The effects of solar radiation on daily and seasonal stem increment of canopy trees in European temperate old-growth forests.

It is well established that solar irradiance greatly influences tree metabolism and growth through photosynthesis, but its effects acting through individual climate metrics have not yet been well quantified. Understanding these effects is crucial for assessing the impacts of climate change on forest ecosystems. To describe the effects of solar irradiance on tree growth, we installed 110 automatic dendrometers in two old-growth mountain forest reserves in Central Europe, performed detailed terrestrial and aerial laser scanning to obtain precise tree profiles, and used these to simulate the sum of solar irradiance received by each tree on a daily basis. Generalized linear mixed-effect models were applied to simulate the probability of growth and the growth intensity over seven growing seasons. Our results demonstrated various contrasting effects of solar irradiance on the growth of canopy trees. On the one hand, the highest daily growth rates corresponded with the highest solar irradiance potentials (i.e. the longest photoperiod). Intense solar irradiance significantly decreased tree growth, through an increase in the vapor pressure deficit. These effects were consistent for all species but had different magnitude. Tree growth is the most effective on long rainy/cloudy days with low solar irradiance.

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 moisture1-11. 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 gain4-6,9-11. 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 °C in temperature increased light-saturated 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 light-saturated 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.

Seasonality of temperate forest photosynthesis and daytime respiration.

Terrestrial ecosystems currently offset one-quarter of anthropogenic carbon dioxide (CO2) emissions because of a slight imbalance between global terrestrial photosynthesis and respiration. Understanding what controls these two biological fluxes is therefore crucial to predicting climate change. Yet there is no way of directly measuring the photosynthesis or daytime respiration of a whole ecosystem of interacting organisms; instead, these fluxes are generally inferred from measurements of net ecosystem-atmosphere CO2 exchange (NEE), in a way that is based on assumed ecosystem-scale responses to the environment. The consequent view of temperate deciduous forests (an important CO2 sink) is that, first, ecosystem respiration is greater during the day than at night; and second, ecosystem photosynthetic light-use efficiency peaks after leaf expansion in spring and then declines, presumably because of leaf ageing or water stress. This view has underlain the development of terrestrial biosphere models used in climate prediction and of remote sensing indices of global biosphere productivity. Here, we use new isotopic instrumentation to determine ecosystem photosynthesis and daytime respiration in a temperate deciduous forest over a three-year period. We find that ecosystem respiration is lower during the day than at night-the first robust evidence of the inhibition of leaf respiration by light at the ecosystem scale. Because they do not capture this effect, standard approaches overestimate ecosystem photosynthesis and daytime respiration in the first half of the growing season at our site, and inaccurately portray ecosystem photosynthetic light-use efficiency. These findings revise our understanding of forest-atmosphere carbon exchange, and provide a basis for investigating how leaf-level physiological dynamics manifest at the canopy scale in other ecosystems.

Diffuse light and wetting differentially affect tropical tree leaf photosynthesis.

Most ecosystems experience frequent cloud cover resulting in light that is predominantly diffuse rather than direct. Moreover, these cloudy conditions are often accompanied by rain that results in wet leaf surfaces. Despite this, our understanding of photosynthesis is built upon measurements made on dry leaves experiencing direct light. Using a modified gas exchange setup, we measured the effects of diffuse light and leaf wetting on photosynthesis in canopy species from a tropical montane cloud forest. We demonstrate significant variation in species-level response to light quality independent of light intensity. Some species demonstrated 100% higher rates of photosynthesis in diffuse light, and others had 15% greater photosynthesis in direct light. Even at lower light intensities, diffuse light photosynthesis was equal to that under direct light conditions. Leaf wetting generally led to decreased photosynthesis, particularly when the leaf surface with stomata became wet; however, there was significant variation across species. Ultimately, we demonstrate that ecosystem photosynthesis is significantly altered in response to environmental conditions that are ubiquitous. Our results help to explain the observation that net ecosystem exchange can increase in cloudy conditions and can improve the representation of these processes in Earth systems models under projected scenarios of global climate change.

