The temperature dependence of C 3 photosynthesis is known to vary with growth environment and with species. In an attempt to quantify this variability, a commonly used biochemically based photosynthesis model was parameterized from 19 gas exchange studies on tree and crop species. The parameter values obtained described the shape and amplitude of the temperature responses of the maximum rate of Rubisco activity ( V cmax ) and the potential rate of electron transport ( J max ). Original data sets were used for this review, as it is shown that derived values of V cmax and its temperature response depend strongly on assumptions made in derivation. Values of J max and V cmax at 25 °°°° C varied considerably among species but were strongly correlated, with an average J max : V cmax ratio of 1·67. Two species grown in cold climates, however, had lower ratios. In all studies, the J max : V cmax ratio declined strongly with measurement temperature. The relative temperature responses of J max and V cmax were relatively constant among tree species. Activation energies averaged 50 kJ mol − − − − 1 for J max and 65 kJ mol − − − − 1 for V cmax , and for most species temperature optima averaged 33 °°°° C for J max and 40 °°°° C for V cmax . However, the cold climate tree species had low temperature optima for both J max ( 19 °°°° C) and V cmax (29 °°°° C), suggesting acclimation of both processes to growth temperature. Crop species had somewhat different temperature responses, with higher activation energies for both J max and V cmax , implying narrower peaks in the temperature response for these species. The results thus suggest that both growth environment and plant type can influence the photosynthetic response to temperature. Based on these results, several suggestions are made to improve modelling of temperature responses.

Seedlings of seven temperate tree species (Acer pseudoplatanus L., Betula pendula Roth, Fagus sylvatica L., Fraxinus excelsior L., Juglans regia L., Quercus petraea Matt. Liebl. and Quercus robur L.) were grown in a nursery under neutral filters transmitting 45% of incident global irradiance. During the second or third year of growth, leaf photosynthetic capacity (i.e., maximal carboxylation rate, Vcmax, maximal photosynthetic electron transport rate, Jmax, and dark respiration, Rd) was estimated for five leaves from each species at five or six leaf temperatures (10, 18, 25, 32, 36 and 40 degrees C). Values of Vcmax and Jmax were obtained by fitting the equations of the Farquhar model on response curves of net CO2 assimilation (A) to sub-stomatal CO2 mole fraction (ci), at high irradiance. Primary parameters describing the kinetic properties of Rubisco (specificity factor, affinity for CO2 and for O2, and their temperature responses) were taken from published data obtained with spinach and tobacco, and were used for all species. The temperature responses of Vcmax and Jmax, which were fitted to a thermodynamic model, differed. Mean values of Vcmax and Jmax at a reference temperature of 25 degrees C were 77.3 and 139 micromol m(-2) s(-1), respectively. The activation energy was higher for Vcmax than for Jmax (mean values of 73.1 versus 57.9 kJ mol(-1)) resulting in a decrease in Jmax/Vcmax ratio with increasing temperature. The mean optimal temperature was higher for Vcmax than for Jmax (38.9 versus 35.9 degrees C). In addition, differences in these temperature responses were observed among species. Temperature optima ranged between 35.9 and above 45 degrees C for Vcmax and between 31.7 and 43.3 degrees C for Jmax, but because of data scatter and the limited range of temperatures tested (10 to 40 degrees C), there were few statistically significant differences among species. The optimal temperature for Jmax was highest in Q. robur, Q. petraea and J. regia, and lowest in A. pseudoplatanus and F. excelsior. Measurements of chlorophyll a fluorescence revealed that the critical temperature at which basal fluorescence begins to increase was close to 47 degrees C, with no difference among species. These results should improve the parameterization of photosynthesis models, and be of particular interest when adapted to heterogeneous forests comprising mixtures of species with diverse ecological requirements.

