碳供给与碳利用对树木生长的限制机制

树木生长固碳过程使森林生态系统成为减缓大气CO_2浓度升高的一个巨大而持续的碳汇。根据树木可利用碳的状况,限制树木生长的机制可分为碳供给限制和碳利用限制。许多环境因子交互作用,共同影响树木的碳供给与碳利用,因而很难量化碳供给和碳利用活动及其对环境变化敏感性对树木生长的影响。因此,从碳供给与碳利用角度揭示环境变化对树木生长影响的生理机制,对于预测全球变化背景下树木生长及森林碳汇功能至关重要。为此,该文介绍了树木生长碳供给与碳利用限制争议的相关背景;从碳供给与碳利用角度探讨了叶损失、干旱和低温等胁迫条件限制树木生长的生理机制;提出该领域今后应优先研究的3个问题:(1)探索非结构性碳水化合物(NSC)储存形成的调控机制,确定什么情况下以及多大程度上树木通过主动降低生长而将光合产物优先分配给NSC储存;(2)加强碳利用活动研究,系统测定光合产物在其碳利用组分的分配(特别是根系及其共生微生物活动);(3)开展树木碳代谢、矿质营养与水分生理的互作研究,充分认识树木碳、水和养分耦合关系及对树木生长的影响。     

虫害叶损失造成的树木非结构性碳减少与树木生长、死亡的关系研究进展

大规模虫害爆发可造成区域森林死亡, 近年的气候变化进一步增加了虫害的频度和危害程度。森林和林地植物死亡会导致植被生产力降低, 改变生态系统结构和功能, 使森林由一个净的碳汇转变为一个碳源。因此, 加深虫害对树木危害机制的认识有重要意义。虫害造成的叶损失(虫害叶损失)降低树木光合作用能力, 增加非结构性碳(NSC)消耗, 使得树木体内碳储备降低, NSC降低到一定程度会导致树木因碳饥饿而死亡。外部环境和树木自身的补偿性机制也会对这个过程产生正或负的影响。在近年气候变化背景下, 树木死亡在全球尺度上增多, 重新激起了人们对碳饥饿的重视, 碳饥饿被视为解释树木死亡的主要生理机制之一。该文介绍了碳饥饿的定义, 综述了虫害叶损失减少树木NSC储备与树木生长、死亡的关系, 以及树木虫害和叶损失与气候变化之间的关系, 并对今后的研究进行了展望。     

树木非结构性碳水化合物含量多时空尺度变化特征及其影响因素研究进展

非结构性碳水化合物(NSC)是树木生长和代谢的重要物质,对树木适应环境变化等具有重要作用。从时间尺度来看,受树木自身生长和碳储存策略影响,其NSC含量在年际尺度上变化不大。各个气候区树木NSC含量主控因素不同,其在季节尺度上差异明显。从空间尺度来看,在全球或大陆尺度上,受水热梯度影响,树木NSC含量随纬度降低总体下降,但变化不显著。而在地区尺度上,因水热梯度减小,采样频率低,树木NSC含量随纬度降低呈相反趋势。受树种特性及区域微生境影响,树木NSC含量随海拔升高的变化更加复杂。树木NSC含量在不同时空尺度的波动受多个生物和非生物因素影响。光合产物合成、呼吸作用消耗以及生长之间的权衡,决定了树木NSC含量的变化动态。不同研究方法各异,使得树木NSC含量在多时空尺度结论存在很大的不确定性。未来需要统一样品采集与分析标准,提高研究结果的可比性,并从整株入手,对不同树种及各个龄级的树木NSC含量在多时空尺度上进行测定,探讨NSC储存、转化与分配在树木生长和存活过程中的重要作用及其意义。     

