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。     

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。     

植物对气候变化生理生态响应的不确定性分析

生态系统对全球气候变化的响应模式有利于人类预测与适应未来生态环境变化,植物作为陆地生态系统的重要组成部分,对全球气候变化的响应具有重要作用.本文通过对近年来植物对气候变化的生理生态响应研究中(包括定点控制实验,空间代替时间样带),植物响应模式的复杂性、多样性及可变性等诸多不确定性进行分析.以探讨预测未来气候情景下植物的动态变化及生理生态响应过程.分析结果认为.造成这些不确定性的主要原因包括:(1)利用空间代替时间的样带研究中,往往忽略了植物的非线性响应,存在明显的阈值;(2)样带及定点研究中,由于各种气候因子的耦合.很难确定各种气候因子对植物生理生态学特性影响的权重;(3)定点控制实验中往往忽略了植物对气候变化的适应性,使实验结果很难代表更长时间尺度上的反映模式;(4)在相同的气候变化条件下,不同植物的响应有可能存在明显差异.提出了今后植物对气候变化生理生态响应研究的建议.     

中国东北典型森林生态系统植物叶片的非结构性碳水化合物研究

植物叶片的非结构性碳水化合物(NSC)不仅可以反应植物的碳供应状况,也能反应植物对外界环境的适应策略。利用传统的蒽酮比色法测定了东北3个典型森林生态系统(呼中、凉水和长白山)242种常见植物叶片的非结构碳水化合物,探讨了温带主要森林植物叶片NSC沿纬度梯度的变化趋势及其在物种-生活型-群落间的分布规律。实验结果表明:3个典型森林生态系统植物叶片可溶性糖、淀粉和NSC含量均呈偏正态分布,多数物种的含量偏中低水平;242种植物叶片可溶性糖、淀粉和NSC的平均含量分别为63.31、65.66和128.96 mg/g。在所调查的森林生态系统中,叶片可溶性糖、淀粉和NSC含量在不同生活型中表现各异。此外,乔木植物叶片的可溶性糖、淀粉和NSC含量从北到南呈递增趋势,呼中最低,凉水次之,长白山最高。乔木淀粉含量均表现为落叶树种大于常绿树种,可溶性糖和NSC含量变化趋势复杂。研究结果不仅为阐明东北主要森林生态系统植被碳代谢和生长适应对策提供数据基础,而且对理解植物对未来气候变化的响应机理提供数据支撑。     

植物叶片性状对气候变化的响应研究进展

叶片性状反映了植物对环境的高度适应能力及其在复杂生境下的自我调控能力。叶片性状如何响应和适应气候变化是植物适应性研究的重点内容。该文系统综述了叶片大小、比叶质量、叶片氮含量、碳同位素等指标对气候变化响应的最新研究结果。不同叶片性状对气候变化的响应结果存在差异,所指示的生态学含义也有所不同。单一叶片性状不能全面地反映植物对气候变化的响应;不同尺度的研究(如环境的修饰或筛选作用的研究)还存在很多不确定性。高寒地区的研究工作相对缺乏。该文有助于理解植物与气候之间的相互关系、植物对气候变化的响应与适应对策,对了解植物演化、预测植物在未来气候变化条件下的变化特征具有一定意义。     

气候变化对雌雄异株植物影响的研究进展

雌雄异株植物作为陆地生态系统的重要组成部分,在维持生态系统结构和功能稳定性方面发挥着关键作用。但是,由于雌雄异株植物的不同性别植株在形态特征、生理特征和资源分配等方面均存在明显差异,至今对全球气候变化背景下雌雄植物的性别差异响应机制的认识仍十分有限。本文综述了全球变暖、降水变化和大气CO2浓度升高等主要气候因子变化对雌雄异株植物的影响,探讨了雌雄异株植物对气候变化响应的性别差异,提出了未来雌雄异株植物与气候变化关系研究的重点,以期为揭示陆地生态系统结构和功能稳定性对气候变化的响应机制提供参考依据。     

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。     

阿尔泰山西伯利亚落叶松径向生长对气候变化的分异响应

在全球气候变暖的背景下, 北半球中高纬度地区出现了树轮径向生长对气候变化的分异响应现象, 但是阿尔泰山优势针叶树种对气候因子响应的稳定性还存在不确定性。该研究选择阿尔泰山中段高海拔西伯利亚落叶松(Larix sibirica)样本建立了树轮宽度年表, 并对年表特征及树木径向生长-气候的动态关系进行了分析。结果表明: 生长季初期和中期的气温是研究区树木生长的主控气候因子; 树木径向生长与当年4月的气温显著负相关, 与当年6-7月的气温显著正相关; 研究区西伯利亚落叶松径向生长与当年4月和6-7月的气温发生了分异现象, 表现为随着气候变化, 树木径向生长对生长季初期由高温引起的干旱的响应敏感性越来越强, 而对生长季中期气温的敏感性表现出先减弱再增强的趋势。阿尔泰山西伯利亚落叶松径向生长对气候变化的响应比较敏感, 适合开展树木生长-气候变化的研究; 检验树木径向生长对气候变化分异响应为该区域基于树木年轮开展历史气候重建和提高未来森林生态系统发展趋势预测的准确性提供了科学依据。     

