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.     

阔叶红松林下6种早夏草本不同生长期生物量分配及模型构建

早夏草本植物作为阔叶红松林林下的一类优势物种,对整个生长季林下草本物种多样性和生物量具有重要影响,对其不同生长阶段生物量分配特征及生物量模型的研究有助于了解该类植物生存策略和碳汇储量。以吉林省蛟河地区阔叶红松林林下的白花碎米荠(Cardamine leucantha)、北重楼(Paris verticillata)、鹿药(Smilacina japonica)、美汉草(Meehania fargesii)、山茄子(Anisodus acutangulus)和紫花变豆菜(Sanicula rubriflora)6种早夏草本植物为研究对象,从4月初到8月底对其物候期进行观察记录,定期取样,从而对6种植物不同生长时期各组分生物量分配特征,地上与地下生物量相对生长关系进行分析研究,并以株高级为自变量建立5种形式(一元线性模型、一元二次模型、指数模型、幂函数模型、对数模型)的单种和混种生物量模型,选取最优模型进行验证。结果表明,6种早夏草本植物花期一般开始于4月底结束于6月中旬,果期开始于5月底结束于8月中旬,不同植物的花、果期持续时间差异较大。生长期内,随着植物生长,株高、生物量和根冠比(R/S)变化明显,但变化趋势不一致。不同物种各组分生物量分配存在差异,用于繁殖的生物量分配比例较小,通常不超过5%。所有物种AGB和BGB间均具有明显的相对生长关系(P0.0001),且均表现为异速生长(相关生长指数a≠1)。根据R~2和SEE选取最优生物量模型,其中幂函数模型形式最常用,其次是一元二次和指数模型。所有最优模型的R~2均较高且SEE较低,拟合效果较好,其中AGB和TB的最优模型要优于BGB,单种模型优于混种模型;通过验证,除混合模型BGB的RMA(30.679%)稍大于30%外,所有模型的RS、EE和RMA均小于30%,P值均大于80%,说明所建立的最优模型能够用来对该地区阔叶红松林林下早夏草本植物生物量进行估算。     

A review of belowground biomass allocation and its response to global climatic change in grassland ecosystems

Biomass allocations between aboveground and belowground organs provide pivotal information for connecting aboveground productivity and belowground carbon sequestration. As accurate measurement of belowground biomass is essential for determining the biomass allocation, we first reviewed the methods in quantifying belowground biomass and their merits. We then presented the major advances on plant biomass allocations between aboveground and belowground organs, as well as the potential drivers such as precipitation, warming, atmospheric CO₂ concentration, and nitrogen deposition. We finally provided a list of challenges in studying belowground biomass allocation for the future. This review has important implications for studies on carbon cycling in grassland ecosystems under the changing climate.     

Climate factors affect forest biomass allocation by altering soil nutrient availability and leaf traits

Biomass in forests sequesters substantial amounts of carbon; although the contribution of aboveground biomass has been extensively studied, the contribution of belowground biomass remains understudied. Investigating the forest biomass allocation is crucial for understanding the impacts of global change on carbon allocation and cycling.Moreover, the question of how climate factors affect biomass allocation in natural and planted forests remains unresolved. Here, we addressed this question by coll...     

温度和降水对森林生物量分配的影响研究进展

森林生物量分配策略是全球变化背景下群落保持生产力的重要机制.温度和降水会影响森林生物量的分配格局.文章基于文献分析,总结了增温、低温和降水对森林地上、地下生物量分配的影响机制,以及温度和降水对森林生物量分配的交互作用,并对未来温度和降水影响森林生物量分配的研究进行了展望,提出该领域今后的研究重点为:(1)加强生物量分配...     

