Effect of soil moisture on leaf ecophysiology of <Emphasis Type="Italic">Parasenecio yatabei</Emphasis>, a summer-green herb in a cool–temperate forest understory in Japan

Leaf physiological and gas-exchange traits of a summer-green herbaceous perennial, Parasenecio yatabei, growing along a stream were examined in relation to leaf age. In its vegetative phase, the aerial part of this plant consists of only one leaf and provides an ideal system for the study of leaf longevity. Volumetric soil water content (SWC) decreased with increasing distance from the stream, whereas relative light intensity was nearly constant. The light-saturated net CO₂ assimilation rate (A _sat) and leaf stomatal conductance (gs) were approximately 1.5-fold and 1.4-fold higher, respectively, in the lower slope near the mountain stream than in the upper slope far from the mountain stream. The lifespan of aerial parts of vegetative plants significantly increased with decreasing SWC. The leaf mass-based nitrogen content of the leaves (N _mass) was almost constant (ca. 2.2%); however, the maximum carboxylation rate by ribulose-1,5-biphosphate carboxylase/oxygenase (rubisco) (V _cmax) and photosynthetic nitrogen use efficiency (PNUE, A _sat/N _area) decreased more slowly in the upper slope than in the lower slope. The higher leaf photosynthetic activity of P. yatabei plants growing lower on the slope leads to a decrease in V _cmax and PNUE in the early growing season, and to a shorter leaf lifespan.     

Photosynthetic acclimation to elevated CO2 in Phaseolus vulgaris L. is altered by growth response to nitrogen supply

Abstract Long‐term exposure of plants to elevated CO₂ often leads to downward photosynthetic acclimation. Nitrogen (N) deficiency could potentially exacerbate this response by reducing growth rate and the sink for photosynthates, but this has not always been observed. Experimentally, the interpretation of N effects on CO₂ responses can be confounded by increasing severity of tissue N deficiency over time when N supply is not adjusted as demand increases. In this study, N supply ranged from sub‐ to supra‐optimal (20–540 kgN ha^–l equivalent), and relatively stable levels of tissue N concentration were obtained in all treatments by varying twice‐weekly application rates in proportion to plant growth. The effects of N on photosynthesis and growth of beans (Phaseolus vulgaris L.) raised at ambient (35 Pa) and three elevated (70, 105, 140 Pa) CO₂ partial pressures (pCO₂) were evaluated. Averaging across N treatments, leaf total non‐structural carbohydrates (TNC) were 2.5‐ to 3‐fold higher and leaf N concentrations were 31–35% lower at elevated compared to ambient pCO₂. Light‐saturated net CO₂ assimilation rates measured at growth pCO₂ (A_satg) were significantly higher (26–40% depending on N supply) in plants grown at elevated compared to ambient pCO₂. When measured at a common pCO₂ of 35 Pa, the A_sat of plants grown at elevated CO₂ was 15–29% less than that of plants grown at 35 Pa, indicative of downward photosynthetic acclimation. The magnitude of downward photosynthetic acclimation to elevated CO₂ was greater in plants grown at high (180 and 540 kgN ha^–l) compared to low (20 and 60 kgN ha^–l) N supply, and this was associated with a higher A_sat at growth pCO₂, higher leaf area ratio (leaf area/total biomass), and higher TNC in leaves of high‐N plants. Our results indicate that the effect of N on acclimation to CO₂ will depend on the balance between supply and demand for N during the growing period, and the effect this has on biomass allocation and source‐sink C balance at the whole‐plant level.     