植物物候研究进展

植物物候直接反映了气候变化的影响,是植被动态模拟的关键.在遥感和模型技术的推动下,植物物候与全球变化关系的研究日益受到人们的关注.文中从植物物候与环境因子的相互关系、植物物候对全球变化的响应以及植物物候的遥感监测方面,综合论述了植物物候的研究进展,找出植被物候研究的不足,进而提出未来植被物候的研究方向.     

阿尔泰山西伯利亚落叶松径向生长对气候变化的分异响应

在全球气候变暖的背景下,北半球中高纬度地区出现了树轮径向生长对气候变化的分异响应现象,但是阿尔泰山优势针叶树种对气候因子响应的稳定性还存在不确定性。该研究选择阿尔泰山中段高海拔西伯利亚落叶松(Larixsibirica)样本建立了树轮宽度年表,并对年表特征及树木径向生长–气候的动态关系进行了分析。结果表明:生长季初期和中期的气温是研究区树木生长的主控气候因子;树木径向生长与当年4月的气温显著负相关,与当年6–7月的气温显著正相关;研究区西伯利亚落叶松径向生长与当年4月和6–7月的气温发生了分异现象,表现为随着气候变化,树木径向生长对生长季初期由高温引起的干旱的响应敏感性越来越强,而对生长季中期气温的敏感性表现出先减弱再增强的趋势。阿尔泰山西伯利亚落叶松径向生长对气候变化的响应比较敏感,适合开展树木生长–气候变化的研究;检验树木径向生长对气候变化分异响应为该区域基于树木年轮开展历史气候重建和提高未来森林生态系统发展趋势预测的准确性提供了科学依据。     

Large seasonal fluctuations in whole-tree carbohydrate reserves: is storage more dynamic in boreal ecosystems?

Background and AimsCarbon reserves are a critical source of energy and substrates that allow trees to cope with periods of minimal carbon gain and/or high carbon demands, conditions which are prevalent in high-latitude forests. However, we have a poor understanding of carbon reserve dynamics at the whole-tree level in mature boreal trees. We therefore sought to quantify the seasonal changes in whole-tree and organ-level carbon reserve pools in mature boreal Betula papyrifera.MethodsNon-structural carbohydrate (NSC; soluble sugars and starch) tissue concentrations were measured at key phenological stages throughout a calendar year in the roots, stem (inner bark and xylem), branches and leaves, and scaled up to estimate changes in organ and whole-tree NSC pool sizes. Fine root and stem growth were also measured to compare the timing of growth processes with changes in NSC pools.Key ResultsThe whole-tree NSC pool increased from its spring minimum to its maximum at bud set, producing an average seasonal fluctuation of 0.96 kg per tree. This fluctuation represents a 72 % change in the whole-tree NSC pool, which greatly exceeds the relative change reported for more temperate conspecifics. At the organ level, branches accounted for roughly 48–60 % of the whole-tree NSC pool throughout the year, and their seasonal fluctuation was four to eight times greater than that observed in the stemwood, coarse roots and inner bark.ConclusionsBranches in boreal B. papyrifera were the largest and most dynamic storage pool, suggesting that storage changes at the branch level largely drive whole-tree storage dynamics in these trees. The greater whole-tree seasonal NSC fluctuation in boreal vs. temperate B. papyrifera may result from (1) higher soluble sugar concentration requirements in branches for frost protection, and/or (2) a larger reliance on reserves to fuel new leaf and shoot growth in the spring.     

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.     