全球森林土壤微生物生物量碳氮磷化学计量的季节动态

土壤微生物生物量在森林生态系统中充当具有生物活性的养分积累和储存库。土壤微生物转化有机质为植物提供可利用养分, 与植物的相互作用维系着陆地生态系统的生态功能。同时, 土壤微生物也与植物争夺营养元素, 在季节交替过程和植物的生长周期中呈现出复杂的互利-竞争关系。综合全球数据对温带、亚热带和热带森林土壤微生物生物量碳(C)、氮(N)、磷(P)含量及其化学计量比值的季节动态进行分析, 发现温带和亚热带森林的土壤微生物生物量C、N、P含量均呈现夏季低、冬季高的格局。热带森林四季的土壤微生物生物量C、N、P含量都低于温带和亚热带森林, 且热带森林土壤微生物生物量C含量、N含量在秋季相对最低, 土壤微生物生物量P含量四季都相对恒定。温带森林的土壤微生物生物量C:N在春季显著高于其他两个森林类型; 热带森林的土壤微生物生物量C:N在秋季显著高于其他2个森林类型。温带森林土壤微生物生物量N:P和C:P在四季都保持相对恒定, 而热带森林土壤微生物生物量N:P和C:P在夏季高于其他3个季节。阔叶树的土壤微生物生物量C含量、N含量、N:P、C:P在四季都显著高于针叶树; 而针叶树的土壤微生物生物量P含量在四季都显著高于阔叶树。在春季和冬季时, 土壤微生物生物量C:N在阔叶树和针叶树之间都没有显著差异; 但是在夏季和秋季, 针叶树的土壤微生物生物量C:N显著高于阔叶树。对于土壤微生物生物量的变化来说, 森林类型是主要的显著影响因子, 季节不是显著影响因子, 暗示土壤微生物生物量的季节波动是随着植物其内在固有的周期变化而变化。植物和土壤微生物密切作用表现出来的对养分的不同步吸收是保留养分和维持生态功能的一种权衡机制。     

青藏高原东部高寒灌丛生物量分配对模拟增温的响应

植物生物量分配特征的变化反映了不同环境条件下植物的适应策略,全球气候变暖正在改变青藏高原高寒生态系统植被动态和生物量分配格局。然而,到目前为止,有关青藏高原高寒灌丛生物量分配特征对气候变暖的响应研究较少。为了探究气候变暖对高寒灌丛生物量分配的影响,以青藏高原东部典型的窄叶鲜卑花高寒灌丛为研究对象,分析了高寒灌丛灌木层、草本层和群落水平生物量分配特征对开顶式生长室（OTC）模拟增温的响应。研究结果表明：整个生长季节,模拟增温使空气温度和表层土壤温度分别升高0.6℃和1.2℃,使表层土壤水分含量下降2.7%。模拟增温使草本层和群落地上生物量显著增加57.8%和7.2%,使灌木层、草本层和群落根系生物量显著增加42.5%、105.6%和45.6%。然而,模拟增温没有显著影响灌木层地上生物量。同时,模拟增温使灌木层、草本层和群落总生物量显著增加25.6%、85.7%和28.4%,使灌木层、草本层和群落根冠比显著增加33.2%、30.4%和36.0%。由此可见,模拟增温在促进高寒灌丛生物量生产的同时将显著提高向地下根系部分的分配比例。Pearson相关分析表明,高寒灌丛生物量分配与空气温度、土壤温度和土壤硝态氮含量呈显著正相关关系;多元线性回归分析结果也表明,空气温度、土壤温度和土壤硝态氮含量解释了高寒灌丛生物量分配变异的50.8%以上。这些结果表明,青藏高原东部高寒灌丛植被能够通过调节生物量分配模式应对未来气候变暖。     

Estimation of biomass allocation and carbon density of Rhododendron simsii shrubland in the subtropical mountainous areas of China

Aims As an important potential carbon sink, shrubland ecosystem plays a vital role in global carbon balance and climate regulation. Our objectives were to derive appropriate regression models for shrub biomass estimation, and to reveal the biomass allocation pattern and carbon density in Rhododendron simsii shrubland.
Methods We conducted investigations in 27 plots, and developed biomass regression models for shrub species to estimate shrub biomass. The biomass of herb and litterfall were obtained through harvesting. Plant samples were collected from each plot to measure carbon content in different organs.
Important findings The results showed that the power and linear models were the most appropriate equation forms. The D and D²H (where D was the basal diameter (cm) and H was the shrub height (m)) were good predictors for organ biomass and total biomass of shrubs. All of the biomass models reached extremely significant level, and could be used to estimate shrub biomass with high accuracy. It was more difficult to predict leaf and annual branch biomass than stem biomass, because leaf and annual branch were susceptible to herbivores and inter-plant competition. The mean biomass of the shrub layer was 20.78 Mg·hm^-2, in which Rhododendron simsii and Symplocos paniculata biomass accounted for 93.63%. Influenced by both environment and species characteristics, the biomass of the shrub layer organs was in the order of stem > root > leaf > annual branch. The root:shoot ratio of the shrub layer was 0.32, which was less than other shrubs in subtropical regions. The relative higher aboveground biomass allocation reflected the adaptation of plants to the warm and humid environment for more photosynthesis. The mean total community biomass was 26.26 Mg·hm^-2, in which shrub layer, herb layer and litter layer accounted for 79.14%, 7.62% and 13.25%, respectively. Litter biomass was relatively high, which suggested that this community had high nutrient return. There were significant correlations among aboveground biomass, belowground biomass and total biomass of shrub layer and herb layer. The mean biomass carbon density of the community was 11.70 Mg·hm^-2 and the carbon content ratio was 44.55%. The carbon density was usually obtained using the conversion coefficient of 0.5 in previous studies, which could overestimate carbon density by 12.22%.     

