Effect of Nitrogen and Water Stress on Photosynthesis and Nitrogen Content in Wheat

The relationships between photosynthesis, leaf nitrogen content and water stress were studied in ten genotypes of wheat differing in the presence of dwarfing genes. Net photosynthetic rate (P_N) was mostly higher at ear emergence stage than at anthesis stage. P_N decreased with water stress (leaf water potential from –2.0 to –2.5 MPa), and with reduced leaf N content in all genotypes studied. Among the various genotypes, single dwarf and wild types showed higher P_N rate and maintained higher leaf N content under different N doses and water supply as compared to the other types studied.     

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

Background and Aims

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

Methods

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

Key Results

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

Conclusions

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

Photosynthetic variation and photosynthetic nitrogen use efficiency in <Emphasis Type="Italic">Brassica</Emphasis> species with different genetic constitution of ribulose-1,5-bisphosphate carboxylase

Net photosynthetic rate (P _N) was high in genotypes with ‘C’ genome both in the nucleus and cytoplasm. This may be attributed to the co-ordinated manner of acting of both genome sources. Leaf mass per area (LMA) and chlorophyll content increased with leaf nitrogen (N) content but did not show any correlation with P _N. The factors which affected P _N had the same effect on photosynthetic nitrogen use efficiency (pNUE). Thus, differential allocation of N to the various components influences plant pNUE which is not significantly affected by genome constitution.     

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     

云南文山石漠化区车桑子叶脉密度与叶氮含量关系对生境的响应

植物叶脉特征和叶氮含量的变化影响着叶片经济谱的形成,为验证叶片结构中叶脉网络构建提供了理论依据。该文以单位质量叶氮含量(N_(mass))和单位面积叶氮含量(N_(area))分别表示叶氮含量,采取主成分分析、线性回归分析的方法,研究了云南文山石漠化区旷地(Ⅰ)、林缘(Ⅱ)和林下(Ⅲ)3种自然生境下车桑子的叶脉密度(Vein density,VD)与N_(mass)和N_(area)的异速关系。结果表明:从乔灌群落的旷地到林下,车桑子的比叶面积、叶绿素总含量、光能利用率和N_(mass)逐渐增大,光饱和点、光补偿点、水分利用效率、VD、N_(area)逐渐减小,净光合速率、蒸腾速率、气孔导度呈先增大后减小的趋势。VD与叶氮含量呈不同程度的相关性,在生境I和III,VD与N_(mass)和N_(area)分别具有显著的负相关(P0.05)和正相关(P0.05);在生境Ⅱ,VD与N_(mass)和N_(area)分别呈不显著负相关(P0.05)和正相关(P0.05)。车桑子在旷地强光生境,高VD的叶片含有低N_(mass)高N_(area),而林下荫蔽生境偏向于相反的配置模式,反映了石漠化区植物较强的叶脉可塑性及其与氮利用性状的权衡机制。     

Changes in leaf characteristics as indicator of the alteration of functional types of steppe plants along the aridity gradient

Changes in some leaf characteristics: leaf mass area (LMA), content of photosynthetic pigments and nitrogen in the leaves, leaf mass ratio (LMR) and leaf area ratio (LAR) were investigated in steppe plants of the Volga land along the gradient of aridity. When drought stress became stronger, the content of chlorophylls in the leaves, LMR and LAR decreased, whereas LMA and the proportion of carotenoids in the leaves rose. In the North to South direction, the content of pigments and nitrogen per unit whole plant weight considerably decreased (4 and 2 times, respectively). The relationship between leaf indices (chlorophyll and nitrogen contents and LMA) differed along this gradient. It was concluded that adaptation of steppe plants to drought stress generally depended on predominant development of heterotrophic tissues in the leaf and the whole plant. During aridization, the stress-tolerant species became more numerous.     

Cotton genotypic variation in the photosynthetic response to irradiance

The photosynthetic response of 8 cotton (Gossypium hirsutum L.) genotypes to changing irradiance was investigated under field conditions during the 1998 through 2000 growing seasons. Equations developed to describe the response of net photosynthetic rate (P_N) to photosynthetic photon flux density (PPFD) demonstrated that, across all irradiances, the two okra leaf-type genotypes photosynthesized at a greater rate per unit leaf area than all of the six normal leaf-type genotypes. This superior photosynthetic performance of the okra leaf-type genotypes can be partially explained by their 13 % greater leaf chlorophyll content relative to that of the normal leaf-type genotypes. The 37 % reduction in leaf size brought upon by the okra leaf trait may have concentrated the amount of photosynthetic machinery per unit leaf area. Nevertheless, the lack of sufficient canopy leaf surface area suppressed the potential yield development that could accompany the higher P_N per unit leaf area.This revised version was published online in March 2005 with corrections to the page numbers.     