Patterns and dynamics of canopy-root coupling in tropical tree saplings vary with light intensity but not with root depth.

Carbon (C) dynamics in canopy and roots influence whole-tree carbon fluxes, but little is known about canopy regulation of tree-root activity. Here, the patterns and dynamics of canopy-root C coupling are assessed in tropical trees. Large aeroponics facility was used to study the root systems of Ceiba pentandra and Khaya anthotheca saplings directly at different light intensities. In Ceiba, root respiration (Rr ) co-varied with photosynthesis (An ) in large saplings (3-to-7-m canopy-root axis) at high-light, but showed no consistent pattern at low-light. At medium-light and in small saplings (c. 1-m axis), Rr tended to decrease transiently towards midday. Proximal roots had higher Rr and nonstructural carbohydrate concentrations than distal roots, but canopy-root coupling was unaffected by root location. In medium-sized Khaya, no Rr pattern was observed, and in both species, Rr was unrelated to temperature. The early-afternoon increase in Rr suggests that canopy-root coupling is based on mass flow of newly fixed C in the phloem, whereas the early-morning rise in Rr with An indicates an additional coupling signal that travels faster than the phloem sap. In large saplings and potentially also in higher trees, light and possibly additional environmental factors control the diurnal patterns of canopy-root coupling, irrespective of root location.

A comparative analysis of the photoprotection hypothesis for the evolution of autumn colours.

The adaptive value of autumn colours-the seasonal production of red anthocyanins observed in many species of trees and shrubs-is still debated. According to the photoprotection hypothesis, anthocyanins protect leaves from photo-inhibition and photo-oxidation at low temperatures, enabling the tree to reabsorb nutrients more efficiently before leaf fall. Hence, the hypothesis predicts that autumn colours are more likely to evolve in species growing in colder environments. We tested this prediction by comparing the climatic parameters of 237 North American tree species. We found that, although species with yellow autumn leaves grow under lower minimum temperatures than species with green leaves, there is no significant difference in temperature between species with red autumn leaves and species with green or yellow autumn leaves. We conclude that, although reabsorbing chlorophyll in autumn, and the consequent unmasking of yellow carotenoids, may be an adaptation to cold temperatures, the production of red anthocyanins is not. Hence, our interspecific comparative analysis does not support the photoprotection hypothesis as an explanation for the evolution of autumn colours.

Responses of respiration in the light to warming in field-grown trees: a comparison of the thermal sensitivity of the Kok and Laisk methods.

The Kok and Laisk techniques can both be used to estimate light respiration Rlight . We investigated whether responses of Rlight to short- and long-term changes in leaf temperature depend on the technique used to estimate Rlight . We grew Eucalyptus tereticornis in whole-tree chambers under ambient temperature (AT) or AT + 3°C (elevated temperature, ET). We assessed dark respiration Rdark and light respiration with the Kok (RKok ) and Laisk (RLaisk ) methods at four temperatures to determine the degree of light suppression of respiration using both methods in AT and ET trees. The ET treatment had little impact on Rdark , RKok or RLaisk . Although the thermal sensitivities of RKok or RLaisk were similar, RKok was higher than RLaisk . We found negative values of RLaisk at the lowest measurement temperatures, indicating positive net CO2 uptake, which we propose may be related to phosphoenolpyruvate carboxylase activity. Light suppression of Rdark decreased with increasing leaf temperature, but the degree of suppression depended on the method used. The Kok and Laisk methods do not generate the same estimates of Rlight or light suppression of Rdark between 20 and 35°C. Negative rates of RLaisk imply that this method may become less reliable at low temperatures.

Size-dependent variations in individual traits and trait scaling relationships within a shade-tolerant evergreen tree species.

