Atmospheric CO(2) and mycorrhiza effects on biomass allocation and nutrient uptake of nodulated pea (Pisum sativum L.) plants

The effect of ambient and elevated atmospheric CO(2) on biomass partitioning and nutrient uptake of mycorrhizal and non-mycorrhizal pea plants grown in pots in a controlled environment was studied. The hypothesis tested was that mycorrhizae would increase C assimilation by increasing photosynthetic rates and reduce below-ground biomass allocation by improving nutrient uptake. This effect was expected to be more pronounced at elevated CO(2) where plant C supply and nutrient demand would be increased. The results showed that mycorrhizae did not interact with atmospheric CO(2) concentration in the variables measured. Mycorrhizae did not affect photosynthetic rates, had no effect on root weight or root length density and almost no effect on nutrient uptake, but still significantly increased shoot weight and reduced root/shoot ratio at harvest. Elevated CO(2) increased photosynthetic rates with no evidence for down-regulation, increased shoot weight and nutrient uptake, had no effect on root weight, and actually reduced root/shoot ratio at harvest. Non-mycorrhizal plants growing at both CO(2) concentrations had lower shoot weight than mycorrhizal plants with similar nutritional status and photosynthetic rates. It is suggested that the positive effect of mycorrhizal inoculation was caused by an enhanced C supply and C use in mycorrhizal plants than in non-mycorrhizal plants. The results indicate that plant growth was not limited by mineral nutrients, but partially source and sink limited for carbon. Mycorrhizal inoculation and elevated CO(2) might have removed such limitations and their effects on above-ground biomass were independent, positive and additive.     

水稻生长及其体内C、N、P组成对开放式空气CO2浓度增高和N、P施肥的响应

采用FACE（Free Air Carbon-dioxide Enrichment)技术,研究了不同N、P施肥水平下,水稻分蘖期、拔节期、抽穗期和成熟期根、茎、穗生长,C/N比、N、P含量及N、P吸收对大气CO2浓度升高的响应,结果表明,高CO2促进水稻茎、穗和根的生长,增加分蘖期叶干重,对拔节期、抽穗期的成熟期叶干重没有显著增加,降低茎、叶N含量;增加抽穗期穗N含量;降低成熟期穗N含量;对分蘖期根N含量影响不显著,而降低拔节期,抽穗期和成熟期根N含量,增加拔节期、抽穗期和成熟期叶P含量,对茎、穗、根P含量影响不显著,水稻各组织C含量变化不显著,C/N比增加,显著增加水稻地上部分P吸收;增加N吸收,但没有统计显著性,N、P施用对水稻各组织生物量没有显著影响,高N（HN）比低N（LN）增加组织中N含量,而不同P肥水平间未表现出明显差异,高N条件下高CO2增加水稻成熟期地下部分/地上部分比,文中还讨论了高CO2对N、P含量及地下部分/地上部分比的影响机制。     

大气CO2浓度和温度升高对作物生理生态的影响

论述了大气CO2浓度和温度升高下的植物生长,光合作用,产量以及水分养分利用效率等方面的研究进展,未来高CO2浓度下,光合作用速率有不同程度的提高,生物量和产量增加;气孔导度降低,水分利用效率（WUE）提高,一般地上部分和根系尤其是细根生物量增加,凋落物量随之增加,C/N比率提高,植物残体的腐解速率降低,CO2浓度升高后,会给根际微生物带来更多的底物,从而提高了微生物活性,加速养分的矿化过程,改善植物的养分状况。     

水稻光合碳在植物-土壤系统中的分配及其对CO2升高和施氮的响应

采用¹³C-CO₂进行连续标记,研究水稻分蘖期和孕穗期光合碳在植株-土壤系统中的分配及其对大气CO₂浓度升高(800 μL·L^-1)和施氮(100 mg·kg^-1)的响应.结果表明: CO₂浓度升高显著提高分蘖期根系生物量和孕穗期地上部生物量,并使生物量根冠比在分蘖期增加,而在孕穗期减小.CO₂浓度升高条件下,施氮使水稻地上部分生物量增加,却显著降低了孕穗期水稻根系生物量.CO₂浓度升高使光合¹³C在孕穗期向土壤的输入显著增加,然而施肥并没有促进由CO₂浓度升高驱动的光合¹³C在土壤中的积累,而且还降低了土壤中的光合¹³C的分配比例.综上,CO₂浓度升高显著提高了稻田土壤光合碳输入,促进稻田有机碳周转;施氮促进了水稻地上部的生长,却降低了光合碳向地下的分配比例.     

