agricultural and forest meteorology 149 (2009) 51–58
available at www.sciencedirect.com
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Interactions of elevated [CO2] and night temperature
on rice growth and yield
Weiguo Cheng *, Hidemitsu Sakai, Kazuyuki Yagi, Toshihiro Hasegawa
National Institute for Agro-Environmental Sciences, Kannondai 3-1-3, Tsukuba, Ibaraki 305-8604, Japan
article info
abstract
Article history:
To understand how a combination of high night temperature and elevated [CO2] during the
Received 14 March 2008
reproductive growth period affects rice growth and yield, we conducted a pot experiment in
Received in revised form
four controlled-environment chambers with two levels of [CO2] (ambient, 380 ppm; and
11 July 2008
elevated, 680 ppm) and two levels of night temperature (22 and 32 8C). The day temperature
Accepted 12 July 2008
was 32 8C in all treatments. As results, the whole plant and stem dry weight were significantly increased by both elevated [CO2] and high night temperature (P < 0.01), while the
ear dry weight was significantly increased by elevated [CO2] and decreased by high night
Keywords:
temperature (both P < 0.01). The fertilized spikelet number was significantly decreased by
Dry weight
high night temperature (P < 0.01), but not affected by elevated [CO2] (P = 0.18). The individual
Elevated [CO2]
grain weight was significantly increased by elevated [CO2] (P = 0.03). Consequently, brown
Interaction
rice yield was significantly increased by elevated [CO2] (P < 0.01) but decreased by high night
Night temperature
temperature (P < 0.01), with a significant interaction of [CO2] and night temperature
Rice growth
(P < 0.05). The results indicate that high night temperature will reduce the stimulatory
Yield
effect of elevated [CO2] on rice production in the future if both continue to increase.
# 2008 Elsevier B.V. All rights reserved.
1.
Introduction
Depending on population growth and energy use scenarios,
atmospheric CO2 concentration ([CO2]) is expected to rise from
380 ppm currently to between 485 and 1000 ppm by 2100 (IPCC,
2001). The projected rise in the global average surface air
temperature at the end of the 21st Century relative to 1980–
1999 will be around 1.8–4.0 8C, with a likely range of 1.1–6.4 8C
(IPCC, 2007). The daily minimum (or night-time) temperature
has increased faster than the daily maximum (daytime)
temperature in the past century (Kukla and Karl, 1993;
Easterling et al., 1997).
Rice (Oryza sativa L.) is one of the most important crops in
the world and the most important food in Asia. Projected
changes in rice production in Asian countries have quite a
wide range, depending on the crop models and global climate
change scenarios (Matthews et al., 1997; Ohta and Kimura,
2007; Tao et al., 2008). Because both elevated [CO2] and air
temperature can have marked effects on rice growth and yield,
it is important to quantify these effects. A number of studies
have examined the effects of elevated atmospheric [CO2] or
combinations of elevated air temperature and [CO2] on rice
yield and growth during the last several decades (Yoshida,
1973a,b; Imai et al., 1985; Baker et al., 1992; Ziska et al., 1996;
Horie et al., 2000; Kim et al., 2003; Baker, 2004; Yang et al., 2006;
Sakai et al., 2006; Sasaki et al., 2007). Most results showed that
elevated [CO2] increased yield. Conversely, several studies
have shown that high air temperatures can reduce grain yield
even under CO2 enrichment (Baker et al., 1992; Ziska et al.,
1996; Matsui et al., 1997; Horie et al., 2000, Prasad et al., 2006)
owing to increased spikelet sterility (Satake and Yoshida, 1978;
Kim et al., 1996; Matsui et al., 1997; Ohe et al., 2007; Jagadish
* Corresponding author at: Agro-Meteorology Division, National Institute for Agro-Environmental Sciences, Kannondai 3-1-3, Tsukuba,
Ibaraki 305-8604, Japan. Tel.: +81 29 838 8205; fax: +81 29 838 8211.
E-mail address: [email protected] (W. Cheng).
