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Received: 11 April 2018 Revised: 27 March 2019 Accepted: 24 April 2019

DOI: 10.1002/met.1804

RESEARCH ARTICLE

Analysis of growth functions that can increase irrigated wheat


yield under climate change

Hamed Eyni-Nargeseh1 | Reza Deihimfard2 | Sajjad Rahimi-Moghaddam2 |


Ali Mokhtassi-Bidgoli1

1
Department of Agronomy, Faculty of
Agriculture, Tarbiat Modares University, Abstract
Tehran, Iran This study assessed the growth of wheat under climate change conditions
2
Department of Agroecology, (increases in CO2 and temperature) in southwestern Iran. Future climate data were
Environmental Sciences Research Institute,
projected using long-term climate data for radiation and minimum and maximum
Shahid Beheshti University, Tehran, Iran
temperatures for the baseline period 1980–2010. The future climate scenarios made
Correspondence use of delta changes from the Coupled Model Intercomparison Project Phase
Ali Mokhtassi-Bidgoli, Department of
Agronomy, Faculty of Agriculture, Tarbiat
5 (CMIP5) in the R package of the Agricultural Model Intercomparison and
Modares University, PO Box 14115-336, Improvement Project for 2040–2070. After generating the weather data, the Agri-
Tehran, Iran. cultural Production System Simulator wheat model was applied to assess the poten-
Email: mokhtassi@modares.ac.ir
tial impact of climate change on wheat yield and the relative growth rate, crop
growth rate, total dry weight and leaf area index. Averaged across all locations, an
increase of 1.6 and 2.3 C in annual temperature was projected under representative
concentration pathway 4.5 (RCP4.5) and RCP8.5, respectively. The decrease in the
average length of the growing season was 7.5 and 9.3% under RCP4.5 and
RCP8.5, respectively, in the mid-21st century with respect to baseline. The mean
cumulative radiation will decrease by 11.6 and 14.3% averaged across all sites for
RCP4.5 and RCP8.5, respectively. Grain yield will decrease at all sites for both
RCPs. The greatest decrease in wheat yield was projected for Izeh under RCP4.5
and RCP8.5 (7.9 and 9.3%, respectively) for 2040–2070 relative to baseline. A
greater grain yield at baseline than for the RCP8.5 and RCP4.5 scenarios was
closely related to higher leaf area index, relative growth rate and crop growth rate
values during the reproductive phase. Thus, these parameters can be defined as
indicators of crop yield.

KEYWORDS
AgMIP, APSIM, CO2, growth indices, rising temperature

1 | INTRODUCTION average temperature will rise by 3.7–4.8 C by the end of the


current century (IPCC, 2014). The global average increase
According to the Fifth Assessment Report of the Intergov- in surface temperatures in the last 50 years has been 0.18 C
ernmental Panel on Climate Change (IPCC), global per decade. The IPCC Fifth Assessment has projected that
warming, which is primarily a result of the increase in green- the increase in the global mean temperature from 2018 to
house gases, will continue. It is projected that the global 2100 will probably be 1.8 ± 0.5 C for representative

Meteorol Appl. 2019;1–10. wileyonlinelibrary.com/journal/met © 2019 Royal Meteorological Society 1


