Temperature dependence of two parameters in a photosynthesis model

The temperature dependence of the photosynthetic parameters V_cmax, the maximum catalytic rate of the enzyme Rubisco, and J_max, the maximum electron transport rate, were examined using published datasets. An Arrehenius equation, modified to account for decreases in each parameter at high temperatures, satisfactorily described the temperature response for both parameters. There was remarkable conformity in V_cmax and J_max between all plants at T_leaf < 25 °C, when each parameter was normalized by their respective values at 25 °C (V_cmax0 and J_max0), but showed a high degree of variability between and within species at T_leaf > 30 °C. For both normalized V_cmax and J_max, the maximum fractional error introduced by assuming a common temperature response function is < ± 0·1 for most plants and < ± 0·22 for all plants when T_leaf < 25 °C. Fractional errors are typically < ± 0·45 in the temperature range 25–30 °C, but very large errors occur when a common function is used to estimate the photosynthetic parameters at temperatures > 30 °C. The ratio J_max/V_cmax varies with temperature, but analysis of the ratio at T_leaf = 25 °C using the fitted mean temperature response functions results in J_max0/V_cmax0 = 2·00 ± 0·60 (SD, n = 43).     

Temperature response of parameters of a biochemically based model of photosynthesis. II. A review of experimental data

The temperature dependence of C₃ 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.     

Acclimation of photosynthesis in a boreal grass (Phalaris arundinacea L.) under different temperature, CO2, and soil water regimes

The aim of this work was to study the acclimation of photosynthesis in a boreal grass (Phalaris arundinacea L.) grown in controlled environment chambers under elevated temperature (ambient + 3.5°C) and CO₂ (700 μmol mol⁻¹) with varying soil water regimes. More specifically, we studied, during two development stages (early: heading; late: florescence completed), how the temperature response of light-saturated net photosynthetic rate (P _sat), maximum rate of ribulose-1,5-bisphosphate carboxylase/oxygenase activity (V _cmax) and potential rate of electron transport (J _max) acclimatized to the changed environment. During the early growing period, we found a greater temperature-induced enhancement of P _sat at higher measurement temperatures, which disappeared during the late stage. Under elevated growth temperature, V _cmax and J _max at lower measurement temperatures (5–15°C) were lower than those under ambient growth temperature during the early period. When the measurements were done at 20–30°C, the situation was the opposite. During the late growing period, V _cmax and J _max under elevated growth temperature were consistently lower across measurement temperatures. CO₂ enrichment significantly increased P _sat with higher intercellular CO₂ compared to ambient CO₂ treatment, however, elevated CO₂ slightly decreased V _cmax and J _max across measurement temperatures, probably due to down-regulation acclimation. For two growing periods, soil water availability affected the variation in photosynthesis and biochemical parameters much more than climatic treatment did. Over two growing periods, V _cmax and J _max were on average 36.4 and 30.6%, respectively, lower with low water availability compared to high water availability across measurement temperatures. During the late growing period, elevated growth temperature further reduced the photosynthesis under low water availability. V _cmax and J _max declined along with the decrease in nitrogen content of leaves as growing period progressed, regardless of climatic treatment and water regime. We suggest that, for grass species, seasonal acclimation of the photosynthetic parameters under varying environmental conditions needed to be identified to fairly estimate the whole-life photosynthesis.     

Photosynthetic parameters,dark respiration and leaf traits in the canopy of a Peruvian tropical montane cloud forest

Few data are available describing the photosynthetic parameters of the leaves of tropical montane cloud forests (TMCF). Here, we present a study of photosynthetic leaf traits (V _cmax and J _max), foliar dark respiration (R _d), foliar nitrogen (N) and phosphorus (P), and leaf mass per area (LMA) throughout the canopy for five different TMCF species at 3025 m a.s.l. in Andean Peru. All leaf traits showed a significant relationship with canopy height when expressed on an area basis, and V _cmax-area and J _max-area almost halved when descending through the TMCF canopy. When corrected to a common temperature, average V _cmax and J _max on a leaf area basis were similar to lowland tropical values, but lower when expressed on a mass basis, because of the higher TMCF LMA values. By contrast, R _d on an area basis was higher than found in tropical lowland forests at a common temperature, and similar to lowland forests on a mass basis. The TMCF J _max–V _cmax relationship was steeper than in other tropical biomes, and we propose that this can be explained by either the light conditions or the relatively low VPD in the studied TMCF. Furthermore, V _cmax had a significant—though relatively weak and shallow—relationship with N on an area basis, but not with P, which is consistent with the general hypothesis that TMCFs are N rather than P limited. Finally, the observed V _cmax–N relationship (i.e., maximum photosynthetic nitrogen use efficiency) was distinctly different from those in tropical and temperate regions, probably because the TMCF leaves compensate for reduced Rubisco activity in cool environments.     