青藏高原高寒灌丛生态系统草本层生物量分配格局

青藏高原高寒灌丛生态系统生物量分配的研究相对较少,尤其是其草本层。为了探究高寒灌丛生态系统草本层生物量分配特征及其影响因素,分析了青藏高原东北部灌丛生态系统的49个高寒灌丛样地的草本层地上与地下生物量特征及其气候因子之间的关系。结果表明1)草本层地上生物量与地下生物量分别为121.1,342.8 g/m2均大于高寒草地的地上生物量与地下生物量。2)草本层的根冠比为3.6低于高寒草地的根冠比。3)地上生物量与地下生物量之间呈现幂函数的关系y=8.0x0.83(R2=0.48,P0.001)。4)根冠比与年均温度、年均降雨量之间没有显著的相关关系。     

Toward an allocation scheme for global terrestrial carbon models

The distribution of assimilated carbon among the plant parts has a profound effect on plant growth, and at a larger scale, on terrestrial biogeochemistry. Although important progress has been made in modelling photosynthesis, less effort has been spent on understanding the carbon allocation, especially at large spatial scales. Whereas several individual-level models of plant growth include an allocation scheme, most global terrestrial models still assume constant allocation of net primary production (NPP) among plant parts, without any environmental coupling. Here, we use the CASA biosphere model as a platform for exploring a new global allocation scheme that estimates allocation of photosynthesis products among leaves, stems, and roots depending on resource availability. The philosophy underlying the model is that allocation patterns result from evolved responses that adjust carbon investments to facilitate capture of the most limiting resources, i.e. light, water, and mineral nitrogen. In addition, we allow allocation of NPP to vary in response to changes in atmospheric CO₂. The relative magnitudes of changes in NPP and resource-use efficiency control the response of root:shoot allocation. For ambient CO₂, the model produces realistic changes in above-ground allocation along productivity gradients. In comparison to the CASA standard estimate using fixed allocation ratios, the new allocation scheme tends to favour root allocation, leading to a 10% lower global biomass. Elevated CO₂, which alters the balance between growth and available resources, generally leads to reduced water stress and consequently, decreased root:shoot ratio. The major exception is forest ecosystems, where increased nitrogen stress induces a larger root allocation.     

Modelling below- and above-ground biomass for non-woody and woody plants

BACKGROUND AND AIMS: Intraspecific relationships between below- and above-ground biomass (MB and MA, respectively) have been studied extensively to evaluate environmental effects on growth and development at the level of the individual plant. However, no current theoretical model for this relationship exists for broad interspecific trends. The aims of this paper are to provide a model and to test its predictions using a recently assembled, large database (1406 data entries for 257 species). METHODS: An allometric model was derived to predict the relationship between MB and MA for non-woody and woody plants based on previously developed scaling relationships for leaf, stem and root standing biomass and annual growth rates. The predictions of this model were tested by comparing the numerical values of predicted scaling exponents (the slopes of log-log regression curves) with those observed for the database. KEY RESULTS AND CONCLUSIONS: For non-woody plants and the juveniles of woody species, the model predicts an isometric scaling relationship (i.e. MB proportional, variant MA). For woody plants, a complex scaling function is predicted. But, for a particular set of biologically reasonable conditions, the model predicts MB proportional, variant MA across woody plants. These predictions accord reasonably well with observed statistical trends when non-woody and woody plants are studied separately (n=1061 and 345 data entries, respectively). Although the reliability of regression formulas to estimate MB based on MA measurements increased with increasing plant size, estimates of MB can be as much as two orders of magnitude off, even when using regression formulas with r2 >0.90 and F >53,000.     

