Importance of the gradient in photosynthetically active radiation in a vegetation stand for leaf nitrogen allocation in two monocotyledons

Carex acutiformis and Brachypodium pinnatum were grown with a uniform distribution of photosynthetic photon flux density (PFD) with height, and in a vertical PFD gradient similar to the PFD gradient in a leaf canopy. Distribution of organic leaf N and light-saturated rates of photosynthesis were determined. These parameters were also determined on plants growing in a natural vegetation stand. The effect of a PFD gradient was compared with the effect of a leaf canopy. In Brachypodium, plants growing in a vegetation stand had increasing leaf N with plant height. However, distribution of leaf N was not influenced by the PFD gradient treatment. The gradient of leaf N in plants growing in a leaf canopy was not due to differences within the long, mostly erect, leaves but to differences between leaves. In Carex, however, the PFD gradient caused a clear increase of leaf N with height in individual leaves and thus also in plants. The leaf N gradient was similar to that of plants growing in a leaf canopy. Leaf N distribution was not affected by nutrient availability in Carex. In most cases, photosynthesis was positively related to leaf N. Hence, lightsaturated rates of photosynthesis increased towards the top of the plants growing in leaf canopies in both species and, in Carex, also in the PFD gradient, thus contributing to increased N use efficiency for photosynthesis of the whole plant. It is concluded that in Carex the PFD gradient is the main environmental signal for leaf N allocation in response to shading in a leaf canopy, but one or more other signals must be involved in Brachypodium.     

Nitrogen allocation in response to partial shading of a plant: Possible mechanisms

The significance of photosynthetic and transpiration rates for the perception by plants of light gradients in leaf canopies has been investigated with regard to nitrogen allocation and re-allocation. A gradient of photon flux density (PFD) over a plant's foliage was simulated by shading one leaf of a pair of primary leaves of bean ( Phaseolus vulgaris L. cv. Rentegever). Photosynthetic rate was manipulated independently of PFD and, to some extent, also of transpiration, by subjecting the leaf to different CO₂ concentrations. Transpiration rate was changed independently of PFD and photosynthetic rate by subjecting the leaf to different vapour pressure differences (VPD). A reduced partial pressure of CO₂ reduced specific leaf mass (SLM) as did a decreased PFD, but did not change leaf N per unit area (N_LA) and light saturated rate of photosynthesis (A_max). A reduced VPD caused several effects consistent with the effect of PFD. It decreased N_LA and A_max and increased the chlorophyll to N ratio in old and young leaves. Furthermore, it decreased the chlorophyll a to b ratio and inhibited leaf growth in young leaves. The transpiration stream is partitioned among the leaves of a plant according to their transpiration rates. The results suggest that relative rates of import of xylem sap into leaves of a plant play an important role in the perception of partial shading of a plant, a situation normally found in dense vegetations. The possible role of cytokinin influx into leaves as controlled by transpiration rate, is discussed.     

Nitrogen allocation in response to partial shading of a plant: Possible mechanisms

The significance of photosynthetic and transpiration rates for the perception by plants of light gradients in leaf canopies has been investigated with regard to nitrogen allocation and re-allocation. A gradient of photon flux density (PFD) over a plant's foliage was simulated by shading one leaf of a pair of primary leaves of bean ( Phaseolus vulgaris L. cv. Rentegever). Photosynthetic rate was manipulated independently of PFD and, to some extent, also of transpiration, by subjecting the leaf to different CO₂ concentrations. Transpiration rate was changed independently of PFD and photosynthetic rate by subjecting the leaf to different vapour pressure differences (VPD). A reduced partial pressure of CO₂ reduced specific leaf mass (SLM) as did a decreased PFD, but did not change leaf N per unit area (N_LA) and light saturated rate of photosynthesis (A_max). A reduced VPD caused several effects consistent with the effect of PFD. It decreased N_LA and A_max and increased the chlorophyll to N ratio in old and young leaves. Furthermore, it decreased the chlorophyll a to b ratio and inhibited leaf growth in young leaves. The transpiration stream is partitioned among the leaves of a plant according to their transpiration rates. The results suggest that relative rates of import of xylem sap into leaves of a plant play an important role in the perception of partial shading of a plant, a situation normally found in dense vegetations. The possible role of cytokinin influx into leaves as controlled by transpiration rate, is discussed.     

