Physiological and production responses of plant growth forms to increases in limiting resources in alpine tundra: implications for differential community response to environmental change

Physiological and growth measurements were made on forbs and graminoids following additions of water and N+water in a graminoid-dominated dry meadow and a forb-dominated moist meadow, to determine if the community-level response was related to differential responses between the growth forms. Graminoids had higher photosynthetic rates and lower transpiration rates and foliar N concentrations than forbs, and consequently maintained higher photosynthetic N- and water-use efficiencies. Photosynthetic rates, stomatal conductance, and transpiration rates increased significantly only in response to N fertilization and only in moist meadow species. The increase in photosynthetic rates was unrelated to variation in foliar N concentration, but instead correlated with variation in stomatal conductance. Growth based N-use efficiency was higher in moist meadow graminoids than in moist meadow forbs, but did not differ between the growth forms in the dry meadow. The moist meadow community had higher biomass and N standing crops, but the relative increase in these factors in response to N fertilization was greater in the dry meadow. Graminoids had a greater relative increase in biomass and N accumulation than forbs following N fertilization, but moist meadow graminoids exhibited a greater response than dry meadow graminoids. The difference in the growth response between the dry meadow and moist meadow graminoids to N fertilization was correlated with more conservative leaf gas exchange responses in dry meadow species, presumably related to a higher frequency of soil water deficits in this community. Community-level response to the resource additions was therefore mediated by the plant growth form response, corresponding with differences between the growth forms in physiological factors related to resource acquisition and use.     

Light-exposed shoots of seven coexisting deciduous species show common photosynthetic responses to tree height

Functional traits of light-exposed leaves have been reported to show tree height-dependent change. However, it remains unknown how plastic response of leaf traits to tree height is linked with shoot-level carbon gain. To answer this question, we examined the photosynthetic properties of fully lit current-year shoots in crown tops with various heights for seven deciduous broad-leaved species dominated in a cool–temperate forest in northern Japan. We measured leaf mass, stomatal conductance, nitrogen content, light-saturated net photosynthetic rate (all per leaf lamina area), foliar stable carbon isotope ratio, and shoot mass allocation to leaf laminae. We employed hierarchical Bayesian models to simultaneously quantify inter-trait relationships for all species. We found that leaf and shoot traits were co-varied in association with height, and that there was no quantitative inter-specific difference in leaf- and shoot-level plastic responses to height. Nitrogen content increased and stomatal conductance decreased with height. Reflecting these antagonistic responses to height, photosynthetic rate was almost unchanged with height. Photosynthetic rate divided by stomatal conductance as a proxy of photosynthetic water use efficiency sufficiently explained the variation of foliar carbon isotope ratio. The increase in mass allocation to leaves in a shoot compensated for the height-dependent decline in photosynthetic rate per leaf lamina mass. Consequently, photosynthetic gain at the scale of current-year shoot mass was kept unchanged with tree height. We suggest that the convergent responses of shoot functional traits across species reflect common requirements for trees coexisting in a forest.     

Ozone exposure over two growing seasons alters root-to-shoot ratio and chemical composition of birch (Betula pendula Roth)

Physiological and chemical responses of 17 birch (Betula pendula Roth) clones to 1.5–1.7 × ambient ozone were studied in an open‐field experiment over two growing seasons. The saplings were studied for growth, foliar visible injuries, net photosynthesis, stomatal conductance, and chlorophyll, carotenoid, Rubisco, total soluble protein, macronutrient and phenolic concentrations in leaves. Elevated ozone resulted in growth enhancement, changes in shoot‐to‐root (s/r) ratio, visible foliar injuries, reduced stomatal conductance, lower late‐season net photosynthesis, foliar nutrient imbalance, changes in phenolic composition, and reductions in pigment, Rubisco and soluble protein contents indicating accelerated leaf senescence. Majority of clones responded to ozone by changing C allocation towards roots, by stomatal closure (reduced ozone uptake), and by investment in low‐cost foliar antioxidants to avoid and tolerate ozone stress. A third of clones, showing increased s/r ratio, relied on inducible efficient high‐cost antioxidants, and enhanced leaf production to compensate ozone‐caused decline in leaf‐level net photosynthesis. However, the best ozone tolerance was found in two s/r ratio‐unaffected clones showing a high constitutive amount of total phenolics, investment in low‐cost antioxidants and N distribution to leaves, and lower stomatal conductance under ozone stress. The results highlight the importance of phenolic compounds in ozone defence mechanisms in the birch population. Depending on the genotype, ozone detoxification was improved by an increase in either efficient high‐cost or less efficient low‐cost antioxidative phenolics, with close connections to whole‐plant physiology.     