PREMISE OF STUDY: The plant size-trait relationship is a fundamental dimension in the spectrum of plant form and function. However, it remains unclear whether the trait scaling relationship within species is modified by tree size. Investigating size-dependent trait covariations within species is crucial for understanding the ontogenetic constraints on the intraspecific economic spectrum and, more broadly, the structure and causes of intraspecific trait variations. METHODS: We measured eight morphological, stoichiometric, and hydraulic traits for 604 individual plants of a shade-tolerant evergreen tree species, Litsea elongata, in a subtropical evergreen forest of eastern China. Individual trait values were regressed against tree basal diameter to evaluate size-dependent trait variations. Standardized major axis regression was employed to examine trait scaling relationships and to test whether there was a common slope and elevation in the trait scaling relationship across size classes. KEY RESULTS: Small trees tended to have larger, thinner leaves and longer, slenderer stems than larger trees, which indicates an acquisitive economic strategy in juvenile trees. Leaf nitrogen concentrations increased with plant size, which was likely due to a high ratio of structural to photosynthetic nitrogen in the evergreen leaves of large trees. Bivariate trait scaling was minimally modified by tree size, although the elevation of some relationships differed between size classes. CONCLUSIONS: Our results suggest that there are common economic and biophysical constraints on intraspecific trait covariation, independent of tree size. Small and large trees tend to be located at opposite ends of an intraspecific plant economic spectrum.

In situ temperature response of photosynthesis of 42 tree and liana species in the canopy of two Panamanian lowland tropical forests with contrasting rainfall regimes.

Tropical forests contribute significantly to the global carbon cycle, but little is known about the temperature response of photosynthetic carbon uptake in tropical species, and how this varies within and across forests. We determined in situ photosynthetic temperature-response curves for upper canopy leaves of 42 tree and liana species from two tropical forests in Panama with contrasting rainfall regimes. On the basis of seedling studies, we hypothesized that species with high photosynthetic capacity - light-demanding, fast-growing species - would have a higher temperature optimum of photosynthesis (TOpt ) than species with low photosynthetic capacity - shade-tolerant, slow-growing species - and that, therefore, TOpt would scale with the position of a species on the slow-fast continuum of plant functional traits. TOpt was remarkably similar across species, regardless of their photosynthetic capacity and other plant functional traits. Community-average TOpt was almost identical to mean maximum daytime temperature, which was higher in the dry forest. Photosynthesis above TOpt appeared to be more strongly limited by stomatal conductance in the dry forest than in the wet forest. The observation that all species in a community shared similar TOpt values suggests that photosynthetic performance is optimized under current temperature regimes. These results should facilitate the scaling up of photosynthesis in relation to temperature from leaf to stand level in species-rich tropical forests.

Increased light-use efficiency sustains net primary productivity of shaded coffee plants in agroforestry system.

In agroforestry systems, shade trees strongly affect the physiology of the undergrown crop. However, a major paradigm is that the reduction in absorbed photosynthetically active radiation is, to a certain extent, compensated by an increase in light-use efficiency, thereby reducing the difference in net primary productivity between shaded and non-shaded plants. Due to the large spatial heterogeneity in agroforestry systems and the lack of appropriate tools, the combined effects of such variables have seldom been analysed, even though they may help understand physiological processes underlying yield dynamics. In this study, we monitored net primary productivity, during two years, on scales ranging from individual coffee plants to the entire plot. Absorbed radiation was mapped with a 3D model (MAESPA). Light-use efficiency and net assimilation rate were derived for each coffee plant individually. We found that although irradiance was reduced by 60% below crowns of shade trees, coffee light-use efficiency increased by 50%, leaving net primary productivity fairly stable across all shade levels. Variability of aboveground net primary productivity of coffee plants was caused primarily by the age of the plants and by intraspecific competition among them (drivers usually overlooked in the agroforestry literature) rather than by the presence of shade trees.

In situ temperature relationships of biochemical and stomatal controls of photosynthesis in four lowland tropical tree species.