Light,leaf age,and leaf nitrogen concentration in a tropical vine

Summary Tropical vines in the Araceae family commonly exhibit alternating periods of upward and downward growth, decoupling the usual relationship between decreasing light environment with increasing age among the leaves on a shoot. In this study I examined patterns of light, leaf specific mass, and leaf nitrogen concentration in relation to leaf position, a measure of developmental age, in field collected shoots of Syngonium podophyllum. These data were analyzed to test the hypothesis that nitrogen allocation parallels within-shoot gradients of light availability, regardless of the relationship between light and leaf age. I found that leaf nitrogen concentration, on a mass basis, was weakly correlated with leaf level light environment. However, leaf specific mass, and consequently nitrogen per unit leaf area, were positively correlated with gradients of light within the shoot, and either increased or decreased with leaf age, providing support for the hypothesis that nitrogen allocation parallels gradients of light availability.     

Seedling growth of five tropical dry forest tree species in relation to light and nitrogen gradients

Aims Increasing anthropogenic nitrogen (N) deposition has been claimed to induce changes in species composition and community dynamics. A greenhouse experiment was conducted to examine the effect of increased N availability on growth and functional attributes of seedlings of five tree species with different life history characteristics under varying irradiances. The following questions have been addressed: (i) how do the pioneer and non-pioneer species respond in absolute growth and relative growth rate (RGR) to the interaction of light and nitrogen? (ii) how does the interaction between irradiance and nitrogen availability modulate growth attributes (i.e. functional attributes)? (iii) is there any variation in growth responses between leguminous and non-leguminous species along the light and nitrogen gradients?Methods Seedlings of five tree species (Acacia catechu, Bridelia retusa, Dalbergia sissoo, Lagerstroemia parviflora and Terminalia arjuna) were subjected to twelve combinations of irradiance and N levels. Various growth traits, including height (HT), basal area (BA), whole plant dry biomass (M D), leaf mass per unit area (LMA), leaf area ratio (LAR), net assimilation rate (NAR), RGR, biomass fractions, root-to-shoot ratio (R:S) and leaf nitrogen content, were studied to analyse intra- and inter-specific responses to interacting light and N gradients.Important findings Significant interactions for irradiance and N availability for majority of growth attributes indicates that growth and biomass allocation of seedlings were more responsive to N availability under high irradiance. However, species responded differentially to N addition and they did not follow successional status. Slow growers (B. retusa, a shade-tolerant species and L. parviflora, a light demander) exhibited greater response to N enrichment than the fast growers (A. catechu, D. sissoo and T. arjuna). However, N-mediated increment in growth traits was greater in non-legumes (B. retusa, L. parviflora and T. arjuna) compared with that of legumes (A. catechu and D. sissoo). Allocation of biomass to root was strongly suppressed at the highest N supply across species; however, at high irradiance and high N availability, a greater suppression in R:S ratio was observed for B. retusa. NAR was a stronger determinant of RGR relative to LAR, suggesting its prominent role in increased RGR along increasing irradiances. Overall, a higher growth response of slow-growing species to elevated N levels, particularly the non-pioneers (B. retusa and L. parviflora) suggests that future N deposition may lead to perturbations in competition hierarchies and species composition, ultimately affecting community dynamics in nutrient-poor tropical dry forests.     

Modest enhancement of nitrogen conservation via retranslocation in response to gradients in N supply and leaf N status

Plant nutrient resorption, a ubiquitous mechanism of nutrient conservation, has often been proposed to be more pronounced in infertile than fertile habitats, and in species common to infertile compared to fertile habitats, because of the presumed advantage when nutrients are scarce. However, previous studies provide weak and inconsistent empirical support for these hypotheses, although few have examined intraspecific variation across well-quantified resource gradients. This study addresses intraspecific patterns of nutrient resorption for eight species across two N availability gradients on similar soils in an N-limited oak savanna ecosystem: a long-term fire frequency gradient with a negatively correlated N fertility gradient and a long-term N fertilization gradient. We hypothesized that both resorption proficiency (the minimum nutrient level retained in a senesced leaf) and efficiency (the proportional change in leaf nutrient concentration) would decrease with increasing soil N availability and plant N status. For the seven non-N fixers, either resorption proficiency or efficiency decreased modestly in treatments with higher N availability. In contrast, the legume Amorpha canescens Pursh had higher N levels in green and senesced leaves, and resorbed N much more weakly than the non-fixers, and did not respond in terms of proficiency or efficiency to soil N availability. Across all species and sites in each N fertility gradient, a scaling analysis showed greater resorption efficiency in plants with lower N concentrations. Our data suggest that species can have modest resorption responses reflective of soil nutrient availability and differences in resorption related to their N economy that represent mechanisms of nutrient conservation in nutrient-limited soils.     

