Carbon and nutrient physiology in shrubs at the upper limits: a multispecies study

高山分布上限灌木的碳与养分生理碳水化合物不足是高山林线树种生长限制假说之一,国内外以碳水化合物为基础的高山林线研究已有很多,而与高山林线相比,人们对碳水化合物在灌丛线形成中的作用知之甚少。除此之外,土壤养分亦被视为限制高山树种向上分布的重要因素之一。本研究将探究欧亚多种高山灌木不同器官中非结构性碳水化合物(NSCs)、氮(N)和磷(P)在不同季节及高低海拔上的含量变化规律。研究结果显示,除了与夏季相比冬季细根中具有较低的P含量以外,不同海拔与季节对灌木不同器官中的N和P含量均无显著影响。冬季灌木枝条中的NSCs和可溶性糖含量显著高于夏季。海拔与季节对细根中NSCs、淀粉、可溶性糖和糖与淀粉比值的影响均有显著的交互作用。在冬季,灌木细根中的可溶性糖与淀粉含量在海拔上限处要显著低于其在低海拔处;而在夏季,这些指标在高低海拔间均无显著差异。本研究结果表明,与高山林线树种相似,海拔分布上限的灌木冬季细根中较低的非结构性碳水化合物含量可能限制了灌木的向上分布。     

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。     

模拟增温和功能群去除对岷江冷杉幼苗存活和生长的影响

通过模拟增温和功能群去除试验,研究不同功能群的草本植物在岷江冷杉幼苗更新定植过程的作用,分析其在温度升高条件下生理指标的变化.结果表明:增温提高了岷江冷杉幼苗的存活率和植物体内非结构性碳水化合物含量,但抑制了幼苗的生长和根冠比.岷江冷杉幼苗的非结构性碳水化合物含量,尤其是可溶性糖含量与存活率呈显著正相关.不增温处理中,草本植物降低了幼苗的存活率,促进幼苗的株高生长和地上生物量积累,禾本科植物和杂类草抑制了幼苗根系生长和地下生物量的积累;增温处理中,杂类草的存在提高了幼苗的存活率,莎草科则显著抑制了幼苗根系生长和地下生物量的积累,禾本科植物和杂类草抑制了幼苗的高生长和地上生物量积累.模拟增温提高了林线树种岷江冷杉幼苗的存活率,但使其面临来自草本植物更剧烈的竞争,抑制其生长.     

川西高山林线交错带土壤动物对岷江冷杉和高山杜鹃凋落物分解的贡献

土壤动物是调控凋落物分解的重要生物因素.为了探究川西高山林线交错带土壤动物对两个优势物种岷江冷杉和高山杜鹃凋落物分解的贡献,在3个海拔梯度(针叶林-林线-高山草甸)采用凋落物分解袋试验,通过不同孔径的网袋(0.04 mm,基本排除土壤动物;3 mm,允许土壤动物通过),研究了分解554 d(2013年5月—2014年11月)土壤动物对凋落物的影响.结果表明: 在整个林线交错带上,岷江冷杉的分解速率(k)为0.209～0.243,高山杜鹃的k为0.173～0.189,岷江冷杉的k大于高山杜鹃.土壤动物的参与显著加速了两种凋落叶分解,同时土壤动物对两种凋落物分解的作用和贡献随海拔升高而降低.自针叶林、高山林线至高山草甸,土壤动物对岷江冷杉分解的质量损失率为15.2%、13.2%、9.8%,对高山杜鹃分解的质量损失率为20.1%、17.5%、12.4%;土壤动物对岷江冷杉分解的平均日贡献率为0.17%、0.13%、0.12%,对高山杜鹃分解的平均日贡献率为0.26%、0.25%、0.23%,土壤动物对高山杜鹃的分解影响相对较大.海拔、凋落物自身性质及其交互作用对土壤动物作用下凋落物的质量损失率和贡献率均表现出显著影响.土壤动物的作用于岷江冷杉和高山杜鹃分解的平均日贡献率在当年生长季(0.25%和0.44%)和次年生长季(0.10%和0.19%)均高于雪被期(0.07%和0.12%).回归分析表明,环境因子(日平均气温、冻融循环次数以及雪被厚度)可以解释土壤动物作用于岷江冷杉和高山杜鹃质量损失率的42.7%和50.9%,贡献率的43.2%和55.6%,这对了解土壤动物在凋落物分解中的作用和深入认识高山生态系统物质循环具有重要意义.     