Genotype Variability in Photosynthetic Characteristics in Finger Millet

High variability in leaf gas exchange and related traits were found in 30 genotypes of field grown finger millet. The variability in carbon exchange rate per unit leaf area (P _N) can be partly attributed to the differences in the stomatal conductance (g_s) and area leaf mass (ALM). The P _N was positively correlated with total dry matter (TDM). However, no relationship between P _N and seed yield was found. The leaf area showed a positive and significant correlation with total biomass. None of the other gas-exchange traits had significant relationship either with TDM or with seed yield. The ALM showed a strong positive association with P _N. However, it was not correlated with either total biomass or seed yield. As a result, the use of ALM as surrogate for P _N for identifying high biomass producing genotypes only had a limited value. Hence selection for high P _N would result in higher biomass producing types.     

Genotypic variation in photosynthesis in cacao is correlated with stomatal conductance and leaf nitrogen

Variation in photosynthetic parameters was observed between eight contrasting cacao (Theobroma cacao) genotypes. Net photosynthetic rate (P_N) ranged from 3.4 to 5.7 μmol(CO₂) m⁻² s⁻¹ for the genotypes IMC 47 and SCA 6, respectively. Furthermore, genotypic differences were detected in quantum efficiency ranging from 0.020 to 0.043 μmol(CO₂) μmol⁻¹(photon) for UF 676 and AMAZ 15/15, respectively. Differences in PN were correlated with both stomatal conductance (g_s) and leaf nitrogen per unit area. Some variation in water use efficiency was observed between genotypes, both intrinsic (P_N/g_s) and instantaneous (P_N/transpiration rate). Both measures of water use efficiency were a negative function of specific leaf area. Evidence was found for a trade-off mechanism between cacao genotypes in photosynthesis and leaf structure. High photosynthetic rate, expressed on a mass basis was associated with smaller leaves. Furthermore, thinner leaves were compensated for by a higher nitrogen content per unit mass.     

Reduced nitrogen allocation to expanding leaf blades suppresses ribulose-1,5-bisphosphate carboxylase/oxygenase synthesis and leads to photosynthetic acclimation to elevated CO<Subscript>2</Subscript> in rice

Net photosynthetic rate (P _N) measured at elevated CO₂ concentration (C _e), ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), and nitrogen (N) content in rice leaves decreased significantly after exposure to long term C _e. The reduction in P _N, Rubisco, and leaf N at C _e was similar for the last fully expanded leaf blade (LFELB) and expanding leaf blade (ELB). Spatial leaf N content in the ELB was highest in the zone of cell division, sharply declined as cell expansion progressed and gradually increased with cell maturation. Maximum reduction in spatial leaf N and Rubisco content was found at C _e only within cell expansion and maturation zones. The spatial leaf N content correlated well with the amount of Rubisco synthesized during leaf expansion, suggesting that N deposition into the expanding leaf blade may be the key for Rubisco synthesis and possibly photosynthetic acclimation to C _e.     

Photosynthetic Performance of Ginkgo biloba L. Grown Under High and Low Irradiance

Diurnal variation in net photosynthetic rate (P _N) of three-year-old plants of Ginkgo biloba was studied under open, O (receiving full sunlight), net-shade, NS (40 % of photosynthetically active radiation, PAR), or greenhouse, G (25 % PAR) conditions. In all three conditions, P _N was higher in morning along with stomatal conductance (g _s), and intercellular CO₂ concentration (C _i), while leaf temperature and vapour pressure deficit were low. The O-plants exhibited a typical decline in P _N during midday, which was not observed in NS-plants. This indicated a possible photoinhibition in O-plants as the ratio of variable to maximum fluorescence (F_v/F_m) and photosystem 2 (PS2) yield (_PS2) values were higher in the NS- and G-plants. On the contrary, stomatal density and index, chlorophyll a/b ratio, leaf thickness, and density of mesophyll cells were greater in O-plants. Further, higher P _N throughout the day along with higher relative growth rate under NS as compared to O and G suggested the better efficiency of Ginkgo plants under NS conditions. Therefore, this plant species could be grown at 40 % irradiance to meet the ever-increasing demand of leaf and also to increase its export potential.     

