海拔梯度对长白山北坡岳桦水分利用效率的影响

以分布于长白山北坡海拔1800～2050 m的岳桦林为对象,以叶片碳稳定同位素(δ13C)值作为岳桦长期水分利用效率指示值,探讨海拔梯度对岳桦林水分利用效率的影响.结果表明:随海拔升高,岳桦林土壤体积含水量、比叶质量极显著增加,而叶片含水量和土壤温度显著降低;岳桦叶片δ13C值与海拔呈极显著正相关,增幅为1.013‰·(100 m)-1,与土壤体积含水量、比叶质量呈显著正相关,与生长季土壤平均温度、叶片含水量呈显著负相关.温度不是长白山林线的唯一限制性因子,海拔梯度上水热条件的差异及其交互作用可能造成岳桦生长期间的生理干旱.     

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。     

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。     

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(650m和970m),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。     

长白山林线树种岳桦幼树叶功能型性状随海拔梯度的变化

通过研究沿不同海拔岳桦幼树叶功能型性状,揭示其对环境的响应机制.结果表明:①随海拔升高,岳桦叶面积(LA)逐渐降低,比叶重(LMA)增加,但LMA较高的可塑性指数表明其适应更依赖叶片的薄厚变化;②岳桦叶绿素含量随海拔升高而显著下降,但类胡萝卜素Car和Car/Chl显著升高,Chlb和Car/Chl表现出较高的可塑性指数,更倾向于吸收蓝紫光和保护光合器官;③岳桦叶氮含量(Narea和Nmass)在海拔1800-1900m间最低,在低海拔和高海拔均表现较高,但Chl/Nmass却随海拔升高而显著增加,Nmasss比Narea具有较高的可塑性指数,对光能的吸收更依赖Nmass对Chl的贡献,高海拔主要将更多的氮投资于光合器官的保护(1900m以上),低海拔则更倾向于光合生产(1800m以下);④随海拔升高,MDA增加,但随之抗氧化物质DS、Pro和APX活性增加,负责对活性氧的抵御和清除,但APX活性最大的可塑性指数表明活性氧的清除更依赖于酶促系统,但在海拔1900m以上,APX活性差异不显著,生理抗性逐渐下降,限制岳桦继续向高海拔生长;⑤抗氧化物质可塑性指数最高,叶绿素和叶形态次之,叶氮最低,表明随海拔升高,岳桦林以保护自身的生存为最主要的适应策略机制,然后以增加吸收光能的Chlb及LMA指标为主要生长策略.     

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。     

巴郎山大叶醉鱼草叶片非结构性碳水化合物和氮分配的海拔响应

为深入认识和探讨植物对环境变化的生理生态响应和适应,以分布在川西巴郎山的大叶醉鱼草(Buddleja davidii)为研究对象,沿海拔梯度对植物叶片中非结构性碳水化合物(NSC)、可溶性糖和淀粉含量,氮含量和氮分配比例(光合系统氮分配比例P_P、细胞壁氮分配比例P_CW和其他组分氮分配比例P_other)等参数进行对比分析,探讨其沿海拔的变化趋势,以及叶片NSC、可溶性糖和淀粉含量与氮分配间的相关关系。结果显示：大叶醉鱼草叶片NSC、可溶性糖、淀粉和单糖含量随海拔的升高而增加,而可溶性糖/淀粉比值未发生显著变化,表明高海拔较高的NSC含量的累积是由可溶性糖和淀粉含量共同决定的,而可溶性糖含量的增加主要由单糖含量的变化引起。叶片氮含量和P_P在海拔间差异不显著,但P_CW和P_other分别随海拔升高而降低和升高。此外,随海拔升高,叶片NSC/N比值随之增加,这主要归因于随海拔升高而增加的NSC含量而非海拔间差异不显著的氮含量。NSC含量和可溶性糖含量均与P_...     