Differential coordination of stomatal conductance,mesophyll conductance,and leaf hydraulic conductance in response to changing light across species

Stomatal conductance (g_s) and mesophyll conductance (g_m) represent major constraints to photosynthetic rate (A), and these traits are expected to coordinate with leaf hydraulic conductance (K_leaf) across species, under both steady‐state and dynamic conditions. However, empirical information about their coordination is scarce. In this study, K_leaf, gas exchange, stomatal kinetics, and leaf anatomy in 10 species including ferns, gymnosperms, and angiosperms were investigated to elucidate the correlation of H₂O and CO₂ diffusion inside leaves under varying light conditions. Gas exchange, K_leaf, and anatomical traits varied widely across species. Under light‐saturated conditions, the A, g_s, g_m, and K_leaf were strongly correlated across species. However, the response patterns of A, g_s, g_m, and K_leaf to varying light intensities were highly species dependent. Moreover, stomatal opening upon light exposure of dark‐adapted leaves in the studied ferns and gymnosperms was generally faster than in the angiosperms; however, stomatal closing in light‐adapted leaves after darkening was faster in angiosperms. The present results show that there is a large variability in the coordination of leaf hydraulic and gas exchange parameters across terrestrial plant species, as well as in their responses to changing light.     

Photosynthetic rates in relation to leaf phosphorus content in pioneer versus climax tropical rainforest trees

In Guyana dense rainforest occurs on intensely weathered acid soils, low in soil phosphorus. To investigate whether low P availability limits photosynthesis of trees growing on these soils more than N does, leaf P and N content, and their relationship with the photosynthetic capacity (A _sat, mol CO₂ m^-2 s^-1) were studied for nine pioneer and climax tree species in a range of light climates. Light environment was described using hemispherical photographs. For both pioneer and climax species, leaf P content (r ²=0.71 and 0.23, respectively) is a more important determinant of A _sat than leaf N content (r ²=0.54 and 0.12, respectively). Pioneer species have a higher leaf P and N content than climax species. At similar P or N content, pioneers have a higher A _sat than climax species. The saplings studied had a relatively high A _sat, considering their low P concentration (15–30 mol P g^-1). All species studied had a constant leaf P and N concentration and photosynthetic capacity across light climates, because specific leaf mass (g m^-2) increased similarly with light availability. This acclimation to a change in light environment makes a possible limitation of A _sat by P or N independent of light environment.     

Photosynthesis, carboxylation and leaf nitrogen responses of 16 species to elevated pCO2 across four free-air CO2 enrichment experiments in forest, grassland and desert

The magnitude of changes in carboxylation capacity in dominant plant species under long‐term elevated CO₂ exposure (elevated pC_a) directly impacts ecosystem CO₂ assimilation from the atmosphere. We analyzed field CO₂ response curves of 16 C₃ species of different plant growth forms in favorable growth conditions in four free‐air CO₂ enrichment (FACE) experiments in a pine and deciduous forest, a grassland and a desert. Among species and across herb, tree and shrub growth forms there were significant enhancements in CO₂ assimilation (A) by +40±5% in elevated pC_a (49.5–57.1 Pa), although there were also significant reductions in photosynthetic capacity in elevated pC_a in some species. Photosynthesis at a common pC_a (A_a) was significantly reduced in five species growing under elevated pC_a, while leaf carboxylation capacity (V_cmax) was significantly reduced by elevated pC_a in seven species (change of ?19±3% among these species) across different growth forms and FACE sites. Adjustments in V_cmax with elevated pC_a were associated with changes in leaf N among species, and occurred in species with the highest leaf N. Elevated pC_a treatment did not affect the mass‐based relationships between A or V_cmax and N, which differed among herbs, trees and shrubs. Thus, effects of elevated pC_a on leaf C assimilation and carboxylation capacity occurred largely through changes in leaf N, rather than through elevated pC_a effects on the relationships themselves. Maintenance of leaf carboxylation capacity among species in elevated pC_a at these sites depends on maintenance of canopy N stocks, with leaf N depletion associated with photosynthetic capacity adjustments. Since CO₂ responses can only be measured experimentally on a small number of species, understanding elevated CO₂ effects on canopy N_m and N_a will greatly contribute to an ability to model responses of leaf photosynthesis to atmospheric CO₂ in different species and plant growth forms.     