实验增温对陆地生态系统根系生物量的影响

根系是植物吸收土壤水分和养分的重要器官, 驱动着多个生态系统过程, 该研究揭示了实验增温对根系生物量的影响及机制, 可为气候变暖背景下土壤碳动态和生态系统过程的变化提供理论依据。该研究从已发表的151篇国内外研究论文中收集到611组数据, 通过整合分析(meta-analysis)方法研究了实验增温对根系生物量(根系总生物量、粗根生物量、细根生物量、根冠比)的影响, 并探讨了增温幅度、增温年限、增温方式的影响, 以及根系生物量对增温的响应与本底环境条件(生态系统类型、年平均气温、年降水量、干旱指数)的关系。结果表明: (1)模拟增温使细根生物量显著增加8.87%, 而对根系总生物量、粗根生物量、根冠比没有显著影响; (2)中等强度增温(1-2 ℃)使得细根生物量和根冠比分别提高14.57%和23.63%; 中短期增温实验(<5年)对细根生物量具有促进影响, 而长期增温实验(≥5年)使细根生物量有降低的趋势; 开顶箱增温和红外辐射增温分别使细根生物量显著提高了17.50%和12.16%, 而电缆加热增温使细根生物量和粗根生物量显著降低了23.44%和43.23%; (3)不同生态系统类型对于增温响应不一致, 模拟增温使苔原生态系统细根生物量显著提高了21.03%, 细根生物量对增温的响应与本底年平均气温、年降水量、干旱指数均呈显著负相关关系。     

帽儿山温带落叶阔叶林通量塔风浪区生物量空间格局

采用网格法在帽儿山温带落叶阔叶林通量塔风浪区(1500 m×400 m)内设置直径为20 m的圆形样地106个,运用地统计学方法和回归分析法研究了乔木生物量空间格局及其驱动因子。结果表明,风浪区总生物量平均值为153.63 Mg/hm~2,变异系数为37.89%;根冠比平均0.25(变化范围0.18—0.36)。总生物量、地上生物量和地下生物量的空间自相关显著,半方差模型的结构比分别为0.50、0.61和0.50,空间异质性尺度分别为276 m、198 m和375 m。硬阔叶林与杂木林的生物量组分和根冠比差异均不显著,但以胸高断面积(BA)为协变量,生物量组分差异显著。硬阔叶林和杂木林生物量组分与BA均呈极显著的线性正相关关系,BA可以解释总生物量和地上生物量空间变异的85%以上,表明局域尺度上BA可作为森林乔木生物量的预测因子。两种林型的生物量与优势高呈对数线性关系,但相关程度较低(R~20.41)。杂木林的各生物量组分与坡度显著正相关,但硬阔叶林的关系不显著。帽儿山落叶阔叶林乔木生物量受BA、优势高、林型、坡度和坡向共同驱动而存在空间变异,因此在整合通量塔与地面碳汇测量时需要考虑空间异质性。     

森林土壤呼吸及其对全球变化的响应

森林土壤呼吸是全球碳循环的重要流通途径之一 ,其动态变化将直接影响全球 C平衡。森林土壤呼吸由自养呼吸和异养呼吸组成 ,不同森林类型、测定季节和测定方法等直接影响其所占比例。土壤温度和湿度是影响森林土壤呼吸的最主要因素 ,共同解释了森林土壤呼吸变化的大部分。因树种组成、生产力和枯落物数量等不同而使不同森林类型土壤呼吸速率表现出明显差异。采伐对森林土壤呼吸的影响结果有增加、降低或无影响 ,因采伐方式、森林类型、采伐迹地上植被恢复进程和气候条件等而异。火烧一般导致土壤呼吸速率降低。因肥料种类、施用剂量和立地条件不同 ,施肥对森林土壤呼吸的影响出现增加、降低或无影响等不同结果。大气 CO2 浓度升高和升温均可促进森林土壤呼吸。 N沉降有可能刺激了土壤呼吸 ,而酸沉降则可能降低了土壤呼吸。臭氧浓度和 UV-B辐射强度亦会在一定程度上影响森林土壤呼吸。但目前全球变化对森林土壤呼吸的综合影响尚不清楚 ,深入探讨森林土壤呼吸的调控因素及其对全球变化和营林措施的响应等仍是今后努力的主要方向。     

