中国4种典型森林中常见乔木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(non-structural carbohydrates,NSC)不仅为植物的代谢过程提供重要能量,还能一定程度上反映植物对外界环境的适应策略。以温带针阔混交林(长白山)、温带阔叶林(东灵山)、亚热带常绿阔叶林(神农架)和热带雨林(尖峰岭)4种森林类型的树种为研究对象,利用蒽酮比色法测定了163种常见乔木叶片可溶性糖、淀粉和NSC(可溶性糖+淀粉)含量,探讨了不同森林类型植物叶片NSC的差异及其地带性变化规律。结果显示：（1）从森林类型上看,植物叶片NSC含量从北到南递减,即温带针阔混交林(170.79 mg/g)>温带阔叶林(100.27 mg/g)>亚热带常绿阔叶林(91.24 mg/g)>热带雨林(80.13 mg/g)。（2）从生活型上看,无论是落叶树还是阔叶树,其叶片可溶性糖、淀粉和NSC含量均表现为：温带针阔混交林>温带阔叶林>亚热带常绿阔叶林>热带雨林;北方森林叶片可溶性糖、淀粉和NSC含量均表现为落叶树种>常绿树种,或阔叶树种>针叶树种。（3）森林植物叶片NSC含量、可溶性糖与淀粉含量比值与年均温和年均降水量均呈显著负相关。研究表明,森林植物叶片可溶性糖、淀粉和NSC含量以及可溶性糖与淀粉含量比值均具有明显的从北到南递减的地带性规律;其NSC含量以及可溶性糖与淀粉含量比值与温度和水分均呈显著负相关的变化规律可能是植物对外界环境适应的重要机制之一。该研究结果不仅为阐明中国主要森林树种碳代谢和生长适应对策提供了数据基础,而且为理解区域尺度森林植被对未来气候变化的响应机理提供新的视角。     

人工油松林中不同植物叶片非结构性碳水化合物含量对氮添加的响应

全球氮沉降通过影响树木叶片的生理过程影响森林生态系统的结构与组成,但目前关于氮沉降对森林生态系统中不同植物叶片非结构性碳水化合物(NSC)含量的影响还不十分清楚.本研究选取黄土丘陵区人工油松林中油松、辽东栎、忍冬、绣线菊、黄刺玫、茜草、披针苔草7种主要高等植物,比较了3年氮添加(0、3、6、9 g N·m^-2·a^-1,分别用N₀、N₃、N₆、N₉表示)对7种植物叶片中NSC的影响.结果表明: 不同植物叶片可溶性糖、淀粉含量变异很大,二者变异最高的均为黄刺玫,最低的分别为绣线菊和披针苔草;不同植物可溶性糖、淀粉对氮添加的响应存在明显差异.N₆处理下茜草的可溶性糖和淀粉的变异高于其他物种,N₃和N₉处理下绣线菊的变异高于其他物种,可溶性糖和淀粉含量变异最小的物种在不同氮添加水平下不同.随着氮添加水平的增加,油松和披针苔草的可溶性糖含量持续升高,绣线菊持续降低,辽东栎、忍冬和黄刺玫的可溶性糖含量先降低后增加,在N₆处理达到最小,而茜草的可溶性糖含量呈现较复杂的变化趋势.氮添加对植物叶片中淀粉含量的影响表现为油松、忍冬和披针苔草的淀粉含量持续增加,绣线菊先降低后增加,在N₃处理达到最小,而黄刺玫和茜草则呈现较复杂的变化趋势,辽东栎淀粉含量变化趋势平缓.在氮添加后,土壤pH、有机碳、全氮、全磷与植物叶片NSC含量无相关关系,仅N₀和N₃水平下上述土壤理化指标会显著影响可溶性糖/淀粉.表明林地不同植物叶片中NSC含量对氮添加的响应明显不同,研究全球氮沉降或氮添加对林地生态系统的影响需要考虑不同植物,尤其不同生活型植物的不同响应.     