施肥方式对幼龄楸树非结构性碳器官分配和生长季动态的影响

以6年生楸树无性系‘9 1’为试验材料,采用水肥一体化、穴施和不施肥(对照)3种施肥方法,分析施肥对楸树生长、各器官非结构性碳(NSC)含量及时间动态变化的影响,探究楸树NSC分配对施肥的响应机制,为评估楸树单株和林分碳储量提供理论依据。结果表明：(1)连续施肥4年后,与对照相比,穴施处理的树高和胸径分别提高了4.7%和7.1%,水肥一体化处理则分别提高了7.1%和20.5%,水肥一体化的施肥效果更佳。(2)不同施肥方式并未显著改变楸树各器官中可溶性糖含量,但是水肥一体化施肥明显提高了根中淀粉和总NSC的积累;可溶性糖和总NSC在叶和粗根中分布较多,淀粉则在根中含量较高。(3)不同施肥方式并未显著改变楸树各生长时期可溶性糖含量,但水肥一体化施肥明显提高了生长初期和末期淀粉和总NSC含量。研究发现,在整个生长季中,楸树叶中可溶性糖和总NSC一直被消耗,枝中可溶性糖一部分被消耗用于支持叶片生长,一部分以淀粉的形式进行储存,而根系接受源自叶和枝的NSC后转化为淀粉储存起来帮助树木抵抗低温环境;楸树遵循“碳消耗(初期) 碳消耗(中期) 碳积累(末期)”的NSC分配策略;应用水肥一体化技术可显著提高楸树生物量和生产力,在今后林木施肥试验中值得优先考虑。     

木本植物非结构性碳水化合物变化及其影响因子研究进展

非结构性碳水化合物(NSC)是植物新陈代谢过程中重要的供能物质,NSC库的动态变化不仅反映了植物体内的碳收支状况,还影响着植物的生理代谢活动.为了预测树木甚至整个森林生态系统对未来气候变化的响应和适应能力,本文综述了树木NSC研究的最新进展,重点介绍了NSC库的季节和区域性变化,分析总结了影响树木NSC含量和分配格局变化的主要环境因素.最后还对未来气候变化背景下树木NSC库可能的响应策略和适应状况进行了讨论,展望了未来树木NSC领域的研究方向.     

森林生态系统可溶性碳和颗粒碳通量

可溶性碳(Dissolved carbon,DC)和颗粒碳(particulate carbon,PC)通量作为森林生态系统碳收支的重要组分,在森林固碳功能的评价和模型预测中具有重要意义,但常因认识不足、测定困难等而在森林碳汇研究中被忽略。综述了森林生态系统DC和PC的组成、作用、相关生态过程及其影响因子,并展望了该领域应该优先考虑的研究问题。森林生态系统DC和PC主要包括可溶性有机碳、可溶性无机碳和颗粒有机碳,主要来源于生态系统的净初级生产量。DC和PC是森林土壤的活性碳库,主要以大气沉降、穿透雨和凋落物的形式输入森林土壤系统,并通过土壤呼吸、侧向运输及渗透流失的方式输出生态系统。从局域尺度看,DC和PC通量受根系分泌、细根分解、微生物周转等生物过程的影响较大;从区域尺度看,它们受土壤和植被特性、生态过程耦联关系、气候因子以及全球变化的综合影响。该领域应该优先考虑:(1)探索不同时空尺度下森林生态系统DC和PC通量的控制因子及其耦联关系,揭示其中的驱动机理;(2)探索DC和PC与其它森林生态系统碳组分的相互关系及转化,阐明DC和PC通量与其它养分之间潜在的生态化学计量关系;(3)探索全球变化,特别是人类活动(如森林经营)和极端干扰事件(如林火、旱涝、冰冻、冻融交替等)对森林生态系统DC和PC通量的影响。     