水稻光合碳在植物-土壤系统中的分配及其对CO2升高和施氮的响应

采用¹³C-CO₂进行连续标记,研究水稻分蘖期和孕穗期光合碳在植株-土壤系统中的分配及其对大气CO₂浓度升高(800 μL·L^-1)和施氮(100 mg·kg^-1)的响应.结果表明: CO₂浓度升高显著提高分蘖期根系生物量和孕穗期地上部生物量,并使生物量根冠比在分蘖期增加,而在孕穗期减小.CO₂浓度升高条件下,施氮使水稻地上部分生物量增加,却显著降低了孕穗期水稻根系生物量.CO₂浓度升高使光合¹³C在孕穗期向土壤的输入显著增加,然而施肥并没有促进由CO₂浓度升高驱动的光合¹³C在土壤中的积累,而且还降低了土壤中的光合¹³C的分配比例.综上,CO₂浓度升高显著提高了稻田土壤光合碳输入,促进稻田有机碳周转;施氮促进了水稻地上部的生长,却降低了光合碳向地下的分配比例.     

氮添加对中国陆地植被地上-地下生物量分配的影响

大气氮沉降水平持续升高导致的外源氮输入增加,强烈影响了陆地生态系统的碳循环。目前,已有大量报道证实了氮沉降升高对全球陆地植被固碳的积极影响。虽然之前大部分研究将这一结果归因于光合作用增强导致的地上生物量增加,但最近的研究发现长期氮添加对植物地下根系的影响也同样重要。归纳整理了181篇公开发表的我国野外模拟氮沉降试验结果,采用整合分析(Meta-analysis)方法,定量评估了氮添加对我国陆地植被地上-地下生物量分配的影响特征和不同生态系统类型及施氮方式之间的影响差异。通过分析地上-地下生物量分配对氮添加的响应差异来探究植被碳增益对长期大气氮沉降增加的潜在响应机制。结果表明,氮添加显著增强了我国陆地植被的光合作用及碳固存,且植物碳增益在不同生态系统类型及施氮制度间有所差异。植物叶片的氮含量显著增加,使得叶片碳氮比及凋落物碳氮比显著降低,但并未显著影响细根的碳氮比。氮添加总体上显著提高了植物的净光合速率,但降低了光合利用效率。地上生物量,凋落物产量和根生物量平均分别显著增加了38%,17%和18%,总体上植物地上部分对氮添加的响应程度比地下部分更高。然而,不同生态系统类型的地上-地下生物...     

大气CO2浓度升高对红桦幼苗根系的影响

应用封闭式生长室系统,研究了CO2浓度升高对红桦(Betula albosinensis)幼苗的根/冠、粗根和细根的干质量、非结构性碳水化合物类含量、碳含量和碳/氮、氮和磷的含量及氮磷吸收量的影响。结果表明:CO2浓度升高使红桦幼苗粗根和细根的干质量增加,同时根/冠值显著升高,表明CO2浓度升高使红桦幼苗生物量向根系的分配增加;与对照相比,粗根的还原糖、蔗糖和总可溶性糖含量显著增加,而在细根中没有显著变化;粗根、细根的淀粉和总的非结构性碳水化合物含量显著增加;CO2浓度升高下粗根和细根的碳含量有升高的趋势但未达到显著水平,同时氮含量降低,碳/氮值升高;氮的吸收量在粗根和细根中均无显著变化。上述结果表明,CO2浓度升高下红桦幼苗根系氮含量下降是由非结构性碳水化合物(主要是淀粉)含量升高和(或)根系生物量增加产生的稀释效应引起的。     

