experiment in 2011. Maize is sown in early
June and harvested in early October in this region. Rainfall is mainly
concentrated in July and August and solar radiation is presented in
The row number and kernel number per row in 2011 were irregular and the
total kernels were recorded. Different letters in each stage mean significant at
P < 0.05. CK, unshaded condition; S1, 70% of solar radiation was intercepted;
S2, 97% of solar radiation was intercepted; Det, detasseling under unshaded
condition; S1Det, S1+ detasseling; S2Det, S2+ detasseling.
Table S.1.
2.2. Treatment and experimental design
Three shade treatments were set up: unshaded control (CK); 70%
shading (S1, which referred to the meteorological data as shown in
Table S.1); and 97% shading (S2). Shade treatments were conducted at
the 14-leaf (V14) stage and tasseling (VT) stage for 6 days, respectively.
At V14 stage, two rows of maize tassels were manually pulled out in
each plot; at VT stage, two rows of maize tassels were cut from the
bottom in each plot when shading started. This experiment was laid out
as split block design with three replicates. Shade experiment in 2011
proved that shade reduced photosynthesis and kernel number; de-
tasseling was only conducted in shade treatments, which revealed that
tassel removal could mitigate kernel abortion under moderate shade
stress in 2012; in 2017, S2 was discontinued owing to its strong in-
hibition, and detasseling treatments were conducted in unshaded and
shaded treatments. Scaffolding and black polypropylene fabric was
employed to build a detachable shed that was 3.5 m high, 10 m long
and 7 m wide. The fabric was 2 m longer at periphery to intercept the

Table 1
Effects of different shading treatments on the light intensity (μmol m−2 s−1) on the maize canopy.
Treatment Measuring Light intensity Percent of intercepted radiation (%)

position 8:00 11:30 14:30 18:00 8:00 11:30 14:30 18:00

CK Top 651.0 1061.7 963.6 865.5 0.0 0.0 0.0 0.0

Ear 117.7 402.8 331.5 260.2 81.9 62.1 65.6 69.9
S1 Top 192.3 320.3 264.2 280.0 70.5 69.8 72.6 67.6
Ear 61.3 103.8 75.6 89.9 68.1 67.6 71.4 67.9
S2 Top 20.0 22.3 21.2 21.2 96.9 97.9 97.8 97.6
Ear 7.1 8.7 7.9 6.8 64.8 61.1 62.8 67.7

CK, unshaded condition; S1, 70% of solar radiation was intercepted at the top position of maize canopy; S2, 97% of solar radiation was intercepted at the top position
of maize canopy.

2
Z. Gao, et al. Agricultural and Forest Meteorology 280 (2020) 107811

Fig. 2. Harvested ears at maturity of different treatments in 2012 (A) and 2017 (B). The top figures indicate treatments at the 14-leaf stage (V14), and the bottom
figures indicate treatments at tasseling (VT).

Fig. 3. Photosynthetic rate of ear leaf as affected by shading treatments at V14 (A) and tasseling (B) in 2012. S1, 70% of solar radiation was intercepted; S2, 97% of
solar radiation was intercepted. Data are means, with error bars showing the standard error of mean (SEM, n = 3).

slanting sunlight.
The commonly planted summer maize variety, Zhengdan958, was
sown manually at a density of 75,000 plants ha−1 with a row spacing of
0.6 m on June 19, 2011, June 15, 2012 and June 17, 2017. After
sowing, 70 mm of irrigation water was immediately applied to guar-
antee germination using the surface flood method. During all the
growing seasons, basal fertilizer was applied at the rate of 72 kg N ha−1,
105 kg P2O5 ha−1 and 105 kg K2O ha−1 with sowing, and
108 kg N ha−1 was top-dressed at the 12-leaf (V12) stage. Pest, disease
and weed management followed common agricultural practice. Maize
was harvested on October 11, October 6, and October 2 in 2011, 2012,
and 2017, respectively.
2.3. Sampling and measurements
2.3.1. Light intensity and photosynthesis
Light intensity was determined at 8:00, 11:30, 14:30 and 18:00
during shading at the top and ear positions for three times in each plot
by using LI-250A Light Meter (Li-Cor Biosciences, Lincoln, Nebraska,
USA). Net photosynthetic rate (Pn) of ear leaf (n = 3) was measured
using a Li-Cor 6400 (Li-Cor Biosciences) under ambient conditions at
9:30–11:30 on 2, 4, 6 and 8 days after shading.
2.3.2. Dry matter accumulation
Three representative plants were sampled per plot on 2, 4, 6 and 8
days after shading in 2011 and 2012. Maize plants were taken every 10
days from silking in 2017. Maize was divided into the stem, leaf, ear to
be dried at 80 °C for 48 h to a constant weight in 2011 and 2012. In
2017, maize stem was divided into upper stem, ear section (ear inter-
node as well as the internodes above and below ear) and lower stem.
2.3.3. Phenology
Twenty successive plants were selected, and their dates of tasseling
(50% of plants with tassels), anthesis (50% of main tassel branches with