Net photosynthetic carbon uptake of Panamanian lowland tropical forest species is typically optimal at 30-32 °C. The processes responsible for the decrease in photosynthesis at higher temperatures are not fully understood for tropical trees. We determined temperature responses of maximum rates of RuBP-carboxylation (VCMax ) and RuBP-regeneration (JMax ), stomatal conductance (Gs ), and respiration in the light (RLight ) in situ for 4 lowland tropical tree species in Panama. Gs had the lowest temperature optimum (TOpt ), similar to that of net photosynthesis, and photosynthesis became increasingly limited by stomatal conductance as temperature increased. JMax peaked at 34-37 °C and VCMax ~2 °C above that, except in the late-successional species Calophyllum longifolium, in which both peaked at ~33 °C. RLight significantly increased with increasing temperature, but simulations with a photosynthesis model indicated that this had only a small effect on net photosynthesis. We found no evidence for Rubisco-activase limitation of photosynthesis. TOpt of VCMax and JMax fell within the observed in situ leaf temperature range, but our study nonetheless suggests that net photosynthesis of tropical trees is more strongly influenced by the indirect effects of high temperature-for example, through elevated vapour pressure deficit and resulting decreases in stomatal conductance-than by direct temperature effects on photosynthetic biochemistry and respiration.

Competition for light in forest population dynamics: From computer simulator to mathematical model.

In this article we build a mathematical model for forest growth and we compare this model with a computer forest simulator named SORTIE. The main ingredient taken into account in both models is the competition for light between trees. The parameters of the mathematical model are estimated by using SORTIE model, when the parameter values of SORTIE model correspond to the ones previously evaluated for the Great Mountain Forest in USA. We see that the best fit of the parameters of the mathematical model is obtained when the competition for light influences only the growth rate of trees. We construct a size structured population dynamics model with one and two species and with spatial structure.

Effects of soil type and light on height growth, biomass partitioning, and nitrogen dynamics on 22 species of tropical dry forest tree seedlings: Comparisons between legumes and nonlegumes.

PREMISE OF THE STUDY: The seedling stage is particularly vulnerable to resource limitation, with potential consequences for community composition. We investigated how light and soil variation affected early growth, biomass partitioning, morphology, and physiology of 22 tree species common in tropical dry forest, including eight legumes. Our hypothesis was that legume seedlings are better at taking advantage of increased resource availability, which contributes to their successful regeneration in tropical dry forests. METHODS: We grew seedlings in a full-factorial design under two light levels in two soil types that differed in nutrient concentrations and soil moisture. We measured height biweekly and, at final harvest, biomass partitioning, internode segments, leaf carbon, nitrogen, δ13C, and δ15N. KEY RESULTS: Legumes initially grew taller and maintained that height advantage over time under all experimental conditions. Legumes also had the highest final total biomass and water-use efficiency in the high-light and high-resource soil. For nitrogen-fixing legumes, the amount of nitrogen derived from fixation was highest in the richer soil. Although seed mass tended to be larger in legumes, seed size alone did not account for all the differences between legumes and nonlegumes. Both belowground and aboveground resources were limiting to early seedling growth and function. CONCLUSIONS: Legumes may have a different regeneration niche, in that they germinate rapidly and grow taller than other species immediately after germination, maximizing their performance when light and belowground resources are readily available, and potentially permitting them to take advantage of high light, nutrient, and water availability at the beginning of the wet season.

[Spatial Distribution of ¹³7Cs in Soil of Spruce Forest in the Distant Zone of Chernobyl Fallout].

The vertical distribution of ¹³7Cs in Albeluvisols of the spruce forest in the Roslavl district of Smolensk region was investigated. The statistical characteristics of spatial variability and distribution laws for the activity con- centrations and activity depositions of ¹³7Cs in upper soil layers (forest litter, 0-5, 5-10, 10-19 cm) were de- termined. Positive correlations between the height, crown length of spruce trees and the content of ¹³7Cs in the soil under them were revealed. The regularities of spatial configuration of ¹³7Cs contamination in soil were related to the parcel structure of spruce forest. The lateral trends for distribution of 137Cs along the tessers un- der spruce and during the transfer to the open positions between the trees were recorded.

Functional ratios among leaf, xylem and phloem areas in branches change with shade tolerance, but not with local light conditions, across temperate tree species.