Developmental changes in shoot N dynamics of lucerne (Medicago sativa L.) in relation to leaf growth dynamics as a function of plant density and hierarchical position within the canopy

Shoot N concentration in plants decreases as they get bigger, due to the fact that N accumulates less rapidly than dry matter in plants during the plant growth process, leading to an allometric relationship between shoot N content (N(sh)) and shoot mass (W(sh)): N(sh)=a(W(sh))b. The results obtained on lucerne plants growing either under controlled low density conditions or in dense stands under field conditions show that the value of the allometric coefficient b that represents the ratio between the relative N accumulation rate in shoots [dN(sh)/(N(sh)dt)] and the relative growth rate [dW(sh)/(W(sh)dt)], decreases from 0.88 for a low plant density to 0.72 for a dense stand. Therefore, the fractional increase of shoot N per unit of shoot dry matter is lower when plants are in competition for light in dense canopies. This decrease can be entirely explained by the parallel decline in the leaf area per unit of shoot mass. Thus, a remarkably constant linear relationship can be established between N(sh) and leaf area (LA): N(sh)=1.7 g m(-2) LA, regardless of the conditions (low versus high density, controlled versus field conditions). Moreover, in a field dense stand, the comparison of plants with contrasting positions between the top and the bottom of the canopy (dominant, intermediate or suppressed plants), also shows that the difference in N(sh) at similar shoot mass is explained by the proportion of leaf mass to shoot mass. These data support the idea that leaf growth drives the dynamics of shoot N accumulation. These results also indicate that competition for light among individual plants within a dense canopy induces developmental changes in plant morphology (leaf:stem ratio) that explain the differences observed in shoot N concentration. This last observation could be extrapolated to multi-specific plant stands. Therefore, the sharing of N resources among plant species could partially be the result of the sharing of light within the canopy.     

Shoot growth dynamics and photosynthetic response to increased nitrogen availability in the alpine willow Salix glauca

Plants subjected to increases in the supply of resource(s) limiting growth may allocate more of those resources to existing leaves, increasing photosynthetic capacity, and/or to production of more leaves, increasing whole-plant photosynthesis. The responses of three populations of the alpine willow, Salix glauca, growing along an alpine topographic sequence representing a gradient in soil moisture and organic matter, and thus potential N supply, to N amendments, were measured over two growing seasons, to elucidate patterns of leaf versus shoot photosynthetic responses. Leaf-(foliar N, photosynthesis rates, photosynthetic N-use efficiency) and shoot-(leaf area per shoot, number of leaves per shoot, stem weight, N resorption efficiency) level measurements were made to examine the spatial and temporal variation in these potential responses to increased N availability. The predominant response of the willows to N fertilization was at the shoot-level, by production of greater leaf area per shoot. Greater leaf area occurred due to production of larger leaves in both years of the experiment and to production of more leaves during the second year of fertilization treatment. Significant leaflevel photosynthetic response occurred only during the first year of treatment, and only in the dry meadow population. Variation in photosynthesis rates was related more to variation in stomatal conductance than to foliar N concentration. Stomatal conductance in turn was significantly related to N fertilization. Differences among the populations in photosynthesis, foliar N, leaf production, and responses to N fertilization indicate N availability may be lowest in the dry meadow population, and highest in the ridge population. This result is contrary to the hypothesis that a gradient of plant available N corresponds with a snowpack/topographic gradient.     

The response of isoprene emission rate and photosynthetic rate to photon flux and nitrogen supply in aspen and white oak trees

Isoprene is the primary biogenic hydrocarbon emitted from temperate deciduous forest ecosystems. The effects of varying photon flux density (PFD) and nitrogen growth regimes on rates of isoprene emission and net photosynthesis in potted aspen and white oak trees are reported. In both aspen and oak trees, whether rates were expressed on a leaf area or dry mass basis, (1) growth at higher PFD resulted in significantly higher rates of isoprene emission, than growth at lower PFD, (2) there is a significant positive relationship between isoprene emission rate and leaf nitrogen concentration in both sun and shade trees, and (3) there is a significant positive correlation between isoprene emission rate and photosynthetic rate in both sun and shade trees. The greater capacity for isoprene emission in sun leaves was due to both higher leaf mass per unit area and differences in the biochemical and/or physiological properties that influence isoprene emission. Positive correlations between isoprene emission rate and leaf nitrogen concentration support the existence of mechanisms that link leaf nitrogen status to isoprene synthase activity. Positive correlations between isoprene emission rate and photosynthesis rate support previous hypotheses that isoprene emission plays a role in protecting photosynthetic mechanisms during stress.     

