Leaf dynamics in growth and reproduction of Xanthium canadense as influenced by stand density.

BACKGROUND AND AIMS: Leaf longevity is controlled by the light gradient in the canopy and also by the nitrogen (N) sink strength in the plant. Stand density may influence leaf dynamics through its effects on light gradient and on plant growth and reproduction. This study tests the hypothesis that the control by the light gradient is manifested more in the vegetative period, whereas the opposite is true when the plant becomes reproductive and develops a strong N sink. METHODS: Stands of Xanthium canadense were established at two densities. Emergence, growth and death of every leaf on the main stem and branches, and plant growth and N uptake were determined from germination to full senescence. Mean residence time and dry mass productivity were calculated per leaf number, leaf area, leaf mass and leaf N (collectively termed 'leaf variables') in order to analyse leaf dynamics and its effect on plant growth. KEY RESULTS: Branching and reproductive activities were higher at low than at high density. Overall there was no significant difference in mean residence time of leaf variables between the two stands. However, early leaf cohorts on the main stem had a longer retention time at low density, whereas later cohorts had a longer retention time at high density. Branch leaves emerged earlier and tended to live longer at low than at high density. Leaf efficiencies, defined as carbon export per unit investment of leaf variables, were higher at low density in all leaf variables except for leaf number. CONCLUSIONS: In the vegetative phase of plant growth, the light gradient strongly controls leaf longevity, whereas later the effects of branching and reproductive activities become stronger and over-rule the effect of light environment. As leaf N supports photosynthesis and also works as an N source for plant development, N use is pivotal in linking leaf dynamics with plant growth and reproduction.

Stem extension and mechanical stability of Xanthium canadense grown in an open or in a dense stand.

BACKGROUND AND AIMS: Plants in open, uncrowded habitats typically have relatively short stems with many branches, whereas plants in crowded habitats grow taller and more slender at the expense of mechanical stability. There seems to be a trade-off between height growth and mechanical stability, and this study addresses how stand density influences stem extension and consequently plant safety margins against mechanical failure. METHODS: Xanthium canadense plants were grown either solitarily (S-plants) or in a dense stand (D-plants) until flowering. Internode dimensions and mechanical properties were measured at the metamer level, and the critical buckling height beyond which the plant elastically buckles under its own weight and the maximum lateral wind force the plant can withstand were calculated. KEY RESULTS: Internodes were longer in D- than S-plants, but basal diameter did not differ significantly. Relative growth rates of internode length and diameter were negatively correlated to the volumetric solid fraction of the internode. Internode dry mass density was higher in S- than D-plants. Young's modulus of elasticity and the breaking stress were higher in lower metamers, and in D- than in S-plants. Within a stand, however, both moduli were positively related to dry mass density. The buckling safety factor, a ratio of critical buckling height to actual height, was higher in S- than in D-plants. D-plants were found to be approaching the limiting value 1. Lateral wind force resistance was higher in S- than in D-plants, and increased with growth in S-plants. CONCLUSIONS: Critical buckling height increased with height growth due mainly to an increase in stem stiffness and diameter and a reduction in crown/stem mass ratio. Lateral wind force resistance was enhanced due to increased tissue strength and diameter. The increase in tissue stiffness and strength with height growth plays a crucial role in maintaining a safety margin against mechanical failure in herbaceous species that lack the capacity for secondary growth.

Mean residence time of leaf number, area, mass, and nitrogen in canopy photosynthesis.

Mean residence time (MRT) of plant nitrogen (N), which is an indicator of the expected length of time N newly taken up is retained before being lost, is an important component in plant nitrogen use. Here we extend the concept MRT to cover such variables as leaf number, leaf area, leaf dry mass, and nitrogen in the canopy. MRT was calculated from leaf duration (i.e., time integral of standing amount) divided by the total production of leaf variables. We determined MRT in a Xanthium canadense stand established with high or low N availability. The MRT of leaf number may imply longevity of leaves in the canopy. We found that the MRT of leaf area and dry mass were shorter than that of leaf number, while the MRT of leaf N was longer. The relatively longer MRT of leaf N was due to N resorption before leaf shedding. The MRT of all variables was longer at low N availability. Leaf productivity is the rate of canopy photosynthesis per unit amount of leaf variables, and multiplication of leaf productivity by MRT gives the leaf photosynthetic efficiency (canopy photosynthesis per unit production of leaf variables). The photosynthetic efficiency of leaf number implies the lifetime carbon gain of a leaf in the canopy. The analysis of plant-level N use efficiency by evaluating the N productivity and MRT is a well-established approach. Extension of these concepts to leaf number, area, mass, and N in the canopy will clarify the underlying logic in the study of leaf life span, leaf area development, and dry mass and N use in canopy photosynthesis.