Retrieval of seasonal variation in photosynthetic capacity from multi-source vegetation indices

Seasonal information on photosynthetic-capacity parameters (maximum carboxylation velocity, V_cmax; and maximum rate of electron transport, J_max) plays an important role in accurate simulation of carbon fixation in gas-exchange models. Exact inclusion of seasonal information on photosynthetic-capacity parameters into the models has been an irresolvable challenge. This paper investigated the relationships between vegetation indices (from multiple sources) and photosynthetic-capacity parameters of three beech forest stands (Fagus crenata) along an elevation gradient in the cold-temperate zone of Japan, over the entire growing season of 2006. Diverse vegetation indices were examined in terms of spectral, spatial and temporal scales; ranging from meteorological sensor-based broadband indices to hyperspectral data-based narrowband indices, to simulated MODIS (MODerate-resolution Imaging Spectroradiometer) indices based on hyperspectral data, and finally satellite-borne MODIS vegetation indices. Regression analysis revealed that all examined indices, with the exception of the downloaded MODIS products, had significant regression relationships with photosynthetic parameters (P < 0.001) when all data were pooled. Among the different indices, the simulated MODIS NDVI (Normalized Difference Vegetation Index) performed the best for both V_cmax and J_max (R² = 0.81 and 0.73, respectively). Site differences were apparent, as the simulated MODIS NDVI performed the best in exponential regressions for the 550 m site, while broadband NDVI performed best in exponential regression models for the 900 m site. The broadband SR (Simple Ratio) in relation to V_cmax performed best with respect to a linear model, whereas the broadband NDVI with J_max performed the best in an exponential model for the 1500 m site. The results reveal that vegetation indices which are obtained across different scales nevertheless retain tight relationships with canopy-scale photosynthetic-capacity parameters. The established relationships were inversely applicable to derive seasonal trajectories of photosynthetic-capacity parameters. Thus, new insight and confidence is gained for using remotely estimated photosynthetic parameters, even though most previous research works were limited on linking of vegetation indices with biophysical parameters. The control effect of physiological capacity on reflectance and further on vegetation indices has not been adequately established and thus needs further orientation for rigorous research work.     

Comparison of the A–Cc curve fitting methods in determining maximum ribulose 1·5-bisphosphate carboxylase/oxygenase carboxylation rate, potential light saturated electron transport rate and leaf dark respiration

A review of the literature revealed that a variety of methods are currently used for fitting net assimilation of CO₂–chloroplastic CO₂ concentration (A–C_c) curves, resulting in considerable differences in estimating the A–C_c parameters [including maximum ribulose 1·5‐bisphosphate carboxylase/oxygenase (Rubisco) carboxylation rate (V_cmax), potential light saturated electron transport rate (J_max), leaf dark respiration in the light (R_d), mesophyll conductance (g_m) and triose‐phosphate utilization (TPU)]. In this paper, we examined the impacts of fitting methods on the estimations of V_cmax, J_max, TPU, R_d and g_m using grid search and non‐linear fitting techniques. Our results suggested that the fitting methods significantly affected the predictions of Rubisco‐limited (A_c), ribulose 1,5‐bisphosphate‐limited (A_j) and TPU‐limited (A_p) curves and leaf photosynthesis velocities because of the inconsistent estimate of V_cmax, J_max, TPU, R_d and g_m, but they barely influenced the J_max : V_cmax, V_cmax : R_d and J_max : TPU ratio. In terms of fitting accuracy, simplicity of fitting procedures and sample size requirement, we recommend to combine grid search and non‐linear techniques to directly and simultaneously fit V_cmax, J_max, TPU, R_d and g_m with the whole A–C_c curve in contrast to the conventional method, which fits V_cmax, R_d or g_m first and then solves for V_cmax, J_max and/or TPU with V_cmax, R_d and/or g_m held as constants.     

