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。     

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。     

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.     

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。     

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.     

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.     

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.     

Dendroecological analysis of defoliator outbreaks on Nothofagus pumilio and their relation to climate variability in the Patagonian Andes

In the temperate forests of the southern Andes, Nothofagus pumilio, the dominant species of the most extensive forest type, experiences severe defoliation caused by caterpillars of the Ormiscodes genus (Lepidoptera: Saturniidae). This study uses tree rings to reconstruct the history of Ormiscodes outbreaks for the 1850–2005 period and examines relationships between outbreaks and climate variability. We used local climate records to compare outbreak–climate relationships in the northern Patagonian Andes (c. 41°S) and the cooler southern Patagonian Andes (c. 49°S). We also examined relationships between outbreak events and regional climate variability driven by variability in the Southern Annular Mode (SAM) and the El Niño‐Southern Oscillation. Although relationships between Ormiscodes outbreaks and climate proved to be complex, in northern Patagonia defoliation events are associated with drier and warmer than average growing seasons. Warming and drying trends in Patagonia during the latter part of the 20th century have been linked to a positive trend in SAM. During the post‐1976 period of accelerated warming in Patagonia, widespread defoliation outbreaks have occurred in both northern and southern Patagonia but the increase in frequency of events has been greater in the south. In southern Patagonia the increases in frequency of outbreaks in the late 20th century appear to be unprecedented over the c. 150 year tree‐ring record of reconstructed outbreaks. These results are consistent with the greater magnitude of recent warming in southern Patagonia, and suggest that under predicted warmer and drier climates in the 21st century, defoliator outbreaks may continue to increase in frequency. This study is the first systematic reconstruction of past insect outbreaks in South America and provides a preliminary understanding of how climate variability affects defoliator outbreaks in Patagonian Nothofagus forests.     

Tracing the influence of larch-bud-moth insect outbreaks and weather conditions on larch tree-ring growth in Engadine (Switzerland).

Tree-ring analysis of insect-defoliated trees has so far been used for detecting past insect outbreaks only. We hypothesize that the impact of the larch-bud-moth (LBM) Zeiraphera dinian outbreaks on the growth of larch Larix decidua in the Engadine (Switzerland) is closely coupled to the spatial development of the outbreak and the ecological characteristics of the respective sites and weather conditions. We tested this hypothesis by reviewing data sets available in the literature and by analysing original data. We monitored LBM population densities and the needle phenology, growth and defoliation of larch over 28 years, i.e. over four outbreak cycles. In addition, information on defoliation patterns covering six earlier outbreaks over 50 years was matched with tree-ring information. Tree-ring chronologies of 18 larch stands were analysed with regard to abrupt growth changes and latewood events. Defoliation induces an immediate reduction in latewood, followed by a reduction in needle length and a significant decrease in radial growth in the subsequent year. We have called this tree-ring pattern the "larch-bud-moth syndrome". A careful analysis of the various parameters of the LBM syndrome for two specific population cycles enabled us to define different interaction patterns between weather conditions and tree growth. These can then be included in climate change models to help disentangle the impact of insect defoliation from that of adverse climatic conditions.     

How drought events during the last century have impacted biomass carbon in Amazonian rainforests

During the last two decades, inventory data show that droughts have reduced biomass carbon sink of the Amazon forest by causing mortality to exceed growth. However, process-based models have struggled to include drought-induced responses of growth and mortality and have not been evaluated against plot data. A process-based model, ORCHIDEE-CAN-NHA, including forest demography with tree cohorts, plant hydraulic architecture and drought-induced tree mortality, was applied over Amazonia rainforests forced by gridded climate fields and rising CO₂ from 1901 to 2019. The model reproduced the decelerating signal of net carbon sink and drought sensitivity of aboveground biomass (AGB) growth and mortality observed at forest plots across selected Amazon intact forests for 2005 and 2010. We predicted a larger mortality rate and a more negative sensitivity of the net carbon sink during the 2015/16 El Niño compared with the former droughts. 2015/16 was indeed the most severe drought since 1901 regarding both AGB loss and area experiencing a severe carbon loss. We found that even if climate change did increase mortality, elevated CO₂ contributed to balance the biomass mortality, since CO₂-induced stomatal closure reduces transpiration, thus, offsets increased transpiration from CO₂-induced higher foliage area.     