Growth and nitrogen use in Xanthium canadense grown in an open or in a dense stand.

Plants develop branches profusely when grown solitarily, while less so when grown in a dense stand. Such changes in architecture are associated with changes in dry mass allocation and nitrogen use. Here, we studied what traits in plant growth and nitrogen use were influenced by different light climates in the stand. Annual plants (Xanthium canadense) were grown solitarily or in a dense stand. Dry mass growth was analyzed as the product of the net assimilation rate (NAR) and leaf area (LA). Nitrogen use efficiency (NUE) was analyzed as the product of nitrogen productivity (NP) and the mean residence time (MRT) of nitrogen. These growth variables were further factorized into their components. Solitary plants maintained a high NAR, whereas plants in the dense stand decreased the NAR due to mutual shading. Plants in the dense stand developed a larger LA with a higher specific leaf area than solitary plants. Solitary plants had higher NUE due to higher NP. A temporal increase in NUE was attributed to the increase in MRT of nitrogen. Light climate was different between solitary and dense-stand plants, but they took up a comparable amount of nitrogen and used it differently in response to the given light climate. NUE was thus demonstrated to be a useful tool for analyzing the mechanism leading to different N use in plant growth.

Demand and supply of N in seed production of soybean (Glycine max) at different N fertilization levels after flowering.

Nitrogen (N) has been suggested as a determinant of seed production especially in species with high seed N content. Assuming that seed yield was determined as the balance between N demand and supply for seed production, we studied the effect of N fertilization after flowering on soybean (Glycine max L. Merr.) yield. Seed N concentration was nearly constant irrespective of N fertilization, indicating that seed production was proportional to the amount of N available for seed growth. N demand for seed production was analyzed as the product of seed number, the rate of N filling in individual seeds, and the length of the reproductive period. N fertilization increased seed number and the reproductive period, but did not influence the N filling rate. Seed number was positively correlated with dry mass productivity after flowering. Three N sources were distinguished: mineral N uptake, symbiotic N(2) fixation and N remobilization from vegetative body. N fertilization increased N uptake and N remobilization, but lowered N(2) fixation. We concluded that N availability in the reproductive period determined seed yield directly through increasing N supply for seed growth and indirectly through increasing seed N demand with enhanced plant dry mass productivity.

Leaf-level nitrogen use efficiency: definition and importance.

Nitrogen use efficiency (NUE) has been widely used to study the relationship between nitrogen uptake and dry mass production in the plant. As a subsystem of plant nitrogen use efficiency (NUE), I have defined leaf-level NUE as the surplus production (gross production minus leaf respiration) per unit amount of nitrogen allocated to the leaf, with factorization into leaf nitrogen productivity (NP) and mean residence time of leaf nitrogen (MRT). These concepts were applied to two herbaceous stands: a perennial Solidago altissima stand and an annual Amaranthus patulus stand. S. altissima had more than three times higher leaf NUE than A. patulus due to nearly three times longer MRT of leaf N. In both species, NUE and NP were higher at the leaf level than at the plant level, because most leaf N is involved directly in the photosynthetic activity and because leaf surplus production is higher than the plant net production. MRT was longer at the plant level. The more than twice as long MRT at the plant level as at the leaf level in S. altissima was due to a large contribution of nitrogen storage belowground in the winter in this species. Thus, comparisons between a perennial and an annual system and between plant- and leaf-level NUE with their components revealed the importance of N allocation, storage, recycling, and turnover of organs for leaf photosynthetic production and plant dry mass growth.

Optimal use of leaf nitrogen explains seasonal changes in leaf nitrogen content of an understorey evergreen shrub.

