Becoming less tolerant with age: sugar maple, shade, and ontogeny

Although shade tolerance is often assumed to be a fixed trait, recent work suggests ontogenetic changes in the light requirements of tree species. We determined the influence of gas exchange, biomass distribution, and self-shading on ontogenetic variation in the instantaneous aboveground carbon balance of Acer saccharum. We quantified the aboveground biomass distributions of 18 juveniles varying in height and growing in low light in a temperate forest understory in Minnesota, USA. Gas exchange rates of leaf and stem tissues were measured, and the crown architecture of each individual was quantified. The YPLANT program was used to estimate the self-shaded fraction of each crown and to model net leaf-level carbon gain. Leaf respiration and photosynthesis per gram of leaf tissue increased with plant size. In contrast, stem respiration rates per gram of stem tissue declined, reflecting a shift in the distribution of stem diameter sizes from smaller (with higher respiration) to larger diameter classes. However, these trends were outweighed by ontogenetic increases in self-shading (which reduces the net photosynthesis realized) and stem mass fraction (which increases the proportion of purely respiratory tissue) in terms of influence on net carbon exchange. As a result, net carbon gain per gram of aboveground plant tissue declined with increasing plant size, and the instantaneous aboveground light compensation point increased. When estimates of root respiration were included to model whole-plant carbon gain and light compensation points, relationships with plant size were even more pronounced. Our findings show how an interplay of gas exchange, self-shading, and biomass distribution shapes ontogenetic changes in shade tolerance

Variation in leaf and twig CO2 flux as a function of plant size : a comparison of seedlings, saplings and trees

Rates of tissue-level function have been hypothesized to decline as trees grow older and larger, but relevant evidence to assess such changes remains limited, especially across a wide range of sizes from saplings to large trees. We measured functional traits of leaves and twigs of three cold-temperate deciduous tree species in Minnesota, USA, to assess how these vary with tree height. Individuals ranging from 0.13 to 20 m in height were sampled in both relatively open and closed canopy environments to minimize light differences as a potential driver of size-related differences in leaf and twig properties. We hypothesized that (H1) gas-exchange rates, tissue N concentration and leaf mass per unit area (LMA) would vary with tree size in a pattern reflecting declining function in taller trees, yet maintaining (H2) bivariate trait relations, common among species as characterized by the leaf economics spectrum. Taking these two ideas together yielded a third, integrated hypothesis that (H3) nitrogen (N) content and gas-exchange rates should decrease monotonically with tree size and LMA should increase. We observed increasing LMA and decreasing leaf and twig Rd with increasing size, which matched predictions from H1 and H3. However, opposite to our predictions, leaf and twig N generally increased with size, and thus had inverse relations with respiration, rather than the predicted positive relations. Two exceptions were area-based leaf N of Prunus serotina Ehrh. in gaps and mass-based leaf N of Quercus ellipsoidalis E. J. Hill in gaps, both of which showed qualitatively hump-shaped patterns. Finally, we observed hump-shaped relationships between photosynthetic capacity and tree height, not mirroring any of the other traits, except in the two cases highlighted above. Bivariate trait relations were weak intra-specifically, but were generally significant and positive for area-based traits using the pooled dataset. Results suggest that different traits vary with tree size in different ways that are not consistent with a universal shift towards a lower 'return on investment' strategy. Instead, species traits vary with size in patterns that likely reflect complex variation in water, light, nitrogen and carbon availability, storage and use

Georgia Southern Walking Tour of Trees: A self-regulated learning opportunity to better engage students in the importance of tree species in urban environments (Phase 2)

Dr. Kerrie Sendall, Assistant Professor, Dr. Alan Harvey, Professor and Dr. John Schenk, Assistant Professor, Department of Biology ($27,400.00) Continued working on identifying and cataloging trees along additional major walking paths on the GS (Statesboro) campus. Information was used to develop a virtual campus tree walk

Size-related shifts in carbon gain and growth responses to light differ among rainforest evergreens of contrasting shade tolerance

Recent work suggests that plant size affects light requirements and carbon balance of juvenile trees, and such shifts may be greater in light-demanding species than in their more shade-tolerant associates. To explore the physiological basis of such shifts, we measured juvenile light interception, carbon gain and growth of four subtropical Australian rainforest trees differing in shade tolerance, comparing individuals ranging from 13 to 238 cm in height, across a wide range of understory environments. We hypothesized that even in a standardized light environment, increasing sapling size would lead to declines in net daily carbon gain of foliage and relative growth rates (RGR) of all species, with declines more pronounced in light demanding species. Crown architecture of individuals was recorded using a 3-dimensional digitizer, and the YPLANT program was used to estimate the self-shaded fraction of each crown and model net carbon gain. Increased sapling size caused a significant increase in self-shading, and significant declines in net daily carbon gain and RGR of light-demanding species, while such ontogenetic variations were minimal or absent in shade-tolerant species. Additionally, differences in the slope of the relationship between light and RGR led to crossovers in RGR among shade-tolerant and light-demanding species at low light. Our results show that the magnitude of ontogenetic variation in net daily carbon gain and RGR can be substantial and may depend on successional status, making it unsafe to assume that young seedling performance can be used to predict or model responses of larger juvenile trees