模拟氮沉降对长白山岳桦林下草本植物和土壤肥力的短期影响

选取长白山岳桦林中的岳桦-蟹甲草群落(Comm. Betula ermanii-Parasenecio forrestii)、岳桦-藜芦群落(Comm. Betula ermanii-Veratrum nigrum)和岳桦-小叶章群落(Comm. Betula ermanii-Deyeuxia purpurea)开展野外模拟氮沉降实验,采用野外原位模拟实验方法,设置对照(0 kg·hm-2·a-1)、低氮(30 kg·hm-2·a-1)、中氮(50 kg·hm-2·a-1)和高氮(100 kg·hm-2·a-1)4个氮处理水平,测定草本植物生长状况和土壤肥力,研究岳桦林下草本层植物和土壤肥力对氮沉降的短期响应。结果显示:(1)岳桦林下草本植物随氮沉降量的增加而加速生长,小叶章对氮沉降的响应较为敏感,藜芦次之,蟹甲草最弱;(2)氮添加造成林下土壤肥力发生变化,有机质含量下降,特别是岳桦-小叶章群落下的土壤有机质含量下降最明显;土壤总氮和速效氮含量增大,岳桦-蟹甲草群落下的土壤总氮和速效氮增加最多;土壤总磷和速效磷含量减小,岳桦-小叶章群落下的土壤总磷和速效磷含量的减少最多。本研究结果表明氮添加在短期内会促进长白山岳桦林下草本植物生长,尤其是小叶章的生长,加快土壤有机质的分解和磷的释放,逐步改变土壤肥力并反馈给植物,促使其进一步变化。     

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...     

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。     

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.     

Temporal variations of mobile carbohydrates in Abies fargesii at the upper tree limits

Low temperatures are associated high‐altitude treelines, but the functional mechanism of treeline formation remains controversial. The relative contributions of carbon limitation (source activity) and growth limitation (sink activity) require more tests across taxa and regions. We examined temporal variations of mobile carbon supply in different tissues of Abies fargesii across treeline ecotones on north‐ and south‐facing slopes of the Qinling Mountains, China. Non‐structural carbohydrate (NSC) concentrations in tissues along the altitudinal gradient on both slopes changed significantly in the early and late growing season, but not in the mid‐growing season, indicating the season‐dependent carbon supply status. Late in the growing season on both slopes, trees at the upper limits had the highest NSC concentrations and total soluble sugars and lowest starch concentrations compared to trees at the lower elevations. NSC concentrations tended to increase in needles and branches throughout the growing season with increasing elevation on both slopes, but declined in roots and stems. NSC concentrations across sampling dates also indicated increases in needles and branches, and decreases in roots and stem with increasing elevation. Overall altitudinal trends of NSC in A. fargesii revealed no depletion of mobile carbon reserves at upper elevation limits, suggesting limitation of sink activity dominates tree life across treeline ecotones in both north‐ and south‐facing slopes. Carbon reserves in storage tissues (especially roots) in the late growing season might also play an important role in winter survival and early growth in spring at upper elevations on both slopes, which define the uppermost limit of A. fargesii.     

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.     

Variations in carbon source–sink relationships in subalpine fir across elevational gradients

• Cold‐adapted trees display acclimation in both carbon source and carbon sink capacity to low‐temperature stress at their upper elevational range limits. Hence a balanced carbon source–sink capacity might be required for their persistence and survival at the elevational tree limits.
• The present study examined the spatial dynamics of carbon source–sink relationship in subalpine fir (Abies fargesii) trees along elevational gradients in the northern slope of the temperate region and in the southern slope of the subtropics in terms of climate in the Qinling Mountain range, north‐central China.
• The results showed that non‐structural carbohydrate (NSC) concentrations in both the source and sink tissues increased with the increase in elevation. The ratio of carbon source–sink displayed a consistent decreasing trend with the increase in elevation and during growing season, showing that it was lowest at a ratio of 2.93 in the northern slope and at a ratio of 2.61 in the southern slope at the upper distribution elevations in the late growing season. Such variations of carbon source–sink ratio might be attributable to the balance between carbon source and sink activities, which changed seasonally across the elevational distribution range.
• We concluded that a ratio of carbon source–sink of at least 2.6 might be essential for subalpine fir trees to persist at their upper range limits. Therefore, a sufficient source–sink ratio and a balanced source–sink relationship might be required for subalpine fir trees to survive and develop at their upper elevational distribution limits.

