Carbon and nitrogen economy of four Triticum aestivum cultivars differing in relative growth rate and water use efficiency

We investigated physiological and morphological traits underlying variation in relative growth rate (KGR) among wheat cultivars. Subsequently, we determined whether higher RGR is correlated with higher water demand and lower plant water use efficiency (WUE_p). Further, the correlation between water use efficiency and leaf nitrogen concentration was examined. For this purpose we chose lour cultivars contrasting in RGR or WUE_p. Gas exchange of shoots and respiration of roots were measured on intact plants over a 24 h period, and total carbon and nitrogen concentrations of all plant parts were determined. The highest RGR was achieved by the cultivars with the highest leaf area ratio. WUE_p was strongly dependent on photosynthetic water use efficiency and was highest for the cultivars with the highest rate of photosynthesis, which achieved higher rates of photosynthesis per unit leaf nitrogen. We found no evidence for a functional or genetic link between the physiological traits underlying differences in RGR (specific leaf area and leaf area ratio) and those causing variation in water use efficiency (photosynthetic rate and transpiration rate). These results indicate that, in wheat, it may be possible to select simultaneously for traits associated with a high WUE_p and a high RGR.     

A genetic link between leaf carbon isotope composition and whole‐plant water use efficiency in the C4 grass Setaria

Genetic selection for whole‐plant water use efficiency (yield per transpiration; WUE_plant) in any crop‐breeding programme requires high‐throughput phenotyping of component traits of WUE_plant such as intrinsic water use efficiency (WUE_i; CO₂ assimilation rate per stomatal conductance). Measuring WUE_i by gas exchange measurements is laborious and time consuming and may not reflect an integrated WUE_i over the life of the leaf. Alternatively, leaf carbon stable isotope composition (δ¹³C_leaf) has been suggested as a potential time‐integrated proxy for WUE_i that may provide a tool to screen for WUE_plant. However, a genetic link between δ¹³C_leaf and WUE_plant in a C₄ species has not been well established. Therefore, to determine if there is a genetic relationship in a C₄ plant between δ¹³C_leaf and WUE_plant under well watered and water‐limited growth conditions, a high‐throughput phenotyping facility was used to measure WUE_plant in a recombinant inbred line (RIL) population created between the C₄ grasses Setaria viridis and S. italica. Three quantitative trait loci (QTL) for δ¹³C_leaf were found and co‐localized with transpiration, biomass accumulation, and WUE_plant. Additionally, WUE_plant for each of the δ¹³C_leaf QTL allele classes was negatively correlated with δ¹³C_leaf, as would be predicted when WUE_i influences WUE_plant. These results demonstrate that δ¹³C_leaf is genetically linked to WUE_plant, likely to be through their relationship with WUE_i, and can be used as a high‐throughput proxy to screen for WUE_plant in these C₄ species.     

Photosynthesis, water use efficiency and stable carbon isotope composition are associated with anatomical properties of leaf and xylem in six poplar species

Although fast‐growing Populus species consume a large amount of water for biomass production, there are considerable variations in water use efficiency (WUE) across different poplar species. To compare differences in growth, WUE and anatomical properties of leaf and xylem and to examine the relationship between photosynthesis/WUE and anatomical properties of leaf and xylem, cuttings of six poplar species were grown in a botanical garden. The growth performance, photosynthesis, intrinsic WUE (WUE_i), stable carbon isotope composition (δ¹³C) and anatomical properties of leaf and xylem were analysed in these poplar plants. Significant differences were found in growth, photosynthesis, WUE_i and anatomical properties among the examined species. Populus cathayana was the clone with the fastest growth and the lowest WUE_i/δ¹³C, whereas P. × euramericana had a considerable growth increment and the highest WUE_i/δ¹³C. Among the analysed poplar species, the highest total stomatal density in P. cathayana was correlated with its highest stomatal conductance (g_s) and lowest WUE_i/δ¹³C. Moreover, significant correlations were observed between WUE_i and abaxial stomatal density and stem vessel lumen area. These data suggest that photosynthesis, WUE_i and δ¹³C are associated with leaf and xylem anatomy and there are tradeoffs between growth and WUE_i. It is anticipated that some poplar species, e.g. P. × euramericana, are better candidates for water‐limited regions and others, e.g. P. cathayana, may be better for water‐abundant areas.     

