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CO₂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。     

干旱导致树木死亡机制研究进展

随着全球气候变化,气温上升,干旱发生的可能性增加,干旱导致树木死亡现象的发生也越来越普遍,但干旱导致树木死亡确切的生理学机制还未完全解决,目前仍是研究的热点。近来提出的干旱导致树木死亡的假说主要有:水力学失败假说,认为干旱导致的低水势阻碍植物体内水分的长距离运输,使植物因组织脱水干燥而死亡;碳饥饿假说,认为干旱过程中发生的气孔关闭会限制碳摄取,导致植物体内可利用碳的耗竭;生物攻击假说,认为干旱中的植物易受生物因子(昆虫和病原体)的攻击而死亡。3种假说各自都能解释一定的现象,但又都存在缺陷,主要问题是不能解释所有干旱导致树木死亡的现象。近来的研究越来越关注这些机制之间的共同作用或相互作用,水力学失败、碳饥饿和生物因子之间会相互影响而加重干旱对植物的作用,尤其是水力学失败和碳饥饿相耦合已得到越来越多的认可。本文主要综述这几种致死机制的试验证据及存在的问题,同时综述当前研究存在的局限性,并讨论了未来可能的研究方向。     

摘叶造成的碳限制对刺槐碳素分配和水力学特性的影响

以3年生刺槐（Robinia pseudoacacia Linn.）为研究对象,通过对其进行连续3次摘叶造成严重碳限制,检测摘叶后刺槐的生物量分配、叶片形态和不同部位的非结构性碳（NSC）浓度,同时检测其根压和根系导水率、枝条水势和导水率损失值（PLC）及茎的抗栓塞能力,研究摘叶造成的碳限制对刺槐碳素分配和水力学特性的影响。结果显示,摘叶显著降低了刺槐不同部位的生物量,其中细根生物量降低程度最大;摘叶还造成了刺槐不同部位NSC浓度显著降低,茎韧皮部、茎木质部、根韧皮部和根木质部的NSC浓度分别为对照的29.6%、20.2%、10.2%和8.7%,且根部NSC的降低程度显著高于茎;碳限制显著降低了刺槐苗木的根压和根系导水率,增加了枝条凌晨和正午的PLC,降低了其抗栓塞能力。研究结果表明摘叶造成的碳限制改变了刺槐的碳素分配模式,限制了碳素向根的分配,抑制细根的发生,进而限制根的水分吸收能力,加重枝条栓塞程度,同时还会导致枝条抗栓塞能力下降,从而降低植物水分输导的安全性。     

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。     

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.     

造林再造林、森林采伐、气候变化、CO2浓度升高、火灾和虫害对森林固碳能力的影响

森林生态系统具有吸收大气CO_2、缓解气候变化的作用。造林再造林作为京都议定书认可的大气CO_2减排途径,是提高森林固碳能力的低成本、有效策略。森林生态系统固碳能力还受森林采伐、气候变化、大气CO_2浓度升高、火灾以及虫害等自然因素和人为因素的强烈影响。综述了全球和区域造林再造林的固碳能力,以及目前较受重视的一些因素(森林采伐、气候变化、大气CO_2浓度升高、火灾以及虫害)对森林生态系统固碳能力的影响。结果表明,全球造林再造林固碳能力为148—2400TgC/a;采伐造成的全球森林碳损失最大为900 TgC/a,其次是火灾为300 TgC/a,虫害造成森林碳释放最小在2—107 TgC/a之间。建议在今后的研究中,应关注固碳措施和多种环境因素对森林生态系统固碳能力,尤其是对森林土壤固碳能力的影响,严格控制森林采伐和火灾发生,以及减少或避免造林再造林活动引起的碳泄漏。     

