Short-term effects of fertilization on loblolly pine (Pinus taeda L.) physiology

Fertilization commonly increases biomass production in loblolly pine (Pinus taeda L.). However, the sequence of short‐term physiological adjustments allowing for the establishment of leaf area and enhanced growth is not well understood. The effects of fertilization on photosynthetic parameters, root respiration, and growth for over 200 d following the application of diammonium phosphate were intensively investigated in an effort to establish a relative sequence of events associated with improved growth. Root respiration, foliar nitrogen concentration [N]_f, and light‐saturated net photosynthesis (A_sat) temporarily increased following fertilization. A_sat was correlated positively with [N]_f when non‐fertilized and fertilized treatments were pooled (R² = 0.47). Increased photosynthetic capacity following fertilization was due to both improved photochemical efficiency and capacity and enhanced carboxylation capacity of Rubisco. Positive effects of fertilization on growth were observed shortly after A_sat increased. Fertilized seedlings had 36.5% more leaf area and 36.5% greater total dry weight biomass at 211 d following fertilization. It is concluded that fertilization temporarily increased photosynthetic capacity, which resulted in a pool of photo‐assimilate used to build leaf area. The N from fertilizer initially invested in photosynthetic structures and enzymes probably re‐translocated to newly developing foliage, explaining the reduction in [N]_f and A_sat that was observed after peak levels were achieved following fertilization.     

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.     

Responses of the functional traits in Cleistogenes squarrosa to nitrogen addition and drought

植物功能性状被广泛地用于研究植物对环境变化的响应。糙隐子草(Cleistogenes squarrosa)是内蒙古草原重要的C4物种,其功能性状是如何对水氮环境的变化做出响应的,还不十分清楚。该文采用盆栽实验的方法,进行氮添加(0,10.5,35.0和56.0 g·m–2·a–1)和降水(自然降水和70%平均月降水量)处理,研究糙隐子草整株性状、叶形态性状和叶生理性状对氮添加和干旱的响应。结果表明,氮添加显著影响了糙隐子草的整株性状,氮、水处理及它们的交互作用显著影响了糙隐子草的叶形态性状和叶生理性状。各功能性状对氮添加的响应格局在自然降水和干旱处理下是不同的。根深、茎生物量和茎叶比在干旱条件下低和中氮添加处理中较高,而在自然降水下无明显变化;比叶面积在干旱条件下随氮添加量的增加而增加,而在自然降水下无增加趋势;自然降水下,高氮添加显著刺激了光合速率和蒸腾速率,增加了水分利用效率,而在干旱条件下氮添加对它们没有显著影响;叶片单位面积的氮含量在自然降水下随氮添加量的增加有增加趋势,而在干旱条件下显著降低。在自然降水下,氮添加主要影响糙隐子草的叶形态和生理性状,而在干旱条件下,氮添加主要影响糙隐子草的整株性状和形态性状。总之,糙隐子草的功能性状对氮添加表现出明显的响应,响应格局在不同的水分条件下不同,反映了其对氮水环境变化的弹性适应。     

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.     

Temperature and precipitation controls over leaf‐ and ecosystem‐level CO2 flux along a woody plant encroachment gradient