The relationship of leaf photosynthetic traits – Vcmax and Jmax – to leaf nitrogen,leaf phosphorus,and specific leaf area: a meta‐analysis and modeling study

Great uncertainty exists in the global exchange of carbon between the atmosphere and the terrestrial biosphere. An important source of this uncertainty lies in the dependency of photosynthesis on the maximum rate of carboxylation (V_cmax) and the maximum rate of electron transport (J_max). Understanding and making accurate prediction of C fluxes thus requires accurate characterization of these rates and their relationship with plant nutrient status over large geographic scales. Plant nutrient status is indicated by the traits: leaf nitrogen (N), leaf phosphorus (P), and specific leaf area (SLA). Correlations between V_cmax and J_max and leaf nitrogen (N) are typically derived from local to global scales, while correlations with leaf phosphorus (P) and specific leaf area (SLA) have typically been derived at a local scale. Thus, there is no global-scale relationship between V_cmax and J_max and P or SLA limiting the ability of global-scale carbon flux models do not account for P or SLA. We gathered published data from 24 studies to reveal global relationships of V_cmax and J_max with leaf N, P, and SLA. V_cmax was strongly related to leaf N, and increasing leaf P substantially increased the sensitivity of V_cmax to leaf N. J_max was strongly related to V_cmax, and neither leaf N, P, or SLA had a substantial impact on the relationship. Although more data are needed to expand the applicability of the relationship, we show leaf P is a globally important determinant of photosynthetic rates. In a model of photosynthesis, we showed that at high leaf N (3 gm⁻²), increasing leaf P from 0.05 to 0.22 gm⁻² nearly doubled assimilation rates. Finally, we show that plants may employ a conservative strategy of J_max to V_cmax coordination that restricts photoinhibition when carboxylation is limiting at the expense of maximizing photosynthetic rates when light is limiting.     

Temperature response of parameters of a biochemically based model of photosynthesis. I. Seasonal changes in mature maritime pine (Pinus pinaster Ait.)

Responses of plant processes to temperature may vary according to the time scale on which they are measured. In this study, both short‐term and seasonal responses of photosynthesis to temperature were examined. A field study of seasonal changes in the temperature response of photosynthesis was conducted on two provenances, French and Moroccan, of mature maritime pine (Pinus pinaster Ait.). Measurements were made every 2 months over a 1‐year period and used to parameterize a mechanistic model of photosynthesis. Temperature responses of maximum Rubisco activity, V_cmax, and potential electron transport rate, J_max, were obtained for each measurement period, as was the response of stomatal conductance, g_s, to water vapour pressure deficit (VPD). Absolute values of V_cmax and J_max at 25 °C were related to needle nitrogen content, N_area.N_area, and thus V_cmax and J_max, were negatively correlated with the mean minimum temperature in the month preceding measurements. The ratio of J_max : V_cmax at 25 °C varied between 1 and 1·7 but did not show any seasonal trend. Nor was there any seasonal trend in the relative temperature response of V_cmax, which had an activation energy H_a of approximately 57 kJ mol^?1 throughout the experiment. The activation energy of J_max was also close to constant throughout the experiment, averaging 39 kJ mol^?1. For the French provenance, the optimal temperature of J_max was positively correlated with the maximum temperature of the previous day, but no such correlation was found for the Moroccan provenance. The response of g_s to VPD also varied seasonally, with much stronger stomatal closure in winter months. Taken together, these results implied a translational shift downwards of the photosynthetic temperature response curve with increasing T_prev, and a shift in the temperature optimum of photosynthesis of 5–10 °C between summer and winter. These results illustrate that the short‐term temperature response of photosynthesis varies significantly on a seasonal basis.     