BACKGROUND AND AIMS: Understorey evergreen species commonly have a higher leaf nitrogen content in winter than in summer. Tested here is a hypothesis that such changes in leaf nitrogen content maximize nitrogen-use efficiency, defined as the daily carbon gain per unit nitrogen, under given temperature and irradiance levels. METHODS: The evergreen shrub Aucuba japonica growing naturally at three sites with different irradiance regimes in Japan was studied. Leaf photosynthetic characteristics, Rubisco and leaf nitrogen with measurements of temperature and irradiance monthly at each site were determined. Daily carbon gain was determined as a function of leaf nitrogen content to calculate the optimal leaf nitrogen content that maximized daily nitrogen-use efficiency. KEY RESULTS: As is known, the optimal leaf nitrogen content increased with increasing irradiance. The optimal leaf nitrogen content also increased with decreasing temperature because the photosynthetic capacity per Rubisco decreased. Across sites and months, the optimal leaf nitrogen content was close to the actual leaf nitrogen content and explained the variation in actual leaf nitrogen by 64 %. Sensitivity analysis showed that the effect of temperature on optimal nitrogen content was similar in magnitude to that of irradiance. CONCLUSIONS: Understorey evergreen species regulate leaf nitrogen content so as to maximize nitrogen-use efficiency in daily carbon gain under changing irradiance and temperature conditions.

Nitrogen use efficiency revisited.

Nitrogen use efficiency (NUE) was originally defined as the dry mass productivity per unit N taken up from soil. The term was subsequently redefined as the product of nitrogen productivity (NP) and mean residence time of nitrogen (MRT). However, this redefinition was found to contradict the original definition under certain conditions, and confusion arose when the MRT defined for a steady-state system was applied to a system that was actually not at steady state. As MRT is the expected length of time that a unit of N newly taken up from soil is retained before being lost, it can be translated into the plant nitrogen duration (PND) divided by the total N uptake. This MRT is determined equally well for a steady state- and a non-steady state system and is in accordance with the original definition of NUE. It can be applied to a herbaceous perennial stand (that was at a steady state) and to an annual stand (that was not at a steady state) to determine NUE. NUE is also applicable when plant growth and reproduction are analyzed in relation to N use.

Effects of elevated CO2 concentration on seed production in C3 annual plants.

The response of seed production to CO(2) concentration ([CO(2)]) is known to vary considerably among C(3) annual species. Here we analyse the interspecific variation in CO(2) responses of seed production per plant with particular attention to nitrogen use. Provided that seed production is limited by nitrogen availability, an increase in seed mass per plant results from increase in seed nitrogen per plant and/or from decrease in seed nitrogen concentration ([N]). Meta-analysis reveals that the increase in seed mass per plant under elevated [CO(2)] is mainly due to increase in seed nitrogen per plant rather than seed [N] dilution. Nitrogen-fixing legumes enhanced nitrogen acquisition more than non-nitrogen-fixers, resulting in a large increase in seed mass per plant. In Poaceae, an increase in seed mass per plant was also caused by a decrease in seed [N]. Greater carbon allocation to albumen (endosperm and/or perisperm) than the embryo may account for [N] reduction in grass seeds. These differences in CO(2) response of seed production among functional groups may affect their fitness, leading to changes in species composition in the future high-[CO(2)] ecosystem.

The leaf anatomy of a broad-leaved evergreen allows an increase in leaf nitrogen content in winter.

In temperate regions, evergreen species are exposed to large seasonal changes in air temperature and irradiance. They change photosynthetic characteristics of leaves responding to such environmental changes. Recent studies have suggested that photosynthetic acclimation is strongly constrained by leaf anatomy such as leaf thickness, mesophyll and chloroplast surface facing the intercellular space, and the chloroplast volume. We studied how these parameters of leaf anatomy are related with photosynthetic seasonal acclimation. We evaluated differential effects of winter and summer irradiance on leaf anatomy and photosynthesis. Using a broad-leaved evergreen Aucuba japonica, we performed a transfer experiment in which irradiance regimes were changed at the beginning of autumn and of spring. We found that a vacant space on mesophyll surface in summer enabled chloroplast volume to increase in winter. The leaf nitrogen and Rubisco content were higher in winter than in summer. They were correlated significantly with chloroplast volume and with chloroplast surface area facing the intercellular space. Thus, summer leaves were thicker than needed to accommodate mesophyll surface chloroplasts at this time of year but this allowed for increases in mesophyll surface chloroplasts in the winter. It appears that summer leaf anatomical characteristics help facilitate photosynthetic acclimation to winter conditions. Photosynthetic capacity and photosynthetic nitrogen use efficiency were lower in winter than in summer but it appears that these reductions were partially compensated by higher Rubisco contents and mesophyll surface chloroplast area in winter foliage.

Does leaf photosynthesis adapt to CO2-enriched environments? An experiment on plants originating from three natural CO2 springs.