Photosynthetic nitrogen-use efficiency of species that differ inherently in specific leaf area

Factors that contribute to interspecific variation in photosynthetic nitrogen-use efficiency (PNUE, the ratio of CO₂ assimilation rate to leaf organic nitrogen content) were investigated, comparing ten dicotyledonous species that differ inherently in specific leaf area (SLA, leaf area:leaf dry mass). Plants were grown hydroponically in controlled environment cabinets at two irradiances (200 and 1000 μmol m^–2 s^–1). CO₂ and irradiance response curves of photosynthesis were measured followed by analysis of the chlorophyll, Rubisco, nitrate and total nitrogen contents of the leaves. At both irradiances, SLA ranged more than twofold across species. High-SLA species had higher in situ rates of photosynthesis per unit leaf mass, but similar rates on an area basis. The organic N content per unit leaf area was lower for the high-SLA species and consequently PNUE at ambient light conditions (PNUE_amb) was higher in those plants. Differences were somewhat smaller, but still present, when PNUE was determined at saturating irradiances (PNUE_max). An assessment was made of the relative importance of the various factors that underlay interspecific variation in PNUE. For plants grown under low irradiance, PNUE_amb of high-SLA species was higher primarily due to their lower N content per unit leaf area. Low-SLA species clearly had an overinvestment in photosynthetic N under these conditions. In addition, high SLA-species allocated a larger fraction of organic nitrogen to thylakoids and Rubisco, which further increased PNUE_amb. High-SLA species grown under high irradiance showed higher PNUE_amb mainly due to a higher Rubisco specific activity. Other factors that contributed were again their lower contents of N_org per unit leaf area and a higher fraction of photosynthetic N in electron transport and Rubisco. For PNUE_max, differences between species in organic leaf nitrogen content per se were no longer important and higher PNUE_max of the high SLA species was due to a higher fraction of N in␣photosynthetic compounds (for low-light plants) and a higher Rubisco specific activity (for high-light grown plants). Received: 11 October 1997 / Accepted: 9 April 1998     

Contrasting growth response of an N2-fixing and non-fixing forb to elevated CO2: dependence on soil N supply

With the ability to symbiotically fix atmospheric N₂, legumes may lack the N-limitations thought to constrain plant response to elevated concentrations of atmospheric CO₂. The growth and photosynthetic responses of two perennial grassland species were compared to test the hypotheses that (1) the CO₂ response of wild species is limited at low N availability, (2) legumes respond to a greater extent than non-fixing forbs to elevated CO₂, and (3) elevated CO₂ stimulates symbiotic N₂ fixation, resulting in an increased amount of N derived from the atmosphere. This study investigated the effects of atmospheric CO₂ concentration (365 and 700 mol mol^–1) and N addition on whole plant growth and C and N acquisition in an N₂-fixing legume (Lupinus perennis) and a non-fixing forb (Achillea millefolium) in controlled-chamber environments. To evaluate the effects of a wide range of N availability on the CO₂ response, we incorporated six levels of soil N addition starting with native field soil inherently low in N (field soil + 0, 4, 8, 12, 16, or 20 g N m^–2 yr^–1). Whole plant growth, leaf net photosynthetic rates (A), and the proportion of N derived from N₂ fixation were determined in plants grown from seed over one growing season. Both species increased growth with CO₂enrichment, but this response was mediated by N supply only for the non-fixer, Achillea. Its response depended on mineral N supply as growth enhancements under elevated CO₂ increased from 0% in low N soil to +25% at the higher levels of N addition. In contrast, Lupinus plants had 80% greater biomass under elevated CO₂ regardless of N treatment. Although partial photosynthetic acclimation to CO₂ enrichment occurred, both species maintained comparably higher A in elevated compared to ambient CO₂ (+38%). N addition facilitated increased A in Achillea, however, in neither species did additional N availability affect the acclimation response of A to CO₂. Elevated CO₂ increased plant total N yield by 57% in Lupinus but had no effect on Achillea. The increased N in Lupinus came from symbiotic N₂ fixation, which resulted in a 47% greater proportion of N derived from fixation relative to other sources of N. These results suggest that compared to non-fixing forbs, N₂-fixers exhibit positive photosynthetic and growth responses to increased atmospheric CO₂ that are independent of soil N supply. The enhanced amount of N derived from N₂ fixation under elevated CO₂ presumably helps meet the increased N demand in N₂-fixing species. This response may lead to modified roles of N₂-fixers and N₂-fixer/non-fixer species interactions in grassland communities, especially those that are inherently N-poor, under projected rising atmospheric CO₂.     