3
Z. Gao, et al. Agricultural and Forest Meteorology 280 (2020) 107811

Fig. 4. Ear biomass affected by treatments at tasseling (2011and 2012, A) and on 10 days after silking (2017, B) in summer maize. CK, unshaded condition; Det,
detasseling under unshaded condition; S1det, 70% shade + detasseling. Data are means, with error bars showing SEM (n = 3), and the same letter above bar means
not significantly different at P < 0.05.

Table 3 2.3.4. Carbohydrates and nitrogen analysis

The anthesis-silking interval (ASI) as affected by the shading and detasseling Oven-dried organs were milled for 5 min using a LAIFU mill (LTP-
treatments in summer maize in 2017. 400A, Ningbo, China) and particle size was less than 0.3 mm. Amounts
Treatment Shading at V14 Shading at VT of 0.2 g dry matter were extracted with distilled water, and the ex-
traction solution was prepared for soluble carbohydrate determination.
ASI (days) ASI (days) Insoluble debris was extracted with HCl (3 M), and the extraction so-
CK 1c 1a
lution was used in starch analysis. Starch and soluble sugars were
CKDet 1c 1a
S1 7a 1a measured with anthranone-H2SO4 using a UV–vis spectrophotometer
S1Det 2b 1a (TU-1901, PERSEE, China) at a wavelength of 625 nm. The total N
content was determined using the Kjeldahl method (Dordas and
CK, normal plant under unshaded condition; CKDet, CK+ detasseling; S1, 70% Sioulas, 2009).
of solar radiation was intercepted; S1Det, S1+ detasseling. Different letters in
each column mean significant at P < 0.05.
2.3.5. Yield components
Six square meters of maize (2 rows × 5 m row length) were hand
pollen), and silking (50% of plants with visible silks) were recorded.
harvested per plot at maturity. All harvested areas were surrounded by
Anthesis-silking interval (ASI) was calculated as the interval between
at least two guard rows. All the ears of detasseling treatments were
anthesis date and silking date. At V14, VT and anthesis (pollen shedding
harvested. Ear length and kernel number per ear were measured on 20
period, PS), tassels of normally growing plants were collected and dried
representative ears per treatment.
at 80 °C for 48 h to a constant weight to evaluate the allocation of
carbohydrates and nitrogen to tassel.
2.4. Statistical analyses

