巴郎山糙皮桦叶片光合氮利用效率的海拔响应

为深入认识植物对环境变化的响应和适应,以分布在川西巴郎山的糙皮桦为研究对象,选择海拔2200、2500、3100和3400 m 4个分布点,测定计算了各分布点叶片光合氮利用效率(PNUE)、CO₂扩散导度(叶肉细胞导度g_m与气孔导度g_s)和氮分配比例(Rubisco氮分配比例P_R、生物力能学组分氮分配比例P_B、捕光组分氮分配比例P_L与细胞壁氮分配比例P_CW)等参数,分析了其沿海拔的变化趋势以及叶片PNUE与其他参数的相关关系.结果表明: 糙皮桦叶片PNUE、P_R和P_B在海拔2500和3100 m相对较高;叶片g_s和g_m则随海拔升高而增加,P_L随海拔升高而降低.糙皮桦叶片P_R和P_B与PNUE呈显著正相关关系,说明P_R和P_B是PNUE随海拔变异的重要内部因素.糙皮桦叶片光合系统氮分配比例P_P在海拔2500和3100 m相对较高,叶片P_CW随海拔升高而降低,叶片其他组分氮分配比例P_other随海拔升高而增加,说明随海拔的升高,糙皮桦叶片趋向将更大比例的氮分配于除光合系统和细胞壁外的其他组分中.     

Photosynthesis and photosynthetic efficiencies along the terrestrial plant’s phylogeny: lessons for improving crop photosynthesis

Photosynthesis is the basis of all life on Earth. Surprisingly, until very recently, data on photosynthesis, photosynthetic efficiencies, and photosynthesis limitations in terrestrial land plants other than spermatophytes were very scarce. Here we provide an updated data compilation showing that maximum photosynthesis rates (expressed either on an area or dry mass basis) progressively scale along the land plant’s phylogeny, from lowest values in bryophytes to largest in angiosperms. Unexpectedly, both photosynthetic water (WUE) and nitrogen (PNUE) use efficiencies also scale positively through the phylogeny, for which it has been commonly reported that these two efficiencies tend to trade‐off between them when comparing different genotypes or a single species subject to different environmental conditions. After providing experimental evidence that these observed trends are mostly due to an increased mesophyll conductance to CO₂ – associated with specific anatomical changes – along the phylogeny, we discuss how these findings on a large phylogenetic scale can provide useful information to address potential photosynthetic improvements in crops in the near future.     

Photorespiration in the context of Rubisco biochemistry,CO2 diffusion and metabolism

Photorespiratory metabolism is essential for plants to maintain functional photosynthesis in an oxygen‐containing environment. Because the oxygenation reaction of Rubisco is followed by the loss of previously fixed carbon, photorespiration is often considered a wasteful process and considerable efforts are aimed at minimizing the negative impact of photorespiration on the plant’s carbon uptake. However, the photorespiratory pathway has also many positive aspects, as it is well integrated within other metabolic processes, such as nitrogen assimilation and C₁ metabolism, and it is important for maintaining the redox balance of the plant. The overall effect of photorespiratory carbon loss on the net CO₂ fixation of the plant is also strongly influenced by the physiology of the leaf related to CO₂ diffusion. This review outlines the distinction between Rubisco oxygenation and photorespiratory CO₂ release as a basis to evaluate the costs and benefits of photorespiration.     

Strong photosynthetic acclimation and enhanced water‐use efficiency in grassland functional groups persist over 21 years of CO2 enrichment,independent of nitrogen supply

Uncertainty about long‐term leaf‐level responses to atmospheric CO₂ rise is a major knowledge gap that exists because of limited empirical data. Thus, it remains unclear how responses of leaf gas exchange to elevated CO₂ (eCO₂) vary among plant species and functional groups, or across different levels of nutrient supply, and whether they persist over time for long‐lived perennials. Here, we report the effects of eCO₂ on rates of net photosynthesis and stomatal conductance in 14 perennial grassland species from four functional groups over two decades in a Minnesota Free‐Air CO₂ Enrichment experiment, BioCON. Monocultures of species belonging to C₃ grasses, C₄ grasses, forbs, and legumes were exposed to two levels of CO₂ and nitrogen supply in factorial combinations over 21 years. eCO₂ increased photosynthesis by 12.9% on average in C₃ species, substantially less than model predictions of instantaneous responses based on physiological theory and results of other studies, even those spanning multiple years. Acclimation of photosynthesis to eCO₂ was observed beginning in the first year and did not strengthen through time. Yet, contrary to expectations, the response of photosynthesis to eCO₂ was not enhanced by increased nitrogen supply. Differences in responses among herbaceous plant functional groups were modest, with legumes responding the most and C₄ grasses the least as expected, but did not further diverge over time. Leaf‐level water‐use efficiency increased by 50% under eCO₂ primarily because of reduced stomatal conductance. Our results imply that enhanced nitrogen supply will not necessarily diminish photosynthetic acclimation to eCO₂ in nitrogen‐limited systems, and that significant and consistent declines in stomatal conductance and increases in water‐use efficiency under eCO₂ may allow plants to better withstand drought.     

