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Current status and prospects of rice canopy temperature research

Min Jiang

Jiangsu Key Laboratory of Crop Genetics and Physiology/Jiangsu Key Laboratory of Crop Cultivation and Physiology, Agricultural College of Yangzhou University, Yangzhou, China

Jiangsu Co-Innovation Center for Modern Production Technology of Grain Crops, Yangzhou University, Yangzhou, China

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Kefan Guo,

Kefan Guo

orcid.org/0000-0003-4719-1062

Jiangsu Key Laboratory of Crop Genetics and Physiology/Jiangsu Key Laboratory of Crop Cultivation and Physiology, Agricultural College of Yangzhou University, Yangzhou, China

Jiangsu Co-Innovation Center for Modern Production Technology of Grain Crops, Yangzhou University, Yangzhou, China

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Jiaqi Wang,

Jiaqi Wang

Jiangsu Key Laboratory of Crop Genetics and Physiology/Jiangsu Key Laboratory of Crop Cultivation and Physiology, Agricultural College of Yangzhou University, Yangzhou, China

Jiangsu Co-Innovation Center for Modern Production Technology of Grain Crops, Yangzhou University, Yangzhou, China

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Yunfei Wu,

Yunfei Wu

orcid.org/0000-0002-2771-9133

Jiangsu Key Laboratory of Crop Genetics and Physiology/Jiangsu Key Laboratory of Crop Cultivation and Physiology, Agricultural College of Yangzhou University, Yangzhou, China

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Xinping Shen,

Xinping Shen

Jiangsu Key Laboratory of Crop Genetics and Physiology/Jiangsu Key Laboratory of Crop Cultivation and Physiology, Agricultural College of Yangzhou University, Yangzhou, China

Jiangsu Co-Innovation Center for Modern Production Technology of Grain Crops, Yangzhou University, Yangzhou, China

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Lifen Huang,

Corresponding Author

Lifen Huang

[email protected]

Jiangsu Key Laboratory of Crop Genetics and Physiology/Jiangsu Key Laboratory of Crop Cultivation and Physiology, Agricultural College of Yangzhou University, Yangzhou, China

Jiangsu Co-Innovation Center for Modern Production Technology of Grain Crops, Yangzhou University, Yangzhou, China

Correspondence

Lifen Huang, Agricultural College of Yangzhou University, Yangzhou 225009, China.

Email: [email protected]

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Min Jiang,

Min Jiang

Jiangsu Key Laboratory of Crop Genetics and Physiology/Jiangsu Key Laboratory of Crop Cultivation and Physiology, Agricultural College of Yangzhou University, Yangzhou, China

Jiangsu Co-Innovation Center for Modern Production Technology of Grain Crops, Yangzhou University, Yangzhou, China

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Kefan Guo,

Kefan Guo

orcid.org/0000-0003-4719-1062

Jiangsu Key Laboratory of Crop Genetics and Physiology/Jiangsu Key Laboratory of Crop Cultivation and Physiology, Agricultural College of Yangzhou University, Yangzhou, China

Jiangsu Co-Innovation Center for Modern Production Technology of Grain Crops, Yangzhou University, Yangzhou, China

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Jiaqi Wang,

Jiaqi Wang

Jiangsu Key Laboratory of Crop Genetics and Physiology/Jiangsu Key Laboratory of Crop Cultivation and Physiology, Agricultural College of Yangzhou University, Yangzhou, China

Jiangsu Co-Innovation Center for Modern Production Technology of Grain Crops, Yangzhou University, Yangzhou, China

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Yunfei Wu,

Yunfei Wu

orcid.org/0000-0002-2771-9133

Jiangsu Key Laboratory of Crop Genetics and Physiology/Jiangsu Key Laboratory of Crop Cultivation and Physiology, Agricultural College of Yangzhou University, Yangzhou, China

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Xinping Shen,

Xinping Shen

Jiangsu Key Laboratory of Crop Genetics and Physiology/Jiangsu Key Laboratory of Crop Cultivation and Physiology, Agricultural College of Yangzhou University, Yangzhou, China

Jiangsu Co-Innovation Center for Modern Production Technology of Grain Crops, Yangzhou University, Yangzhou, China

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Lifen Huang,

Corresponding Author

Lifen Huang

[email protected]

Jiangsu Key Laboratory of Crop Genetics and Physiology/Jiangsu Key Laboratory of Crop Cultivation and Physiology, Agricultural College of Yangzhou University, Yangzhou, China

Jiangsu Co-Innovation Center for Modern Production Technology of Grain Crops, Yangzhou University, Yangzhou, China

Correspondence

Lifen Huang, Agricultural College of Yangzhou University, Yangzhou 225009, China.