Conversion of grasslands to woodlands may alter the sensitivity of CO₂ exchange of individual plants and entire ecosystems to air temperature and precipitation. We combined leaf‐level gas exchange and ecosystem‐level eddy covariance measurements to quantify the effects of plant temperature sensitivity and ecosystem temperature responses within a grassland and mesquite woodland across seasonal precipitation periods. In so doing, we were able to estimate the role of moisture availability on ecosystem temperature sensitivity under large‐scale vegetative shifts. Optimum temperatures (T_opt) for net photosynthetic assimilation (A) and net ecosystem productivity (NEP) were estimated from a function fitted to A and NEP plotted against air temperature. The convexities of these temperature responses were quantified by the range of temperatures over which a leaf or an ecosystem assimilated 50% of maximum NEP (Ω₅₀). Under dry pre‐ and postmonsoon conditions, leaf‐level Ω₅₀ in C₃ shrubs were two‐to‐three times that of C₄ grasses, but under moist monsoon conditions, leaf‐level Ω₅₀ was similar between growth forms. At the ecosystems‐scale, grassland NEP was more sensitive to precipitation, as evidenced by a 104% increase in maximum NEP at monsoon onset, compared to a 57% increase in the woodland. Also, woodland NEP was greater across all temperatures experienced by both ecosystems in all seasons. By maintaining physiological function across a wider temperature range during water‐limited periods, woody plants assimilated larger amounts of carbon. This higher carbon‐assimilation capacity may have significant implications for ecosystem responses to projected climate change scenarios of higher temperatures and more variable precipitation, particularly as semiarid regions experience conversions from C₄ grasses to C₃ shrubs. As regional carbon models, CLM 4.0, are now able to incorporate functional type and photosynthetic pathway differences, this work highlights the need for a better integration of the interactive effects of growth form/functional type and photosynthetic pathway on water resource acquisition and temperature sensitivity.     

Photosynthetic and gas exchange characteristics of dominant woody plants on a moisture gradient in an African savanna

We determined key photosynthetic gas exchange parameters, and their temperature dependence, in dominant woody plants at four savanna sites on a moisture gradient in Botswana, southern Africa. Leaf stable carbon and nitrogen (N) isotope and morphological measures were made concurrently. Sampling of these predominantly non‐N‐fixing species took place during an exceptional rainfall season, representing near‐optimum conditions for primary production at these sites. The mean specific leaf area and leaf size were positively related to mean annual rainfall (MAR); species with larger leaves of lower density were more abundant in wetter sites. Almost all species at all sites showed high net light‐saturated photosynthetic rates (A_max?10 μmol CO₂ m^?2 s^?1) due both to high CO₂ carboxylation (V_c,max) and RubP‐regeneration capacity (J_max). These high rates were associated with high values of leaf [N]. Across all sites, the temperature response of A_max showed no clear optimum, and a gradual drop from 25°C to 35°C, without notable temperature limitation at leaf temperatures in excess of 35°C. Dark respiration rate (R_day) across all species and sites increased exponentially with increasing leaf temperature. Species sampled at selected sites revealed a negative relationship between leaf δ¹³C (stable carbon isotope ratio) and MAR, suggesting higher leaf‐level water‐use efficiency at drier sites when integrated over the life of the leaf. At wetter sites, specific leaf [N] was lower and photosynthetic nitrogen‐use efficiency increased, a pattern reflected at the ecosystem level by less ¹⁵N enrichment of leaves at these sites. Taken together, the results suggest a switch from water‐use to nitrogen‐use efficiency constraints with increasing moisture availability. These constraints impact leaf form and function significantly, and may emerge at the ecosystem level in aspects of water and N cycling.     

The growth of soybean under free air [CO<Subscript>2</Subscript>] enrichment (FACE) stimulates photosynthesis while decreasing in vivo Rubisco capacity

