Leaf nitrogen distribution and whole canopy photosynthetic carbon gain in herbaceous stands

The amount of photosynthetically-active photon flux density incident upon a leaf and the nitrogen content of that leaf strongly affect the photosynthetic carbon gain of that leaf. Therefore, the canopy structure of a stand, affecting the light climate in the canopy, and the leaf nitrogen distribution pattern in the canopy, affect the carbon gain of the whole canopy. This review discusses the results of studies directed to this problem and obtained so far in open and in dense canopies of stands of herbaceous, monocotyledonous or dicotyledonous, plants in their growing or flowering stages. It is found that the leaf nitrogen distribution pattern in the canopy is vertically non-uniform, and in dense stands more strongly so than in open stands. The leaf nitrogen distribution pattern in most canopies closely approaches an optimal pattern in that it maximizes whole canopy potential carbon gain as calculated for the actual total leaf nitrogen content and leaf area index of the stand. The resulting increase in potential carbon gain as compared to a uniform leaf nitrogen distribution pattern is considerable and it is larger in dense stands than in open stands. For at least some dense stands simulation studies show that with the available total leaf nitrogen content, whole canopy carbon gains could still be considerable higher had a lower leaf area index been developed.     

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.     

Functional correlates of leaf demographic response to gap release in saplings of a shade-tolerant tree,<Emphasis Type="Italic"> Elateriospermum tapos</Emphasis>

For a shade-tolerant SE Asian tropical tree, Elateriospermum tapos (Euphorbiaceae), we studied field-established saplings in gaps and the shaded understory to test the hypothesis that differences in leaf demography and leaf life span under contrasting light regimes should be functionally correlated with architecture, self-shading and nitrogen distribution within the sapling crown. Rates of leaf production and net leaf gain were greater for saplings in gaps than those in the understory. Median leaf life span was approximately 26 months in the gap saplings, while it was estimated to be greater than 38 months in the understory saplings. Consequently, gap saplings had a greater standing leaf number and experienced greater degrees of self-shading than understory saplings. Light availability at individual leaves, estimated by a combination of canopy photos and a three-dimensional architecture model, were negatively correlated with leaf age in gap saplings but not so in understory saplings. Leaf nitrogen content per unit area (N_area) was influenced more by light availability than by leaf age in the gap saplings. In contrast, in understory saplings, N_area was neither correlated with light availability nor with leaf age, and did not decrease significantly before 38 months in leaf age. We conclude that saplings of this shade-tolerant species apparently prolong their leaf life span in the shaded understory through slower rates of leaf production, lower standing number of leaves and lower degrees of self shading than in gap, and that the rate of decline of N_area with leaf age depends on architecture and self-shading regimes that respond to changes in light regimes.     

Canopy structure and leaf nitrogen distribution in a stand of Lysimachia vulgaris L. as influenced by stand density

Summary A hypothesis that a dense stand should develop a less uniform distribution of leaf nitrogen through the canopy than an open stand to increase total canopy photosynthesis was tested with experimentally established stands of Lysimachia vulgaris L. The effect of stand density on spatial variation of photon flux density, leaf nitrogen and specific leaf weight within the canopy was examined. Stand density had little effect on the value of the light extinction coefficient, but strongly affected the distribution of leaf nitrogen per unit area within a canopy. The open stand had more uniform distribution of leaf nitrogen than the dense stand. However, different light climates between stands explained only part of the variation of leaf nitrogen in the canopy. The specific leaf weight in the canopy increased with increasing relative photon flux density and with decreasing nitrogen concentration.     

Evolutionarily stable leaf area production in plant populations

Using an analytical model, it was shown that for a given amount of nitrogen in the canopy of a stand (N(T)), there exists an evolutionarily stable leaf area index (ES-LAI), and therefore an evolutionarily stable average leaf nitrogen content (n(ES)(av);n(ES)(av) =N(T)/ES-LAI), at which no individual plant in the stand can increase its photosynthesis by changing its leaf area. It was also shown that this ES-LAI is always greater than the optimal LAI that maximizes photosynthesis per unit N(T) of the stand. This illustrates that the canopy structure that maximizes photosynthesis of a population is not the same as the canopy structure that maximizes photosynthesis of individuals within a population. It was further derived that the ES-LAI at given N(T) increases with the ratio between the light-saturated photosynthesis and the N content per unit leaf area (leaf-PPNUE) and that it decreases with the canopy extinction coefficient for light (K(L)), the light availability and the apparent quantum yield (phi). These hypotheses were tested by comparing calculated ES-LAI and n(ES)(av) values to actual LAIs and leaf N contents measured for stands of a large variety of herbaceous plants. There was a close correspondence between the calculated and measured values. As predicted by the model, plants with high leaf-PPNUEs produced more leaf area per unit nitrogen than those with low leaf-PPNUEs while plants with horizontal leaves, forming stands with higher K(L) values, produced less leaf area than those with more vertically inclined leaves. These results suggest that maximization of individual plant photosynthesis per unit of nitrogen plays an important role in determining leaf area production of plants and the resulting canopy structure of stands of vegetation. They further suggest this optimization to be a mechanism by which leaf traits such as leaf-PPNUE and leaf inclination angle are causally related to structural characteristics of the population, i.e. the leaf area index of the stand.     

