Herbivory and leaf traits of Amazonian tree species as affected by irradiance.

Herbivory is one of the major biotic stress factors that affect the establishment of plants. However, the main factors that drive herbivory in seedlings of Amazonian tree species are still not well understood. Here we investigated whether contrasting levels of irradiance influence herbivory according to different herbivory indicators and which leaf traits are most related to interspecific variation in herbivory under contrasting irradiance conditions. We measured the leaf area lost as a result of insect herbivory in five tree species planted in a silvicultural system of secondary forest enrichment according to two indicators, herbivore damage (accumulated since plant germination) and herbivory rate (measured over time), and two irradiance conditions, understorey PPFD 2.6 mol·m-2 ·day-1 ) and gap PPFD 33.1 mol·m-2 ·day-1 . Furthermore, we related the interspecific variation in herbivory to a set of leaf traits: SLA, RWC, sclerophylly, phenolic compound content, tannins, condensed tannins and non-structural carbohydrates. Herbivore damage was significantly affected by light availability and species, with the highest percentage variation observed in the Meliaceae (Carapa guianensis and Swietenia macrophylla). For the herbivory rate, only the interspecific variation was significant, with Bertholletia excelsa having the lowest rates. Chemical characteristics (phenolic compounds and tannins) were most related to herbivory rates, as well as highly influenced by light conditions. Non-structural carbohydrates (starch and sucrose) were also related to the interspecific variation in herbivory. The phenolic compounds and starch, as affected by light quantity, are species dependent. Thus, the selective pressure on herbivores may be driven by species-dependent responses to light conditions.

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.

Determination of the most effective design for the measurement of photosynthetic light-response curves for planted Larix olgensis trees.

A photosynthetic light-response (PLR) curve is a mathematical description of a single biochemical process and has been widely applied in many eco-physiological models. To date, many PLR measurement designs have been suggested, although their differences have rarely been explored, and the most effective design has not been determined. In this study, we measured three types of PLR curves (High, Middle and Low) from planted Larix olgensis trees by setting 31 photosynthetically active radiation (PAR) gradients. More than 530 million designs with different combinations of PAR gradients from 5 to 30 measured points were conducted to fit each of the three types of PLR curves. The influence of different PLR measurement designs on the goodness of fit of the PLR curves and the accuracy of the estimated photosynthetic indicators were analysed, and the optimal design was determined. The results showed that the measurement designs with fewer PAR gradients generally resulted in worse predicted accuracy for the photosynthetic indicators. However, the accuracy increased and remained stable when more than ten measurement points were used for the PAR gradients. The mean percent error (M%E) of the estimated maximum net photosynthetic rate (Pmax) and dark respiratory rate (Rd) for the designs with less than ten measurement points were, on average, 16.4 times and 20.1 times greater than those for the designs with more than ten measurement points. For a single tree, a unique PLR curve design generally reduced the accuracy of the predicted photosynthetic indicators. Thus, three optimal measurement designs were provided for the three PLR curve types, in which the root mean square error (RMSE) values reduced by an average of 8.3% and the coefficient of determination (R2) values increased by 0.3%. The optimal design for the High PLR curve type should shift more towards high-intensity PAR values, which is in contrast to the optimal design for the Low PLR curve type, which should shift more towards low-intensity PAR values.

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.

Effects of nitric oxide-releasing nanoparticles on neotropical tree seedlings submitted to acclimation under full sun in the nursery.

Polymeric nanoparticles have emerged as carrier systems for molecules that release nitric oxide (NO), a free radical involved in plant stress responses. However, to date, nanoencapsulated NO donors have not been applied to plants under realistic field conditions. Here, we verified the effects of free and nanoencapsulated NO donor, S-nitroso-mercaptosuccinic acid (S-nitroso-MSA), on growth, physiological and biochemical parameters of neotropical tree seedlings kept under full sunlight in the nursery for acclimation. S-nitroso-MSA incorporation into chitosan nanoparticles partially protected the NO donor from thermal and photochemical degradation. The application of nanoencapsulated S-nitroso-MSA in the substrate favoured the growth of seedlings of Heliocarpus popayanensis, a shade-intolerant tree. In contrast, free S-nitroso-MSA or nanoparticles containing non-nitrosated mercaptosuccinic acid reduced photosynthesis and seedling growth. Seedlings of Cariniana estrellensis, a shade-tolerant tree, did not have their photosynthesis and growth affected by any formulations, despite the increase of foliar S-nitrosothiol levels mainly induced by S-nitroso-MSA-loaded nanoparticles. These results suggest that depending on the tree species, nanoencapsulated NO donors can be used to improve seedling acclimation in the nursery.