Community succession and photosynthetic physiological characteristics of pasture plants in a sub-alpine meadow in Gannan,China

 为了解甘南亚高山草甸围封地恢复演替动态, 探究围封恢复进程中植物光合生理特征的变化规律及其影响因子, 对围封试验地内5个典型群落样地进行样方调查, 测定了各群落优势种及3个共有种的光合参数和叶性状参数, 并测定了群落表层土壤(0–20 cm)的水分含量及全氮含量。结果显示: 该围封地内形成一个以草本植物→半灌木→灌木的演替序列, 群落表层土壤含水量及全氮含量随着演替的进行逐渐增加; 在演替的时间尺度上, 各演替阶段优势种光合生理特征间存在明显差异, 随着演替的进行, 群落优势种的净光合速率(A_area)、光合水分利用效率(WUE)、相对叶绿素含量(SPAD)呈降低趋势, 其叶片氮含量(N_mass)、光合氮利用效率(PNUE)、比叶面积(SLA)随演替变化没有严格一致的规律, 而更多地表现为不同植物功能型之间的差异; 从演替前期到后期, 同种植物的A_area、SPAD值逐渐降低, 非豆科植物披碱草(Elymus dahuricus)、老鹳草(Geranium wilfordii)的PNUE、WUE随演替进行呈降低趋势, 其N_mass、SLA随演替进行却呈增加趋势, 然而豆科植物紫苜蓿(Medicago sativa)由于具有固氮能力, 受养分限制不明显, 这些光合生理特征值没有随演替发生明显的变化。这些结果表明, 在恢复演替过程中, 该围封地由一个物质获得能力强的群落向物质保持能力强的群落过渡, 土壤水分含量及全氮含量是推动这种过渡发生的主要因子。掌握围封地群落演替过程中的光合生理动态对于亚高寒退化草甸恢复具有一定的理论指导意义。     

不同组成群落3种共有植物光合生理特征研究

为研究物种在不同群落中光合生理特征的变化,以亚高寒草甸围封恢复地为研究对象,对样地内3个不同组成群落进行样方调查,测定了物种高度及各群落垂直方向上光照强度以及群落中3个共有种披碱草(Elymus dahuricus)、刺儿菜(Cirsium setosum)和紫花苜蓿(Medicago sativa)的净光合速率(Aarea)、叶片氮含量(Nmass)、比叶重(LMA)及光合氮利用效率(PNUE)。结果表明:(1)3个样地的群落组成有明显的差异,豆科植物的增多可以一定程度上改善群落氮养分状况,但植物叶片Nmass还受到群落优势种竞争的影响。(2)同一物种在不同群落的高度不同,不同群落垂直方向上光照强度也不相同,导致同一物种在不同群落中能够获得的光照强度有一定差异。(3)在养分、光照强度有差异的情况下,不同植物的Aarea、LMA及PNUE在不同群落中的变化趋势不尽相同,而Narea与Aarea的关系在总体上、群落间及物种间变化不大,基本上显示了较强的正相关关系。由此可见,群落组成、结构引起的光照及氮素差异是导致同一物种光合生理特征在不同群落中变化的重要因素,但不同物种光合生理特征对光照及氮素变化的响应不同。     

Acclimation response of spring wheat in a free-air CO2 enrichment (FACE) atmosphere with variable soil nitrogen regimes. 1. Leaf position and phenology determine acclimation response

