森林生态系统细根周转规律及影响因素

根系周转是陆地生态系统碳循环的关键过程,对研究土壤碳库变化及全球气候变化均具有重要意义。然而由于根系周转率的测量计算方法较多,不同方法得出的结果差异较大,且目前对全球区域尺度上森林生态系统根系周转的研究还不够充分,使得全球森林生态系统根系周转变化规律仍不清楚。该研究通过收集文献数据并统一周转率计算方法,对全球5种森林类型的细根周转空间格局进行整合,同时结合土壤理化性质和气候数据,得出影响森林生态系统细根周转的因子。结果表明,不同森林类型细根周转率存在显著差异,且随着纬度的升高逐渐降低;森林生态系统细根周转率与年平均温度和年平均降水量呈正相关;森林生态系统细根周转率与土壤有机碳含量呈正相关但与土壤pH值呈负相关。该研究为揭示森林生态系统细根周转规律及机制提供了科学依据。     

施氮肥对水曲柳人工林细根生产和周转的影响

细根周转与土壤养分密切相关,但由于根系研究方法的差异以及研究对象的不同,土壤养分对细根周转影响的研究存在不一致的结论。本文以水曲柳（Fraxinus mandshurica）人工林为对象,应用3种方法研究施氮肥对细根生产和周转的影响。结果表明：施肥降低了活细根现存生物量,但施氮肥样地细根年生产量平均值（93．105g·m^-2·a^-1）与对照样地（93．505g·m^-2·a^-1）没有差异,不同方法得出施氮肥样地细根平均周转率（0．917次·a^-1）大于对照样地（0．710次·a^-1）;不同土层内细根的生产量显著不同,表层生产量最大,土层越深细根生产量越低,但细根周转率一般随土壤加深而加快;不同的研究方法得出细根的年生产量和周转率差异较大,分室模型法最高,其次是内生长土芯法,极差法和积分法最低。     

中国草地生态系统根系周转的空间格局和驱动因子

根系周转是陆地生态系统物质循环的关键指标, 也是陆地生态系统净初级生产力及碳固持潜力估算的核心参数。然而, 由于地下净初级生产力数据获取困难, 区域和全球尺度上的相关研究十分有限, 尤其是分布广泛的中国草地, 区域尺度上的整合研究几乎为空白。基于样地实测数据、已发表文献和在线数据库数据, 对中国草地5种植被类型、共计154个草地生态系统根系周转的空间格局进行整合分析, 并结合气象和土壤数据, 揭示了草地生态系统根系周转的关键驱动因子。研究发现: (1)根系周转速率随纬度升高而降低, 低纬度温暖地区根系周转更快; (2)气候因子(年平均气温、年降水量)和土壤理化性质(砾石含量、容重、pH值)共同影响根系周转, 对周转变异性的解释度为44%, 其中气候因子的相对贡献率为57%, 土壤理化性质的相对贡献率为43%; (3)中国草地根系周转的格局和驱动因子与全球尺度的研究结果不尽相同。该研究对根系周转的驱动因子提出了新的观点和证据, 为全球尺度上的整合研究提供了关键数据。     

Spatial patterns and drivers of root turnover in grassland ecosystems in China

根系周转是陆地生态系统物质循环的关键指标, 也是陆地生态系统净初级生产力及碳固持潜力估算的核心参数。然而, 由于地下净初级生产力数据获取困难, 区域和全球尺度上的相关研究十分有限, 尤其是分布广泛的中国草地, 区域尺度上的整合研究几乎为空白。基于样地实测数据、已发表文献和在线数据库数据, 对中国草地5种植被类型、共计154个草地生态系统根系周转的空间格局进行整合分析, 并结合气象和土壤数据, 揭示了草地生态系统根系周转的关键驱动因子。研究发现: (1)根系周转速率随纬度升高而降低, 低纬度温暖地区根系周转更快; (2)气候因子(年平均气温、年降水量)和土壤理化性质(砾石含量、容重、pH值)共同影响根系周转, 对周转变异性的解释度为44%, 其中气候因子的相对贡献率为57%, 土壤理化性质的相对贡献率为43%; (3)中国草地根系周转的格局和驱动因子与全球尺度的研究结果不尽相同。该研究对根系周转的驱动因子提出了新的观点和证据, 为全球尺度上的整合研究提供了关键数据。     