Leaf, xylem and phloem areas drive the water and carbon fluxes within branches and trees, but their mutual coordination is poorly understood. We test the hypothesis that xylem and phloem areas increase relative to leaf area when species are selected for, or branches are exposed to, higher levels of light intensity. Trees of 10 temperate, broadleaved and deciduous, tree species were selected. Fifty-centimetre-long branches were collected from shaded and exposed conditions at a height of 3-4 m. We measured the total leaf area, xylem area, phloem area and leaf traits, as well as the area of the constituent cell types, for a stem section at the branch base. Xylem area : leaf area and phloem area : leaf area ratios did not differ consistently between sun and shade branches, but, as expected, they decreased with species' shade tolerance. Similar trends were observed for conductive cell areas in xylem and phloem. Trees of light-demanding species maintain higher water loss and carbon gain rates per leaf area by producing more xylem area and phloem area than shade-tolerant species. We call for more comparative branch studies as they provide an integrated biological perspective on functional traits and their role in the ecology of tree species.

Improvement of water and light availability after thinning at a xeric site: which matters more? A dual isotope approach.

Thinning fosters individual tree growth by increasing the availability of water, light and nutrients. At sites where water rather than light is limiting, thinning also enhances soil evaporation and might not be beneficial. Detailed knowledge of the short- to long-term physiological response underlying the growth responses to thinning is crucial for the management of forests already suffering from recurrent drought-induced dieback. We applied a dual isotope approach together with mechanistic isotope models to study the physiological processes underlying long-term growth enhancement of heavily thinned Pinus sylvestris in a xeric forest in Switzerland. This approach allowed us to identify and disentangle thinning-induced changes in stomatal conductance and assimilation rate. At our xeric study site, the increase in stomatal conductance far outweighed the increase in assimilation, implying that growth release in heavily thinned trees is primarily driven by enhanced water availability rather than increased light availability. We conclude that in forests with relatively isohydric species (drought avoiders) that are growing close to their physiological limits, thinning is recommended to maintain a less negative water balance and thus foster tree growth, and ultimately the survival of forest trees under drought.

Convergent acclimation of leaf photosynthesis and respiration to prevailing ambient temperatures under current and warmer climates in Eucalyptus tereticornis.

Understanding physiological acclimation of photosynthesis and respiration is important in elucidating the metabolic performance of trees in a changing climate. Does physiological acclimation to climate warming mirror acclimation to seasonal temperature changes? We grew Eucalyptus tereticornis trees in the field for 14 months inside 9-m tall whole-tree chambers tracking ambient air temperature (Tair ) or ambient Tair  + 3°C (i.e. 'warmed'). We measured light- and CO2 -saturated net photosynthesis (Amax ) and night-time dark respiration (R) each month at 25°C to quantify acclimation. Tree growth was measured, and leaf nitrogen (N) and total nonstructural carbohydrate (TNC) concentrations were determined to investigate mechanisms of acclimation. Warming reduced Amax and R measured at 25°C compared to ambient-grown trees. Both traits also declined as mean daily Tair increased, and did so in a similar way across temperature treatments. Amax and R (at 25°C) both increased as TNC concentrations increased seasonally; these relationships appeared to arise from source-sink imbalances, suggesting potential substrate regulation of thermal acclimation. We found that photosynthesis and respiration each acclimated equivalently to experimental warming and seasonal temperature change of a similar magnitude, reflecting a common, nearly homeostatic constraint on leaf carbon exchange that will be important in governing tree responses to climate warming.

Transcriptomic analysis suggests a key role for SQUAMOSA PROMOTER BINDING PROTEIN LIKE, NAC and YUCCA genes in the heteroblastic development of the temperate rainforest tree Gevuina avellana (Proteaceae).