Email: [email protected]

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First published: 21 October 2022

https://doi.org/10.1002/fes3.424

Citations: 4

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Abstract

Canopy temperature is an important physiological and ecological characteristic of rice plants. It can comprehensively reflect their individual characteristics, such as their adaptability to the environment and the quality of the population. This paper reviews influencing factors, variation patterns, practical applications, formation processes, and measurement techniques of rice canopy temperature. Moreover, we analyzed the weaknesses of current studies on this topic, proposed improvements to the existing methods and techniques used to measure rice canopy temperature, and systematically assessed the effects of canopy temperature on rice production. This study provides a theoretical support for the construction of an auxiliary decision-making system to guide rice crop production practices.

1 INTRODUCTION

The canopy is where matter and energy exchanges occur between plants and the atmosphere. These exchanges determine the yield and quality of rice. Canopy temperature is a typically mean value that reflects the surface temperature of a plant and is related to canopy function (Dong, 1984). Therefore, the canopy temperature is an important index that intuitively reflects the growth characteristics of multiple rice varieties. To improve the quality and yield of cultivated rice, the biological basis for changes in canopy temperature and the effects thereof must be determined (Gabaldón-Leal et al., 2016; Siebert et al., 2014). Previous studies on Gramineae and Leguminosae crops have identified substantial non-climate-induced differences in the canopy temperature, even under the same external environmental conditions (Kumar et al., 2017; Tafesse et al., 2019; Takai et al., 2010). This demonstrates that surrounding air temperature does not reflect the actual canopy temperature. The adaptability of rice in response to environmental changes, such as variations in rice canopy temperature, different patterns of variation, and regulatory mechanisms must be investigated; this could provide new insights into rice variety selection (breeding), cultivation management, and novel methods to monitor rice growth and predict yield quality (Cai et al., 2020; Wang et al., 2020).

2 VARIATIONS IN RICE CANOPY TEMPERATURE

The canopy temperature of rice is controlled by its thermal properties and physiological responses to the environment (Schymanski et al., 2013). The thermal properties include the heat conduction between rice and the environment through four processes (conduction, convection, radiation, and evapotranspiration of latent heat), and the heat conduction within tissues. Rice plants primarily absorb heat through solar radiation and long-wave radiation from surrounding objects, including the soil and the sky, and generate heat energy via metabolic processes (Pieruschka et al., 2010). Heat dissipation occurs primarily through the emission of long-wave radiation from the plant, transpiration, and convection with the environment (Figure 1). For the leaves to achieve a steady-state temperature equilibrium, the sum of the energy balance components must equal zero (Lambers et al., 2008):

{SR}_{net} + {LR}_{net} + C + λE + M = 0,

Details are in the caption following the image — **FIGURE 1**
Open in figure viewer PowerPoint

Schematic representation of the components of energy balance of a leaf consisting of short-wave radiation (SR), long-wave radiation (LR), both incident (in) and emitted (em), convective heat transfer (C), and evaporative heat loss (lE). Reflection (r), transmission (tr), and fluorescent emission (FL) are only given for the SR incidence on the upper side of the leaf. A and M represent CO₂ assimilation and heat-producing metabolic processes, respectively.

where SR_net is the net absorption of solar radiation, LR_net is the net effect of emission and absorption, C is the convective heat transfer, λE is the evaporative heat loss, and M is the heat generated by respiration and other metabolic processes.

{SR}_{net} = {SR}_{abs} - {SR}_{A} - {SR}_{FL},

where SR_abs is the absorbed solar radiation, SR_A can be used to drive photochemical processes, and SR_FL is a small percentage of SR_abs emitted as fluorescence.

{LR}_{net} = {LR}_{in} - {LR}_{r} - {LR}_{em},

where LR_in is incident radiation, LR_r is reflected radiation, and LR_em is the emission of long-wave radiation.

The canopy temperature of rice is controlled by continuous changes in the heat balance between the environment and the heat conduction within the plant (Michaletz et al., 2016). Under consistent external environmental conditions, canopy temperature is primarily determined by the characteristics of the rice varieties and cultivation measures. It is an important characteristic of rice physiology and biochemistry that affects plant growth, leaf function, transpiration rate, photosynthetic capacity, and ultimately affects rice yield and quality (Balota et al., 2008; Jensen et al., 1990).

3 FACTORS INFLUENCING RICE CANOPY TEMPERATURE

The primary factors influencing rice canopy temperature are the natural environmental conditions in which the rice grows, rice variety characteristics and cultivation measures (Penfield, 2008). Natural environmental conditions mainly consist of meteorological factors, including atmospheric temperature, solar radiation, air humidity, wind speed, cloud cover, and the physical and chemical properties of the soil (e.g., moisture, nutrient content, and pH). The characteristics of a rice variety primarily include its individual morphological characteristics, physiological structure, and metabolic capacity (Jones et al., 2003). Factors associated with artificial cultivation include fertilizer application (Mon et al., 2016), irrigation (Garrity & O'Toole, 1994), pest and weed control, and resultant microenvironmental changes in light penetration in the field (Pinter et al., 1979). Multiple factors determine the rice canopy temperature, and thus it is insufficient to study the effect of one factor alone. However, the variability and interconnectedness of environmental factors, different rice varieties, and the integrated regulatory mechanisms make it difficult to determine the effects of multiple factors when studying rice canopy temperatures. Therefore, how these factors collectively influence rice canopy temperature together requires additional study.