湖南省森林植被碳储量、碳密度动态特征

利用湖南省4次(1983—1987年、1990—1995年、2003—2004年和2009年)森林资源清查数据,采用材积源-生物量法,结合湖南省现有森林植被主要树种碳含量实测数据,研究近20多年来湖南省森林植被碳储量、碳密度的动态特征。结果表明:从1987年到2009年,湖南省乔木林植被碳汇为66.40×106tC,碳密度提高了5.65 tC/hm~2,阔叶林碳汇最大(48.43×10~6tC),其次是杉木林(9.54×10~6tC)和松木林(6.68×10~6tC),各乔木林植被碳密度波动较大;除过熟林外,各龄组乔木林均为碳汇,中龄林碳汇最大,幼龄林、中龄林、近熟林植被碳密度依次提高了4.75、4.09、0.83 tC/hm~2,成熟林、过熟林分别下降了6.87、13.88 tC/hm~2;天然林、人工林植被碳汇分别为41.01×10~6tC、25.39×10~6tC,碳密度分别提高了7.19、4.91 tC/hm~2。湖南省森林植被(包括疏林)碳汇为84.87×10~6tC,乔木林碳汇最大,其次是竹林,分别占湖南省森林植被碳汇的78.24%和33.31%,碳密度提高了6.24 tC/hm~2,各森林类型植被碳储量随其面积变化而变化。表明近20多年来,湖南省乔木林植被单位面积储碳能力明显提高,天然林在湖南省乔木林植被碳储量占有重要地位。     

中国陆地生态系统土壤有机碳变化研究进展

通过文献资料, 对中国陆地生态系统土壤有机碳变化研究进行评述. 20世纪80年代初至21世纪初, 中国森林、草地、灌丛和农田土壤有机碳库合计年均增加(71±19) Tg/a, 三江平原沼泽湿地垦殖导致土壤有机碳损失(6±2) Tg/a. 该结果存在极大的不确定性, 尤其是对森林、灌丛和草地碳库变化的估计. 未来研究需重点关注土地利用变化及其碳源、碳汇效应, 放牧管理对草地土壤有机碳库的影响, 灌丛和非森林树木(经济林、防护林及四旁绿化造林)土壤有机碳变化估算, 深层土壤有机碳变化的测定和估算, 中国土壤的固碳潜力及陆地生态系统碳收支模型开发.     

芦芽山不同海拔白杄非结构性碳水化合物含量动态

高山林线对环境变化具有高度的敏感性, 但林线形成机制仍然没有明确的结论。为了检验高山林线形成是由碳限制还是生长限制决定, 并探讨林线树种适应高山环境的生理生态机制, 选择山西省吕梁山脉北端芦芽山, 沿3个海拔梯度测定了林线树种白杄(Picea meyeri)各组织非结构性碳水化合物(NSC)及其组分含量。结果表明: 白杄总体及各组织NSC含量均随海拔升高而增加, 林线树木不存在碳限制; 白杄NSC源、汇均随海拔升高而增加, 源-汇比在3个海拔之间没有差异, 表明源-汇平衡关系对海拔的适应性, 林线树木碳源活动没有受到限制; 各组织中可溶性糖与淀粉的比值随海拔升高呈增大趋势, 说明树木生长的环境越寒冷, 树木组织中表现出越明显的保护策略, 也可能暗示林线区域的树木更多地受到生长限制。研究结果在一定程度上支持“生长限制”假说。     

长白山阔叶红松林3个主要树种的非结构性碳储存特征

非结构性碳水化合物(NSC,包括可溶性糖和淀粉)作为树木生命代谢的关键物质之一,在树木器官中的储存特征受到了广泛关注,但NSC在器官内部具有不同功能的组织间(韧皮部与木质部)的分配和权衡还不清楚.本研究以长白山阔叶红松林的3个优势树种——红松、水曲柳和紫椴为研究对象,对比分析NSC在根、树干韧皮与木质部中的浓度和分配特征.结果表明: 树木韧皮部和木质部间的NSC浓度差异显著,总体分配趋势为韧皮部以可溶性糖为主,而木质部以淀粉为主.树干外侧(以年轮划分,0～20年)、中段(20～40年)和内侧(>40年)的NSC浓度在不同树种间差异显著,而根中的差异不显著.红松和水曲柳树干韧皮部可溶性总糖浓度显著高于紫椴,在木质部中差异不显著.本研究结果表明,树体内部NSC在韧皮部和木质部上的分配存在明显分异,这与树种的演替阶段及组织的功能进化有关.研究结果对于深入理解温带树木的碳储存特征和分配机理具有参考意义.     