高寒草甸植被细根生产和周转的比较研究

植物根系是陆地生态系统重要的碳汇和养分库,细根周转过程是陆地生态系统地下部分碳氮循环的核心环节,在陆地生态系统如何响应全球变化中起着关键作用。在全球变化敏感地区之一的青藏高原,对该地区的主要植被类型矮嵩草草甸同时采用根钻法、内生长袋法和微根管法3种观测方法研究细根生产和周转速率,并探讨了极差法、积分法、矩阵法和Kaplan-Meier法等数据处理方法对计算值的影响。研究结果显示:在估算细根净初级生产力时,根钻法宜采用积分法,内生长袋法宜选用矩阵法;由此进一步以最大细根生物量为基础,根钻法和内生长袋法估测的细根年周转速率分别为0.36 a-1和0.52 a-1,内生长袋法的估算结果是根钻法的1.44倍。对于微根管法,将其观测得到的细根长度转换为单位面积的生物量值后,采用积分法计算出细根周转速率为0.84 a-1,远高于传统方法的估算结果;若采用Kaplan-Meier生存分析方法,则计算出的细根周转速率更高达3.41 a-1。     

养分输入方式对森林生态系统土壤养分循环的影响研究进展

土壤养分循环对森林生态系统稳定性维持、树种选择及更新、可持续经营具有重要意义,掌握养分输入来源及过程可指导森林生态系统物质循环与能量流动分析及生态功能评估。凋落物、根系周转、根系分泌物是森林土壤养分的主要来源,是土壤养分循环的重要组成部分。本文分析了3种主要养分输入方式及其影响因素,总结了凋落物组成及理化性质、生物因子、环境因子等对凋落物分解及土壤养分循环的影响;综述了细根底物性质、树种组成、土壤生物、环境因子变化对细根周转及土壤养分循环的影响;探讨了根系分泌物对土壤养分循环过程的作用,基于此,提出了该领域亟需深入研究的重要方向,以期为相关研究及森林生态系统养分管理提供参考。     

林木细根寿命及其影响因子研究进展

 细根周转要消耗大量的C,它影响森林生态系统C分配格局与过程和养分循环,对生态系统生产力具有重要意义。细根的周转取决于细根的寿命,细根寿命越短,周转越快,根系对C的消耗也越多。大量研究表明,细根的寿命与地上部分C向根系供应的多少有密切关系,同时也与细根直径大小、土壤中N和水分的有效性、土壤温度以及根际周围的土壤动物和微生物的活动有关。本文综述了国外近年来在该领域里的研究进展,特别是对控制细根寿命的机理和主要影响因子进行了评述,目的是引起国内研究者的关注,促进我国根系生态学的研究与发展。     

微根管法和同位素法在细根寿命研究中的应用及比较

细根的生产和周转在陆地生态系统的碳和养分循环中起着重要作用,并且对全球环境变化具有一定的指示意义。细根寿命是估计细根周转的关键,其长短决定了养分和碳消耗与循环的速度。由于采用的研究方法不同,导致所得细根寿命估计值存在较大差异,目前最新的同位素和微根管2种方法之间寿命估计值差异可达10倍以上。本文对这2种研究方法的原理和优点进行了阐述,并从细根定义、细根寿命理论分布假设、细根取样误差等方面对导致这2种方法研究结果存在差异的原因进行分析,以期有助于今后根系研究的发展。     

应用微根管法测定细根指标方法评述

树木细根(直径<2mm)在森林生态系统能量流动和物质循环中起着重要的作用。原有的细根生产周转研究中常采用的土钻法、内生长法、挖掘法、根室法和土柱法等,均不能直接观察到细根的动态变化。微根管法是一种非破坏性、可定点直接观察和研究植物根系的方法,为研究细根的生长、衰老、死亡、分解和再生长的过程提供了有效的工具,尤其适用于细根周转、寿命和分解等方面的研究。但该技术不能直接测定单位面积的细根生物量、细根化学组成及细根周转对土壤碳和养分循环的影响,需要与土钻法结合。本文就运用微根管法对细根生物量、生产、周转和寿命等指标的研究方法进行了评述。     