3.1 Natural environmental conditions

Ambient temperature, light, water, air, and fertilizers in the natural environment affect the rice canopy temperature. Among these, air temperature presents with similar temperature changes to that of the rice canopy, and therefore can reflect such fluctuations. The difference between the rice canopy temperature and the ambient temperature causes heat convection and conduction. The extent of convection and conduction is related to the temperature difference between the leaves and the air, and the thermal conductivity of the boundary layer where the leaves exchange heat with the environment (Huband & Monteith, 1986).

The effects of light on the rice canopy temperature are primarily caused by radiant energy, rising air temperature, and changes in physiological and biochemical processes, such as transpiration and photosynthesis. Light acts directly on the rice canopies in the form of radiant energy, increasing its temperature, and is also an important energy source for atmospheric warming (Jones & Rotenberg, 2001; Monteith, 1962). When light satisfies the normal photosynthesis of the rice plants through short-wavelength, blue-violet light, large amounts of long-wavelength, red light are absorbed by the water in the rice plants, warming the plants. Light is also a major factor that directly affects rice transpiration, promotes stomatal opening, and reduces stomatal resistance. Strong light typically leads to a high temperature, which tends to cause water deficiency, stomatal closure, and an insufficient supply of CO₂, in turn leading to a decrease in the photosynthetic rate, thereby affecting plant growth; however, insufficient light causes chlorophyll decomposition and the possible yellowing of rice leaves (Dai & Sun, 2006). Environmental conditions, such as humidity, wind speed, and cloud cover, also indirectly affect the canopy temperature of rice by influencing photosynthesis and transpiration (Gautam et al., 2015).

Physicochemical properties, such as soil nutrient content, moisture content, and pH, can significantly affect rice growth, thereby affecting canopy temperature. Soil water deficit has a significant effect on leaf physiology, organ temperature, and canopy. A mild water deficit (soil water potential maintained at −15 to −20 kPa, which was at 0–20 cm below the surface) could produce a water-saving rice population that has resistance to heat stress, which would also result in a high yield (Yan et al., 2012). Stable soil nutrient content promotes metabolism and transpiration in rice plants, increases the heat dissipation ability of the population, and reduces the rice canopy temperature (Garrity et al., 1986). When the soil is water deficient, rice leaves undergo transient rolling and stomatal closure, and both photosynthesis and transpiration are temporarily suspended, resulting in an increase in canopy temperature (Followell et al., 2020).

3.2 Rice variety characteristics

Rice variety characteristics directly affect the thermal properties and physiological responses of rice plants. Increasing the rice tiller number, leaf area index, and plant height can result in a denser population niche, thereby decreasing ground heat dissipation, light penetration in the field, and canopy temperature (Yang et al., 2019). Different varieties of rice have different canopy temperatures with highly heritable characteristics. For example, qCTd11 associated with rice canopy temperature difference affects stomatal conductance and leaf photosynthesis. Therefore, canopy temperature can be a good indicator of stomatal conductance (Fukuda et al., 2018). In summary, the effects of rice variety characteristics on canopy temperature are primarily caused by (1) the thermal conductivity of the boundary layer of heat exchange between rice plants and the environment; (2) the metabolic heat generated during the physiological and biochemical reactions in rice plants; and (3) microclimate differences in the canopy caused by different qualities of rice populations.

3.2.1 Morphological structure

Rice leaves receive solar radiation directly and are an important location for heat exchange with the external environment. Leaves are the most sensitive organ for the short-term regulation of canopy temperature; and characteristics such as leaf size, shape, and angle toward the sun affect heat conduction between the rice plant and its environment, which affects canopy temperature (Parkhurst & Loucks, 1972; Smith, 1978). The area around the leaf boundary facilitates convective heat exchange and increasing this area can help reduce the canopy temperature to improve endurance during dry and hot conditions (Sikma et al., 2020). Mathematical analysis has revealed that cumulative temperature rather than non-cumulative photosynthetically active radiation can trigger exponential canopy growth (Fukuda et al., 2021).