We have examined the photosynthetic acclimation of wheat leaves grown at an elevated CO₂ concentration, and ample and limiting N supplies, within a field experiment using free-air CO₂ enrichment (FACE). To understand how leaf age and developmental stage affected any acclimation response, measurements were made on a vertical profile of leaves every week from tillering until maturity. The response of assimilation (A) to internal CO₂ concentration (C_i) was used to estimate the in vivo carboxylation capacity (Vc_max) and maximum rate of ribulose-1,5-bisphosphate limited photosynthesis (A _sat). The total activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), and leaf content of Rubisco and the Light Harvesting Chlorophyll a/b protein associated with Photosystem II (LHC II), were determined. Elevated CO₂ did not alter Vc_max in the flag leaf at either low or high N. In the older shaded leaves lower in the canopy, acclimatory decline in Vc_max and A _sat was observed, and was found to correlate with reduced Rubisco activity and content. The dependency of acclimation on N supply was different at each developmental stage. With adequate N supply, acclimation to elevated CO₂ was also accompanied by an increased LHC II/Rubisco ratio. At low N supply, contents of Rubisco and LHC II were reduced in all leaves, although an increased LHC II/Rubisco ratio under elevated CO₂ was still observed. These results underscore the importance of leaf position, leaf age and crop developmental stage in understanding the acclimation of photosynthesis to elevated CO₂ and nutrient stress. This revised version was published online in June 2006 with corrections to the Cover Date.     

Interspecific difference in the photosynthesis–nitrogen relationship: patterns,physiological causes,and ecological importance

The photosynthesis–nitrogen relationship is significantly different among species. Photosynthetic capacity per unit leaf nitrogen, termed as photosynthetic nitrogen-use efficiency (PNUE), has been considered an important leaf trait to characterise species in relation to their leaf economics, physiology, and strategy. In this review, I discuss (1) relations between PNUE and species ecology, (2) physiological causes and (3) ecological implications of the interspecific difference in PNUE. Species with a high PNUE tend to have high growth rates and occur in disturbed or high productivity habitats, while those with a low PNUE occur in stressful or low productivity habitats. PNUE is an important leaf trait that correlates with other leaf traits, such as leaf mass per area (LMA) and leaf life span, irrespective of life form, phylogeny, and biomes. Various factors are involved in the interspecific difference. In particular, nitrogen allocation within leaves and the mesophyll conductance for CO₂ diffusion are important. To produce tough leaves, plants need to allocate more biomass and nitrogen to make thick cell walls, leading to a reduction in the mesophyll conductance and in nitrogen allocation to the photosynthetic apparatus. Allocation of biomass and nitrogen to cell walls may cause the negative relationship between PNUE and LMA. Since plants cannot maximise both PNUE and leaf toughness, there is a trade-off between photosynthesis and persistence, which enables the existence of species with various leaf characteristics on the earth.     

Responses of leaf nitrogen concentration and specific leaf area to atmospheric CO2 enrichment: a retrospective synthesis across 62 species

Knowledge of leaf responses to elevated atmospheric [CO₂] (CO₂ concentration) is integral to understanding interactions between vegetation and global change. This work deals with responses of leaf mass‐based nitrogen concentration (N_m) and specific leaf area (SLA). It assesses the statistical significance of factors perceived as influential on the responses, and quantifies how the responses vary with the significant factors identified, based on 170 data cases of 62 species compiled from the literature. Resultant equations capture about 41% of the variance in the data for percent responses of N_m and SLA, or about 95% of the variance for N_m and SLA at 57–320% normal [CO₂]; these performance statistics also hold for leaf area‐based N concentration and specific leaf weight. The equations generalize that: (i) both N_m and SLA decline as [CO₂] increases; (ii) proportional decline of N_m is greater with deciduous woody species and with plants of normally low N_m, increases with pot size in growth chamber and greenhouse settings and with temperature and photosynthetic photon flux density (PPFD), and is mitigated by N fertilization; and (iii) proportional decline of SLA depends on pot size and PPFD similarly to N_m, increases with leaf life span and water vapour pressure deficit in enclosed experiments, and decreases with prolonged exposure to elevated [CO₂] among broadleaf woody species in field conditions. The results highlight great uncertainty in the percent‐response data and reveal the potential feasibility to estimate N_m and SLA at various magnitudes of elevated [CO₂] from a few key plant and environmental factors of broad data bases.     