Nitrogen Deficiency in Small Closed Communities of S24 Ryegrass. I. Photosynthesis, Respiration, Dry Matter Production and Partition

Small communities of S24 ryegrass were grown under supplementarylights in a glasshouse at 20°C, and abundantly suppliedwith a complete nutrient solution containing 300 p.p.m. of nitrogen,until they had a leaf area index of 5 and fully interceptedthe light. Half were then given a solution containing only 3p.p.m. of nitrogen (LN) while the rest were kept at 300 p.p.m.(HN). The LN plants had a rate of single leaf photosynthesis lowerthan that of the HN plants at all but the lowest light intensities(33 per cent lower at the saturating irradiance of 170 W m^–2).Similarly, the LN communities had rates of canopy gross photosynthesis(P_sc) markedly lower than those of the HN communities. A comparisonof the observed rates of P_sc with those predicted by a mathematicalmodel of canopy photosynthesis indicated that it was the effectof nitrogen on single leaf photosynthesis, rather than differencesbetween the communities in leaf area, which led to the observeddifferences in P_sc. The superiority of the HN communities in terms of P_sc was partlyoffset by a higher rate of respiration so that they only exceededthe LN communities in terms of canopy net photosynthesis atirradiances in excess of 180 W m_–2, and produced only15 per cent more total dry matter. Nevertheless, the HN plantsdirected less of that dry matter into root and more into topsso that they came to possess twice the weight of live laminae,and the HN communities twice the leaf area, of their nitrogendeficient counterparts. Lolium perenne, S24 ryegrass, photosynthesis, respiration, dry matter production and partition, nitrogen dekieacy     

Branching, nitrogen distribution, and carbon gain in solitary plants of an annual herb, Xanthium canadense

The theory of optimal nitrogen (N) distribution predicts that the carbon gain of plants will be maximised when leaves of higher irradiance have higher N content per area (N _area). Most previous studies have examined optimal N distribution without explicitly considering the branching status of plants. I investigated light environment, N distribution and photosynthetic traits of individual leaves of an herbaceous species, Xanthium canadense. X. canadense was grown solitary under high (HN) and low nutrients (LN). Light availability, leaf mass per unit area and N _area were measured for all leaves within plants. Daily photosynthesis of the plants of actual N distribution was compared with those of optimal and constant N distribution. Branch production was facilitated in HN but not in LN plants. N _area was correlated more with leaf order than with leaf light environment. Although N was more limited and the light environment was less heterogeneous within crowns in LN than in HN plants, leaf N distribution was closer to optimal in the latter. These results suggest that leaf N distribution was not optimised in solitary plants of X. canadense. Because this species often regenerates in a dense stand, leaf N distribution might be selected to maximise carbon gain only in such a stand. Leaf N distribution might thus be constrained by the regeneration strategy of the species.     

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.     

Canopy structure,nitrogen distribution and whole canopy photosynthetic carbon gain in growing and flowering stands of tall herbs

Using a combination of mathematical modeling and field studies we showed that in dense stands of growing herbaceous plants the vertical pattern of leaf nitrogen distribution resembles the pattern of mean light attenuation in the stand and hence tends to maximize total daily photosynthetic carbon gain of the whole stand. Flowering represents a strong sink of nitrogen away from the photosynthetic apparatus and in herbs like Solidago altissima it induces leaf shedding. We studied both the effect of nitrogen reallocation and leaf shedding on the whole canopy photosynthesis and changes in leaf nitrogen distributions in stands moving from the growing to the flowering stage. Despite a decrease in leaf area index and total nitrogen available for photosynthesis in the flowering stand, the leaf nitrogen distribution here also leads to an almost maximum canopy photosynthesis. In both the growing and the flowering stands the leaf area index was higher than calculated optimum values. It is pointed out that this should not necessarily be interpreted as non-adaptive.     

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.     

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.     

