坡向对暗紫贝母生长和繁殖特征的影响

在青藏高原东部的牛牛山和卡卡山,在其东、南、西、北4个坡面各设置一个海拔相同(3800m)的样地,每个样地中随机选择30株同龄级的暗紫贝母植株作为研究对象,然后比较各坡面的生态因子以及暗紫贝母的生长和繁殖特征。结果表明:1融雪时间、气温、土壤含水量在南北坡面间有显著的差异,但土壤p H在各坡面间无明显差异。2开始生长期、始花期和盛花期等物候阶段都是北坡最迟,南坡最早,而枯黄期在各坡面间无显著差异。3单叶面积、比叶面积和植株株高以北坡最大,南坡最小。4鳞茎生物量以南坡最大,北坡最小,东、西坡面的值介于南、北坡之间。5单粒果实的生物量在各坡面之间无明显的差异,果实生物量在地上部分中所占的比例以南坡最大,北坡最小。6单粒果实平均种子数以北坡最多,南坡最少;而种子千粒重则以南坡最重,北坡最轻。可见,坡面对暗紫贝母的生长和繁殖特征有显著的影响,尤其是在南北坡面之间大多数性状差异显著。     

2000-2016年秦岭山地植被覆盖变化地形分异效应

利用2000-2016年MODIS NDVI数据,采用趋势分析及地形差异修正法,探讨秦岭山地植被覆盖变化在南北坡、不同海拔以及不同坡度坡向下的空间分异性。结果表明：近17年来,秦岭山地植被覆盖度良好,整体呈上升趋势,南北坡、不同海拔、不同坡度、不同坡向下植被覆盖度有所差异,植被变化趋势也不同。（1）就南北坡而言,近17年来秦岭南坡植被覆盖度上升趋势大于北坡,南坡植被覆盖以上升趋势为主,而北坡以稳定为主。（2）不同的海拔高度上秦岭山地植被覆盖变化在存在分异性：低海拔区域呈减少趋势,中海拔区呈明显的上升趋势,2000 m以上的高海拔区域北坡的植被覆盖度较为稳定,而南坡的2500到3100 m区域内有较明显的减小趋势。（3）从坡度来看,随着坡度的增加秦岭山地植被覆盖度由减少转为增加再转为稳定,南北坡植被变化分异性不明显。（4）不同坡向上,秦岭南北坡植被覆盖度变化差异明显,由阴坡转为阳坡时,北坡植被覆盖有明显的增长趋势,而南坡则不明显,植被覆盖度减小区在南北坡的分布呈相反趋势,分别分布在南坡的阳坡以及北坡的阴坡。     

山西恒山温带草原与暖温带落叶阔叶林交错区植被生态研究

在中国植被分区上,山西恒山是温带草原地带与暖温带落叶阔叶林地带的交错区.采用TWINSPAN分类和DCA排序相结合的方法,对恒山南北坡植被类型进行了比较研究.TWINSPAN分类结果将南北坡的植被分别划分为16个群丛和20个群丛,分别隶属于16个和20个群系.DCA排序较好的印证了其分类结果：DCA排序第一轴反映海拔、温度的变化,南坡植被随海拔的升高,从耐旱的山蒿灌丛向寒温性的针叶林过渡,北坡由旱生的长芒草草原也向寒温性的针叶林过渡; 第二轴反映湿度的变化,排序轴从上到下,南坡由低海拔的山蒿群丛向高海拔的华北落叶松群丛,北坡是由长芒草、碱茅群丛向青杨群丛过渡.DCA排序结果显示,南北坡基带植被的群丛组成迥然不同,南坡基带为山蒿群丛、三裂叶绣线菊-山蒿＋赖草群丛、虎榛子-披针叶苔草群丛,是落叶阔叶林破坏后形成的次生植被类型;北坡基带为长芒草群丛、碱茅＋千里光群丛、三裂叶绣线菊-硬质早熟禾群丛、山菊＋硬质早熟禾群丛,具有典型的草原植被特征,这与它们所处的植被带有较高的吻合度.南北坡随海拔升高,植被类型表现出明显的趋同性.对南北坡群系的比较结果表明：有4个群系为南北坡共有,但群丛数量存在较大的差异,三裂叶绣线菊群系、山蒿群系在南坡的群丛数要比北坡多,披针叶苔草群系在北坡的群丛数要比南坡多,华北落叶松群系分布大致相同.     

