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The productivity and climate feedbacks of tropical forests depend on tree physiological responses to warmer and, over large areas, seasonally drier conditions. However, knowledge regarding such responses is limited due to data scarcity. We studied the impact of growth temperature on net photosynthesis (An), maximum rates of Rubisco carboxylation at 25 °C (Vcmax25), stomatal conductance (gs) and the slope parameter of the stomatal conductance-photosynthesis model (g1), in 10 early successional (ES) and 8 late-successional (LS) tropical tree species grown at three sites along an elevation gradient in Rwanda, differing by 6.8 °C in daytime ambient air temperature. The effect of seasonal drought on An was also investigated. We found that warm climate decreased wet-season An in LS species, but not in ES species. Values of Vcmax25 were lower at the warmest site across both successional groups, and An and Vcmax25 were higher in ES compared with LS species. Stomatal conductance exhibited no significant site differences and g1 was similar across both sites and successional groups. Drought strongly reduced An at warmer sites but not at the coolest montane site and this response was similar in both ES and LS species. Our results suggest that warming has negative effects on leaf-level photosynthesis in LS species, while both LS and ES species suffer photosynthesis declines in a warmer climate with more pronounced droughts. The contrasting responses of An between successional groups may lead to shifts in species’ competitive balance in a warmer world, to the disadvantage of LS trees.

The effect of temperature change on leaf physiology has been extensively studied in temperate trees and to some extent in boreal and tropical tree species. While increased temperature typically stimulates leaf CO2 assimilation and tree growth in high-altitude ecosystems, tropical species are often negatively affected. These trees may operate close to their temperature optima and have a limited thermal acclimation capacity due to low seasonal and historical variation in temperature. To test this hypothesis, we studied the extent to which the temperature sensitivities of leaf photosynthesis and respiration acclimate to growth temperature in four common African tropical tree species. Tree seedlings native to different altitudes and therefore adapted to different growth temperatures were cultivated at three different temperatures in climate-controlled chambers. We estimated the acclimation capacity of the temperature sensitivities of light-saturated net photosynthesis, the maximum rates of Rubisco carboxylation (Vcmax) and thylakoid electron transport (J), and dark respiration. Leaf thylakoid membrane lipid composition, nitrogen content and leaf mass per area were also analyzed. Our results showed that photosynthesis in tropical tree species acclimated to higher growth temperatures, but that this was weakest in the species originating from the coolest climate. The temperature optimum of J acclimated significantly in three species and variation in J was linked to changes in the thylakoid membrane lipid composition. For Vcmax, there was only evidence of significant acclimation of optimal temperature in the lowest elevation species. Respiration acclimated to maintain homeostasis at growth temperature in all four species. Our results suggest that the lowest elevation species is better physiologically adapted to acclimate to high growth temperatures than the highest elevation species, indicating a potential shift in competitive balance and tree community composition to the disadvantage of montane tree species in a warmer world.

Several studies have suggested that CO2 transport in the transpiration stream can considerably bias estimates of root and stem respiration in ring-porous and diffuse-porous tree species. Whether this also happens in species with tracheid xylem anatomy and lower sap flow rates, such as conifers, is currently unclear. We infused 13C-labelled solution into the xylem near the base of two 90-year-old Pinus sylvestris L. trees. A custom-built gas exchange system and an online isotopic analyser were used to sample the CO2 efflux and its isotopic composition continuously from four positions along the bole and one upper canopy shoot in each tree. Phloem and needle tissue 13C enrichment was also evaluated at these positions. Most of the 13C label was lost by diffusion within a few metres of the infusion point indicating rapid CO2 loss during vertical xylem transport. No 13C enrichment was detected in the upper bole needle tissues. Furthermore, mass balance calculations showed that c. 97% of the locally respired CO2 diffused radially to the atmosphere. Our results support the notion that xylem CO2 transport is of limited magnitude in conifers. This implies that the concerns that stem transport of CO2 derived from root respiration biases chamber-based estimates of forest carbon cycling may be unwarranted for mature conifer stands.