The vertical distribution of nitrogen and photosynthetic activity at different plant densities in Carex acutiformis

Shoots of the monocotyledonous perennial Carex acutiformis were grown in open (28 shoots m⁻²) and dense stands (280 shoots m⁻²). For fully grown stands the distribution of relative PPFD and leaf nitrogen per unit leaf area over canopy depth was determined. Light response of photosynthesis was measured on leaf segments sampled at various heights in the stands. Relations between parameters of these curves and leaf nitrogen were investigated. Simulations showed that in the open stand daily canopy photosynthesis was not affected by nitrogen redistribution in the canopy. For the dense stand however, a uniform nitrogen distribution would lead to only 73% of the maximum net carbon gain by the stand under optimal nitrogen distribution. The actual canopy photosynthesis was only 7% less than this theoretical maximum; the actual nitrogen distribution of the dense stand clearly tended to the optimal distribution. The vertical pattern of the nitrogen distribution was to a large extent determined by the minimum leaf nitrogen content. The relatively high minimum leaf nitrogen content found for Carex leaves may perhaps be necessary to maintain the physiological function of the basal parts of the leaves.     

Evaluation of secondary dispersal in a large-seeded tree Aesculus turbinata: a test of directed dispersal

Gap characteristics and gap phase replacement of major tree species were examined in two primary old-growth (mean DBHs of the canopy trees were 45.2 and 56.1 cm) and four secondary developing (range of mean DBH of the canopy trees was 23.5–39.9 cm) beech (Fagus crenata) stands in the Daisen Forest Reserve, southwestern Japan, and these were analyzed in relation to stand development as expressed by the difference of mean DBH of canopy trees. Percentage gap area (percentage of total gap area to total surveyed area) and mean and maximum gap size varied widely and ranged from 1.7 to 20.0%, from 19.4 to 162.8 m2, and from 35.7 to 585.1 m2, respectively. Mean percentage gap area and mean gap size were significantly greater in old-growth than in developing stands. However, they and maximum gap size might not increase linearly with stand development, and the gap feature of less developed stands was greater than that of later stages in developing stands. The cause was a higher formation rate, in younger developing stands, of gaps formed by simultaneous death (multiple trees falling down in domino fashion) which tends to produce larger gaps. In developing stands mean DBH of gapmakers (canopy trees that formed a gap) was smaller than that of canopy trees, though the inverse trend might be found in old-growth stands. Three typical types of death or injury states of gapmakers such as standing dead, trunk broken and uprooted were found in every study stand and the difference in stand development may not cause stand-to-stand variations for them. Importance of F. crenata (the most dominant species) in the canopy layer increased and its importance in the understory layer decreased with stand development. Shade-intolerant Quercus mongolica in the canopy layer was more important in younger than in old-growth stands, and there was no Quercus regeneration in old-growth stands. Acer mono consistently appeared, though in much less abundance than other species, in both canopy and understory layers of all study stands. Sub-canopy layers, which are mainly formed by sub-canopy tree species such as Acanthopanax sciadophylloides and Acer japonicum, may gradually develop with stand development.p>     

Leaf dynamics,self-shading and carbon gain in seedlings of a tropical pioneer tree

We examined leaf dynamics and leaf age gradients of photosynthetic capacity and nitrogen concentration in seedlings of the tropical pioneer tree, Heliocarpus appendiculatus, grown in a factorial design under controlled conditions with two levels each of nutrients, ambient light (light levels incident above the canopy), and self-shading (the gradient of light levels from upper to lower leaves on the shoot). Correlations among these parameters were examined in order to determine the influence of self-shading, and the regulation of standing leaf numbers, on leaf longevity and its association with leaf photosynthetic capacity. Leaf longevity and the number of leaves on the main shoot were both reduced in high light, while in the low light environment, they were reduced in the steeper self-shading gradient. In high nutrients, leaf longevity was reduced whereas leaf number increased. Leaf initiation rates were higher in the high nutrient treatment but were not influenced by either light treatment. Maximum-light saturated photosynthetic rate, on an area basis, was greater in the high light and nutrient treatments, while the decline in photosynthetic capacity in realtion to leaf position on the shoot was more rapid in high light and in low nutrients. Leaf longevity was negatively correlated among treatments with initial photosynthetic capacity. The leaf position at which photosynthetic capacity was predicted to reach zero was positively correlated with the number of leaves on the shoot, supporting the hypothesis that leaf numbers are regulated by patterns of self-shading. The negative association of longevity and initial photosynthetic capacity apparently arises from different associations among gradients of photosynthetic capacity, leaf numbers and leaf initiation rates in relation to light and nutrient availability. The simultaneous consideration of age and position of leaves illuminates the role of self-shading as an important factor influencing leaf senescence and canopy structure and dynamics.     