植物干旱胁迫下水分代谢、碳饥饿与死亡机理

植物在生长发育过程中受众多环境因子共同作用。随着全球气候变化,气温升高、降水量下降等问题频繁出现。目前气象学家一致预测未来环境变暖会使干旱更加频繁剧烈,这一环境改变使植物死亡更加严重。植物在水分胁迫、特别是干旱胁迫条件下,体内水分代谢与碳代谢会发生失衡现象:光合速率降低、蒸腾速率降低,带来生长降低;为维持植物新陈代谢,植物呼吸作用必然下调。在长期干旱胁迫条件下植物体内碳水化合物储存发生失衡现象,这种失衡使植物陷入碳饥饿现象。另外,由于水分失衡而出现的木质部栓塞和空穴会进一步加剧水分运输障碍,而修复空穴则需要大量非结构性碳水化合物(NSC),这使植物陷入两难选择。总结了植物干旱胁迫下,碳饥饿与水分代谢、植物死亡关系的相关研究,对未来的研究方向和重点提出建议,以期对未来的植物死亡研究提供帮助。     

林火干扰下的树木生理及主要影响因素

随着全球逐渐暖干化,林火不仅驱动着森林生态系统结构和功能的变化,同时也影响树木的生理和生长。林火导致的热损伤引发树木一系列复杂生理响应。揭示火后树木生理的响应机制,对于进一步理解树木碳水关系和火后恢复生长限制,以及提高火后树木死亡预测准确性具有重要指导意义。该文从林火对树木的作用途径和方式着手,基于不同形式林火(树冠火、地表火、地下火)对树木各部分(树冠、树干、根系)造成的损伤,综述了林火对树木生理的直接影响和间接影响,以及火后树木生理与非生物和生物因素的互作关系。热损伤诱导的形成层、韧皮部坏死和木质部水力失衡是火后树木生理的主要响应机制,二者导致的两个生理功能限制——“碳饥饿”和水力失效——严重影响树木的碳水关系,也决定了火后树木是恢复生长还是延迟死亡。火后树木生理机制还与干旱、昆虫攻击和微生物入侵等其他因素密切相关。该文强调了对林火强度的定量分析和对植物组织死亡阈值的准确判断的迫切性,同时提出了探究火后树木生理与功能性状和其他因素的互作关系的必要性。精确评估树木生理机制间关系对于深入理解林火如何影响树木功能完整性极为关键,有助于完善林火风险评估和树木死亡模型预测。在未来气候暖干化驱...     

去叶对水曲柳苗木根系非结构性碳水化合物分配的影响

全球气候变化有促进食叶害虫爆发的趋势.叶片被食会导致光合产物的生产降低,进一步影响非结构性碳水化合物(NSC)在树木体内的储存和分配.本研究以水曲柳2年生苗木根系为研究对象,通过40%去叶处理,于6-10月对根系取样,研究地上碳(C)供应不足条件下主根、粗根和1～5级细根NSC及其组分的分配格局.结果表明: 对照和去叶处理根系NSC浓度及其组分浓度均具有明显的季节动态;与对照相比,去叶处理苗木主根和粗根NSC浓度分别降低3.8%和30.7%,而1～5级细根NSC浓度增加1.2%～23.5%,这主要受淀粉浓度变化的影响;去叶处理苗木主根和粗根可溶性糖浓度增加7.1%和62.3%,而1～5级根可溶性糖浓度显著降低2.7%～42.8%;去叶对苗木根系可溶性糖和淀粉浓度的不同影响,导致二者的比值在主根和粗根中增加,而在1～5级细根中降低.去叶引起光合产物的生产减少,导致水曲柳苗木主根和粗根中淀粉活化后流向细根并以淀粉的形式储存,这可能有利于提高细根对冬季低温胁迫的抵抗力.     

去叶对青杨雌雄植株生长和非结构性碳水化合物的影响

近年来,森林食叶害虫在全世界呈爆发趋势.树木的非结构性碳水化合物(NSC)如何响应叶片损失对其生长和生存至关重要.雌雄异株植物在维持森林生态系统稳定性方面扮演着重要角色.然而,目前对该类植物性别之间如何响应去叶的研究还比较少.本文以我国重要的经济和生态恢复树种青杨(Populus cathayana)为研究材料,比较了...     

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.     