Trees are able to reduce their carbon (C) losses by refixing some of the CO2 diffusing out of their stems through corticular photosynthesis. Previous studies have shown that under ideal conditions the outflowing CO2 can be completely assimilated in metabolically active, young stem and branch tissues. Fewer studies have, however, been carried out on the older stem sections of large trees and, accordingly, the importance of refixation is still unclear under natural environmental conditions. We investigated the spatial and temporal variation in refixation in ~90-year-old boreal Scots pine (Pinus sylvestris L.) trees by utilizing month-long continuous measurements of stem CO2 efflux (Ec) made at four heights along the bole. Refixation rates were found to vary considerably along the bole, leading to a 28% reduction in long-term Ec in the upper stem compared with a negligible reduction at breast height. This vertical pattern correlated with variation in light availability, bark chlorophyll content and bark type. Analysis of the vertical and diurnal patterns in Ec further suggested that the influence of sap flow on the observed daytime reduction in Ec was small. The areal rates of corticular photosynthesis were much lower than previous estimates of photosynthetic rates per unit leaf area from the same trees, implying that the impact of refixation on tree-scale C uptake was small. However, upscaling of refixation indicated that 23-27% of the potential Ec was refixed by the bole and the branches, thereby significantly reducing the woody tissue C losses. Thus, our results suggest that refixation needs to be considered when evaluating the aboveground C cycling of mature P. sylvestris stands and that breast-height estimates should not be extrapolated to the whole tree.

Stem CO2 efflux is known to vary seasonally and vertically along tree stems. However, annual tree- and stand-scale efflux estimates are commonly based on measurements made only a few times a year, during daytime and at breast height. In this study, the effect of these simplifying assumptions on annual efflux estimates and their influence on the estimates of the importance of stems in stand-scale carbon cycling are evaluated. In order to assess the strength of seasonal, diurnal and along-stem variability in CO2 efflux, half-hourly measurements were carried out at three heights on three mature Norway spruce (Picea abies (L.) Karst.) trees over a period of 3 years. Making the common assumption of breast height efflux rates being representative of the entire stem was found to result in underestimations of 10-17% in the annual tree-scale CO2 efflux. Upscaling using only daytime measurements from breast height increased the underestimation to 15-20%. Furthermore, the results show that the strength of the vertical gradient varies seasonally, being strongest in the early summer and non-existent during the cool months. The observed seasonality in the vertical CO2 efflux gradient could not be explained by variation in stem temperature, temperature response of the CO2 efflux (Q(10)), outer-bark permeability, CO2 transport in the xylem or CO2 release from the phloem. However, the estimated CO2 concentration immediately beneath the bark was considerably higher in the upper stem during the main period of diameter growth, coinciding with the strongest vertical efflux gradient. These results suggest that higher growth rates in the upper stem are the main cause for the observed vertical variation in the stem CO2 effluxes. Furthermore, the results indicate that accounting for the vertical efflux variation is essential for assessments of the importance of stems in stand-scale carbon cycling.

Photosynthesis is highly responsive to environmental and physiological variables, including phenology, foliage nitrogen (N) content, atmospheric CO2 concentration ([CO2]), irradiation (Q), air temperature (T) and vapour pressure deficit (D). Each of these responses is likely to be modified by long-term changes in climatic conditions such as rising air temperature and [CO2]. When modelling photosynthesis under climatic changes, which parameters are then most important to calibrate for future conditions? To assess this, we used measurements of shoot carbon assimilation rates and microclimate conditions collected at Flakaliden, northern Sweden. Twelve 40-year-old Norway spruce trees were enclosed in whole-tree chambers and exposed to elevated [CO2] and elevated air temperature, separately and in combination. The treatments imposed were elevated temperature, +2.8 °C in July/August and +5.6 °C in December above ambient, and [CO2] (ambient CO2 ∼370 μ mol mol(-1), elevated CO2 ∼700 μ mol mol(-1)). The relative importance of parameterization of Q, T and D responses for effects on the photosynthetic rate, expressed on a projected needle area, and the annual shoot carbon uptake was quantified using an empirical shoot photosynthesis model, which was developed and fitted to the measurements. The functional form of the response curves was established using an artificial neural network. The [CO2] treatment increased annual shoot carbon (C) uptake by 50%. Most important was effects on the light response curve, with a 67% increase in light-saturated photosynthetic rate, and a 52% increase in the initial slope of the light response curve. An interactive effect of light saturated photosynthetic rate was found with foliage N status, but no interactive effect for high temperature and high CO2. The air temperature treatment increased the annual shoot C uptake by 44%. The most important parameter was the seasonality, with an elongation of the growing season by almost 4 weeks. The temperature response curve was almost flat over much of the temperature range. A shift in temperature optimum had thus an insignificant effect on modelled annual shoot C uptake. The combined temperature and [CO2] treatment resulted in a 74% increase in annual shoot C uptake compared with ambient conditions, with no clear interactive effects on parameter values. (Less)