3.2.2 Physiological structure

The energy generated by physiological metabolism in rice plants can properly regulate canopy temperature and avoid plant temperature imbalances via processes such as transpiration (Jones, 1999). Leaf physiology significantly influences canopy temperature. For example, higher vein density and stomatal pore area index facilitate higher transpiration capacity of the plants (Figure 2). Transpiration and physical traits of leaves provide more insurance against overheating, although transpiration is the more effective method cooling when water is sufficient (Hill et al., 2014; Lin et al., 2017). Physical leaf traits affect leaf temperature by influencing heat transfer, storage, and radiation (Levizou & Kyparissis, 2016). However, while increased stomatal conductance in rice leaves at high temperatures can increase the transpiration rate to reduce leaf temperature (Lo et al., 2020), changes in the external temperature affects photosynthetic intensity, thereby influencing stomatal opening and closure. These results indicated that leaf movements also have influence on leaf temperature. Transpiration is an important indicator of lower surface temperature in rice plants, as the evaporation of water transfers heat away from the plant surface. Under high temperature or water stress, stomatal closure reduces transpiration to preserve the existing water content of the plant.

3.3 Cultivation measures

In addition to climate and variety characteristics, cultivation practices also affect the rice canopy temperature. Common field water, fertilizer, and density management practices directly affect rice population growth and development, thereby affecting the canopy temperature (Perdomo et al., 2017). For example, nitrogen fertilizer application can significantly increase rice population biomass and leaf area index, cause population shading, reduce light energy reception within the rice plants (Ju et al., 2015), and reduce its canopy temperature. However, extremely high nitrogen fertilizer application tends to increase the ineffective leaf area index, the risk of disease in the rice plants (Ata-Ul-Karim et al., 2017; Xing et al., 2019), and canopy temperature. Proper planting density is necessary to create an adequate field microclimate. Low density is detrimental to achieving high yield, whereas high density leads to competition for nutrients and water within the rice population, making the formation of a suitable population structure difficult. Therefore, planting density along with water and fertilizer management can change the ventilation and light penetration in the paddy fields, and thus, are important measures to control paddy field microclimate and regulate rice canopy temperature (Liu et al., 2013).

4 VARIATIONS IN CANOPY TEMPERATURE OF RICE PLANTS AND THEIR RELATIONSHIP WITH YIELD

4.1 Rice canopy temperature variation patterns

Rice plants grow well at suitable temperatures and humidity. Under different growth conditions, rice has the ability to optimize its canopy energy balance, revealing that the canopy and air temperature changes are not completely synchronized. From the research (Figure 3), which was conducted during the rice-growing season (May to October) at research farm of Yangzhou University, Jiangsu Province, China (32°30′-N, 119°25′-E), we observed that the variations in rice canopy temperature were significantly smaller than those of the air temperature and could be maintained in a relatively stable state (Liu et al., 2018).

The differences between the rice canopy temperature and air temperature were the largest between 12 PM and 2 PM, which was the best time to observe the rice canopy temperature (Zheng et al., 2020). Rice can maintain certain canopy temperatures via physiological metabolism during different reproductive stages (Figure 4). While the changes in rice canopy temperature were consistent with changes in ambient or air temperature, they occurred slightly later and to a lesser extent than those in the air temperature.

4.2 Relationship between canopy temperature and yield of rice

Rice canopy temperature is the result of a comprehensive reaction between the growth environment and the self-regulation of rice plants. Air temperature affects the growth, yield and quality of rice by influencing the canopy temperature. Peng et al. (2004) reported that rice yield decreases because of the global warming induced increase in nighttime temperature, at a potential rate of 10% decrease for every 1°C increase in the minimum nighttime temperature. Significantly different effects of temperature have been observed among rice varieties (Shi et al., 2016), which may be caused by inherent differences in canopy temperature. Crop yield estimates are possible, from remotely acquired canopy temperatures and auxiliary air temperature measurements obtained during the period from head emergence until the cessation of head growth (Idso et al., 1977). The mean canopy temperature of different rice varieties has been observed to be significantly negatively correlated with grain yield (R² = −0.63**) and spikelet fertility (R² = −0.51**) (Garrity & O'Toole, 1995; O'Toole & Real, 1984).

All physiological and biochemical processes in rice are carried out within plant organs and interact to some extent with the canopy temperature. For instance, canopy temperature affects the leaf functional phase, chlorophyll content, transpiration, photosynthetic capacity, sucrose synthase activity, and internal anti-aging mechanisms that involve superoxide dismutase, peroxide dismutase, and malondialdehyde (Labuschagne et al., 2009; Wu et al., 2016). These effects resulted in different degrees of adaptability to temperature change with changes in canopy temperature for different rice varieties. However, there is a lack of studies that identify rice environmental temperature adaptation from the perspective of canopy temperature variability.

5 PRACTICAL APPLICATION OF THE CANOPY TEMPERATURE OF RICE PLANTS

Rice plants have the certain ability to regulate canopy energy balance. Numerous recent studies have used canopy temperature as an indicator for studying resistances to drought, heat, and other stressors (Ahmed et al., 2020; Carvalho et al., 2020). When rice plants respond to adverse external environments, such as drought and heat stress, metabolism changes and significantly increased canopy temperatures are observed before wilting, lesions, and death (Lawas et al., 2018). An infrared thermometer can be used as an effective and non-destructive method to monitor rice growth.