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。     

圆叶玉兰叶片非结构性碳水化合物与氮、磷含量对海拔的响应

非结构性碳水化合物(non-structural carbohydrates, NSC)、氮(N)和磷(P)是植物生长的重要能源物质和影响植物分布的限制生长因子,圆叶玉兰(Magnolia sinensis)是四川省特有的珍稀濒危极小种群野生植物,研究其NSC、N和P可以反映它的营养供应水平及对环境的适应策略。选取芦山6个海拔梯度(1840,1960,2070,2170,2270,2390 m)的圆叶玉兰为对象,研究不同海拔下圆叶玉兰叶片中NSC与N、P及其化学计量间的关系。结果表明,圆叶玉兰叶片可溶性糖含量在2390 m处显著高于1840 m处, NSC含量在不同海拔差异极显著,随海拔增加呈"低-高-低"的单峰变化,2170 m处叶片NSC含量最高,碳水化合物供应充足;可溶性糖/淀粉的比值随海拔升高呈增大趋势,N含量和N/P比都随海拔上升而下降,且N/P比在各海拔上均小于14,NSC/N比在2390 m处显著高于1840 m处。总之,圆叶玉兰叶片的可溶性糖和NSC含量显著不受海拔的影响,较高的可溶性糖含量有利于抵御低温环境,其生长主要受氮元素限制而不受碳限制,反映了濒危植物圆叶玉兰在其有限的分布范围内NSC及N、P的保护策略,为圆叶玉兰的碳代谢和生长适应对策提供数据基础。     

云南普洱季风常绿阔叶林非结构性碳水化合物及其组分随环境的变化

为探索植物非结构性碳水化合物(NSC)组分随环境的变化特征,在云南普洱季风常绿阔叶林原始林选取6块60m×60m样地进行野外调查,各样地分别选择重要值前10位的物种进行取样,通过测定不同样地主要物种的NSC及其组分和样地环境因素,分析其非结构性碳水化合物与环境的关系。结果显示:(1)云南普洱季风常绿阔叶林植物中总体淀粉、可溶性糖、NSC含量分别为13.91%、3.31%、17.21%,淀粉和NSC含量在各器官中排序依次为根干枝叶,而可溶性糖含量排序依次为叶根枝干;淀粉、可溶性糖、NSC含量变异最大值分别为叶片19.09%、干15.32%和根16.30%。(2)云南普洱季风常绿阔叶林植物不同器官中淀粉含量仅与凋落物厚度、土壤pH及速效钾、全钾存在显著正或负相关关系,可溶性糖含量与海拔高度及土壤理化性质存在显著正或负相关关系;但茶梨与红椎中淀粉含量与土壤有效磷或全磷存在显著正相关关系,茶梨各器官中可溶性糖含量与所有环境因子均无显著相关性,红椎根中可溶性糖含量与pH值、枝中可溶性糖含量与全钾分别呈显著负相关关系;短刺椎各器官中淀粉含量与所有环境因子均无显著相关性,可溶性糖含量则与水解性氮、有效磷含量呈显著负相关关系,与坡度、全钾含量、凋落物厚度呈显著正相关关系。(3)单因素回归分析显示,海拔高度、有机质、水解性氮、全氮影响季风常绿阔叶林的总体可溶性糖含量,而其淀粉含量则主要受pH值影响。研究发现,季风常绿阔叶林NSC受环境因子影响,但不同NSC组分的环境影响因子不同。     

岳桦幼苗光合特性和非结构性碳水化合物积累对干旱胁迫的响应

以长白山林线树种岳桦为对象,利用生长控制试验进行干旱处理,研究干旱对岳桦幼苗光合特性及非结构性碳水化合物(NSC)积累的影响。结果表明:干旱显著降低了岳桦幼苗的净光合速率和气孔导度,提高了其水分利用效率;干旱显著增加了岳桦幼苗叶、皮、干和根中的可溶性糖和总NSC的含量,但显著降低了淀粉含量;随着干旱的持续,叶片的气孔导度、光合速率和瞬时水分利用效率迅速降低,而可溶性糖、淀粉和NSC则是先增后减;在试验末期,叶片90%发黄,岳桦幼苗干、皮和根中可溶性糖与淀粉含量的比值均显著高于对照。这表明岳桦在受到干旱胁迫时,迅速降低气孔导度以减少水分散失,提高水分利用效率,它属于避旱型植物;岳桦通过优先储存策略来提高组织器官中可溶性糖含量、增加淀粉与糖之间的转化率来应对水分亏缺的不利环境;在遭受持续干旱,幼苗面临死亡的时候,干旱胁迫可能超过了植物自我调节的阈值,但此时其组织器官中NSC含量并未降低,这说明岳桦最终的死亡可能不是碳饥饿导致的。     