2 EYNI-NARGESEH ET AL.

concentration pathway 4.5 (RCP4.5) and 3.7 ± 0.7 C for parameters showing the positive response of plants to an
RCP8.5 relative to 1986–2005 (Wang et al., 2015). The increase in CO2 will be compensated. Depending on the cul-
most important greenhouse gas is CO2, which is released tivar sensitivity to temperature, the yield response of a crop
into the atmosphere through human activity. to CO2 can decrease or increase. In general, an increase in
Climate changes will have different effects in different CO2 at low temperatures will have less effect on crop pro-
areas, although it is expected that the negative effects will be duction (Bannayan, 2009).
greater than the positive effects (Porter et al., 2014). In this Under climate change, new crop varieties should be
context, agriculture is vulnerable to climate change because developed with physiological and agronomic traits that are
it is linked to weather conditions. The optimal growth and suited to future conditions (Fang et al., 2017). Growth analy-
yield of crops are possible under a range of climate vari- sis is frequently used by plant physiologists and agronomists
ables, but climate change can change the optimal range of to address this issue (Gul et al., 2013). Growth indicator
temperature required for plant growth (Ranuzzi and analysis interprets the response of plant species and varieties
Srivastava, 2012). Climate change includes increases in to environmental conditions. Growth analysis is a valuable
atmospheric CO2 and temperature and rainfall fluctuations, method for study of the growth capability of plants under
which directly affect crop growth and yield (Bannayan different climate conditions and management (Hunt, 1991).
et al., 2011). Iran has a Mediterranean climate and long dry In order to analyse growth, leaf area and dry weight traits
summers that make it vulnerable to climate change. These are required, while other growth indicators can be calculated
changes could have profound effects on the sustainability of (Buttery, 1969). The optimum leaf area index (LAI) is an
crops in arid and semi-arid areas such as Iran (Bannayan and important factor in grain yield. If the LAI reaches an opti-
Eyshi Rezaei, 2014). mum level within a shorter time, a higher grain yield could
In the last decade, much research has been conducted on result. The slow development of the leaf area could weaken
the effects of climate change on crop production (Koocheki development of the canopy, decreasing radiation absorption
et al., 2006; Deihimfard et al., 2018; Rahimi-Moghaddam and the growth rate (Darzi-Naftchali et al., 2017). Nouri
et al., 2018). Eyshi Rezaie and Bannayan (2012) studied et al. (2017) investigated the effects of climate change on
northeastern Iran and showed that the greatest increase and rainfed wheat production in western and northwestern Iran.
decrease in grain yield of rainfed wheat compared to base- They concluded that crop yield and maximum LAI will
line was from 2010 to 2039 (+15%) under the B2 scenario decrease chiefly because of precipitation deficits from
and from 2040 to 2069 (−50%) under the A2 scenario. You October through December and January through March in
et al. (2009) revealed that a 1 C increase during the wheat 2071–2100.
growing season would reduce wheat yields by 3–10%. Many Growth functions to increase yield may vary in different
of these studies used crop simulation models and weather regions and when different general circulation models
generators (Bannayan et al., 2014; Deihimfard et al., 2018). (GCMs) and emission pathways are used. Thus, growth
Simulation research using crop models enhances under- analysis should be conducted for each specific region to
standing about plant response to weather conditions reduce projection uncertainty. CO2 and temperature are cur-
(Deihimfard et al., 2019) and soil and their interaction with rently increasing globally (IPCC, 2014), including in Iran
various scenarios of crop management (Manschadi et al., (Bannayan et al., 2014; Deihimfard et al., 2018; Rahimi-
2010). Crop growth and development as affected by future Moghaddam et al., 2018). These two parameters directly
climate change can be estimated using crop models such as affect the rate of development, growth and crop yield. The
the Decision Support System for Agrotechnology Transfer, current study aimed to determine the effect of climate
the World Food Studies Crop Simulation Model, Simple and change (increase in CO2 and temperature) on irrigated wheat
Universal Crop Growth Simulation, the Agricultural Produc- yield, biomass, LAI, crop growth rate (CGR), relative
growth rate (RGR) and distance between planting and har-
tion System Simulator (APSIM), the Simple Simulation
vest from 2040 to 2070 relative to current climatology
Modelling Legume and WheatGrow (Lv et al., 2013;
(1980–2010) in southwestern Iran.
Moradi et al., 2013; Amiri et al., 2016).
Wheat is a major crop globally and is grown on more
than 220 million ha (5.7 million ha in Iran) annually (FAO, 2 | MATERIALS AND METHODS
2016). Any environmental change will affect wheat growth
and development, which will affect wheat production and This study was conducted at three locations in Khuzestan
yield (Meza et al., 2008). In the future, under climate change province; Ahwaz, Izeh and Ramhormoz (Figure 1).
conditions extreme values of climate variables, particularly Khuzestan province comprises about 64,057 km2 in south-
of temperature, will threaten crop production. However, it is western Iran. The study area is one of the most productive
possible that the negative effects of extreme values of agricultural regions in Iran, accounting for more than
EYNI-NARGESEH ET AL. 3

48 ° 0 ′ 0 ′′ E 49 ° 0 ′ 0 ′′ E 50 ° 0 ′ 0 ′′ E
33 ° 0 ′ 0 ′′ N

33 ° 0 ′ 0 ′′ N
Elevation= 21 m Caspian Sea
32 ° 0 ′ 0 ′′ N

32 ° 0 ′ 0 ′′ N
Max T= 28.47°C
Min T= 13.93°C
Elevation= 18 m Period= 1980–2010
Max T= 33.23°C
Min T= 18.76°C Izeh
Period= 1980–2010