Growth, light and trait data

Relative growth rate, light availability and leaf trait data for saplings growing in subtropical rainforest sites in Australia

Surprising lack of sensitivity of biochemical limitation of photosynthesis of nine tree species to open‐air experimental warming and reduced rainfall in a southern boreal forest

Photosynthetic biochemical limitation parameters (i.e., Vcmax, Jmax and Jmax:Vcmax ratio) are sensitive to temperature and water availability, but whether these parameters in cold climate species at biome ecotones are positively or negatively influenced by projected changes in global temperature and water availability remains uncertain. Prior exploration of this question has largely involved greenhouse based short-term manipulative studies with mixed results in terms of direction and magnitude of responses. To address this question in a more realistic context, we examined the effects of increased temperature and rainfall reduction on the biochemical limitations of photosynthesis using a long-term chamber-less manipulative experiment located in northern Minnesota, USA. Nine tree species from the boreal-temperate ecotone were grown in natural neighborhoods under ambient and elevated (+3.4°C) growing season temperatures and ambient or reduced (≈40% of rainfall removed) summer rainfall. Apparent rubisco carboxylation and RuBP regeneration standardized to 25°C (Vcmax25°C and Jmax25°C, respectively) were estimated based on ACi curves measured in situ over three growing seasons. Our primary objective was to test whether species would downregulate Vcmax25°C and Jmax25°C in response to warming and reduced rainfall, with such responses expected to be greatest in species with the coldest and most humid native ranges, respectively. These hypotheses were not supported, as there were no overall main treatment effects on Vcmax25°C or Jmax25°C (p > .14). However, Jmax:Vcmax ratio decreased significantly with warming (p = .0178), whereas interactions between warming and rainfall reduction on the Jmax25°C to Vcmax25°C ratio were not significant. The insensitivity of photosynthetic parameters to warming contrasts with many prior studies done under larger temperature differentials and often fixed daytime temperatures. In sum, plants growing in relatively realistic conditions under naturally varying temperatures and soil moisture levels were remarkably insensitive in terms of their Jmax25°C and Vcmax25°C when grown at elevated temperatures, reduced rainfall, or both combined

Surprising lack of sensitivity of biochemical limitation of photosynthesis of nine tree species to open-air experimental warming and reduced rainfall in a southern boreal forest

Photosynthetic biochemical limitation parameters (i.e., Vcmax, Jmax and Jmax:Vcmax ratio) are sensitive to temperature and water availability, but whether these parameters in cold climate species at biome ecotones are positively or negatively influenced by projected changes in global temperature and water availability remains uncertain. Prior exploration of this question has largely involved greenhouse based short-term manipulative studies with mixed results in terms of direction and magnitude of responses. To address this question in a more realistic context, we examined the effects of increased temperature and rainfall reduction on the biochemical limitations of photosynthesis using a long-term chamber-less manipulative experiment located in northern Minnesota, USA. Nine tree species from the boreal-temperate ecotone were grown in natural neighborhoods under ambient and elevated (+3.4°C) growing season temperatures and ambient or reduced (≈40% of rainfall removed) summer rainfall. Apparent rubisco carboxylation and RuBP regeneration standardized to 25°C (Vcmax25°C and Jmax25°C, respectively) were estimated based on ACi curves measured in situ over three growing seasons. Our primary objective was to test whether species would downregulate Vcmax25°C and Jmax25°C in response to warming and reduced rainfall, with such responses expected to be greatest in species with the coldest and most humid native ranges, respectively. These hypotheses were not supported, as there were no overall main treatment effects on Vcmax25°C or Jmax25°C (p > .14). However, Jmax:Vcmax ratio decreased significantly with warming (p = .0178), whereas interactions between warming and rainfall reduction on the Jmax25°C to Vcmax25°C ratio were not significant. The insensitivity of photosynthetic parameters to warming contrasts with many prior studies done under larger temperature differentials and often fixed daytime temperatures. In sum, plants growing in relatively realistic conditions under naturally varying temperatures and soil moisture levels were remarkably insensitive in terms of their Jmax25°C and Vcmax25°C when grown at elevated temperatures, reduced rainfall, or both combined

Effects of climate warming on photosynthesis in boreal tree species depend on soil moisture

Climate warming will influence photosynthesis via thermal effects and by altering soil moisture(1-11.) Both effects may be important for the vast areas of global forests that fluctuate between periods when cool temperatures limit photosynthesis and periods when soil moisture may be limiting to carbon gain(4-6,9-11). Here we show that the effects of climate warming flip from positive to negative as southern boreal forests transition from rainy to modestly dry periods during the growing season. In a three-year open-air warming experiment with juveniles of 11 temperate and boreal tree species, an increase of 3.4 degrees C in temperature increased lightsaturated net photosynthesis and leaf diffusive conductance on average on the one-third of days with the wettest soils. In all 11 species, leaf diffusive conductance and, as a result, light-saturated net photosynthesis decreased during dry spells, and did so more sharply in warmed plants than in plants at ambient temperatures. Consequently, across the 11 species, warming reduced lightsaturated net photosynthesis on the two-thirds of days with driest soils. Thus, low soil moisture may reduce, or even reverse, the potential benefits of climate warming on photosynthesis in mesic, seasonally cold environments, both during drought and in regularly occurring, modestly dry periods during the growing season