Outbreaks of forest defoliating insects in Japan, 1950-2000

In Japan, several forest-defoliating insects reach outbreak levels and cause serious defoliation. Stand mortality sometimes occurs after severe defoliation. However, in general, tree mortality caused by insect defoliation is low because of the prevailing moist climate in Japan. Evergreen conifers are more susceptible to tree mortality as a result of insect defoliation whereas deciduous broad-leaved trees are seldom killed. Insect defoliation occurs more frequently in man-made environments such as among shade trees, orchards, and plantations than in natural habitats. Outbreaks of some defoliators tend to occur in stands of a particular age: e.g. outbreaks of the pine caterpillar, Dendrolimus spectabilis Butler (Lepidoptera: Lasiocampidae) occur more frequently in young pine plantations. In contrast, defoliation caused by outbreaks of lepidopterous and hymenopterous pests in larch plantations is more frequent with stand maturation. There is a relationship between outbreaks of some defoliators and altitude above sea level. Most outbreaks of forest defoliators were terminated by insect pathogens that operated in a density-dependent fashion. Since the 1970s, Japan has been prosperous and can afford to buy timber from abroad. More recently, there has been an increasing demand for timber in Japan, that coincides with a huge demand internationally, so that the country will need to produce more timber locally in the future. The increasing pressure on the forestry industry to meet this demand will require more sophisticated methods of pest control coupled with more sustainable methods of silviculture.     

Interactions Among Fuel Management,Species Composition,Bark Beetles,and Climate Change and the Potential Effects on Forests of the Lake Tahoe Basin

Climate-driven increases in wildfires, drought conditions, and insect outbreaks are critical threats to forest carbon stores. In particular, bark beetles are important disturbance agents although their long-term interactions with future climate change are poorly understood. Droughts and the associated moisture deficit contribute to the onset of bark beetle outbreaks although outbreak extent and severity is dependent upon the density of host trees, wildfire, and forest management. Our objective was to estimate the effects of climate change and bark beetle outbreaks on ecosystem carbon dynamics over the next century in a western US forest. Specifically, we hypothesized that (a) bark beetle outbreaks under climate change would reduce net ecosystem carbon balance (NECB) and increase uncertainty and (b) these effects could be ameliorated by fuels management. We also examined the specific tree species dynamics—competition and release—that determined NECB response to bark beetle outbreaks. Our study area was the Lake Tahoe Basin (LTB), CA and NV, USA, an area of diverse forest types encompassing steep elevation and climatic gradients and representative of mixed-conifer forests throughout the western United States. We simulated climate change, bark beetles, wildfire, and fuels management using a landscape-scale stochastic model of disturbance and succession. We simulated the period 2010–2100 using downscaled climate projections. Recurring droughts generated conditions conducive to large-scale outbreaks; the resulting large and sustained outbreaks significantly increased the probability of LTB forests becoming C sources over decadal time scales, with slower-than-anticipated landscape-scale recovery. Tree species composition was substantially altered with a reduction in functional redundancy and productivity. Results indicate heightened uncertainty due to the synergistic influences of climate change and interacting disturbances. Our results further indicate that current fuel management practices will not be effective at reducing landscape-scale outbreak mortality. Our results provide critical insights into the interaction of drivers (bark beetles, wildfire, fuel management) that increase the risk of C loss and shifting community composition if bark beetle outbreaks become more frequent.     