0168-1923/$ – see front matter # 2008 Elsevier B.V. All rights reserved.
doi:10.1016/j.agrformet.2008.07.006
52
agricultural and forest meteorology 149 (2009) 51–58
et al., 2007). A correlation study showed a yield decline with
higher night temperatures due to global warming from 1979 to
2003 (Peng et al., 2004). However, no study has examined the
interactions of increased [CO2] and increased night air
temperature on rice growth and yield, especially during rice
reproductive stage. A theoretical analysis of the effect of day
temperature on photosynthesis indicated that the enhancement of photosynthesis due to elevated [CO2] is larger at
higher temperature (Long, 1991; Morison and Lawlor, 1999).
However, increased night temperature can counter this
because of the heavier respiratory cost, increased by the
larger biomass. Therefore, we hypothesized that an increased
night temperature would reduce the stimulatory effect of
elevated [CO2] on both rice biomass and grain yield. We tested
this hypothesis by examining growth, respiration, yield and
yield components of rice grown at two CO2 concentrations and
two night air temperatures during the reproductive stage. Our
objective was to understand how rice growth, fertilized
spikelet percentage and yield respond to increased [CO2]
and high night temperature and their interaction.
2.
thickness) filled with 7.4 kg grey sandy soil. The plants growing
in each pot was defined as one hill for the whole rice growing
period. The sandy soil was collected from the plough layer
(15 cm of the top layer) of a rice field in Kujukuri, Chiba
Prefecture, Japan, that contained 8.1 g kg1 organic C and
0.9 g kg1 total N, and the average pH was 6.3. Before
transplanting, 9.0 kg (7.4 kg dry soil equivalent) of soil was
mixed with 1.53 g NH4Cl and 0.87 g KH2PO4 and placed in each
pot. The NPK levels were 0.40:0.20:0.25 g per pot. At 35 and 49
days after transplanting (DAT), we top-dressed with
0.62:0.10:0.13 g per pot. The water depth was maintained at
about 5 cm throughout the cultivation period.
We applied the treatments during the reproductive stage
only, because the climatic factors, such as high or low
temperatures during this stage strongly affect rice yield
(Matsui et al., 1997). Treatments were started at 59 DAT, when
three pots were randomly moved to each of four controlledenvironment chambers (see below) and plants in the other
three pots were destructively sampled. Grain was harvested at
107 and 114 DAT in the high and low night temperature
treatments, respectively, depending on the physiological
maturity.
Materials and methods
2.1.
Experimental schedule, cultural practices, rice cultivar
and soil
This research was conducted at the National Institute for
Agro-Environmental Sciences, Tsukuba, Japan (368010 N,
1408070 E). Seedlings were raised in an outdoor water tank
and then grown in controlled-environment chambers during
the reproductive growth stage (Fig. 1). We grew a semi-dwarf
rice, cv. IR72, an indica cultivar popularly grown in South-East
Asia (Peng et al., 1999). On 5 June 2006, germinated seeds were
sown in a seedling tray (three seeds per cell). At 3 weeks after
sowing, the seedlings (three plants) were transplanted to 15
plastic pots (19.5 cm inside diameter, 27.0 cm height, 0.2 cm
2.2.
Experimental design and controlled-environment
chambers
We used four controlled-environment chambers (Climatrons,
made by Shimadzu, Kyoto, Japan) to apply the two [CO2] and
two night temperature treatments. Each chamber measured
4 m 2 m 2 m (L W H). Each housed two stainless-steel
containers (1.5 m 1.5 m 0.3 m; L W D) filled with water
to hold the pots. We have used these chambers since 1996 to
carry out elevated CO2 experiments with rice (Cheng et al.,
2001, 2006; Sakai et al., 2001, 2006). Because there were no
chamber replicates (i.e., true replicates) in this study, we
rotated the treatments among the four chambers every 3
weeks (changed the environmental settings and moved the
Fig. 1 – Experimental schedule, changes in daily average air temperature (before treatment in outdoor water tank), and daily
air temperature variation (during treatment in controlled-environment chambers).
agricultural and forest meteorology 149 (2009) 51–58
pots so that the pots received the same treatment) during the
treatment period, so as to minimize variation due to the
controlling system.