氮沉降对森林生态系统土壤碳库的影响

森林土壤碳库是陆地生态系统碳库的重要组成部分,对维持全球碳平衡具有重要意义。不断加剧的全球氮沉降有可能改变森林生态系统中碳元素的地球化学循环过程,从而引起森林土壤碳储量的变化。本文从森林土壤碳收支的角度,将氮沉降对森林生态系统土壤碳库影响的复杂过程划分为凋落物分解、细根周转、土壤呼吸和土壤可溶性有机碳淋失4个相对独立的过程。综合国内外研究现状,对其进行了简要评述,指出了目前研究的不足,并探讨了这一研究领域的发展方向。     

Fine Root Biomass,Production, Turnover Rates,and Nutrient Contents in Boreal Forest Ecosystems in Relation to Species,Climate, Fertility,and Stand Age: Literature Review and Meta-Analyses

Fine roots <2 mm in diameter play a key role in regulating the biogeochemical cycles of ecosystems and are important to our understanding of ecosystem responses to global climate changes. Given the sensitivity of fine roots, especially in boreal region, to climate changes, it is important to assess whether and to what extent fine roots in this region change with climates. Here, in this synthesis, a data set of 218 root studies were complied to examine fine root patterns in the boreal forest in relation to site and climatic factors. The mean fine root biomass in the boreal forest was 5.28 Mg ha^?1, and the production of fine roots was 2.82 Mg ha^?1 yr^?1, accounting for 32% of annual net primary production of the boreal forest. Fine roots in the boreal forest on average turned over 1.07 times per year. Fine roots contained 50.9 kg ha^?1 of nitrogen (N) and 3.63 kg ha^?1 of phosphorous (P). In total, fine roots in the boreal forest ecosystems contain 6.1 × 10⁷ Mg N and 4.4×10⁶Mg P pools, respectively, about 10% of the global nutrients of fine roots. Fine root biomass, production, and turnover rate generally increased with increasing mean annual temperature and precipitation. Fine root biomass in the boreal forest decreased significantly with soil N and P availability. With increasing stand age, fine root biomass increased until about 100 years old for forest stands and then leveled off or decreased thereafter. These results of meta analysis suggest that environmental factors strongly influence fine root biomass, production, and turnover in boreal forest, and future studies should place a particular emphasis on the root-environment relationships.     

Global patterns of root turnover for terrestrial ecosystems

Root turnover is a critical component of ecosystem nutrient dynamics and carbon sequestration and is also an important sink for plant primary productivity. We tested global controls on root turnover across climatic gradients and for plant functional groups by using a database of 190 published studies. Root turnover rates increased exponentially with mean annual temperature for fine roots of grasslands ( r ² = 0.48) and forests ( r ² = 0.17) and for total root biomass in shrublands ( r ² = 0.55). On the basis of the best-fit exponential model, the Q ₁₀ for root turnover was 1.4 for forest small diameter roots (5 mm or less), 1.6 for grassland fine roots, and 1.9 for shrublands. Surprisingly, after accounting for temperature, there was no such global relationship between precipitation and root turnover. The slowest average turnover rates were observed for entire tree root systems (10% annually), followed by 34% for shrubland total roots, 53% for grassland fine roots, 55% for wetland fine roots, and 56% for forest fine roots. Root turnover decreased from tropical to high-latitude systems for all plant functional groups. To test whether global relationships can be used to predict interannual variability in root turnover, we evaluated 14 yr of published root turnover data from a shortgrass steppe site in northeastern Colorado, USA. At this site there was no correlation between interannual variability in mean annual temperature and root turnover. Rather, turnover was positively correlated with the ratio of growing season precipitation and maximum monthly temperature ( r ² = 0.61). We conclude that there are global patterns in rates of root turnover between plant groups and across climatic gradients but that these patterns cannot always be used for the successful prediction of the relationship of root turnover to climate change at a particular site.     