西藏色季拉山林线冷杉种群结构与动态

急尖长苞冷杉(Abies georgei var.smithii)是西藏东南部地区高山林线森林群落的主要建群树种,主要分布在色季拉山海拔3600～4400m区域,并成为阴坡高山林线的优势树种。通过对色季拉山林线群落交错区域的定位调查,分析了急尖长苞冷杉的个体生长、种群结构与动态以及林线特征。分析结果表明:(1)西藏色季拉山海拔4320m处为森林郁闭上限,该区域存在两种类型的高山林线,阳坡为渐变型林线,林线树种为方枝柏(Sabina saltuaria);阴坡为急变型林线,林线树种为急尖长苞冷杉。阳坡与阴坡林线分布海拔上限分别为4570m和4390m,阳坡高于阴坡180m;阳坡与阴坡林线群落交错区垂直宽度分别为250m和70m,阳坡比阴坡宽180m。(2)阴坡海拔3700～3800m属急尖长苞冷杉分布的最适范围,种群径级结构表现为典型的反"J"型,种群密度约380株/hm2;种群年龄结构表现为"金字塔"型,属于扩展型种群。(3)静态生命表和种群存活曲线反映了急尖长苞冷杉种群在形成初期的20a和生长发育的60～160a分别经历了强烈的环境筛选和竞争自疏,以及后期与环境变化相关的死亡波动,200a左右为急尖长苞冷杉的生理寿命,种群后期基本稳定,400a左右为极限寿命。     

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。     

高山林线形成机理研究进展

高山林线是郁闭森林与高山植被之间的分布界限,作为重要的生态过渡带,对全球和区域性气候变化的反应极为敏感,被认为是气候变化的理想监测器.高山林线研究从最初的形态描述到林线成因假说都是为了寻找高山林线形成的原因.迄今出现的高山林线成因假说都能够在局地尺度解释高山林线成因,但仍然缺乏可以普遍解释全球高山林线现象的假说.温度是林线分布的限制因子,低温限制了林线树种的生存及生长,但是低温影响了哪一个生化过程仍不明确,其影响机理还需进一步探讨.本文对高山林线形成机理,特别是对低温对高山林线植物光合特性、养分特征、非结构性碳水化合物和抗氧化系统的影响等研究进展进行综述,并提出了未来林线研究应该关注的问题.     

升温突变对川西不同坡向和海拔岷江冷杉径向生长的影响

坡向和海拔作为重要的地形因子决定着林木立地条件的水热分配，为揭示川西地区升温突变前后制约岷江冷杉(Abies fargesii var.faxoniana)径向生长的气候因子变化及其在坡向和海拔上的响应格局，通过树木年轮学方法构建了4个坡向(NE、N、NW和W)的3个海拔梯度(≥3650 m)共12个岷江冷杉的标准年表，采用皮尔逊相关分析和回归分析的方法分析了升温突变前后(1980年)限制岷江冷杉径向生长主要气候因子的变化及径向生长的变化趋势。结果表明：升温前，生长季(7月)低温和降水、前一年冬季(10—11月)最高温和平均温制约岷江冷杉的生长，而当年春季(3月)温度的升高和生长季前(5月)较多的降水不利于其生长。升温后，4个坡向的林线、东北坡中海拔，西坡中、低海拔岷江冷杉的径向生长与大部分月温度表现为正相关关系，且上述样点树木径向生长明显加速。1980年升温前，制约不同样点岷江冷杉径向生长的气候因子具有一致性——坡向和海拔对其径向生长与温度相关关系的干扰和影响较小。增温促进了各坡向林线岷江冷杉的径向生长，且偏阳坡在更大的海拔范围内对增温表现出正反馈。研究对未来气候变化背景下川西不同坡...     