The four treatments and their abbreviations were as
follows:
(1) Elevated [CO2] at 680 ppm and High night temperature
32 8C (EH);
(2) Ambient [CO2] at 380 ppm and High night temperature
32 8C (AH);
(3) Elevated [CO2] at 680 ppm and Low night temperature
22 8C (EL);
(4) Ambient [CO2] at 380 ppm and Low night temperature
22 8C (AL).
at
at
at
at
As implied in Fig. 1, the temperature was maintained at a
constant 32 8C in the high night air temperature treatment. In
the low night air temperature treatment, it was maintained at
32 8C from 08:00 h until 16:00 h, decreased at 2.5 8C h1 to
20:00 h, maintained at 22 8C from 20:00 h until 04:00 h, then
increased at 2.5 8C h1 to 08:00 h. Air and soil (at 10 cm depth)
temperatures during 9–11 September 2006 in both temperature treatments were recorded every 10 min by a Thermo
Recorder (TR-71U, T&D Corp., Tokyo, Japan) (Fig. 2). The actual
average air temperatures were 30.3 and 26.5 8C, respectively;
and the soil temperatures were 28.4 and 24.2 8C for high and
low night temperature treatments, respectively. The differences of daily average air and soil temperatures between the
high and low night temperature treatments were 3.8 and
4.2 8C, respectively, close to values predicted in global climate
change scenarios at the end of the 21st century (IPCC, 2007).
53
2.3.
Growth, destructive sampling and yield
measurements
Plant height and tiller number were measured once a week. At
the beginning of treatment, 59 DAT and final harvest, we
collected plants from three pots, carefully washed the soil
from the roots in running water, then separated the plants into
stems, leaves, roots and ears (final harvest only). The parts
were then oven-dried at 80 8C for 3 days and weighed.
To measure yield, we air-dried the ears for 4 weeks,
counted the number of panicles and carefully threshed the
grain. Grain was soaked in tap water and the numbers of
sunken and floating grains were counted to determine the
grain-filling rate. The dry weight of sunken grain was
determined after drying at 80 8C in a forced-air oven for 1
week. Gross yield was defined as the dry weight of filled grain.
After hulling, the weight of the edible portion was defined as
the brown rice yield (expressed on the basis of 15% moisture
content). The floating grain was used to determine grain
fertility by iodine reaction with starch (Matsushima and
Tanaka, 1960). Harvest index (HI) was calculated from the
gross yield and the plant total dry weight.
2.4.
Night-time respiration measurements
Night respiration was measured as the CO2 flux in a cylindrical
closed-top flux chamber (20.5 cm inside diameter, 100.0 cm
high, 0.3 cm thick) each week during the treatment period at
around 22:00 h. We covered each pot within the growth
chamber for 30 min. At 0, 15 and 30 min after the chamber
was put in place, we withdrew about 30 ml of gas with a 50 ml
plastic syringe through a capillary tube at the top of the chamber
and injected it into a 19 ml vacuum bottle with a rubber stopper
and screw cap. Back at the laboratory, the CO2 was measured by
gas chromatograph (Shimadzu GC-7A, Kyoto, Japan) with a
thermal conductivity detector (TCD). The flux rate was
calculated from the increase in CO2 concentration inside the
flux chamber for each pot or hill (Cheng et al., 2006).
2.5.
Statistical analysis
We calculated the analysis of variance (ANOVA) of plant
height, tiller number and night respiration for each week
during the treatment period. [CO2] and night temperature
were main-plot factors, and DAT was a split-plot factor. We
excluded the data on 113 DAT because the high night
temperature treatment had finished. We conducted a twoway ANOVA of the dry weight of each organ and whole plants,
grain yield and yield components at harvest for effects of
[CO2], night temperature and [CO2] night temperature. The
analysis was carried out with the statistical package SPSS 14
(SPSS Inc., Chicago, IL, USA).
Fig. 2 – Air temperature and soil temperature at 10 cm over
72 h (recorded every 10 min) in (a) low night temperature
and (b) high night temperature treatments during 9–11
September 2006.
3.
Results
3.1.