Declining carbohydrate content of Sitka-spruce treesdying from seawater exposure

Increasing sea levels associated with climate change threaten the survival of coastal forests, yet the mechanisms by which seawater exposure causes tree death remain poorly understood. Despite the potentially crucial role of nonstructural carbohydrate (NSC) reserves in tree survival, their dynamics in the process of death under seawater exposure are unknown. Here we monitored progressive tree mortality and associated NSC storage in Sitka-spruce (Picea sitchensis) trees dying under ecosystem-scale increases in seawater exposure in western Washington, USA. All trees exposed to seawater, because of monthly tidal intrusion, experienced declining crown foliage during the sampling period, and individuals with a lower percentage of live foliated crown (PLFC) died faster. Tree PLFC was strongly correlated with subsurface salinity and needle ion contents. Total NSC concentrations in trees declined remarkably with crown decline, and reached extremely low levels at tree death (2.4% and 1.6% in leaves and branches, respectively, and 0.4% in stems and roots). Starch in all tissues was almost completely consumed, while sugars remained at a homeostatic level in foliage. The decreasing NSC with closer proximity to death and near zero starch at death are evidences that carbon starvation occurred during Sitka-spruce mortality during seawater exposure. Our results highlight the importance of carbon storage as an indicator of tree mortality risks under seawater exposure.

Decline in carbohydrate storage is strongly associated with progressive mortality of trees under frequent seawater exposure, and starch is almost completely consumed at crown death.     

Extreme defoliation reduces tree growth but not C and N storage in a winter-deciduous species

Background and Aims There is a growing concern about how forests will respond to increased herbivory associated with climate change. Carbon (C) and nitrogen (N) limitation are hypothesized to cause decreasing growth after defoliation, and eventually mortality. This study examines the effects of a natural and massive defoliation by an insect on mature trees’ C and N storage, which have rarely been studied together, particularly in winter-deciduous species.Methods Survival, growth rate, carbon [C, as non-structural carbohydrate (NSC) concentration] and nitrogen (N) storage, defences (tannins and total polyphenols), and re-foliation traits were examined in naturally defoliated and non-defoliated adult trees of the winter-deciduous temperate species Nothofagus pumilio 1 and 2 years after a massive and complete defoliation caused by the caterpillar of Ormiscodes amphimone (Saturniidae) during summer 2009 in Patagonia.Key Results Defoliated trees did not die but grew significantly less than non-defoliated trees for at least 2 years after defoliation. One year after defoliation, defoliated trees had similar NSC and N concentrations in woody tissues, higher polyphenol concentrations and lower re-foliation than non-defoliated trees. In the second year, however, NSC concentrations in branches were significantly higher in defoliated trees while differences in polyphenols and re-foliation disappeared and decreased, respectively.Conclusions The significant reduction in growth following defoliation was not caused by insufficient C or N availability, as frequently assumed; instead, it was probably due to growth limitations due to factors other than C or N, or to preventative C allocation to storage. This study shows an integrative approach to evaluating plant growth limitations in response to disturbance, by examining major resources other than C (e.g. N), and other C sinks besides storage and growth (e.g. defences and re-foliation).     

Insect‐induced tree mortality of boreal forests in eastern Canada under a changing climate

Forest insects are major disturbances that induce tree mortality in eastern coniferous (or fir-spruce) forests in eastern North America. The spruce budworm (SBW) (Choristoneura fumiferana [Clemens]) is the most devastating insect causing tree mortality. However, the relative importance of insect-caused mortality versus tree mortality caused by other agents and how this relationship will change with climate change is not known. Based on permanent sample plots across eastern Canada, we combined a logistic model with a negative model to estimate tree mortality. The results showed that tree mortality increased mainly due to forest insects. The mean difference in annual tree mortality between plots disturbed by insects and those without insect disturbance was 0.0680 per year (P < 0.0001, T-test), and the carbon sink loss was about 2.87t C ha⁻¹ year⁻¹ larger than in natural forests. We also found that annual tree mortality increased significantly with the annual climate moisture index (CMI) and decreased significantly with annual minimum temperature (T_min), annual mean temperature (T_mean) and the number of degree days below 0°C (DD0), which was inconsistent with previous studies (Adams et al. 2009; van Mantgem et al. 2009; Allen et al. 2010). Furthermore, the results for the trends in the magnitude of forest insect outbreaks were consistent with those of climate factors for annual tree mortality. Our results demonstrate that forest insects are the dominant cause of the tree mortality in eastern Canada but that tree mortality induced by insect outbreaks will decrease in eastern Canada under warming climate.     