Statistical analyses were carried out with SPSS 17.0 (SPSS Inc.,

4
Z. Gao, et al.
Fig. 5. The biomass, sugar and nitrogen content in the tassel at different stages in 2017. V14, 14-leaf stage; VT, tasseling stage; PS, pollen shedding stage. Values
plotted are means, with error bars showing SEM (n = 3), and the different letters indicate significantly different at P < 0.05.
Chicago, IL, USA). Significant differences were determined using the
least significant differences test at a 0.05 probability level. All figures
were created using SigmaPlot 12.5 (Systat Software Inc., San Jose, CA,
USA).
3. Results
3.1. Effects of detasseling on kernel set under shading
Table 1 summarizes light intensity and intercepted radiation under
different shade treatments. Treatment S1 intercepted 68–73% of solar
radiation while nearly 100% of radiation was blocked under S2. As
shown in Table 2 and Fig. 2, all of the shading treatments at the V14
and VT significantly reduced ear length. However, detasseling at V14
and VT did reverse the decline to some extent under S1. Under S2 even
detasseling did not change the ear length reduction at either the V14 or
VT. At the V14 stage treatment, the kernel number of S1 was reduced
by 28.7–44.7% compared with the CK in the three years. The S2
treatment decreased kernel number by 37.8% and 51.1% than CK in
2011 and 2012, respectively. At the VT stage treatment, kernel number
of S1 was 43.7–50.5% lower than that of CK across the three years. S2
treatment significantly decreased kernel number by 70.1% and 68.1%
compared with that of CK in 2011 and 2012, respectively. Under
moderate shade condition (S1), detasseling treatments at V14 increased
kernel number by 36.1% and 49.2% compared to S1 in 2012 and 2017,
respectively; detasselig at VT elevated kernel number by 36.2% and
61.3% compared to S1 in 2012 and 2017, respectively. However, the
removal of tassel did not mitigate kernel abortion caused by S2 treat-
ment. In addition, under unshaded condition, detasseling at V14 did not
significantly increase kernel number, perhaps owing to the removal of
2–3 leaves that occurred with detasseling. However, detasseling at VT
significantly increase kernel number compared with the control.
5
Agricultural and Forest Meteorology 280 (2020) 107811
3.2. Effects of shading on ear leaf photosynthesis and ear biomass
Shade stress inhibited net photosynthesis of ear leaf, especially
under S2 treatment. After removing shade, net photosynthesis on day 2
recovered completely to the control rate observed in S1 plants.
However, a wide gap still existed between CK and S2. The shading
treatment at V14 and VT had similar effects on photosynthesis (Fig. 3).
With the decline in photosynthesis, stem biomass under S1 and S2
decreased owing to respiration. Moreover, ear biomass remained low
under shading. Naturally, ear rapidly grew without shading (Fig. 4A).
As indicated in Fig. 4B, shading at V14 significantly reduced ear bio-
mass. However, detasseling obviously reversed ear growth. Similarly,
the inhibition to ear growth was also serious under VT shading, and
tassel removal significantly increased ear biomass. Moreover, shading
stress in the most sensitive stage (VT) directly suppressed the devel-
opment of the middle and apical silks, which resulted in more ovaries
abortion compared to shading at V14 (Fig. 2).
3.3. Relationship between tassel and ear
ASI greatly increased up to 7 days under shading; however, de-
tasseling significantly decreased ASI by 5 days at V14 (Table 3). In
addition, shading at VT did not change ASI but obviously inhibited
middle and upper silks growth and prevented ear biomass accumula-
tion, which resulted in more ovaries abortion compared to the shade
stress at V14.
The biomass, total N content, starch content, and soluble sugar
content of tassel were measured at V14, VT and PS (Fig. 5). The tassels
at PS did not exhibit a significant difference in biomass, N content or
starch compared to those at VT, but the soluble sugar content at PS was
higher than that of VT. In addition, all of the indexes at VT were sig-
nificantly higher compared to those at V14. These results suggested that
Z. Gao, et al.
Fig. 6. The soluble sugar in the lower portion (A, B), ear portion (C, D) and upper portion (E, F) of stem affected by shading and detasseling at V14 (left) and VT
(right) in 2017. CK, normal plant under unshaded condition; CKdet, CK+ detasseling; Sdet, 70% shade+ detasseling. The upper stem of detasseling treatment at V14
was removed. Values plotted are means, with error bars showing SEM (n = 3).
more N and carbohydrates shifted from tassel to other growth center
under detasseling at V14. Additionally, detasseling at VT can only save
a part of soluble sugar for the growth of other organs (e.g. ear, stem).
As shown in Fig. 6, shading significantly reduced the soluble sugar
concentration at silking, especially in the ear section; however, de-
tasseling under shade significantly increased the soluble sugar con-
centration of ear portion at silking, which increased photosynthate
availability for ear growth. With the kernel abortion, assimilates could
not be used and accumulated rapidly in the stem. As described above