Down-regulation of light-saturated photosynthesis (A_sat) at elevated atmospheric CO₂ concentration, [CO₂], has been demonstrated for many C₃ species and is often associated with inability to utilize additional photosynthate and/or nitrogen limitation. In soybean, a nitrogen-fixing species, both limitations are less likely than in crops lacking an N-fixing symbiont. Prior studies have used controlled environment or field enclosures where the artificial environment can modify responses to [CO₂]. A soybean free air [CO₂] enrichment (FACE) facility has provided the first opportunity to analyze the effects of elevated [CO₂] on photosynthesis under fully open-air conditions. Potential ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) carboxylation (V_c,max) and electron transport through photosystem II (J_max) were determined from the responses of A_sat to intercellular [CO₂] (C_i) throughout two growing seasons. Mesophyll conductance to CO₂ (g_m) was determined from the responses of A_sat and whole chain electron transport (J) to light. Elevated [CO₂] increased A_sat by 15–20% even though there was a small, statistically significant, decrease in V_c,max. This differs from previous studies in that V_c,max/J_max decreased, inferring a shift in resource investment away from Rubisco. This raised the C_i at which the transition from Rubisco-limited to ribulose-1,5-bisphosphate regeneration-limited photosynthesis occurred. The decrease in V_c,max was not the result of a change in g_m, which was unchanged by elevated [CO₂]. This first analysis of limitations to soybean photosynthesis under fully open-air conditions reveals important differences to prior studies that have used enclosures to elevate [CO₂], most significantly a smaller response of A_sat and an apparent shift in resources away from Rubisco relative to capacity for electron transport.Abbreviations FACE Free air [CO₂] enrichment - Rubisco Ribulose-1,5-bisphosphate carboxylase/oxygenase - RuBP Ribulose-1,5-bisphosphate - SoyFACE Soybean free air [CO₂] enrichment - VPD Vapor pressure deficit     

不同降水条件下荒漠草原植物的养分含量及回收对增温和氮素添加的响应

为了探讨荒漠草原植物养分回收特征对长期增温和氮素添加的响应以及自然降水变异对其的调控作用,该研究依托实施12年的模拟增温和氮素添加实验平台,在相对多雨的2016年(超过长期均值52%)和相对少雨的2017年(低于长期均值16%),以常见C_3植物银灰旋花(Convolvulus ammannii)和C_4植物木地肤(Kochia prostrata)为研究对象,测定分析绿叶和枯叶的氮磷含量及回收效率。结果表明:(1)在相对多雨年(2016年),增温使2种植物的绿叶氮、枯叶氮、绿叶磷、枯叶磷含量分别增加了14.32%、25.45%、17.97%和46.47%,氮、磷回收效率分别显著减小了9.41%和16.99%(P0.05);氮素添加使2种植物的绿叶氮、枯叶氮、绿叶磷、枯叶磷含量分别提高了17.32%、25.62%、20.21%和51.41%,而氮、磷回收效率显著降低了9.33%和18.89%(P0.05);增温+氮素添加共同处理显著增加了植物氮磷含量、降低了氮磷回收效率。(2)在相对少雨年(2017年),增温、氮素添加、增温+氮素添加处理对植物叶片氮磷含量、回收效率均无显著影响。(3)叶片氮磷含量在物种间差异极显著(P0.000 1),而氮磷回收效率在物种间无显著差异。(4)回归分析表明,植物叶片氮磷含量随着土壤无机氮、有效磷及含水量的增加而增加,植物氮磷回收效率则随着土壤养分和水分的可利用性的增加而降低。研究认为,荒漠草原植物养分回收对全球变化的响应受自然降水变异的调控。     

Maintenance of leaf N controls the photosynthetic CO2 response of grassland species exposed to 9 years of free‐air CO2 enrichment

Determining underlying physiological patterns governing plant productivity and diversity in grasslands are critical to evaluate species responses to future environmental conditions of elevated CO₂ and nitrogen (N) deposition. In a 9‐year experiment, N was added to monocultures of seven C₃ grassland species exposed to elevated atmospheric CO₂ (560 μmol CO₂ mol^?1) to evaluate how N addition affects CO₂ responsiveness in species of contrasting functional groups. Functional groups differed in their responses to elevated CO₂ and N treatments. Forb species exhibited strong down‐regulation of leaf N_mass concentrations (?26%) and photosynthetic capacity (?28%) in response to elevated CO₂, especially at high N supply, whereas C₃ grasses did not. Hence, achieved photosynthetic performance was markedly enhanced for C₃ grasses (+68%) in elevated CO₂, but not significantly for forbs. Differences in access to soil resources between forbs and grasses may distinguish their responses to elevated CO₂ and N addition. Forbs had lesser root biomass, a lower distribution of biomass to roots, and lower specific root length than grasses. Maintenance of leaf N, possibly through increased root foraging in this nutrient‐poor grassland, was necessary to sustain stimulation of photosynthesis under long‐term elevated CO₂. Dilution of leaf N and associated photosynthetic down‐regulation in forbs under elevated [CO₂], relative to the C₃ grasses, illustrates the potential for shifts in species composition and diversity in grassland ecosystems that have significant forb and grass components.     