Effects of simulated herbivory on photosynthesis and N resorption efficiency in <Emphasis Type="Italic">Quercus</Emphasis><Emphasis Type="Italic">pyrenaica</Emphasis> Willd. saplings

We examined the effects of simulated folivory by caterpillars on photosynthetic parameters and nitrogen (N) resorption efficiency in Quercus pyrenaica saplings. We analyzed the differences between intact leaves in control plants, punched leaves in damaged plants, and intact leaves in damaged plants. We then established two levels of simulated folivory: low (≈13% of the leaf area of one main branch removed per plant) and high (≈26% of the leaf area of one main branch removed per plant) treatments. No differences were found in net assimilation rate and conductance between either leaf type or treatment during the most favourable period for photosynthesis. However, the N content was lower in punched than in intact leaves, and as a result PNUE was higher in damaged leaves from treated trees. In leaf-litter samples, N mass was significantly higher in punched than in intact leaves in treated plants, and LMA was significantly higher in damaged than in intact leaves of both the treated and control plants. Consequently, N resorption efficiency was around 15% lower in damaged leaves as compared with intact leaves from treated and control plants. Mechanical injury to leaves not only triggered no compensatory photosynthetic response to compensate a lower carbon uptake due to leaf area loss, but also affected the resorption process that characterizes leaf senescence.     

Photosynthesis in leaves and siliques of winter oilseed rape (Brassica napus L.)

The rate of photosynthesis and its relation to tissue nitrogen content was studied in leaves and siliques of winter oilseed rape (Brassica napus L.) growing under field conditions including three rates of nitrogen application (0, 100 or 200 kg N ha^-1) and two levels of irrigation (rainfed or irrigated at a deficit of 20 mm). The predominant effect of increasing N application under conditions without water deficiency was enhanced expansion of photosynthetically active leaf and silique surfaces, while the rate of photosynthesis per unit leaf or silique surface area was similar in the different N treatments. Thus, oilseed rape did not increase N investment in leaf area expansion before a decline in photosynthetic rate per unit leaf area due to N deficiency could be avoided. Much less photosynthetically active radiation penetrated into high-N canopies than into low-N canopies. The specific leaf area increased markedly in low light conditions, causing leaves in shade to be less dense than leaves exposed to ample light. In both leaves and siliques the photosynthetic rate per unit surface area responded linearly to increasing N content up to about 2 g m^-2, thus showing a constant rate of net CO₂ assimilation per unit increment in N (constant photosynthetic N use efficiency). At higher tissue N contents, photosynthetic rate responded less to changes in N status. Expressed per unit N, light saturated photosynthetic rate was three times higher in leaves than in silique valves, indicating a more efficient photosynthetic N utilization in leaves than in siliques. Nevertheless, from about two weeks after completion of flowering and onwards total net CO₂ fixation in silique valves exceeded that in leaves because siliques received much higher radiation intensities than leaves and because the leaf area declined rapidly during the reproductive phase of growth. Water deficiency in late vegetative and early reproductive growth stages reduced the photosynthetic rate in leaves and, in particular, siliques of medium- and high-N plants, but not of low-N plants.     

Hydraulic limitation not declining nitrogen availability causes the age‐related photosynthetic decline in loblolly pine (Pinus taeda L.)

Declining net primary production (NPP) with forest age is often attributed to a corresponding decline in gross primary production (GPP). We tested two hypotheses explaining the decline of GPP in ageing stands (14–115 years old) of Pinus taeda L.: (1) increasing N limitation limits photosynthetic capacity and thus decreases GPP with increasing age; and (2) hydraulic limitations increasingly induce stomatal closure, reducing GPP with increasing age. We tested these hypotheses using measurements of foliar nitrogen, photosynthesis, sap‐flow and dendroclimatological techniques. Hypothesis (1) was not supported; foliar N retranslocation did not increase and declines were not observed in foliar N, leaf area per tree or photosynthetic capacity. Hypothesis (2) was supported; declines were observed in light‐saturated photosynthesis, leaf‐ and canopy‐level stomatal conductance, concentration of CO₂ inside leaf air‐spaces (corroborated by an increase in wood δ¹³C) and specific leaf area (SLA), while stomatal limitation and the ratio of sapwood area (SA) to leaf area increased. The sensitivity of radial growth to inter‐annual variation in temperature and drought decreased with age, suggesting that tree water use becomes increasingly conservative with age. We conclude that hydraulic limitation increasingly limits the photosynthetic rates of ageing loblolly pine trees, possibly explaining the observed reduction of NPP.     