高浓度二氧化碳对植物影响的研究进展

工业革命后全球大气CO2浓度持续上升,不仅对全球气候的变迁产生重大影响,而且对植物的形态、水分利用、蛋白质合成、光合、抗性、生长及生物量等都有不同程度的影响。高浓度CO2促进植物根、幼苗的生长,叶片增厚,降低气孔密度、气孔导度及蒸腾速率,增加水分利用效率、作物的产量及生物量,促进乙烯生物合成,增强植物的抗氧化能力。不同光合途径(C3、C4及CAM)及不同植被类型的植物对高浓度CO2的响应不同。长期和短期的高浓度CO2处理,植物响应方式有很大的差异,如短期高CO2处理使光合能力增强,而长期处理则使光合能力下调。     

Variation in acclimation of photosynthesis in Trifolium repens after eight years of exposure to Free Air CO2 Enrichment (FACE)

The initial stimulation of photosynthesis observed on elevation of [CO2] in grasslands has been predicted to be a transient phenomenon constrained by the loss of photosynthetic capacity due to other limitations, notably nutrients and sinks for carbohydrates. Legumes might be expected partially to escape these feedbacks through symbiotic N2 fixation. The Free-Air Carbon dioxide Enrichment (FACE) experiment at Eschikon, Switzerland, has been the longest running investigation of the effects of open-air elevation of [CO2] on vegetation. The prediction of a long-term loss of photosynthetic capacity was tested by analysing photosynthesis in Trifolium repens L. (cv. Milkanova) in the spring and autumn of the eighth, ninth and tenth years of treatment. A high and low N treatment also allowed a test of the significance of exogenous N-supply in maintaining a stimulation of photosynthetic capacity in the long-term. Prior work in this Free Air CO2 Enrichment (FACE) experiment has revealed that elevated [CO2] increased both vegetative and reproductive growth of T. repens independent of N treatment. It is shown here that the photosynthetic response of T. repens was also independent of N fertilization under both current ambient and elevated (600 micro mol mol-1) [CO2]. There was a strong effect of season on photosynthesis, with light-saturated rates (Asat) 37% higher in spring than in autumn. Higher Asat in the spring was supported by higher maximum Rubisco carboxylation rates (Vc,max) and maximum rates of electron transport (Jmax) contributing to RuBP regeneration. Elevated [CO2] increased Asat by 37% when averaged across all measurement periods and both N fertilization levels, and decreased stomatal conductance by 25%. In spring, there was no effect of elevated [CO2] on photosynthetic capacity of leaves, but in autumn both Vc,max and Jmax were reduced by approximately 20% in elevated [CO2]. The results show that acclimation of photosynthetic capacity can occur in a nitrogen-fixing species, in the field where there are no artificial restrictions on sink capacity. However, even with acclimation there was a highly significant increase in photosynthesis at elevated [CO2].     

植物生理生态指标对大气CO2浓度倍增响应的整合分析

对 8 4篇文献有关植物对大气CO2 浓度倍增响应进行整合分析(一种对同一主题下多个独立实验进行综合的统计学方法),发现环境因素(土壤水分亏缺、土壤低氮、高温和高浓度O3 )显著地影响植物对高CO2 浓度的响应。无任何环境胁迫时,高CO2 浓度对C3 植物的 12个植物生理生态指标产生负效应,对另 12个则表现正效应,负响应最强的前 5个指标为:气孔导度(gs) >暗呼吸速率(Rd) >单位叶重中的氮含量(Nm) >单位叶重中蛋白质含量(Prm) >单位叶结构重量中氮含量(Ns);正响应最强烈的前 5个指标为:根生物量(Br) >地上部生物量(Bs) >单位叶重中淀粉含量(St) >光饱和时的光合速率(A) >总生物量(Bt)。可见植物的气体交换和生物量受高CO2 浓度影响较大,叶化学成分的变化则以淀粉、单位叶重含氮量和单位叶重蛋白质含量较为明显。无任何胁迫时,C3 植物的总生物量和光饱和时的光合速率分别提高 30.0 1%和 40.36 %;气孔导度下降 30.39%。     