沙地樟子松幼苗干旱致死过程中非结构性碳水化合物的变化

以2年生沙地樟子松幼苗为对象,通过持续自然干旱处理,研究当土壤含水量下降到田间持水量的60%、40%、30%、20%和15%时幼苗叶片水势及不同器官(一年生叶、当年生叶、茎、粗根和细根)的可溶性糖、淀粉和非结构性碳水化合物(NSC)的含量,分析沙地樟子松幼苗在干旱致死过程中各器官NSC的分配规律及其适应机制.结果表明: 土壤含水量从田间持水量的40%下降到15%,幼苗叶片凌晨及正午水势无显著变化.当土壤含水量从田间持水量的60%下降到30%,各器官可溶性糖、淀粉、NSC含量和可溶性糖/淀粉先下降后上升.从30%下降到20%,当年生叶、一年生叶、茎和细根可溶性糖、淀粉和NSC含量降低,而粗根可溶性糖含量增加,淀粉和NSC含量减少.从20%下降到15%,当年生叶、一年生叶和茎可溶性糖、淀粉和NSC含量降低,粗根可溶性糖和NSC含量下降,淀粉含量上升,细根可溶性糖含量减少,淀粉和NSC含量增加.沙地樟子松幼苗通过不断调整各器官NSC及其组分含量变化以适应不同干旱环境,土壤含水量下降到田间持水量的30%后,幼苗可溶性糖和NSC含量总体呈下降趋势,淀粉在粗根和细根中积累,幼苗可能因碳耗竭而死亡.     

隔离降雨对马尾松针叶非结构性碳水化合物和碳、氮、磷的影响

植物叶片中的非结构性碳水化合物（NSC）和碳（C）、氮（N）、磷（P）含量可反映植物和生态系统对水分亏缺环境的响应及适应程度。以福建省长汀县25a马尾松（Pinus massoniana）为对象,原位观测分析了3年持续100%隔离降雨对成年马尾松针叶NSC与C、N、P的影响。结果表明：（1）持续隔离降雨导致马尾松针叶NSC含量先显著增加后减少,最终导致针叶NSC含量季节变化消失;且针叶可溶性糖含量、可溶性糖/淀粉比值在后期显著增加;（2）持续隔离降雨使马尾松针叶N含量、P含量、N ： P均表现为前期与对照组差异不显著,后期显著高于对照组;（3）马尾松针叶NSC含量、可溶性糖含量、P含量、N ： P与土壤水分含量呈显著或极显著相关,且均与P含量显著或极显著相关,与N素无相关性。以上研究结果表明,随着土壤水分含量的持续减少马尾松可通过调整针叶中NSC含量的积累及分配和提高针叶N、P含量来适应缺水环境,P含量的增加对NSC含量波动及可溶性糖和淀粉的相互转化起促进作用。     

平潭岛海岸三种典型沙生植物非结构性碳水化合物含量特征研究

非结构性碳水化合物(non-structural carbonhydrate,NSC)作为光合作用的主要产物,其含量多少可反映植物整体的碳收支平衡关系及对外界环境胁迫的适应策略。该研究采用改进的苯酚-浓硫酸法对福建省平潭岛海岸处于生长旺盛期的3种典型沙生植物木麻黄、单叶蔓荆和老鼠艻茎叶的NSC及其组分含量进行测定。结果表明:(1)单叶蔓荆叶片、茎干可溶性糖含量均最大,分别为(137.83±3.75)、(90.74±2.65)mg·g~(-1),老鼠艻叶片、茎干淀粉含量均最大,分别为(105.41±2.49)、(120.48±7.67) mg·g~(-1),二者茎叶NSC含量均大于木麻黄,且全部达到显著水平(P0.05)。(2)沙堆表面老鼠艻叶片NSC及组分含量从雏形、发育到稳定阶段变化不明显,但茎干NSC及组分含量在雏形阶段沙堆表面显著大于其他两个演替阶段(P0.05),且依次呈现下降趋势。(3)从幼龄林、中龄林到成龄林阶段,木麻黄NSC含量呈现先上升后下降趋势,其中成龄林叶片可溶性糖、淀粉和NSC含量、茎干可溶性糖和NSC含量均显著低于其他两个生长阶段(P0.05)。(4)不同沙生植物对环境胁迫的适应策略各异,海岸沙丘表面植被恢复应充分考虑物种特性及其更新。     