Photosynthetic capacity and temperature responses of photosynthesis of rubber trees (Hevea brasiliensis Müll. Arg.) acclimate to changes in ambient temperatures

The aim of this study was to assess the temperature response of photosynthesis in rubber trees (Hevea brasiliensis Müll. Arg.) to provide data for process-based growth modeling, and to test whether photosynthetic capacity and temperature response of photosynthesis acclimates to changes in ambient temperature. Net CO₂ assimilation rate (A) was measured in rubber saplings grown in a nursery or in growth chambers at 18 and 28°C. The temperature response of A was measured from 9 to 45°C and the data were fitted to an empirical model. Photosynthetic capacity (maximal carboxylation rate, V _cmax, and maximal light driven electron flux, J _max) of plants acclimated to 18 and 28°C were estimated by fitting a biochemical photosynthesis model to the CO₂ response curves (A–C _i curves) at six temperatures: 15, 22, 28, 32, 36 and 40°C. The optimal temperature for A (T _opt) was much lower in plants grown at 18°C compared to 28°C and nursery. Net CO₂ assimilation rate at optimal temperature (A _opt), V _cmax and J _max at a reference temperature of 25°C (V _cmax25 and J _max25) as well as activation energy of V _cmax and J _max (E _aV and E _aJ) decreased in individuals acclimated to 18°C. The optimal temperature for V _cmax and J _max could not be clearly defined from our response curves, as they always were above 36°C and not far from 40°C. The ratio J _max25/V _cmax25 was larger in plants acclimated to 18°C. Less nitrogen was present and photosynthetic nitrogen use efficiency (V _cmax25/N _a) was smaller in leaves acclimated to 18°C. These results indicate that rubber saplings acclimated their photosynthetic characteristics in response to growth temperature, and that higher temperatures resulted in an enhanced photosynthetic capacity in the leaves, as well as larger activation energy for photosynthesis.     

The response of photosynthetic model parameters to temperature and nitrogen concentration in Pinus radiata D. Don

Responses of photosynthesis (A) to intercellular CO₂ concentration (c_i) in 2-year-old Pinus radiata D. Don seedlings were measured at a range of temperatures in order to parametrize a biophysical model of leaf photosynthesis. Increasing leaf temperature from 8 to 30°C caused a 4-fold increase in V_cmax, the maximum rate of carboxylation (10.7–43.3 μol m^?2 s^?1 and a 3-fold increase in J_max, the maximum electron transport rate (20.5–60.2 μmol m ^?2 s^?1). The temperature optimum for J_max was lower than that for V_cmax, causing a decline in the ratio J_max:V_cmax from 2.0 to 1.4 as leaf temperature increased from 8 to 30°C. To determine the response of photosynthesis to leaf nitrogen concentration, additional measurements were made on seedlings grown under four nitrogen treatments. Foliar N concentrations varied between 0.36 and 1.27 mol kg^?1, and there were linear relationships between N concentration and both V_cmax and J_max. Measurements made throughout the crown of a plantation forest tree, where foliar N concentrations varied from 0.83 mol kg^?1 near the base to 1.54 mol kg^?1 near the leader, yielded similar relationships. These results will be useful in scaling carbon assimilation models from leaves to canopies.     

Estimation of vegetation photosynthetic capacity from space‐based measurements of chlorophyll fluorescence for terrestrial biosphere models

Photosynthesis simulations by terrestrial biosphere models are usually based on the Farquhar's model, in which the maximum rate of carboxylation (V_cmax) is a key control parameter of photosynthetic capacity. Even though V_cmax is known to vary substantially in space and time in response to environmental controls, it is typically parameterized in models with tabulated values associated to plant functional types. Remote sensing can be used to produce a spatially continuous and temporally resolved view on photosynthetic efficiency, but traditional vegetation observations based on spectral reflectance lack a direct link to plant photochemical processes. Alternatively, recent space‐borne measurements of sun‐induced chlorophyll fluorescence (SIF) can offer an observational constraint on photosynthesis simulations. Here, we show that top‐of‐canopy SIF measurements from space are sensitive to V_cmax at the ecosystem level, and present an approach to invert V_cmax from SIF data. We use the Soil‐Canopy Observation of Photosynthesis and Energy (SCOPE) balance model to derive empirical relationships between seasonal V_cmax and SIF which are used to solve the inverse problem. We evaluate our V_cmax estimation method at six agricultural flux tower sites in the midwestern US using spaced‐based SIF retrievals. Our V_cmax estimates agree well with literature values for corn and soybean plants (average values of 37 and 101 μmol m^?2 s^?1, respectively) and show plausible seasonal patterns. The effect of the updated seasonally varying V_cmax parameterization on simulated gross primary productivity (GPP) is tested by comparing to simulations with fixed V_cmax values. Validation against flux tower observations demonstrate that simulations of GPP and light use efficiency improve significantly when our time‐resolved V_cmax estimates from SIF are used, with R² for GPP comparisons increasing from 0.85 to 0.93, and for light use efficiency from 0.44 to 0.83. Our results support the use of space‐based SIF data as a proxy for photosynthetic capacity and suggest the potential for global, time‐resolved estimates of V_cmax.     