The interspecific variability of sunlit leaf carbon isotope composition (δC), an indicator of leaf intrinsic water-use efficiency (WUE, CO assimilation rate/leaf conductance for water vapour), was investigated in canopy trees of three lowland rainforest stands in French Guiana, differing in floristic composition and in soil drainage characteristics, but subjected to similar climatic conditions. We sampled leaves with a rifle from 406 trees in total, representing 102 species. Eighteen species were common to the three stands. Mean species δC varied over a 6.0‰ range within each stand, corresponding to WUE varying over about a threefold range. Species occurring in at least two stands displayed remarkably stable δC values, suggesting a close genetic control of species δC. Marked differences in species δC values were found with respect to: (1) the leaf phenology pattern (average δC=-29.7‰ and -31.0‰ in deciduous-leaved and evergreen-leaved species, respectively), and (2) different types of shade tolerance defined by features reflecting the plasticity of growth dynamics with respect to contrasting light conditions. Heliophilic species exhibited more negative δC values (average δC=-30.5‰) (i.e. lower WUE) than hemitolerant species (-29.3‰). However, tolerant species (-31.4‰) displayed even more negative δC values than heliophilic ones. We could not provide a straightforward ecophysiological interpretation of this result. The negative relationship found between species δC and midday leaf water potential (Ψ) suggests that low δC is associated with high whole tree leaf specific hydraulic conductance. Canopy carbon isotope discrimination (Δ ) calculated from the basal area-weighed integral of the species δC values was similar in the three stands (average Δ =23.1‰), despite differences in stand species composition and soil drainage type, reflecting the similar proportions of the three different shade-tolerance types among stands.

Chlorophyll (Chl) and epidermal polyphenol (EPhen) contents were estimated in vivo using two optical leaf-clips, SPAD-502 and Dualex, respectively. The area-based measurements were transformed into mass-based data by taking into account the leaf dry mass per area (LMA). Measurements were performed on forest trees and on saplings grown under controlled conditions. While LMA increased with irradiance along a vertical transect in a beech canopy or in saplings grown under different and increasing irradiance levels, mass-based EPhen (EPhen m ) increased, whereas mass-based Chl (Chl m ) decreased. This was a signature of a gradual switch of investment from protein into polyphenol production. A similar signature was obtained in saplings grown on nitrogen-deficient soil with respect to fertilized controls. However, nitrogen effects remained moderate compared to irradiance-induced effects. EPhen m and Chl m both declined with plant ageing-induced increases in LMA, under all tested growth conditions. This was a signature of an accumulation of dry matter that diluted Chl and EPhen. The described competition between Chl and EPhen in leaves fits well with the predictions of the Protein Competition Model (PCM), that is, that the total leaf mass-based polyphenols content (Phen t ) is controlled by the competition between protein and polyphenol biosynthetic pathways and its metabolic regulation.Key-words : acclimation to light; chlorophyll; Dualex; epidermal polyphenols; leaf; LMA; nitrogen; phenylpropanoids; SPAD.Abbreviations : Chl, chlorophyll; Chl a , area-based chlorophyll content; Chl m , mass-based chlorophyll content; EPhen, epidermal polyphenols; EPhen a , area-based epidermal polyphenols content; EPhen m , mass-based epidermal polyphenol content; LMA, leaf dry mass per area; N m , mass-based leaf nitrogen content; PAL, phenylalanine ammonia lyase; PCM, Protein Competition Model; Phen, polyphenols; Phen t , total leaf mass-based polyphenols content; Prot t , total leaf mass-based protein content; SLA, specific leaf area ( = 1/LMA).

Aim Climate warming reshuffles biological assemblages towards less cold‐adapted but more warm‐adapted species, a process coined thermophilization. However, the velocity at which this process is happening generally lags behind the velocity of climate change, generating a climatic debt the temporal dynamics of which remain misunderstood. Relying on high‐resolution time series of vegetation data from a long‐term monitoring network of permanent forest plots, we aim at quantifying the temporal dynamics – up to a yearly resolution – of the climatic debt in the understorey of temperate forests before identifying the key determinants that modulate it. Location France. Time period 1995–2017. Taxa studied Vascular plants. Methods We used the community temperature index (CTI) to produce a time series of understorey plant community thermophilization, which we subsequently compared to a time series of mean annual temperature changes over the same period and for the same sites. The direction and magnitude of the difference (i.e., the climatic debt) was finally analysed using linear mixed‐effect models to assess the relative contributions of abiotic and biotic determinants, including forest stand characteristics. Results We found a significant increase in CTI values over time (0.08–0.09 °C/decade), whereas the velocity of mean annual temperature changes was three times higher over the same period (0.22–0.28 °C/decade). Hence, the climatic debt increased over time and was greater in forest stands with higher basal area or older trees as well as under warmer macroclimate. By contrast, a greater frequency of anthropogenic disturbances decreased the climatic debt, while natural disturbances and herbivory had no impact. Conclusions Although often overlooked in understanding the climatic debt of forest biodiversity, changes in forest stand characteristics may modulate the climatic debt by locally modifying microclimatic conditions. Notably, the buffering effect of the upper canopy layer implies microclimate dynamics that may provide more time for understorey plant communities to locally adapt.