Optimizing nitrogen economy under drought: increased leaf nitrogen is an acclimation to water stress in willow (Salix spp.)

Background and Aims

The major objective was to identify plant traits functionally important for optimization of shoot growth and nitrogen (N) economy under drought. Although increased leaf N content (area basis) has been observed in dry environments and theory predicts increased leaf N to be an acclimation to drought, experimental evidence for the prediction is rare.

Methods

A pedigree of 200 full-sibling hybrid willows was pot-grown in a glasshouse in three replicate blocks and exposed to two water regimes for 3 weeks. Drought conditions were simulated as repeated periods of water shortage. The total leaf mass and area, leaf area efficiency (shoot growth per unit leaf area, E_A), area-based leaf N content (N_A), total leaf N pool (N_L) and leaf N efficiency (shoot growth per unit leaf N, E_N) were assessed.

Key Results

In the water-stress treatment, shoot biomass growth was N limited in the genotypes with low N_L, but increasingly limited by other factors in the genotypes with greatest N_L. The N_A was increased by drought, and drought-induced shift in N_A varied between genotypes (significant G × E). Judged from the E_A–N_A relationship, optimal N_A was 16 % higher in the water-stress compared with the well-watered treatment. Biomass allocation to leaves and shoots varied between treatments, but the treatment response of the leaf : shoot ratio was similar across all genotypes.

Conclusions

It is concluded that N-uptake efficiency and leaf N efficiency are important traits to improve growth under drought. Increased leaf N content (area basis) is an acclimation to optimize N economy under drought. The leaf N content is an interesting trait for breeding of willow bioenergy crops in a climate change future. In contrast, leaf biomass allocation is a less interesting breeding target to improve yield under drought.     

Development of a nutrient pollution indicator using the seagrass,Zostera marina,along nutrient gradients in three New England estuaries

Worldwide, seagrasses provide important habitats in coastal ecosystems, but seagrass meadows are often degraded or destroyed by cultural eutrophication. Presently, there are no available tools for early assessment of nutrient over-enrichment; direct measurements of water column nutrients are ineffective since the nutrients typical of early enrichment are rapidly taken up by plants within the ecosystem. We investigated whether, in a gradient of nutrient availability but prior to actual habitat loss, eelgrass (Zostera marina L.) plant morphology and tissue nutrients might reflect environmental nutrient availability. Eelgrass responses to nitrogen along estuarine gradients were assessed; two of these plant responses were combined to create an early indicator of nutrient over-enrichment. Eelgrass plant morphology and leaf tissue nitrogen (N) were measured along nutrient gradients in three New England estuaries: Great Bay Estuary (NH), Narragansett Bay (RI) and Waquoit Bay (MA). Eelgrass leaf N was significantly higher in up-estuary sampling stations than stations down-estuary, reflecting environmental nitrogen gradients. Leaf N content showed high variance, however, limiting its ability to discriminate the early stages of eutrophication. To find a stronger indicator, plant morphological characteristics such as number of leaves per shoot, blade width, and leaf and sheath length were examined, but they only weakly correlated with leaf tissue N. Area normalized leaf mass (mg dry weight cm⁻²), however, exhibited a strong and consistently negative relationship with leaf tissue N and a significant response to the estuarine nutrient gradients. We found the ratio of leaf N to leaf mass to be a more sensitive and consistent indicator of early eutrophication than either characteristic alone. We suggest the use of this ratio as a nutrient pollution indicator (NPI).     