Plant height and tiller number
Plant height increased steadily until the grain-filling stage
(90 DAT) in all four treatments. It was significantly increased
54
**
ns
ns
**
**
*
ns
*
ns
**
**
**
**
+
ns
**
**
+
**
ns
ns
**
ns
ns
ns
ns
ns
ns: no significance; +, P < 0.1; *, P < 0.05; **, P < 0.01.
ns
ns
ns
36.9
34.0
8.5
42.5
33.5
26.9
19.0
18.1
5.3
97.9
87.4
12.0
84.4
73.9
14.1
2306
1933
19.3
88.4
84.6
4.5
2356
2213
6.5
2667
2619
1.8
97.6
100.1
2.6
27.3
26.3
3.8
34.4
31.7
8.5
19.4
18.5
5.1
97.3
96.2
1.1
66.5
66.6
0.1
1854
1777
4.4
69.8
70.0
0.3
1905
1846
3.2
2747
2642
3.9
Filled/total
spikelet (%)
No. per
hill
Fertile/total
spikelet (%)
No. per
hill
Fertile spikelet
Total spikelet
no. per hill
Spikelet
no. per
panicle
98.0
99.0
1.0
28.0
26.7
5.0
ns
ns
ns
ANOVA results
Temperature
CO2
Temperature CO2
Leaf and root dry weights were not affected by either [CO2] or
night temperature, since they were determined largely during
Elevated (EL)
Ambient (AL)
% change
Dry weights
Low (22 8C)
3.3.
Elevated (EH)
Ambient (AH)
% change
Night respiration ranged between 4 and 18 mg C per hill per hour
throughout the treatment period and changed significantly
with growth stage among treatments and with daily solar
radiation (Fig. 4). ANOVA results show that night respiration
was significantly increased by elevated [CO2] (by 12% before
heading and 13% after) and high night temperature (by 35%
before heading and 17% after) (both P < 0.01, Fig. 4). The effect of
temperature was notable before heading, but decreased after.
There was no significant interaction between elevated [CO2] and
night temperature in night respiration (P = 0.16), but after
heading the effect of temperature tended to be smaller.
High (32 8C)
Night respiration
Panicle
no. per
hill
3.2.
CO2
(abbrev.)
(by 7 cm) by high night temperature (P < 0.01, Fig. 3a), but
there was no significant effect of elevated [CO2] (P = 0.26).
The tiller number increased to a maximum at 35 DAT, 24
days before treatment start, then decreased until heading
stage. The tiller number (=panicle number) at harvest was not
affected by either elevated [CO2] or high night temperature
(Fig. 3b, Table 1).
Night
temperature
Fig. 3 – Changes in (a) plant heights and (b) tiller numbers of
rice plants throughout the experiment. Bars indicate
standard deviation (n = 3). Bef: before treatment; EH:
elevated [CO2] and high night temperature; AH: ambient
[CO2] and high night temperature; EL: elevated [CO2] and
low night temperature; AL: ambient [CO2] and low night
temperature. ANOVA results are inset. ns: not significant;
**, P < 0.01.
Table 1 – Effects of elevated CO2 and night temperature on yield and its components of IR72 rice crop
Filled spikelet
Filled/fertile
spikelet (%)
Individual
grain
weight (mg)
Brown
rice yield
(g per hill)
29.6
28.4
4.2
HI (%)
agricultural and forest meteorology 149 (2009) 51–58
agricultural and forest meteorology 149 (2009) 51–58
55
the vegetative growth stage (Fig. 5d and e). Ear dry weight was
significantly increased by elevated [CO2] and decreased by
high night temperature (both P < 0.01), and stem dry weight
was increased by both treatments (both P < 0.01, Fig. 5b and c).
The whole plant dry weight at harvest was significantly
increased by both treatments (P < 0.01, Fig. 5a). There were no
interactions between [CO2] and night temperature in dry
weight of organs or whole plants. (Note that ear dry weight is
different from rice yield, where there was an interaction; see
Section 3.4.) The net whole plant dry weight increase during
treatment was due to both ear formation and stem biomass
accumulation.
3.4.