Effect of nitrogen application on growth and photosynthetic nitrogen use efficiency in two ecotypes of wild strawberry, Fragaria chiloensis

The relationships between increasing nitrogen fertilization and growth, maximum CO₂ assimilation and the initial slope of the CO₂ response curve were studied in 2 ecotypes of wild strawberry, Fragaria chiloensis (L.) Duchn. Nitrogen accumulation of CA11, an ecotype from a low-nutrient dune site, was greater at all nitrogen concentrations than that of RCP37, an ecotype from a higher-nutrient strand site. Maximum CO₂ assimilation, total Rubisco activity, dry weight, and initiation of leaves and crowns were higher in CAI1 than RCP37 as nitrogen treatment was increased from 0 to 200 mg ^l-1, whereas these parameters were lower in CAl1 when fertilized at 300 mg T¹, but not in RCP37. The mean leaf area of CA11 was greater than RCP37 when grown with no supplemental nitrogen, but mean leaf area of the 2 lines was similar under nitrogen fertilization. Maximum CO₂ assimilation and carboxylation efficiency increased with increasing leaf nitrogen in both clones. At equivalent concentrations of leaf nitrogen, RCP37 had higher CO₂ assimilation and carboxylation efficiency than CA11 and the difference between the 2 clones increased as ieaf nitrogen increased. Thus, RCP37 had a higher photosynthetic nitrogen use efficiency than CA11. However, at a given applied nitrogen level, CA11 allocated more nitrogen to a unit of leaf area so that photosynthetic rates were higher than RCP37, except at the highest application of 300 mg ^l-1. The high nitrogen accumulation capacity and resource allocation to fruiting structures (crowns) in CA11 leads us to suggest that this clone may possess genes that could increase fruit yield in cultivated strawberry.     

The effects of increasing CO2 on crop photosynthesis and productivity: a review of field studies

Abstract. Only a small proportion of elevated CO₂ studies on crops have taken place in the field. They generally confirm results obtained in controlled environments: CO₂ increases photosynthesis, dry matter production and yield, substantially in C₃ species, but less in C₄, it decreases stomatal conductance and transpiration in C₃ and C₄ species and greatly improves water-use efficiency in all plants. The increased productivity of crops with CO₂ enrichment is also related to the greater leaf area produced. Stimulation of yield is due more to an increase in the number of yield-forming structures than in their size. There is little evidence of a consistent effect of CO₂ on partitioning of dry matter between organs or on their chemical composition, except for tubers. Work has concentrated on a few crops (largely soybean) and more is needed on crops for which there are few data (e.g. rice). Field studies on the effects of elevated CO₂ in combination with temperature, water and nutrition are essential; they should be related to the development and improvement of mechanistic crop models, and designed to test their predictions.     

Changes in Stomatal Conductance,Transpiration and Water Use Efficiency of Ten Species Experienced in High CO2 Concentrations in Biosphere 2