Atmospheric CO2 elevation may act as a selective agent, which consequently may alter plant traits in the future. We investigated the adaptation to high CO2 using transplant experiments with plants originating from natural CO2 springs and from respective control sites. We tested three hypotheses for adaptation to high-CO2 conditions: a higher photosynthetic nitrogen use efficiency (PNUE); a higher photosynthetic water use efficiency (WUE); and a higher capacity for carbohydrate transport from leaves. Although elevated growth CO2 enhanced both PNUE and WUE, there was no genotypic improvement in PNUE. However, some spring plants had a higher WUE, as a result of a significant reduction in stomatal conductance, and also a lower starch concentration. Higher natural variation (assessed by the coefficient of variation) within populations in WUE and starch concentration, compared with PNUE, might be responsible for the observed population differentiation. These results support the concept that atmospheric CO2 elevation can act as a selective agent on some plant traits in natural plant communities. Reduced stomatal conductance and reduced starch accumulation are highlighted for possible adaptation to high CO2.

Reproductive yield of individuals competing for light in a dense stand of an annual, Xanthium canadense.

In a dense stand, individuals compete with each other for resources, especially for light. Light availability decreases with increasing depth in the canopy, thus light competition becoming stronger with time in the vegetative phase. In the reproductive phase, on the other hand, leaves start senescing, and the light environment, particularly of smaller individuals, will be improved. To study the effect of change in light climate on reproduction of individuals, we established an experimental stand of an annual, Xanthium canadense, and assessed temporal changes in whole plant photosynthesis through the reproductive phase with particular reference to light availability of individuals. At flowering, 83% of individuals were still alive, but only 27% survived to set seeds. Most of the individuals that died in the reproductive phase were smaller than those that produced seeds. Individuals that died at the early stage of the reproductive phase had a lower leaf to stem mass ratio, suggesting that the fate of individuals was determined partly by the pattern of biomass allocation in this period. At the early stage of the reproductive phase, leaf area index (LAI) of the stand was high and larger individuals had higher whole plant photosynthesis than smaller individuals. Although light availability at later stages was improved with reduction in LAI, whole plant photosynthesis was very low in all individuals due to a lower light use efficiency, which was caused by a decrease in photosynthetic N use efficiency. We conclude that light competition was still strong at the early stage of the reproductive phase and that later improvement of light availability did not ameliorate the photosynthesis of smaller individuals.

Does leaf shedding increase the whole-plant carbon gain despite some nitrogen being lost with shedding?

When old leaves are shed, part of the nitrogen in the leaf is retranslocated to new leaves. This retranslocation will increase the whole-plant carbon gain when daily C gain : leaf N ratio (daily photosynthetic N-use efficiency, NUE) in the old leaf, expressed as a fraction of NUE in the new leaf, becomes lower than the fraction of leaf N that is resorbed before shedding (R(N)). We examined whether plants shed their leaves to increase the whole-plant C gain in accord with this criterion in a dense stand of an annual herb, Xanthium canadense, grown under high (HN) and low (LN) nitrogen availability. The NUE of a leaf at shedding expressed as a fraction of NUE in a new leaf was nearly equal to the R(N) in the LN stand, but significantly lower than the R(N) in the HN stand. Thus shedding of old leaves occurred as expected in the LN stand, whereas in the HN stand, shedding occurred later than expected. Sensitivity analyses showed that the decline in NUE of a leaf resulted primarily from a reduction in irradiance in the HN stand. On the other hand, it resulted from a reduction in irradiance and also in light-saturated photosynthesis : leaf N content ratio (potential photosynthetic NUE) in the LN stand.

Nitrogen resorption and protein degradation during leaf senescence in Chenopodium album grown in different light and nitrogen conditions.

The extent of nitrogen (N) resorption and the degradability of different protein pools were examined in senescing leaves of an annual herb, Chenopodium album L., grown in two light and N conditions. Both N resorption efficiency (REFF; the proportion of green-leaf N resorbed) and proficiency (RPROF; the level to which leaf N content is reduced by resorption) varied among different growth conditions. During leaf senescence, the majority of soluble and membrane proteins was degraded in all growth conditions. Structural proteins were also highly degradable, implying that no particular protein pool constitutes a non-retranslocatable N pool in the leaf. Leaf carbon/N ratio affected the timing and duration of senescing processes, but it did not regulate the extent of protein degradation or N resorption. Sink-source relationships for N in the plant exerted a more direct influence, depressing N resorption when N sink strength was weakened in the low-light and high-N condition. N resorption was, however, not enhanced in high-light and low-N plants with the strongest N sinks, possibly because it reached an upper limit at some point. We conclude that a combination of several physiological factors determines the extent of N resorption in different growth conditions.