(Table 2), the kernel number in the S1+detasseling (Sdet) treatment at
V14 exhibited no differences from the CK. The soluble sugar trans-
ported to the ear and did not accumulate in the stem under Sdet
treatment at V14. In contrast, Sdet treatment at tasseling had fewer
kernels, which resulted in more soluble sugar stored in the stem com-
pared with CK plants. Moreover, the correlation analysis suggested that
higher soluble sugar concentration in the ear section stem, more kernels
could be fed (Fig. 7), i.e. detasseling under shade could promote pho-
tosynthates being allocated to ear at sensitive stage, thereby increasing
kernel number.
6
Agricultural and Forest Meteorology 280 (2020) 107811
4. Discussion
4.1. Apical dominance in maize
In maize, tassel, stem and ear organs act as sinks for photosynthates
synchronously at flowering. Accordingly, each of these sinks would
compete for assimilations during this sensitive period. Apical dom-
inance resulted in protandry, which decreased allocation of assimilates
to ear, favoring tassel development (Monneveux et al., 2008). In the
present experiment, apical dominance of maize under shading treat-
ment at V14 delayed silks growth. After tasseling, apical dominance
was still manifested owing to shade avoidance and inhibited ear
growth. Although the silking date was not delayed by shading,
abounding apical and middle silks were inhibited. Detasseling elimi-
nated the apical dominance of tassel and thus increased the soluble
sugar concentration of ear section stem at silking (Fig. 6). Conse-
quently, ear growth was accelerated and kernel number was greatly
increased by tassel removal. Hunter et al. (1973) and Cabrera-
Bosquet et al. (2017) also found that detasseling caused a visible
Z. Gao, et al.
Fig. 7. Relationship between the soluble sugar at silking in the ear section and
the kernel number per plant. Red and green circles mean treatment at 14-leaf
stage (V14) and tasseling (VT), respectively. CK, normal plant under unshaded
condition; Det, detasseling under unshaded condition; Shade+Det, detasseling
under shade treatment.
increase in grain yield under stressful environments. On the other hand,
several research results suggested that the positive effect of detasseling
on yield just before anthesis is not caused by altered apical dominance
but rather by reduced shading of tassels, which increased light inter-
ception by the top leaves especially under high plant densities
(Duncan et al., 1967; Hunter et al., 1969). However, modern temperate
maize has reached progressively smaller tassel size along with the in-
direct selection by maize breeders over the past 50 years (Duvick and
Cassman, 1999). Therefore, we concluded that apical dominance
played a critical role in dominating ear growth. Detasseling could
eliminate the apical dominance, thus promoting assimilates being
partitioned to the ear and facilitating ear growth, especially under
stressful conditions. Additionally, when growth was focused on ear and
other organs (leaf and stem) did not compete with ear for assimilates,
appropriate stress could increase the remobilization of carbohydrates
from vegetative organs to kernel (Yang and Zhang, 2006). Similarly, in
this detasseling experiment, assimilates were more easily transported to
the sole growth center (i.e. ear).
7
Agricultural and Forest Meteorology 280 (2020) 107811
4.2. The role of assimilates in kernel set
Shade stress usually lowered leaf photosynthesis and sugars avail-
ability, which resulted in kernel abortion (Hiyane et al., 2010). Our
results also demonstrated that shading treatments reduced ear leaf
photosynthesis rate and diminished soluble sugar concentration of
stem, which resulted in longer ASI and lower kernel number compared
to control plants. Moreover, although shading at VT did not change the
ASI, stress did inhibit amount of apical and middle silks growth, re-
sulting in more ovaries abortion compared with shade treatment at
V14. However, detasseling under the moderate shading (S1) did con-
spicuously increase soluble sugar in the ear section stem at silking.
Consequently, ASI was shortened and the kernel number was reversed
partially. Additionally, the carbon and nitrogen content of tassel in-
dicated that detasseling at V14 saved a few sugars and nitrogen for ear;
however, the assimilates of tassel at VT reached the peak, i.e. de-
tasseling at VT mainly changed the assimilates allocation for ear.
Slewinski (2012) also indicated that manipulating the allocation of
stem non-structural carbohydrates provided one avenue for stabilizing
and increasing productivity. The stored redundancy carbohydrates in
crop stem to buffer source–sink interactions during different periods
under changing environments (Ruuska et al., 2006; Schnyder, 1993). In
maize, buffering confers an ability to mitigate the detrimental effects of
seasonal instability on maize yields in unpredictable regions
(Shiferaw et al., 2011). The assimilate reserves in stem have been
shown to contribute little to the final grain weight under normal field
conditions (Cliquet et al., 1990). However, during the critical stage