The stomotal conductance, transpiration and water use efficiency (WUE) were measured using a LI-6400 portable photosynthesis system for 5 tropical rain forest species and 5 desert species in Biosphere 2, USA. All the species have experienced in very high CO2 ( > 2 200 μmol• mol- 1 ) for more than 4.5 years. The results showed that the stomatal conductance and transpiration of rain forest species decreased from ( 127.4 ± 65.6) and (2.04 ± 0.61 ) mmol• m- 2•s- 1 to (61.3 + 30.5) and ( 1.54 ± 0.65 ) mmol• m-2• s -1 respectively, while WUE increased from (2.90 ± 0.55) to (8.45 ± 2.71) μmol CO2 •mmo1-1 H2O, with CO2 increasing from 350 – 400 to 700 – 820 μmol• mol-l. For the desert species, stomatal conductance and transpiration decreased from respectively (142.8±94.6) and (2.09±0.71) mmol•m-2•s-1 to (57.7±35.8) and (1.36±0.52) mmolm-2•s-l, but WUE increased from (4.69 ± 1.39) to (9.68 ± 1.61) μmol CO2•mmo1-1 H2O, with the CO2 increase from 320 - 400 to 820 – 850 μtmol• mol- 1. The stomatal conductance, transpiration and WUE were less influenced by light intensity under high CO2 than low CO2 concentrations. Most rain forest species reached their light saturation points at light intensity of 500 μmol• m-2•s-1, while desert species at 1 000 μmol•m-2•s-1. Among different species, the desert C3 tree, Nicotiana glauca Grah., had the highest decrease in stomatal conductance and transpiration and the highest increase in WUE, by 78%, 69% and 310% respectively. The enhancement of increasing CO2 to the stomatal, transpiration and WUE of species with different photosynthesis pathway and life forms in Biosphere 2 could be concluded as: C3 species > C4 species, and desert C3 species > rain forest C3 species.     

施氮和大气CO2浓度升高对小麦旗叶光合电子传递和分配的影响

采用开顶式气室盆栽培养小麦,设计2个大气CO₂浓度（正常：400 μmol·mol^-1;高：760 μmol·mol^-1）、2个氮素水平（0和200 mg·kg^-1土）的组合处理,通过测定小麦抽穗期旗叶氮素和叶绿素浓度、光合速率（P_n）-胞间CO₂浓度（C_i）响应曲线及荧光动力学参数,来测算小麦叶片光合电子传递速率等,研究了高大气CO₂浓度下施氮对小麦旗叶光合能量分配的影响.结果表明：与正常大气CO₂浓度相比,高大气CO₂浓度下小麦叶片氮浓度和叶绿素浓度降低,高氮处理的小麦叶片叶绿素a/b升高.施氮后小麦叶片PSⅡ最大光化学效率（F_v/F_m）、PSⅡ反应中心最大量子产额（F_v′/F_m′）、PSⅡ反应中心的开放比例（q_p）和PSⅡ反应中心实际光化学效率（Φ_PSⅡ）在大气CO₂浓度升高后无明显变化,虽然叶片非光化学猝灭系数（NPQ）显著降低,但PSⅡ总电子传递速率（J_F）无明显增加;不施氮处理的F_v′/F_m′、Φ_PSⅡ和NPQ在高大气CO₂浓度下显著降低,尽管F_v/F_m和q_P无明显变化,J_F仍显著下降.施氮后小麦叶片J_F增加,参与光化学反应的非环式电子流传递速率(J_C)明显升高.大气CO₂浓度升高使参与光呼吸的非环式电子流传递速率(J₀)、Rubisco氧化速率（V₀）、光合电子的光呼吸/光化学传递速率比（J₀/J_C）和Rubisco氧化/羧化比（V₀/V_C）降低,但使J_C和Rubisco羧化速率（V_C）增加.因此,高大气CO₂浓度下小麦叶片氮浓度和叶绿素浓度降低,而增施氮素使通过PSⅡ反应中心的电子流速率显著增加,促进了光合电子流向光化学方向的传递,使更多的电子进入Rubisco羧化过程,P_n显著升高.     

常绿阔叶林8种乔灌木叶片氮含量及其分配与光合能力的关系

以浙江天童国家森林公园常绿阔叶林为研究对象,采用空间代替时间的方法,研究了5个不同演替阶段常见的4种乔木以及4种灌木叶片的光饱和速率(Pmax)、光合氮素利用效率(PNUE)及其与叶片氮含量(NL)、叶片氮素在细胞壁的分配比例(细胞壁N/叶片总N,NCW/NL)、氮素在光合酶中的分配比例(NR/NL)、单位面积叶干重(LMA)的相互关系。结果表明:(1)演替系列4种乔木和4种灌木各种间指标除NL外均表现出显著差异,前期种较后期种具有更高的NR/NL、PNUE、Pmax,而后期种LMA、NCW/NL、MCW/ML(细胞壁干重/叶片总干重)更大,NL在乔木各种间差异不明显,在灌木种间则差异显著;乔木种较灌木种具有更大的LMA、NCW/NL、MCW/ML,而NR/NL则较灌木小;8种植物的Pmax与NL以杨梅为最高,连蕊茶最低;苦槠具有最高的PNUE,而栲树最低。(2)随着演替的进行,前期种的NR/NL、PNUE、Pmax有减小趋势,而LMA、NCW/NL、MCW/ML逐渐增大,后期种则表现出相反的趋势。(3)NR/NL与Pmax、PNUE之间呈显著正相关关系,而LMA、NCW/NL、MCW/ML则与Pmax、PNUE、NR/NL显著负相关。研究认为,NR/NL与NCW/NL之间的负相关性及其对PNUE的影响可以在一定程度上解释树木在光合与维持两方面的权衡关系以及演替的生理机制。     

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

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

Specific leaf area, leaf nitrogen content, and photosynthetic acclimation of Trifolium repens L. seedlings grown at different irradiances and nitrogen concentrations

Clover seedlings were grown at different nitrogen concentrations (5, 10, 15, 20, 25 mM NO₃ ⁻, i.e. N₅ to N₂₅) and two irradiances, I (200 and 400 μmol m⁻² s⁻¹ of photon flux density, i.e. I ₂₀₀ and I ₄₀₀). Net photosynthetic rate (P _N), photosynthetic nitrogen use efficiency (PNUE), leaf chlorophyll (Chl) content, maximum photochemical efficiency (F_v/F_m), and actual photochemical efficiency of photosystem 2 (PS2) (Φ_PS2) increased from N₅ to N₁₅ and decreased with N₁₅ to N₂₅. P _N, PNUE, and Φ_PS2 were higher at I ₄₀₀ than at I ₂₀₀, but F_v/F_m and leaf Chl contents at I ₄₀₀ were lower than at I ₂₀₀. The effects of the N and I on specific leaf area (SLA) and N contents per unit dry mass (N_m) were similar, the SLA and N_m increased from N₅ to N₂₅ and they were higher at I ₂₀₀ than at I ₄₀₀. The nitrogen contents per unit area (N_a) increased from N₅ to N₂₀, but decreased from N₂₀ to N₂₅. The N_a was higher at I ₂₀₀ than at I ₄₀₀ when Trifolium repens grew at N₅ and N₁₀, but it was higher at I ₄₀₀ than at I ₂₀₀ at N₁₅ to N₂₅.     

Growth, gas exchange and water use efficiency of the facultative hemiparasite Rhinanthus minor associated with hosts differing in foliar nitrogen concentration

The facultative hemiparasitic angiosperm Rhinanthus minor was grown on 11 different host species, and in the absence of a host. The height of R. minor parasitising the legume Trifolium pratense exceeded that of unattached plants by more than an order of magnitude, with performance on grasses and non-legume dicotlyedonous hosts between these two extremes. Light saturated rates of photosynthesis in R. minor on different hosts ranged from 1. 5 to 22. 5 μmol g⁻¹ dry weight min⁻¹, and were positively correlated with growth. Foliar nitrogen concentrations in the parasite exceeded those of the hosts. The former were positively related to light saturated rates of photosynthesis, and inversely related to photosynthetic nitrogen use efficiency. There was no relationship between leaf nitrogen concentration and water use efficiency in R. minor. The data are discussed in relation to studies of nitrogen and water use in mistletoes.     

Interrelationship between Leaf Gas-Exchange Characteristics,Area Leaf Mass,and Yield in Soybean (Glycine max L. Merr) Genotypes

Variability in leaf gas-exchange traits in thirteen soybean (Glycine max L. Merr) genotypes was assessed in a field experiment conducted at high altitude (1 950 m). Leaf net photosynthetic rate (P ^N) exhibited a high degree of variability at all the growth stages studied. P _N and other gas-exchange parameters exhibited a seasonal pattern that was similar for all the genotypes. P _N rate was highest at seed filling stage. P _N was positively and significantly associated with aboveground dry matter and seed yield. The area leaf mass (ALM) exhibited a strong positive association with leaf P _N, aboveground dry matter, and seed yield. The positive association between ALM, P _N, and seed yield suggests that this simple and easy to measure character can be used in breeding programmes as a surrogate for higher photosynthetic efficiency and eventually higher yield.     

