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• The drier climates predicted for many regions will result in reduced evaporative cooling, leading to leaf heat stress and enhanced mortality. The extent to which nonevaporative cooling can contribute to plant resilience under these increasingly stressful conditions is not well known at present. • Using a novel, high accuracy infrared system for the continuous measurement of leaf temperature in mature trees under field conditions, we assessed leaf-to-air temperature differences (ΔT leaf–air) of pine needles during drought. • On mid-summer days, ΔT leaf–air remained < 3°C, both in trees exposed to summer drought and in those provided with supplemental irrigation, which had a more than 10-fold higher transpiration rate. The nonevaporative cooling in the drought-exposed trees must be facilitated by low resistance to heat transfer, generating a large sensible heat flux, H. ΔT leaf–air was weakly related to variations in the radiation load and mean wind speed in the lower part of the canopy, but was dependent on canopy structure and within-canopy turbulence that enhanced the H. • Nonevaporative cooling is demonstrated as an effective cooling mechanism in needle-leaf trees which can be a critical factor in forest resistance to drying climates. The generation of a large H at the leaf scale provides a basis for the development of the previously identified canopy-scale ‘convector effect’.

• Temperature is a key control over biological activities from the cellular to the ecosystem scales. However, direct, high-precision measurements of surface temperature of small objects, such as leaves, under field conditions with large variations in ambient conditions remain rare. Contact methods, such as thermocouples, are prone to large errors. The use of noncontact remote-sensing methods, such as thermal infrared measurements, provides an ideal solution, but their accuracy has been low (c. 2°C) owing to the necessity for corrections for material emissivity and fluctuations in background radiation L bg. • A novel ‘dual-reference’ method was developed to increase the accuracy of infrared needle-leaf surface temperature measurements in the field. It accounts for variations in L bg and corrects for the systematic camera offset using two reference plates. • We accurately captured surface temperature and leaf-to-air temperature differences of needle-leaves in a forest ecosystem with large diurnal and seasonal temperature fluctuations with an uncertainty of ± 0.23°C and ± 0.28°C, respectively. • Routine high-precision leaf temperature measurements even under harsh field conditions, such as demonstrated here, opens the way for investigating a wide range of leaf-scale processes and their dynamics.

This article is a Commentary on Drake et al. (2020), 228: 1511–1523.

Current climate change scenarios indicate warmer temperatures and the potential for more extreme droughts in the tropics, such that a mechanistic understanding of the water cycle from individual trees to landscapes is needed to adequately predict future changes in forest structure and function. In this study, we contrasted physiological responses of tropical trees during a normal dry season with the extreme dry season due to the 2015-2016 El Niño-Southern Oscillation (ENSO) event. We quantified high resolution temporal dynamics of sap velocity (V ), stomatal conductance (g ) and leaf water potential (Ψ ) of multiple canopy trees, and their correlations with leaf temperature (T ) and environmental conditions [direct solar radiation, air temperature (T ) and vapor pressure deficit (VPD)]. The experiment leveraged canopy access towers to measure adjacent trees at the ZF2 and Tapajós tropical forest research (near the cities of Manaus and Santarém). The temporal difference between the peak of g (late morning) and the peak of VPD (early afternoon) is one of the major regulators of sap velocity hysteresis patterns. Sap velocity displayed species-specific diurnal hysteresis patterns reflected by changes in T . In the morning, T and sap velocity displayed a sigmoidal relationship. In the afternoon, stomatal conductance declined as T approached a daily peak, allowing Ψ to begin recovery, while sap velocity declined with an exponential relationship with T . In Manaus, hysteresis indices of the variables T -T and Ψ -T were calculated for different species and a significant difference ( < 0.01, α = 0.05) was observed when the 2015 dry season (ENSO period) was compared with the 2017 dry season (\"control scenario\"). In some days during the 2015 ENSO event, T approached 40°C for all studied species and the differences between T and T reached as high at 8°C (average difference: 1.65 ± 1.07°C). Generally, T was higher than T during the middle morning to early afternoon, and lower than T during the early morning, late afternoon and night. Our results support the hypothesis that partial stomatal closure allows for a recovery in Ψ during the afternoon period giving an observed counterclockwise hysteresis pattern between Ψ and T .

Climate change is contributing to an increased risk of flower damage by late spring frosts. Monitoring flower temperature is critical for the timely start of frost protection systems. However, there are many weak points that complicate the use of this method. The aims of this study were to: i) find the method of air temperature measurement with the best relationship to the surface temperature of plant tissues and ii) quantify the differences between plant tissues surface temperature and ambient temperature during different weather situations. The surface temperature of plant tissues (budding leaves of grapevine, apricot flower, and unripe pear fruit), air temperature and humidity in the radiation shield, wet bulb temperature and air temperature with an unsheltered thermometer were measured at ten-minute intervals in the spring months. The average temperatures obtained by the individual methods as well as the lowest temperatures were determined from each measurement. Differences between air temperatures and plant surface temperatures, including variation ranges, were also determined. An unsheltered thermometer, in which the energy balance corresponds approximately to that of the evaluated plant surfaces, provided the best relationship with plant tissue temperature. The air temperature measured by the standard method (in a Stevenson screen or in the radiation shield) was almost always higher than the temperature of the plant tissue during periods of negative energy balance. The difference between the minimum temperatures was approximately 0.5 °C. Temperatures more than 1.5 °C higher than the actual temperature of plant tissues were measured in extreme cases.