Light adaptation and the role of autotrophic epiphytes in primary production of the temperate seagrass, Zostera marina L.

Photosynthetic features of Zostera marina L. and its autotrophic epiphyte community were investigated in a population inhabiting a shallow (1.3 m depth) water meadow in Great Harbor, Woods Hole, MA (U.S.A.). Photosynthesis versus irradiance (P-I) relationships were measured with respect to leaf age determined by the leaf position in the shoot bundle and by location of the tissue along the leaf axis. Therefore both age and light intensity gradients along the leaf axis were considered. The maximum photosynthesis (P_max) per dm² typically increased nearly two-fold along the leaf axis from leaf bases to apices. Photosynthetic rate on a chlorophyll (Chl) basis did not increase as dramatically along the leaf axis, and rates were usually lowest in tissues with the highest Chl content. The P-I relationships of leaves of different ages did not reveal photoinhibition even at light intensities > 1400 μE • m⁻² • s ⁻¹. Furthermore, no photoinhibition was observed in tissues from leaf blade bases, which never experienced high light levels (> 500 μE • m ⁻² • s⁻¹) in situ in Great Harbor. The initial slopes of the P-I curves and light compensation and saturation values varied along the leaf axis in relation to in situ light intensity gradients and in relation to leaf or tissue age. It appeared that leaf and/or tissue age was more important than light environment in determining P-I responses. The contribution of the autotrophic epiphyte community on Z. marina leaves to total photosynthesis per dm² was between 27 and 50%, and between 10 and 44% per mg chlorophyll. These levels of epiphyte photosynthesis can double the primary production of Z. marina leaves. No detrimental effects of epiphyte cover were realized in leaf maximal photosynthesis or P-I relationships. Non-epiphytized leaves and leaves from which epiphytes were removed showed essentially identical photosynthetic features. Light intensity and age gradients along the leaf axis control both the photosynthetic performance of the leaves and epiphyte biomass and photosynthesis.     

Effects of altitudinal gradient on Salix atopantha foliar delta13C

In 2010, measurements were conducted on the foliar delta13C, photosynthesis, CO2 diffusive conductivity, nitrogen content, photosynthetic nitrogen use efficiency (PNUE), and special leaf area (SLA) of Salix atopantha at different altitudes (2350 m, 2700 m, 3150 m, and 3530 m) in Wolong Natural Reserve. With the increase of altitude, the foliar nitrogen content (especially the nitrogen content per unit leaf area, N(area)) and the PNUE increased, and the foliar delta13C had a significant increase, with an increment of 1.4 per thousand per 1000 m altitude. The stomatal and mesophyll CO2 diffusion conductance also increased with increasing altitude, which had definite negative effect on the increase of foliar delta13C, but the effect was not strong enough. Comparing with CO2 diffusion conductance, carboxylation capacity was a more important factor limiting the P(c)/P(a), and even, the foliar delta13C. At altitude 2350-2700 m, air temperature was the main factor affecting the allocation of nitrogen in S. atopantha photosynthetic system, whereas at altitude 2700-3530 m, light could be the main affecting factor. No significant difference was observed in the SLA at different altitudes.     

Relationships between photosynthetic activity and silica accumulation with ages of leaf in Sasa veitchii (Poaceae, Bambusoideae)