Quantifying the adjustments of leaf respiration in response to seasonal temperature variation and climate warming is crucial because carbon loss from vegetation is a large but uncertain part of the global carbon cycle. We grew fast-growing Eucalyptus globulus Labill. trees exposed to +3 °C warming and elevated CO2 in 10-m tall whole-tree chambers and measured the temperature responses of leaf mitochondrial respiration, both in light (RLight) and in darkness (RDark), over a 20-40 °C temperature range and during two different seasons. RLight was assessed using the Laisk method. Respiration rates measured at a standard temperature (25 °C - R25) were higher in warm-grown trees and in the warm season, related to higher total leaf nitrogen (N) investment with higher temperatures (both experimental and seasonal), indicating that leaf N concentrations modulated the respiratory capacity to changes in temperature. Once differences in leaf N were accounted for, there were no differences in R25 but the Q10 (i.e., short-term temperature sensitivity) was higher in late summer compared with early spring. The variation in RLight between experimental treatments and seasons was positively correlated with carboxylation capacity and photorespiration. RLight was less responsive to short-term changes in temperature than RDark, as shown by a lower Q10 in RLight compared with RDark. The overall light inhibition of R was ∼40%. Our results highlight the dynamic nature of leaf respiration to temperature variation and that the responses of RLight do not simply mirror those of RDark. Therefore, it is important not to assume that RLight is the same as RDark in ecosystem models, as doing so may lead to large errors in predicting plant CO2 release and productivity.

According to well-known biochemical and biophysical mechanisms, the stimulation of C(3) photosynthesis by elevated atmospheric CO(2) concentration ([CO(2)]) is strongly modified by changes in temperature and radiation. In order to investigate whether a static parameterization of the commonly used Farquhar et al. model of photosynthesis (i.e., without CO(2)-induced seasonal or thermal acclimation of photosynthetic capacity) can accurately predict these interactions in mature boreal Norway spruce (Picea abies (L.) Karst.) during the frost-free part of the growing season, shoot gas exchange was continuously measured on trees during their second/third year of exposure to ambient or doubled [CO(2)] inside whole-tree chambers. The relative CO(2)-induced enhancement of net photosynthesis (A(n)) at a given temperature remained stable over the study period, but increased strongly with temperature and radiation, in agreement with predictions by the model. Light-saturated A(n) (+67% at 20 °C), dark respiration (+36%) and intercellular to ambient [CO(2)] ratio (c(i)/c(a); +27%) were significantly increased by CO(2) treatment. Stomatal conductance (g(s)) was not significantly affected. Our results demonstrate that the Farquhar et al. model of photosynthesis has the capability to predict interactions between [CO(2)] and seasonal weather variability on A(n) in Norway spruce during the non-frost growing season without accounting for CO(2)-induced seasonal and/or thermal photosynthetic acclimation. However, stomatal model assumptions of reduced g(s) and constant c(i)/c(a) under rising atmospheric [CO(2)] did not hold.