Temperature responses of photosynthetic capacity parameters were not affected by foliar nitrogen content in mature Pinus sylvestris

A key weakness in current Earth System Models is the representation of thermal acclimation of photosynthesis in response to changes in growth temperatures. Previous studies in boreal and temperate ecosystems have shown leaf‐scale photosynthetic capacity parameters, the maximum rates of carboxylation (V_cmax) and electron transport (J_max), to be positively correlated with foliar nitrogen (N) content at a given reference temperature. It is also known that V_cmax and J_max exhibit temperature optima that are affected by various environmental factors and, further, that N partitioning among the foliar photosynthetic pools is affected by N availability. However, despite the strong recent anthropogenic influence on atmospheric temperatures and N deposition to forests, little is known about the role of foliar N contents in controlling the photosynthetic temperature responses. In this study, we investigated the temperature dependencies of V_cmax and J_max in 1‐year‐old needles of mature boreal Pinus sylvestris (Scots pine) trees growing under low and high N availabilities in northern Sweden. We found that needle N status did not significantly affect the temperature responses of V_cmax or J_max when the responses were fitted to a peaked function. If such N insensitivity is a common tree trait it will simplify the interpretation of the results from gradient and multi‐species studies, which commonly use sites with differing N availabilities, on temperature acclimation of photosynthetic capacity. Moreover, it will simplify modeling efforts aimed at understanding future carbon uptake by precluding the need to adjust the shape of the temperature response curves to variation in N availability.     

Effect of leaf age and position on light-saturated CO<Subscript>2</Subscript> assimilation rate,photosynthetic capacity,and stomatal conductance in rubber trees

Shoots of the tropical latex-producing tree Hevea brasiliensis (rubber tree) grow according to a periodic pattern, producing four to five whorls of leaves per year. All leaves in the same whorl were considered to be in the same leaf-age class, in order to assess the evolution of photosynthesis with leaf age in three clones of rubber trees, in a plantation in eastern Thailand. Light-saturated CO₂ assimilation rate (A _max) decreased more with leaf age than did photosynthetic capacity (maximal rate of carboxylation, V _cmax , and maximum rate of electron transport, J _max), which was estimated by fitting a biochemical photosynthesis model to the CO₂-response curves. Nitrogen-use efficiency (A _max/N_a, N_a is nitrogen content per leaf area) decreased also with leaf age, whereas J _max and V _cmax did not correlate with N _a. Although measurements were performed during the rainy season, the leaf gas exchange parameter that showed the best correlation with A _max was stomatal conductance (g _s). An asymptotic function was fitted to the A _max-g _s relationship, with R ² = 0.85. A _max, V _cmax, J _max and g _s varied more among different whorls in the same clone than among different clones in the same whorl. We concluded that leaf whorl was an appropriate parameter to characterize leaves for the purpose of modelling canopy photosynthesis in field-grown rubber trees, and that stomatal conductance was the most important variable explaining changes in A _max with leaf age in rubber trees.     

Seasonal variation in leaf properties and ecosystem carbon budget in a cool-temperate deciduous broad-leaved forest: simulation analysis at Takayama site, Japan