Mesophyll CO2 conductance and leakiness are not responsive to short‐ and long‐term soil water limitations in the C4 plant Sorghum bicolor

Breeding economically important C₄ crops for enhanced whole‐plant water‐use efficiency (WUE_plant) is needed for sustainable agriculture. WUE_plant is a complex trait and an efficient phenotyping method that reports on components of WUE_plant, such as intrinsic water‐use efficiency (WUE_i, the rate of leaf CO₂ assimilation relative to water loss via stomatal conductance), is needed. In C₄ plants, theoretical models suggest that leaf carbon isotope composition (δ¹³C), when the efficiency of the CO₂‐concentrating mechanism (leakiness, ?) remains constant, can be used to screen for WUE_i. The limited information about how ? responds to water limitations confines the application of δ¹³C for WUE_i screening of C₄ crops. The current research aimed to test the response of ? to short‐ or long‐term moderate water limitations, and the relationship of δ¹³C with WUE_i and WUE_plant, by addressing potential mesophyll CO₂ conductance (g_m) and biochemical limitations in the C₄ plant Sorghum bicolor. We demonstrate that g_m and ? are not responsive to short‐ or long‐term water limitations. Additionally, δ¹³C was not correlated with gas‐exchange estimates of WUE_i under short‐ and long‐term water limitations, but showed a significant negative relationship with WUE_plant. The observed association between the δ¹³C and WUE_plant suggests an intrinsic link of δ¹³C with WUE_i in this C₄ plant, and can potentially be used as a screening tool for WUE_plant in sorghum.     

Water-use efficiency and carbon isotope discrimination in two cultivars of upland rice during different developmental stages under three water regimes

A pot experiment was conducted in a glasshouse to clarify and quantify the effect of plant part, water regime, growth period, and cultivar on carbon isotope discrimination (CID), and to analyze the relationship between CID, stomatal behavior and water-use efficiency (WUE). The experiment was comprised of two upland rice (Oryza sativa L.) cultivars and three water regimes (100, 70, and 40% of saturation moisture) in a completely randomized design. Plants were harvested at tillering, flowering, and maturity. No significant cultivar differences in above-ground dry matter-based WUE (WUE_A) and total dry matter-based WUE (WUE_T) were observed. WUE_A (and WUE_T) increased with water stress up to tillering, but decreased with water stress after tillering. Significant cultivar differences in CID in all the analyzed plant parts were observed at all harvest times. Reduction in CID with water stress was greatest at tillering, and the effect was less pronounced at flowering and at maturity. At each harvest, the effect was most pronounced in newly developed plant parts. Root and grain tended to have the lowest CID values, and stem the highest, at all harvest times. A negative relationship was observed between CID measured at tillering and WUE_A (and WUE_T) measured over the period from seedling to tillering, whereas a reverse relationship was obtained between CID measured at flowering and WUE_A (and WUE_T) measured over the period from tillering to flowering, and an unclear relationship between CID measured at maturity and WUE_A (and WUE_T) measured over the period from flowering to maturity. The ratio of the intercellular and atmospheric concentration of CO₂ (C_i/C_a) were closely associated with CID throughout the water regimes when one cultivar was considered, however, cultivar differences in CID were not related to variations in C_i/C_a. The results indicate that significant cultivar difference existed in CID in all the analyzed plant parts at all harvest times, while corresponding difference in WUE_A (and WUE_T) between the cultivars was not necessarily consistent. Abbreviations: WUE – water-use efficiency; WUEi – instantaneous WUE (or leaf transpiration efficiency); ADM – above-ground dry matter; TDM – total dry matter; WUE_A– ADM-based WUE; WUE_T– TDM-based WUE} CID – carbon isotope discrimination; NL – the newest leaves; FEL – recently fully expanded leaves; FL – flag leaves; P – photosynthesis rate; g – leaf stomatal conductance to water vapor; C_i– intercellular CO₂ concentration; C_a– atmospheric CO₂ concentration; T – transpiration rate; gs – total conductance of CO₂     

Nitrogen, phosphorus and potassium nutritional status of semiarid steppe grassland in Inner Mongolia