The influence of spectral composition on spring and autumn phenology in trees.

Several recent reviews highlight the molecular mechanisms that underpin phenological responses to temperature and photoperiod; however, these have mostly overlooked the influence of solar radiation and its spectral composition on these processes. For instance, solar radiation in the blue and ultraviolet (UV) regions of the spectrum, as well as the red/far-red (R:FR) ratio, can influence spring and autumn phenology. Solar radiation reaching the Earth changes diurnally and seasonally; however, rising global temperatures, latitudinal range shifts and light pollution are likely to produce novel combinations of phenological cues for tree species. Here, we review the literature on phenological responses to spectral composition. Our objective was to explore the natural variation in spectral composition using radiative transfer models and to reveal any species-specific or ecotype-specific responses relating to latitudinal origin. These responses are likely to be most pronounced at high latitudes where spectral composition varies most throughout the year. For instance, trees from high latitudes tend to be more sensitive to changes in R:FR than those from low latitudes. The effects of blue light and UV radiation on phenology have not been studied as much as those of R:FR, but the limited results available suggest both could be candidate cues affecting autumn leaf colouration and senescence. Failure of more-southern species and ecotypes to adapt and use spectral cues during northwards range shifts could result in mistimed phenology, potentially resulting in frost damage, reduced fitness and limited range expansion. Future areas for research should look to establish how consistently different functional types of tree respond to spectral cues and identify photoreceptor-mediated mechanisms that allow plants to combine information from multiple light cues to coordinate the timing of phenological events. It should then be feasible to consider the synchronous or sequential action of light cues within a hierarchy of environmental factors regulating phenology.

Anthocyanin synthesis and light utilisation can be enhanced by reflective mulch - Visualisation of light penetration into a tree canopy.

With the increasing use of hailnets and decrease in light availability in the ripening period of apple fruit, insufficient light exposure often causes poor colouration viz anthocyanin synthesis on certain parts of the fruit and on certain fruit within the tree. The aim of this study was to investigate the potential of improving anthocyanin synthesis, in terms of fruit colouration, the major incentive for a consumer. Therefore, the reflective ground cover Lumilys® was spread between 'Braeburn Mariri Red' apple rows under a crystalline hailnet seven weeks prior to harvest and colour measured on 240 attached apple fruit. The reflective ground cover increased the reflected light by 1.6 to 3.9 times 1.0 m above ground. The improved light utilization led to an improved peel colouration, especially on the shaded side of the apple fruit and fruit in the lower inner part of the canopy, A coloured visualization from orange (high light intensity), yellow (medium) to green (low light intensity) as a result of the individual PAR measurements every 20 cm inside the canopy showed how the reflective mulch influences the light penetration into the different parts of the tree canopy.

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.

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.

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.

Nitrogen deposition does not affect the impact of shade on Quercus acutissima seedlings.