Seasonal changes in gross primary production (GPP) and net ecosystem production (NEP) in temperate deciduous forests are mostly driven by environmental conditions and the phenology of leaf demography. This study addresses another factor, temporal changes in leaf properties, i.e., leaf aging from emergence to senescence. A process-based model was used to link the ecosystem-scale carbon budget with leaf-level properties on the basis of field observation and scaling procedures; temporal variations in leaf thickness (leaf mass per area, LMA), photosynthetic rubisco (V_cmax) and electron-transport (J_max) capacity, and dark respiration (R_d) were empirically parameterized. The model was applied to a cool-temperate deciduous broad-leaved forest at Takayama, in central Japan, and validated with data of net ecosystem CO₂ exchange (NEE=–NEP) measured using the eddy-covariance method. NEP of the Takayama site varied seasonally from 3 g C m^–2 day^–1 net source in late winter to 5 g C m^–2 day^–1 net sink in early to mid-summer. A sensitivity experiment showed that removing the leaf-aging effect changed the seasonal CO₂ exchange pattern, and led to overestimation of annual GPP by 6% and annual NEP by 38%. We found that seasonal variation in V_cmax affected the seasonal pattern and annual budget of CO₂ exchange most strongly; LMA and R_d had moderate influences. The rapid change in V_cmax and R_d during leaf emergence and senescence was important in evaluating GPP and NEP of the temperate deciduous forest.     

Polygonum sachalinense alters the balance between capacities of regeneration and carboxylation of ribulose‐1,5‐bisphosphate in response to growth CO2 increment but not the nitrogen allocation within the photosynthetic apparatus

The limiting step of photosynthesis changes depending on CO₂ concentration and, in theory, photosynthetic nitrogen use efficiency at a respective CO₂ concentration is maximized if nitrogen is redistributed from non‐limiting to limiting processes. It has been shown that some plants increase the capacity of ribulose‐1,5‐bisphoshate (RuBP) regeneration (evaluated as J_max) relative to the RuBP carboxylation capacity (evaluated as V_cmax) at elevated CO₂, which is in accord with the theory. However, there is no study that tests whether this change is accompanied by redistribution of nitrogen in the photosynthetic apparatus. We raised a perennial plant, Polygonum sachalinense, at two nutrient availabilities under two CO₂ concentrations. The J_max to V_cmax ratio significantly changed with CO₂ increment but the nitrogen allocation among the photosynthetic apparatus did not respond to growth CO₂. Enzymes involved in RuBP regeneration might be more activated at elevated CO₂, leading to the higher J_max to V_cmax ratio. Our result suggests that nitrogen partitioning is not responsive to elevated CO₂ even in species that alters the balance between RuBP regeneration and carboxylation. Nitrogen partitioning seems to be conservative against changes in growth CO₂ concentration.     

Changes in needle nitrogen partitioning and photosynthesis during 80 years of tree ontogeny in Picea abies

Needle nitrogen partitioning and photosynthesis of Norway spruce were studied in a forest chronosequence in Järvselja Experimental Forest, Estonia. Current- and previous-year shoots were sampled from upper and lower canopy positions in four stands, ranging in age from 13 to 82 years. A/c _i curves were determined to obtain maximum carboxylation rate (V _cmax) and maximum rate of electron transport (J _max), whereas needle nitrogen partitioning into carboxylation (P _R), bioenergetics associated with electron transport (P _B) and thylakoid light harvesting components (P _L) was calculated from the values of V _cmax, J _max and leaf chlorophyll concentration. The greatest changes in studied needle characteristics took place between tree ages of 13 and 26 years, and this pattern was independent of needle age and canopy position. Needle mass per projected area (LMA) was lowest in the 13-year-old stand and mass-based nitrogen concentration (N^M) was generally highest in that stand. The values of LMA were significantly higher and those of N^M lower in the 26-year-old stand. Mass-based V _cmax and J _max were highest in the 13-year-old stand. Area-based photosynthetic capacity was independent of tree age. The proportion of photosynthetic nitrogen (P _R, P _B and P _L) was highest and that of non-photosynthetic nitrogen lowest in the 13-year-old stand. Current-year needles had lower LMA and P _L, but higher photosynthetic capacity compared to 1-year-old foliage. Needles from lower canopy positions exhibited lower LMA, area-based nitrogen concentration and photosynthetic capacity than needles from upper canopy. The period of substantial reductions in needle photosynthetic capacity and changes in nitrogen partitioning coincides with the onset of reproductive phase during tree ontogeny.     

岩溶与非岩溶区典型植物最适光合模型和光合特征研究

为探讨不同生境典型植物光合特征对主要环境因子的响应,选择Ethier&Livingston、Ellsworth和Sharkey等3种光合生化模型拟合我国西南岩溶与非岩溶区8种典型植物的A-Ci曲线.结果表明,用Ethier&Livingston模型拟合的A-Ci曲线要优于Ellsworth模型和Sharkey模型,拟合...     