when maize was sensitive to environmental stress, the maize suffered
the brunt of lack of assimilations caused by varying environmental
stressors. Then the stem assimilation reserves could play a critical role
in kernel set and yield stability (Setter et al., 2001; Setter and
Meller, 1984). All of the studies proved that the assimilates in stem
were essential for kernel setting, which were consistent with our results.
Additionally, even though sink activity was not measured in this ex-
periment, abiotic stress usually inhibited enzyme activity in kernel
(Hiyane et al. 2010; McLaughlin and Boyer, 2004). There were no
differences between shade+detasseling and shade treatment except
that detasseling changed the carbon allocation. Similarly, sucrose
feeding under stressful conditions could partially reverse the activity of
sucrose processing enzymes and recover kernel growth (Hiyane et al.
2010; McLaughlin and Boyer, 2004). Accordingly, we suggested that
allocating assimilates to ear determined the kernel number under
stressful environments. As shown in Fig. 8, the stem, tassel and ear sink
would compete for assimilations at the same time. When solar radiation
Fig. 8. The assimilate allocation was
affected by detasseling under shading
in maize. Under normal light radiation,
enough assimilate can simultaneously
support tassel and ear growth. Under
shade, apical dominance resulted in
assimilates being preferentially allo-
cated to tassel. However, the de-
tasseling under shading condition
eliminated the apical dominance and
the assimilates were partitioned to ear
for kernel setting.
Z. Gao, et al.
was sufficient, the photosynthate was able to sustain tassel and ear
growth together, resulting in high yield. In contrast, shade stress would
reduce assimilates availability, which resulted in tassel apical dom-
inance and assimilates were preferentially allocated to tassel. However,
detasseling under stress condition removed the apical dominance, and
thus more assimilates were apportioned to ear. Hence, kernel losses
could be reversed to some extent.
4.3. Strategy to maintain maize yield stability under stressful environments
The North China Plain (NCP), as other areas in the world, was hotter
and more prone to drought, also had continuous overcast and rainy
stress, especially during maize growing seasons. Such adverse en-
vironmental conditions have limited maize productivity of the NCP
(Gao et al., 2018). Low-yield maize varieties tend to have high sugar
concentration in the stem, which displayed yield stability even when
challenged by environmental stressors. In contrast, high-yield maize
varieties showed lower stem carbohydrate and were sensitive to
stressful environments (Van Reen and Singleton, 1952). To meet the
demand of increasing human populations, high-yield varieties have
become pivotal in modern agricultural systems. The allocation of as-
similates is an avenue to mitigate environmental stress and stabilize
yield productivity. As presented in our results and other studies
(Cabrera-Bosquet et al., 2017; Monneveux et al., 2008), detasseling
could increase grain yield under abiotic stress. Therefore, we suggested
detasseling could be employed in the field production when maize
suffered stressors around tasseling. To ensure pollination, interlaced
detasseling should be used, i.e. half of plants left to provide pollens
while the other half were detasseled to reverse kernels losses. However,
mechanical detasseling was labor intensive, time-consuming, and often
imperfect. Most importantly, nuclear genetic male sterility (i.e. Ms 44)
has displayed to be steady across different germplasms and various
environments (Fox et al., 2017). More assimilations and nitrogen
shifted from tassel to ear growth in male sterile maize plant compared
with wild type. The blends of Ms44 male sterile F1 hybrids and normal
hybrids could increase maize yield under stressful environments
(Fox et al., 2017), which could serve as a strategy to improve kernel
number and grain yield in response to stressful environments around
silking.
5. Conclusions
Our hypothesis that detasseling removed tassel apical dominance
hence more assimilates are partitioned to the ear for kernel setting was
confirmed under moderate shade condition, but under severe shade
condition the hypothesis was rejected. The removal of tassel could
mitigate shade stress and reverse shade induced kernel abortion. As a
consequence, we concluded that interlaced detasseling or nuclear ge-
netic male sterility could serve as ways to maintain yield stability under
shade stress in countless areas.
Declaration of Competing Interest
None.
Acknowledgements
This work was supported by the National Key Research and
Development Program of China (2016YFD0300301), the China
Agriculture Research System (CARS-02-13), National Science Fund of
Hebei Province (C2019204250), and the Startup Fund of Hebei
Agricultural University (YJ201827).
Supplementary materials
Supplementary material associated with this article can be found, in
8
Agricultural and Forest Meteorology 280 (2020) 107811
the online version, at doi: 10.1016/j.agrformet.2019.107811.
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