南京地区落叶栎林主要木本植物的展叶动态研究

 植物的展叶过程是由自身遗传因子决定的,同时又受到多种生态因子的调节,反映了植物的生活史对策和群落物种多样性的维持机制。在2001和2002年的3～6月间,不定期记录了南京地区三个落叶栎(Quercus spp.)林中主要木本植物的展叶情况,包括被标记标准枝的叶数、叶的长度、宽度、叶面积、叶干重等参数。结果表明在所调查的落叶栎林中,林冠层物种的成熟叶面积和单位叶面积干重都显著大于林下层物种;最早展叶的物种为林下层物种,但林冠层展叶顺序与林下层无显著差异。叶面积越大、单位叶面积干重越小的物种展叶越晚;林冠层物种展叶较林下层快,物种成熟叶面积越大,展叶速率越大。最后对展叶顺序和展叶速度的生态学意义作了讨论。     

Nitrogen distribution and leaf area indices in relation to photosynthetic nitrogen use efficiency in savanna grasses

Leaf photosynthetic characteristics, distribution patterns of nitrogen content per unit leaf area (n_L) and leaf area production per unit n_Lwere measured in natural stands of a C₄ grass (Hyparrhenia rufa) from the seasonal savannas and of a C₄grass (Paspalum fasciculatum) and two C₃grasses (Leersia hexandra and Hymenachne amplexicaulis) from the flooded savannas in central Venezuela. Daily rates of canopy photosynthesis (P_cD) as well as the optimal leaf area production per unit n_Lat which P_cDfor a given total amount of nitrogen in the canopy (i.e., canopy-PNUE) is maximized were also calculated. The C₃and C₄species from the flooded savannas had similar light saturated rates of photosynthesis per unit n_L(i.e. leaf-PNUE) and similar canopy-PNUEs which was in strong contrast with previous studies. Especially H. rufa but also L. hexandra and H. amplexicaulis had leaf- and canopy-PNUEs which were considerably higher than the values calculated for most other species with the same photosynthetic pathway (i.e., C₃or C₄). In contrast to previous studies, differences in the light gradient in the canopy between stands only partially explained differences in N distribution. Measured leaf area indices were greater and the average n_L values were consequently smaller than the calculated optima. There was, however, a very strong linear correlation between the optimal and actual average n_Lindicating that even though the model overestimated average n_L, it did predict the differences in leaf area production per unit nitrogen – the inverse average n_L– very well. This result strongly indicates that leaf area production per unit of leaf nitrogen increases with leaf-PNUE and decreases with the extinction coefficient for light. Grass species from seasonal savannas have extremely high leaf-PNUEs and thus optimally produce large amounts of leaf area per unit n_L. This helps explain how stands of these species may have high leaf area indices and achieve high photosynthetic productivity despite the very low nutrient availability at which they grow.     

大气CO2浓度和温度升高对水稻叶片及群体光合作用的影响

大气ＣＯ２浓度升高对植物光合作用的影响研究多集中在单叶水平,在高ＣＯ２及高温下对植物单叶及群体光合进行比较的研究少有报道,而群体水平的研究则是预测生态系统反应所不可缺少的。采用田间开顶式培养室研究了大气ＣＯ２浓度和温度升高对水稻（ＯｒｙｚａｓａｔｉｖａＬ．）叶片及群体光合作用的影响。发现ＣＯ２浓度和温度对水稻叶片光合作用有协同促进作用,而对群体光合作用的促进则随时间的推移而减弱;单叶光合受到的促进作用大于群体光合;叶面积指数只在营养生长期受到促进,冠层叶片含氮量受ＣＯ２影响降低。群体呼吸（包括茎杆）增加及冠层叶片早衰可能是后期ＣＯ２对群体光合促进作用下降的原因。     