贡嘎山雅家埂峨眉冷杉林线种群的时空动态

通过对贡嘎山雅家埂峨眉冷杉种群林线附近6个3000 m2样地(阴阳坡各3个)中峨眉冷杉(Abies fabri Craib)种群的定位调查,分析了过去100a间该区峨眉冷杉种群的时间-空间动态。结果表明:1)雅家埂林线附近峨眉冷杉种群密度在过去100 a(主要是近50 a)有显著的升高,但树线的海拔位置并无明显的爬升;2)阴阳坡林线格局存在显著的坡向分异:阴坡林线和树线的海拔高度显著高于阳坡(分别比阳坡高152.5 m和135.8 m),阳坡林线附近峨眉冷杉早期的生长速率在大于阴坡,但后期的生长速率却低于阴坡;3)热量(温度)控制假说不能完全解释雅家埂目前的树线格局,除气候因素之外,其它因素也限制了雅家梗地区树线位置的变化。     

川西亚高山林线过渡带及邻近植被土壤性质

研究了川西亚高山林线过渡带及邻近植被上缘流石滩草甸与下缘冷杉林残积母质土壤物理、化学和生物学特性.结果表明:从流石滩草甸→林线过渡带→冷杉林,土层逐渐增厚,表层土壤(0～30 cm)粉粒、粘粒、物理性粘粒含量、团聚度、结构系数和自然含水量逐渐增高,砂粒含量逐渐降低,阳离子交换量(CEC)、交换性盐基含量、盐基饱和度以及水解性酸含量逐渐增高,pH值逐渐降低,有机质及养分库全P、全K、有效N、有效P和速效K含量逐渐增高,土壤物理结构和化学性质逐步有所改善,但亚高山林线过渡带区域残积母质土壤成土过程缓慢.林线过渡带表层土壤细菌、真菌、放线菌及微生物总量,土壤脲酶、蔗糖酶、酸性磷酸酶、中性磷酸酶、多酚氧化酶及过氧化氢酶活性高于邻近植被上缘流石滩草甸与下缘冷杉林,体现了作为生态交错带,林线过渡带比邻近植被土壤具有相对较强的生物学活性.     

Spatio-temporal dynamics of two alpine treeline ecotones and ecological characteristics of their dominate species at the eastern margin of Qinghai-Xizang Plateau

热量匮乏是高山树线的主要成因, 在全球变暖趋势下对高山树线及其建群种的生态学过程及特征的研究具有重要意义。该文以青藏高原东缘的折多山和剪子弯山两处高山树线(海拔分别为4 265 m和4 425 m)作为研究对象, 通过设置垂直样带, 同时结合区域温度、降水的长时间序列分析, 探究两处树线的时空动态过程, 并明确了建群种冷杉(Abies spp.)的生态学特征。结果表明: 1)折多山和剪子弯山区域的气温在过去58年均存在显著的上升趋势(分别上升了0.72和0.91 ℃), 而折多山和剪子弯山区域降水均存在微弱的降低趋势。2)折多山的峨眉冷杉(A. fabri)龄级结构呈反J形, 剪子弯山的鳞皮冷杉(A. squamata)龄级结构呈双峰形, 二者种群结构均相对稳定。3)在小尺度上, 种子扩散限制使得两处树线的冷杉聚集分布。在大尺度上, 折多山峨眉冷杉亦呈聚集分布, 而剪子弯山鳞皮冷杉受生长环境以及种内或种间关系的影响呈随机分布。4)两处样地建群树种的树高和基径均随海拔升高而降低, 位于树线交错带上部的冷杉均呈现树高生长大于径向生长的异速生长关系, 而位于样地中、下部位的冷杉大部分呈等速生长关系。5)相比10年前, 折多山和剪子弯山的树线及树种线位置均无明显变化, 剪子弯山鳞皮冷杉种群的树木密度亦无明显变化, 而折多山的树木个体数提高了约25%; 相比20年前, 折多山和剪子弯山的树种线分别上移了50和30 m, 树线位置分别升高了75和40 m, 树木个体数亦明显增加, 分别提高了约220%和100%。树线及其建群种在较大时空尺度上主要受热量的控制, 而在较小时空尺度上受温度及生长环境共同作用的影响。     