Effects of elevated CO2 and temperature on leaf characteristics,photosynthesis and carbon storage in aboveground biomass of a boreal bioenergy crop (Phalaris arundinacea L.) under varying water regimes

This work examined the effects of elevated CO₂ and temperature and water regimes, alone and in interaction, on the leaf characteristics [leaf area (LA), specific leaf weight (SLW), leaf nitrogen content (N_L) based on LA], photosynthesis (light‐saturated net carbon fixation rate, P_sat) and carbon storage in aboveground biomass of leaves (C_l) and stem (C_s) for a perennial reed canary grass (Phalaris arundinacea L., Finnish local cultivar). For this purpose, plants were grown under different water regimes (ranging from high to low soil moisture) in climate‐controlled growth chambers under the elevated CO₂ and/or temperature (following a factorial design) over a whole growing season (May–September in 2009). The results showed that the elevated temperature increased the leaf growth, photosynthesis and carbon storage of aboveground biomass the most in the early growing periods, compared with ambient temperature. However, the plant growth declined rapidly thereafter with a lower carbon storage at the end of growing season. This was related to the accelerated phenology regulation and consequent earlier growth senescence. Consequently, the elevation of CO₂ increased the P_sat, LA and SLW during the growing season, with a significant concurrent increase in the carbon storage in aboveground biomass. Low soil moisture decreased the P_sat, leaf stomatal conductance, LA and carbon storage in above ground biomass compared with high and normal soil moisture. This water stress effect was the largest under the elevated temperature. The elevated CO₂ partially mitigated the adverse effects of high temperature and low soil moisture. However, the combination of elevated temperature and CO₂ did not significantly increase the carbon storage in aboveground biomass of the plants.     

Nitrogen use strategy drives interspecific differences in plant photosynthetic CO2 acclimation

Rising atmospheric CO₂ concentration triggers an emergent phenomenon called plant photosynthetic acclimation to elevated CO₂ (PAC). PAC is often characterized by a reduction in leaf photosynthetic capacity (A_sat), which varies dramatically along the continuum of plant phylogeny. However, it remains unclear whether the mechanisms responsible for PAC are also different across plant phylogeny, especially between gymnosperms and angiosperms. Here, by compiling a dataset of 73 species, we found that although leaf A_sat increased significantly from gymnosperms to angiosperms, there was no phylogenetic signal in the PAC magnitude along the phylogenetic continuum. Physio-morphologically, leaf nitrogen concentration (N_m), photosynthetic nitrogen-use efficiency (PNUE), and leaf mass per area (LMA) dominated PAC for 36, 29, and 8 species, respectively. However, there was no apparent difference in PAC mechanisms across major evolutionary clades, with 75% of gymnosperms and 92% of angiosperms regulated by the combination of N_m and PNUE. There was a trade-off between N_m and PNUE in driving PAC across species, and PNUE dominated the long-term changes and inter-specific differences in A_sat under elevated CO₂. These findings indicate that nitrogen-use strategy drives the acclimation of leaf photosynthetic capacity to elevated CO₂ across terrestrial plant species.     