The photosynthesis – leaf nitrogen relationship at ambient and elevated atmospheric carbon dioxide: a meta-analysis

Estimation of leaf photosynthetic rate (A) from leaf nitrogen content (N) is both conceptually and numerically important in models of plant, ecosystem, and biosphere responses to global change. The relationship between A and N has been studied extensively at ambient CO₂ but much less at elevated CO₂. This study was designed to (i) assess whether the A–N relationship was more similar for species within than between community and vegetation types, and (ii) examine how growth at elevated CO₂ affects the A–N relationship. Data were obtained for 39 C3 species grown at ambient CO₂ and 10 C3 species grown at ambient and elevated CO₂. A regression model was applied to each species as well as to species pooled within different community and vegetation types. Cluster analysis of the regression coefficients indicated that species measured at ambient CO₂ did not separate into distinct groups matching community or vegetation type. Instead, most community and vegetation types shared the same general parameter space for regression coefficients. Growth at elevated CO₂ increased photosynthetic nitrogen use efficiency for pines and deciduous trees. When species were pooled by vegetation type, the A–N relationship for deciduous trees expressed on a leaf-mass basis was not altered by elevated CO₂, while the intercept increased for pines. When regression coefficients were averaged to give mean responses for different vegetation types, elevated CO₂ increased the intercept and the slope for deciduous trees but increased only the intercept for pines. There were no statistical differences between the pines and deciduous trees for the effect of CO₂. Generalizations about the effect of elevated CO₂ on the A–N relationship, and differences between pines and deciduous trees will be enhanced as more data become available.     

Quantifying the response of photosynthesis to changes in leaf nitrogen content and leaf mass per area in plants grown under atmospheric CO2 enrichment

Previous modelling exercises and conceptual arguments have predicted that a reduction in biochemical capacity for photosynthesis (A_area) at elevated CO₂ may be compensated by an increase in mesophyll tissue growth if the total amount of photosynthetic machinery per unit leaf area is maintained (i.e. morphological upregulation). The model prediction was based on modelling photosynthesis as a function of leaf N per unit leaf area (N_area), where N_area = N_mass×LMA. Here, N_mass is percentage leaf N and is used to estimate biochemical capacity and LMA is leaf mass per unit leaf area and is an index of leaf morphology. To assess the relative importance of changes in biochemical capacity versus leaf morphology we need to control for multiple correlations that are known, or that are likely to exist between CO₂ concentration, N_area, N_mass, LMA and A_area. Although this is impractical experimentally, we can control for these correlations statistically using systems of linear multiple-regression equations. We developed a linear model to partition the response of A_area to elevated CO₂ into components representing the independent and interactive effects of changes in indexes of biochemical capacity, leaf morphology and CO₂ limitation of photosynthesis. The model was fitted to data from three pine and seven deciduous tree species grown in separate chamber-based field experiments. Photosynthetic enhancement at elevated CO₂ due to morphological upregulation was negligible for most species. The response of A_area in these species was dominated by the reduction in CO₂ limitation occurring at higher CO₂ concentration. However, some species displayed a significant reduction in potential photosynthesis at elevated CO₂ due to an increase in LMA that was independent of any changes in N_area. This morphologically based inhibition of A_area combined additively with a reduction in biochemical capacity to significantly offset the direct enhancement of A_area caused by reduced CO₂ limitation in two species. This offset was 100% for Acer rubrum, resulting in no net effect of elevated CO₂ on A_area for this species, and 44% for Betula pendula. This analysis shows that interactions between biochemical and morphological responses to elevated CO₂ can have important effects on photosynthesis.     