Light and atmospheric nitrogen (N) deposition are among the important environmental factors influencing plant growth and forest regeneration. We used Quercus acutissima, a dominant broadleaf tree species native to the deciduous forests of Northern China, to study the combined effects of light exposure and N addition on leaf physiology and individual plant growth. In the greenhouse, we exposed Quercus acutissima seedlings to one of two light conditions (8% and 80% of full irradiation) and one of three N treatments (0, 6, and 12 g N m-2 y-1). After 87 d, we observed that nitrogen deposition had no significant effects on the seedlings regardless of light exposure. In addition, shade significantly reduced plant height, basal diameter, leaf number, total biomass, gas exchange capacity, and carbohydrate content. In contrast, however, shade significantly increased the amount of photosynthetic pigment, above-ground biomass allocation, and specific leaf area. There was also a hierarchical plasticity among the different seedling characteristics. Compared to traits of growth, biomass, biomass allocation and leaf morphology, the leaf physiology, including photosynthetic pigment, gas exchange, carbohydrate, and PUNE, is more sensitive to light conditions. Among the biomass allocation parameters, the leaf and root mass ratios had a relatively low phenotypic plasticity. The seedlings had high foliar physiological plasticity under various light conditions. Nevertheless, we recommend high irradiance to maintain vigorous seedling growth and, in turn, promote the restoration and reconstruction of vegetation.

Wind loads and competition for light sculpt trees into self-similar structures.

Trees are self-similar structures: their branch lengths and diameters vary allometrically within the tree architecture, with longer and thicker branches near the ground. These tree allometries are often attributed to optimisation of hydraulic sap transport and safety against elastic buckling. Here, we show that these allometries also emerge from a model that includes competition for light, wind biomechanics and no hydraulics. We have developed MECHATREE, a numerical model of trees growing and evolving on a virtual island. With this model, we identify the fittest growth strategy when trees compete for light and allocate their photosynthates to grow seeds, create new branches or reinforce existing ones in response to wind-induced loads. Strikingly, we find that selected trees species are self-similar and follow allometric scalings similar to those observed on dicots and conifers. This result suggests that resistance to wind and competition for light play an essential role in determining tree allometries.

Short-term effects of light quality on leaf gas exchange and hydraulic properties of silver birch (Betula pendula).

Leaves have to acclimatize to heterogeneous radiation fields inside forest canopies in order to efficiently exploit diverse light conditions. Short-term effects of light quality on photosynthetic gas exchange, leaf water use and hydraulic traits were studied on Betula pendula Roth shoots cut from upper and lower thirds of the canopy of 39- to 35-year-old trees growing in natural forest stand, and illuminated with white, red or blue light in the laboratory. Photosynthetic machinery of the leaves developed in different spectral conditions acclimated differently with respect to incident light spectrum: the stimulating effect of complete visible spectrum (white light) on net photosynthesis is more pronounced in upper-canopy layers. Upper-canopy leaves exhibit less water saving behaviour, which may be beneficial for the fast-growing pioneer species on a daily basis. Lower-canopy leaves have lower stomatal conductance resulting in more efficient water use. Spectral gradients existing within natural forest stands represent signals for the fine-tuning of stomatal conductance and tree water relations to afford lavish water use in sun foliage and enhance leaf water-use efficiency in shade foliage sustaining greater hydraulic limitations. Higher sensitivity of hydraulic conductance of shade leaves to blue light probably contributes to the efficient use of short duration sunflecks by lower-canopy leaves.

Leaf thermotolerance in tropical trees from a seasonally dry climate varies along the slow-fast resource acquisition spectrum.

Knowledge of the upper limits of temperature tolerance is essential to understand how tropical trees will respond to global warming. We quantified leaf thermotolerance in 41 tree species growing in a seasonally dry tropical region of the Indian subcontinent to examine: (1) differences between evergreen and deciduous species; (2) relationships with leaf mass per area (LMA) and leaf size; and, (3) seasonal variation in thermotolerance. Thermotolerance ranged from 45.5 °C to 50.5 °C among species, was higher for evergreen than deciduous species, and was negatively related to a continuous estimate of deciduousness. Species with higher LMA had higher thermotolerance, but we did not detect any relationship between leaf size and thermotolerance. Seasonal changes in thermotolerance varied among species implying that species' capacity to acclimate may differ. Thermal safety margins, the difference between thermotolerance and maximum habitat temperatures indicate that most species may be highly vulnerable to future warming. Overall our results show that deciduous, and fast growing species with low LMA are likely to be more negatively affected by global warming. This differential vulnerability may lead to directional changes in composition in dry tropical forests, and such changes could alter vegetation-atmosphere feedbacks and further exacerbate global warming.

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.

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.

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.

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.

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.