Sensitivity and threshold dynamics of Pinus strobus and Quercus spp. in response to experimental and naturally occurring severe droughts.

Increased drought frequency and severity are a pervasive global threat, yet the capacity of mesic temperate forests to maintain resilience in response to drought remains poorly understood. We deployed a throughfall removal experiment to simulate a once in a century drought in New Hampshire, USA, which coupled with the region-wide 2016 drought, intensified moisture stress beyond that experienced in the lifetimes of our study trees. To assess the sensitivity and threshold dynamics of two dominant northeastern tree genera (Quercus and Pinus), we monitored sap flux density (Js), leaf water potential and gas exchange, growth and intrinsic water-use efficiency (iWUE) for one pretreatment year (2015) and two treatment years (2016-17). Results showed that Js in pine (Pinus strobus L.) declined abruptly at a soil moisture threshold of 0.15 m3 m-3, whereas oak's (Quercus rubra L. and Quercus velutina Lam.) threshold was 0.11 m3 m-3-a finding consistent with pine's more isohydric strategy. Nevertheless, once oaks' moisture threshold was surpassed, Js declined abruptly, suggesting that while oaks are well adapted to moderate drought, they are highly susceptible to extreme drought. The radial growth reduction in response to the 2016 drought was more than twice as great for pine as for oaks (50 vs 18%, respectively). Despite relatively high precipitation in 2017, the oaks' growth continued to decline (low recovery), whereas pine showed neutral (treatment) or improved (control) growth. The iWUE increased in 2016 for both treatment and control pines, but only in treatment oaks. Notably, pines exhibited a significant linear relationship between iWUE and precipitation across years, whereas the oaks only showed a response during the driest conditions, further underscoring the different sensitivity thresholds for these species. Our results provide new insights into how interactions between temperate forest tree species' contrasting physiologies and soil moisture thresholds influence their responses and resilience to extreme drought.

Tropical rainforest carbon sink declines during El Niño as a result of reduced photosynthesis and increased respiration rates.

Changes in tropical forest carbon sink strength during El Niño Southern Oscillation (ENSO) events can indicate future behavior under climate change. Previous studies revealed ˜6 Mg C ha-1  yr-1 lower net ecosystem production (NEP) during ENSO year 1998 compared with non-ENSO year 2000 in a Costa Rican tropical rainforest. We explored environmental drivers of this change and examined the contributions of ecosystem respiration (RE) and gross primary production (GPP) to this weakened carbon sink. For 1998-2000, we estimated RE using chamber-based respiration measurements, and we estimated GPP in two ways: using (1) the canopy process model MAESTRA, and (2) combined eddy covariance and chamber respiration data. MAESTRA-estimated GPP did not statistically differ from GPP estimated using approach 2, but was ˜ 28% greater than published GPP estimates for the same site and years using eddy covariance data only. A 7% increase in RE (primarily increased soil respiration) and a 10% reduction in GPP contributed equally to the difference in NEP between ENSO year 1998 and non-ENSO year 2000. A warming and drying climate for tropical forests may yield a weakened carbon sink from both decreased GPP and increased RE. Understanding physiological acclimation will be critical for the large carbon stores in these ecosystems.

Canopy gradients in leaf functional traits for species that differ in growth strategies and shade tolerance.

In temperate deciduous forests, vertical gradients in leaf mass per area (LMA) and area-based leaf nitrogen (Narea) are strongly controlled by gradients in light availability. While there is evidence that hydrostatic constraints on leaf development may diminish LMA and Narea responses to light, inherent differences among tree species may also influence leaf developmental and morphological response to light. We investigated vertical gradients in LMA, Narea and leaf carbon isotope composition (δ13C) for three temperate deciduous species (Carpinus caroliniana Walter, Fagus grandifolia Ehrh., Liriodendron tulipifera L.) that differed in growth strategy (e.g., indeterminate and determinate growth), shade tolerance and leaf area to sapwood ratio (Al:As). Leaves were sampled across a broad range of light conditions within three vertical layers of tree crowns to maximize variation in light availability at each height and to minimize collinearity between light and height. All species displayed similar responses to light with respect to Narea and δ13C, but not for LMA. Light was more important for gradients in LMA for the shade-tolerant (C. caroliniana) and -intolerant (L. tulipifera) species with indeterminate growth, and height (e.g., hydrostatic gradients) and light were equally important for the shade-tolerant (F. grandifolia) species with determinate growth. Fagus grandifolia had a higher morphological plasticity in response to light, which may offer a competitive advantage in occupying a broader range of light conditions throughout the canopy. Differences in responses to light and height for the taller tree species, L. tulipifera and F. grandifolia, may be attributed to differences in growth strategy or Al:As, which may alter morphological and functional responses to light availability. While height was important in F. grandifolia, height was no more robust in predicting LMA than light in any of the species, confirming the strong role of light availability in determining LMA for temperate deciduous species.

Vertical leaf mass per area gradient of mature sugar maple reflects both height-driven increases in vascular tissue and light-driven increases in palisade layer thickness.