Ahwaz Ramhormoz
31 ° 0 ′ 0 ′′ N

31 ° 0 ′ 0 ′′ N
Elevation= 179 m
Max T= 32.79°C
Min T= 19.54°C
Period= 1980–2010
30 ° 0 ′ 0 ′′ N

30 ° 0 ′ 0 ′′ N
N
Persian Gulf
0 37.5 75 150
Kilometers
29 ° 0 ′ 0 ′′ N

29 ° 0 ′ 0 ′′ N

48 ° 0 ′ 0 ′′ E 49 ° 0 ′ 0 ′′ E 50 ° 0 ′ 0 ′′ E

FIGURE 1 Selected meteorological stations for the different locations in Khuzestan province, Iran. Max T, maximum temperature; Min T,
minimum temperature

11.08% of the country’s annual irrigated wheat production. 2.1 | Scenarios, period and climate model
It covers 6.89% of the total arable land (Agricultural Jihad
Forecasting future agricultural yield requires attention to cli-
Ministry of Iran, 2018).
Historical weather data for daily solar radiation mate variability and conditions (Schoof and Pryor, 2001).
(MJ m−2 day−1) and maximum and minimum temperature Projections from GCMs are commonly used as scenarios of
( C) for the baseline period (Figure 1) were collected for future climate conditions for the 21st century (Cheng et al.,
each study location from the respective climatological sta- 2008). To study the local impact of climate, spatial scales,
tions. The geographical details of the study locations are which are much smaller than those offered by the most
shown in Figure 1 and were used as inputs for the climate highly resolved GCMs, are used. In order to relate the large-
model and crop growth simulation model. The Weather- scale weather patterns to the local scale, downscaling is nec-
Man programme was used to modify and restore outliers essary. In the present study, the relationship between these
and missing data in the stations (Hoogenboom et al., scales was determined using the R tool in the Agricultural
2003). The Ångström formula was used to calculate the Model Intercomparison and Improvement Project (AgMIP)
daily solar radiation from the duration of sunlight to downscale daily station-scale meteorological variables
(Ångström, 1924): from outputs of the large-scale GCM of Miroc5 (Watanabe
et al., 2010). Climate data at baseline were used as the basis
Rs = ða + b × n=N ÞRa ð1Þ of future climate scenario analysis. The future climate sce-
narios made use of delta changes of the Coupled Model
where Rs is the daily solar radiation (MJ m−2 day−1), n is the Intercomparison Project Phase 5 (CMIP5) from the climate
actual duration of sunlight (h), N is the maximum possible scenario generation tools in R (Ruane et al., 2013).
duration of sunlight or daylight (h), Ra is the extraterrestrial There is a need to study future regional climate changes
radiation (MJ m−2 day−1), a is a regression constant and uncertainties using up-to-date climate models (Tian
expressing the fraction of extraterrestrial radiation reaching et al., 2015). In the current study, the effect of future climate
the Earth on an overcast day (n = 0) and a + b is the fraction change on wheat growth from 2040 to 2070 was analysed
of extraterrestrial radiation reaching the Earth on a clear day relative to the current climate (1980–2010). Outputs of two
(n = N). The values for a and b were 0.25 and 0.50, respec- emission scenarios, RCP4.5 and RCP8.5, were used. The
tively, in this study. average temperature for RCP4.5 quickly rises to 2050 and
4 EYNI-NARGESEH ET AL.

from 2050 to 2100 the increase slows. In RCP8.5, the tem- (nRMSE) was applied (Wallach and Goffinet, 1987). This
perature increase from 2080 to 2100 is faster than from statistic shows the relative difference between observed and
2035–2080. Additional details about these scenarios can be simulated data:
found in Moss et al. (2010) and Wayne (2013).
sffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi
X n