The relationship between the reduction of nonstructural carbohydrate induced by defoliator and the growth and mortality of trees

Large scale herbivorous insect outbreaks can cause death of regional forests, and the events are expected to be exacerbated with climate change. Mortality of forest and woodland plants would cause a series of serious consequences, such as decrease in vegetation production, shifts in ecosystem structure and function, and transformation of forest function from a net carbon sink into a net carbon source. There is thus a need to better understand the impact of insects on trees. Defoliation by insect pests mainly reduces photosynthesis (source decrease) and increases carbon consumption (sink increase), and hence causes reduction of nonstructural carbohydrate (NSC). When the reduction in NSC reaches to a certain level, trees would die of carbon starvation. External environment and internal compensatory mechanisms can also positively or negatively influence the process of tree death. At present, the research of carbon starvation is a hotspot because the increase of tree mortality globally with climate change, and carbon starvation is considered as one of the dominating physiological mechanisms for explaining tree death. In this study, we reviewed the definition of carbon starvation, and the relationships between the reduction of NSC induced by defoliation and the growth and death of trees, and the relationships among insect outbreaks, leaf loss and climate change. We also presented the potential directions of future studies on insect-caused defoliation and tree mortality.     

Advancement in studies of tree growth and ecophysiology incorporating micro-sampling approach

The recently developed micro-sampling approach has been widely used to extract micro-tree-cores at weekly intervals to monitor the process of stem cambial activity and xylem formation. Compared with the traditional dendrochronology, the micro-sampling approach enables us to better understand the inherent physiological processes in tree growth and their relationships with the environment at a more precise level. This review article aims to: 1) summarize the progresses in the micro-sampling approach-based studies published over recent years and its potential applications, and 2) elucidate the relationships between primary growth and secondary growth and the response mechanisms of radial growth of trees to global change (global warming, drought, and carbon and nitrogen fertilization effects) based on information from literature. It is anticipated that this review will assist with predicting productivity and carbon sink potential of forests, and help policy-makers with sustainable forest management decisions.     

Height-related growth declines in ponderosa pine are not due to carbon limitation

Decreased gas exchange as trees grow tall has been proposed to explain age-related growth declines in trees. We examined changes of mobile carbon stores (starch, sugars and lipids) with tree height in ponderosa pine ( Pinus ponderosa ) at two sites differing in water availability, and tested the following hypotheses: (1) carbon supply does not become increasingly limited as trees grow tall; rather, the concentration of mobile carbon compounds increases with tree height reflecting greater reductions of carbon sink activities relative to carbon assimilation; and (2) increases of stored mobile carbon compounds with tree height are greater in drier sites. Height-related growth reductions were associated with significant increases of non-structural carbohydrates (NSC) and lipid concentrations in all tissues in the upper canopy and of NSC in the bole. Lipid concentrations in the bole decreased with tree height, but such decrease is not necessarily inconsistent with non-limiting carbon supply in tall trees. Furthermore, we found stronger increases of mobile carbon stores with tree height at the dry site relative to the moist site. Our results provide first direct evidence that carbon supply does not limit growth in tall trees and that decreases of water availability might negatively impact growth processes more than net-photosynthesis.     

Season dynamics of carbon use efficiency and its influencing factors in the old-growth Abies fabri forest in Gongga Mountain,western Sichuan,China