A model separating leaf structural and physiological effects on carbon gain along light gradients for the shade-tolerant species Acer saccharum

A process-based leaf gas exchange model for C₃ plants was developed which specifically describes the effects observed along light gradients of shifting nitrogen investment in carboxylation and bioenergetics and modified leaf thickness due to altered stacking of photosynthetic units. The model was parametrized for the late-successional, shade-tolerant deciduous species Acer saccharum Marsh. The specific activity of ribulose-1,5-bisphosphate carboxylase (Rubisco) and the maximum photosynthetic electron transport rate per unit cytochrome f (cyt f) were used as indices that vary proportionally with nitrogen investment in the capacities for carboxylation and electron transport. Rubisco and cyt f per unit leaf area are related in the model to leaf dry mass per area (M_A), leaf nitrogen content per unit leaf dry mass (N_m), and partitioning coefficients for leaf nitrogen in Rubisco (P_R) and in bioenergetics (P_B). These partitioning coefficients are estimated from characteristic response curves of photosynthesis along with information on lear structure and composition. While P_R and P_B determine the light-saturated value of photosynthesis, the fraction of leaf nitrogen in thylakoid light-harvesting components (P_L) and the ratio of leaf chlorophyll to leaf nitrogen invested in light harvesting (C_B), which is dependent on thylakoid stoichiometry, determine the initial photosynthetic light utilization efficiency in the model. Carbon loss due to mitochondrial respiration, which also changes along light gradients, was considered to vary in proportion with carboxylation capacity. Key model parameters - N_m, P_R, P_B, P_LC_B and stomatal sensitivity with respect to changes in net photosynthesis (G_r) – were examined as a function of M_A, which is linearly related to irradiance during growth of the leaves. The results of the analysis applied to A. saccharum indicate that P_B and P_R increase, and G_f, P_L and C_B decrease with increasing M_A. As a result of these effects of irradiaiice on nitrogen partitioning, the slope of the light-saturated net photosynthesis rate per unit leaf dry mass (A^m_max) versus N_m relationship increased with increasing growth irradiance in mid-season. Furthermore, the nitrogen partitioning coefficients as well as the slopes of A^m_max versus N_m were independent of season, except during development of the leaf photosynthetic apparatus. Simulations revealed that the acclimation to high light increased A^m_max by 40% with respect to the low light regime. However, light-saturated photosynthesis per leaf area (A^a_max) varied 3-fold between these habitats, suggesting that the acclimation to high light was dominated by adjustments in leaf anatomy (A^a_max=A^m_maxM_A) rather than in foliar biochemistry. This differed from adaptation to low light, where the alterations in foliar biochemistry were predicted to be at least as important as anatomical modifications. Due to the light-related accumulation of photosynthetic mass per unit area, A^a_max depended on M_A and leaf nitrogen per unit area (N_a). However, N_a conceals the variation in both M_A and N_m (N_a=N_mM_A), and prevents clear separation of anatomical adjustments in foliage structure and biochemical modifications in foliar composition. Given the large seasonal and site nutrient availability-related variation in N_m, and the influences of growth irradiance on nitrogen partitioning, the relationship between A^a_max and N_a is universal neither in time nor in space and in natural canopies at mid-season is mostly driven by variability in M_A. Thus, we conclude that analyses of the effects of nitrogen investments on potential carbon acquisition should use mass-based rather than area-based expressions.     

Effects of elevated CO<Subscript>2</Subscript> and nitrogen on wheat growth and photosynthesis

The effects of nitrogen [75 and 150 kg (N) ha⁻¹] and elevated CO₂ on growth, photosynthetic rate, contents of soluble leaf proteins and activities of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) and nitrate reductase (NR) were studied on wheat (Triticum aestivum L. cv. HD-2285) grown in open top chambers under either ambient (AC) or elevated (EC) CO₂ concentration (350 ± 50, 600 ± 50 μmol mol⁻¹) and analyzed at 40, 60 and 90 d after sowing. Plants grown under EC showed greater photosynthetic rate and were taller and attained greater leaf area along with higher total plant dry mass at all growth stages than those grown under AC. Total soluble and Rubisco protein contents decreased under EC but the activation of Rubisco was higher at EC with higher N supply. Nitrogen increased the NR activity whereas EC reduced it. Thus, EC causes increased growth and P_N ability per unit uptake of N in wheat plants, even if N is limiting.     

