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1. Thermal sensitivity across forest vertical profiles: patterns, mechanisms, and ecological implications

   https://doi.org/10.1111/nph.18539  

   Vinod, Nidhi; Slot, Martijn; McGregor, Ian_R; Ordway, Elsa_M; Smith, Marielle_N; Taylor, Tyeen_C; Sack, Lawren; Buckley, Thomas_N; Anderson‐Teixeira, Kristina_J (December 2022, New Phytologist)

   Summary Rising temperatures are influencing forests on many scales, with potentially strong variation vertically across forest strata. Using published research and new analyses, we evaluate how microclimate and leaf temperatures, traits, and gas exchange vary vertically in forests, shaping tree, and ecosystem ecology. In closed‐canopy forests, upper canopy leaves are exposed to the highest solar radiation and evaporative demand, which can elevate leaf temperature (T_leaf), particularly when transpirational cooling is curtailed by limited stomatal conductance. However, foliar traits also vary across height or light gradients, partially mitigating and protecting against the elevation of upper canopyT_leaf. Leaf metabolism generally increases with height across the vertical gradient, yet differences in thermal sensitivity across the gradient appear modest. Scaling from leaves to trees, canopy trees have higher absolute metabolic capacity and growth, yet are more vulnerable to drought and damagingT_leafthan their smaller counterparts, particularly under climate change. By contrast, understory trees experience fewer extreme highT_leaf's but have fewer cooling mechanisms and thus may be strongly impacted by warming under some conditions, particularly when exposed to a harsher microenvironment through canopy disturbance. As the climate changes, integrating the patterns and mechanisms reviewed here into models will be critical to forecasting forest–climate feedback. 

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2. The capacity to emit isoprene differentiates the photosynthetic temperature responses of tropical plant species

   https://doi.org/10.1111/pce.13564  

   Taylor, Tyeen C.; Smith, Marielle N.; Slot, Martijn; Feeley, Kenneth J. (June 2019, Plant, Cell & Environment)

   Abstract Experimental research shows that isoprene emission by plants can improve photosynthetic performance at high temperatures. But whether species that emit isoprene have higher thermal limits than non‐emitting species remains largely untested. Tropical plants are adapted to narrow temperature ranges and global warming could result in significant ecosystem restructuring due to small variations in species' thermal tolerances. We compared photosynthetic temperature responses of 26 co‐occurring tropical tree and liana species to test whether isoprene‐emitting species are more tolerant to high temperatures. We classified species as isoprene emitters versus non‐emitters based on published datasets. Maximum temperatures for net photosynthesis were ~1.8°C higher for isoprene‐emitting species than for non‐emitters, and thermal response curves were 24% wider; differences in optimum temperatures (T_opt) or photosynthetic rates at T_optwere not significant. Modelling the carbon cost of isoprene emission, we show that even strong emission rates cause little reduction in the net carbon assimilation advantage over non‐emitters at supraoptimal temperatures. Isoprene emissions may alleviate biochemical limitations, which together with stomatal conductance, co‐limit photosynthesis above T_opt. Our findings provide evidence that isoprene emission may be an adaptation to warmer thermal niches, and that emitting species may fare better under global warming than co‐occurring non‐emitting species. 

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3. Thermal acclimation of leaf respiration of tropical trees and lianas: response to experimental canopy warming, and consequences for tropical forest carbon balance

   https://doi.org/10.1111/gcb.12563  

   Slot, Martijn; Rey‐Sánchez, Camilo; Gerber, Stefan; Lichstein, Jeremy W; Winter, Klaus; Kitajima, Kaoru (September 2014, Global Change Biology)

   Abstract Climate warming is expected to increase respiration rates of tropical forest trees and lianas, which may negatively affect the carbon balance of tropical forests. Thermal acclimation could mitigate the expected respiration increase, but the thermal acclimation potential of tropical forests remains largely unknown. In a tropical forest in Panama, we experimentally increased nighttime temperatures of upper canopy leaves of three tree and two liana species by on average 3 °C for 1 week, and quantified temperature responses of leaf dark respiration. Respiration at 25 °C (R₂₅) decreased with increasing leaf temperature, but acclimation did not result in perfect homeostasis of respiration across temperatures. In contrast, Q₁₀of treatment and control leaves exhibited similarly high values (range 2.5–3.0) without evidence of acclimation. The decrease inR₂₅was not caused by respiratory substrate depletion, as warming did not reduce leaf carbohydrate concentration. To evaluate the wider implications of our experimental results, we simulated the carbon cycle of tropical latitudes (24°S–24°N) from 2000 to 2100 using a dynamic global vegetation model (LM3VN) modified to account for acclimation. Acclimation reduced the degree to which respiration increases with climate warming in the model relative to a no‐acclimation scenario, leading to 21% greater increase in net primary productivity and 18% greater increase in biomass carbon storage over the 21st century. We conclude that leaf respiration of tropical forest plants can acclimate to nighttime warming, thereby reducing the magnitude of the positive feedback between climate change and the carbon cycle. 

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4. No evidence of canopy-scale leaf thermoregulation to cool leaves below air temperature across a range of forest ecosystems

   https://doi.org/10.1073/pnas.2205682119  

   Still, Christopher J.; Page, Gerald; Rastogi, Bharat; Griffith, Daniel M.; Aubrecht, Donald M.; Kim, Youngil; Burns, Sean P.; Hanson, Chad V.; Kwon, Hyojung; Hawkins, Linnia; et al (September 2022, Proceedings of the National Academy of Sciences)

   Understanding and predicting the relationship between leaf temperature ( T leaf ) and air temperature ( T air ) 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 T leaf is maintained near photosynthetic temperature optima and below damaging temperature thresholds. Specifically, leaves should cool below T air at higher temperatures (i.e., > ∼25–30°C) leading to slopes <1 in T leaf / T air 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 ( T can ) 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 T can / T air 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 T can / T air relationship. Canopy structure also plays an important role in T can dynamics. Future climate warming is likely to lead to even greater T can , with attendant impacts on forest carbon cycling and mortality risk. 

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5. Optimal carbon partitioning helps reconcile the apparent divergence between optimal and observed canopy profiles of photosynthetic capacity

   https://doi.org/10.1111/nph.17199  

   Buckley, Thomas N. (March 2021, New Phytologist)

   Summary Photosynthetic capacity per unit irradiance is greater, and the marginal carbon revenue of water (∂A/∂E) is smaller, in shaded leaves than sunlit leaves, apparently contradicting optimization theory. I tested the hypothesis that these patterns arise from optimal carbon partitioning subject to biophysical constraints on leaf water potential.In a whole plant model with two canopy modules, I adjusted carbon partitioning, nitrogen partitioning and leaf water potential to maximize carbon profit or canopy photosynthesis, and recorded how gas exchange parameters compared between shaded and sunlit modules in the optimum.The model predicted that photosynthetic capacity per unit irradiance should be larger, and ∂A/∂Esmaller, in shaded modules compared to sunlit modules. This was attributable partly to radiation‐driven differences in evaporative demand, and partly to differences in hydraulic conductance arising from the need to balance marginal returns on stem carbon investment between modules. The model verified, however, that invariance in the marginal carbon revenue of N (∂A/∂N) is in fact optimal.The Cowan–Farquhar optimality solution (invariance of ∂A/∂E) does not apply to spatial variation within a canopy. The resulting variation in carbon–water economy explains differences in capacity per unit irradiance, reconciling optimization theory with observations. 

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