Functional diversity of photosynthesis during drought in a model tropical rainforest – the contributions of leaf area, photosynthetic electron transport and stomatal conductance to reduction in net ecosystem carbon exchange

The tropical rainforest mesocosm within the Biosphere 2 Laboratory, a model system of some 110 species developed over 12 years under controlled environmental conditions, has been subjected to a series of comparable drought experiments during 2000–2002. In each study, the mesocosm was subjected to a 4–6 week drought, with well‐defined rainfall events before and after the treatment. Ecosystem CO₂ uptake rate (A_eco) declined 32% in response to the drought, with changes occurring within days and being reversible within weeks, even though the deeper soil layers did not become significantly drier and leaf‐level water status of most large trees was not greatly affected. The reduced A_eco during the drought reflected both morphological and physiological responses. It is estimated that the drought‐induced 32% reduction of A_eco has three principal components: (1) leaf fall increased two‐fold whereas leaf expansion growth of some canopy dominants declined to 60%, leading to a 10% decrease in foliage coverage of the canopy. This might be the main reason for the persistent reduction of A_eco after rewatering. (2) The maximum photosynthetic electron transport rate at high light intensities in remaining leaves was reduced to 71% for three of the four species measured, even though no chronic photo‐inhibition occurred. (3) Stomata closed, leading to a reduced ecosystem water conductance to water vapour (33% of pre‐drought values), which not only reduced ecosystem carbon uptake rate, but may also have implications for water and energy budgets of tropical ecosystems. Additionally, individual rainforest trees responded differently, expressing different levels of stress and stress avoiding mechanisms. This functional diversity renders the individual response heterogeneous and has fundamental implications to scale leaf level responses to ecosystem dynamics.     

Gas exchange and photosynthetic acclimation over subambient to elevated CO2 in a C3–C4 grassland

Atmospheric CO₂ (C_a) has risen dramatically since preglacial times and is projected to double in the next century. As part of a 4‐year study, we examined leaf gas exchange and photosynthetic acclimation in C₃ and C₄ plants using unique chambers that maintained a continuous C_a gradient from 200 to 550 µmol mol^?1 in a natural grassland. Our goals were to characterize linear, nonlinear and threshold responses to increasing C_a from past to future C_a levels. Photosynthesis (A), stomatal conductance (g_s), leaf water‐use efficiency (A/g_s) and leaf N content were measured in three common species: Bothriochloa ischaemum, a C₄ perennial grass, Bromus japonicus, a C₃ annual grass, and Solanum dimidiatum, a C₃ perennial forb. Assimilation responses to internal CO₂ concentrations (A/C_i curves) and photosynthetically active radiation (A/PAR curves) were also assessed, and acclimation parameters estimated from these data. Photosynthesis increased linearly with C_a in all species (P < 0.05). S. dimidiatum and B. ischaemum had greater carboxylation rates for Rubisco and PEP carboxylase, respectively, at subambient than superambient C_a (P < 0.05). To our knowledge, this is the first published evidence of A up‐regulation at subambient C_a in the field. No species showed down‐regulation at superambient C_a. Stomatal conductance generally showed curvilinear decreases with C_a in the perennial species (P < 0.05), with steeper declines over subambient C_a than superambient, suggesting that plant water relations have already changed significantly with past C_a increases. Resource‐use efficiency (A/g_s and A/leaf N) in all species increased linearly with C_a. As both C₃ and C₄ plants had significant responses in A, g_s, A/g_s and A/leaf N to C_a enrichment, future C_a increases in this grassland may not favour C₃ species as much as originally thought. Non‐linear responses and acclimation to low C_a should be incorporated into mechanistic models to better predict the effects of past and present rising C_a on grassland ecosystems.     