Leaf lifespan and lifetime carbon balance of individual leaves in a stand of an annual herb, Xanthium canadense.

Leaf lifespan in response to resource availability has been documented in many studies, but it still remains uncertain what determines the timing of leaf shedding. Here, we evaluate the lifetime carbon (C) balance of a leaf in a canopy as influenced by nitrogen (N) availability. Stands of Xanthium canadense were established with high-nitrogen (HN) and low-nitrogen (LN) treatments and temporal changes of C gain of individual leaves were investigated with a canopy photosynthesis model. Daily C gain of a leaf was maximal early in its development and subsequently declined. Daily C gain at shedding was nearly zero in HN, while it was still positive in LN. Sensitivity analyses showed that the decline in the daily C gain resulted primarily from the reduction in light level in HN and by the reduction in leaf N in LN. Smaller leaf size in LN than in HN led to higher light levels in the canopy, which helped leaves of the LN stand maintain for a longer period. These results suggest that the mechanism by which leaf lifespan is determined changes depending on the availability of the resource that is most limiting to plant growth.

Seasonal changes in photosynthesis, nitrogen content and nitrogen partitioning in Lindera umbellata leaves grown in high or low irradiance.

Seasonal changes in photosynthetic capacity, leaf nitrogen (N) content and N partitioning were studied from before leaf maturation (spring) until death (autumn) in high- and low-light-exposed leaves of a deciduous shrub, Lindera umbellata var. membranacea (Maxim.) Momiyama growing in a natural forest in northeast Japan. In spring, light-saturated photosynthetic rate (Pmax) was low despite high leaf N and Rubisco contents, indicating that the photosynthetic apparatus was not yet functionally developed. Rubisco seemed to be only partially active. In summer and autumn, Pmax per unit leaf N increased and changes in Pmax were correlated with changes in leaf N and two photosynthetic components, Rubisco and chlorophyll. Changes in these components paralleled the changes in leaf N. During leaf senescence, about 70% of leaf N was resorbed. Metabolic proteins that accounted for the majority of leaf N in summer were highly degradable and more than sufficient to explain the high N-resorption efficiency. Structural proteins represented only a small part of leaf N and were relatively resistant to degradation and thus contributed little to N resorption. Leaf N partitioning between metabolic and structural proteins determined the amount of retranslocatable N, but did not strictly determine the N content of a dead leaf or N-resorption efficiency.

Seasonal changes in temperature dependence of photosynthetic rate in rice under a free-air CO(2) enrichment.

BACKGROUND AND AIMS: Influences of rising global CO(2) concentration and temperature on plant growth and ecosystem function have become major concerns, but how photosynthesis changes with CO(2) and temperature in the field is poorly understood. Therefore, studies were made of the effect of elevated CO(2) on temperature dependence of photosynthetic rates in rice (Oryza sativa) grown in a paddy field, in relation to seasons in two years. METHODS: Photosynthetic rates were determined monthly for rice grown under free-air CO(2) enrichment (FACE) compared to the normal atmosphere (570 vs 370 micromol mol(-1)). Temperature dependence of the maximum rate of RuBP (ribulose-1,5-bisphosphate) carboxylation (V(cmax)) and the maximum rate of electron transport (J(max)) were analysed with the Arrhenius equation. The photosynthesis-temperature response was reconstructed to determine the optimal temperature (T(opt)) that maximizes the photosynthetic rate. KEY RESULTS AND CONCLUSIONS: There was both an increase in the absolute value of the light-saturated photosynthetic rate at growth CO(2) (P(growth)) and an increase in T(opt) for P(growth) caused by elevated CO(2) in FACE conditions. Seasonal decrease in P(growth) was associated with a decrease in nitrogen content per unit leaf area (N(area)) and thus in the maximum rate of electron transport (J(max)) and the maximum rate of RuBP carboxylation (V(cmax)). At ambient CO(2), T(opt) increased with increasing growth temperature due mainly to increasing activation energy of V(cmax). At elevated CO(2), T(opt) did not show a clear seasonal trend. Temperature dependence of photosynthesis was changed by seasonal climate and plant nitrogen status, which differed between ambient and elevated CO(2).

Respiration and reproductive effort in Xanthium canadense.