A long-established theoretical result states that, for a given total canopy nitrogen (N) content, canopy photosynthesis is maximized when the within-canopy gradient in leaf N per unit area (N(a)) is equal to the light gradient. However, it is widely observed that N(a) declines less rapidly than light in real plant canopies. Here we show that this general observation can be explained by optimal leaf acclimation to light subject to a lower-bound constraint on the leaf mass per area (m(a)). Using a simple model of the carbon-nitrogen (C-N) balance of trees with a steady-state canopy, we implement this constraint within the framework of the MAXX optimization hypothesis that maximizes net canopy C export. Virtually all canopy traits predicted by MAXX (leaf N gradient, leaf N concentration, leaf photosynthetic capacity, canopy N content, leaf-area index) are in close agreement with the values observed in a mature stand of Norway spruce trees (Picea abies L. Karst.). An alternative upper-bound constraint on leaf photosynthetic capacity (A(sat)) does not reproduce the canopy traits of this stand. MAXX subject to a lower bound on m(a) is also qualitatively consistent with co-variations in leaf N gradient, m(a) and A(sat) observed across a range of temperate and tropical tree species. Our study highlights the key role of constraints in optimization models of plant function.

Accumulated carbon uptake, apparent quantum yield (AQY) and light-saturated net CO2 assimilation (A(sat)) were used to assess the responses of photosynthesis to environmental conditions during spring for three consecutive years. Whole-tree chambers were used to expose 40-year-old field-grown Norway spruce trees in northern Sweden to an elevated atmospheric CO2 concentration, [CO2], of 700 mu mol CO2 mol(-1) (C-E) and an air temperature (T) between 2.8 and 5.6 degrees C above ambient T (T-E), during summer and winter. Net shoot CO2 exchange (A(net)) was measured continuously on 1-year-old shoots and was used to calculate the accumulated carbon uptake and daily A(sat) and AQY. The accumulated carbon uptake, from 1 March to 30 June, was stimulated by 33, 44 and 61% when trees were exposed to C-E, T-E, and C-E and T-E combined, respectively. Air temperature strongly influenced the timing and extent of photosynthetic recovery expressed as AQY and A(sat) during the spring. Under elevated T (T-E), the recovery of AQY and A(sat) commenced similar to 10 days earlier and the activity of these parameters was significantly higher throughout the recovery period. In the absence of frost events, the photosynthetic recovery period was less than a week. However, frost events during spring slowed recovery so that full recovery could take up to 60 days to complete. Elevated [CO2] stimulated AQY and A(sat) on average by similar to 10 and similar to 50%, respectively, throughout the recovery period, but had minimal or no effect on the onset and length of the photosynthetic recovery period during the spring. However, AQY, A(sat) and A(net) all recovered at significantly higher T (average +2.2 degrees C) in T-E than in T-A, possibly caused by acclimation or by shorter days and lower light levels during the early part of the recovery in T-E compared with T-A. The results suggest that predicted future climate changes will cause prominent stimulation of photosynthetic CO2 uptake in boreal Norway spruce forest during spring, mainly caused by elevated T, but also elevated [CO2]. However, the effects of elevated T may not be linearly extrapolated to future warmer climates. (Less)

The growth responses of mature Norway spruce (Picea abies (L.) Karst.) trees exposed to elevated [CO(2)] (CE; 670-700 ppm) and long-term optimized nutrient availability or elevated air temperature (TE; ±3.9 °C) were studied in situ in northern Sweden in two 3 year field experiments using 12 whole-tree chambers in ca. 40-year-old forest. The first experiment (Exp. I) studied the interactions between CE and nutrient availability and the second (Exp. II) between CE and TE. It should be noted that only air temperature was elevated in Exp. II, while soil temperature was maintained close to ambient. In Exp. I, CE significantly increased the mean annual height increment, stem volume and biomass increment during the treatment period (25, 28, and 22%, respectively) when nutrients were supplied. There was, however, no significant positive CE effect found at the low natural nutrient availability. In Exp. II, which was conducted at the natural site fertility, neither CE nor TE significantly affected height or stem increment. It is concluded that the low nutrient availability (mainly nitrogen) in the boreal forests is likely to restrict their response to the continuous rise in [CO(2)] and/or TE.