云南普洱季风常绿阔叶林不同林层非结构性碳水化合物特征

在聚类分析的基础上,研究云南普洱季风常绿阔叶林主要物种非结构性碳水化合物(NSC)及其组分浓度、分配和季节性动态在林冠层、亚冠层和林下层间的变化特征.结果表明:亚冠层中可溶性糖及NSC浓度最高,分别为3.9%和13.3%,可溶性糖淀粉比在林下层最低,为0.76,而淀粉浓度则在各林层间无显著性差异.3个林层的可溶性糖均主要分配在叶片中,淀粉和NSC主要分配在根中.亚冠层中叶片和树干的可溶性糖浓度显著高于林冠层和林下层,枝和根的可溶性糖浓度在3个林层间无显著性差异;叶片的淀粉浓度则随林层高度降低而增加,但根淀粉浓度则是在林下层最低,为10.7%,枝和树干的淀粉浓度在3个林层间无显著差异;叶片NSC浓度为林冠层(10.7%)显著低于亚冠层(12.3%)和林下层(12.0%),但根的NSC浓度在林下层中最低,为14.2%;林下层叶片、枝、树干中可溶性糖淀粉比值均最低,但根的可溶性糖淀粉比值最低值出现在林冠层(0.79).3个林层NSC及其组分均存在显著的季节性变化,可溶性糖及可溶性糖淀粉比均为雨季显著高于旱季,而淀粉和NSC浓度则均在旱季中较高.不同林层NSC及其组分浓度的差异反映了不同高度树种碳利用策略的差异,部分地解释了物种的共存机制.     

Patterns in nonstructural carbohydrate contents at the tree organ level in response to drought duration

Nonstructural carbohydrates (NSCs) facilitate the adaptation of trees to drought stress. There have been a large number of studies exploring NSC changes in individual plant species and individual organ under drought and showed different trends; however, an understanding of the universal pattern of the plant NSCs responses to drought, particularly to drought duration, is still lacking. Here, we compiled data from 47 experimental studies on 52 tree species and conducted a meta‐analysis to evaluate the responses of soluble sugars, starch, and TNSC (total nonstructural carbohydrates including both soluble sugars and starch) concentrations in different tree organs (leaf, stem, and root) to drought intensity and duration. We found that starch in all organs decreased and soluble sugars in leaf increased with prolonged experiment time, and the changes in soluble sugars in all organs were stronger under severe drought than under slight‐to‐moderate drought. Under slight‐to‐moderate drought, the NSC content of each organ varied with time, while with the extension of the drought duration, the NSCs gradually approached the control value (no drought stress); this trend remained in the late drought, which means that trees activated physiological regulation processes to increase carbon storage and reduce the risks of carbon starvation. In contrast, long‐term severe drought could lead to a net loss of carbohydrates, especially in the root, implying that prolonged severe drought could lead to NSC depletion in the whole plant. As prolonged drought duration has occurred in and is projected for many regions, this paper could shed light into studies on how trees respond and adapt extending drought duration through nonstructural carbon production, transportation, and reallocation.     

Leaf non-structural carbohydrates regulated by plant functional groups and climate: Evidences from a tropical to cold-temperate forest transect

Non-structural carbohydrates (NSCs), e.g., glucose and starch, play important roles in metabolic processes of plants and represent important functional traits in plant's adaptation to external environment. To explore the variations in leaf NSCs among species and communities at a large scale and their influencing factors, we investigated the contents of leaf NSCs among 890 plant species in nine typical forests along the north–south transect of eastern China. The results showed that the contents of leaf soluble sugars, starch, and NSCs (sugars + starch) were highly variable among different plant species on the site scale, and their mean values for the 890 plant species were 45.7 mg g⁻¹, 47.5 mg g⁻¹, and 93.2 mg g⁻¹, respectively. All three metrics varied markedly across plant functional groups in the order of trees < shrubs < herbs. Weak latitudinal patterns of leaf soluble sugars, starch, and NSCs were observed from tropical to cold-temperate forests at the levels of species and plant functional groups. The contents of leaf soluble sugars, starch, and NSCs decreased with increasing temperature and precipitation which supports the growth limitation hypothesis at a large scale. In trees, leaf soluble sugars, starch, and NSCs increased with increasing photosynthetic active radiation (PAR); and were positively correlated with specific leaf area (SLA). The spatial patterns of leaf NSCs in forests along the north–south transect of eastern China and their relationships with temperature, precipitation, PAR, and SLA illustrate an important adaptation of plant communities to environmental changes at the continental scale.     