碳利用效率(CUE)是植被生态系统的一个重要功能参数, 反映了植被生态系统的固碳能力, 适用于分析不同时间段内器官、个体和群落等不同层次的碳收支趋势, 因而有助于对陆地生态系统碳功能的确定与预测, 引起了广泛关注。该研究采用生物计量法, 测定和计算了川西贡嘎山东坡峨眉冷杉(Abies fabri)成熟林树木不同器官的呼吸与净生产力动态, 分析了乔木层及其各器官CUE动态及主要影响因子, 并估算了乔木层不同径级树木CUE。主要结果: (1)乔木层各器官月呼吸速率与温度呈正相关关系, 以细根月呼吸速率为最大; 不同径级树木年呼吸量无显著差异, 以小径级树木树干的年呼吸量为最小。(2)乔木层细根和树干月净初级生产力(NPP)均随温度增加而增加, 以细根月NPP为最大。小径级树木年NPP最大, 其针叶年NPP也显著高于中径级和大径级树木。(3)林分乔木层及其各器官CUE大多集中在0.30-0.60之间, 其中细根、树干CUE具有相似的月变化动态, 均随温度的升高而上升。不同径级树木CUE及树干和针叶CUE均随树木个体的增大而明显下降。(4)气温和土壤温度与乔木层树干和细根CUE呈正相关关系, 而降水量与针叶CUE呈负相关关系。细根CUE与树干CUE呈正相关关系,与针叶CUE呈负相关关系。峨眉冷杉成熟林乔木层CUE主要取决于树干和细根CUE。该研究证实了川西亚高山暗针叶成熟林仍具有较强的碳汇功能, 在区域碳储存和森林生态系统碳循环中发挥着极其重要的作用。     

Nitrogen and carbon source-sink relationships in trees at the Himalayan treelines compared with lower elevations

No single hypothesis or theory has been widely accepted for explaining the functional mechanism of global alpine/arctic treeline formation. The present study tested whether the alpine treeline is determined by (1) the needle nitrogen content associated with photosynthesis (carbon gain); (2) a sufficient source-sink ratio of carbon; or (3) a sufficient C-N ratio. Nitrogen does not limit the growth and development of trees studied at the Himalayan treelines. Levels of non-structural carbohydrates (NSC) in trees were species-specific and site-dependent; therefore, the treeline cases studied did not show consistent evidence of source/carbon limitation or sink/growth limitation in treeline trees. However, results of the combined three treelines showed that the treeline trees may suffer from a winter carbon shortage. The source capacity and the sink capacity of a tree influence its tissue NSC concentrations and the carbon balance; therefore, we suggest that the persistence and development of treeline trees in a harsh alpine environment may require a minimum level of the total NSC concentration, a sufficiently high sugar:starch ratio, and a balanced carbon source-sink relationship.     

A review on the effects of nitrogen and phosphorus addition on tree growth and productivity in forest ecosystems

Nitrogen (N) and phosphorus (P) inputs induced by anthropogenic activities and atmospheric N and P deposition have largely increased the availability of soil N and P in terrestrial ecosystems, which have considerably affected terrestrial carbon cycling processes. Tree growth and productivity in forest ecosystems play an important role in global carbon cycling, and determine the magnitude and direction of terrestrial carbon sequestration. Currently, a large number of field manipulation experiments have been conducted to investigate the effects of N and/or P addition on tree growth and forest productivity, but the results from these studies were inconsistent. Such inconsistent results might be affected by multiple factors, including biological, environmental and experimental variables. Here, we reviewed the present research status of the effects of N and P addition on tree growth and forest productivity in forest ecosystems based on three aspects, including the number of publications and experiments with field N and P addition, and the global distributions of these experiments. Then, we summarized the methods for assessing tree growth and forest productivity at ecosystem level in forest ecosystems, including relative growth rate and absolute increment. According to the related results, we reviewed the regulating factors that affect tree growth and productivity, and the potential mechanisms for such factors, including climate, tree size and stand age, plant functional traits (including type of tree-associated mycorrhizal fungi, N-fixation property of trees, and conservative and acquisitive functional traits), plant-microbe interaction, ambient nutrient (i.e., N and P) deposition rate, and experimental variables. Finally, we summarized the current studies, and pointed out five aspects that are urgently needed to provide further insights in future studies, including the physiological mechanism of how tree growth responds to N and P addition, the tradeoff and allocation among growth of various parts of tree under N and P addition, the role of plant functional traits in regulating and predicting the responses of tree growth to N and P addition, how the competition among trees regulates the responses of tree growth to N and P addition, and conducting long-term and coordinated distributed field experiments investigating the effects of N and P addition on tree growth and forest productivity at the global scale.     