In grazed semiarid steppe ecosystems, much attention has been paid to aspects of growth limitation by water. So far, potential limitation of primary production by plant nutrients was rarely considered. This knowledge is essential for identification of sustainable land-use practices in these large and important ecosystems on the background of over-exploitation and climate change. In the present study plant nutrient concentrations and ratios were investigated with factorial additions of water and N fertilizer at two sites with contrasting soil nutrient availability. Combined analysis of nutrient concentrations, contents, biomass production, and plant N:P ratios consistently confirmed primary growth limitation by water and a strong N limitation when sufficient amounts of water were supplied. P limitation only occurred at the site with low P availability when in addition to the natural supply, water and N fertilizer were given. According to reported thresholds of N:K and K:P ratios, K was not limiting in any plot. The observed nutritional patterns in the plant community were related to the dynamics of species composition and their specific nutrient status. Stipa grandis had the highest N:P ratio whereas Artemisia frigida showed lowest N:P. These nutrient characteristics were related to growth strategies of dominant species. Accordingly, the relative biomass contribution of S. grandis and A. frigida strongly affected the nutrient status of the plant community. Plant N:P ratios indicate the relative limitation by N or P in the semiarid grasslands under sufficient water supply, but other methods of nutritional diagnosis should be used when plant N:P ratios remain below critical values.     

Leaf-level nitrogen use efficiency: definition and importance

Nitrogen use efficiency (NUE) has been widely used to study the relationship between nitrogen uptake and dry mass production in the plant. As a subsystem of plant nitrogen use efficiency (NUE), I have defined leaf-level NUE as the surplus production (gross production minus leaf respiration) per unit amount of nitrogen allocated to the leaf, with factorization into leaf nitrogen productivity (NP) and mean residence time of leaf nitrogen (MRT). These concepts were applied to two herbaceous stands: a perennial Solidago altissima stand and an annual Amaranthus patulus stand. S. altissima had more than three times higher leaf NUE than A. patulus due to nearly three times longer MRT of leaf N. In both species, NUE and NP were higher at the leaf level than at the plant level, because most leaf N is involved directly in the photosynthetic activity and because leaf surplus production is higher than the plant net production. MRT was longer at the plant level. The more than twice as long MRT at the plant level as at the leaf level in S. altissima was due to a large contribution of nitrogen storage belowground in the winter in this species. Thus, comparisons between a perennial and an annual system and between plant- and leaf-level NUE with their components revealed the importance of N allocation, storage, recycling, and turnover of organs for leaf photosynthetic production and plant dry mass growth.     

Fluctuations in nitrate reductase activity,and nitrate and organic nitrogen concentrations of succulent plants under different nitrogen and water regimes

The CAM (Crassulacean acid metabolism) succulent species Kalanchoe daigremontiana, K. tubiflora and Crassula argentea, and the succulent C₃ species Peperomia obtusifolia, were cultivated in pure culture in open-air conditions under two different regimes of nitrogen and water supply. At specified intervals during the course of vegetative growth, biomass, nitrate reductase activity (NRA), nitrate concentration, and organic nitrogen concentration of whole plants were measured. After 100 days of cultivation the leaf conductance of Crassula and Peperomia was measured at intervals for the duration of a day. Behaviour of all four species was strongly influenced by the cultivation regime. This was apparent in terms of productivity and variable flucturations in NRA, nitrate concentration, and organic nitrogen concentration during the vegetative period. Increase in biomass was mostly connected with a decrease in all other investigated parameters, especially under conditions of water and/or nitrogen deficiency. The typical reaction of the CAM species Crassula to limited netrogen but adequate soil water was to reduce leaf conductance during light, whereas the C₃ plant Peperomia increased conductance in comparison with plants having a nitrogen suppy. The NRA of all plant species was reduced by both soil nitrate deficiency and drought. The succulent plant species, which are specially adapted to drought, neither took up nor used nitrate when water was limited. This was particularly the case for the CAM species, but less so for the C₃ Peperomia, which showed very high concentrations of nitrate and organic nitrogen, but low NRA and biomass gain. A formula was derived to express the nitrogen use efficiency (NUE) of the species, i.e. the ability of a plant to use nitrogen over a specific period of growth. NUE was shown to increase with age for the crassulacean species but to decrease for the C₃ Peperomia. Furthermore, NUE varied with the different nutrient levels in a species-specific manner, with high values for NUE not necessarily coupled to high productivity, and with NUE of the C₃ species generally higher than that of CAM species.     