Summary Effects of ambient and elevated temperature and atmospheric carbon dioxide concentration ([CO2]) on CO2 assimilation rate and the structural and phenological development of shoots during their first growing season were studied in 45-year-old Norway spruce trees (Picea abies (L.) Karst.) enclosed in whole-tree chambers. Continuous measurements of net assimilation rate (NAR) in individual buds and shoots were made from early bud development to late August in two consecutive years. The largest effect of elevated temperature (TE) was manifest early in the season as an earlier start and completion of shoot length development, and a 1–3-week earlier shift from negative to positive NAR compared with the ambient temperature (TA) treatments. The largest effect of elevated [CO2 ]( CE) was found later in the season, with a 30% increase in maximum NAR compared with trees in the ambient [CO2] treatments (CA), and shoots assimilating their own mass in terms of carbon earlier in the CE treatments than in the CA treatments. Once the net carbon assimilation compensation point (NACP) had been reached, TE had little or no effect on the development of NAR performance, whereas CE had little effect before the NACP. No interactive effects of TE and CE on NAR were found. We conclude that in a climate predicted for northern Sweden in 2100, current-year shoots of P. abies will assimilate their own mass in terms of carbon 20–30 days earlier compared with the current climate, and thereby significantly contribute to canopy assimilation during their first year.

Effects of elevated temperature and atmospheric CO2 concentration ([CO2]) on spring phenology of mature field-grown Norway spruce (Picea abies (L.) Karst.) trees were followed for three years. Twelve whole-tree chambers (WTC) were installed around individual trees and used to expose the trees to a predicted future climate. The predicted climate scenario for the site, in the year 2100, was 700 micromol mol-1 [CO2], and an air temperature 3 degrees C higher in summer and 5 degrees C higher in winter, compared with current conditions. Four WTC treatments were imposed using combinations of ambient and elevated [CO2] and temperature. Control trees outside the WTCs were also studied. Bud development and shoot extension were monitored from early spring until the termination of elongation growth. Elevated air temperature hastened both bud development and the initiation and termination of shoot growth by two to three weeks in each study year. Elevated [CO2] had no significant effect on bud development patterns or the length of the shoot growth period. There was a good correlation between temperature sum (day degrees>or=0 degrees C) and shoot elongation, but a precise timing of bud burst could not be derived by using an accumulation of temperature sums.

Net CO2 exchange in a 35-year-old boreal Norway spruce (Picea abies (L.) Karst.) forest in northern Sweden was measured at the shoot (NSE), tree (NTE) and ecosystem levels (NEE) by means of shoot cuvettes, whole-tree chambers and the eddy covariance technique, respectively. We compared the dynamics of gross primary production (GPP) at the three levels during the course of a single week. The diurnal dynamics of GPP at each level were estimated by subtracting half-hourly or hourly model-estimated values of total respiration (excluding light-dependent respiration) from net CO(2) exchange. The relationship between temperature and total respiration at each level was derived from nighttime measurements of NSE, NTE and NEE over the course of 1 month. There was a strong linear relationship (r2 = 0.93) between the hourly estimates of GPP at the shoot and tree levels, but the correlation between shoot- and ecosystem-level GPP was weaker (r2 = 0.69). However, the correlation between shoot- and ecosystem-level GPP was improved (r2 = 0.88) if eddy covariance measurements were restricted to periods when friction velocity was > or = 0.5 m s(-1). Daily means were less dependent on friction velocity, giving an r2 value of 0.94 between shoot- and ecosystem-level GPP. The correlation between shoot and tree levels also increased when daily means were compared (r2 = 0.98). Most of the measured variation in carbon exchange rate among the shoot, tree and ecosystem levels was the result of periodic low coupling between vegetation and the atmosphere at the ecosystem level. The results validate the use of measurements at the shoot and tree level for analyzing the contribution of different compartments to net ecosystem CO2 exchange.