A key trait used in canopy and ecosystem function modeling, leaf mass per area (LMA), is influenced by changes in both leaf thickness and leaf density (LMA = Thickness × Density). In tall trees, LMA is understood to increase with height through two primary mechanisms: (i) increasing palisade layer thickness (and thus leaf thickness) in response to light and/or (ii) reduced cell expansion and intercellular air space in response to hydrostatic constraints, leading to increased leaf density. Our objective was to investigate within-canopy gradients in leaf anatomical traits in order to understand environmental factors that influence leaf morphology in a sugar maple (Acer saccharum Marshall) forest canopy. We teased apart the effects of light and height on anatomical traits by sampling at exposed and closed canopies that had different light conditions at similar heights. As expected, palisade layer thickness responded strongly to cumulative light exposure. Mesophyll porosity, however, was weakly and negatively correlated with light and height (i.e., hydrostatic gradients). Reduced mesophyll porosity was not likely caused by limitations on cell expansion; in fact, epidermal cell width increased with height. Palisade layer thickness was better related to LMA, leaf density and leaf thickness than was mesophyll porosity. Vein diameter and fraction of vascular tissue also increased with height and LMA, density and thickness, revealing that greater investment in vascular and support tissue may be a third mechanism for increased LMA with height. Overall, decreasing mesophyll porosity with height was likely due to palisade cells expanding into the available air space and also greater investments in vascular and support tissue, rather than a reduction of cell expansion due to hydrostatic constraints. Our results provide evidence that light influences both palisade layer thickness and mesophyll porosity and indicate that hydrostatic gradients influence leaf vascular and support tissues in mature Acer saccharum trees.

How vertical patterns in leaf traits shift seasonally and the implications for modeling canopy photosynthesis in a temperate deciduous forest.

Leaf functional traits are used in modeling forest canopy photosynthesis (Ac) due to strong correlations between photosynthetic capacity, leaf mass per area (LMA) and leaf nitrogen per area (Narea). Vertical distributions of these traits may change over time in temperate deciduous forests as a result of acclimation to light, which may result in seasonal changes in Ac To assess both spatial and temporal variations in key traits, we measured vertical profiles of Narea and LMA from leaf expansion through leaf senescence in a sugar maple (Acer saccharum Marshall) forest. To investigate mechanisms behind coordinated changes in leaf morphology and function, we also measured vertical variation in leaf carbon isotope composition (δ(13)C), predawn turgor pressure, leaf water potential and osmotic potential. Finally, we assessed potential biases in Ac estimations by parameterizing models with and without vertical and seasonal Narea variations following leaf expansion. Our data are consistent with the hypothesis that hydrostatic constraints on leaf morphology drive the vertical increase in LMA with height early in the growing season; however, LMA in the upper canopy continued to increase over time during light acclimation, indicating that light is primarily driving gradients in LMA later in the growing season. Models with no seasonal variation in Narea overestimated Ac by up to 11% early in the growing season, while models with no vertical variation in Narea overestimated Ac by up to 60% throughout the season. According to the multilayer model, the upper 25% of leaf area contributed to over 50% of Ac, but when gradients of intercellular CO2, as estimated from δ(13)C, were accounted for, the upper 25% of leaf area contributed to 26% of total Ac Our results suggest that ignoring vertical variation of key traits can lead to considerable overestimation of Ac.

Light acclimation optimizes leaf functional traits despite height-related constraints in a canopy shading experiment.

Within-canopy gradients of leaf functional traits have been linked to both light availability and vertical gradients in leaf water potential. While observational studies can reveal patterns in leaf traits, within-canopy experimental manipulations can provide mechanistic insight to tease apart multiple interacting drivers. Our objectives were to disentangle effects of height and light environment on leaf functional traits by experimentally shading branches along vertical gradients within a sugar maple (Acer saccharum) forest. Shading reduced leaf mass per area (LMA), leaf density, area-based leaf nitrogen (N(area)), and carbon:nitrogen (C:N) ratio, and increased mass-based leaf nitrogen (N(mass)), highlighting the importance of light availability on leaf morphology and chemistry. Early in the growing season, midday leaf water potential (Ψ(mid)), LMA, and N(area) were driven primarily by height; later in the growing season, light became the most important driver for LMA and Narea. Carbon isotope composition (δ(13)C) displayed strong, linear correlations with height throughout the growing season, but did not change with shading, implying that height is more influential than light on water use efficiency and stomatal behavior. LMA, leaf density, N(mass), C:N ratio, and δ(13)C all changed seasonally, suggesting that leaf ageing effects on leaf functional traits are equally as important as microclimatic conditions. Overall, our results indicate that: (1) stomatal sensitivity to vapor pressure deficit or Ψ(mid) constrains the supply of CO2 to leaves at higher heights, independent of light environment, and (2) LMA and N(area) distributions become functionally optimized through morphological acclimation to light with increasing leaf age despite height-related constraints.

Light drives vertical gradients of leaf morphology in a sugar maple (Acer saccharum) forest.

Leaf mass per area (LMA, g m(-2)) is an essential trait for modeling canopy function due to its strong association with photosynthesis, respiration and leaf nitrogen. Leaf mass per area, which is influenced by both leaf thickness and density (LMA = thickness × density), generally increases from the bottom to the top of tree canopies, yet the mechanisms behind this universal pattern are not yet resolved. For decades, the light environment was assumed to be the most influential driver of within-canopy variation in LMA, yet recent evidence has shown hydrostatic gradients to be more important in upper canopy positions, especially in tall evergreen trees in temperate and tropical forests. The aim of this study was to disentangle the importance of various environmental drivers on vertical LMA gradients in a mature sugar maple (Acer saccharum Marshall) forest. We compared LMA, leaf density and leaf thickness relationships with height, light and predawn leaf water potential (ΨPre) within a closed and an exposed canopy to assess leaf morphological traits at similar heights but different light conditions. Contrary to our expectations and recent findings in the literature, we found strong evidence that light was the primary driver of vertical gradients in leaf morphology. At similar heights (13-23 m), LMA was greater within the exposed canopy than the closed canopy, and light had a stronger influence over LMA compared with ΨPre. Light also had a stronger influence over both leaf thickness and density compared with ΨPre; however, the increase in LMA within both canopy types was primarily due to increasing leaf thickness with increasing light availability. This study provides strong evidence that canopy structure and crown exposure, in addition to height, should be considered as a parameter for determining vertical patterns in LMA and modeling canopy function.