Temporal development of the barley leaf metabolic response to Pi limitation

The response of plants to P_i limitation involves interplay between root uptake of P_i, adjustment of resource allocation to different plant organs and increased metabolic P_i use efficiency. To identify potentially novel, early‐responding, metabolic hallmarks of P_i limitation in crop plants, we studied the metabolic response of barley leaves over the first 7 d of P_i stress, and the relationship of primary metabolites with leaf P_i levels and leaf biomass. The abundance of leaf P_i, Tyr and shikimate were significantly different between low Pi and control plants 1 h after transfer of the plants to low P_i. Combining these data with ¹⁵N metabolic labelling, we show that over the first 48 h of P_i limitation, metabolic flux through the N assimilation and aromatic amino acid pathways is increased. We propose that together with a shift in amino acid metabolism in the chloroplast a transient restoration of the energetic and redox state of the leaf is achieved. Correlation analysis of metabolite abundances revealed a central role for major amino acids in P_i stress, appearing to modulate partitioning of soluble sugars between amino acid and carboxylate synthesis, thereby limiting leaf biomass accumulation when external P_i is low.     

The combined effect of constant water deficit and nitrogen supply on WUE, NUE and Δ13C in durum wheat potted plants

Water scarcity and nitrogen shortage are the main constraints on durum wheat productivity. This paper examines the combined effects of a constant water deficit and nitrogen supply (NS) on growth, photosynthesis, stomatal conductance (g_s) and transpiration, instantaneous and time‐integrated water use efficiency (WUE) and nitrogen use efficiency (NUE) and carbon isotope discrimination (Δ¹³C) in durum wheat genotypes grown in pots under greenhouse conditions. Three water levels (40%, 70% and 100% container capacity), two nitrogen doses (high and low N) and four genotypes were assayed in a total of 24 experimental treatments. Water and nitrogen treatments were imposed 2 weeks after plant emergence. The growth, nitrogen content and Δ¹³C of the shoot and the gas exchange in the flag leaf were determined about 2 weeks after anthesis. As expected, both water and NS had a strong positive effect on growth. However, a reduction in water supply had low effect decreasing photosynthesis and transpiration, Δ¹³C and NUE and increasing WUE. On the contrary, increasing the level of nitrogen supplied had a significant negative effect on g_s, which decreased significantly the ratio of intercellular to ambient CO₂ concentrations and Δ¹³C, and increased both instantaneous and time‐integrated WUE. In addition, a higher N level also negatively affected the instantaneous and time‐integrated NUE. The Δ¹³C of shoots correlated significantly and negatively with either instantaneous or time‐integrated measurements of WUE. Moreover, within each NS, Δ¹³C also correlated negatively with the integrated NUE. We concluded that under our experimental conditions, Δ¹³C gives information about the efficiency with which not just water but also nitrogen are used by the plant. In addition, this study illustrates that a steady water limitation may strongly affect biomass without consistent changes in WUE. The lack of effect of the different water regimes on gas exchange, WUE and Δ¹³C illustrate the importance of how stress is imposed during growth.     

Photosynthetic performance of invasive Pinus ponderosa and Juniperus virginiana seedlings under gradual soil water depletion

Changes in climate, land management and fire regime have contributed to woody species expansion into grasslands and savannas worldwide. In the USA, Pinus ponderosa P.&C. Lawson and Juniperus virginiana L. are expanding into semiarid grasslands of Nebraska and other regions of the Great Plains. We examined P. ponderosa and J. virginiana seedling response to soil water content, one of the most important limiting factors in semiarid grasslands, to provide insight into their success in the region. Photosynthesis, stomatal conductance, maximum photochemical efficiency of PSII, maximum carboxylation velocity, maximum rate of electron transport, stomatal limitation to photosynthesis, water potential, root‐to‐shoot ratio, and needle nitrogen content were followed under gradual soil water depletion for 40 days. J. virginiana maintained lower L_s, higher A, g_s, and initial F_v/F_m, and displayed a more gradual decline in V_cmax and J_max with increasing water deficit compared to P. ponderosa. J. virginiana also invested more in roots relative to shoots compared to P. ponderosa. F_v/F_m showed high PSII resistance to dehydration in both species. Photoinhibition was observed at ～30% of field capacity. Soil water content was a better predictor of A and g_s than Ψ, indicating that there are other growth factors controlling physiological processes under increased water stress. The two species followed different strategies to succeed in semiarid grasslands. P. ponderosa seedlings behaved like a drought‐avoidant species with strong stomatal control, while J. virginiana was more of a drought‐tolerant species, maintaining physiological activity at lower soil water content. Differences between the studied species and the ecological implications are discussed.     