Leaf nonstructural carbohydrate concentrations of understory woody species regulated by soil phosphorus availability in a tropical forest

Leaf soluble sugars and starch are important components of nonstructural carbohydrates (NSCs), which are crucial for plant growth, development, and reproduction. Although there is a large body of research focusing on the regulation of plant NSC (soluble sugars and starch) concentrations, the response of foliar NSC concentrations to continuous nitrogen (N) and phosphorus (P) addition is still unclear, especially in tropical forests. Here, we used a long‐term manipulative field experiment to investigate the response of leaf NSC concentrations to continuous N and P addition (3‐, 5‐, and 8‐year fertilization) in a tropical forest in southern China. We found significant species‐specific variation in leaf NSC concentrations in this tropical forest. Phosphorus addition dramatically decreased both leaf soluble sugar and starch concentrations, while N addition had no significant effects on leaf soluble sugar and starch concentrations. These results suggest that, in plants growing in P‐limiting tropical soil, leaf NSC concentrations are regulated by soil P availability rather than N availability. Moreover, the negative relationships between NSC concentrations and leaf mass per area (LMA) revealed that NSCs could supply excess carbon (C) for leaf expansion under P addition. This was further supported by the increased structural P fraction after P fertilization in our previous study at the same site. We conclude that soil P availability strongly regulates leaf starch and soluble sugar concentrations in the tropical tree species included in this study. The response of leaf NSC concentrations to long‐term N and P addition can reflect the close relationships between plant C dynamics and soil nutrient availability in tropical forests. Maintaining relatively higher leaf NSC concentrations in tropical plants can be a potential mechanism for adapting to P‐deficient conditions.     

热带次生林不同林层植物叶片非结构性碳水化合物的季节变化及其对氮磷添加的响应

非结构性碳水化合物(Non-structural Carbohydrates, NSCs)是植物生长代谢过程中重要的能量来源。通过在华南热带次生林进行氮磷添加试验,探究不同林层植物叶片NSCs的季节变化及其对氮磷添加的响应,取样时间为2019年1月、4月、7月和10月。结果表明:1)植物叶片NSCs存在显著的种间差异,磷(P)添加对叶片淀粉和NSCs含量具有显著影响,且物种与磷添加的交互作用显著影响叶片淀粉含量。2)黑嘴蒲桃和紫玉盘叶片NSCs含量对氮(N)添加的响应较为敏感,而白车和竹节叶片NSCs含量对P添加的响应较为敏感,氮磷同时添加(+NP)对植物叶片NSCs的增效作用最好。3)植物叶片NSCs存在显著的季节性变化,且季节与林层间的交互作用对叶片可溶性糖和NSCs含量具有显著影响。4)不同林层植物对氮磷添加的响应不同,氮磷添加使林下层植物叶片可溶性糖含量增高,林冠层降低,在干季,N添加会使林下层植物叶片淀粉含量增高,林冠层降低。P添加的影响恰好与之相反。在湿季,氮磷添加使林下层和林冠层植物叶片的淀粉含量增加。5)林冠层植物叶片NSCs含量高于林下层,且林下层植物叶片NSCs含量...     

Homoeostatic maintenance of nonstructural carbohydrates during the 2015–2016 El Niño drought across a tropical forest precipitation gradient

Nonstructural carbohydrates (NSCs) are essential for maintenance of plant metabolism and may be sensitive to short‐ and long‐term climatic variation. NSC variation in moist tropical forests has rarely been studied, so regulation of NSCs in these systems is poorly understood. We measured foliar and branch NSC content in 23 tree species at three sites located across a large precipitation gradient in Panama during the 2015–2016 El Niño to examine how short‐ and long‐term climatic variation impact carbohydrate dynamics. There was no significant difference in total NSCs as the drought progressed (leaf P = 0.32, branch P = 0.30) nor across the rainfall gradient (leaf P = 0.91, branch P = 0.96). Foliar soluble sugars decreased while starch increased over the duration of the dry period, suggesting greater partitioning of NSCs to storage than metabolism or transport as drought progressed. There was a large variation across species at all sites, but total foliar NSCs were positively correlated with leaf mass per area, whereas branch sugars were positively related to leaf temperature and negatively correlated with daily photosynthesis and wood density. The NSC homoeostasis across a wide range of conditions suggests that NSCs are an allocation priority in moist tropical forests.     

