Within-canopy experimental leaf warming induces photosynthetic decline instead of acclimation in two northern hardwood species

Northern hardwood forests are experiencing higher temperatures and more extreme heat waves, potentially altering plant physiological processes. We implemented in-situ leaf-level warming along a vertical gradient within a mature forest canopy to investigate photosynthetic acclimation potential of two northern hardwood species, Acer saccharum and Tilia americana. After 7 days of +3°C warming, photosynthetic acclimation was assessed by measuring differences between heated and control photosynthetic rates (Aopt) at leaf optimum temperatures (Topt). We also measured the effects of warming and height on maximum rates of Rubisco carboxylation, stomatal conductance, transpiration, and leaf traits: leaf area, leaf mass per area, leaf nitrogen, and leaf water content. We found no evidence of photosynthetic acclimation for either species, but rather Aopt declined with warming overall. We found slight shifts in LMA and Narea, leaf traits associated with photosynthetic capacity, after 1 week of experimental warming. T. americana LMA and Narea was lower in the upper canopy heated leaves than in the control leaves, contributing a shift in Narea height distribution in the heated leaves. T. americana showed evidence of greater resiliency to warming, with greater thermoregulation, physiological plasticity, and evapotranspiration. As expected, Aopt of A. saccharum increased with height, but Aopt of T. americana was highest in the sub canopy, possibly due to constraints on leaf water balance and photosynthetic capacity in the upper canopy. Thus, models relying on canopy height or light environment may incorrectly estimate vertical variation of photosynthetic capacity. If these species are not able to acclimate to warmer temperatures, we could see alteration of plant carbon balance of these two key northern hardwood species

Temperate and tropical forest canopies are already functioning beyond their thermal thresholds for photosynthesis

Tropical tree species have evolved under very narrow temperature ranges compared to temperate forest species. Studies suggest that tropical trees may be more vulnerable to continued warming compared to temperate species, as tropical trees have shown declines in growth and photosynthesis at elevated temperatures. However, regional and global vegetation models lack the data needed to accurately represent such physiological responses to increased temperatures, especially for tropical forests. To address this need, we compared instantaneous photosynthetic temperature responses of mature canopy foliage, leaf temperatures, and air temperatures across vertical canopy gradients in three forest types: tropical wet, tropical moist, and temperate deciduous. Temperatures at which maximum photosynthesis occurred were greater in the tropical forests canopies than the temperate canopy (30 ± 0.3 °C vs. 27 ± 0.4 °C). However, contrary to expectations that tropical species would be functioning closer to threshold temperatures, photosynthetic temperature optima was exceeded by maximum daily leaf temperatures, resulting in sub-optimal rates of carbon assimilation for much of the day, especially in upper canopy foliage (\u3e10 m). If trees are unable to thermally acclimate to projected elevated temperatures, these forests may shift from net carbon sinks to sources, with potentially dire implications to climate feedbacks and forest community composition

On the shoulders of giants: Continuing the legacy of large-scale ecosystem manipulation experiments in Puerto Rico

There is a long history of experimental research in the Luquillo Experimental Forest in Puerto Rico. These experiments have addressed questions about biotic thresholds, assessed why communities vary along natural gradients, and have explored forest responses to a range of both anthropogenic and non-anthropogenic disturbances. Combined, these studies cover many of the major disturbances that affect tropical forests around the world and span a wide range of topics, including the effects of forest thinning, ionizing radiation, hurricane disturbance, nitrogen deposition, drought, and global warming. These invaluable studies have greatly enhanced our understanding of tropical forest function under different disturbance regimes and informed the development of management strategies. Here we summarize the major field experiments that have occurred within the Luquillo Experimental Forest. Taken together, results from the major experiments conducted in the Luquillo Experimental Forest demonstrate a high resilience of Puerto Rico’s tropical forests to a variety of stressors