BACKGROUND AND AIMS: Bamboos have long-lived, evergreen leaves that continue to accumulate silica throughout their life. Silica accumulation has been suggested to suppress their photosynthetic activity. However, nitrogen content per unit leaf area (N(area)), an important determinant of maximum photosynthetic capacity per unit leaf area (P(max)), decreases as leaves age and senescence. In many species, P(max) decreases in parallel with the leaf nitrogen content. It is hypothesized that if silica accumulation affects photosynthesis, then P(max) would decrease faster than N(area), leading to a decrease in photosynthetic rate per unit leaf nitrogen (photosynthetic nitrogen use efficiency, PNUE) with increasing silica content in leaves. METHODS: The hypothesis was tested in leaves of Sasa veitchii, which have a life span of 2 years and accumulate silica up to 41 % of dry mass. Seasonal changes in P(max), stomatal conductance, N(area) and silica content were measured for leaves of different ages. KEY RESULTS: Although P(max) and PNUE were negatively related with silica content across leaves of different ages, the relationship between PNUE and silica differed depending on leaf age. In second-year leaves, PNUE was almost constant although there was a large increase in silica content, suggesting that leaf nitrogen was a primary factor determining the variation in P(max) and that silica accumulation did not affect photosynthesis. PNUE was strongly and negatively correlated with silica content in third-year leaves, suggesting that silica accumulation affected photosynthesis of older leaves. CONCLUSIONS: Silica accumulation in long-lived leaves of bamboo did not affect photosynthesis when the silica concentration of a leaf was less than 25 % of dry mass. Silica may be actively transported to epidermal cells rather than chlorenchyma cells, avoiding inhibition of CO2 diffusion from the intercellular space to chloroplasts. However, in older leaves with a larger silica content, silica was also deposited in chlorenchyma cells, which may relate to the decrease in PNUE.     

Growth response of barley and tomato to nitrogen stress and its control by abscisic acid,water relations and photosynthesis

Barley (Hordeum vulgare L.) and tomato Lycopersicon esculentum Mill.) were grown hydroponically and examined 2, 5, and 10 d after being deprived of nitrogen (N) supply. Leaf elongation rate declined in both species in response to N stress before there was any reduction in rate of dryweight accumulation. Changes in water transport to the shoot could not explain reduced leaf elongation in tomato because leaf water content and water potential were unaffected by N stress at the time leaf elongation began to decline. Tomato maintained its shoot water status in N-stressed plants, despite reduced water absorption per gram root, because the decline in root hydraulic conductance with N stress was matched by a decline in stomatal conductance. In barley the decline in leaf elongation coincided with a small (8%) decline in water content per unit area of young leaves; this decline occurred because root hydraulic conductance was reduced more strongly by N stress than was stomatal conductance. Nitrogen stress caused a rapid decline in tissue NO ₃ ^- pools and in NO ₃ ^- flux to the xylem, particularly in tomato which had smaller tissue NO ₃ ^- reserves. Even in barley, tissue NO ₃ ^- reserves were too small and were mobilized too slowly (60% in 2 d) to support maximal growth for more than a few hours. Organic N mobilized from old leaves provided an additional N source to support continued growth of N-stressed plants. Abscisic acid (ABA) levels increased in leaves of both species within 2 d in response to N stress. Addition of ABA to roots caused an increase in volume of xylem exudate but had no effect upon NO ₃ ^- flux to the xylem. After leaf-elongation rate had been reduced by N stress, photosynthesis declined in both barley and tomato. This decline was associated with increased leaf ABA content, reduced stomatal conductance and a decrease in organic N content. We suggest that N stress reduces growth by several mechanisms operating on different time scales: (1) increased leaf ABA content causing reduced cell-wall extensibility and leaf elongation and (2) a more gradual decline in photosynthesis caused by ABA-induced stomatal closure and by a decrease in leaf organic N.Abbreviation and symbols ABA abscisic acid - c_i leaf internal CO₂ concentration - L_p root hydraulic conductance     

Complex adjustments of photosynthetic potentials and internal diffusion conductance to current and previous light availabilities and leaf age in Mediterranean evergreen species Quercus ilex