Water‐use efficiency in response to climate change: from leaf to ecosystem in a temperate steppe

Water‐use efficiency (WUE) has been recognized as an important characteristic of ecosystem productivity, which links carbon (C) and water cycling. However, little is known about how WUE responds to climate change at different scales. Here, we investigated WUE at leaf, canopy, and ecosystem levels under increased precipitation and warming from 2005 to 2008 in a temperate steppe in Northern China. We measured gross ecosystem productivity (GEP), net ecosystem CO₂ exchange (NEE), evapotranspiration (ET), evaporation (E), canopy transpiration (T_c), as well as leaf photosynthesis (P_max) and transpiration (T_l) of a dominant species to calculate canopy WUE (WUE_c=GEP/T), ecosystem WUE (WUE_gep=GEP/ET or WUE_nee=NEE/ET) and leaf WUE (WUE_l=P_max/T_l). The results showed that increased precipitation stimulated WUE_c, WUE_gep and WUE_nee by 17.1%, 10.2% and 12.6%, respectively, but decreased WUE_l by 27.4%. Climate warming reduced canopy and ecosystem WUE over the 4 years but did not affect leaf level WUE. Across the 4 years and the measured plots, canopy and ecosystem WUE linearly increased, but leaf level WUE of the dominant species linearly decreased with increasing precipitation. The differential responses of canopy/ecosystem WUE and leaf WUE to climate change suggest that caution should be taken when upscaling WUE from leaf to larger scales. Our findings will also facilitate mechanistic understanding of the C–water relationships across different organism levels and in projecting the effects of climate warming and shifting precipitation regimes on productivity in arid and semiarid ecosystems.     

中国东部南北样带森林优势植物叶片的水分利用效率和氮素利用效率

通过测定中国东部南北样带主要森林生态系统中10种优势植物(兴安落叶松、蒙古栎、水曲柳、紫椴、色木槭、红松、杉木、木荷、马尾松、锥栗)叶片的碳氮含量(Cmass、Nmass)、同位素丰度(δ13C、δ15N)以及光合响应曲线,分析了不同优势植物叶片的水分利用效率和氮素利用效率之间的差异及其相互关系.结果表明： 不同生活型植物叶片的Nmass和δ15N差异显著,表现为阔叶植物>针叶植物,落叶植物>常绿植物;最大光合速率(Pn max)表现为针叶植物>阔叶植物,落叶植物>常绿植物;植物叶片的瞬时水分利用效率(WUEi)和长期水分利用效率(WUE)均表现为阔叶植物>针叶植物,常绿植物>落叶植物;植物叶片的瞬时氮素利用效率(NUEi)和长期氮素利用效率(NUE)则表现出相反的规律,且常绿植物和落叶植物叶片的NUE差异显著;WUEi和WUE之间相关性不显著,而NUEi和NUE之间呈显著正相关.植物叶片的水分利用效率与氮素利用效率显著负相关.两种资源利用效率均受植物生活型的影响,并且存在一定的制约关系.     