落叶松和水曲柳人工林细根生长、死亡和周转

 细根周转是陆地生态系统碳分配格局与过程的核心环节,而细根周转估计的关键是了解细根的生长和死亡动态。该研究以18年生落叶松(Larix gmelinii)和水曲柳(Fraxi nus mandshurica)人工林为对象,采用微根管（Minirhizotron）技术对两树种0～40 cm深度的细根生长和死亡动态进行了为期1年的观测,研究了两树种细根在不同土层深度的生长与死亡动态、细根周转以及与土壤有效氮含量、土壤温度、大气温度和降水的关系。结果表明：1） 落叶松平均细根生长（Root length density production, RLDP）0.0045 mm•cm^－2•d^－1）明显低于水曲柳RLDP（0.0077 mm•cm^－2•d^－1）。两个树种细根平均RLDP在表层（0～10 cm）最大,而底层（30～40 cm）最小 ,两树种平均细根死亡（Root length density mortality, RLDM）也表现同样规律 。水曲柳春季生长的细根占41.7%,夏季占39.7%,而落叶松细根生长分别是24.0%和51.2%,水曲柳细根死亡主要发生在春季（34.3%） 和夏季（34.0%）,而落叶松细根死亡主要发生在夏季和秋季（分别占28.5%和32.3%）,两 树种细根生长与死亡在冬季均较小;2）落叶松细根年生长量（0.94 mm•cm^－2•a^－1）和年死亡量（0.72 mm•cm^－2•a^－1）明显低于水曲柳（1.52和1.21 mm•cm^－2•a^－1）,两树种细根表层年生长量和年死亡量均最高,底层最低。落叶松细根年周转为3.1次•a^－1（按年生长量计算）和2.4次•a^－1（按年死亡量计算）,相比较,水曲柳细根年周转分别为2.7次•a^－1和2.2次•a^－1;3）土壤有效氮含量、土壤温度、大气温度和降水综合作用影响细根生长和死亡动态,可以解释细根生长80%的变异和细根死亡95%以上的变异。     

Increased topsoil carbon stock across China's forests

Biomass carbon accumulation in forest ecosystems is a widespread phenomenon at both regional and global scales. However, as coupled carbon–climate models predicted, a positive feedback could be triggered if accelerated soil carbon decomposition offsets enhanced vegetation growth under a warming climate. It is thus crucial to reveal whether and how soil carbon stock in forest ecosystems has changed over recent decades. However, large‐scale changes in soil carbon stock across forest ecosystems have not yet been carefully examined at both regional and global scales, which have been widely perceived as a big bottleneck in untangling carbon–climate feedback. Using newly developed database and sophisticated data mining approach, here we evaluated temporal changes in topsoil carbon stock across major forest ecosystem in China and analysed potential drivers in soil carbon dynamics over broad geographical scale. Our results indicated that topsoil carbon stock increased significantly within all of five major forest types during the period of 1980s–2000s, with an overall rate of 20.0 g C m^?2 yr^?1 (95% confidence interval, 14.1–25.5). The magnitude of soil carbon accumulation across coniferous forests and coniferous/broadleaved mixed forests exhibited meaningful increases with both mean annual temperature and precipitation. Moreover, soil carbon dynamics across these forest ecosystems were positively associated with clay content, with a larger amount of SOC accumulation occurring in fine‐textured soils. In contrast, changes in soil carbon stock across broadleaved forests were insensitive to either climatic or edaphic variables. Overall, these results suggest that soil carbon accumulation does not counteract vegetation carbon sequestration across China's forest ecosystems. The combination of soil carbon accumulation and vegetation carbon sequestration triggers a negative feedback to climate warming, rather than a positive feedback predicted by coupled carbon–climate models.     

Allometric constraints on,and trade‐offs in,belowground carbon allocation and their control of soil respiration across global forest ecosystems

To fully understand how soil respiration is partitioned among its component fluxes and responds to climate, it is essential to relate it to belowground carbon allocation, the ultimate carbon source for soil respiration. This remains one of the largest gaps in knowledge of terrestrial carbon cycling. Here, we synthesize data on gross and net primary production and their components, and soil respiration and its components, from a global forest database, to determine mechanisms governing belowground carbon allocation and their relationship with soil respiration partitioning and soil respiration responses to climatic factors across global forest ecosystems. Our results revealed that there are three independent mechanisms controlling belowground carbon allocation and which influence soil respiration and its partitioning: an allometric constraint; a fine‐root production vs. root respiration trade‐off; and an above‐ vs. belowground trade‐off in plant carbon. Global patterns in soil respiration and its partitioning are constrained primarily by the allometric allocation, which explains some of the previously ambiguous results reported in the literature. Responses of soil respiration and its components to mean annual temperature, precipitation, and nitrogen deposition can be mediated by changes in belowground carbon allocation. Soil respiration responds to mean annual temperature overwhelmingly through an increasing belowground carbon input as a result of extending total day length of growing season, but not by temperature‐driven acceleration of soil carbon decomposition, which argues against the possibility of a strong positive feedback between global warming and soil carbon loss. Different nitrogen loads can trigger distinct belowground carbon allocation mechanisms, which are responsible for different responses of soil respiration to nitrogen addition that have been observed. These results provide new insights into belowground carbon allocation, partitioning of soil respiration, and its responses to climate in forest ecosystems and are, therefore, valuable for terrestrial carbon simulations and projections.     