Productivity and thallus toughness trade‐off relationship in marine macroalgae from the Japan Sea

Trade‐offs are considered key to understanding mechanisms supporting the coexistence of multiple plant species. Thus, understanding the mechanisms underlying trade‐offs is expected to contribute to conservation and management of macroalgal beds composed of diverse macroalgae of rocky shore ecosystems. To test the occurrence of trade‐offs between productivity and thallus toughness as well as pair‐wise thallus trait relationships that are expected to indirectly relate to any trade‐offs, traits and relationships for 13 species of macroalgae from the central area along the Japan Sea coast of Honshu, Japan were examined. In each species we examined for photosynthetic capacity per unit biomass (as A_mass) and nitrogen (i.e., photosynthetic nitrogen‐use efficiency, PNUE), nitrogen content (as N_mass), thallus mass per unit thallus area (as TMA) and force required to penetrate the thallus (as F_p, a common index of leaf toughness in land plants by punch test). A significant negative correlation indicating a trade‐off between productivity and thallus toughness was found between A_mass or PNUE and F_p. Pair‐wise relationships that were expected to indirectly relate to the trade‐off were as follows. A_mass was positively correlated with N_mass. Thalli with high N_mass extensively utilizing nitrogen in the photosynthetic parts, and consequently exhibiting elevated metabolic rates. Moreover, thalli with high N_mass tended to be associated with low TMA, and N_mass decreased with increasing TMA. A significant negative correlation was observed between TMA and A_mass or PNUE because of the linkage of high A_mass or PNUE with high N_mass and high N_mass associated with low TMA, while a significant positive correlation was observed between TMA and F_p. The two correlations indicate a physiological and structural trade‐off, which underlies the interdependency of thallus traits. Results of multivariate analyses also indicated that the thallus traits interdependently vary across a single axis based on the trade‐off.     

Photosynthesis–nitrogen relationships in species at different altitudes on Mount Kinabalu, Malaysia

In situ photosynthetic nitrogen-use efficiency (PNUE, photosynthetic capacity per unit leaf nitrogen) was investigated in species that commonly distributed at different altitudes (600–3700m above sea level) on Mount Kinabalu. Photosynthetic nitrogen-use efficiency was lower in species at higher altitudes, with a mean PNUE at 3700m being one-third as large as that at 600m. This difference in PNUE was larger than that explained by the biochemical response to lower air pressures only. Across altitudes a negative correlation between ¹³C abundance (¹³C) and PNUE was found. Species at higher altitudes tended to have higher ¹³C, suggesting that they had a lower conductance for CO₂ diffusion from the air to chloroplasts. The lower conductance might be responsible for the lower PNUE in species at higher altitudes. Although leaf nitrogen content per unit area tended to be higher at higher altitudes, it did not seem to contribute to increasing photosynthetic rates. Thus, the idea that a higher nitrogen content at higher altitudes is a compensation for a lower PNUE was not supported. In contrast to the large difference in PNUE among altitudes, PNUE tended to converge within a narrow range among species growing at the same altitude.     

Acclimation of photosynthesis to irradiance and spectral quality in British plant species: chlorophyll content, photosynthetic capacity and habitat preference

Twenty-two common British angiosperms were examined for their ability to acclimate photosynthetically to sun and shade conditions. Plants were grown under low irradiance, far-red enriched light (50 μmol m^?2 s^?1), selected to mimic as closely as possible natural canopy shade, and moderately high light of insufficient irradiance to induce photoinhibitory or photoprotective responses (300 μmol m^?2 s^?1). Light-and CO²-saturated photosynthetic rates of oxygen evolution (P_max) and chlorophyll content were measured. Large variation was found in both parameters, and two ‘strategies’ for long-term acclimation were identified: firstly a change in chlorophyll per unit leaf area which was found to correlate positively with photosynthetic capacity, and secondly changes in chlorophyll alb ratio and P_max, indicative of alterations at the chloroplast level, which were not associated with a change in chlorophyll content per unit leaf area. Combinations of these two strategies may occur, giving rise to the observed diversity in photosynthetic acclimation. The extent and nature of photosynthetic acclimation were compared with an index of shade association, calculated from the association each species has with woodland. It was found that the greatest flexibility for change at the chloroplast level was found in those species possessing an intermediate shade association, whilst acclimation in ‘sun’ species proceeded by a change in chlorophyll content; obligate shade species showed little capacity for acclimation at either the chloroplast or leaf level. A framework for explaining the variation between plant species in leaf-level photosynthetic capacity, in relation to the natural light environment, is presented. This is the first time the potential for light acclimation of photosynthesis in different plant species has been satisfactorily linked to habitat distribution.     