Fig. 4 – Changes in night respiration of rice plants. Bars
indicate standard deviation (n = 3). Arrows indicate
heading date in each treatment. EH: elevated [CO2] and
high night temperature; AH: ambient [CO2] and high night
temperature; EL: elevated [CO2] and low night
temperature; AL: ambient [CO2] and low night
temperature. ANOVA results are inset. ns: not significant;
**, P < 0.01. The solar radiation on sampling days is shown
in the bottom part of the figure.
Grain yield and yield components
The number of spikelets per panicle and the total number per
pot were not affected by either elevated [CO2] or night
temperature (Table 1). The number of fertile spikelets was
not affected by elevated [CO2] but was significantly decreased
by high night temperature (P < 0.01): from 87% at low night
temperature to 70% at high night temperature. The number of
filled spikelets was significantly affected by both elevated
[CO2] and night temperature (both P < 0.01): it was increased
by elevated [CO2] by 4.4% at high night temperature and by 19%
at low night temperature; and decreased by high night
temperature by 8.1% under ambient [CO2] and by 20% under
elevated [CO2]. There was a slight interaction between [CO2]
and night temperature (P = 0.06). Individual grain weight was
significantly increased by elevated [CO2] (by 5.2% on average),
but was not affected by night temperature. Overall, brown rice
yield was significantly increased by elevated [CO2] and
Fig. 5 – Dry weight of (a) whole plant, (b) ear, (c) stem, (d) leaf and (e) root before treatment (Bef.) and at harvest. EH: elevated
[CO2] and high night temperature; AH: ambient [CO2] and high night temperature; EL: elevated [CO2] and low night
temperature; AL: ambient [CO2] and low night temperature. Bars indicate standard deviation (n = 3). ANOVA results are
inset. ns: not significant; **, P < 0.01. The units of Y-axes were similar.
56
agricultural and forest meteorology 149 (2009) 51–58
decreased by elevated night temperature (both P < 0.01).
Although the interaction of [CO2] and night temperature in
terms of ear dry weight was not significant (Fig. 5b; P = 0.12),
the interaction in brown rice yield was significant (Table 1;
P < 0.05).
4.
Discussion
4.1.
Effects of elevated [CO2] and high night temperature
on rice growth
Although many elevated [CO2] experiments were carried out
on rice during the last several decades, no paper reported that
the plant height of rice was increased or decreased by elevated
[CO2]. This lack of attention in the literature implies that
elevated [CO2] likely does not affect the plant height of rice, as
our results show in Fig. 3a. However, plant height of rice was
significantly increased by high night temperature, which is
similar to the height increases due to high daily temperature
as reported by Osada et al. (1973) and Ohe et al. (2007). The
elongated rice stems retained more photosynthate than did
ears under high night temperature (Fig. 5). The tiller (=panicle)
number at harvest was not affected by either elevated [CO2] or
high night temperature in this present study. However, the
apparent lack of tiller response undoubtedly was due to our
not imposing the CO2 and temperature treatments until after
the rice vegetative stage, which is the stage for tiller formation
(Fig. 3b).
Rice biomass production is determined by the balance
between net photosynthesis rate and night respiration
(Yamagishi, 1994; Sakai et al., 2001). Previous studies have
shown that elevated [CO2] promotes photosynthesis and
thus biomass production, whereas high temperatures
increase respiration, potentially reducing biomass production (Sakai et al., 2001; Peng et al., 2004). In the present
study, however, both elevated [CO2] and high night
temperature increased biomass production (both P < 0.01,
Fig. 5a). The total dry weight increase under high night
temperature resulted from the increase in stem biomass
(Fig. 5c), because the ear dry weight decreased under high
night temperature (P < 0.01, Fig. 5b). Three reasons might
explain this result. First, as shown in Fig. 4, night respiration
increased at high night temperature, but the magnitude was
modest, particularly after heading (17%), and was much
smaller than is generally expected from the temperature
dependence of respiration (typical Q10 values of two:
Yamagishi, 1994). Secondly, high night temperature might
have decreased N accumulation in the panicles through
lower N translocation from the vegetative organs under
high night temperature, thus resulting in a higher N
concentration in the leaves and stems (P < 0.01, data not
shown). These higher N concentrations might have
enhanced photosynthesis and maintained vegetative
growth during the later growth stage, and thus increased
the total dry weight in the high night temperature
treatment. Thirdly, the elongated rice stem under high
night temperature lead to a more efficient plant architecture
for rice photosynthesis where mutual shading between
leaves was reduced.