Understanding and predicting the relationship between leaf temperature (Tleaf ) and air temperature (Tair ) is essential for projecting responses to a warming climate, as studies suggest that many forests are near thermal thresholds for carbon uptake. Based on leaf measurements, the limited leaf homeothermy hypothesis argues that daytime Tleaf is maintained near photosynthetic temperature optima and below damaging temperature thresholds. Specifically, leaves should cool below Tair at higher temperatures (i.e., > ∼25–30°C) leading to slopes <1 in Tleaf/Tair relationships and substantial carbon uptake when leaves are cooler than air. This hypothesis implies that climate warming will be mitigated by a compensatory leaf cooling response. A key uncertainty is understanding whether such thermoregulatory behavior occurs in natural forest canopies. We present an unprecedented set of growing season canopy-level leaf temperature (Tcan ) data measured with thermal imaging at multiple well-instrumented forest sites in North and Central America. Our data do not support the limited homeothermy hypothesis: canopy leaves are warmer than air during most of the day and only cool below air in mid to late afternoon, leading to Tcan/Tair slopes >1 and hysteretic behavior. We find that the majority of ecosystem photosynthesis occurs when canopy leaves are warmer than air. Using energy balance and physiological modeling, we show that key leaf traits influence leaf-air coupling and ultimately the Tcan/Tair relationship. Canopy structure also plays an important role in Tcan dynamics. Future climate warming is likely to lead to even greater Tcan , with attendant impacts on forest carbon cycling and mortality risk.

As evaporation of water is an energy-demanding process, increasing evapotranspiration rates decrease the surface temperature (Ts) of leaves and plants. Based on this principle, ground-based thermal remote sensing has become one of the most important methods for estimating evapotranspiration and drought stress and for irrigation. This paper reviews its application in agriculture. The review consists of four parts. First, the basics of thermal remote sensing are briefly reviewed. Second, the theoretical relation between Ts and the sensible and latent heat flux is elaborated. A modelling approach was used to evaluate the effect of weather conditions and leaf or vegetation properties on leaf and canopy temperature. Ts increases with increasing air temperature and incoming radiation and with decreasing wind speed and relative humidity. At the leaf level, the leaf angle and leaf dimension have a large influence on Ts; at the vegetation level, Ts is strongly impacted by the roughness length; hence, by canopy height and structure. In the third part, an overview of the different ground-based thermal remote sensing techniques and approaches used to estimate drought stress or evapotranspiration in agriculture is provided. Among other methods, stress time, stress degree day, crop water stress index (CWSI), and stomatal conductance index are discussed. The theoretical models are used to evaluate the performance and sensitivity of the most important methods, corroborating the literature data. In the fourth and final part, a critical view on the future and remaining challenges of ground-based thermal remote sensing is presented.

Leaf temperature exerts an important impact on the microenvironment and physiological processes of leaves. Plants from different habitats have different strategies to regulate leaf temperature. The relative importance of physical traits and transpiration for leaf temperature regulation in the hot habitat is still unclear. We investigated 22 leaf physical traits, transpiration, and thermal properties of 38 canopy species of seedlings in a greenhouse, including 18 dominant species from a hot wet habitat (HW) and 20 dominant species from a hot dry habitat (HD). To separate the impact of transpiration and leaf physical traits on leaf temperature, we measured the diurnal courses of leaf temperatures with and without transpiration. The temperature of a reference leaf beside each individual was measured simultaneously to render temperatures comparable. Generally, the species from HD showed lower leaf temperatures than the species from HW under the same conditions. Both transpiration capacity and cooling effect of leaf physical traits were stronger for the plants from HD. Active transpiration provides a suitable thermal environment for photosynthesis, while xeromorphic leaves can dampen heat stress when transpiration is suppressed. Higher vein density and stomatal pore area index (SPI) facilitated higher transpiration capacity of the plants from HD. Meanwhile, shorter leaves and thinner lower epidermis of the plants from HD were more efficient in heat transfer, although relationships were much weaker than the synergic effect of all the physical traits. Our results confirmed that transpiration and leaf physical traits provided double insurance for avoiding overheating, particularly for plant from HD. We emphasize that transpiration is a more effective way to cool leaves than physical traits when water is sufficient, which may be an important adaptation for plant from HD where rainfall is sporadic. Our results provide further insight into the relationship between physical traits and transpiration for the regulation of leaf temperature, and the co‐evolution of gas exchange and thermal regulation of leaves. A plain language summary is available for this article. Plain Language Summary

The thermal limit of ectotherms provides an estimate of vulnerability to climate change. It differs between contrasting microhabitats, consistent with thermal ecology predictions that a species’ temperature sensitivity matches the microclimate it experiences. However, observed thermal limits may differ between ectotherms from the same environment, challenging this theory. We resolved this apparent paradox by showing that ectotherm activity generates microclimatic deviations large enough to account for differences in thermal limits between species from the same microhabitat. We studied upper lethal temperature, effect of feeding mode on plant gas exchange, and temperature of attacked leaves in a community of six arthropod species feeding on apple leaves. Thermal limits differed by up to 8 °C among the species. Species that caused an increase in leaf transpiration (+182%), thus cooling the leaf, had a lower thermal limit than those that decreased leaf transpiration (−75%), causing the leaf to warm up. Therefore, cryptic microclimatic variations at the scale of a single leaf determine the thermal limit in this community of herbivores. We investigated the consequences of these changes in plant transpiration induced by plant–insect feedbacks for species vulnerability to thermal extremes. Warming tolerance was similar between species, at ±2 °C, providing little margin for resisting increasingly frequent and intense heat waves. The thermal safety margin (the difference between thermal limit and temperature) was greatly overestimated when air temperature or intact leaf temperature was erroneously used. We conclude that feedback processes define the vulnerability of species in the phyllosphere, and beyond, to thermal extremes.