Diurnal and seasonal variations in photosynthetic and morphological traits of the tree ferns Dicksonia antarctica (Dicksoniaceae) and Cyathea australis (Cyatheaceae) in wet sclerophyll forests of Australia

Steady state and dynamic responses of two tree fern species of contrasting origins, Dicksonia antarctica (of Gondwanan origin) and Cyathea australis (Pan-tropical), were studied over two consecutive years under field conditions in a wet sclerophyll forest of south-east Australia. Irrespective of their different origins, there were no significant differences in photosynthetic performance between the two species. Growth irradiance and leaf temperature, but not plant water status, was significantly related to photosynthetic and morphological traits. At a common leaf temperature, maximum light-use efficiency of photosystem II (F_v/F_m) was significantly lower in winter than in summer, suggesting some limitation to PSII efficiency potentially associated with cold winter mornings. Both species displayed seasonal acclimation in a number of measured photosynthetic parameters and frond traits (i.e. F_v/F_m, A_sat, g_s, N_A, total chlorophyll, SLA). Acclimation of stomatal density to spatial variation in growth irradiance seemed limited in both species, although stomatal pattern differed between species. Because there were no significant differences between the two species in photosynthetic parameters, both species can be described by common carbon gain and water use models at the leaf scale.     

Photosynthetic nitrogen and water use efficiency of acacia and eucalypt seedlings as afforestation species

The ecophysiological traits of acacia and eucalypt are important in assessing their suitability for afforestation. We measured the gas-exchange rate, the leaf dry mass per area (LMA) and the leaf nitrogen content of two acacia and four eucalypt species. Relative to the eucalypts, the acacias had lower leaf net photosynthetic rate (P _N), lower photosynthetic nitrogen-use efficiency (PNUE), higher water-use efficiency (WUE), higher LMA and higher leaf nitrogen per unit area (N _area). No clear differences were observed within or between genera in the maximum rate of carboxylation (V _cmax) or the maximum rate of electron transport (J _max), although these parameters tended to be higher in eucalypts. PNUE and LMA were negatively correlated. We conclude that acacias with higher LMA do not allocate nitrogen efficiently to photosynthetic system, explaining why their P _N and PNUE were lower than in eucalypts.     

Irradiance acclimation,capture ability,and efficiency in invasive and non-invasive alien plant species

We tested the hypothesis that invasive (IN) species could capture resources more rapidly and efficiently than noninvasive (NIN) species. Two IN alien species, Ageratina adenophora and Chromolaena odorata, and one NIN alien species, Gynura sp. were compared at five irradiances. Photon-saturated photosynthetic rate (P _max), leaf mass (LMA) and nitrogen content (N_A) per unit area, and photosynthetic nitrogen utilization efficiency (PNUE) increased significantly with irradiance. LMA, N_A, and PNUE all contributed to the increased P _max, indicating that both morphological and physiological acclimation were important for the three alien species. Under stronger irradiance, PNUE was improved through changes in N allocation. With the increase of irradiance, the amount of N converted into carboxylation and bioenergetics increased, whereas that allocated to light-harvesting components decreased. The three alien species could adequately acclimate to high irradiance by increasing the ability to utilize and dissipate photon energy and decreasing the efficiency of photon capture. The two IN species survived at 4.5 % irradiance while the NIN species Gynura died, representing their different invasiveness. Ageratina generally exhibited higher respiration rate (R _D) and N_A. However, distinctly higher P _max, PNUE, P _max/R _D, or P _max/LMA were not detected in the two invasive species, nor was lower LMA. Hence the abilities to capture and utilize resources were not always associated with invasiveness of the alien species.