Dryland Wheat Domestication Changed the Development of Aboveground Architecture for a Well-Structured Canopy

We examined three different-ploidy wheat species to elucidate the development of aboveground architecture and its domesticated mechanism under environment-controlled field conditions. Architecture parameters including leaf, stem, spike and canopy morphology were measured together with biomass allocation, leaf net photosynthetic rate and instantaneous water use efficiency (WUE_i). Canopy biomass density was decreased from diploid to tetraploid wheat, but increased to maximum in hexaploid wheat. Population yield in hexaploid wheat was higher than in diploid wheat, but the population fitness and individual competition ability was higher in diploid wheats. Plant architecture was modified from a compact type in diploid wheats to an incompact type in tetraploid wheats, and then to a more compact type of hexaploid wheats. Biomass accumulation, population yield, harvest index and the seed to leaf ratio increased from diploid to tetraploid and hexaploid, associated with heavier specific internode weight and greater canopy biomass density in hexaploid and tetraploid than in diploid wheat. Leaf photosynthetic rate and WUE_i were decreased from diploid to tetraploid and increased from tetraploid to hexaploid due to more compact leaf type in hexaploid and diploid than in tetraploid. Grain yield formation and WUE_i were closely associated with spatial stance of leaves and stems. We conclude that the ideotype of dryland wheats could be based on spatial reconstruction of leaf type and further exertion of leaf photosynthetic rate.     

The greening of the Northern Great Plains and its biogeochemical precursors

Vegetation greenness has increased across much of the global land surface over recent decades. This trend is projected to continue—particularly in northern latitudes—but future greening may be constrained by nutrient availability needed for plant carbon (C) assimilation in response to CO₂ enrichment (eCO₂). eCO₂ impacts foliar chemistry and function, yet the relative strengths of these effects versus climate in driving patterns of vegetative greening remain uncertain. Here we combine satellite measurements of greening with a 135 year record of plant C and nitrogen (N) concentrations and stable isotope ratios (δ¹³C and δ¹⁵N) in the Northern Great Plains (NGP) of North America to examine N constraints on greening. We document significant greening over the past two decades with the highest proportional increases in net greening occurring in the dries and warmest areas. In contrast to the climate dependency of greening, we find spatially uniform increases in leaf‐level intercellular CO₂ and intrinsic water use efficiency that track rising atmospheric CO₂. Despite large spatial variation in greening, we find sustained and climate‐independent declines in foliar N over the last century. Parallel declines in foliar δ¹⁵N and increases in C:N ratios point to diminished N availability as the likely cause. The simultaneous increase in greening and decline in foliar N across our study area points to increased N use efficiency (NUE) over the last two decades. However, our results suggest that plant NUE responses are likely insufficient to sustain observed greening trends in NGP grasslands in the future.     

Influence of phosphorus and nitrogen on photosynthetic parameters and growth in <Emphasis Type="Italic">Vicia faba</Emphasis> L.

The influence of phosphorus (P) and nitrogen (N) supply on biomass, leaf area, photon saturated photosynthetic rate (P_max), quantum yield efficiency (α), intercellular CO₂ concentration (C_i), and carboxylation efficiency (CE) was investigated in Vicia faba. The influence of P on N accumulation, biomass, and leaf area production was also investigated. An increase in P supply was consistently associated with an increase in N accumulation and N productivity in terms of biomass and leaf area production. Furthermore, P increased the photosynthetic N use efficiency (NUE) in terms of P_max and α. An increase in P supply was also associated with an increase in CE and a decrease in C_i. Under variable daily meteorological conditions specific leaf nitrogen content (N_L), specific leaf phosphorus content (P_L), specific leaf area (δ_L), root mass fraction (R_f), P_max, and α remained constant for a given N and P supply. A monotonic decline in the steady-state value of R_f occurred with increasing N supply. δ_L increased with increasing N supply or with increasing N_L. We tested also the hypothesis that P supply positively affects both N demand and photosynthetic NUE by influencing the upper limit of the asymptotic values for P_max and CE, and the lower limit for C_i in response to increasing N.     

Nitrogen response efficiency increased monotonically with decreasing soil resource availability: a case study from a semiarid grassland in northern China