Effect of Elevated CO2 and Drought on Soil Microbial Communities Associated with Andropogon gerardii

Our understanding of the effects of elevated atmospheric CO2, singly and In combination with other environmental changes,on plant-soil interactions is incomplete. Elevated CO2 effects on C4 plants, though smaller than on C3 species, are mediated mostly via decreased stomatal conductance and thus water loss. Therefore, we characterized the interactive effect of elevated CO2 and drought on soil microbial communities associated with a dominant C4 prairie grass, Andropogon gerardii Vitman. Elevated CO2 and drought both affected resources available to the soil microbial community. For example, elevated CO2 increased the soil C:N ratio and water content during drought, whereas drought alone decreased both. Drought significantly decreased soil microbial biomass. In contrast, elevated COz increased biomass while ameliorating biomass decreases that were induced under drought. Total and active direct bacterial counts and carbon substrate use (overall use and number of used sources) increased significantly under elevated CO2. Denaturing gradient gel electrophoresis analysis revealed that drought and elevated CO2, singly and combined, did not affect the soil bacteria community structure.We conclude that elevated CO2 alone increased bacterial abundance and microbial activity and carbon use, probably in response to increased root exudation. Elevated CO2 also limited drought-related impacts on microbial activity and biomass,which likely resulted from decreased plant water use under elevated CO2. These are among the first results showing that elevated CO2 and drought work in opposition to modulate plant-associated soil-bacteria responses,which should then Influence soil resources and plant and ecosystem function.     

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 nitrogen on growth ofPoa pratensis (L.)

The growth responses of a grass,Poa pratensis, to elevated CO₂ and nitrogen were investigated. Light-saturated photosynthetic rate per unit leaf area increased with exposure to elevated CO₂, while dry weight did not respond to increased CO₂. Patterns of biomass allocation within plants, including leaf area, leaf area ratio, specific leaf area, and root to shoot ratios, were not altered by elevated CO₂, but changed considerably with N treatment Shoot and whole-plant tissue N concentrations were significantly diluted by elevated CO₂ (Tukey test, P < 0.05). Total N content did not differ significantly among CO₂ treatments. The absence of a concomitant increase in N uptake under elevated CO₂ may have caused a dilution in plant tissue [N], probably negating the positive effects of increased photosynthesis on biomass accumulation.     

Growth under elevated atmospheric CO(2) concentration accelerates leaf senescence in sunflower (Helianthus annuus L.) plants

Some morphogenetic and metabolic processes were sensitive to a high atmospheric CO(2) concentration during sunflower primary leaf ontogeny. Young leaves of sunflower plants growing under elevated CO(2) concentration exhibited increased growth, as reflected by the high specific leaf mass referred to as dry weight in young leaves (16days). The content of photosynthetic pigments decreased with leaf development, especially in plants grown under elevated CO(2) concentrations, suggesting that high CO(2) accelerates chlorophyll degradation, and also possibly leaf senescence. Elevated CO(2) concentration increased the oxidative stress in sunflower plants by increasing H(2)O(2) levels and decreasing activity of antioxidant enzymes such as catalase and ascorbate peroxidase. The loss of plant defenses probably increases the concentration of reactive oxygen species in the chloroplast, decreasing the photosynthetic pigment content as a result. Elevated CO(2) concentration was found to boost photosynthetic CO(2) fixation, especially in young leaves. High CO(2) also increased the starch and soluble sugar contents (glucose and fructose) and the C/N ratio during sunflower primary leaf development. At the beginning of senescence, we observed a strong increase in the hexoses to sucrose ratio that was especially marked at high CO(2) concentration. These results indicate that elevated CO(2) concentration could promote leaf senescence in sunflower plants by affecting the soluble sugar levels, the C/N ratio and the oxidative status during leaf ontogeny. It is likely that systemic signals produced in plants grown with elevated CO(2), lead to early senescence and a higher oxidation state of the cells of these plant leaves.     

Elevated [CO2] and increased N supply reduce leaf disease and related photosynthetic impacts on Solidago rigida

To evaluate whether leaf spot disease and related effects on photosynthesis are influenced by increased nitrogen (N) input and elevated atmospheric CO₂ concentration ([CO₂]), we examined disease incidence and photosynthetic rate of Solidago rigida grown in monoculture under ambient or elevated (560 μmol mol⁻¹) [CO₂] and ambient or elevated (+4 g N m⁻² year⁻¹) N conditions in a field experiment in Minnesota, USA. Disease incidence was lower in plots with either elevated [CO₂] or enriched N (−57 and −37%, respectively) than in plots with ambient conditions. Elevated [CO₂] had no significant effect on total plant biomass, or on photosynthetic rate, but reduced tissue%N by 13%. In contrast, N fertilization increased both biomass and total plant N by 70%, and as a consequence tissue%N was unaffected and photosynthetic rate was lower on N fertilized plants than on unfertilized plants. Regardless of treatment, photosynthetic rate was reduced on leaves with disease symptoms. On average across all treatments, asymptomatic leaf tissue on diseased leaves had 53% lower photosynthetic rate than non-diseased leaves, indicating that the negative effect from the disease extended beyond the visual lesion area. Our results show that, in this instance, indirect effects from elevated [CO₂], i.e., lower disease incidence, had a stronger effect on realized photosynthetic rate than the direct effect of higher [CO₂].     

Meta-analysis*of*the Response*of*Plant Ecophysiological Variables to Doubled Atmospheric CO2 Concentrations

Eighty-four published papers were synthesized using meta-analysis (a statistical method to summarize the different individual studies under the same subject), with results of the environmental factors (soil water deficit, low soil nitrogen, high temperature and high concentration O3) affecting significantly the response of plant to elevated atmospheric CO2 concentrations. Under unstressed condition, the overall effect sizes of twelve ecophysiological variables of C3 plants were negative, other twelve positive. For negative effect group, the first five variables ranked as: stomatal conductance ( g s)> leaf dark respiration rate （Rd）>leaf nitrogen content on a mass basis ( N m)>leaf protein content on a mass basis ( Prm)>leaf nitrogen content on a structural mass basis ( Ns); for positive one, root biomass ( Bt)>shoot biomass ( Bs)>leaf starch content on a mass basis ( St )>light saturated net photosynthetic rate ( A )>total biomass ( Bt). The Bt and A of C3 plants increased 30.01% and 40.36% respectively under unstressed condition, while g s decreased 30.39%.     

CO2浓度升高对沉水克隆植物生长速率及营养元素积累的影响

 研究在不同CO2浓度下水生克隆植物刺苦草（Vallisneria spinulosa）整个生活周期中生长的动态变化及对营养元素积累情况。在不损伤植物体的前提下,采用刺苦草形态学指标组合史估计了植株生物量的动态变化。结果表明：刺苦草鳞茎的萌发不受CO2浓度变化的影响。在高浓度CO2即(1000±50) μmol·mol-1下,刺苦草源株地上部分生长速率在整个生长前期和中期都远远高于低浓度CO2即(400±20) μmol·mol-1,而在后期则出现相反的现象,其中一个原因是因为高浓度CO2下后期光合物质向地下大量转移形成鳞茎引起地上部分生长减慢。但高浓度CO2下克隆株中的初级和次级分株生长速率均高于低浓度CO2。在两种CO2浓度下相同克隆植株构件中的总碳含量没有明显差异;除鳞茎外,根、叶、匍匐茎中的总磷含量随CO2浓度升高显著增加;由于各构件生物量增加有明显差异,导致叶和鳞茎因为生物稀释作用而使其含氮量降低了12%～14%,但根和茎中含量基本保持不变。在高浓度CO2中植株总生物量显著升高,所以总碳、总氮和总磷吸收量均显著大于在低浓度CO2中的吸收量。研究结果揭示,大气CO2浓度升高对沉水克隆植物生长的促进,有利于提高水生克隆植物在群落中的竞争能力;水生植物克隆生长将增加水生生态系统中碳的沉积;水环境中N、P含量将直接影响到水生克隆植物生长。     