不同种源五角枫幼苗叶片N、P化学计量学特征对氮添加的响应

不同土壤氮环境下植物对氮沉降的响应趋势不同,因此研究不同种源地的五角枫(Acer mono)对氮沉降的响应机理具有重要的意义。为了深入了解不同种源五角枫对氮沉降的适应和响应机制,主要通过3个种源的五角枫幼苗氮添加控制实验,分析不同种源的五角枫幼苗比叶面积(SLA)和叶片干物质含量(LMDC)以及叶片N、P含量和N∶P对氮添加的响应规律。结果表明:(1)不同氮添加处理下,内蒙古种源五角枫幼苗(NW)和山西种源五角枫幼苗(SW)的LDMC均显著低于对照,NW和SW在N4条件下表现为显著的促进作用。随着氮添加浓度的增加,NW与北京种源的五角枫幼苗(BW)的SLA在N4条件下显著增加;SW的SLA在N2的水平下显著增加。(2)氮添加下,只有SW的P_(mass)在N1水平下显著增加,N_(mass)在N1和N3水平下表现为显著增加。说明在低氮水平下,能够促进SW对P的吸收,从而增加植物体内对P的积累。同时在低氮和中高氮条件下,能够促进SW对N的吸收,说明SW能较好的适应低氮和高氮环境,使其生存能力更强。(3)分析变异来源及各指标的相关性得出,氮添加条件下,五角枫的SLA、LDMC、P_(area)及N∶P与氮添加处理呈显著的相关关系,而SLA、LDMC、N_(mass)、P_(mass)、Narea、P_(area)和N∶P对氮添加处理和种源的交互作用均未表现出相关关系,说明引起五角枫各性状主要受氮添加处理影响。(4)综合各指标的变化,氮添加条件下,NW的LDMC显著降低,SLA显著增大,同时叶N含量、叶P含量以及N∶P比较高;SW的LDMC显著降低,SLA增大,同时叶N含量、叶P含量表现为显著增大,结合叶经济谱理论分析得出,SW和NW属于"快速投资-收益"型,而BW倾向于"缓慢投资-收益"型。因此,不同种源的五角枫对于氮沉降的响应存在差异,并且持续的氮添加已经改变了五角枫的适应性。     

Monitoring leaf photosynthesis with canopy spectral reflectance in rice

Non-destructive and rapid method for assessment of leaf photosynthetic characteristics is needed to support photosynthesis modelling and growth monitoring in crop plants. We determined the quantitative relationships between leaf photosynthetic characteristics and canopy spectral reflectance under different water supply and nitrogen application rates. The responses of reflectance at red radiation (wavelength 680 nm) to different water contents and nitrogen rates were parallel to those of leaf net photosynthetic rate (P _N). The relationships of reflectance at 680 nm and ratio index of R(810,680) (near infrared/red, NIR/R) to P _N of different leaf positions and leaf layers in rice indicated that the top two full leaves were the best leaf positions for quantitative monitoring of leaf P _N with remote sensing technique, and the ratio index R(810,680) was the best ratio index for evaluating leaf photosynthetic characteristics in rice. Testing of the models with independent data sets indicated that R(810,680) could well estimate P _N of top two leaves and canopy leaf photosynthetic potential in rice, with the root mean square error of 0.25, 0.16, and 4.38, respectively. Hence R(810,680) can be used to monitor leaf photosynthetic characteristics at different growth stages of rice under diverse growing conditions.     

Variations in the relationship between maximum leaf carboxylation rate and leaf nitrogen concentration

