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.

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.

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.

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.

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.

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.

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.

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.

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 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.

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.

Leaf discs floated on water are different from intact leaves in photosynthesis and photoinhibition.

Photoinhibition has been often evaluated with leaf discs floated on water or placed on wet papers to prevent desiccation. Under these conditions, there is a possibility that CO(2) diffusion is blocked by water, which may lead to reduction in photosynthetic CO(2) assimilation. Using Chenopodium album L. grown at two irradiances, photosynthesis, quantum yield of Photosystem II (DeltaF/F (m)'), non-photochemical quenching (qN), and photoinhibition were compared between detached leaves and leaf discs. In low-light-grown plants, photoinhibition was greater in leaf discs than in detached leaves, while in high-light-grown plants, there was little difference. Leaf discs showed lower rates of photosynthesis and DeltaF/F (m)', and higher qN. The DeltaF/F (m)' in leaf discs increased when leaf discs were exposed to high concentration of CO(2), suggesting that CO(2) diffusion to chloroplasts was limited in leaf discs floated on water.

Nitrogen uptake and use by competing individuals in a Xanthium canadense stand.

We studied differences in nitrogen uptake and use for plant growth among individuals competing in a natural dense stand of an annual herb, Xanthium canadense. Larger individuals took up more nitrogen than proportionately to their size, indicating that the competition for soil nitrogen was asymmetric among individuals, although it was more symmetric than the competition for light. The rate of nitrogen loss of individuals also increased with plant size. While smaller individuals shared smaller fractions of total plant nitrogen in the stand, they had higher nitrogen concentrations per unit mass. "Turnover" rates of nitrogen influx (rin) and outflux (rout) were defined as the rates of nitrogen uptake and loss per unit aboveground nitrogen, respectively. rin was higher in larger individuals, whereas rout was higher in smaller individuals. Consequently, the relative rate of nitrogen increment (rin-rout) was higher in larger individuals, whereas it was around zero in the smallest individuals. The mean residence time of nitrogen (MRT), defined as the inverse of rout, was longer in larger individuals. Nitrogen productivity (NP), i.e. the growth rate per unit aboveground nitrogen, was higher in larger individuals. As the product of lifetime MRT and NP gives the nitrogen use efficiency (NUE), defined as biomass production per unit flux of nitrogen, higher MRT and NP observed in larger individuals would have contributed to their higher lifetime NUE. Shorter MRT in smaller individuals was caused by the abscission of leaves which contained relatively large fractions of total plant nitrogen. Xanthium canadense, as a competitive ruderal, tended to produce leaves at higher positions to acquire higher light levels at the expense of older leaves rather than to modify their productive structure to efficiently use low light levels as observed in shade-tolerant species.

Light acquisition and use by individuals competing in a dense stand of an annual herb, Xanthium canadense.

The importance of light acquisition and utilization by individuals in intraspecific competition was evaluated by determining growth and photosynthesis of individual plants in a dense monospecific stand of an annual, Xanthium canadense. Photosynthesis of individual plants in the stand was calculated using a canopy photosynthesis model in which leaf photosynthesis was assumed to be function of leaf nitrogen content and light availability. The estimated photosynthetic rates of individuals were strongly correlated with the measured growth rates. Photosynthetic rates per unit aboveground mass (RPR, relative photosynthetic rate) increased with increasing aboveground mass, suggesting asymmetric (one-sided) competition in the stand. However, larger individuals had similar RPRs, suggesting symmetric (two-sided) competition. These results were consistent with the observation that size inequality over the whole stand increased with growth, but it remained stable among the larger individuals. The RPR of an individual was calculated as the product of absorbed photon flux per unit aboveground mass (Φmass) and light use efficiency (LUE, photosynthesis per unit absorbed photon flux). Φmass indicates the efficiency of light acquisition, and was higher in larger individuals in the stand, while LUE was highest in individuals with intermediate aboveground mass. LUE depends on leaf nitrogen content. At an early stage, leaf nitrogen contents of smaller individuals were similar to those that maximize LUE. Light availability to smaller individuals decreased as they grew, while their nitrogen contents did not change markedly, which decreased their LUE. We concluded that asymmetric competition among individuals in the stand resulted mainly from lower efficiencies in both light acquisition and light use by smaller individuals.