Temperature effects on photosynthesis were studied in seedlings of evergreen Mediterranean cork oak (Quercus suber L.). Responses to changes in temperature and the temperature optima of maximal carboxylation rate (V(cmax)) and maximal light-driven electron flux (J(max)) were estimated from gas exchange measurements and a leaf-level photosynthesis model. The estimated temperature optima were approximately 34 and 33 degrees C for V(cmax) and J(max), respectively, which fall within the lower range of temperature optima previously observed in deciduous tree species. The thermostability of the photosynthetic apparatus was estimated according to the temperature at which basal chlorophyll a fluorescence begins to increase (T(c)). The T(c) was highly variable, increasing from 42 to 51 degrees C when ambient temperature rose from 10 to 40 degrees C, and increasing from 44 to 54 degrees C with decreasing soil water availability while net CO(2) assimilation rate dropped to almost zero. When a heat shock was imposed, an additional small increase in T(c) was observed in drought-stressed and control seedlings. Maximal T(c) values following heat shock were about 56 degrees C, which, to our knowledge, are the highest values that have been observed in tree species. In conclusion, the intrinsic temperature responses of cork oak did not differ from those of other species (similar T(c) under ambient temperature and water availability, and relatively low thermal optima for photosynthetic capacity in seedlings grown at cool temperatures). However, the large ability of cork oak to acclimate to drought and elevated temperature may be an important factor in the tolerance of this evergreen Mediterranean species to summer drought and high temperatures.

Large wild ungulates are a major biotic factor shaping plant communities. They influence species abundance and occurrence directly by herbivory and plant dispersal, or indirectly by modifying plant-plant interactions and through soil disturbance. In forest ecosystems, researchers' attention has been mainly focused on deer overabundance. Far less is known about the effects on understory plant dynamics and diversity of wild ungulates where their abundance is maintained at lower levels to mitigate impacts on tree regeneration. We used vegetation data collected over 10 years on 82 pairs of exclosure (excluding ungulates) and control plots located in a nation-wide forest monitoring network (Renecofor). We report the effects of ungulate exclusion on (i) plant species richness and ecological characteristics, (ii) and cover percentage of herbaceous and shrub layers. We also analyzed the response of these variables along gradients of ungulate abundance, based on hunting statistics, for wild boar (Sus scrofa), red deer (Cervus elaphus) and roe deer (Capreolus capreolus). Outside the exclosures, forest ungulates maintained higher species richness in the herbaceous layer (+15%), while the shrub layer was 17% less rich, and the plant communities became more light-demanding. Inside the exclosures, shrub cover increased, often to the benefit of bramble (Rubus fruticosus agg.). Ungulates tend to favour ruderal, hemerobic, epizoochorous and non-forest species. Among plots, the magnitude of vegetation changes was proportional to deer abundance. We conclude that ungulates, through the control of the shrub layer, indirectly increase herbaceous plant species richness by increasing light reaching the ground. However, this increase is detrimental to the peculiarity of forest plant communities and contributes to a landscape-level biotic homogenization. Even at population density levels considered to be harmless for overall plant species richness, ungulates remain a conservation issue for plant community composition.

Leaf photosynthesis is known to acclimate to the actual irradiance received by the different layers of a canopy. This acclimation is usually described in terms of changes in leaf structure, and in photosynthetic capacity. Photosynthetic capacity is likely to be affected by mesophyll conductance to CO(2) which has seldom been assessed in tree species, and whose plasticity in response to local irradiance is still poorly known. Structural [N and chlorophyll content, leaf mass to area ratio (LMA)] and functional leaf traits [maximum carboxylation rate (V(cmax)), maximum light-driven electron flux (J(max)), and mesophyll conductance (g(i))] were assessed by measuring leaf response curves of net CO(2) assimilation versus intercellular CO(2) partial pressure, along a vertical profile across a beech canopy, and by fitting a version of the Farquhar model including g(i). The measurements were repeated five times during a growth season to catch potential seasonal variation. Irradiance gradients resulted in large decreasing gradients of LMA, g(i), V(cmax), and J(max). Relative allocation of leaf N to the different photosynthetic processes was only slightly affected by local irradiance. Seasonal changes after leaf expansion and before induction of leaf senescence were only minor. Structural equation modelling confirmed that LMA was the main driving force for changes in photosynthetic traits, with only a minor contribution of leaf Nitrogen content. In conclusion, mesophyll conductance to CO(2) displays a large plasticity that scales with photosynthetic capacity across a tree canopy, and that it is only moderately (if at all) affected by seasonal changes in the absence of significant soil water depletion.