叶片最大羧化速率是表征植物光合能力的关键参数, 受到光照、温度、水分、CO₂浓度、叶片氮含量等多个要素的控制。准确地模拟植物叶片最大羧化速率对环境因子的响应是预测未来植被生产力和碳循环过程的前提。目前大多数陆地碳循环过程模型以Farqhuar光合作用模型为基础模拟植物的光合作用, 关于植物叶片的最大羧化速率与叶氮含量关系的模拟方法却各不相同。该文汇总了1990-2013年国内外植物叶片光合速率观测研究文献中叶片最大羧化速率与叶氮含量的关系式及相关数据, 分析了叶片最大羧化速率与叶氮含量关系随不同植被功能型和时间的变化特征, 以及环境因子变化条件下最大羧化速率与叶氮含量关系的变化特征, 探讨了二者关系变异性的可能原因以及影响因子。结果表明: 1)不同功能型植物叶片的最大羧化速率和叶氮含量的关系存在较大差异, 二者线性关系式的斜率平均值变化范围为16.29-50.25 μmol CO₂·g N^-1·s^-1。落叶植被叶片的最大羧化速率随叶氮含量的变化率和光合氮利用效率一般都高于常绿植被, 其变异主要源于植物的比叶重和叶片内部氮素分配的差异。2)叶片最大羧化速率随叶氮含量的变化存在季节和年际变异。在没有受到水分胁迫的年份中, 叶片最大羧化速率随叶氮含量变化的速率一般在春季或夏季最高, 其季节变异与比叶重和叶氮在Rubisco的分配比例的季节变化有关。受到干旱的影响, 叶片最大羧化速率随叶氮含量的变化率会升高。3)当大气CO₂浓度增加时, 由于叶片中Rubisco含量的降低, 多年生针叶叶片最大羧化速率和叶氮关系斜率值会出现降低; 当供氮水平增加时, 叶片最大羧化速率和叶片氮含量均表现出增加趋势, 二者线性关系的斜率也相应增加。在此基础上, 该文指出在模拟叶片最大羧化速率与叶氮含量的关系时, 应考虑叶片比叶重和叶氮在Rubisco中的分配比例的季节变异、水分胁迫、大气CO₂浓度和供氮水平变化对二者关系的影响。囿于数据的有限性, 今后应进一步加强多因子控制实验研究, 深入探讨叶片最大羧化速率与叶氮含量关系的变异性机理, 并获得更系统的观测数据, 以助生态系统过程模型的改进, 提高模型的模拟精度。     

Nitrogen enhanced photosynthesis of Miscanthus by increasing stomatal conductance and phosphoenolpyruvate carboxylase concentration

Miscanthus is one of the most promising bioenergy crops with high photosynthetic nitrogen-use efficiency (PNUE). It is unclear how nitrogen (N) influences the photosynthesis in Miscanthus. Among three Miscanthus genotypes, the net photosynthetic rate (P _N) under the different light intensity and CO₂ concentration was measured at three levels of N: 0, 100, and 200 kg ha^?1. The concentrations of chlorophyll, soluble protein, phosphoenolpyruvate carboxylase (PEPC), ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) large subunit, leaf anatomy and carbon isotope discrimination (Δ) in the leaf were analyzed to probe the response of photosynthesis in Miscanthus genotypes to N levels. P _N in all genotypes rose significantly as N application increased. The initial slope of response curves of P _N to C _i was promoted by N application in all genotypes. Both stomatal conductance and C _i were increased with increased N supply, indicating that stomatal factors played an important role in increasing P _N. At a given C _i, P _N in all genotypes was enhanced by N, implying that nonstomatal factors might also play an important role in increasing P _N. Miscanthus markedly regulated N investment into PEPC rather than the Rubisco large subunit under higher N conditions. Bundle sheath leakiness of CO₂ was constant at about 0.35 for all N levels. Therefore, N enhanced the photosynthesis of Miscanthus mainly by increasing stomatal conductance and PEPC concentration.     