Soil drying and nitrogen availability modulate carbon and water exchange over a range of annual precipitation totals and grassland vegetation types

Increased intensity in precipitation events and longer periods of water deficit are predicted as a general trend under future climate scenarios with potentially large effects on terrestrial ecosystem function. The primary objective of this study was to understand how variation in the intensity of precipitation inputs followed by intermittent soil drying events influence leaf and ecosystem carbon dioxide (CO₂) and water exchange in a California annual grassland mesocosm experiment. We further examined how nitrogen (N) availability, and differences in plant community composition (grass-forb combinations) affected gas exchange responses to the precipitation treatments. Net ecosystem CO₂ exchange (NEE) and evapotranspiration (ET) increased significantly with greater precipitation and were positively correlated with soil moisture. A repeated 10-day soil drying period, following 11 days of watering, strongly depressed NEE over a range of annual precipitation totals (297, 657 and 987 mm), and plant community types. Ecosystem dark respiration ( R _e) and leaf level photosynthesis ( A _max) showed greater sensitivity to periods of soil drying in the low precipitation plots (297 mm). N additions significantly increased NEE and R _e, particularly as water availability was increased. Across the range of precipitation totals and plant community types, intermittent periods of soil moisture deficit and native soil N availability constrained leaf and ecosystem level CO₂ exchange, while the influence on water vapor exchange was less pronounced.     

Influence of summer rainfall on root and shoot growth of a cold-winter desert shrub,Atriplex confertifolia

Summary The influence of irrigation and nitrogen fertilization in early summer on root and shoot growth of Atriplex confertifolia, a C₄ shrub species, was examined in a cold-winter desert community in northern Utah. Soil water and xylem pressure potentials were monitored during the summer period.At the time of watering the surface soil (0–30 cm) was dry but there were turgid fine roots in this horizon. Watering of the soil reduced plant water stress from-30 to-15 bars (dawn values) indicating that roots near the surface were capable of absorbing water, and induced root growth in the 0–30 cm zone. The addition of N to the water treatment did not further increase root production. However, watering and watering +N fertilizer failed to stimulate shoot elongation or any dry weight increase of shoots. This shoot dormancy during summer is not typical of C₄ plants and is probably associated with adaptation to the cool arid environment.This work was carried out while the senior author was on study leave from CSIRO     

No cumulative effect of 10 years of elevated [CO2] on perennial plant biomass components in the Mojave Desert

Elevated atmospheric CO₂ concentrations ([CO₂]) generally increase primary production of terrestrial ecosystems. Production responses to elevated [CO₂] may be particularly large in deserts, but information on their long‐term response is unknown. We evaluated the cumulative effects of elevated [CO₂] on primary production at the Nevada Desert FACE (free‐air carbon dioxide enrichment) Facility. Aboveground and belowground perennial plant biomass was harvested in an intact Mojave Desert ecosystem at the end of a 10‐year elevated [CO₂] experiment. We measured community standing biomass, biomass allocation, canopy cover, leaf area index (LAI), carbon and nitrogen content, and isotopic composition of plant tissues for five to eight dominant species. We provide the first long‐term results of elevated [CO₂] on biomass components of a desert ecosystem and offer information on understudied Mojave Desert species. In contrast to initial expectations, 10 years of elevated [CO₂] had no significant effect on standing biomass, biomass allocation, canopy cover, and C : N ratios of above‐ and belowground components. However, elevated [CO₂] increased short‐term responses, including leaf water‐use efficiency (WUE) as measured by carbon isotope discrimination and increased plot‐level LAI. Standing biomass, biomass allocation, canopy cover, and C : N ratios of above‐ and belowground pools significantly differed among dominant species, but responses to elevated [CO₂] did not vary among species, photosynthetic pathway (C₃ vs. C₄), or growth form (drought‐deciduous shrub vs. evergreen shrub vs. grass). Thus, even though previous and current results occasionally show increased leaf‐level photosynthetic rates, WUE, LAI, and plant growth under elevated [CO₂] during the 10‐year experiment, most responses were in wet years and did not lead to sustained increases in community biomass. We presume that the lack of sustained biomass responses to elevated [CO₂] is explained by inter‐annual differences in water availability. Therefore, the high frequency of low precipitation years may constrain cumulative biomass responses to elevated [CO₂] in desert environments.     