开放系统中农作物对空气CO2浓度增加的响应

FACE试验（free-air CO2 enrichment)开展的10多年中,供试农作物主要有：C3禾本科作物小麦（Triticum aestivum L.）、多年生黑麦草（Lolium perenne)和水稻（Oryza sativaL.),C4禾本科类高梁（Sorghum bicolor(L.)Moench),C3豆科植物白三叶草（Trifolium repens ),C3非禾本科块茎状作物马铃薯（Solanum tuberosum L.）,以及多年生C3类木作物棉花（Gossypium hirsutum L.)和葡萄（Vitisvinifera l.)。本文系统整理和分析了以下各项参数的结果;光合作用、气孔导度、冠层温度、水分利用、水势、叶面积指数、根茎生物量累积、作物产量、辐射利用率,比叶面积、N含量、N收益、碳水化合物含量、物候变化、土壤微生物、土壤呼吸、痕量气体交换以及土壤碳固定,CO2浓度升高对农作物的影响作用主要表现在以下方面：（1）促进了植物光合作用,增加了其生物量累积;（2）显著提高C3作物产量,但对C4作物产量的影响很小;（3）降低了C3和C4作物气孔导度,非常显著地提高了所有作物的水分利用率;（4）对植物生长的促进作用在水分不足与水分充中时二者相当或前者大于后者;（6）对根系生长的促进作用要大于地上部分;（7）对多年生植物气孔导度的影响较小,但对其生长的促进作用仍很高;（8）降低了植物体内N含量,但作物体内碳水化合物及某些其他含碳化合物含量增加,且叶部含量要明显高于植物其他器官;（9）对大多数作物的物候略有加速;（10）对某些土壤微生物具显著影响,而对有些则无,但都增加了微生物活性;（11）综合多年、多地点的试验结果表明土壤对大气CO2的固定增加,但单独一个试验无法观测到SOC的显著性变化,对FACE和前期的熏气室试验结果都进行了尽可能的对比研究,除了二例以外,发现在大多数情况下二者的结果基本一致,其中,FACE使气孔导度降低的1.5倍,明显高于前期熏气室试验的结果;其二,相对于熏气室,FACE条件下CO2倍增对根的相对促进作用要高于地上部分,因此,我们对基于这二者的结论的准确性和可靠性是充满信心的,不过,更接近自然环境和具更大小区面积的FACE试验仍是必需的,它可以为我们提供在CO2升高条件下更具代表性的田间试验条件,从而为我们提供更多、更有益的多学科交叉的试验数据和研究结果。     

Short-term exposure to elevated atmospheric CO2 benefits the growth of a facultative annual root hemiparasite,Rhinanthus minor (L.), more than that of its host,Poa pratensis (L.)

The effects of elevated CO2 (650 ppm) on interactions between a chlorophyllous parasitic angiosperm, Rhinanthus minor (L.) and a host, Poa pratensis (L.) were investigated. R. minor benefited from elevated CO2, with both photosynthesis and biomass increasing, and transpiration and tissue N concentration remaining unaffected. However, this did not alleviate the negative effect of the parasite on the host; R. minor reduced host photosynthesis, transpiration, leaf area and biomass, irrespective of CO2 concentration. Elevated CO2 resulted in increased host photosynthesis, but there was no concomitant increase in biomass and foliar N decreased. It appears that the parasite may reduce host growth more by competition for nitrogen than for carbon. Contrary to expectation, R. minor did not reduce the productivity of the host-parasite association, and it actually contributed to the stimulation of productivity of the association by elevated CO2.