植物叶片最大羧化速率与叶氮含量关系的变异性

叶片最大羧化速率是表征植物光合能力的关键参数, 受到光照、温度、水分、CO₂浓度、叶片氮含量等多个要素的控制。准确地模拟植物叶片最大羧化速率对环境因子的响应是预测未来植被生产力和碳循环过程的前提。目前大多数陆地碳循环过程模型以Farqhuar光合作用模型为基础模拟植物的光合作用, 关于植物叶片的最大羧化速率与叶氮含量关系的模拟方法却各不相同。该文汇总了1990-2013年国内外植物叶片光合速率观测研究文献中叶片最大羧化速率与叶氮含量的关系式及相关数据, 分析了叶片最大羧化速率与叶氮含量关系随不同植被功能型和时间的变化特征, 以及环境因子变化条件下最大羧化速率与叶氮含量关系的变化特征, 探讨了二者关系变异性的可能原因以及影响因子。结果表明: 1)不同功能型植物叶片的最大羧化速率和叶氮含量的关系存在较大差异, 二者线性关系式的斜率平均值变化范围为16.29-50.25 μmol CO₂·g N^-1·s^-1。落叶植被叶片的最大羧化速率随叶氮含量的变化率和光合氮利用效率一般都高于常绿植被, 其变异主要源于植物的比叶重和叶片内部氮素分配的差异。2)叶片最大羧化速率随叶氮含量的变化存在季节和年际变异。在没有受到水分胁迫的年份中, 叶片最大羧化速率随叶氮含量变化的速率一般在春季或夏季最高, 其季节变异与比叶重和叶氮在Rubisco的分配比例的季节变化有关。受到干旱的影响, 叶片最大羧化速率随叶氮含量的变化率会升高。3)当大气CO₂浓度增加时, 由于叶片中Rubisco含量的降低, 多年生针叶叶片最大羧化速率和叶氮关系斜率值会出现降低; 当供氮水平增加时, 叶片最大羧化速率和叶片氮含量均表现出增加趋势, 二者线性关系的斜率也相应增加。在此基础上, 该文指出在模拟叶片最大羧化速率与叶氮含量的关系时, 应考虑叶片比叶重和叶氮在Rubisco中的分配比例的季节变异、水分胁迫、大气CO₂浓度和供氮水平变化对二者关系的影响。囿于数据的有限性, 今后应进一步加强多因子控制实验研究, 深入探讨叶片最大羧化速率与叶氮含量关系的变异性机理, 并获得更系统的观测数据, 以助生态系统过程模型的改进, 提高模型的模拟精度。     

敦煌阳关湿地芦苇叶性状对土壤水分的响应

为了了解干旱区湿地不同水分梯度下芦苇叶性状的构建模式及对不同水分环境的适应策略差异性,以敦煌阳关渥洼池湿地植物芦苇(Phragmites australis)为研究对象,分析不同水分梯度下芦苇叶性状对土壤水分的响应。结果表明:(1)低水分梯度下的叶厚度(LT)和叶片磷含量(LPC)显著高于中、高水分梯度(P0.05),高水分梯度下的叶碳含量(LCC)也显著高于中、低水分梯度(P0.05)。(2)芦苇叶性状之间的关系也因水分的变化而有所不同。就整体而言,叶片碳含量与叶干物质含量(LDMC)极显著正相关(P0.01);叶厚度与叶片磷含量极显著正相关(P0.01),比叶面积(SLA)与叶氮含量(LNC)极显著正相关(P0.01)、与叶碳氮比(C/N)极显著负相关(P0.01)。(3)在水分作为限制因素的条件下,叶片碳含量与比叶面积对水分的响应最为突出。土壤含水量与叶厚度、叶碳含量和叶片磷含量均显著负相关(P0.05)。芦苇在低水分环境下采取高收入低投入的积极型生存策略,高水分下则采取低获取高消耗的保守型生存策略,说明了芦苇在适应异质生境的自我调节机制。     

Suboptimal nitrogen status sensitizes the photosynthetic apparatus in willow leaves to long term but not short term water stress

The response to drought was compared for willow plants of optimal leaf nitrogen content (100 N) and those of 86% of this content (86 N). Gas exchange measurements revealed that the carboxylation efficiency (CE) of photosynthesis was more sensitive to drought than the photosynthetic capacity in both N regimes. Since the leaf content of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) was found to be much more resistant it is suggested that a decreased specific activity of Rubisco underlies the decreased CE. Although the rate of water consumption was the same for 86 N and 100 N plants the photosynthetic apparatus responded much more rapidly in the 86 N leaves. This increased sensitivity of 86 N leaves was not due to accelerated senescence as judged by comparison with parallel plants subjected to discontinued fertilization; the two categories of treatments resulted in the same loss of leaf nitrogen and Rubisco but drought induced a much more rapid photosynthetic depression. In contrast to the drought situation, 86 N and 100 N plants behaved similarly when compared under short term water stress. First, when single attached leaves were exposed to a sudden drop in air humidity the capacity of CO₂ uptake in both N regimes decreased about 20% over 10 min while the leaf water potential remained high. Second, in freely transpiring leaf discs cut from 86 N and 100 N leaves the same relationship between capacity of O₂ evolution and extent of dehydration was observed. The possible mechanisms underlying the increased susceptibility of 86 N leaves to drought is discussed; the water status of the roots not the leaves is suggested to be the determining factor.Abbreviations CE carboxylation efficiency - 100 N optimal nitrogen regime - 86 N suboptimal nitrogen regime with 86% of the optimal leaf nitrogen content, Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase