植物同化物分配及其模型研究综述

目前生态系统模型模拟中所用的大多数同化物分配模型是经验性的。同化物分配对植物的生长、竞争及结构的形成有重要的影响,是植物生长的关键,也是植物生长模型中的薄弱环节。总结了影响同化物分配的因素：生理过程和环境因子。指出植物作为一个有机的整体,所有的生理过程都对其有影响,维管束作为各器官间的连接系统,以及同化物的运输管道,其性质对同化物分配有重要的影响。综述了环境因子特别是环境水分条件对同化物分配的影响。总结了以往研究中发现的、主要的同化物分配规律,指出同化物分配的模式极其复杂,分配过程完全是根据环境以及生长阶段变化而变化的、随机应变的过程。 对于同化物分配模型按照经验模型,目的性模型,源汇关系模型进行了总结归纳,分析指出：经验性模型应用最多但机理性差;功能平衡模型在模拟营养生长阶段同化物在条与根之间的分配很成功,但应用于其它器官之间很困难;最优化模型适于模拟平衡态下同化物的分配;源汇关系模型机理性最强,可模拟任何器官间的同化物分配,应用范围最广泛。 同化物研究取得了很大的进展,但研究中仍存在很多不足：对于各相关过程的研究存在不平衡性;整体水平上同化物分配的机理仍缺乏深入研究;同化物分配对于环境的响应方面的研究相对较弱;缺乏多环境因素的研究;缺乏长期的实验观测研究。作者认为环境与同化物分配相互关系的研究将成为日后研究中的热点问题。     

氮输入对滨海盐沼湿地碳循环关键过程的影响及机制

滨海盐沼湿地是缓解全球变暖的有效蓝色碳汇, 但是近岸海域富营养化导致的大量氮输入对盐沼湿地稳定性和碳汇功能构成严重威胁。潮汐作用下大量氮输入对盐沼湿地植物光合碳输入、植物-土壤碳分配和土壤碳输出等碳循环关键过程产生深刻影响, 进而影响盐沼湿地碳汇功能评估的准确性。该文从植物光合固碳、植物-土壤系统碳分配、土壤有机碳分解、土壤可溶性有机碳释放、盐沼湿地土壤碳库5个方面综述了氮输入对盐沼湿地碳循环关键过程的影响。在此基础上, 针对当前研究的不足, 提出今后的研究中, 需要进一步探究氮输入对盐沼湿地植物光合固碳及碳分配过程的影响、盐沼湿地土壤有机碳分解的微生物机制、盐沼湿地土壤可溶性有机碳产生和横向流动的影响、以及氮类型对盐沼湿地土壤碳库的影响。以期为揭示氮输入对盐沼湿地碳汇形成过程与机制提供基础资料和理论依据, 为评估未来近岸海域水体富营养化影响下滨海盐沼湿地碳库的潜在变化提供新思路。     

森林土壤碳氮循环过程的新视角:丛枝与外生菌根树种的作用

几乎所有树木的根系都能与丛枝菌根(AM)真菌或外生菌根(EM)真菌形成共生关系,从而调节森林生态系统土壤碳(C)、氮(N)循环等功能过程。深入理解不同菌根类型森林土壤C、N循环的差异及其影响机制是重要的生态研究命题。该文明晰了AM与EM森林土壤C、N循环的差异;基于森林土壤C、N输入、稳定和输出等3个过程剖析了AM和EM树种对土壤C、N循环的影响机制;比较了不同菌根类型森林土壤C、N循环过程对全球变化的响应;指出了该研究领域所面临的主要挑战:(1)全面比较研究不同菌根类型森林土壤C、N循环及其相关联的生态系统结构和功能特征,为提高森林生产力、发挥生态系统服务功能提供理论基础和数据;(2)深入认知不同菌根树种地上凋落物及地下菌根与自由微生物间相互作用对土壤C、N循环的影响,以阐明不同菌根类型森林土壤C、N循环的潜在机制;(3)改进研究方法,应用新技术手段,充分考虑时空尺度效应,以便能用小尺度的研究结果合理地解释和预测生态系统C、N循环;(4)加强不同菌根类型森林土壤C、N稳定性差异的研究,以准确评价森林结构和功能对全球变化的响应。     

磷影响陆地生态系统碳循环过程及模型表达方法

全球气候变暖已大大改变了陆地植物碳吸收能力, 提高了全球植被净初级生产力。随着气候变暖的加剧, 磷对植物生长的限制作用逐渐显现且不断增强, 磷影响陆地生态系统碳循环的机理和模型研究已成为研究热点。该文系统分析了磷影响陆地生态系统碳循环的相关机理以及模型对相关过程的定量化表达方法。综合对比分析了国际上的Carnegie- Ames-Stanford Approach-CNP (CASA-CNP)、Community Land Model-CNP (CLM-CNP)和Jena Scheme for Biosphere-Atmosphere Coupling in Hamburg-CNP (JSBACH-CNP)等碳、氮、磷耦合模型中磷影响植物光合作用与同化物分配过程、植物对磷的吸收过程、土壤中磷的转化过程以及生态系统磷输入与输出等过程的相关数学表达方法, 指出了模型算法的局限与不确定性以及未来模型发展与改进的方向。同时综合对比分析了CASA-CNP、CLM-CNP、JSBACH-CNP模型的基本特征, 总结了磷循环模型的建模方法, 为未来开展磷影响陆地生态系统碳循环的模型模拟研究提供了借鉴方法与参考思路。     

磷影响陆地生态系统碳循环过程及模型表达方法

全球气候变暖已大大改变了陆地植物碳吸收能力,提高了全球植被净初级生产力。随着气候变暖的加剧,磷对植物生长的限制作用逐渐显现且不断增强,磷影响陆地生态系统碳循环的机理和模型研究已成为研究热点。该文系统分析了磷影响陆地生态系统碳循环的相关机理以及模型对相关过程的定量化表达方法。综合对比分析了国际上的CarnegieAmes-Stanford Approach-CNP (CASA-CNP)、Community Land Model-CNP (CLM-CNP)和Jena Scheme for Biosphere-Atmosphere Coupling in Hamburg-CNP (JSBACH-CNP)等碳、氮、磷耦合模型中磷影响植物光合作用与同化物分配过程、植物对磷的吸收过程、土壤中磷的转化过程以及生态系统磷输入与输出等过程的相关数学表达方法,指出了模型算法的局限与不确定性以及未来模型发展与改进的方向。同时综合对比分析了CASA-CNP、CLM-CNP、JSBACH-CNP模型的基本特征,总结了磷循环模型的建模方法,为未来开展磷影响陆地生态系统碳循环的模型模拟研究提供了借鉴方法与参考思路。     

全球气候变化对陆地植物碳氮磷生态化学计量学特征的影响

全球气候变化背景下生物地球化学循环的响应规律和陆地植物适应对策已受到广泛关注.本文在分析气候变暖和降水变化对不同生态系统植物C∶N∶P的影响、CO₂浓度升高对不同光合途径物种元素的影响,以及氮沉降对土壤 植物元素影响的短期和长期效应等基础上,从植物生理特性和土壤有效营养元素变化等方面揭示了其可能存在的内在机理,以期为研究C、N、P化学元素在土壤 植物之间传递与调节机制、陆地生态系统结构和功能,以及生物地球化学元素循环对气候变化的响应提供理论依据.最后提出了该领域研究中存在的问题及对今后研究的展望.     

植物近期光合碳分配及转化

碳 (C) 既是生命基础元素,又是与大气环境及全球气候变化密切相关的基本成分。近期 (系指数星期的时间尺度,约相当于植物某一生长期) 光合C是“大气-植物-土壤”系统C循环的重要组成部分;定量近期光合C在植物组织、土壤和呼吸损失的分配,对于理解全球C循环是必不可少的。植物近期光合C分结构性C和代谢C;由于代谢C周转快和结构性C尚未及时形成有机质等,近期光合C容易被忽视。应用同位素技术,可从多方面研究植物近期光合C分配及转化特征。这些方面主要包括根际区光合C代谢、近期光合C对CO₂ 和CH₄排放的贡献、C₃植物与C₄ 植物对全球C循环模式的作用、人类活动对近期光合C分配的影响等。我国鲜见植物近期光合C分配及转化的研究,开展此方面的工作,对我国从生命代谢角度研究C循环具有积极的推动作用。     

营养生长期植物冠根比及其对环境因子的响应

将光强、温度和土壤水势等环境因子对植物光合、呼吸、同化物运输及生长等生理过程的影响结合起来,并考虑到各生理过程之间的交互作用,建立了一个营养生长期内植物条根比变化及对环境因子综合响应的模型。模型的运行结果表明,该模型与许多实验现象均相吻合。     

植物源挥发性有机物对氮沉降响应研究展望

植物排放的挥发性有机物(Biogenic volatile organic compounds, BVOCs),属于植物次生代谢物质,是植物重要的防御物质,亦是全球碳(C)素循环的一个重要组成部分。它们具有很高的化学活性,参与对流层大气化学过程,并对全球变化和碳氮(N)循环等具有潜在的影响。尽管N沉降全球化已严重干扰了生态系统的碳氮循环,并且已威胁到生态系统的健康和安全。然而N沉降对BVOCs影响的研究报道十分缺乏。综述了BVOCs影响因素的基础上,重点论述了N素对BVOCs的影响,提出了N沉降对植物BVOCs影响的趋势模型:在N素不足的系统中,N沉降的增加补充了系统所需的N素,有利于植物的生长,大量BVOCs的排放会受到抑制;在N素丰富或过量的系统中,N沉降导致系统N素过饱和或富营养化,不利于植物的生长,刺激BVOCs的排放增加。此外,还探讨了研究N沉降对BVOCs影响的可行性方法,强调开展我国N沉降对BVOCs的影响研究的重要性和紧迫性。为我国开展N沉降对BVOCs的影响研究以及加深了解生态系统CN循环及其藕合提供参考。     

荒漠生态系统磷循环及其驱动机制研究进展

磷(P)循环在维持荒漠生态系统的生物多样性水平、结构和功能的稳定性、元素的动态平衡,以及荒漠自然资源的可持续利用方面有重要作用。通过查阅国内外有关P循环的文献资料,发现当前国内尚缺乏针对荒漠生态系统P循环的系统研究,特别是P循环的生物和非生物驱动机制。综述了荒漠生态系统P的输入-输出过程,植物对P的吸收转运机制以及对P循环的作用,生物土壤结皮(BSC)有机分泌物对P循环的贡献,以及荒漠生态系统P循环过程对气候变化的响应机制等。文末展望了荒漠生态系统P循环的一些重要研究方向和亟需解决的科学问题,包括(1)P在荒漠生态系统中的存在形态、分配及动态平衡;(2)土壤微生物对荒漠植物获取土壤有效P的驱动作用;(3)入侵植物对P循环的影响与潜在生态风险评估;(4)利用分子生物学和基因组学手段揭示真菌-荒漠植物根系系统P循环的基因调控机制;(5)微生物分泌物、土壤磷酸酶类(包括磷酸单酯酶、磷酸二酯酶和三磷酸单酯水解酶)和作用于含磷酸酐和N—P键的酶对土壤P循环的调控;(6)气候变化(干旱、高温和降水节律变化等)如何影响P的生物和非生物转化过程;(7)基于同位素示踪和生态化学计量学理论解释荒漠生态系统...     

Warming prevents the elevated CO2-induced reduction in available soil nitrogen in a temperate, perennial grassland

Rising atmospheric carbon dioxide concentration ([CO₂]) has the potential to stimulate ecosystem productivity and sink strength, reducing the effects of carbon (C) emissions on climate. In terrestrial ecosystems, increasing [CO₂] can reduce soil nitrogen (N) availability to plants, preventing the stimulation of ecosystem C assimilation; a process known as progressive N limitation. Using ion exchange membranes to assess the availability of dissolved organic N, ammonium and nitrate, we found that CO₂ enrichment in an Australian, temperate, perennial grassland did not increase plant productivity, but did reduce soil N availability, mostly by reducing nitrate availability. Importantly, the addition of 2 °C warming prevented this effect while warming without CO₂ enrichment did not significantly affect N availability. These findings indicate that warming could play an important role in the impact of [CO₂] on ecosystem N cycling, potentially overturning CO₂‐induced effects in some ecosystems.     

模拟大气氮沉降对中国森林生态系统影响的研究进展

人类活动加剧了活性氮的生产和排放,并导致氮沉降日益增加并全球化。目前,人类活动对全球氮循环的干扰已经超出了地球系统安全运行的界限。中国已成为全球氮沉降的高发区域,高氮沉降已经威胁到生态系统的健康和安全,并成为生态文明建设过程中亟待理清和解决的热点问题。对国际上和中国森林生态系统模拟氮沉降研究的概况进行了综述,并从生物学和非生物学两大过程重点阐述模拟氮沉降增加对中国主要森林生态系统影响的研究进展。中国自2000年以后才开始重视大气氮沉降产生的生态环境问题,中国科学院华南植物园在国内森林生态系统模拟氮沉降试验研究上做出了开创性的贡献。模拟氮沉降研究表明,持续高氮输入将会显著改变森林生态系统的结构和功能,并威胁生态系统的健康发展,特别是处于氮沉降热点区域的中国中南部。森林生态系统的氮沉降效应依赖于系统的氮状态、土地利用历史、气候特征、林型和林龄等。最后,对未来的研究提出了一些建议,包括加强长期跟踪研究和不同气候带站点之间的联网研究,特别是在森林生态系统对长期氮沉降响应与适应的过程机制、地下碳氮吸存潜力研究、以及与其他全球变化因子的耦合研究等方面,以期为森林生态系统的可持续发展提供理论基础和管理依据。     

陆地生态系统氮沉降增加的生态效应

 人类活动在全球范围内极大地改变着氮素从大气向陆地生态系统输入的方式和速率，人为固定的氮素正在不断积累，并对生态系统的结构和功 能产生显著影响。该文从以下几个方面综述了大气氮沉降增加对陆地生态系统的影响：1)氮输入增加可能影响植物生产力和生态系统碳蓄积能 力，生态系统响应的方向和程度取决于系统的初始氮状况(氮限制或氮饱和)以及当地的植被和土壤特征；2)持续氮输入有可能改变土壤氮循环 过程，降低土壤固持氮的能力，甚至导致土壤酸化、盐基离子损耗，进而影响到土壤有机碳的分解；3)高的氮沉降速率和持续氮输入都可能加 速含氮痕量气体的释放，但其影响程度受生态系统初始状态的影响(例如磷限制和氮限制)；4)氮沉降增加会影响生态系统的物种丰富度、植物 群落结构和动态，促进森林扩张，改变菌根真菌的物种多样性；5)持续氮输入带来的植物群落结构和植物生理特征的变化可能影响昆虫取食特 性，进而通过食物链改变生态系统的营养结构；6) 氮沉降增加对生态系统的影响并不是孤立存在的，它与CO₂浓度升高和O₃浓度变化有协同作 用，但难以从其协同效应中区分出各自的影响。最后，该文总结了我国的氮沉降研究现状，并对今后的研究前景提出了展望。     

The effects of elevated CO2 on symbiotic N2 fixation: a link between the carbon and nitrogen cycles in grassland ecosystems

The response of plants to elevated CO₂ is dependent on the availability of nutrients, especially nitrogen. It is generally accepted that an increase in the atmospheric CO₂ concentration increases the C:N ratio of plant residues and exudates. This promotes temporary N-immobilization which might, in turn, reduce the availability of soil nitrogen. In addition, both a CO₂ stimulated increase in plant growth (thus requiring more nitrogen) and an increased N demand for the decomposition of soil residues with a large C:N will result under elevated CO₂ in a larger N-sink of the whole grassland ecosystem. One way to maintain the balance between the C and N cycles in elevated CO₂ would be to increase N-import to the grassland ecosystem through symbiotic N₂ fixation. Whether this might happen in the context of temperate ecosystems is discussed, by assessing the following hypothesis: i) symbiotic N₂ fixation in legumes will be enhanced under elevated CO₂, ii) this enhancement of N₂ fixation will result in a larger N-input to the grassland ecosystem, and iii) a larger N-input will allow the sequestration of additional carbon, either above or below-ground, into the ecosystem. Data from long-term experiments with model grassland ecosystems, consisting of monocultures or mixtures of perennial ryegrass and white clover, grown under elevated CO₂ under free-air or field-like conditions, supports the first two hypothesis, since: i) both the percentage and the amount of fixed N increases in white clover grown under elevated CO₂, ii) the contribution of fixed N to the nitrogen nutrition of the mixed grass also increases in elevated CO₂. Concerning the third hypothesis, an increased nitrogen input to the grassland ecosystem from N₂ fixation usually promotes shoot growth (above-ground C storage) in elevated CO₂. However, the consequences of this larger N input under elevated CO₂ on the below-ground carbon fluxes are not fully understood. On one hand, the positive effect of elevated CO₂ on the quantity of plant residues might be overwhelming and lead to an increased long-term below-ground C storage; on the other hand, the enhancement of the decomposition process by the N-rich legume material might favour carbon turn-over and, hence, limit the storage of below-ground carbon.     

Rising sea level,temperature, and precipitation impact plant and ecosystem responses to elevated CO2 on a Chesapeake Bay wetland: review of a 28‐year study

An ongoing field study of the effects of elevated atmospheric CO₂ on a brackish wetland on Chesapeake Bay, started in 1987, is unique as the longest continually running investigation of the effects of elevated CO₂ on an ecosystem. Since the beginning of the study, atmospheric CO₂ increased 18%, sea level rose 20 cm, and growing season temperature varied with approximately the same range as predicted for global warming in the 21st century. This review looks back at this study for clues about how the effects of rising sea level, temperature, and precipitation interact with high atmospheric CO₂ to alter the physiology of C3 and C4 photosynthetic species, carbon assimilation, evapotranspiration, plant and ecosystem nitrogen, and distribution of plant communities in this brackish wetland. Rising sea level caused a shift to higher elevations in the Scirpus olneyi C3 populations on the wetland, displacing the Spartina patens C4 populations. Elevated CO₂ stimulated carbon assimilation in the Scirpus C3 species measured by increased shoot and root density and biomass, net ecosystem production, dissolved organic and inorganic carbon, and methane production. But elevated CO₂ also decreased biomass of the grass, S. patens C4. The elevated CO₂ treatment reduced tissue nitrogen concentration in shoots, roots, and total canopy nitrogen, which was associated with reduced ecosystem respiration. Net ecosystem production was mediated by precipitation through soil salinity: high salinity reduced the CO₂ effect on net ecosystem production, which was zero in years of severe drought. The elevated CO₂ stimulation of shoot density in the Scirpus C3 species was sustained throughout the 28 years of the study. Results from this study suggest that rising CO₂ can add substantial amounts of carbon to ecosystems through stimulation of carbon assimilation, increased root exudates to supply nitrogen fixation, reduced dark respiration, and improved water and nitrogen use efficiency.     

短期氮素添加和模拟放牧对青藏高原高寒草甸生态系统呼吸的影响

氮沉降和放牧是影响草地碳循环过程的重要环境因子,但很少有研究探讨这些因子交互作用对生态系统呼吸的影响。在西藏高原高寒草甸地区开展了外源氮素添加与刈割模拟放牧实验,测定了其对植物生物量分配、土壤微生物碳氮和生态系统呼吸的影响。结果表明:氮素添加显著促进生态系统呼吸,而模拟放牧对其无显著影响,且降低了氮素添加的刺激作用。氮素添加通过提高微生物氮含量和土壤微生物代谢活性,促进植物地上生产,从而增加生态系统的碳排放;而模拟放牧降低了微生物碳含量,且降低了氮素添加的作用,促进根系的补偿性生长,降低了氮素添加对生态系统碳排放的刺激作用。这表明,放牧压力的存在会抑制氮沉降对高寒草甸生态系统碳排放的促进作用,同时外源氮输入也会缓解放牧压力对高寒草甸生态系统生产的负面影响。     

Mechanisms of plant species impacts on ecosystem nitrogen cycling

Plant species are hypothesized to impact ecosystem nitrogen cycling in two distinctly different ways. First, differences in nitrogen use efficiency can lead to positive feedbacks on the rate of nitrogen cycling. Alternatively, plant species can also control the inputs and losses of nitrogen from ecosystems. Our current understanding of litter decomposition shows that most nitrogen present within litter is not released during decomposition but incorporated into soil organic matter. This nitrogen retention is caused by an increase in the relative nitrogen content in decomposing litter and a much lower carbon‐to‐nitrogen ratio of soil organic matter. The long time lag between plant litter formation and the actual release of nitrogen from the litter results in a bottleneck, which prevents feedbacks of plant quality differences on nitrogen cycling. Instead, rates of gross nitrogen mineralization, which are often an order of magnitude higher than net mineralization, indicate that nitrogen cycling within ecosystems is dominated by a microbial nitrogen loop. Nitrogen is released from the soil organic matter and incorporated into microbial biomass. Upon their death, the nitrogen is again incorporated into the soil organic matter. However, this microbial nitrogen loop is driven by plant‐supplied carbon and provides a strong negative feedback through nitrogen cycling on plant productivity. Evidence supporting this hypothesis is strong for temperate grassland ecosystems. For other terrestrial ecosystems, such as forests, tropical and boreal regions, the data are much more limited. Thus, current evidence does not support the view that differences in the efficiency of plant nitrogen use lead to positive feedbacks. In contrast, soil microbes are the dominant factor structuring ecosystem nitrogen cycling. Soil microbes derive nitrogen from the decomposition of soil organic matter, but this microbial activity is driven by recent plant carbon inputs. Changes in plant carbon inputs, resulting from plant species shifts, lead to a negative feedback through microbial nitrogen immobilization. In contrast, there is abundant evidence that plant species impact nitrogen inputs and losses, such as: atmospheric deposition, fire‐induced losses, nitrogen leaching, and nitrogen fixation, which is driven by carbon supply from plants to nitrogen fixers. Additionally, plants can influence the activity and composition of soil microbial communities, which has the potential to lead to differences in nitrification, denitrification and trace nitrogen gas losses. Plant species also impact herbivore behaviour and thereby have the potential to lead to animal‐facilitated movement of nitrogen between ecosystems. Thus, current evidence supports the view that plant species can have large impacts on ecosystem nitrogen cycling. However, species impacts are not caused by differences in plant quantity and quality, but by plant species impacts on nitrogen inputs and losses.     

Carbon dioxide increase: the implications at the ecosystem level*

Abstract Possible effects of increased atmospheric concentrations of CO₂ on forest ecosystems are discussed and as an example a simulation case study using a set of mixed-age and mixed-species forest stand models is presented. The responses of the models to a simple scenario (uniform growth increase of all trees as a response to CO₂ enrichment) include increases in biomass that are considerably less than the increases in growth rate of the trees. These simulations and more general discussion of the possible effects of increased photosynthetic production identify the problem of scaling-up small time-scale and space-scale measurements of plant responses to CO₂ enrichment to the ecosystem level.     

基于稳定碳同位素技术研究大气CO2浓度升高对植物-土壤系统碳循环的影响

大气CO₂浓度升高影响植物光合作用过程和生物量积累,改变植物地上和地下生物量的动态分配.土壤有机质的形成和周转依赖于植物组分的输入,因此,CO₂浓度升高所造成的植物生理和代谢的变化对土壤碳库收支平衡具有重要影响.采用稳定碳同位素(¹³C)技术研究土壤-植物系统的碳循环可阐明大气CO₂浓度升高条件下光合碳在植物各器官的分配特征和时间动态,明确光合碳在土壤中的积累、分解与迁移转化过程以及对土壤有机碳库周转的影响.本文综述了基于¹³C自然丰度法或¹³C示踪技术研究大气CO₂浓度升高对土壤-植物系统碳循环的影响,主要包括:1)对植物光合作用的同位素分馏的影响;2)对植物光合碳(新碳)分配动态的影响;3)对土壤有机碳新老碳库动态以及微生物转化过程的影响.明确上述过程及其调控机制可为预测CO₂浓度升高对陆地生态系统碳循环及源汇效应的长期影响奠定基础.