蒙古莸幼苗干旱致死过程中非结构性碳水化合物的变化

以一年生蒙古莸幼苗为对象,设置适宜水分、慢速干旱致死和快速干旱致死3个处理,研究不同干旱强度致死下蒙古莸幼苗各器官中非结构性碳水化合物(NSC,包括可溶性糖和淀粉)的含量变化及其分配规律.结果表明:慢速干旱致死胁迫下各器官可溶性糖含量与适宜水分组无显著差异.随时间的推移,茎可溶性糖含量先增加后减少,淀粉和NSC含量增加;粗根可溶性糖含量减少,淀粉和NSC含量增加;叶可溶性糖含量增加,淀粉和NSC含量减少.致死时(80 d),叶、茎、粗根和细根的NSC含量分别为6.2%、7.8%、8.3%和7.4%.快速干旱致死胁迫下,各器官可溶性糖含量均高于适宜水分处理组,而淀粉和NSC含量均低于适宜水分组.随时间的推移,根可溶性糖含量下降,淀粉和NSC含量上升;茎可溶性糖、淀粉和NSC含量均上升;叶可溶性糖含量上升,淀粉和NSC含量下降.致死时(30 d),叶、茎、粗根和细根的NSC含量分别为5.9%、6.6%、8.9%和7.7%.应对不同的干旱致死情况,蒙古莸幼苗各器官间非结构性碳水化合物呈现出不同的动态变化.在慢速干旱致死胁迫下,NSC优先为维持各器官生理代谢活动提供能量;而在快速干旱致死下,NSC主要以可溶性糖形式维持植物代谢,调节渗透势,促进吸水,应对急剧的干旱胁迫.     

三个亚热带森林优势种凋落物非结构性碳水化合物含量的季节动态

非结构性碳水化合物(NSC)是凋落物中的易分解组分,在凋落物分解早期快速释放进入土壤并被微生物利用,参与森林土壤生物地球化学循环,因此新鲜凋落物中NSC变化规律是认识森林土壤碳和养分循环的关键之一。选取亚热带常绿阔叶林优势树种米槠(Castanopsis carlesii)和主要造林树种杉木(Cunninghamia lanceolata)、马尾松(Pinus massoniana)为研究对象,分析其新鲜凋落叶和凋落枝中NSC(可溶性糖和淀粉)含量的动态变化规律。结果表明：凋落物中NSC含量在不同月份表现出明显的时间动态,米槠、杉木和马尾松凋落叶和凋落枝中NSC含量总体上在11—12月呈上升趋势,而在2—6月呈缓慢下降趋势。不同类型的凋落物NSC含量存在显著差异,米槠、杉木和马尾松凋落叶中NSC含量分别为3.03%—3.56%、2.18%—4.37%、3.38%—4.89%,凋落枝中NSC含量分别为1.87%—4.22%、2.88%—4.28%、2.75%—5.27%,米槠和马尾松凋落叶中NSC含量高于凋落枝,而杉木凋落枝中NSC含量高于凋落叶。不同树种凋落物NSC含量差异显著,米槠和...     

Seasonal dynamics of non-structural carbohydrates in two species of mediterranean sub-shrubs with different leaf phenology

The seasonal dynamics of non-structural carbohydrates in the woody organs of two co-existing mediterranean sub-shrubs were analyzed. The two species show different leaf phenology during summer: Linum suffruticosum, maintains many of its green leaves, while Lepidium subulatum sheds most of its leaves. These different leaf phenologies are related to different strategies with regard to summer stress. The maintenance of leaves in Linum is related to its stress tolerance while Lepidium avoids stress by shedding its leaves. The main objectives were to: (1) determine the differences in the seasonal dynamics of non-structural carbohydrates among the main woody organs of both species; (2) verify if differences in the leaf phenology, and hence in the strategy with regard to summer drought, lead to different seasonal patterns of carbohydrate storage and use between the two species; (3) compare the seasonal dynamics of carbohydrates of the two studied sub-shrubs with those of mediterranean trees and shrubs previously reported in the literature. The concentration of soluble sugars (SS), starch and total non-structural carbohydrates (TNC) were assessed monthly, over 17 months, in the main roots, stems and the transition zone between root and shoot systems of both species. Starch storage capacity and SS, starch and TNC pools were calculated. The seasonal pattern of carbohydrate accumulation was similar among the woody organs analyzed, but it differed with those reported for mediterranean trees and shrubs. The two species showed different pools and seasonal patterns of non-structural carbohydrate concentrations in its woody organ, which corresponded to their different extent of leaf shedding. The stress-avoider Lepidium accumulated starch during spring shoot growth as a carbon store for summer respiration and had low pools of SS, whereas the stress-tolerant Linum increased SS during summer drought to maintain photosynthetic activity during summer and had low starch pools and storage capacity. However, irrespective of their different leaf shedding patterns, both species had a similar relative variation of their TNC concentration, which contrasts with previous results on deciduous and evergreen woody species.