BACKGROUND AND AIMS: The proportion of resources devoted to reproduction in the plant is called the reproductive effort (RE), which is most commonly expressed as the proportion of reproductive biomass to total plant biomass production (RE(W)). Reproductive yield is the outcome of photosynthates allocated to reproductive structures minus subsequent respiratory consumption for construction and maintenance of reproductive structures. Thus, RE(W) can differ from RE in terms of photosynthates allocated to reproductive structures (RE(P)). * METHODS: Dry mass growth and respiration of vegetative and reproductive organs were measured in Xanthium canadense and the amount of photosynthates and its partitioning to dry mass growth and respiratory consumption were determined. Differences between RE(W) and RE(P) were analysed in terms of growth and maintenance respiration. * KEY RESULTS: The fraction of allocated photosynthates that was consumed by respiration was smaller in the reproductive organ than in the vegetative organs. Consequently, RE(P) was smaller than RE(W). The smaller respiratory consumption in the reproductive organ resulted from its shorter period of existence and a seasonal decline in temperature, as well as a slower rate of maintenance respiration, although the fraction of photosynthates consumed by growth respiration was larger than in the vegetative organs. * CONCLUSIONS: Reproductive effort in terms of photosynthates (RE(P)) was smaller than that in terms of biomass (RE(W)). This difference resulted from respiratory consumption for maintenance, which was far smaller in the reproductive organ than in vegetative organs.

Biomass allocation and leaf chemical defence in defoliated seedlings of Quercus serrata with respect to carbon-nitrogen balance.

BACKGROUND AND AIMS: Both nutrient availability and defoliation affect the carbon-nutrient balance in plants, which in turn influences biomass allocation (e.g. shoot-to-root ratio) and leaf chemical composition (concentration of nitrogen and secondary compounds). In this study it is questioned whether defoliation alters biomass allocation and chemical defence in a similar fashion to the response to nutrient deficiency. METHODS: Current-year seedlings of Quercus serrata were grown with or without removal of all leaves at three levels of nutrient availability. KEY RESULTS: Plant nitrogen concentration (PNC), a measure of the carbon-nutrient balance in the plant, significantly decreased immediately after defoliation because leaves had higher nitrogen concentrations than stems and roots. However, PNC recovered to levels similar to or higher than that of control plants in 3 or 6 weeks after the defoliation. Nitrogen concentration of leaves produced after defoliation was significantly higher than leaf nitrogen concentration of control leaves. Leaf mass per plant mass (leaf mass ratio, LMR) was positively correlated with PNC but the relationship was significantly different between defoliated and control plants. When compared at the same PNC, defoliated plants had a lower LMR. However, the ratio of the leaf to root tissues that were newly produced after defoliation as a function of PNC did not differ between defoliated and control plants. Defoliated plants had a significantly lower concentration of total phenolics and condensed tannins. Across defoliated and control plants, the leaf tannin concentration was negatively correlated with the leaf nitrogen concentration, suggesting that the amount of carbon-based defensive compounds was controlled by the carbon-nutrient balance at the leaf level. CONCLUSIONS: Defoliation alters biomass allocation and chemical defence through the carbon-nutrient balance at the plant and at the leaf level, respectively.

Seasonal changes in light and temperature affect the balance between light harvesting and light utilisation components of photosynthesis in an evergreen understory shrub.

In a temperate climate, evergreen species in the understory are exposed to large changes in photosynthetic photon flux density (PPFD) and temperature over the year. We determined the photosynthetic traits of leaves of an evergreen understory shrub Aucuba japonica at three sites at monthly intervals: understorys of a deciduous forest; an evergreen forest; and a gap in a mixed forest. This set up enabled us to separate the effects of seasonal change in PPFD and temperature on photosynthetic acclimation under natural conditions. The effects of PPFD and temperature were analysed by simple and multiple regression analyses. The amounts of light utilisation components (LU), represented by nitrogen and rubisco contents per area, were higher in winter, when temperature was low and PPFD was high. The LU relative to the amount of light harvesting components (LH), represented by chlorophyll a/b and rubisco/chlorophyll ratios, and the inverse of chlorophyll/nitrogen ratio were also higher in winter. We quantified the effects of PPFD and temperature on the LU and LH components. Across sites PPFD had stronger effects than air temperature, while within a site temperature had stronger effects on photosynthetic acclimation. We concluded that the photosynthetic apparatus is strongly affected by the prevailing PPFD at the time of leaf development. Within a given light regime, however, plants acclimated by increasing LU relative to LH primarily in response to temperature and to a lesser extent to PPFD.