Characterization of the canopy structure of Dutch grasslands

In order to characterize the canopy structure of different grassland types 50 stands, representing 15 syntaxonomically distinct types, were examined by harvesting the standing crop during the main flowering period. The types differ in water and nutrient conditions and vary largely in aboveground phytomass (0.5–23 t·ha^?1) and Leaf Area Index (0.4–21: bifacial). Living aboveground phytomass and canopy structure are almost entirely determined by phanerogams. The graminoids generally dominate over the forbs. Variation in aboveground phytomass is related to foliage characteristics such as Aboveground Leaf Area Ratio, Specific Foliage Weight and Leaf Area Development. A PCA with these canopy variables and the variables canopy height, phytomass density and the phytomass ratios stem-leaf-inflorescence shows a clear arrangement related to aboveground phytomass and LAI. The grassland stands can be divided into four groups of different productivity levels for which various combinations of canopy variables are characteristic. Leaf size and leaf inclination are also used for characterization of the different grasslands. Ordinations with these variables by means of PCA and CCA resemble partly to those computed with the canopy variables of aboveground phytomass and LAI. Small leaf sizes are characteristic for low productive grasslands, while the largest leaves occur in high productive grasslands, although they mostly do not belong to the species contributing most strongly to the phytomass of the stand. The leaf inclinations erect and erecto-patent are most common in each grassland. Horizontal leaf areas occur less frequent, but they are relatively well presented in some high productive grasslands. Ranges in leaf size vary more than ranges in leaf inclination do for this series of grasslands. However, leaf size and leaf inclination are useful variables for characterization of grassland canopies in a hierarchical way, where phytomass and leaf area are the first criteria for such a characterization.     

Plant responses to elevated CO<Subscript>2</Subscript> concentration at different scales: leaf,whole plant,canopy, and population

Elevated CO₂ enhances photosynthesis and growth of plants, but the enhancement is strongly influenced by the availability of nitrogen. In this article, we summarise our studies on plant responses to elevated CO₂. The photosynthetic capacity of leaves depends not only on leaf nitrogen content but also on nitrogen partitioning within a leaf. In Polygonum cuspidatum, nitrogen partitioning among the photosynthetic components was not influenced by elevated CO₂ but changed between seasons. Since the alteration in nitrogen partitioning resulted in different CO₂-dependence of photosynthetic rates, enhancement of photosynthesis by elevated CO₂ was greater in autumn than in summer. Leaf mass per unit area (LMA) increases in plants grown at elevated CO₂. This increase was considered to have resulted from the accumulation of carbohydrates not used for plant growth. With a sensitive analysis of a growth model, however, we suggested that the increase in LMA is advantageous for growth at elevated CO₂ by compensating for the reduction in leaf nitrogen concentration per unit mass. Enhancement of reproductive yield by elevated CO₂ is often smaller than that expected from vegetative growth. In Xanthium canadense, elevated CO₂ did not increase seed production, though the vegetative growth increased by 53%. As nitrogen concentration of seeds remained constant at different CO₂ levels, we suggest that the availability of nitrogen limited seed production at elevated CO₂ levels. We found that leaf area development of plant canopy was strongly constrained by the availability of nitrogen rather than by CO₂. In a rice field cultivated at free-air CO₂ enrichment, the leaf area index (LAI) increased with an increase in nitrogen availability but did not change with CO₂ elevation. We determined optimal LAI to maximise canopy photosynthesis and demonstrated that enhancement of canopy photosynthesis by elevated CO₂ was larger at high than at low nitrogen availability. We also studied competitive asymmetry among individuals in an even-aged, monospecific stand at elevated CO₂. Light acquisition (acquired light per unit aboveground mass) and utilisation (photosynthesis per unit acquired light) were calculated for each individual in the stand. Elevated CO₂ enhanced photosynthesis and growth of tall dominants, which reduced the light availability for shorter subordinates and consequently increased size inequality in the stand.     

Nitrogen remobilization response to current supply in young citrus trees

Internal nitrogen (N) storage and remobilization processes support seasonal growth (flowering/fructification and subsequent leaf development) in particular in early spring, when soil temperatures are unfavourable for adequate N uptake. Storage nitrogen mobilization in young citrus trees was studied under two contrasting N supplies; high N (HN) and low N dose (LN) in the critical period of flowering and fruit set. ¹⁵N labelling technique was used to distinguish N derived from internal remobilization from that taken up by the roots. Regardless N supply, the greatest N remobilization took place from the beginning of the vegetative activity until flowering. Low N availability significantly increased (+14%) N retranslocation at the end of June drop agreeing with the hypothesis that reserve mobilization depends on soil N availability during flowering and fruit set. At the end of fruit drop, N remobilization contributed up to 70% and 61% of total N of young organs for LN and HN, respectively. Remobilized N was mainly recovered in abscised organs of both HN and LN trees and to a lesser extent in new flush leaves; however a greater percentage partitioned to abscised organs of LN as a consequence of the greater remobilization rate and the increased fruit abscission. Old leaves of LN remobilized significantly higher N, while woody organs and root system did not show differences between HN and LN supplied trees. The results presented in this paper demonstrate that the amount of N remobilized by young citrus plants depends on external N availability. Thus, low N application rates in early stages (flowering and fruit set) lead to higher translocation of N stored during the previous cycle to developing new organs.