Change of growth characters and carbon stocks in plantations of Pinus sylvestris var. mongolica in Saihanba,Hebei, China

Aims Pinus sylvestris var. mongolica is one of the main afforestation tree species in North China. It is important to study the characters of growth and carbon (C) sequestration, which can provide scientific basis for the sustainable management. Therefore, our study aims at quantifying the growth characters and C sequestration in these middle-aged plantations, and to investigate the effect of diameter at breast height (DBH) on those dynamics. Methods We selected a middle-aged P. sylvestris var. mongolica plantation as our permanent experimental plot, which is located in Saihanba, Hebei Province, China. DBH and height of all stands in this plot were measured in 2006 and 2016. Based on the anatomical trees and allometric equation, we calculated C density and sequestration from 2006 to 2016. We also analyzed C sequestration in different DBH groups in the study area. Important findings Our results showed that the carbon sink of those middle-age (age between 28 and 38 years old) plantation would be enhanced in future, and there were differences in characters of growth and C sequestration among DBH groups. The decadal increment rate of DBH and height were 4.19% and 1.97%, and the increment rate was the lowest in the 0-10 cm DBH class. The mortality rate of the plantation was 8.39%, with 7.82% mortality occurred in 0-10 cm tree size class. The forest stands biomass carbon stocks in 2006 and 2016 were 59.04 and 109.64 t?hm^-2, respectively, and almost 87.1% of the carbon stocks were in the middle DBH-class, even though the number of trees only accounted for nearly 59.2%. The small class’s number of trees accounted for 39.1%, while the carbon stocks accounted for 8.3%. Our results also demonstrate that forests in Saihanba would continue to act as a carbon sink in the coming years. The variations among DBH groups highlights that the diameter class should be taken into consideration while assess the ecological efficiency and carbon sequestration capacity in a certain area.     

秦岭东段栓皮栎枝条非结构性碳水化合物含量的季节动态

枝条是碳供应器官和碳需求器官的连接者, 研究其非结构性碳水化合物(NSC)含量的季节变化对理解树木体内的碳分配至关重要。该研究以秦岭东段栓皮栎(Quercus variabilis)优势群落为研究对象, 于2016年5月至2017年5月, 在其分布的海拔上下限(650 m和970 m), 通过在展叶期采用旬尺度和在非展叶期采用月尺度相结合的周期性取样方法(共计12次), 测定栓皮栎枝条NSC组分及含量, 并观测同期叶片物候变化。结果表明: (1)栓皮栎枝条NSC含量随季节波动较小, 变化差异不显著。但枝条可溶性糖含量(高海拔)或淀粉含量(低海拔)在一定生境条件下, 均存在明显的季节波动, 说明栓皮栎枝条可溶性糖和淀粉之间存在动态转化过程。(2)栓皮栎枝条NSC组成以可溶性糖为主(61%), 这可能是该树种在暖温带季风气候区所采取的生长策略。(3)土壤含水量(正相关)和饱和水汽压差(负相关)分别是在高海拔和低海拔影响栓皮栎枝条NSC含量的主导环境因子, 说明相比高海拔, 低海拔的栓皮栎可能对高温引起的水分胁迫更敏感。(4)结合叶片物候发现, 栓皮栎枝条NSC含量最大值出现在萌芽前(3月中下旬, 11%左右), 最小值出现在展叶后期(4月末, 5%左右), 叶片萌芽展叶后枝条NSC含量下降。总体而言, 枝条NSC含量在高低海拔不存在显著差异, 但春季萌芽前后存在显著差异, 海拔引起的叶片物候时间差极可能是造成这一现象的主要原因。研究结果说明, 栓皮栎叶片物候会直接影响枝条NSC含量的季节变化, 枝条NSC含量对叶片萌芽生长至关重要, 研究结果有助于加深对栓皮栎树体内碳调配机制的理解。