Unfolding Non-structural Carbohydrates from Sapling to Dying Black Locust on China’s Loess Plateau

Non-structural carbohydrates (NSC) play important roles in metabolic processes of plants and represent important functional traits in plant adaptation to the external environment. However, there are few reports concerning intra-annual NSC distribution in temperate deciduous forests, especially for the purpose of comparison among different stand ages on China’s Loess Plateau. Here, NSC allocation dynamics with age ring-porous black locust tree was determined for the growing (May) and dormant (November) seasons over the period from sapling to dying trees—defined as a completely defoliated tree with dried branches in the growing season. It was noted that regardless of tree age, NSC concentration was highest in coarse roots [16.4 g per 100 g of dry mass (16.4% DM)] and stems (15.1% DM). At the tree level, NSC concentration was highest (14.3% DM) in a 30-year-old stand in November and lowest (4.1% DM) in dying stands in May. The pool of NSC at tree level was highest (25.2 kg DM per tree) in 30-year-old stands in November and lowest (0.13 kg DM per tree) in sapling stands in May. The concentration of NSC was significantly lower in May than in November for all tree ages, organs, and biochemical components. The results underscored the importance of NSC in plant growth on China’s Loess Plateau. It also provided a useful insight into the dynamics of NSC from sapling to dying broadleaved tree species.     

Can fungal endophytes fast-track plant adaptations to climate change?

Rapid climate change threatens plant communities. While many studies address the impact of climate change on plants and mechanisms of their resilience to climate stressors, the role of the plant microbiome in aiding plants' adaptation to climate change has been less investigated. We argue here that fungal endophytes, an important constituent of the plant microbiome, may be key to the ability of plants to adapt to climatic stressors. The rapid adaptive response of endophytes coupled with their ability to ‘transfer’ resistance to their hosts may fast-track plants' adaptation to climate change. We briefly review the importance of Class 3 fungal endophytes of terrestrial plants and discuss how they may accelerate adaptations to climate change in crops and natural plant communities and call for efforts directed at improving the understanding of fungal endophyte-facilitated plant health. Such information could aid in devising improved strategies for mitigating climate change effects on plant communities.     

Tree-rings,genetics and the environment: Complex interactions at the rear edge of species distribution range

Under climate change, modifications on plants’ growth are expected to be the strongest at species margins. Therein, tree acclimation could play a key role as migration is predicted to be too slow to track shifts of bioclimatic envelops. A requirement is, however, that intra-population genetic diversity be high enough for allowing such adaptation of tree populations to climate change. In this study, we tested for the existence of relationships between genetic diversity, site environmental conditions, and the response of annual tree growth to climate of Pinus cembra at its southern limit in the Alps. Site-specific climatic and environmental factors predominantly determined the response of trees along the precipitation gradient. The growth-climate interactions were chiefly linked to mean annual precipitation and temperature, slope and tree-size, and less to genetic diversity. We show that genetic background of Pinus cembra has exclusively indirect modulating power with limited effects on tree-ring formation, and within the southern limit in the Alps, genetic variability is not necessarily well expressed in the patterns of annual tree growth. Our results may imply little adaptive capacity of these populations to future changes in the water balance.     

兴安落叶松针叶解剖结构变化及其光合能力对气候变化的适应性

叶片易受环境因子影响,其形态解剖结构特征不但与叶片的生理功能密切相关,而且反映树木对环境变化的响应和适应。叶片结构的改变势必会改变树木的生理功能。同一树种长期生长在异质环境条件下,经过自然选择和适应,会在形态和生理特性等方面产生变异,形成特定的地理种群。另外,母体所经受的环境胁迫也会影响到其子代的生长、发育和生理等特征。因此,了解植物叶片形态结构对环境变化的响应与适应是探索植物对环境变化的响应适应机制的基础。兴安落叶松(Larix gmelinii Rupr.)是我国北方森林的优势树种,主要分布在我国东北地区,但日益加剧的气候变化可能会改变其现有的分布区。为了区分叶片对气候变化的可塑性和适应性,本研究采用同质园法比较测定了6个不同气候条件下的兴安落叶松种源的32年生树木的针叶解剖结构和光合生理相关因子,利用石蜡切片方法分析了针叶的解剖结构特征、光合能力（Pmax-a）、水分利用效率（WUE）之间的关系及其对气候变化的适应性。结果表明：表皮细胞厚度、叶肉细胞厚度、传输组织厚度、维管束厚度、内皮层厚度以及叶片总厚度均存在显著的种源间差异（P < 0.05）。叶肉细胞厚度与Pmax-a、气孔导度和WUE之间均存在显著的正相关关系（P < 0.05）。叶肉细胞厚度、表皮细胞厚度、叶片总厚度以及叶肉细胞厚度和表皮细胞厚度在叶片总厚度中所占比例均与种源地的干燥度指数（即年蒸发量与年降水量之比）呈正线性关系。这些结果说明：不同种源兴安落叶松针叶解剖结构因对种源原地气候条件的长期适应而产生显著的差异,从而引起其针叶光合作用、水分利用等生理功能发生相应的变化,从而有利于该树种在气候变化的情景下得以生存和繁衍。