Co‐limitation of photosynthetic capacity by nitrogen and phosphorus in West Africa woodlands

Photosynthetic leaf traits were determined for savanna and forest ecosystems in West Africa, spanning a large range in precipitation. Standardized major axis fits revealed important differences between our data and reported global relationships. Especially for sites in the drier areas, plants showed higher photosynthetic rates for a given N or P when compared with relationships from the global data set. The best multiple regression for the pooled data set estimated V_cmax and J_max from N_DW and S. However, the best regression for different vegetation types varied, suggesting that the scaling of photosynthesis with leaf traits changed with vegetation types. A new model is presented representing independent constraints by N and P on photosynthesis, which can be evaluated with or without interactions with S. It assumes that limitation of photosynthesis will result from the least abundant nutrient, thereby being less sensitive to the allocation of the non‐limiting nutrient to non‐photosynthetic pools. The model predicts an optimum proportionality for N and P, which is distinct for V_cmax and J_max and inversely proportional to S. Initial tests showed the model to predict V_cmax and J_max successfully for other tropical forests characterized by a range of different foliar N and P concentrations.     

Photosynthetic parameters in seedlings of Eucalyptus grandis as affected by rate of nitrogen supply

Seedlings of Eucalyptus grandis were grown at five different rates of nitrogen supply. Once steady‐state growth rates were established, a detailed set of CO₂ and water vapour exchange measurements were made to investigate the effects of leaf nitrogen content (N), as determined by nitrogen supply rate, on leaf structural, photosynthetic, respiratory and stomatal properties. Gas exchange data were used to parametrize the Farquhar–von Caemmerer photosynthesis model. Leaf mass per area (LMA) was negatively correlated to N. A positive correlation was observed between both day (R_d) and night respiration (R_n) and N when they were expressed on a leaf mass basis, but no correlation was found on a leaf area basis. An R_d/R_n ratio of 0·59 indicated a significant inhibition of dark respiration by light. The maximum net CO₂ assimilation rate at ambient CO₂ concentration (A_max), the maximum rate of potential electron transport (J_max) and the maximum rate of carboxylation (V_cmax) significantly increased with N, particularly when expressed on a mass basis. Although the maximum stomatal conductance to CO₂ (g_scmax) was positively correlated with A_max, there was no relationship between g_scmax and N. Leaf N content influenced the allocation of nitrogen to photosynthetic processes, resulting in a decrease of the J_max/V_cmax ratio with increasing N. It was concluded that leaf nitrogen concentration is a major determinant of photosynthetic capacity in Eucalyptus grandis seedlings and, to a lesser extent, of leaf respiration and nitrogen partitioning among photosynthetic processes, but not of stomatal conductance.     

Season effects on leaf nitrogen partitioning and photosynthetic water use efficiency in mango

The key parameters of photosynthetic capacity (maximum carboxylation rate (V_cmax), electron transport capacity (J_max) and dark respiration rate (R_d)) and the slope (m) of the stomatal conductance model of Ball et al. [Progress in photosynthetic research, Martinus Nijhoff, Dordrecht, 1987] were measured for a whole growing season in fully expanded leaves of 12-year-old mango trees cv. Cogshall in La Réunion island. Leaf nitrogen partitioning into carboxylation (P_c) and bioenergetic (P_b) pools were computed according to the model of Niinemets and Tenhunen [Plant Cell Environ 1997;20: 845–66]. V_cmax, J_max, R_d, P_c and P_b remained relatively stable over the whole study period, with the exception of the period of linear fruit growth when J_max, R_d and P_b were slightly lower, and leaf non-structural carbohydrate content higher. During the pre-floral and floral periods, m decreased by more than 50%, indicating an increase in photosynthetic water use efficiency and m increased again during the period of linear fruit growth. Our results show that, in tropical orchard conditions characterized by mild seasonal climatic changes and non-limiting water supply, leaf nitrogen partitioning is rather stable. Our results also advocate for more studies on the effect of phenology on m and photosynthetic water use efficiency, which is of paramount importance for building coupled biochemical models of photosynthetic carbon assimilation.