skip to main content

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

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 (Tleaf), 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 canopyTleaf. 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 damagingTleafthan their smaller counterparts, particularly under climate change. By contrast, understory trees experience fewer extreme highTleaf'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.

more » « less
Award ID(s):
1951244 2017949
Author(s) / Creator(s):
 ;  ;  ;  ;  ;  ;  ;  ;  
Publisher / Repository:
Date Published:
Journal Name:
New Phytologist
Medium: X Size: p. 22-47
["p. 22-47"]
Sponsoring Org:
National Science Foundation
More Like this
  1. Abstract

    Tropical forest canopies cycle vast amounts of carbon, yet we still have a limited understanding of how these critical ecosystems will respond to climate warming. We implemented in situ leaf‐level + 3°C experimental warming from the understory to the upper canopy of two Puerto Rican tropical tree species,Guarea guidoniaandOcotea sintenisii. After approximately 1 month of continuous warming, we assessed adjustments in photosynthesis, chlorophyll fluorescence, stomatal conductance, leaf traits and foliar respiration. Warming did not alter net photosynthetic temperature response for either species; however, the optimum temperature ofOcoteaunderstory leaf photosynthetic electron transport shifted upward. There was noOcotearespiratory treatment effect, whileGuarearespiratory temperature sensitivity (Q10) was down‐regulated in heated leaves. The optimum temperatures for photosynthesis (Topt) decreased 3–5°C from understory to the highest canopy position, perhaps due to upper canopy stomatal conductance limitations.Guareaupper canopyToptwas similar to the mean daytime temperatures, whileOcoteacanopy leaves often operated aboveTopt. With minimal acclimation to warmer temperatures in the upper canopy, further warming could put these forests at risk of reduced CO2uptake, which could weaken the overall carbon sink strength of this tropical forest.

    more » « less
  2. Summary

    Seasonal dynamics in the vertical distribution of leaf area index (LAI) may impact the seasonality of forest productivity in Amazonian forests. However, until recently, fine‐scale observations critical to revealing ecological mechanisms underlying these changes have been lacking.

    To investigate fine‐scale variation in leaf area with seasonality and drought we conducted monthly ground‐based LiDAR surveys over 4 yr at an Amazon forest site. We analysed temporal changes in vertically structuredLAIalong axes of both canopy height and light environments.

    Upper canopyLAIincreased during the dry season, whereas lower canopyLAIdecreased. The low canopy decrease was driven by highly illuminated leaves of smaller trees in gaps. By contrast, understoryLAIincreased concurrently with the upper canopy. Hence, tree phenological strategies were stratified by height and light environments. Trends were amplified during a 2015–2016 severe El Niño drought.

    Leaf area low in the canopy exhibited behaviour consistent with water limitation. Leaf loss from short trees in high light during drought may be associated with strategies to tolerate limited access to deep soil water and stressful leaf environments. Vertically and environmentally structured phenological processes suggest a critical role of canopy structural heterogeneity in seasonal changes in Amazon ecosystem function.

    more » « less
  3. 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. 
    more » « less
  4. Abstract

    Climate change is stressing many forests around the globe, yet some tree species may be able to persist through acclimation and adaptation to new environmental conditions. The ability of a tree to acclimate during its lifetime through changes in physiology and functional traits, defined here as its acclimation potential, is not well known.

    We investigated the acclimation potential of trembling aspenPopulus tremuloidesand ponderosa pinePinus ponderosatrees by examining within‐species variation in drought response functional traits across both space and time, and how trait variation influences drought‐induced tree mortality. We measured xylem tension, morphological traits and physiological traits on mature trees in southwestern Colorado, USA across a climate gradient that spanned the distribution limits of each species and 3 years with large differences in climate.

    Trembling aspen functional traits showed high within‐species variation, and osmotic adjustment and carbon isotope discrimination were key determinants for increased drought tolerance in dry sites and in dry years. However, trembling aspen trees at low elevation were pushed past their drought tolerance limit during the severe 2018 drought year, as elevated mortality occurred. Higher specific leaf area during drought was correlated with higher percentages of canopy dieback the following year. Ponderosa pine functional traits showed less within‐species variation, though osmotic adjustment was also a key mechanism for increased drought tolerance. Remarkably, almost all traits varied more year‐to‐year than across elevation in both species.

    Our results shed light on the scope and limits of intraspecific trait variation for mediating drought responses in key southwestern US tree species and will help improve our ability to model and predict forest responses to climate change.

    Read the freePlain Language Summaryfor this article on the Journal blog.

    more » « less
  5. Abstract

    Understanding variation in key functional traits across gradients in high diversity systems and the ecology of community changes along gradients in these systems is crucial in light of conservation and climate change. We examined inter‐ and intraspecific variation in leaf mass per area (LMA) of sun and shade leaves along a 3330‐m elevation gradient in Peru, and in sun leaves across a forest–savanna vegetation gradient in Brazil. We also comparedLMAvariance ratios (T‐statistics metrics) to null models to explore internal (i.e., abiotic) and environmental filtering on community structure along the gradients. Community‐weightedLMAincreased with decreasing forest cover in Brazil, likely due to increased light availability and water stress, and increased with elevation in Peru, consistent with the leaf economic spectrum strategy expected in colder, less productive environments. A very high species turnover was observed along both environmental gradients, and consequently, the first source of variation inLMAwas species turnover. Variation inLMAat the genus or family levels was greater in Peru than in Brazil. Using dominant trees to examine possible filters on community assembly, we found that in Brazil, internal filtering was strongest in the forest, while environmental filtering was observed in the dry savanna. In Peru, internal filtering was observed along 80% of the gradient, perhaps due to variation in taxa or interspecific competition. Environmental filtering was observed at cloud zone edges and in lowlands, possibly due to water and nutrient availability, respectively. These results related to variation inLMAindicate that biodiversity in species rich tropical assemblages may be structured by differential niche‐based processes. In the future, specific mechanisms generating these patterns of variation in leaf functional traits across tropical environmental gradients should be explored.

    more » « less