Mature non-senescent leaves of evergreen species become gradually shaded as new foliage develops and canopy expands, but the interactive effects of integrated light during leaf formation (Q(int)G), current light (Q(int)C) and leaf age on foliage photosynthetic competence are poorly understood. In Quercus ilex L., we measured the responses of leaf structural and physiological variables to Q(int)C and Q(int)G for four leaf age classes. Leaf aging resulted in increases in leaf dry mass per unit area (M(A)), and leaf dry to fresh mass ratio (D(F)) and decreases in N content per dry mass (N(M)). N content per area (N(A)) was independent of age, indicating that decreases in N(M) reflected dilution of leaf N because of accumulation of dry mass (NA = N(M) M(A)). M(A), D(F) and N(A) scaled positively with irradiance, whereas these age-specific correlations were stronger with leaf growth light than with current leaf light. Area-based maximum ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) carboxylase activity (V(cmax)A), capacity for photosynthetic electron transport (J(max)A) and the rate of non-photorespiratory respiration in light (R(d)A) were also positively associated with irradiance. Differently from leaf structural characteristics, for all data pooled, these relationships were stronger with current light with little differences among leaves of different age. Acclimation to current leaf light environment was achieved by light-dependent partitioning of N in rate-limiting proteins. Mass-based physiological activities decreased with increasing leaf age, reflecting dilution of leaf N and a larger fraction of non-photosynthetic N in older leaves. This resulted in age-dependent modification of leaf photosynthetic potentials versus N relationships. Internal diffusion conductance (g(m)) per unit area (g(m)A) increased curvilinearly with increasing irradiance for two youngest leaf age classes and was independent of light for older leaves. In contrast, g(m) per dry mass (g(m)M) was negatively associated with light in current-year leaves. Greater photosynthetic potentials and moderate changes in diffusion conductance resulted in greater internal diffusion limitations of photosynthesis in higher light. Both area- and mass-based g(m) decreased with increasing leaf age. The decrease in diffusion conductance was larger than changes in photosynthetic potentials, leading to larger CO2 drawdown from leaf internal air space to chloroplasts (delta(c)) in older leaves. The increases in diffusion limitations in older leaves and at higher light scaled with age- and light-dependent increases in MA and D(F). Overall, our study demonstrates a large potential of foliage photosynthetic acclimation to changes in leaf light environment, but also highlights enhanced structural diffusion limitations in older leaves that result from leaf structural acclimation to previous rather than to current light environment and accumulation of structural compounds with leaf age.     

A high proportion of blue light increases the photosynthesis capacity and leaf formation rate of Rosa × hybrida but does not affect time to flower opening

Alterations in light quality affect plant morphogenesis and photosynthetic responses but the effects vary significantly between species. Roses exhibit an irradiance‐dependent flowering control but knowledge on light quality responses is scarce. In this study we analyzed, the responses in morphology, photosynthesis and flowering of Rosa × hybrida to different blue (B) light proportions provided by light‐emitting diodes (LED, high B 20%) and high pressure sodium (HPS, low B 5%) lamps. There was a strong morphological and growth effect of the light sources but no significant difference in total dry matter production and flowering. HPS‐grown plants had significantly higher leaf area and plant height, yet a higher dry weight proportion was allocated to leaves than stems under LED. LED plants showed 20% higher photosynthetic capacity (A_max) and higher levels of soluble carbohydrates. The increase in A_max correlated with an increase in leaf mass per unit leaf area, higher stomata conductance and CO₂ exchange, total chlorophyll (Chl) content per area and Chl a/b ratio. LED‐grown leaves also displayed a more sun‐type leaf anatomy with more and longer palisade cells and a higher stomata frequency. Although floral initiation occurred at a higher leaf number in LED, the time to open flowers was the same under both light conditions. Thereby the study shows that a higher portion of B light is efficient in increasing photosynthesis performance per unit leaf area, enhancing growth and morphological changes in roses but does not affect the total Dry Matter (DM) production or time to open flower.     

Optimal nitrogen allocation controls tree responses to elevated CO2

Despite the abundance of experimental data, understanding of forest responses to elevated CO2 is limited. Here I show that a key to previously unexplained production and leaf area responses lies in the interplay between whole-plant nitrogen (N) allocation and leaf photosynthesis. A simple tree growth model, controlled by net growth maximization through optimization of leaf area index (LAI) and plant N, is used to analyse CO2 responses in both young, expanding and closed, steady-state canopies. The responses are sensitive to only two independent parameters, the photosynthetic capacity per leaf N (a) and the fine-root N:leaf N ratio. The model explains observed CO2 responses of photosynthesis, production and LAI in four forest free air CO2 enrichment (FACE) experiments. Insensitivity of LAI except at low LAI, increase in light-use efficiency, and photosynthetic down-regulation (as a result of reduced leaf N per area) at elevated CO2 are all explained through the combined effects on a and leaf quantum efficiency. The model bridges the gap between the understanding of leaf-level and plant-level responses and provides a transparent framework for interpreting and linking structural (LAI) and functional (net primary production (NPP):gross primary production (GPP) ratio, light-use efficiency, photosynthetic down-regulation) responses to elevated CO2.     