Turnover of Grassland Roots in Mountain Ecosystems Revealed by Their Radiocarbon Signature: Role of Temperature and Management

Root turnover is an important carbon flux component in grassland ecosystems because it replenishes substantial parts of carbon lost from soil via heterotrophic respiration and leaching. Among the various methods to estimate root turnover, the root’s radiocarbon signature has rarely been applied to grassland soils previously, although the value of this approach is known from studies in forest soils. In this paper, we utilize the root’s radiocarbon signatures, at 25 plots, in mountain grasslands of the montane to alpine zone of Europe. We place the results in context of a global data base on root turnover and discuss driving factors. Root turnover rates were similar to those of a subsample of the global data, comprising a similar temperature range, but measured with different approaches, indicating that the radiocarbon method gives reliable, plausible and comparable results. Root turnover rates (0.06–1.0 y^-1) scaled significantly and exponentially with mean annual temperatures. Root turnover rates indicated no trend with soil depth. The temperature sensitivity was significantly higher in mountain grassland, compared to the global data set, suggesting additional factors influencing root turnover. Information on management intensity from the 25 plots reveals that root turnover may be accelerated under intensive and moderate management compared to low intensity or semi-natural conditions. Because management intensity, in the studied ecosystems, co-varied with temperature, estimates on root turnover, based on mean annual temperature alone, may be biased. A greater recognition of management as a driver for root dynamics is warranted when effects of climatic change on belowground carbon dynamics are studied in mountain grasslands.     

植被恢复对西南喀斯特地区土壤气候韧性的提升作用

土壤质量提升是生态系统应对气候变化能力增强的关键。为探究气候变化背景下不同植被恢复方式对喀斯特地区土壤质量的影响,基于黔桂喀斯特地区气候梯度样带84个样方土壤物理、化学和生物性质综合分析,分别以耕地和次生林作为退化和顶级恢复对照,探讨了自然恢复(灌木林)和人工恢复(人工林)的土壤质量提升效应及其对气候变化的响应。结果表明:(1)植被恢复不仅显著提高了土壤细菌、真菌、放线菌等微生物丰度以及有机碳、全氮、速效氮等养分含量,也对土壤质地有一定改善;(2)自然恢复和人工恢复均提高了土壤质量,两种恢复方式之间土壤质量指数无显著差异,但与次生林依然存在差距。灌木林和人工林的土壤质量仅约为次生林土壤质量的62%-66%;(3)耕地土壤质量随年均温和年平均降雨量的增加而下降,次生林的土壤质量随年平均降雨量的增加而上升,植被恢复对土壤质量的提升率与年均温和年平均降雨量呈正相关关系。阐明了在一定范围的气候变化下进行植被恢复可以显著提升喀斯特地区土壤的气候韧性,揭示了喀斯特地区植被恢复对土壤质量的提升主要是由于提高了土壤碳氮养分含量及改善了土壤微生物群落结构,这为全球气候变化背景下喀斯特退化土地生态恢复和管理提供了理论依据。     

Estimation of fine root production, mortality and turnover with Minirhizotron in Larix gmelinii and Fraxinus mandshurica plantations