Variation in carbohydrate source–sink relations of forest and treeline white spruce in southern, interior and northern Alaska

Two opposing hypotheses have been presented to explain reduced tree growth at the treeline, compared with growth in lower elevation or lower latitude forests: the carbon source and sink limitation hypotheses. The former states that treeline trees have an unfavorable carbon balance and cannot support growth of the magnitude observed at lower elevations or latitudes, while the latter argues that treeline trees have an adequate carbon supply, but that cold temperatures directly limit growth. In this study, we examined the relative importance of source and sink limitation in forest and treeline white spruce (Picea glauca) in three mountain ranges from southern to northern Alaska. We related seasonal changes in needle nonstructural carbohydrate (NSC) content with branch extension growth, an approach we argue is more powerful than using needle NSC concentration. Branch extension growth in the southernmost Chugach Mountains was much greater than in the White Mountains and the Brooks Range. Trees in the Chugach Mountains showed a greater seasonal decline in needle NSC content than trees in the other mountain ranges, and the seasonal change in NSC was correlated with site-level branch growth across mountain ranges. There was no evidence of a consistent difference in branch growth between the forest and treeline sites, which differ in elevation by approximately 100 m. Our results point to a continuum between source and sink limitation of growth, with high-elevation trees in northern and interior Alaska showing greater evidence of sink limitation, and those in southern Alaska showing greater potential for source limitation.     

Infestation and hydraulic consequences of induced carbon starvation

Drought impacts on forests, including widespread die-off, are likely to increase with future climate change, although the physiological responses of trees to lethal drought are poorly understood. In particular, in situ examinations of carbon starvation and its interactions with and effects on infestation and hydraulic vulnerability are largely lacking. In this study, we conducted a controlled, in situ, repeated defoliation experiment to induce carbon stress in isolated trembling aspen (Populus tremuloides) ramets. We monitored leaf morphology, leaves per branch, and multitissue carbohydrate concentrations during canopy defoliation. We examined the subsequent effects of defoliation and defoliation-induced carbon stress on vulnerability to insect/fungus infestation and hydraulic vulnerability the following year. Defoliated ramets flushed multiple canopies, which coincided with moderate drawdown of nonstructural carbohydrate reserves. Infestation frequency greatly increased and hydraulic conductivity decreased 1 year after defoliation. Despite incomplete carbohydrate drawdown from defoliation and relatively rapid carbohydrate recovery, suggesting considerable carbohydrate reserves in aspen, defoliation-induced carbon stress held significant consequences for vulnerability to mortality agents and hydraulic performance. Our results indicate that multiyear consequences of drought via feedbacks are likely important for understanding forests' responses to drought and climate change over the coming decades.     

Determinants of mountain birch growth in situ: effects of temperature and herbivory

Variation in in situ growth performance of the mountain birch as indicated by the widths of annual rings was analysed and related mainly to temperature and herbivory using ring width series from five heath forest sites in the Lake Torneträsk area, northern Sweden. Climate explained 48–64% of the variation in age-corrected mean ring width series. In general, the effect of current year July followed by June temperature was most important at all sites. A warm May resulted in wider rings due to an earlier budburst. Short-term (inter-annual) responses to increased temperature were in most cases not reflected into long-term responses (decades). A large proportion of the variation in stem mean ring width was due to variation among stems within trees (81%) in these polycormic trees, while variation among sites was marginal (0.4%). Within trees, main stems grew faster and were more responsive to climate variation than subordinate stems. No effect of insect herbivory on ring width was found at low defoliation levels (≤12%). At a defoliation level of ca 84% a one-year reduction in stem growth was observed while the growth reduction (ca 50% reduction in ring width) lasted for 4 yr after ca 93% defoliation. After outbreaks resulting in complete defoliation and some stem mortality, ring widths of surviving stems mainly responded with increased growth. Basal sprouts, emerging just after a severe insect outbreak with a high mortality of old stems, grew faster than sprouts occurring during other periods. It is concluded that the mountain birch is well adapted to recover from Epirrita outbreaks; the ability to produce basal sprouts, that can benefit from an existing root system for fast initial growth, is one important mechanism for this.