Divergent patterns of foliar δ13C and δ15N in Quercus aquifolioides with an altitudinal transect on the Tibetan Plateau: an integrated study based on multiple key leaf functional traits

Aims With a close association with plant water availability, foliar δ 13 C had been investigated extensively in alpine regions; however, foliar δ 15 N has rarely been concurrently used as an indicator of plant nitrogen availability. Due to the positive correlations between leaf nitrogen content and foliar δ 13 C and δ 15 N found in previous studies, we expected that they should show consistent patterns along an altitudinal gradient.Methods To test our hypothesis, we measured foliar δ 13 C and δ 15 N in conjunction with multiple key leaf functional traits of Quercus aquifolioides, a dominant species of alpine forest on the eastern slopes of the Sygera Mountains, southeastern Tibetan Plateau from 2500 to 3800 m.Important findings (i) Contrary to our hypothesis, foliar δ 13 C exhibited a significant positive linear relationship with altitude; in contrast, foliar δ 15 N initially increased and subsequently decreased with altitude, the change in trend occurring around 3300 m. (ii) Our analyses indicated that leaf internal resistance and stomatal conductance, rather than photosynthetic capacity indicated by leaf N concentration, apparently explained the altitudinal variation in foliar δ 13 C, while differences in foliar δ 15 N were likely the result of soil N availability. (iii) Principal component analysis revealed a clear association between δ 13 C and a tradeoff between water loss and carbon gain, indicated by traits related to gas exchange such as leaf thickness, density, stomatal properties. In contrast, the second axis was associated with δ 15 N and nitrogen acquisition strategy in Q. aquifolioides across its altitudinal distribution, represented by traits related to nitrogen concentration and stomata per gram of leaf nitrogen.     

红池坝草地常见物种叶片性状沿海拔梯度的响应特征

叶片性状是决定植物光合能力和羧化能力的关键因素,研究叶片性状在海拔梯度上的变化特征是解释植物对于环境变化的适应策略的重要手段。本文以分布于红池坝(10958′E, 3130′ N)草地的5个常见物种红三叶(Trifolium pratense)、老鹳草(Geranium wilfordii)、紫菀(Aster tataricus)、火绒草(Leontopodium leontopodioides)和绣线菊(Spiraea prunifolia)为研究对象,分析了所有物种(n=56)和不同物种的叶片比叶重(LMA)、叶氮含量(单位面积氮含量Narea、单位重量氮含量Nmass)以及叶片δ13C含量沿海拔梯度(815-2545m)的变化趋势及叶片性状之间的关系。研究结果表明：所有物种样品(n=56)的比叶重(LMA)、Narea和δ13C含量沿海拔梯度的增加呈显著增加趋势;Nmass沿海拔梯度的变化趋势不明显;δ13C含量与LMA、Narea呈现极显著正相关关系;不同物种的叶片性状沿着海拔梯度的响应特征有所不同,绣线菊(S. prunifolia)和老鹳草(G. wilfordii)的叶片性状沿海拔梯度的分布规律与所有物种(n=56)样品分布规律一致,红三叶(T. pratense)、紫菀(A. tataricus)、火绒草(L. leontopodioides)的各叶片性状沿海拔梯度的分布特征有所不同。     

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.     

Evidence that abscisic acid does not regulate a centralized whole-plant response to low soil-resource availability

It has been suggested that abscisic acid (ABA) regulates a centralized response of plants to low soil resource availability that is characterized by decreased shoot growth relative to root growth, decreased photosynthesis and stomatal conductance, and decreased plant growth rate. The hypothesis was tested that an ABA-deficient mutant of tomato (flacca; flc) would not exhibit the same pattern of down-regulation of photosynthesis, conductance, leaf area and growth, as well as increased root/shoot partitioning, as its near isogenic wild-type in response to nitrogen or water deficiency, or at least not exhibit these responses to the same degree. Plants were grown from seed in acid-washed sand and exposed to control, nutrient stress, or water stress treatments. Additionally, exogenous ABA was sprayed onto the leaves of a separate group of flc individuals in each treatment. Growth analysis, based on data from frequent harvests of a few individuals, was used to assess the growth and partitioning responses of plants, and gas exchange characteristics were measured on plants throughout the experiment to examine the response of photosynthesis and stomatal conductance. Differences in growth, partitioning and gas exchange variables were found between flc and wild-type individuals, and both nutrient and water treatments caused significant reductions in relative growth rate (RGR) and changes in biomass partitioning. Only the nutrient treatment caused significant reductions in photosynthetic rates. However, flc and wild-type plants responded identically to nutrient and water stress for all but one of the variables measured. The exception was that flc showed a greater decrease in the relative change in leaf area per unit increase of plant biomass (an estimate of the dynamics of leaf area ratio) in response to nutrient stress—a result that is opposite to that predicted by the centralized stress response model. Furthermore, addition of exogenous ABA to flc did not significantly alter any of the responses to nutrient and water stress that we examined. Although it was clear that ABA regulated short-term stomatal responses, we found no evidence to support a pivotal role for ABA, at least absolute amounts of ABA, in regulating a centralized whole-plant response to low soil resource availability.     