青藏高原东缘两处高山树线交错带时空动态及其建群种的生态学特征

热量匮乏是高山树线的主要成因, 在全球变暖趋势下对高山树线及其建群种的生态学过程及特征的研究具有重要意义。该文以青藏高原东缘的折多山和剪子弯山两处高山树线(海拔分别为4 265 m和4 425 m)作为研究对象, 通过设置垂直样带, 同时结合区域温度、降水的长时间序列分析, 探究两处树线的时空动态过程, 并明确了建群种冷杉(Abies spp.)的生态学特征。结果表明: 1)折多山和剪子弯山区域的气温在过去58年均存在显著的上升趋势(分别上升了0.72和0.91 ℃), 而折多山和剪子弯山区域降水均存在微弱的降低趋势。2)折多山的峨眉冷杉(A. fabri)龄级结构呈反J形, 剪子弯山的鳞皮冷杉(A. squamata)龄级结构呈双峰形, 二者种群结构均相对稳定。3)在小尺度上, 种子扩散限制使得两处树线的冷杉聚集分布。在大尺度上, 折多山峨眉冷杉亦呈聚集分布, 而剪子弯山鳞皮冷杉受生长环境以及种内或种间关系的影响呈随机分布。4)两处样地建群树种的树高和基径均随海拔升高而降低, 位于树线交错带上部的冷杉均呈现树高生长大于径向生长的异速生长关系, 而位于样地中、下部位的冷杉大部分呈等速生长关系。5)相比10年前, 折多山和剪子弯山的树线及树种线位置均无明显变化, 剪子弯山鳞皮冷杉种群的树木密度亦无明显变化, 而折多山的树木个体数提高了约25%; 相比20年前, 折多山和剪子弯山的树种线分别上移了50和30 m, 树线位置分别升高了75和40 m, 树木个体数亦明显增加, 分别提高了约220%和100%。树线及其建群种在较大时空尺度上主要受热量的控制, 而在较小时空尺度上受温度及生长环境共同作用的影响。     

Seasonal dynamics of mobile carbon supply in Quercus aquifolioides at the upper elevational limit

Many studies have tried to explain the physiological mechanisms of the alpine treeline phenomenon, but the debate on the alpine treeline formation remains controversial due to opposite results from different studies. The present study explored the carbon-physiology of an alpine shrub species (Quercus aquifolioides) grown at its upper elevational limit compared to lower elevations, to test whether the elevational limit of alpine shrubs (<3 m in height) are determined by carbon limitation or growth limitation. We studied the seasonal variations in non-structural carbohydrate (NSC) and its pool size in Q. aquifolioides grown at 3000 m, 3500 m, and at its elevational limit of 3950 m above sea level (a.s.l.) on Zheduo Mt., SW China. The tissue NSC concentrations along the elevational gradient varied significantly with season, reflecting the season-dependent carbon balance. The NSC levels in tissues were lowest at the beginning of the growing season, indicating that plants used the winter reserve storage for re-growth in the early spring. During the growing season, plants grown at the elevational limit did not show lower NSC concentrations compared to plants at lower elevations, but during the winter season, storage tissues, especially roots, had significantly lower NSC concentrations in plants at the elevational limit compared to lower elevations. The present results suggest the significance of winter reserve in storage tissues, which may determine the winter survival and early-spring re-growth of Q. aquifolioides shrubs at high elevation, leading to the formation of the uppermost distribution limit. This result is consistent with a recent hypothesis for the alpine treeline formation.     