5.1 Detecting water stress

Water stress affects physiological and biochemical processes, such as transpiration, photosynthesis, and accumulation of osmoregulatory substances in rice (Wasaya et al., 2018), which greatly affects the rice canopy temperature. The degree of water stress can be reflected in the rice canopy temperature before phenomena such as leaf curling and wilting occur, which helps to promptly detect water deficiency (Blonquist et al., 2009). Red–Green–Blue images, Near-infrared imaging, infrared, fluorescence, and DroughtSpotter technology can detect the difference between drought-tolerant and susceptible plants, suggesting their value as high-throughput phenotyping methods for short breeding cycles as well as for functional genetic studies of tolerance to drought stress (Ikawa et al., 2018; Kim et al., 2020).

Crop water stress index (CWSI) is a comprehensive parameter reflecting the characteristics of the crop–soil–atmosphere system based on the difference between crop canopy temperature and air temperature (Idso et al., 1981),

CWSI = \frac{(T_{c} - T_{a}) - (T_{cl} - T_{a})}{(T_{cu} - T_{a}) - (T_{cl} - T_{a})}

where T_c is the canopy temperature, T_a is the air temperature, T_cl and T_cu correspond to the lower and upper limits, respectively. Based on the spatial distribution of canopy temperatures measured by thermal imaging cameras, the general sensitive CWSI, which was calculated by removing low temperatures, had a better performance in crop water stress diagnosis (Luan et al., 2021).

Jackson et al. (1988) provided the theoretical formula for canopy temperature–air temperature difference based on canopy surface energy balance and the Penman–Monteith formula, as follows:

T_{c} - T_{a} = \frac{r_{a}}{ρ C_{p}} \frac{γ (R_{n} - G) (1 + \frac{r_{c}}{r_{a}}) - ρ C_{p} \frac{e_{s} (T_{a}) - e_{a}}{r_{a}}}{∆ + γ (1 + \frac{r_{c}}{r_{a}})}

where

∆

is the slope of the saturated vapor pressure–temperature relation.

R_{n}

is the net radiation (

W m^{- 2}

G

is the heat flux into the surface (

W m^{- 2}

ρ

is the density of air

(Pa)

C_{p}

is the specific heat capacity of air at a constant pressure

(J {kg}^{- 1})

e_{s}

is the saturated vapor pressure of the air

(Pa)

T_{c}

e_{a}

is the vapor pressure of the air

(Pa)

γ

is the psychrometric constant

(Pa ° C^{- 1})

r_{c}

is the canopy resistance to water loss

(s m^{- 1}), and r_{a}

is the aerodynamic resistance

(s m^{- 1})

Drought affects the growth and yield of most modern rice varieties (Baisakh et al., 2020; Khahani et al., 2021). Grain yield correlated significantly with root growth along the soil profile, flowering time, and canopy temperature under drought conditions. The QTL lines with highest yield under drought also showed lower canopy temperature (Grondin et al., 2018; Susanto et al., 2019). With the development of remote sensing technology and infrared thermometry, canopy temperatures can be monitored on a large scale. Moran et al. (1994) proposed the water deficit index, for the integrated judgment of surface and air temperatures. This heralded the application of a canopy temperature-based water stress diagnostic index in actual production.

Zhao et al. (2018) analyzed water deficiency using a CWSI at the elongation and flowering stages, which confirmed the relationship between air–canopy temperature difference and field water stress, and established a monitoring system for rice water deficits based on the air–canopy temperature difference. Characterizing the physiological mechanisms behind major-effect drought-yield quantitative trait loci (QTLs) can provide an understanding of the function of the QTLs as well as plant responses to drought in general. The BIL and 2-QTL NILs showed consistently lower canopy temperatures than IR64 in the drought treatments (Henry et al., 2015). Recent studies on irrigation and drought resistance in relation to canopy temperature (Zhang et al., 2019) reported canopy temperature as a good indicator of drought stress in the field, as it indirectly measures stomatal conductance (Melandri et al., 2020). By measuring the rice canopy temperature, the growth status and water deficiency of the rice canopy can be determined to guide rice irrigation management (Mahan & Burke, 2015). Moreover, considering the sensitivity of the rice canopy temperature to soil moisture, establishing a canopy temperature-based field water deficit diagnosis can efficiently and quickly identify the field water status to develop irrigation systems (Akkuzu et al., 2013). According to the measured degree of water stress and canopy temperature performance of local extension varieties, establishing a corresponding water deficiency diagnostic model can enable an efficient and rapid understanding of field moisture conditions (Zheng et al., 2020).