Effects of soil water on photosynthetic characteristics and leaf traits of Cyclobalanopsis glauca seedlings growing under nutrient-rich and -poor soil

Cyclobalanopsis glauca is a dominant species in mid-subtropical forest, and usually plays an important role in forest ecosystems. However, it often suffers redundant precipitation or water stress, which often concurs with high temperature, nutrient depletion and strong irradiance. The study presented in the paper hypothesized that soil water exerted strong influence on leaf gas exchange and traits. The objective of this study was to clarify the effect of soil water changes on photosynthetic characteristics and leaf traits and their relationships of C. glauca seedlings growing on nutrient-rich and nutrient-poor soil at three water levels. The study measured the specific leaf area (SLA), nitrogen content, chlorophyll concentrations and photosynthetic light response curve. Its results showed that there were no differences in leaf size, leaf dry weight, SLA, leaf dry matter content, Leaf nitrogen concentration and Leaf chlorophyll between the two soil nutrient treatments, while these parameters differed significantly among different water levels for either of the treatments. There were large variations in leaf photosynthetic parameters and leaf traits among the different water treatments, indicating different response patterns of C. glauca seedling and its adaptation to the different soil water conditions. There were no significant differences in light-saturated photosynthetic rate (A_max) and apparent quantum yield (Ø) between the nutrient-rich and nutrient-poor soils, which indicated that the C. glauca seedlings could maintain similar capacities in different soils that differed in nutrient condition. As to the relation between the photosynthesis and leaf traits, the A_max and PNUE were positively correlated with the SLA, respectively, but the SLA had significant negative relationship with the leaf N (P<0.01) in nutrient-rich soil. In contrast, both A_max and PNUE were significantly negatively correlated with the SLA, respectively (P<0.01); and the SLA was not significantly positively correlated with the leaf nitrogen concentration of the nutrient-poor soil (P>0.05). The specific leaf areas (SLA), nitrogen and chlorophyll concentrations as well as other photosynthetic features were influenced in a coordinative manner by the soil water. The relation among the A_max, PNUE and the N_mass, SLA could be described as a binomial equation and a liner negative regression for the nutrient-rich and nutrient-poor soil, respectively. In conclusion, soil water was more constraining factor than the soil nutrient to the photosynthesis of C. glauca seedlings, nutrient-rich soil could offset some negative influence resulting from soil water deficit on LSP and LCP. Factors affecting the variations of photosynthetic characteristics and leaf traits of C. glauca seedlings differed between the nutrient-rich and nutrient-poor soils.     

Needle CO2 exchange, structure and defense traits in relation to needle age in Pinus heldreichii Christ – a relict of Tertiary flora

 Pinus heldreichii Christ is a long-lived, slow-growing Tertiary relict from the Balkans. In this study we evaluated the physiological characteristics of eight needle-age classes of P. heldreichii grown at the Arboretum of the Institute of Dendrology in Kórnik, Poland. At the end of the growing season, current-year foliage had the highest rates of mass-based light-saturated net photosynthesis (A_sat) of 33.5 nmol CO₂· g^–1· s^–1. A_sat decreased with needle age, but older needle classes retained from approximately 62 to 26% of the current needles’ rate. The relationship between leaf N and chlorophyll a concentration among all needle-age classes was highly significant (r = 0.96, P = 0.0006). The variation in A_sat of 1- to 7-year-old needles was linearly related to needle N concentration (r = 0.98, P = 0.0001). Needle dark respiration rates among these needle age classes ranged from 0.8 to 2.2 nmol · g^–1· s^–1 and decreased with needle age and nitrogen concentration. Total phenols and glucose concentrations increased linearly with needle age. A similar pattern was observed in acid buffering capacity and the pH of tissue homogenates. The water content ranged from 62% for the current needles to 51% for the 6-year-old needles. Greater investment in leaf structural tissue and increased chemical defense is associated with higher structural cost of older needles and may reduce their photosynthetic activity. Significant declines in water and nitrogen content with needle age and an increase in content of phenolics is most likely a defense adaptation of P. heldreichii related to the species’ long-lived leaves. Received: 8 January 1997 / Accepted: 4 March 1997     