1958-2008年太白山太白红杉林碳循环模拟

太白红杉(Larix chinensis林主要分布于我国秦岭太白山的林线位置,对气候变化的响应十分敏感.为了定量分析太白山太白红杉林在气候变化背景下的碳循环特征,基于模型(MTCLIM)模拟的温度和降水数据,应用植被动态过程模型(LPJ-GUESS)模拟了太白山南北坡1958-2008年太白红杉林的净初级生产力(NPP)、生物量和净生态系统碳交换量(NEE).结果表明:1)太白红杉和巴山冷杉(Abies fargesii)的NPP和生物量在太白红杉林占有优势,太白红杉的NPP和生物量均大于巴山冷杉.1958-2008年间太白红杉南北坡NPP的平均值为0.38 kgC·m-2·a-1,巴山冷杉为0.25 kgC·m-2,a-1,两者之和占整个太白红杉林NPP的86％;1958-2008年间太白红杉南北坡生物量的平均值为2.91 kgC/m2,巴山冷杉为2.02 kgC/m2,两者之和占太白红杉林生物量的94％.2)太白红杉和巴山冷杉的NPP均表现为北坡大于南坡,且南北坡均有逐年增加的趋势,北坡的增幅小于南坡,所以太白山南北坡太白红杉林的NPP差异有逐年减少的趋势.3)太白红杉生物量的年际波动较大,南北坡呈交替上升趋势,南坡的平均值(2.94 kgC/m2)大于北坡(2.89 kgC/m2).巴山冷杉生物量的年际波动相对较小,北坡生物量水平大于南坡.4)1958-2008年南北坡太白红杉林平均NEE均为-0.023 kgC·m-2·a-1,表现为碳汇.南北坡碳汇水平均呈逐年增加趋势,南坡的增加幅度(0.91 g·m-2·a-1)大于北坡(0.42 g·m-2·a-1).以气候和CO2为驱动因子对太白山太白红杉林的长期碳循环动态做了定量分析,从机理上揭示气候变化与生态系统碳循环的关系,还需要做进一步的野外观测和控制实验研究.     

秦岭太白红杉球果与种子特性及其与环境的关系

太白红杉(Larix chinensis)是研究秦岭高山林线与环境因子关系的理想树种, 球果与种子特征易受基因和环境的共同影响, 是决定林线迁移和分布格局的关键因素。应用单因素方差、多重比较分析坡向和居群间的球果与种子差异, 采用变异系数和表型分化系数分析球果与种子特征的变异来源, 运用Pearson相关系数、RDA排序分析球果与种子特征与环境因子的关系。结果表明: (1) 南坡和北坡的球果与种子特征差异不显著, 北坡4个球果与种子特征在不同居群间均存在显著差异(P<0.05), 南坡的球果重和平均种子数在不同居群间差异显著(P<0.05); (2) 南坡的变异系数和表型分化系数均高于北坡; 且南北坡球果与种子特征表型分化系数(Vst)均小于0.5, 说明变异主要存在于种群内, 异质的生境是球果与种子差异的重要因素; (3) 环境因子与太白红杉4个球果与种子特征相关系数的绝对值从大到小依次为: 海拔>土壤速效钾>坡向>pH值>土壤碱解氮>土壤有机质>土壤速效磷>坡度; 海拔与球果与种子特征平均相关性最强(r=-0.475), 是限制太白红杉分布的重要因素。研究表明, 球果与种子差异主要来自太白红杉对环境的响应, 海拔是限制太白红杉分布的重要因素。该研究揭示了林线树种太白红杉分布的限制因素, 为生长抑制假说提供了数据支撑, 也为该物种的保护提供理论依据。     