全球环境变化对典型生态系统的影响研究:现状、挑战与发展趋势

随着全球环境变化和人类活动对生态系统影响的日益加深,生态系统结构和功能发生强烈变化,生态系统提供各类资源和服务的能力在显著下降。在这种背景下,全面认识生态系统的结构功能与全球环境变化的关系已成为当前生态学研究的热点之一。本文综述了全球环境变化对典型生态系统(包括森林生态系统、河口湿地生态系统、城市生态系统)影响以及全球环境变化适应的研究现状,分析研究面临的困难及挑战。在此基础上,提出对未来研究发展趋势的展望。在森林生态系统与全球环境变化研究上,未来应重视能更好模拟现实情景的、多因子、长期的全球环境变化控制试验,并注重不同生物地球化学循环之间的耦合作用。在湿地生态系统与全球环境变化研究上,未来应加强氮沉降、硫沉降及盐水入侵对湿地生态系统碳氮循环的影响,明晰滨海湿地的蓝碳功能,加强极端气候和人类干扰影响下湿地生态系统结构和功能变化及恢复力的研究。在城市生态系统与全球环境变化研究上,未来应深化城市生物地球化学循环机制研究,实现城市生态系统的人本需求侧重与转向,并开展典型地区长期、多要素综合响应研究。在全球环境变化适应研究上,未来应构架定量化、跨尺度的适应性评价体系,加强典型区域/部门的适应性研究以及适应策略实施的可行性研究,注重适应与减缓对策的关联研究及实施的风险评估。期望本综述为我国生态系统与全球环境变化研究提供一些参考。     

菌根真菌影响森林生态系统碳循环研究进展

森林生态系统在全球碳(C)储量中占据极为重要的地位。菌根真菌广泛存在于森林生态系统中,在森林生态系统C循环过程中发挥重要的作用。阐述了不同菌根类型真菌在森林生态系统C循环过程中的功能,对比了温带/北方森林与热带/亚热带森林中菌根真菌介导的C循环研究方面新近取得的研究结果。发现温带和北方森林的外生菌根(EcM)植物对地上生物量C的贡献相对较小,然而是地下C储量的主要贡献者;以丛枝菌根(AM)共生为主的热带/亚热带森林地表生物量占比较高,表明AM植被对热带/亚热带森林地上生物量C的贡献相对较大。我们还就全球变化背景下,菌根真菌及其介导的森林生态系统C汇功能,以及不同菌根类型树种影响C循环的机制等进行了总结。菌根真菌通过影响凋落物分解、土壤有机质形成及地下根系生物量,进而影响整个森林生态系统的C循环功能。菌根介导的森林C循环过程很大程度上取决于(优势)树木的菌根类型和森林土壤中菌根真菌的群落结构。最后指出了当前研究存在的主要问题以及未来研究展望。本文旨在明确菌根真菌在森林生态系统C循环转化过程中的重要生态功能,有助于准确地评估森林生态系统C汇现状,为应对全球变化等提供重要的依据。     

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.     

陆生植物生物量分配对模拟氮沉降响应的Meta分析

分析了陆生植物地上、地下各组织中生物量分配对氮沉降的响应,为研究大气氮沉降背景下陆地生态系统的碳、氮循环过程及植物生物量分配、立木收获、定向培育等相关研究和实践提供参考依据。共收集整理了国内外63篇论文的原始数据资料进行Meta分析(Meta-analysis),用以定量评估氮沉降对植物生物量分配的影响,并通过亚组分析进一步探讨了不同生态系统类型、植物种类、氮肥形式、施氮水平和持续时间对生物量分配的影响。结果表明,总体来看施氮会显著促进植物地上部分生物量分配,植物叶生物量和茎生物量在施氮条件下均显著增加;然而地下生物量所受促进作用要低于地上部分,表现为植物细根生物量和粗根生物量在氮输入下并没有显著变化;植物根冠比在氮沉降下显著降低;叶重比、茎重比和根重比在氮沉降下没有显著变化。此外,亚组分析结果表明生态系统类型和植物类型会显著影响植物总生物量和根冠比对氮沉降的响应,草本植物在氮沉降下的生物量累积明显优于木本,这说明短期氮沉降可能会增加草本的覆盖面积;施肥形式对根冠比的影响存在明显差异,相比于尿素,硝酸铵对植物根冠比的作用更显著;不同施氮水平显著影响地上生物量分配,中氮水平(本研究为60—120 kg hm-2a-1)促进作用最大,高氮水平(本研究为≥120 kg hm-2a-1)促进作用明显减弱,这与总生物量的变化一致,表明过高的氮沉降量将抑制植物生长;氮沉降处理时间长短对植物地上生物量的影响也存在显著差异,当施氮时间高于3年,氮沉降对地上生物量的促进作用几乎消失。总之,短期氮沉降会使植物分配更多生物量给地上部分,且氮沉降对草本植物生物量的累积作用明显优于木本,这些发现可为未来大气氮沉降背景下植物地上、地下部分碳存储、植物群落结构、植被动态等相关研究提供科学依据。