Temporal expression of effects of varying nitrogen supply on canopy growth, photosynthesis and fruit production for Actinidia deliciosa vines in the field

The effects of varying nitrogen supply on canopy leaf area, response of leaf net photosynthesis (A_n) to quantum flux density (Q), and fruit yields of kiwifruit vines (Actinidia deliciosa var. deliciosa) were examined in a two-year field experiment. Vines were grown with 0, 250 or 750 kg N ha^?1 year^?1. The responses to nitrogen supply were compared with responses to shade, to examine the impact of reduced carbon assimilation on canopy leaf area and fruit yields. Nitrogen supply did not affect significantly any of the measured variables during the first season of the experiment. In the second season, canopy leaf area was reduced significantly where nitrogen supply was limited. The quantum efficiency of photosynthesis (φ_q) increased from 0. 03 mol CO₂ mol^?1 Q soon after leaf emergence to more than 0. 05 mol CO₂ mol^?1 Q during the middle of the growing season. The quantum saturated rate of A_n (A_sat) also increased during the season, from 7–10 μmol CO₂ m^?2 s^?1 soon after leaf emergence, to 15–20 (μmol CO₂ m^?2 s^?1 during the middle of the growing season. φ_q and A_sat increased significantly with nitrogen supply at all measurement times during the second season. For vines with high nitrogen, fruit yields in both seasons were similar, averaging 3. 05 kg m^?2. Fruit yields in the second season were reduced significantly where nitrogen supply was limited, due to reduced fruit numbers. The relative effects of reduced leaf area and reduced leaf photosynthesis for carbon assimilation by nitrogen deficient vines were examined using a mathematical model of canopy photosynthesis for kiwifruit vines. Simulations of canopy photosynthesis indicated that effects on leaf area and on leaf photosynthesis were of similar importance in the overall effects of nitrogen deficiency on carbon assimilation. The effects of nitrogen supply on fruit numbers (i. e. flower development) preceded the measured effects on carbon assimilation, indicating that the nitrogen supply affected carbon partitioning to reserves in the first season.     

Stem diameter growth rates in a fire‐prone savanna correlate with photosynthetic rate and branch‐scale biomass allocation,but not specific leaf area

Plant growth rates strongly determine ecosystem productivity and are a central element of plant ecological strategies. For laboratory and glasshouse‐grown seedlings, specific leaf area (SLA; ratio of leaf area to mass) is a key driver of interspecific variation in growth rate (GR). Consequently, SLA is often assumed to drive GR variation in field‐grown adult plants. However, there is an increasing evidence that this is not the general case. This suggests that GR – SLA relationships (and perhaps those for other traits) may vary depending on the age or size of the plants being studied. Here we investigated GR – trait relationships and their size dependence among 17 woody species from an open‐canopy, fire‐prone savanna in northern Australia. We tested the predictions that SLA and stem diameter growth rate would be positively correlated in saplings but unrelated in adults while, in both age classes, faster‐GR species would have higher light‐saturated photosynthetic rate (A_sat), higher leaf nutrient concentrations, higher branch‐scale biomass allocation to leaf versus stem tissues and lower wood density (WD). SLA showed no relationship to stem diameter GR, even in saplings, and the same was true of leaf N and P concentrations, and WD. However, branch‐scale leaf:stem allocation was strongly related to GR in both age groups, as was A_sat. Together, these two traits accounted for up to 80% of interspecific variation in adult GR, and 41% of sapling GR. A_sat is rarely measured in field‐based GR studies, and this is the first report of branch‐scale leaf:stem allocation (analogous to a benefit:cost ratio) in relation to plant growth rate. Our results suggest that we may yet find general trait‐drivers of field growth rates, but SLA will not be one.