高寒草甸不同坡向条件下植物叶片δ~(13)C及水分利用效率的变化

通过对高寒草甸不同坡向条件下25科86种C3植物叶片稳定性碳同位素组成(δ~(13)C)的测定,研究了高寒草甸C3植物δ~(13)C和水分利用效率对坡向变化的响应以及环境因子对其产生的影响,分析了坡向上控制植物δ~(13)C变化的主要环境因子.结果表明:从北坡到南坡的生境梯度上,土壤含水量不断减少,而土壤温度及光照强度不断增加,植物群落结构也发生了相应变化.5个坡向中,植物叶片δ~(13)C值为-31.19‰~-21.80‰,均值为(27.18±0.13)‰;生长季植物叶片δ~(13)C均值在南坡最高,其次为西南坡、西坡、西北坡,最低值出现在北坡.坡向间δ~(13)C值的差异主要是由不同坡向土壤含水量和土壤温度,以及光照强度的不同导致的,其中土壤含水量是主要的限制因子.北坡-南坡梯度上植物叶片δ~(13)C值随土壤含水量下降、土壤温度及光照强度升高而变大,表明不同坡向植物的水分利用状况对干旱胁迫的响应不同,植物逐渐提高了水分利用效率以适应干旱胁迫的生境.     

藏东南色季拉山急尖长苞冷杉(Abies georgei var. smithii)林线的生态气候特征

高山林线因其对气候因子的敏感性和自身结构的不稳定性而被视为气候变化的"监视器",通过自动微气象站对藏东南色季拉山急尖长苞冷杉(Abies georgei var. smithii)林线气象因子的连续监测,探讨了色季拉山高山林线的生态气候特征.结果表明:在色季拉山急尖长苞冷杉林线处,年平均温度为1.3℃,年平均相对湿度为79.4%,全年降水量为892.6 mm,主要集中在6、7、8、9月4个月份.林线最热月平均温度为9.4℃,最冷月平均温度为-6.4℃,年生物温度为3.1℃,温暖指数是11.6℃月,寒冷指数是-55.9℃月,生长季内平均温度为7.7℃,生长季长度为133d,大陆度指数为34.0,湿润指数为217.5.其温暖指数与我国两种山地森林树种祁连圆柏和喜马拉雅冷杉的分布上限较为接近.林线的最热月温度、年生物学温度和生长季平均温度与全国高山林线的平均值十分接近.从全球范围来看,色季拉山林线最热月平均温度9.4℃与全球均值10℃很接近.生长季内10cm深度的土壤均温为6.22℃,接近Krner观测的(6.7±0.8)℃全球均值.     

南亚热带次生林土壤pH 值与含水量的空间异质性

在广西青秀山南坡和北坡的次生林内各建立1 块50 m×50 m 样地, 进行表层土壤(0-10 cm)网格取样, 并利用经典统计学与地统计学方法对土壤pH 值和含水量的空间异质性进行了分析。结果表明, 北坡和南坡的土壤pH 值均属弱变异性, 而土壤含水量均表现为中等程度变异。除北坡土壤水分含量的理论变异模型为指数模型外, 其余土壤指标均为球状模型。两块样地的土壤pH 值和含水量表现为中等程度的空间自相关, 这是由结构性和随机性因素共同作用的结果。Kriging 等值线图显示, 北坡土壤pH 值的空间变异格局呈现由左向右逐渐递增的趋势, 而南坡呈现从左到右逐渐递减的趋势; 北坡表土水分含量的最高值出现在样地的左上角和右下角区域, 而南坡土壤水分为从左到右逐渐递增。微地貌、群落盖度以及物种分布差异等是影响青秀山南亚热带次生林土壤pH 值和含水量空间变异和分布格局的主要因素。     

基于Biome-BGC模型及树木年轮的太白红杉林生态系统对气候变化的响应研究

利用Biome-BGC模型模拟了1960—2013年太白山太白红杉林生态系统的净初级生产力(NPP),对其与太白红杉的径向生长关系进行了探讨,并分析了NPP值对气候变化的响应关系。结果表明:1960—2013年太白山太白红杉林北坡NPP年均值为305.33g C m~(-2)a~(-1),南坡为320.71g C m~(-2)a~(-1),南北坡的NPP值均呈现出一定的上升趋势,北坡的上升速率(0.47g C m~(-2)a~(-1))要小于南坡(1.29g C m~(-2)a~(-1)),但是北坡太白红杉分布下限区NPP值波动浮动较大。且北坡太白红杉NPP值随着海拔高度的上升而逐渐下降,低海拔的变化振幅要大于高海拔地区,南坡无明显变化。多数采样点的模拟NPP值与树轮宽度指数年际变化趋势趋于一致,相关关系呈显著相关。太白红杉标准年表、模型模拟NPP值与气象因子的相关分析均表明太白红杉的生长与生长季气温的相关性显著高于降水,即生长季的气温是太白红杉生长的限制因子。气候的变化作为制约太白红杉生境的重要因素,影响了太白红杉树木的生长,进而对NPP的变化产生了影响。树木年轮很好的检验了Biome-BGC模型模拟结果。     