长白山不同海拔岳桦非结构碳水化合物含量的变化

 通常认为, 随着林木不断接近其海拔分布极限, 光合作用产量不断下降, 导致碳水化合物供应不足(碳供应限制), 或者低温限制了碳投资(生长限制)。植物组织内非结构性碳水化合物(Nonstructural carbohydrates, NSC)的含量反映了植物碳供应与碳吸收的平衡。为了检验“碳供应限制”和“生长抑制”假说, 我们对长白山海拔1 700~ 2 050 m的自然生境下生长的岳桦(Betula ermanii)的叶片和枝条组织的NSC含量进行了比较。结果表明: 岳桦叶片的NSC含量随海拔升高变化不显著, 枝条的NSC含量随海拔升高显著增加; 叶片和枝条中淀粉含量与可溶性总糖含量的比值均随海拔的升高而减小; 林线附近的岳桦林木不存在碳水化合物供应不足的问题, 这在一定程度上表明生长限制导致长白山岳桦林线的形成。     

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.     

End of season carbon supply status of woody species near the treeline in western China

As trees and shrubs approach the high elevation tree limit, it is often assumed that they fall short in photosynthate (source limitation). Alternatively, low temperature may restrict carbon investment (growth, sink limitation). The content of mobile non-structural carbohydrates (NSC) in tissues is considered a measure of the carbon source–sink balance. To test the source vs. sink limitation hypothesis, we compared late-season NSC concentrations of various woody taxa across altitudinal gradients from the subalpine forest to the treeline at the eastern edge of the Tibetan Plateau. Since we were interested in the generality of trends, we present “community” trends across four taxa, namely Quercus aquifolioides, Abies faxoniana, Rhododendron fabri subsp. prattii and Sorbus rufopilosa. NSC concentrations increased significantly with altitude in branch wood, current-year and last-year leaves, while there were no significant trends in stem sapwood and root xylem. The sugar to starch ratio was roughly 1:1 in branches and evergreen leaves, while stems and roots showed a higher starch fraction. Analyses of total nitrogen in leaves and wood tissues indicated no change in the trees’ nitrogen supply with elevation. The overall altitudinal trends of NSC in this group of woody plant species revealed no depletion of carbon reserves near the tree limit, suggesting that sink limitation predominates woody plant life across this treeline ecotone community.     

Growth and mortality of trembling aspen (Populus tremuloides) in response to artificial defoliation

To simulate the effects of forest tent caterpillar (FTC) defoliation on trembling aspen growth and mortality, an artificial defoliation experiment was performed over three years in young aspen stands of northwestern Quebec. Defoliation plots of 15 × 15 m were established on three sites, together with associated control stands of pure trembling aspen. In 2007, root collar diameters were measured and positions of all trees were mapped prior defoliation. Severe FTC defoliation was simulated for three successive years (2007–2009) by manually removing all leaves from all but 7–10% of the trees present in the defoliation plots. Yearly surveys of growth and mortality were conducted until 2010 to evaluate defoliation effects on defoliated as well as surrounding undefoliated trees. In absence of other factors, growth and mortality of trembling aspen decreased and increased, respectively, after defoliation. Our study further revealed that small diameter trees died after one year of artificial defoliation, while larger-diameter trees died after repeated defoliations. Distributions of tree mortality tended to be aggregated at small scales (<5 m), corroborating gap patterns observed in mature stands following FTC outbreaks. This experiment revealed that trembling aspen mortality can be directly attributed solely to defoliation. Repeated defoliations during FTC outbreaks have the potential to profoundly modify stand productivity and structure by reducing tree growth and increasing tree mortality in the absence of predisposing factors.     

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,比较了雌雄青杨幼苗的生长、NSC含量和储量对去叶(0,50%和100%叶片去除)的响应差异。结果表明:随去叶强度的增加,植物的生物量和植株NSC呈降低趋势,且根系(尤其是粗根)的生物量和NSC比地上部分受去叶的影响更大;雌株叶、粗根、细根和植株NSC储量总是高于雄株;随去叶强度增加,雄株的生物量积累和NSC含量和储量降低得比雌株更多。这些结果表明,青杨雌雄植株生长和NSC对不同去叶强度的响应存在性别差异,且去叶对青杨雄株的影响更大。这暗示了雌性青杨对去叶的耐性比雄性强。这些结果有助于理解雌雄异株植株性别水平上的碳平衡机理,也可为杨树人工林的选育提供支撑。