The concept of nutrient use efficiency is central to understanding ecosystem functioning because it is the step in which plants can influence the return of nutrients to the soil pool and the quality of the litter. Theory suggests that nutrient efficiency increases unimodally with declining soil resources, but this has not been tested empirically for N and water in grassland ecosystems, where plant growth in these ecosystems is generally thought to be limited by soil N and moisture. In this paper, we tested the N uptake and the N use efficiency (NUE) of two Stipa species (S. grandis and S. krylovii) from 20 sites in the Inner Mongolia grassland by measuring the N content of net primary productivity (NPP). NUE is defined as the total net primary production per unit N absorbed. We further distinguished NUE from N response efficiency (NRE; production per unit N available). We found that NPP increased with soil N and water availability. Efficiency of whole-plant N use, uptake, and response increased monotonically with decreasing soil N and water, being higher on infertile (dry) habitats than on fertile (wet) habitats. We further considered NUE as the product of the N productivity (NP the rate of biomass increase per unit N in the plant) and the mean residence time (MRT; the ratio between the average N pool and the annual N uptake or loss). The NP and NUE of S. grandis growing usually in dry and N-poor habitats exceeded those of S. krylovii abundant in wet and N-rich habitats. NUE differed among sites, and was often affected by the evolutionary trade-off between NP and MRT, where plants and communities had adapted in a way to maximize either NP or MRT, but not both concurrently. Soil N availability and moisture influenced the community-level N uptake efficiency and ultimately the NRE, though the response to N was dependent on the plant community examined. These results show that soil N and water had exerted a great impact on the N efficiency in Stipa species. The intraspecific differences in N efficiency within both Stipa species along soil resource availability gradient may explain the differences in plant productivity on various soils, which will be conducive to our general understanding of the N cycling and vegetation dynamics in northern Chinese grasslands.     

The coupling effects of water deficit and nitrogen supply on photosynthesis,WUE, and stable isotope composition in <Emphasis Type="Italic">Picea asperata</Emphasis>

Water stress and nitrogen (N) availability are the two main factors limiting plant growth, and the two constrains can interact in intricate ways. Moreover, atmospheric N depositions are altering the availability of these limiting factors in many terrestrial ecosystems. Here, we studied the combined effects of different soil water availability and N supply on photosynthesis and water-use efficiency (WUE) in Picea asperata seedlings cultured in pots, using gas exchange, and stable carbon and nitrogen isotope composition (δ ¹³C and δ ¹⁵N). Photosynthesis under light saturation (A _sat) and stomatal conductance (g _s) of P. asperata decreased as the soil moisture gradually diminished. Under severe water-stress condition, N addition decreased the A _sat and g _s, whereas the positive effects were observed in moderate water-stress and well-watered conditions. The effect of N addition on the intrinsic WUE (WUE_i) deduced from gas exchange was associated with soil water availability, whereas long-term WUE evaluated by leaf δ ¹³C only affected by soil water availability, and it would be elevated with soil moisture gradually diminished. Water deficit would restrict the uptake and further transport of N to the aboveground parts of P. asperata, and then increasing δ ¹⁵N. Therefore, δ ¹⁵N in plant tissues may reflect changes in N allocation within plants. These results indicate that the effect of N enrichment on photosynthesis in P. asperata is largely, if not entirely, dependent on the severity of water stress, and P. asperata would be more sensitive to increasing N enrichment under low soil water availability than under high soil moisture.     

Seasonal water-use efficiency and chlorophyll fluorescence response in alpha grass (<Emphasis Type="Italic">Stipa tenacissima</Emphasis> L.) is affected by tussock size

Abstact  Twelve randomly chosen Stipa tenacissima L. individuals were grouped into three tussock size classes, small (ST), medium (MT), and large (LT) with 5.6±0.8, 34.1±4.2, and 631.9±85.8 g of dry green foliar matter, respectively, in three plots with different S. tenacissima cover. Instantaneous (WUE_i) and long-term (WUE_l) water-use efficiencies were measured in two seasons of contrasting volumetric soil water content (early winter 21.0±0.8 % and summer 5.8±0.3 %). Maximum photochemical efficiency of photosystem 2 and stomatal conductance in summer assessed the extent of water and irradiance stress in tussocks of different size. WUE_i was lower in MT and ST “water spender” strategies than in LT during the high water-availability season. In summer net photosynthetic rate and WUE_i were higher and photoinhibition was lower in LT than in MT and ST. Significant spatial variability was found in WUE_i. Water uptake was competitive in stands with denser alpha grass and more water availability in summer, reducing their WUE_i. However, WUE_l showed a rising tendency when water became scarce. Thus it is important to explicitly account for plant size in ecophysiological studies, which must be combined with demographic information when estimating functional processes at stand level in sequential scaling procedures.