秦岭山地生态分界线的论证

山地自然生态分界线,实际为一条生态过渡带。分界线位置的准确划分,有助于把握现代气候-植被-土壤的敏感变化特征,也是指导当地大农业生产的基础依据。秦岭山地长期以来被认为是中国东部重要的南北生态分界线,即亚热带和暖温带的分界线。但对该界线的具体位置,历来存在着北坡、分水岭和南坡3种争议。根据作者多年的工作积累和对相关问题的思考,以文献评析为基础,以气候-植被参数计算结果为依据,并通过对秦岭山地不同区域植物区系和植被垂直带谱已有研究结果的比较分析,从中得出如下主要结论:(1)根据秦岭地区气候-植被参数等值线的分布,整个秦岭山地都应属于暖温带,这一结论可与国际有关研究相互接轨。(2)根据对秦岭不同区域植物区系调查结果的比较分析,秦岭南坡大部分区域都应属于暖温带,而南麓海拔1000m等高线附近很可能是暖温带的南缘。(3)根据对秦岭南北坡植被垂直带谱的比较分析,南麓海拔1000m等高线应该是一条重要的生态分界线。综合来看,将秦岭山地的南北分界线放在南坡海拔1000m附近比较适宜。上述归纳研究所获结论,将通过在秦岭山地深入开展现代"3S"技术支撑下的植被、土壤、气候等生态与地理方面的综合研究,最终得到完全验证。     

Nitrogen limitation and microbial diversity at the treeline

Tree growth limitation at treeline has mainly been studied in terms of carbon limitation while effects and mechanisms of potential nitrogen (N) limitation are barely known, especially in the southern hemisphere. We investigated how soil abiotic properties and microbial community structure and composition change from lower to upper sites within three vegetation belts (Nothofagus betuloides and N. pumilio forests, and alpine vegetation) across an elevation gradient (from 0 to 650 m a.s.l.) in Cordillera Darwin, southern Patagonia. Increasing elevation was associated with a decrease in soil N‐NH₄⁺ availability within the N. pumilio and the alpine vegetation belt. Within the alpine vegetation belt, a concurrent increase in the soil C:N ratio was associated with a shift from bacterial‐dominated in lower alpine sites to fungal‐dominated microbial communities in upper alpine sites. Lower forested belts (N. betuloides, N. pumilio) exhibited more complex patterns both in terms of soil properties and microbial communities. Overall, our results concur with recent findings from high‐latitude and altitude ecosystems showing decreased nutrient availability with elevation, leading to fungal‐dominated microbial communities. We suggest that growth limitation at treeline may result, in addition to proximal climatic parameters, from a competition between trees and soil microbial communities for limited soil inorganic N. At higher elevation, soil microbial communities could have comparably greater capacities to uptake soil N than trees, and the shift towards a fungal‐dominated community would favour N immobilization over N mineralization. Though evidences of altered nutrient dynamics in tree and alpine plant tissue with increasing altitude remain needed, we contend that the measured residual low amount of inorganic N available for trees in the soil could participate to the establishment limitation. Finally, our results suggest that responses of soil microbial communities to elevation could be influenced by functional properties of forest communities for instance through variations in litter quality.     

Comparison of Southern Appalachian high-elevation outcrop plant communities with their Northern Appalachian counterparts

Abstract. Southern Appalachian high-elevation outcrops harbour six regionally rare Northern Appalachian taxa usually considered relicts of a Pleistocene alpine flora. For five of the six taxa, minimum elevation in the south was 367–1113 m higher than in the north. While habitats compared between the two regions share only 9% of their total flora, individual plots had up to 70% of their species occurring in the opposite region. The northern affinity of southern outcrops increased with elevation, slope steepness, soil Cu, B and SO₄ and decreased with potential solar radiation and soil Na. As a result, communities above 1600 m on felsic bedrock, and above 1350 m on mafic bedrock, were most northern in composition. Northern affinity of southern outcrops also increased with latitude, which may partly result from closer geographic proximity to past communities that provided progenitors for the current northern flora. Northern treeless habitats increased in southern affinity with increased slope steepness, perennial seepage, vegetation height, shade, soil pH, Al, Mn, Na and decreased elevation and organic matter. As a result, northern outcrop communities below treeline were most similar to those on southern outcrops. This suggests that southern outcrop vegetation may be more similar to Pleistocene outcrop vegetation than to Pleistocene alpine vegetation. Partial constrained ordination showed that while compositional differences between the Northern and Southern Appalachian habitats were largely explained by environmental differences, there was a significant component of residual variation explained by north or south position that was unrelated to environment. These residual compositional differences may result from historical influences on community structure involving stochastic extinction and colonization processes.     