Infrared heater system for warming tropical forest understory plants and soils

The response of tropical forests to global warming is one of the largest uncertainties in predicting the future carbon balance of Earth. To determine the likely effects of elevated temperatures on tropical forest understory plants and soils, as well as other ecosystems, an infrared (IR) heater system was developed to provide in situ warming for the Tropical Responses to Altered Climate Experiment (TRACE) in the Luquillo Experimental Forest in Puerto Rico. Three replicate heated 4‐m‐diameter plots were warmed to maintain a 4°C increase in understory vegetation compared to three unheated control plots, as sensed by IR thermometers. The equipment was larger than any used previously and was subjected to challenges different from those of many temperate ecosystem warming systems, including frequent power surges and outages, high humidity, heavy rains, hurricanes, saturated clayey soils, and steep slopes. The system was able to maintain the target 4.0°C increase in hourly average vegetation temperatures to within ± 0.1°C. The vegetation was heterogeneous and on a 21° slope, which decreased uniformity of the warming treatment on the plots; yet, the green leaves were fairly uniformly warmed, and there was little difference among 0–10 cm depth soil temperatures at the plot centers, edges, and midway between. Soil temperatures at the 40–50 cm depth increased about 3°C compared to the controls after a month of warming. As expected, the soil in the heated plots dried faster than that of the control plots, but the average soil moisture remained adequate for the plants. The TRACE heating system produced an adequately uniform warming precisely controlled down to at least 50‐cm soil depth, thereby creating a treatment that allows for assessing mechanistic responses of tropical plants and soil to warming, with applicability to other ecosystems. No physical obstacles to scaling the approach to taller vegetation (i.e., trees) and larger plots were observed

Photosynthetic and Respiratory Acclimation of Understory Shrubs in Response to in situ Experimental Warming of a Wet Tropical Forest

Despite the importance of tropical forests to global carbon balance, our understanding of how tropical plant physiology will respond to climate warming is limited. In addition, the contribution of tropical forest understories to global carbon cycling is predicted to increase with rising temperatures, however, in situ warming studies of tropical forest plants to date focus only on upper canopies. We present results of an in situ field-scale +4°C understory infrared warming experiment in Puerto Rico (Tropical Responses to Altered Climate Experiment; TRACE). We investigated gas exchange responses of two common understory shrubs, Psychotria brachiata and Piper glabrescens, after exposure to 4 and 8 months warming. We assessed physiological acclimation in two ways: (1) by comparing plot-level physiological responses in heated versus control treatments before and after warming, and (2) by examining physiological responses of individual plants to variation in environmental drivers across all plots, seasons, and treatments. P. brachiata has the capacity to up-regulate (i.e., acclimate) photosynthesis through broadened thermal niche and up-regulation of photosynthetic temperature optimum (Topt) with warmer temperatures. P. glabrescens, however, did not upregulate any photosynthetic parameter, but rather experienced declines in the rate of photosynthesis at the optimum temperature (Aopt), corresponding with lower stomatal conductance under warmer daily temperatures. Contrary to expectation, neither species showed strong evidence for respiratory acclimation. P. brachiata down-regulated basal respiration with warmer daily temperatures during the drier winter months only. P. glabrescens showed no evidence of respiratory acclimation. Unexpectedly, soil moisture, was the strongest environmental driver of daily physiological temperature responses, not vegetation temperature. Topt increased, while photosynthesis and basal respiration declined as soils dried, suggesting that drier conditions negatively affected carbon uptake for both species. Overall, P. brachiata, an early successional shrub, showed higher acclimation potential to daily temperature variations, potentially mitigating negative effects of chronic warming. The negative photosynthetic response to warming experienced by P. glabrescens, a mid-successional shrub, suggests that this species may not be able to as successfully tolerate future, warmer temperatures. These results highlight the importance of considering species when assessing climate change and relay the importance of soil moisture on plant function in large-scale warming experiments

Predicting effects of climate change on productivity and persistence of forest trees

Global climate change increases uncertainty in sustained functioning of forest ecosystems. Forest canopies are a key link between terrestrial ecosystems, the atmosphere, and climate. Here, we introduce research presented at the 66th meeting of the Ecological Society of Japan in the symposium “Structure and function of forest canopies under climate change.” Old-growth forest carbon stores are the largest and may be the most vulnerable to climate change as the balance between sequestration and emission could easily be tipped. Detailed structural analysis of individual large, old trees shows they are allocating wood to the trunk and crown in patterns that cannot be deduced from ground, thus can be used to more accurately quantify total forest carbon and sequestration. Slowly migrating species sensitive to novel climatic conditions will have to acclimate at the individual level. Accounting for physiological responses of trees to climate change will improve predictions of future species distributions and subsequent functioning of forest ecosystems. Field experiments manipulating temperature and precipitation show how trees compensate physiologically to mitigate for higher temperatures and drought. However, it is difficult to measure acclimation responses over long timeframes. Intraindividual trait variation is proposed as an indicator of acclimation potential of trees to future conditions and suggests that acclimation potential may vary among regional populations within a species. Integrating whole-tree structural data with physiological data offers a promising avenue for understanding how trees will respond to climatic shifts

Understory plant communities show resistance to drought, hurricanes, and experimental warming in a wet tropical forest