贡嘎山东坡植物区系的垂直分布格局

为了探讨贡嘎山植物区系的垂直分化特征及其与周边地区植物区系的联系,结合样带法与样方法,对贡嘎山东坡垂直植被带进行了调查,统计得出各垂直植被带的科、属的物种数量,分析了科、属、种级区系成分的构成及其沿海拔梯度的分布格局,并对各垂直植被带区系的相似性进行了聚类分析。结果表明：1)贡嘎山植物区系在整体上具有温带性质,但在干旱河谷地带,热带和温带区系成分的比例相当：热带成分的构成和分布反映古热带和古地中海区系的残遗性影响;2)东亚(含亚型)和东亚-北美成分对贡嘎山中部森林植物区系的影响最大,这些成分以温带古老性质为主;3)北温带成分是贡嘎山植物区系的主体之一,对青藏高原隆升以来贡嘎山植物区系进化类群和特有成分的发展有主要贡献,代表区系的年轻组分;4)中国特有种类型多样,占不同垂直植被带物种数量的40％-65％,其比例随海拔上升而增大。各类型比例的垂直变化突出反映了贡嘎山及横断山脉中海拔地段的植物区系与华中地区的联系,以及高海拔地段与青藏高原及东喜马拉雅的区系之间的联系。本文还就贡嘎山在生物地理分布上的意义以及贡嘎山和横断山脉植物区系特有性的性质进行了讨论。     

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.     

Carbon balance of conifer seedlings at timberline: relative changes in uptake,storage, and utilization

Low rates of growth for conifers at high elevations may relate to problems in acquiring or utilizing carbon. A traditional hypothesis for growth limits of trees at alpine treeline is that low photosynthesis (A) leads to insufficient supply of carbon for growth. Alternatively, the sink-limitation hypothesis questions the importance of low A, and suggests that trees at treeline have abundant carbon for growth as a result of greater decreases in carbon use [respiration (R) and growth] than A at higher elevations. Concentrations of carbon intermediates (e.g., nonstructural carbohydrates, NSC) have been used to support the sink-limitation hypothesis, with the assumption that NSC reflects the ratio of carbon source to sinks. Our objective was to determine elevation effects on carbon balance (whole-plant uptake, storage, and efflux of carbon) of tree seedlings during their critical establishment phase at timberline. Changes in A and R (of whole crowns), NSC (starches and simple sugars), and growth were measured in seedlings of a treeline (Abies lasiocarpa) and nontreeline species (Pseudotsuga menziesii). Seedlings were outplanted at the lower (2,450 m) and upper (3,000 m) edges of the timberline zone in the Rocky Mountains, USA. At the upper compared with lower elevation, both species had 10-20% less root and needle growth, 13-15% less A, 35-39% less R, and up to 50% greater NSC. A. lasiocarpa allocated more biomass to needles and had greater A but less NSC than P. menziesii. The greater effects of elevation on R compared with A led to a 1.3-fold increase in A:R at the upper elevation, and a corresponding increase in starch (r2 = 0.42)-a pattern consistent with the predictions of the sink-limitation hypothesis. Nevertheless, A and dry mass gain were positively correlated (r2 = 0.42), indicating that variation in photosynthesis was related to growth of seedlings at timberline.     

Variation of mobile carbon reserves in trees at the alpine treeline ecotone is under environmental control

? In low temperature-adapted plants, including treeline trees, light-saturated photosynthesis is considerably less sensitive to temperature than growth. As a consequence, all plants tested so far show increased nonstructural carbohydrate (NSC) tissue concentrations when exposed to low temperatures. Reduced carbon supply is thus an unlikely cause for low temperature range limits of plants. For altitudinal treeline trees there is, however, a possibility that high NSC genotypes have been selected. ? Here, we explored this possibility using afforestations with single-provenance conifers along elevational gradients in the Southern Chilean Andes and the Swiss Alps. ? Tree growth was measured at each of four approximately equidistant elevations at and below the treeline. Additionally, at the same elevations, needle, branch and stem sapwood tissues were collected to determine NSC concentrations. ? Overall, growth decreased and NSC concentrations increased with elevation. Along with previous empirical and experimental studies, the findings of this study provide no indication of NSC reduction at the treeline; NSC increased in most species (each represented by one common population) towards their upper climatic limit. The disparity between carbon acquisition and structural carbon investment at low temperature (accumulation of NSC) thus does occur even among genotypes not adapted to treeline environments.     

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.