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.     

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.     

甘南高寒草甸植物元素含量与土壤因子对坡向梯度的响应

通过测定甘南高寒草甸不同坡向条件下25科86种植物叶片氮(N)、磷(P)、钾(K)含量、有机碳(C)含量、叶片含水量和相对叶绿素(SPAD)值,以及不同坡向的土壤含水量、有机碳、全氮、全磷含量等土壤指标,分析了不同坡向植物叶片元素含量与土壤环境因子之间的关系。研究结果表明,在南坡-北坡梯度上,随着土壤含水量的增加,植物叶片P含量、叶K含量和叶片含水量显著增加,而相对叶绿素显著降低。土壤养分含量与植物叶片P、叶K含量和叶含水量显著正相关,与叶片相对叶绿素显著负相关。说明不同坡向条件下叶片养分含量受土壤因子的影响显著,土壤的水分及养分状况对植物叶片元素含量的贡献不同。土壤含水量是坡向梯度上影响植物叶片特征的最主要因子。坡向梯度上土壤含水量对植物叶片各种元素含量的影响和植物叶片含水量对不同土壤因子的响应模式支持了生长在南坡的植物能以提高水分和养分利用效率而适应南坡较为干旱和贫瘠的生境。     

Effects of elevation and slope on the alpha and beta diversity of ground-dwelling beetles in Mt. Jirisan National Park,South Korea

We examined the diversity of ground-dwelling (epigaeic) beetles at different elevations of the northern and southern slopes of Mt. Jirisan National Park, South Korea. We selected eight study sites from both slopes and collected the beetles 10 times from May 2018 to August 2019 using pitfall traps. We collected a total of 67 species and 12,304 individuals and found higher species richness and abundance among the beetles from the northern slope (54 species and 6,969 individuals) than the southern slope (46 species and 5,335 individuals). We observed that the proportion of species based on the biogeographic affinity (Palearctic or Oriental) did not depend on the elevation and slope. The species richness increased with elevation and the abundance showed hump-shaped with a peak at 800 m. While the overall beta diversity was similar at the mountain and slope levels, the underlying processes such as spatial turnover and nestedness differed at the mountain and slope levels, respectively. We found that the ground beetle assemblages depended on elevation and soil characteristics such as soil organic matter and pH but were unaffected by the vegetation type.     

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。     

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。     

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.     

Responses of white spruce (Picea glauca) to experimental warming at a subarctic alpine treeline

From 2001 to 2004 we experimentally warmed 40 large, naturally established, white spruce [Picea glauca (Moench) Voss] seedlings at alpine treeline in southwest Yukon, Canada, using passive open‐top chambers (OTCs) distributed equally between opposing north and south‐facing slopes. Our goal was to test the hypothesis that an increase in temperature consistent with global climate warming would elicit a positive growth response. OTCs increased growing season air temperatures by 1.8°C and annual growing degree‐days by one‐third. In response, warmed seedlings grew significantly taller and had higher photosynthetic rates compared with control seedlings. On the south aspect, soil temperatures averaged 1.0°C warmer and the snow‐free period was nearly 1 month longer. These seedlings grew longer branches and wider annual rings than seedlings on the north aspect, but had reduced Photosystem‐II efficiency and experienced higher winter needle mortality. The presence of OTCs tended to reduce winter dieback over the course of the experiment. These results indicate that climate warming will enhance vertical growth rates of young conifers, with implications for future changes to the structure and elevation of treeline contingent upon exposure‐related differences. Our results suggest that the growth of seedlings on north‐facing slopes is limited by low soil temperature in the presence of permafrost, while growth on south‐facing slopes appears limited by winter desiccation and cold‐induced photoinhibition.