Global climate change has led to rising temperatures and to more frequent and intense climatic events, such as storms and droughts. Changes in climate and disturbance regimes can have non-additive effects on plant communities and result in complicated legacies we have yet to understand. This is especially true for tropical forests, which play a significant role in regulating global climate. We used understory vegetation data from the Tropical Responses to Altered Climate Experiment (TRACE) in Puerto Rico to evaluate how plant communities responded to climate warming and disturbance. The TRACE understory vegetation was exposed to a severe drought (2015), 2 years of experimental warming (4°C above ambient in half of the plots, 2016–2017 and 2018–2019), and two major hurricanes (Irma and María, September 2017). Woody seedlings and saplings were censused yearly from 2015 to 2019, with an additional census in 2015 after the drought ended. We evaluated disturbance-driven changes in species richness, diversity, and composition across ontogeny. We then used Bayesian predictive trait modeling to assess how species responded to disturbance and how this might influence the functional structure of the plant community. Our results show decreased seedling richness after hurricane disturbance, as well as increased sapling richness and diversity after warming. We found a shift in species composition through time for both seedlings and saplings, yet the individual effects of each disturbance were not significant. At both ontogenetic stages, we observed about twice as many species responding to experimental warming as those responding to drought and hurricanes. Predicted changes in functional structure point to disturbance-driven functional shifts toward a mixture of fast-growing and drought-tolerant species. Our findings demonstrate that the tropical forest understory community is more resistant to climatic stressors than expected, especially at the sapling stage. However, early signs of changes in species composition suggest that, in a warming climate with frequent droughts and hurricanes, plant communities might shift over time toward fast-growing or drought-tolerant species

Tropical understory herbaceous community responds more strongly to hurricane disturbance than to experimental warming

Ecology and Evolution published by John Wiley & Sons Ltd. The effects of climate change on tropical forests may have global consequences due to the forests’ high biodiversity and major role in the global carbon cycle. In this study, we document the effects of experimental warming on the abundance and composition of a tropical forest floor herbaceous plant community in the Luquillo Experimental Forest, Puerto Rico. This study was conducted within Tropical Responses to Altered Climate Experiment (TRACE) plots, which use infrared heaters under free-air, open-field conditions, to warm understory vegetation and soils + 4°C above nearby control plots. Hurricanes Irma and María damaged the heating infrastructure in the second year of warming, therefore, the study included one pretreatment year, one year of warming, and one year of hurricane response with no warming. We measured percent leaf cover of individual herbaceous species, fern population dynamics, and species richness and diversity within three warmed and three control plots. Results showed that one year of experimental warming did not significantly affect the cover of individual herbaceous species, fern population dynamics, species richness, or species diversity. In contrast, herbaceous cover increased from 20% to 70%, bare ground decreased from 70% to 6%, and species composition shifted pre to posthurricane. The negligible effects of warming may have been due to the short duration of the warming treatment or an understory that is somewhat resistant to higher temperatures. Our results suggest that climate extremes that are predicted to increase with climate change, such as hurricanes and droughts, may cause more abrupt changes in tropical forest understories than longer-term sustained warming

Understory plant communities show resistance to drought, hurricanes, and experimental warming in a wet tropical forest

Global climate change has led to rising temperatures and to more frequent and intense climatic events, such as storms and droughts. Changes in climate and disturbance regimes can have non-additive effects on plant communities and result in complicated legacies we have yet to understand. This is especially true for tropical forests, which play a significant role in regulating global climate. We used understory vegetation data from the Tropical Responses to Altered Climate Experiment (TRACE) in Puerto Rico to evaluate how plant communities responded to climate warming and disturbance. The TRACE understory vegetation was exposed to a severe drought (2015), 2 years of experimental warming (4°C above ambient in half of the plots, 2016–2017 and 2018–2019), and two major hurricanes (Irma and María, September 2017). Woody seedlings and saplings were censused yearly from 2015 to 2019, with an additional census in 2015 after the drought ended. We evaluated disturbance-driven changes in species richness, diversity, and composition across ontogeny. We then used Bayesian predictive trait modeling to assess how species responded to disturbance and how this might influence the functional structure of the plant community. Our results show decreased seedling richness after hurricane disturbance, as well as increased sapling richness and diversity after warming. We found a shift in species composition through time for both seedlings and saplings, yet the individual effects of each disturbance were not significant. At both ontogenetic stages, we observed about twice as many species responding to experimental warming as those responding to drought and hurricanes. Predicted changes in functional structure point to disturbance-driven functional shifts toward a mixture of fast-growing and drought-tolerant species. Our findings demonstrate that the tropical forest understory community is more resistant to climatic stressors than expected, especially at the sapling stage. However, early signs of changes in species composition suggest that, in a warming climate with frequent droughts and hurricanes, plant communities might shift over time toward fast-growing or drought-tolerant species