Effects of leaf age,nitrogen nutrition and photon flux density on the distribution of nitrogen among leaves of a vine (Ipomoea tricolor Cav.) grown horizontally to avoid mutual shading of leaves

Effects of leaf age, nitrogen nutrition and photon flux density (PFD) on the distribution of nitrogen among leaves were investigated in a vine, Ipomoea tricolor Cav., which had been grown horizontally so as to avoid mutual shading of leaves. The nitrogen content was highest in newly developed young leaves and decreased with age of leaves in plants grown at low nitrate concentrations and with all leaves exposed to full sunlight. Thus, a distinct gradient of leaf nitrogen content was formed along the gradient of leaf age. However, no gradient of leaf nitrogen content was formed in plants grown at a high nitrate concentration. Effects of PFD on the distribution of nitrogen were examined by shading leaves in a manner that simulated changes in the light gradient of an erect herbaceous canopy (i.e., where old leaves were placed under increasingly darker conditions with growth of the canopy). This canopy-type shading steepened the gradient of leaf nitrogen content in plants grown at a low nitrogen supply, and created a gradient in plants grown at high concentrations of nitrate. The steeper the gradient of PFD, the larger the gradient of leaf nitrogen that was formed. When the gradient of shading was inverted, that is, younger leaves were subjected to increasingly heavier shade, while keeping the oldest leaves exposed to full sunlight, an inverted gradient of leaf nitrogen content was formed at high nitrate concentrations. The gradient of leaf nitrogen content generated either by advance of leaf age at low nitrogen availability, or by canopy-type shading, was comparable to those reported for the canopies of erect herbaceous plants. It is concluded that both leaf age and PFD have potential to cause the non-uniform distribution of leaf nitrogen. It is also shown that the contribution of leaf age increases with the decrease in nitrogen nutrition level.     

The influence of nitrogen availability on carbon and nitrogen storage in the biennial Cirsium vulgare (Savi) Ten. I. Storage capacity in relation to resource acquisition, allocation and recycling

Plants of Cirsium vulgare (Savi) Ten. were cultivated under five different nitrogen regimes in order to investigate the effects of nitrogen supply on the storage processes in a biennial species during its first year of growth. External N supply increased total biomass production without changing the relationship between ‘productive plant compartments’ (i.e. shoot plus fine roots) and ‘storage plant compartments’ (i.e. structural root dry weight, which is defined as the difference between tap root biomass and the amount of stored carbohydrates and N compounds). The amount of carbohydrates and N compounds stored per unit of structural tap root dry weight was not affected by external N availability during the season, because high rates of N supply increased the concentration of N compounds whilst decreasing the carbohydrate concentration, and low rates of N supply had the opposite effect. Mobilization of N from senescing leaves was not related to the N status of the plants. The relationship between nitrogen compounds stored in the tap root and the maximum amount of nitrogen in leaves was an increasing function with increasing nitrogen supply. We conclude that the allocation between vegetative plant growth and the growth of storage structures over a wide range of N availability seems to follow predictions from optimum allocation theory, whereas N storage responds in a rather plastic way to N availability.     

Photosynthetic acclimation to light in woody and herbaceous species: a comparison of leaf structure, pigment content and chlorophyll fluorescence characteristics measured in the field

Acclimation of foliage photosynthetic properties occurs with varying time kinetics, but structural, chemical and physiological factors controlling the kinetics of acclimation are poorly understood, especially in field environments. We measured chlorophyll fluorescence characteristics, leaf total carotenoid (Car), chlorophyll (Chl) and nitrogen (N) content and leaf dry mass per area (LMA) along vertical light gradients in natural canopies of the herb species, Inula salicina and Centaurea jacea, and tree species, Populus tremula and Tilia cordata, in the middle of the growing season. Presence of stress was assessed on the basis of night measurements of chlorophyll fluorescence. Our aim was to compare the light acclimation of leaf traits, which respond to light availability at long (LMA and N), medium (Chl a/b ratio, Car/Chl ratio) and short time scales (fluorescence characteristics). We found that light acclimation of nitrogen content per unit leaf area (N(area)), chlorophyll content per unit dry mass (Chl(mass)) and Chl/N ratio were related to modifications in LMA. The maximum PSII quantum yield (F(v) /F(m)) increased with increasing growth irradiance in I. salicina and P. tremula but decreased in T. cordata. Leaf growth irradiance, N content and plant species explained the majority of variability in chlorophyll fluorescence characteristics, up to 90% for steady-state fluorescence yield, while the contribution of leaf total carotenoid content was generally not significant. Chlorophyll fluorescence characteristics did not differ strongly between growth forms, but differed among species within a given growth form. These data highlight that foliage acclimation to light is driven by interactions between traits with varying time kinetics.     