Heteroblasty, the temporal development of the meristem, can produce diverse leaf shapes within a plant. Gevuina avellana, a tree from the South American temperate rainforest shows strong heteroblasty affecting leaf shape, transitioning from juvenile simple leaves to highly pinnate adult leaves. Light availability within the forest canopy also modulates its leaf size and complexity. Here we studied how the interaction between the light environment and the heteroblastic progression of leaves is coordinated in this species. We used RNA-seq on the Illumina platform to compare the range of transcriptional responses in leaf primordia of G. avellana at different heteroblastic stages and growing under different light environments. We found a steady up-regulation of SQUAMOSA PROMOTER BINDING PROTEIN LIKE (SPL), NAC, YUCCA and AGAMOUS-LIKE genes associated with increases in age, leaf complexity, and light availability. In contrast, expression of TCP, TPR and KNOTTED1 homeobox genes showed a sustained down-regulation. Additionally, genes involved in auxin synthesis/transport and jasmonate activity were differentially expressed, indicating an active regulation of processes controlled by these hormones. Our large-scale transcriptional analysis of the leaf primordia of G. avellana sheds light on the integration of internal and external cues during heteroblastic development in this species.

The expression of light-related leaf functional traits depends on the location of individual leaves within the crown of isolated Olea europaea trees.

BACKGROUND: The spatial arrangement and expression of foliar syndromes within tree crowns can reflect the coupling between crown form and function in a given environment. Isolated trees subjected to high irradiance and concomitant stress may adjust leaf phenotypes to cope with environmental gradients that are heterogeneous in space and time within the tree crown. The distinct expression of leaf phenotypes among crown positions could lead to complementary patterns in light interception at the crown scale. METHODS: We quantified eight light-related leaf traits across 12 crown positions of ten isolated Olea europaea trees in the field. Specifically, we investigated whether the phenotypic expression of foliar traits differed among crown sectors and layers and five periods of the day from sunrise to sunset. We investigated the consequences in terms of the exposed area of the leaves at the tree scale during a single day. KEY RESULTS: All traits differed among crown positions except the length-to-width ratio of the leaves. We found a strong complementarity in the patterns of the potential exposed area of the leaves among day periods as a result of a non-random distribution of leaf angles across the crown. Leaf exposure at the outer layer was below 60 % of the displayed surface, reaching maximum interception during morning periods. Daily interception increased towards the inner layer, achieving consecutive maximization from east to west positions within the crown, matching the sun's trajectory. CONCLUSIONS: The expression of leaf traits within isolated trees of O. europaea varies continuously through the crown in a gradient of leaf morphotypes and leaf angles depending on the exposure and location of individual leaves. The distribution of light-related traits within the crown and the complementarity in the potential exposure patterns of the leaves during the day challenges the assumption of low trait variability within individuals.

Hyperspectral remote sensing of foliar nitrogen content.

A strong positive correlation between vegetation canopy bidirectional reflectance factor (BRF) in the near infrared (NIR) spectral region and foliar mass-based nitrogen concentration (%N) has been reported in some temperate and boreal forests. This relationship, if true, would indicate an additional role for nitrogen in the climate system via its influence on surface albedo and may offer a simple approach for monitoring foliar nitrogen using satellite data. We report, however, that the previously reported correlation is an artifact--it is a consequence of variations in canopy structure, rather than of %N. The data underlying this relationship were collected at sites with varying proportions of foliar nitrogen-poor needleleaf and nitrogen-rich broadleaf species, whose canopy structure differs considerably. When the BRF data are corrected for canopy-structure effects, the residual reflectance variations are negatively related to %N at all wavelengths in the interval 423-855 nm. This suggests that the observed positive correlation between BRF and %N conveys no information about %N. We find that to infer leaf biochemical constituents, e.g., N content, from remotely sensed data, BRF spectra in the interval 710-790 nm provide critical information for correction of structural influences. Our analysis also suggests that surface characteristics of leaves impact remote sensing of its internal constituents. This further decreases the ability to remotely sense canopy foliar nitrogen. Finally, the analysis presented here is generic to the problem of remote sensing of leaf-tissue constituents and is therefore not a specific critique of articles espousing remote sensing of foliar %N.