Fine root turnover is a major pathway for car-bon and nutrient cycling in forest ecosystems. However, to estimate fine root turnover, it is important to first understand the fine root dynamic processes associated with soil resource availability and climate factors. The objectives of this study were: (1) to examine patterns of fine root production and mortality in different seasons and soil depths in the Larix gmelinii and Fraxinus man-dshurica plantations, (2) to analyze the correlation of fine root production and mortality with environmental factors such as air temperature, precipitation, soil temperature and available nitrogen, and (3) to estimate fine root turn-over. We installed 36 Minirhizotron tubes in six mono-specific plots of each species in September 2003 in the Mao'ershan Experimental Forest Station. Minirhizotron sampling was conducted every two weeks from April 2004 to April 2005. We calculated the average fine root length, annual fine root length production and mortality using image data of Minirhizotrons, and estimated fine root turnover using three approaches. Results show that the average growth rate and mortality rate in L. melinii were markedly smaller than in F. mandshurica, and were high-est in the surface soil and lowest at the bottom among all the four soil layers. The annual fine root production and mortality in F. mandshurica were significantly higher than in L. gmelinii. The fine root production in spring and summer accounted for 41.7% and 39.7% of the total annual production in F. mandshurica and 24.0% and 51.2% in L. gmelinii. The majority of fine root mortality occurred in spring and summer for F. mandshurica and in summer and autumn for L. gmelinii. The turnover rate was 3.1 a-1 for L. gmelinii and 2.7 a-1 for F. mandshurica. Multiple regression analysis indicates that climate and soil resource factors together could explain 80% of the varia-tions of the fine root seasonal growth and 95% of the seasonal mortality. In conclusion, fine root production and mortality in L. gmelinii and F. mandshurica have dif-ferent patterns in different seasons and at different soil depths. Air temperature, precipitation, soil temperature and soil available nitrogen integratively control the dynamics of fine root production, mortality and turnover in both species.     

Measuring Fine Root Turnover in Forest Ecosystems

Development of direct and indirect methods for measuring root turnover and the status of knowledge on fine root turnover in forest ecosystems are discussed. While soil and ingrowth cores give estimates of standing root biomass and relative growth, respectively, minirhizotrons provide estimates of median root longevity (turnover time) i.e., the time by which 50% of the roots are dead. Advanced minirhizotron and carbon tracer studies combined with demographic statistical methods and new models hold the promise of improving our fundamental understanding of the factors controlling root turnover. Using minirhizotron data, fine root turnover (y⁻¹) can be estimated in two ways: as the ratio of annual root length production to average live root length observed and as the inverse of median root longevity. Fine root production and mortality can be estimated by combining data from minirhizotrons and soil cores, provided that these data are based on roots of the same diameter class (e.g., < 1 mm in diameter) and changes in the same time steps. Fluxes of carbon and nutrients via fine root mortality can then be estimated by multiplying the amount of carbon and nutrients in fine root biomass by fine root turnover. It is suggested that the minirhizotron method is suitable for estimating median fine root longevity. In comparison to the minirhizotron method, the radio carbon technique favor larger fine roots that are less dynamics. We need to reconcile and improve both methods to develop a more complete understanding of root turnover.     

Regional scale patterns of fine root lifespan and turnover under current and future climate

Fine root dynamics control a dominant flux of carbon from plants and into soils and mediate potential uptake and cycling of nutrients and water in terrestrial ecosystems. Understanding of these patterns is needed to accurately describe critical processes like productivity and carbon storage from ecosystem to global scales. However, limited observations of root dynamics make it difficult to define and predict patterns of root dynamics across broad spatial scales. Here, we combine species‐specific estimates of fine root dynamics with a model that predicts current distribution and future suitable habitat of temperate tree species across the eastern United States (US). Estimates of fine root lifespan and turnover are based on empirical observations and relationships with fine root and whole‐plant traits and apply explicitly to the fine root pool that is relatively short‐lived and most active in nutrient and water uptake. Results from the combined model identified patterns of faster root turnover rates in the North Central US and slower turnover rates in the Southeastern US. Portions of Minnesota, Ohio, and Pennsylvania were also predicted to experience >10% increases in root turnover rates given potential shifts in tree species composition under future climate scenarios while root turnover rates in other portions of the eastern US were predicted to decrease. Despite potential regional changes, the average estimates of root lifespan and turnover for the entire study area remained relatively stable between the current and future climate scenarios. Our combined model provides the first empirically based, spatially explicit, and spatially extensive estimates of fine root lifespan and turnover and is a potentially powerful tool allowing researchers to identify reasonable approximations of forest fine root turnover in areas where no direct observations are available. Future efforts should focus on reducing uncertainty in estimates of root dynamics by better understanding how climate and soil factors drive variability in root dynamics of different species.