Photosynthesis and resource distribution through plant canopies

Plant canopies are characterized by dramatic gradients of light between canopy top and bottom, and interactions between light, temperature and water vapour deficits. This review summarizes current knowledge of potentials and limitations of acclimation of foliage photosynthetic capacity (A(max)) and light-harvesting efficiency to complex environmental gradients within the canopies. Acclimation of A(max) to high light availability involves accumulation of rate-limiting photosynthetic proteins per unit leaf area as the result of increases in leaf thickness in broad-leaved species and volume: total area ratio and mesophyll thickness in species with complex geometry of leaf cross-section. Enhancement of light-harvesting efficiency in low light occurs through increased chlorophyll production per unit dry mass, greater leaf area per unit dry mass investment in leaves and shoot architectural modifications that improve leaf exposure and reduce within-shoot shading. All these acclimation responses vary among species, resulting in species-specific use efficiencies of low and high light. In fast-growing canopies and in evergreen species, where foliage developed and acclimated to a certain light environment becomes shaded by newly developing foliage, leaf senescence, age-dependent changes in cell wall characteristics and limited foliage re-acclimation capacity can constrain adjustment of older leaves to modified light availabilities. The review further demonstrates that leaves in different canopy positions respond differently to dynamic fluctuations in light availability and to multiple environmental stresses. Foliage acclimated to high irradiance respond more plastically to rapid changes in leaf light environment, and is more resistant to co-occurring heat and water stress. However, in higher light, co-occurring stresses can more strongly curb the efficiency of foliage photosynthetic machinery through reductions in internal diffusion conductance to CO(2). This review demonstrates strong foliage potential for acclimation to within-canopy environmental gradients, but also highlights complex constraints on acclimation and foliage functioning resulting from light x foliage age interactions, multiple environmental stresses, dynamic light fluctuations and species-specific leaf and shoot structural constraints.     

Interactive effects of soil nitrogen and water availability on leaf mass loss in a temperate steppe

Although the link between leaf mass loss and assessment of ecosystem nutrient use efficiency and plant nutrient resorption efficiency has received considerable attention in various ecosystems, there has been relatively little effort to assess plant leaf mass loss during senescence, especially for herbaceous species. We conducted experimental studies to assess leaf mass loss during senescence in five dominant herbaceous species and examined the effects of increasing nitrogen (N) and water availability on leaf mass loss of four species in a temperate steppe in northern China. We nondestructively estimated mature leaf mass based on leaf length and width. Leaf mass loss varied substantially among species, ranging from 20–50%. On average across all species, N and water addition increased leaf mass loss by 30% and 19%, respectively. N and water addition interacted to affect leaf mass loss, as water addition had a significant positive effect on leaf mass loss under enriched N conditions but showed no effect under ambient N levels. We conclude that leaf mass loss of herbaceous plants was considerable and can potentially be more pronounced with increasing N and water availability. It is notable that the responses of plant species to N and water addition were variable. We suggest that leaf mass loss during senescence should be given full consideration in assessing nutrient use and resorption efficiency in semi-arid areas.     

Grain yield and kernel weight of two maize genotypes differing in nitrogen use efficiency at various levels of nitrogen and carbohydrate availability during flowering and grain filling

Grain yield per plant (GYP) and mean kernel weight (KW) of maize (Zea mays L.) are sensitive to changes in the environment during the lag phase of kernel growth (the time after pollination in which the potential kernel size is determined), and during the phase of linear kernel growth. The aim of this study was to assess genotypic differences in the response to environmental stresses associated with N and/or carbohydrate shortage at different phases during plant development. The rate and timing of N and carbohydrate supply were modified by application of fertilizer, shading, and varying the plant density at sowing, at silking or at 14 d after silking. The effects of these treatments on the photosynthetic capacity, grain yield and mean kernel weight were investigated in two hybrids differing in N use efficiency. The total above-ground biomass and grain yield per plant of the efficient hybrid responded little to altered environmental conditions such as suboptimal N supply, enhanced inter-plant competition, and shading for 14 d during flowering, when compared to the less efficient genotype. We conclude that grain yields in the efficient genotype are less sensitive not only to N stress, but also to carbohydrate shortage before grain filling. Shading of N deficient plants from 14 d after silking to maturity did not significantly reduce grain yield in the non-efficient genotype, indicating complete sink limitation of grain yield during grain filling. In the efficient genotype, in contrast, grain yield of N-deficient plants was significantly reduced by shading during grain filling. The rate of photosynthesis declined with decreasing foliar N content. No genotypic differences in photosynthesis were observed at high or low foliar N contents. However, at high plant density and low N supply, the leaf chlorophyll content after flowering in the efficient genotype was higher than that in the non-efficient genotype. Obviously, the higher source capacity of the efficient genotype was not due to higher photosynthetic N use efficiency but due to maintenance of high chlorophyll contents under stressful conditions. In the efficient genotype, the harvest index was not significantly affected by N fertilization, plant density, or shading before the grain filling period. In contrast, in the non-efficient genotype the harvest index was diminished by N deficiency and shading during flowering. We conclude that the high yielding ability of the efficient genotype under stressful conditions was associated with formation of a high sink capacity of the grains under conditions of low carbohydrate and N availability during flowering and with maintenance of high source strength during grain filling under conditions of high plant density and low N availability.     