     

秦岭东段栓皮栎叶片非结构性碳水化合物含量的季节动态

研究树木叶片非结构性碳水化合物(Nonstructural carbohydrate,NSC)组分的季节变化是掌握树木碳代谢规律的基础,也有利于判断以往研究仅凭生长季单次或几次(5次)的取样方法是否存在一定局限性。以秦岭东段栓皮栎(Quercus variabilis Blume)优势群落为研究对象,在其分布的海拔上下限(650 m和970 m),于2016年5月至2017年5月,通过月尺度周期性取样(共计9次),测定栓皮栎叶片NSC及其组分含量,并观测同期叶片物候变化。结果显示:(1)栓皮栎叶片NSC及其组分季节变化差异显著(P0.05),可溶性糖、淀粉和NSC变异系数分别为20.99%、52.28%和25.96%;(2)整体而言,栓皮栎叶片NSC最小值在展叶初期(3月末—4月初,5%左右),最大值在展叶末期(5月上旬,12%左右),之后NSC呈持续下降趋势。不同海拔NSC极值出现时间略有不同,叶片物候可能是影响年内极值的主要原因。(3)栓皮栎叶片NSC组成以可溶性糖为主(65%),这可能是树种在暖温带所采取的生长策略。(4)海拔对栓皮栎叶片NSC及其组分影响差异不显著,低海拔栓皮栎叶片NSC及其组分含量略大于高海拔。研究结果,栓皮栎叶片NSC含量存在明显的季节波动,适当加大NSC采样频率对于正确理解树木碳代谢十分必要。     

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.     

Non-structural carbohydrate pools in a tropical forest

The pool size of mobile, i.e. non-structural carbohydrates (NSC) in trees reflects the balance between net photosynthetic carbon uptake (source) and irreversible investments in structures or loss of carbon (sink). The seasonal variation of NSC concentration should reflect the sink/source relationship, provided all tissues from root to crown tops are considered. Using the Smithsonian canopy crane in Panama we studied NSC concentrations in a semi-deciduous tropical forest over 22 months. In the 9 most intensively studied species (out of the 17 investigated), we found higher NSC concentrations (starch, glucose, fructose, sucrose) across all species and organs in the dry season than in the wet season (NSC 7.2% vs 5.8% of dry matter in leaves, 8.8/6.0 in branches, 9.7/8.5 in stems, 8.3/6.4 in coarse and 3.9/2.2 in fine roots). Since this increase was due to starch only, we attribute this to drought-constrained growth (photosynthesis less affected by drought than sink activity). Species-specific phenological rhythms (leafing or fruiting) did not overturn these seasonal trends. Most of the stem volume (diameter at breast height around 40 cm) stores NSC. We present the first whole forest estimate of NSC pool size, assuming a 200 t ha^–1 forest biomass: 8% of this i.e. ca. 16 t ha^–1 is NSC, with ca. 13 t ha^–1 in stems and branches, ca. 0.5 and 2.8 t ha^–1 in leaves and roots. Starch alone (ca. 10.5 t ha^–1) accounts for far more C than would be needed to replace the total leaf canopy without additional photosynthesis. NSC never passed through a period of significant depletion. Leaf flushing did not draw heavily upon NSC pools. Overall, the data imply a high carbon supply status of this forest and that growth during the dry season is not carbon limited. Rather, water shortage seems to limit carbon investment (new tissue formation) directly, leaving little leeway for a direct CO₂ fertilization effects.