Least-cost input mixtures of water and nitrogen for photosynthesis

In microeconomics, a standard framework is used for determining the optimal input mix for a two-input production process. Here we adapt this framework for understanding the way plants use water and nitrogen (N) in photosynthesis. The least-cost input mixture for generating a given output depends on the relative cost of procuring and using nitrogen versus water. This way of considering the issue integrates concepts such as water-use efficiency and photosynthetic nitrogen-use efficiency into the more inclusive objective of optimizing the input mix for a given situation. We explore the implications of deploying alternative combinations of leaf nitrogen concentration and stomatal conductance to water, focusing on comparing hypothetical species occurring in low- versus high-humidity habitats. We then present data from sites in both the United States and Australia and show that low-rainfall species operate with substantially higher leaf N concentration per unit leaf area. The extra protein reflected in higher leaf N concentration is associated with a greater drawdown of internal CO2, such that low-rainfall species achieve higher photosynthetic rates at a given stomatal conductance. This restraint of transpirational water use apparently counterbalances the multiple costs of deploying high-nitrogen leaves.     

Development of leaf photosynthetic parameters in Betula pendula Roth leaves: correlations with photosystem I density

The global modelling of photosynthesis is based on exact knowledge of the leaf photosynthetic machinery. The capacities of partial reactions of leaf photosynthesis develop at different rates, but it is not clear how the development of photoreactions and the Calvin cycle are co-ordinated. We investigated the development of foliar photosynthesis in the temperate deciduous tree Betula pendula Roth. using a unique integrated optical/gas exchange methodology that allows simultaneous estimation of photosystem I and II (PS I and PS II) densities per leaf area, interphotosystem electron transport activities, and ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) kinetic properties. We combined these measurements with in vitro determinations of Rubisco, soluble protein and chlorophyll contents. We observed a strong increase in leaf photosynthetic capacity in developing leaves per leaf area, as well as per dry mass, that was paralleled by accumulation of leaf Rubisco. Enhanced mesophyll conductance was the outcome of increased carboxylation capacity and increased CO(2) diffusion conductance. However, Rubisco was only partly activated in the leaves, according to in vivo measurements of Rubisco kinetics. The amount of active Rubisco increased in proportion with development of PS I, probably through a direct link between Rubisco activase and PS I electron transport. Since the kinetics for post-illumination P700 re-reduction did not change, the synthesis of cytochrome b(6)f complex was also proportional to PS I. The synthesis of PS II began later and continued for several days after reaching the full PS I activity, but leaf chlorophyll was shared equally between the photosystems. Due to this, the antenna of PS II was very large and not optimally organized, leading to greater losses of excitation and lower quantum yields in young leaves. We conclude that co-ordinated development of leaf photosynthesis is regulated at the level of PS I with subordinated changes in PS II content and Rubisco activation.     

Photosynthetic and biomass allocation responses of Liquidambar styraciflua (Hamamelidaceae) to vine competition

This 2-year field study examined stomatal conductance, photosynthesis, and biomass allocation of Liquidambar styraciflua saplings in response to below- and aboveground competition with the vines Lonicera japonica and Parthenocissus quinquefolia. Vine competition did not affect stomatal conductance of the host trees. The leaf photosynthetic capacity and photosynthetic nitrogen-use efficiency were significantly reduced by root competition with vines, either singly or in combination with aboveground competition, early in the second growing season. However, such differences disappeared by the end of the second growing season. Trees competing below ground with vines also had lower allocation to leaves compared with steins. Aboveground competition with vines resulted in reduced photosynthetic capacity per unit leaf area, but not per unit leaf weight, in trees. No correlation was found between single leaf photosynthetic capacity and tree growth. In contrast, a high positive correlation existed between allocation to leaves and diameter growth. Results from this study suggest that allocation patterns are more affected than leaf photosynthesis in trees competing with vines.