The effects of light and nitrogen on photosynthesis,leaf characteristics,and dry matter allocation in the chaparral shrub,Diplacus aurantiacus

Summary Plants of Diplacus aurantiacus, a successional shrub common in California chaparral, were grown under controlled conditions in which either quantum flux density or nitrogen availability was varied. Photosynthesis and leaf nitrogen content were determined on a leaf area and a leaf weight basis, and whole plant growth was monitored.There was a direct relationship between photosynthesis and leaf nitrogen content on both area and weight bases. Reduced light intensity of the growth environment resulted in reductions in light-saturated photosynthesis and nitrogen content on an area basis, but not on a weight basis. With reduced nitrogen availability, photosynthesis and leaf nitrogen content per unit leaf weight decreased.Resource use efficiency increased as the resource became more limiting. The results are consistent with a model of plant growth in which net carbon gain of the leaf is maximized. Abbreviations. For brevity, the following set of abbreviations is used in presenting and discussing the results. P/area and N/area are, respectively, photosynthesis and leaf nitrogen content per unit leaf area. P/wt and N/wt are the same quantities per unit leaf dry weight. SLW (specific leaf weight) is dry weight per unit leaf area. RGR (relative growth rate) is the relative rate of increase in shoot dry matter per day     

Dependence of starch storage on nutrient availability and photon flux density in small birch Betula pendula Roth)

Abstract Small birch plants (Betula pendula Roth) were grown in a climate chamber at different levels of nutrient availability and at two photon flux densities. The extent to which starch storage was dependent upon nutrient availability and photon flux density was investigated. Acclimated values of starch concentration in leaves were highest at low nutrient availability and high photon flux density. Starch storage in roots was only found at the lowest nutrient availability. However, the relative rate of starch storage (starch stored per unit plant dry weight and time) was higher in plants with good nutrition. The data suggest that, at sub-optimal nutrient availability, the momentary rate of net shoot photosynthesis is unlikely to limit the structural (as opposed to carbon storage) growth of the plant. Although photosynthetic rate per unit leaf area (as measured at the growth climate) was slightly lower in plants with poor nutrient availability, photosynthetic rate per unit leaf nitrogen was higher. These data suggest a priority of leaf nitrogen usage in photosynthesis, with limiting amounts of leaf nitrogen (and possibly other nutrients) for subsequent growth processes. This argument is consistent with the higher concentrations of starch found in plants with poor nutrient availability.     

The role of nitrogen in a simple scheme to scale up photosynthesis from leaf to canopy

A simple analytical scheme, involving the distribution of nitrogen, to scale up photosynthesis from leaf to canopy is proposed. The scheme is based on the assumption that there are two pools of nitrogen in leaves: nitrogen in photosynthetic, degradable structures (N_p) and nitrogen in non-photosynthetic and non-degradable structures (N_s). The rate of photon-saturated photosynthesis, F_m, is assumed to be proportional to N_p and is distributed inside the canopy similarly to photon flux density (PFD). Prior assumptions of an optimum distribution of nitrogen are not a prerequisite. Calculations made with the scheme lead to development of the hypothesis that the canopy can be treated as a ‘big leaf’ on the time scales involved in acclimation of photosynthesis to PFD. Simulations using parameters for tree species with different requirements for PFD show that shade-tolerant species may have denser canopies than sun-demanding species because of smaller amounts of non-photosynthetic structural nitrogen and/or supporting tissue in their leaves.