4.2.
yield
Elevated [CO2] and night temperature affected rice
Numerous controlled-environment experiments have shown
that elevated CO2 increases tiller and panicle numbers in rice
(Baker et al., 1992; Ziska et al., 1997; Moya et al., 1998; Kim et al.,
2003; Yang et al., 2006), but that increased daily mean
temperature does not change the spikelet number per panicle
or yield of rice (Yoshida, 1973a; Ziska et al., 1997). The increase
in air temperature often offsets the stimulation of rice biomass
and grain yield due to elevated [CO2] (Kim et al., 1996; Ziska
et al., 1996; Moya et al., 1998). Only limited information on the
direct effects of night temperature is available, but Morita et al.
(2004) reported that increased night temperature alone
reduced the size and weight of grain. Here, we have
determined the interaction of elevated [CO2] and night
temperature on rice yield.
Rice yield is determined by panicle number per land area,
spikelet number per panicle, filled spikelet percentage and
individual grain weight. Usually, productive tiller number is
determined by the maximum tiller number that is formed
during the vegetative growth period and tiller degeneration
during the reproductive growth period. In rice FACE experiments, yield increases caused by elevated [CO2] are related
most strongly to larger productive panicle number per area
and larger spikelet number per panicle (Kim et al., 2003; Yang
et al., 2006). We applied the [CO2] and night temperature
treatments during the reproductive growth stage, so the
benefit of elevated [CO2] on sink formation was negligible. In
fact, the numbers of tillers (panicles) and total spikelets per pot
at harvest were not different between the treatments (Fig. 3,
Table 1).
Our results must be interpreted as the effects of elevated
[CO2] and night temperature on the grain setting and filling
processes only. The major determinant of grain set is
spikelet fertilization just after flowering. Night temperature
has a strong effect on grain set by increasing spikelet
sterility, whereas elevated [CO2] has virtually no effect.
Satake and Yoshida (1978) showed that night temperature
higher than 30 8C reduced spikelet fertility. Our study shows
that this can occur under both ambient and elevated [CO2].
Kim et al. (1996) and Matsui et al. (1997) showed that spikelet
sterility caused by daytime heat was exacerbated by
elevated [CO2], but the present study shows that sterility
caused by night-time heat does not have a significant
interaction with [CO2]. The reason for this difference is not
clear, but elevated [CO2] may exacerbate daytime-heatinduced sterility because of indirect effects of increased
crop surface and panicle temperatures (Yoshimoto et al.,
2005).
We found a significant interaction between [CO2] and night
temperature in the ratio of filled grain to fertile spikelets: there
was a positive effect of elevated [CO2] only under low night
temperature. This could be related to the fact that the number
of fertile spikelets was significantly reduced by high night
temperature, so enhanced photosynthesis under elevated
[CO2] did not translate into filled grain percentage. This result
reflects the responses of brown rice yield to elevated [CO2].
These results have shown for the first time that limitation of
grain setting under high night temperature limits the sink
agricultural and forest meteorology 149 (2009) 51–58
capacity, reducing the advantage of grain yield due to elevated
[CO2].
5.
Conclusions
Our results show that high night temperature during the
reproductive growth stage reduced the stimulatory effect of
elevated [CO2] on brown rice yield. This was not due to reduced
enhancement of biomass, but due to decreased dry matter
allocation to grain as a result of reduced spikelet fertility. The
enhanced vegetative growth and a smaller-than-expected
temperature effect on night respiration were reasons for
increased total biomass by high night temperature during the
reproductive period. Note, however, that our results are based
on a growth chamber experiment, and the way in which
elevated night temperature decreases the fertile spikelet
percentage is still not understood. Further studies on the
interactive effects of elevated [CO2] and night temperature
should be carried out to confirm our results in the field under
future climate warming scenarios.
Acknowledgements
This research was funded by the Global Environment Research
Program of the Ministry of the Environment, Japan. We thank
Drs M. Yoshimoto and M. Fukuoka of NIAES for their
comments and discussion.
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