Effects of elevated carbon dioxide concentration on photosynthesis and growth of small birch plants (Betula pendula Roth.) at optimal nutrition

Small birch plants (Betula pendula Roth.) were grown from seed for periods of up to 70d in a climate chamber at optimal nutrition and at present (350 μmol mol^?1) or elevated (700 μmol mol^?1) concentrations of atmospheric CO₂. Nutrients were sprayed over the roots in Ingestad-type units. Relative growth rate and net assimilation rate were slightly higher at elevated CO₂, whereas leaf area ratio was slightly lower. Smaller leaf area ratio was associated with lower values of specific leaf area. Leaves grown at elevated CO₂ had higher starch concentrations (dry weight basis) than leaves grown at present levels of CO₂. Biomass allocation showed no change with CO₂, and no large effects on stem height, number of side shoots and number of leaves were found. However, the specific root length of fine roots was higher at elevated CO₂. No large difference in the response of carbon assimilation to intercellular CO₂ concentration (A/C_i curves) were found between CO₂ treatments. When measured at the growth environments, the rates of photosynthesis were higher in plants grown at elevated CO₂ than in plants grown at present CO₂. Water use efficiency of single leaves was higher in the elevated treatment. This was mainly attributable to higher carbon assimilation rate at elevated CO₂. The difference in water use efficiency diminished with leaf age. The small treatment difference in relative growth rate was maintained throughout the experiment, which meant that the difference in plant size became progressively greater. Thus, where plant nutrition is sufficient to maintain maximum growth, small birch plants may potentially increase in size more rapidly at elevated CO₂.     

施氮和大气CO2浓度升高对小麦旗叶光合电子传递和分配的影响

采用开顶式气室盆栽培养小麦,设计2个大气CO2浓度(正常:400 μmol.mol-1;高:760 μmol·mol-1)、2个氮素水平(0和200 mg·kg-1土)的组合处理,通过测定小麦抽穗期旗叶氮素和叶绿素浓度、光合速率(Pn)-胞间CO2浓度(C1)响应曲线及荧光动力学参数,来测算小麦叶片光合电子传递速率等,研究了高大气CO2浓度下施氮对小麦旗叶光合能量分配的影响.结果表明:与正常大气CO2浓度相比,高大气CO2浓度下小麦叶片氮浓度和叶绿素浓度降低,高氮处理的小麦叶片叶绿素a/b升高.施氮后小麦叶片PSⅡ最大光化学效率(Fv/Fm)、PSⅡ反应中心最大量子产额(Fv'/Fm')、PSⅡ反应中心的开放比例(qr)和PSⅡ反应中心实际光化学效率(φPSⅡ)在大气CO2浓度升高后无明显变化,虽然叶片非光化学猝灭系数(NPQ)显著降低,但PSⅡ总电子传递速率(JF)无明显增加;不施氮处理的Fv'/Fm'、φPSⅡ和NPQ在高大气CO2浓度下显著降低,尽管Fv/Fm和qp无明显变化,JF仍显著下降.施氮后小麦叶片JF增加,参与光化学反应的非环式电子流传递速率(Jc)明显升高.大气CO2浓度升高使参与光呼吸的非环式电子流传递速率(J0)、Rubisco氧化速率(V0)、光合电子的光呼吸/光化学传递速率比(J0/Jc)和Rubisco氧化/羧化比(V0/Vc)降低,但使Jc和Rubisco羧化速率(Vc)增加.因此,高大气CO2浓度下小麦叶片氮浓度和叶绿素浓度降低,而增施氮素使通过PSⅡ反应中心的电子流速率显著增加,促进了光合电子流向光化学方向的传递,使更多的电子进入Rubisco羧化过程,Pn显著升高.     