nRMSE = 100=O ðPi − Oi Þ2 =n ð2Þ
2.2 | Modelling i=1

After assessment of the weather data, the APSIM wheat


model (Keating et al., 2003) was used to simulate the growth where Pi is the simulated data, Oi is the observed data, Ō is
and development of wheat under climate change conditions the mean of the observed data and n is the number of obser-
and its potential impact. The model was originally calibrated vations. Model simulation accuracy increases as nRMSE
approaches 0.
and validated for the Chamran cultivar (a prevalent cultivar
in Khuzestan province) (Deihimfard et al., 2015; Deihimfard
et al., 2018). The APSIM model has high power to simulate 2.4 | Growth indicators
crop growth and yield. It simulates the effects of tempera-
It is assumed for calculation of growth analysis (Buttery,
ture, precipitation, radiation, genotype, soil properties and
1969) that changes in the plant total dry weight (TDW) and
management factors on crop growth and development, soil
LAI are second-degree polynomials which can be denoted as
water and nitrogen balance on a daily basis and predicts
Ln in order to reduce the dependence of variance relative to
yield. The outputs of daily climate parameters for baseline
the means:
and future scenarios after preparation were used as inputs in
the APSIM. The other inputs needed to run the model are
genetic co-efficients and management information. These Ln TDW = a + bt + ct2 ð3Þ
inputs included the genetic co-efficients of the Chamran cul-
tivar, plant density (350 plant/ha), row spacing (25 cm) and Ln LAI = a1 + b1 t + c1 t 2 ð4Þ
planting data (November 5).
The estimation of genetic co-efficients is required in the To calculate other growth indicators, these indices were
simulation models because they show the specifications of a used as follows (Goudriaan and Van Laar, 1994):
species or cultivar. There are two types of co-efficients. The
first type, species-specific genetic co-efficients, will be the RGR = b + 2ct ð5Þ
same for all cultivars and the other type, cultivar-specific
genetic co-efficients, are specific to one cultivar (Wallach CGR = ðb + 2ct Þ × TDW ð6Þ
et al., 2011). Genetic co-efficients are used in crop growth
simulation models to explain the key specifications of the where a, a1, b, b1, c and c1 are regression co-efficients, and
cultivars. These co-efficients relate to plant genetics that are t is the number of days after sowing. OriginPro 9.1 software
not affected by the environment. Changes in these are lim- was used to fit the equations and draw the figures.
ited to a specific range for the co-efficient. The yield of a
genotype under different environmental conditions is also
3 | R E S U L T S A N D DI S C U S S I O N
affected by the co-efficients in addition to climate factors
and farm management practices (Roman-Paoli et al., 2000).
3.1 | Model performance
The co-efficient values should be measured before their use
in field experiments or for estimation (Makowski et al., The APSIM performance was evaluated by comparing the
2006). After running the model, the outputs under future cli- simulated data with the data obtained from Deihimfard et al.
mate conditions and baseline in all locations were analysed. (2015) for grain yield, biomass and LAI (Figure 2). For
grain yield, biomass and LAI, the model was able to repli-
cate the trend with R2 values of 0.78, 0.66 and 0.88, respec-
2.3 | Quantitative evaluation of model
tively (Figure 2). The nRMSE values for grain yield,
performance
biomass and LAI were 7.59, 5.56 and 11.45%, respectively.
Comparison of the values simulated by the model and the This indicates that performance was acceptable for the three
observed values is an important step before using the model traits, but less satisfactory for LAI. Model performance is
for simulation studies. Different indicators are used to evalu- considered excellent at nRMSE values of less than 10%,
ate the differences between the observed and simulated data. good for nRMSE values of 10 to 20%, fair for nRMSE
In the present study, the normalized root mean square error values of 20 to 30% and poor for nRMSE values greater than
EYNI-NARGESEH ET AL. 5

FIGURE 2 Scatter plot of simulated versus observed grain yield (yield), total above ground biomass (biomass) and maximum leaf area index
(LAI) of wheat

30% (Nouri et al., 2017). These results are similar to those growing season. These results are supported by Deihimfard
of Bannayan et al. (2014) who reported R2 values of 0.88, et al. (2018), who reported that temperatures during the
0.51 and 0.94 and rRMSE values of 5.31, 4.81 and 8.10% growth period for winter wheat under climate change would
for grain yield, biomass and maximum LAI of wheat, increase significantly compared to baseline, while the length
respectively, using the Cropping System Model Crop Esti- of the growing season and cumulative radiation would
mation through Resource and Environment Synthesis decrease.
(CSM-CERES) wheat model.
3.3 | Grain yield: baseline and future
3.2 | Future changes in climate and phenology
The simulation results using the APSIM showed that, for the
When averaged across all locations, an increase of 1.6 and base time period and RCP scenarios, the highest grain yield
2.3 C in annual temperature was projected under RCP4.5 was in Izeh and the lowest was in Ahwaz (Table 2). This
and RCP8.5, respectively (Table 1). The decrease in the pro- high grain yield in Izeh could relate to the temperature, the
jected average length of the growing season (based on days increased length of the growing season and higher cumula-
after planting) was 7.5 and 9.3% under RCP4.5 and RCP8.5, tive radiation during the growing season (Table 1). Table 1
respectively, at 2050 with respect to baseline. The mean shows that Izeh has a lower average temperature during the
cumulative radiation was projected to decrease by 11.6 and growing season than the other locations (Ahwaz and
14.3%, averaged across all sites, for RCP4.5 and RCP8.5, Ramhormoz); thus, the length of the growing season was
respectively (Table 1). Consequently, it was projected that longer and resulted in more cumulative radiation, photosyn-
irrigated wheat in southwestern Iran will experience a thesis and grain yield. Asseng et al. (2015) reported that
decrease in radiation due a decrease in the length of the higher temperatures decrease the number of days during

TABLE 1 Averaged climatic variables over the wheat growing season under climate change scenarios in the southwest of Iran

Location Scenario Temperature ( C) Days after planting (day) Radiation (MJ/m2)


Ahwaz Baseline 16.16 (±0.14)a 144.46 (±0.89) 1,997.19 (±33)
RCP4.5 17.8 (±0.16) 135.7 (±0.72) 1,818.57 (±27)
RCP8.5 18.45 (±0.17) 133.86 (±0.69) 1,783.3 (±26)
Izeh Baseline 13.25 (±0.08) 167.8 (±0.79) 2,821.58 (±78)
RCP4.5 14.85 (±0.09) 153.33 (±0.78) 2,464.74 (±66)
RCP8.5 15.37 (±0.10) 150.03 (±0.74) 2,384.32 (±63)
Ramhormoz Baseline 16.14 (±0.13) 144.53 (±0.9) 2,285.45 (±61)
RCP4.5 17.8 (±0.15) 135.76 (±0.74) 2,084.96 (±55)
RCP8.5 18.47 (±0.16) 133.96 (±0.69) 2,046.37 (±56)
a
Values in parentheses are standard errors.
6 EYNI-NARGESEH ET AL.

TABLE 2 Wheat grain yield (kg/ha) in the baseline and future climate change scenarios (2040–2070) in the southwest of Iran

Location Baseline RCP4.5 RCP8.5


a
Ahwaz 5,640.7 (±146) 5,527.48 (±134) 5,564.4 (±133)
Izeh 8,757.7 (±329) 8,118.69 (±305) 8,011.4 (±289)
Ramhormoz 6,731.9 (±274) 6,686.81 (±267) 6,716.1 (±271)
a
Values in parentheses are standard errors.

which plants can intercept light for photosynthesis, which this have been provided by Taiz and Zeiger (2010). Wheat is
will reduce biomass and grain yield. a C3 plant; thus, an increase in CO2 concentration can
Grain yield is projected to decrease at all sites for both increase grain yield under appropriate temperature condi-
RCPs (Table 2). The greatest decrease in wheat yield was tions. An excessive increase in temperature, however, will
projected for Izeh under RCP4.5 and RCP8.5 (7.9 and 9.3%, neutralize the positive effects of an increase in CO2. In the
respectively) for 2040–2070 relative to baseline. This can be present study, under climate change the increase in tempera-
attributed to differences in major climate variables of tem- ture is projected to decrease the grain yield. The effect of cli-
perature (+12 to +16%), cumulative radiation (−15 to mate change on wheat strongly depends on the cultivar
−18%) and growth duration (−9 to −12%) for 2040–2070 characteristics and the spatial and temporal patterns of cli-
relative to baseline. Recent studies have shown that climate mate change (Semenov, 2009). Under climate change condi-
warming in the past decades has advanced wheat phenology tions, there are interactions between an increase in
and reduced wheat growth duration and yield (Nouri et al., temperature and an increase in CO2 (Ludwig and Asseng,
2017; Tao et al., 2017). 2006). For example, in the northern part of the Western
Plants can be classified as C3, C4 or CAM according to Australian wheat belt, higher temperatures had a negative
their carbon fixation pathways. This is a standard biological effect on yield, while in the southern cooler part of the region
approach to categorization of plants in terms of mechanisms higher temperatures had a strong positive effect (Ludwig and
for concentrating CO2 at the site of carboxylation. In the C3 Asseng, 2006). Lv et al. (2013) indicated that under climate
carbon fixation pathway, CO2 and ribulose bisphosphate are change rainfed wheat yield in the northern regions of China
converted into 3-phosphoglycerate acid. Further details on will decrease but it will increase in the southern regions.

FIGURE 3 Wheat growth indices (CGR, crop growth rate; LAI, leaf area index; RGR, relative growth rate; TDW, total dry weight) under
climate change in Ahwaz. DAE, days after emergence
EYNI-NARGESEH ET AL. 7

FIGURE 4 Wheat growth indices (CGR, crop growth rate; LAI, leaf area index; RGR, relative growth rate; TDW, total dry weight) under
climate change in Izeh. DAE, days after emergence

3.4 | Growth analysis under climate change under climate change (RCP4.5 and RCP8.5 scenarios in
2040–2070). In all sites, wheat plants exhibited similar
Growth analysis of crops allows integration of physiological
trends of response under projected climate change conditions
and morphological indices. These measurements can help to
(Figures 3–5). At baseline and under climate change condi-
explain yield differences under different conditions. In the
tions, the highest TDW, LAI, RGR and CGR values were
present study, growth indices were calculated at baseline and
observed in Izeh (Figure 4). All these indices showed no

FIGURE 5 Wheat growth indices (CGR, crop growth rate; LAI, leaf area index; RGR, relative growth rate; TDW, total dry weight) under
climate change in Ramhormoz. DAE, days after emergence
8 EYNI-NARGESEH ET AL.

significant differences between the RCP8.5 and RCP4.5 sce- 4 | CONCLUSION


narios over the growing season (Figures 3–5). The indices of
LAI, RGR and CGR were higher at baseline from 100 days The results of the current study show that the temperature
after emergence (DAE) onward (approximately at the during the growing season under two emission scenarios
flowering stage and during grain filling) than in the RCP8.5 (on average) would increase 11.33% compared to baseline at
and RCP4.5 scenarios (Figures 3–5). Conversely, TDW the selected locations, while the length of the growing sea-
showed a continuous increase, although at baseline the son (7.76%) and cumulative radiation (11.44%) would
increase was lower than for the climate change scenarios decrease. As the temperature rises under climate change
(Figures 3–5). (representative concentration pathway 8.5 [RCP8.5] and
As seen, the higher grain yield values at baseline than for RCP4.5 scenarios), the wheat grain yield would decrease 3%
the RCP8.5 and RCP4.5 scenarios are closely related to the compared to baseline, largely because of the negative effects
increases in LAI, RGR and CGR during the reproductive of high temperature and reductions in the length of the grow-
phase. Thus, these parameters can be defined as indicators ing season and cumulative radiation. In the current study,
of crop yield. Similar results were reported by Camargo wheat, as a C3 plant, was affected by temperature and CO2
et al. (2015) in a study on the effects of irrigation treatments concentration under climate change. An excessive increase
on the growth and development of potatoes (Solanum of temperature was shown to neutralize the positive effects
tuberosum L.). Tao et al. (2017) reported that wheat yields of increased CO2 concentration. Consequently, the wheat
significantly and positively correlated with the length of grain yield declined due to the reduction in the length of the
GP2 (from winter dormancy date to greenup date), GP4 growing season and in the cumulative radiation. The gener-
(from greenup date to anthesis date), and GPw (growth ally higher grain yield at baseline than for the RCP8.5 and
period from emergence date to maturity date). They indi- RCP4.5 scenarios was closely related to the increase in leaf
cated that high temperature stress during GP4 is a challenge area index, relative growth rate and crop growth rate during
that may potentially decrease yield. the reproductive phase; thus, these parameters can be
C3 plants seem to benefit more in terms of dry matter defined as indicators of crop yield.
production from a higher CO2 level because of their
increased leaf expansion, photosynthesis rate per unit area, A C KN O W L E D G E M E N T
water use efficiency and photorespiration rates (Attri and
The authors declare that they have no conflict of interest.
Rathore, 2003). Low temperatures limit growth during the
winter; thus, higher temperatures can increase biomass
production. OR CI D
The LAI is the most important determinant of photosyn-
Ali Mokhtassi-Bidgoli https://orcid.org/0000-0002-3330-
thesis and dry matter accumulation. Increases or decreases in
1198
the LAI have a direct effect on plant growth and are the most
important indicators for increasing photosynthesis and pro-
duction of assimilates (Ayisi and Poswall, 1997). Valizadeh REF ER ENC ES
et al. (2013) indicated that the LAI of wheat under climate Agricultural Jihad Ministry of Iran. (2018) Agricultural statistics, Teh-
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