Limitations on photosynthesis of competing individuals in stands and the consequences for canopy structure.

The canopy structure of a stand of vegetation is determined by the growth patterns of the individual plants within the stand and the competitive interactions among them. We analyzed the carbon gain of individuals in two dense monospecific stands of Xanthium canadense and evaluated the consequences for intra-specific competition and whole-stand canopy structure. The stands differed in productivity, and this was associated with differences in nitrogen availability. Canopy structure, aboveground mass, and nitrogen contents per unit leaf area (N area) were determined for individuals, and leaf photosynthesis was measured as a function of N area. These data were used to calculate the daily carbon gain of individuals. Within stands, photosynthesis per unit aboveground mass (P mass) of individual plants increased with plant height, despite the lower leaf area ratios of taller plants. The differences in P mass between the tallest most dominant and shortest most subordinate plants were greater in the high-nitrogen than in the low-nitrogen stand. This indicated that competition was asymmetric and that this asymmetry increased with nitrogen availability. In the high-nitrogen stand, taller plants had a higher P mass than shorter ones, because they captured more light per unit mass and because they had higher photosynthesis per unit of absorbed light. Conversely, in the low-nitrogen stand, the differences in P mass between plants of different heights resulted only from differences in their light capture per unit mass. Sensitivity analyses revealed that an increase in N area, keeping leaf area of plants constant, increased whole-plant carbon gain for the taller more dominant plants but reduced carbon gain in the shorter more subordinate ones, which implies that the N area values of shorter plants were greater than the optimal values for maximum photosynthesis. On the other hand, the carbon gain of all individual plants, keeping their total canopy N constant, was positively related to an increase in their individual leaf area. At the same time, however, increasing the leaf area for all plants simultaneously reduced the carbon gain of the whole stand. This result shows that the optimal leaf area index (LAI), which maximizes photosynthesis of a stand, is not evolutionarily stable because at this LAI, any individual can increase its carbon gain by increasing its leaf area.

Effects of CO2 elevation on canopy development in the stands of two co-occurring annuals.

Elevated CO2 may increase dry mass production of canopies directly through increasing net assimilation rate of leaves and also indirectly through increasing leaf area index (LAI). We studied the effects of CO2 elevation on canopy productivity and development in monospecific and mixed (1:1) stands of two co-occurring C3 annual species, Abutilon theophrasti, and Ambrosia artemisiifolia. The stands were established in the glasshouse with two CO2 levels (360 and 700 µl/l) under natural light conditions. The planting density was 100 per m2 and LAI increased up to 2.6 in 53 days of growth. Root competition was excluded by growing each plant in an individual pot. However, interference was apparent in the amount of photons absorbed by the plants and in photon absorption per unit leaf area. Greater photon absorption by Abutilon in the mixed stand was due to different canopy structures: Abutilon distributed leaves in the upper layers in the canopy while Ambrosia distributed leaves more to the lower layers. CO2 elevation did not affect the relative performance and light interception of the two species in mixed stands. Total aboveground dry mass was significantly increased with CO2 elevation, while no significant effects on leaf area development were observed. CO2 elevation increased dry mass production by 30-50%, which was mediated by 35-38% increase in the net assimilation rate (NAR) and 37-60% increase in the nitrogen use efficiency (NUE, net assimilation rate per unit leaf nitrogen). Since there was a strong overall correlation between LAI and aboveground nitrogen and no significant difference was found in the regression of LAI against aboveground nitrogen between the two CO2 levels, we hypothesized that leaf area development was controlled by the amount of nitrogen taken up from the soil. This hypothesis suggests that the increased LAI with CO2 elevation observed by several authors might be due to increased uptake of nitrogen with increased root growth.

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.

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.