氮沉降增加情景下植物-土壤-微生物交互对自然生态系统土壤有机碳的调控研究进展

大气氮沉降增加倾向于促进受氮限制陆地生态系统地上生物量,但是对地下碳过程和土壤碳截存的影响结果迥异,导致陆地生态系统“氮促碳汇”的评估存在很大的不确定性。大气氮沉降输入直接影响微生物活性或间接影响底物质量,改变凋落物和土壤有机质(SOM)的分解速率和分解程度,进而影响土壤有机碳(SOC)的积累与损耗过程。过去相关研究主要集中在土壤碳转化过程和碳储量动态方面,缺乏植物-微生物-SOM交互作用的理解,对土壤碳截存调控的生物化学和微生物学机理尚不清楚。本文以地下碳循环过程为主线,分别综述了氮沉降增加对植物地下碳分配、SOC激发效应、微生物群落碳代谢过程的影响,深入分析SOM化学稳定性与微生物群落动态的关系。该领域研究的薄弱环节体现在:(1)增氮倾向于降低根系的生长和周转,对根际沉积碳分配(数量和格局)的影响及驱动因素不明确;(2)虽然认识到氮素有效性影响土壤激发效应的方向和强度,但是氧化态NO-3和还原态NH+4输入对有机质激发效应的差异性影响及潜在机理知之甚少;(3)微生物碳利用效率(CUE)是微生物群落碳代谢的关键表征,能够很好地解释土壤碳的积累与损耗过程;由于缺乏适宜的测定方法,难以准确量化土壤微生物的CUE及微生物生物量的周转时间;(4)增氮会抑制土壤真菌群落及其胞外酶活性,对细菌群落组成的影响尚未定论,有关SOM化学质量与土壤微生物群落活性、组成之间的耦合关系尚不清楚。未来研究应基于长期的氮添加控制实验平台,结合碳氧稳定性同位素示踪、有机质化学、分子生物学和宏基因组学等方法,深入分析植物同化碳的地下分配规律、微生物碳代谢和周转、有机质化学结构与功能微生物群落的耦合关系等关键环节。上述研究将有助于揭示植物-土壤-微生物交互作用对SOC动态的调控机制,完善陆地生态系统碳-氮耦合循环模型,有效降低区域陆地碳汇评估的不确定性,并可为陆地生态系统应对全球变化提供科学依据。     

Photosynthetic nitrogen-use efficiency in leaves of woody and herbaceous species

1. Photosynthetic characteristics of an annual herb, Chenopodium album , and an evergreen tree, Quercus myrsinaefolia , were compared to clarify causes of the difference in photosynthetic nitrogen-use efficiency (photosynthetic capacity per unit nitrogen) between leaves of herbaceous and evergreen species.
2. When leaves with the same nitrogen content on an area basis were compared, photosynthetic capacity of C. album was twice as high as that of Q. myrsinaefolia . Gas-exchange measurements showed higher intercellular CO₂ concentration in C. album . Biochemical analyses indicated larger allocation of nitrogen into ribulose-1,5-bisphosphate carboxylase, a key enzyme of photosynthesis, and higher specific activity of ribulose-1,5-bisphosphate carboxylase in C. album . However, these differences were not large.
3. Compositional deviation of ¹³C in leaves of the two species suggested that the drop of CO₂ level between the intercellular space and the chloroplast was slightly larger in Q. myrsinaefolia when compared between the leaves with the same photosynthetic capacity.
4. It is concluded that the difference in photosynthetic nitrogen-use efficiency between C. album and Q. myrsinaefolia is not caused by a sole factor that is markedly different between the two species but by several factors each of which is slightly disadvantageous to Q. myrsinaefolia compared with C. album .     

Seasonal dynamics of above‐ and below‐ground biomass and nitrogen partitioning in Miscanthus × giganteus and Panicum virgatum across three growing seasons

The first replicated productivity trials of the C4 perennial grass Miscanthus × giganteus in the United States showed this emerging ligno‐cellulosic bioenergy feedstock to provide remarkably high annual yields. This covered the 5 years after planting, leaving it uncertain if this high productivity could be maintained in the absence of N fertilization. An expected, but until now unsubstantiated, benefit of both species was investment in roots and perennating rhizomes. This study examines for years 5–7 yields, biomass, C and N in shoots, roots, and rhizomes. The mean peak shoot biomass for M. × giganteus in years 5–7 was 46.5 t ha^?1 in October, declining to 38.1 t ha^?1 on completion of senescence and at harvest in December, and 20.7 t ha^?1 declining to 11.3 t ha^?1 for Panicum virgatum. There was no evidence of decline in annual yield with age. Mean rhizome biomass was significantly higher in M. × giganteus at 21.5 t ha^?1 compared to 7.2 t ha^?1 for P. virgatum, whereas root biomass was similar at 5.6–5.9 t ha^?1. M. × giganteus shoots contained 339 kg ha^?1 N in August, declining to 193 kg ha^?1 in December, compared to 168 and 58 kg ha^?1 for P. virgatum. The results suggest substantial remobilization of N to roots and rhizomes, yet still a substantial loss with December harvests. The shoot and rhizome biomass increase of 33.6 t ha^?1 during the 2‐month period between June and August for M. × giganteus corresponds to a solar energy conversion of 4.4% of solar energy into biomass, one of the highest recorded and confirming the remarkable productivity potential of this plant.