5.2 Assessment of high-temperature heat damage

High-temperature heat damage is a major meteorological hazards experienced by rice, which typically results in reduced fertility and photosynthetic capacity, a shorter reproductive period, and lower yields and quality (Fu et al., 2016). The increased temperature was observed to increase the leaf temperature of the rice population, which responded with higher stomatal conductance, thereby accelerating the transpiration rate and reducing the increase in leaf temperature. When the temperature is extremely high, water rapidly passes through the leaf conduit, and transpiration through the stomata regulates the rice canopy temperature (Luan & Vico, 2021). Transpiration regulates canopy temperature under high temperatures (Karwa et al., 2020), and provides a theoretical basis for assessing high-temperature heat damage. However, this regulation is influenced by several factors, such as stomata closure, which prevents water loss when the soil is water deficient. Stomatal conductance, net photosynthetic rate, and soluble protein content of rice flag leaves decrease under the influence of high temperature. However, heat-tolerant varieties have a higher photosynthetic capacity than more sensitive varieties and have high root vigor and leaf antioxidant protection system capacities, which increase their resistance to high temperatures (Wang et al., 2019; Yang et al., 2017; Yao et al., 2007). Canopy temperature is suggested as a promising approach for detecting both heat and drought stress, which is likely to improve the accuracy of crop model responses to the effects of high temperatures on rice yields (Eyshi Rezaei et al., 2015).

5.3 Selection and breeding of improved rice varieties

The rice canopy temperature varied among different genotypes. For example, the canopy temperatures of drought- and heat-tolerant varieties were relatively constant under drought and high-temperature conditions, whereas the canopy temperature of varieties with poor tolerance increased sharply. Therefore, canopy temperature stability can be used as an indicator to distinguish more resistant varieties (Webber et al., 2018).

Rice canopy temperature is closely correlated with physiological biochemistry, yield, and quality, and is monitored using a high-throughput and non-destructive technique for selecting superior varieties (Prashar & Jones, 2014). Chaudhuri et al. (1986) were the first to use the correlation of canopy microclimate variation to screen drought-resistant genotypes of crops. During rice production, when rice encounters adverse conditions, such as low temperature, low radiation, salt stress, pests, and weeds (Baghel et al., 2020; Tian et al., 2020), its physiological metabolism is primarily affected, canopy temperature increases, and later, leaf spots, leaf wilt, and rice lodging appear. Canopy temperature changes are significantly earlier than those observed phenomena in the field. Monitoring rice growth is the primary measure for responding to stress and selecting varieties, and traditional monitoring methods are time consuming and inefficient (Lindenthal et al., 2005). By measuring canopy temperature, we can determine whether rice is normal in advance and develop corresponding cultivation measures. Therefore, canopy temperature can be used as a monitoring index of rice health, and as an important index of rice stress resistance in the process of variety selection.

6 METHODS FOR DETERMINING CANOPY TEMPERATURE IN RICE

6.1 Thermal infrared temperature measurement techniques

Canopy temperature is difficult to measure directly, but it can be estimated indirectly based on thermal imagery (Kim et al., 2018; Still et al., 2019). All objects constantly emit infrared radiation into the surrounding space, and hence, the surface temperature of an object can be accurately determined by measuring the infrared energy radiated by it (Jin et al., 2017), a technique first applied in measuring plant temperature in 1963 (Tanner, 1963). This technique has the advantages of convenience, speed, non-destructivity, and relatively small error and has been used in several aspects of agricultural studies (Jones et al., 2002). However, it is also influenced by the environment and therefore has the disadvantage of its reading being affected by the temperature around the leaf (Norman & Becker, 1995; Norman et al., 1995). However, with the recent development of related research, canopy temperature monitoring technology has also made great progress (Fuchs, 1990) in terms of accuracy, sensitivity, and ease of use. During its inception, this technology required taking the average value of the fixed orientation, angle, and multiple shots (Fuchs & Tanner, 1966; Lomas et al., 1971). The empirical model was developed with environmental temperature, humidity, and other indicators (Hatfield, 1985), which are powerful tools for exploring how canopy temperature changes with growing conditions and plant traits beyond what is observable in specific experiments. Existing crop canopy temperature models link canopy to growing conditions via simple empirical relations (Shao et al., 2019) or explicitly model the leaf or canopy energy balance (Webber et al., 2018). Mechanistic models representing plant physiology can estimate crop canopy temperature while better reflecting soil water and weather dynamics, as well as plant responses to environmental conditions.

6.2 Drone temperature measurement techniques

Owing to limitations in the efficiency and accuracy of canopy temperature measurement methods, normal infrared thermometers cannot be used to perform large-area canopy temperature measurements and are rarely used in actual production. With the development of drones and thermal imaging technology for agriculture, unmanned aerial vehicles (UAVs) equipped with high-precision infrared thermometry devices can achieve high-throughput, non-destructive, and large-area evaluation (Deery et al., 2016), the technology provides an efficient determination method with production applications and a technical basis for subsequent studies (Sui & Baggard, 2018).

This technology can be divided into two major aspects: the first of which is drone route planning. The lowest point of flight is the height at which the airflow generated by the propeller does not affect the ground environment. The flight height is determined by considering the efficiency of image acquisition and the accuracy of the data. In general, as the flight height increases, measurement accuracy decreases and measurement efficiency increases. The second major aspect involves the extraction of rice canopy temperature image data, which differs before and after rice row closing because of the effects of bare soil (Figure 5). Before rice row closing, the effects of bare soil are extremely significant in the captured thermal infrared images, particularly for the water surface temperature. When extracting the data at a later stage, the effects of bare soil were greatly avoided, and rice surface images in 15 holes with a large coverage area were selected per treatment to measure the canopy temperature of each hole, effectively preventing the effects of bare soil on the rice canopy temperature. The images collected after rice row closing were essentially free of bare soil effects, and five 1-m² areas were selected for each treatment using the five-point method to read the surface canopy temperature data according to the height and resolution of the images taken.

7 RESEARCH SUMMARY AND OUTLOOKS

We divided the relevant studies of canopy temperature into three parts: measurement techniques, regulatory mechanisms, and production applications (Table 1). These three components are interrelated and synergistic. Chronologically, we divided them into two groups.

TABLE 1. The relevant studies on canopy temperature from 1960s to present

Fields	Contents	Relevant studies
Measurement Techniques	Infrared Thermometer	Tanner (1963), Fuchs and Tanner (1966), Fuchs et al. (1967), Lomas et al. (1971), O'Toole and Real (1984), Huband and Monteith (1986), Fuchs (1990), Norman and Becker (1995), Prashar and Jones (2014), Deery et al. (2016), Liu et al. (2018), Sui and Baggard (2018), Zhang et al. (2019), Kim et al. (2020), Fukuda et al. (2021)
Regulatory Mechanism	Energy Transmission	Monteith and Szeicz (1962), Dong (1984), Hatfield (1985), Huband and Monteith (1986), Norman et al. (1995), Dai and Sun (2006), Lambers et al. (2008)
	Physiology & Biochemical Process	Jensen et al. (1990), Jones (1999), Jones et al. (2002), Lindenthal et al. (2005), Balota et al. (2008), Blonquist et al. (2009), Pieruschka et al. (2010), Takai et al. (2010), Maes & Steppe (2012), Mahan and Burke (2015), Lin et al. (2017), Cai et al. (2020), Sikma et al. (2020), Zheng et al. (2020)
	Genetic Research	Chaudhuri et al. (1986), Henry et al. (2015), Fukuda et al. (2018), Grondin et al. (2018), Susanto et al. (2019), Melandri et al. (2020)
Production Application	Water or Drought Stress	Idso et al. (1981), Fuchs (1990), Moran et al. (1994), Jones et al. (2003), Yan et al. (2012), Maes and Steppe (2012), Akkuzu et al. (2013), Mon et al. (2016), Lawas et al. (2018), Wasaya et al. (2018), Zhao et al. (2018), Zhang et al. (2019), Ahmed et al. (2020), Melandri et al. (2020), Lo et al. (2020), Baisakh et al. (2020), Khahani et al. (2021), Luan and Vico (2021), Luan et al. (2021)
	Heat Stress	Eyshi Rezaei et al. (2015), Gautam et al. (2015), Shi et al. (2016), Fu et al. (2016), Wu et al. (2016), Lawas et al. (2018), Webber et al. (2018), Tafesse et al. (2019), Wang et al. (2019), Karwa et al. (2020), Luan and Vico (2021)
	Salt, Heavy Metal Stress Pests & Diseases	Tian et al. (2020), Jin et al. (2017), Lindenthal et al. (2005)
	Yield & Quality	Idso et al. (1977), O'Toole and Real (1984), Garrity and O'Toole (1995), Siebert et al. (2014), Gabaldón-Leal et al. (2016), Kumar et al. (2017), Followell et al. (2020), Zheng et al. (2020), Yang et al. (2021)

The early stages of temperature measurement and energy conduction theory occurred from 1960 to 1999. Using the theory of energy transfer between plants and the environment, and the first adaptation of infrared thermometers in plants, researchers have studied the canopy temperature, mechanism of temperature maintenance in plants, morphological characteristics, and energy exchange with the environment. From the perspective of energy balance, these studies have built a theoretical foundation for canopy temperature research.

Theoretical and practical research has increased dramatically from 2000 to the present year. The measurement technology for canopy temperature has developed rapidly, and its accuracy and maneuverability have been strengthened. In particular, the UAVs carrying infrared high-precision equipment have realized large-scale, high-throughput, and dynamic monitoring of canopy temperatures, which has improved their applicability. Researchers have performed a series of physiological and biochemical studies and practical applications that mainly involved plant morphological characteristics, biochemical processes, QTL gene mapping, and genotypic varieties selection. For example, studies have shown that transpiration is the main driving force for controlling canopy temperature, selecting genotypes of drought and heat resistance, and obtaining high yields. Practical applications have focused on guiding irrigation and predicting yields. Combined with other indicators, canopy temperature can provide useful information for evaluating stress resistance (water, temperature, salt, heavy metal stress, pests, and diseases).

In summary, after over 60 years of theoretical and practical development, monitoring canopy temperature as an indicator of plant health now has a robust research basis. However, its large-scale applications still need to be further explored. Considering the problems and development trends in rice canopy temperature studies, there are still many aspects to investigate.

7.1 Strengthening research on the mechanisms influencing canopy temperature in rice

The mechanism of canopy temperature change in rice is the theoretical basis for its application in production. Previous studies have clarified that transpiration and photosynthesis are the main factors regulating canopy temperature in rice, and temperature stress, water stress, and other stressors lead to differences in canopy temperatures. However, multiple factors affect the changes in canopy temperature. The self-regulation mechanisms in rice canopies of different varieties in relation to physiological structures, such as differences in roots, stems, and leaves, require further clarification. Notably, existing studies typically use canopy temperature as a bridge between stress and rice growth, and there are relatively few studies on the response mechanism of rice canopy temperature to environmental changes. Future research should focus on developing rice crop models using physiological and ecological parameters to monitor changes in canopy temperature and the dynamic response of photosynthetic transpiration. This way, the mechanism influencing canopy temperature can play a greater role in assessing rice growth.

7.2 Improving methods and techniques for determining canopy temperature in rice

The accurate determination of rice canopy temperature is a prerequisite for a comprehensive analysis of its mechanism. Remote sensing and drone technology have been widely applied for temperature measurement; however, thermal imaging systems can only measure the rice surface temperature and not the internal temperature. Thus, the accuracy of thermal infrared cameras needs to be further improved. These weaknesses lead to limitations in perception, and therefore the development of faster, more accurate, and continuous temperature measurement devices is required. The analysis of rice canopy temperature data requires attention on the effects of non-climatic factors and their interactions with climatic factors, the autocorrelation of climatic elements, and the selection of appropriate spatial and temporal study scales. Moreover, the integration of multiple methods is important to improve the analysis of canopy temperature data. Examples include the collective assessment of statistical models, crop growth models, and observation trials, as well as the combination of crop growth models and remote sensing, drone monitoring, and multiple methods of crop phenotype observation, which can help improve the credibility and practicality of rice canopy temperature results.

7.3 Systematic impact assessment of canopy temperature on rice production

The effects of canopy temperature on the microclimate, rice yield and quality, and pest and disease occurrence and the effects of climate factors, such as temperature, light, moisture, CO₂ concentration, and extreme weather on the stabilization of rice canopy temperature should all be clarified according to the representative rice varieties of different rice cropping areas. The mechanism by which extreme weather affects rice canopy temperature and the role of fertilizer application, irrigation, pesticides, and other cultivation measures should also be investigated. An ecological monitoring and assessment system for rice crops should be constructed by studying the characteristics of rice canopy temperature, meteorological factors, and field water and fertilizer conditions to accelerate the refinement and application of canopy temperature-related research results, which will provide new methods for monitoring rice growth and predicting its yield and quality.

ACKNOWLEDGMENTS

We are grateful to Qing Lan Project of Yangzhou University Young Scholars Program and Poject Funded by Priority Academic Program Development of Jiangsu Higher Education Institutions (PAPD) for support to our research. We’d like to thank Hengyang Zhuang, Shiping Liu, and Tao Liu for their thoughts and feedback on this manuscript.

FUNDING INFORMATION

This study was supported by the National Natural Science Foundation of China (NSFC) (31801310); the Open Project Program of Joint International Research Laboratory of Agriculture and Agri-Product Safety, the Ministry of Education of China, Yangzhou University (JILAR-KF202012).

CONFLICT OF INTEREST

The authors have stated explicitly that there are no conflicts of interest in connection with this article.

Open Research

DATA AVAILABILITY STATEMENT

The data that support the findings of this study are openly available in [repository name e.g “figshare”] at (Liu et al., 2018, https://doi.org/10.1016/j.agrformet.2018.01.021; Li et al., 2017, ttps://doi.org/10.1093/jxb/erx101).

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Citing Literature

Volume12, Issue2

Special Issue:Food Security, 2030

March 2023

e424

Current status and prospects of rice canopy temperature research

Abstract

1 INTRODUCTION

2 VARIATIONS IN RICE CANOPY TEMPERATURE