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.     

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.     

Distinguishing local from global climate influences in the variation of carbon status with altitude in a tree line species

Aim Two alternative hypotheses attempt to explain the upper elevation limit of tree lines world‐wide, the carbon‐limitation hypothesis (CLH) and the growth‐limitation hypothesis (GLH); the altitudinal decrease of temperature is considered the driver constraining either carbon gain or growth. Using a widely distributed tree line species (Nothofagus pumilio) we tested whether tree line altitude is explained by the CLH or the GLH, distinguishing local from global effects. We elaborated expectations based on most probable trends of carbon charging with altitude according to both hypotheses, considering the alternative effects of drought. Location Two climatically contrasting tree line ecotones in the southern Andes of Chile: Mediterranean (36°54′ S) and Patagonia (46°04′ S). Methods At both locations, 35–50 trees of different ages were selected at each of four altitudes (including tree line), and stem and root sapwood tissues were collected to determine non‐structural carbohydrate (NSC) concentrations. NSC accumulates whenever growth is more limited than photosynthesis. An altitudinal increase in NSCs means support for the GLH, while the opposite trend supports the CLH. We also determined stable carbon isotope ratios (δ¹³C) to examine drought constraints on carbon gain. Results NSC concentrations were positively correlated with altitude for stem tissue at the Mediterranean and root sapwood tissue at the Patagonia site. No depletion of NSC was found at either site in either tissue type. For both tissues, mean NSC concentrations were higher for the Patagonia site than for the Mediterranean site. Mean root sapwood NSC concentration values were five times higher than those of the corresponding stem sapwood at all altitudes. Values for δ¹³C were positively correlated with altitude in the Mediterranean site only. Main conclusions We found support for the GLH at the site without drought effects (Patagonia) and no support for the CLH at either site. It is suggested that drought moderated the effects of low temperature by masking the expected trend of the GLH at the Mediterranean site.     

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。     

Non-structural carbon compounds in temperate forest trees

The current carbon supply status of temperate forest trees was assessed by analysing the seasonal variation of non‐structural carbohydrate (NSC) concentrations in leaves, branch wood and stem sapwood of 10 tree species (six deciduous broad‐leafed, one deciduous conifer and three evergreen conifer trees) in a temperate forest that is approximately 100 years old. In addition, all woody tissue was analysed for lipids (acylglycerols). The major NSC fractions were starch, sucrose, glucose and fructose, with other carbohydrates (e.g. raffinose and stachyose) and sugar alcohols (cyclitols and sorbitol) playing only a minor quantitative role. The radial distribution of NSC within entire stem cores, assessed here for the first time in a direct interspecific comparison, revealed large differences in the size of the active sapwood fraction among the species, reflecting the specific wood anatomy (ring‐porous versus diffuse‐porous xylem). The mean minimum NSC concentrations in branch wood during the growing season was 55% of maximum, and even high NSC concentrations were maintained during times of extensive fruit production in masting Fagus sylvestris. The NSC in stem sapwood varied very little throughout the season (cross species mean never below 67% of maximum), and the small reductions observed were not significant for any of the investigated species. Although some species contained substantial quantities of lipids in woody tissues (‘fat trees’; Tilia, Pinus, Picea, Larix), the lipid pools did not vary significantly across the growing season in any species. On average, the carbon stores of deciduous trees would permit to replace the whole leave canopy four times. These data imply that there is not a lot of leeway for a further stimulation of growth by ongoing atmospheric CO₂ enrichment. The classical view that deciduous trees rely more on C‐reserves than evergreen trees, seems unwarranted or has lost its justification due to the greater than 30% increase in atmospheric CO₂ concentrations over the last 150 years.     

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.     

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。     

Similar variation in carbon storage between deciduous and evergreen treeline species across elevational gradients

Background and Aims

The most plausible explanation for treeline formation so far is provided by the growth limitation hypothesis (GLH), which proposes that carbon sinks are more restricted by low temperatures than by carbon sources. Evidence supporting the GLH has been strong in evergreen, but less and weaker in deciduous treeline species. Here a test is made of the GLH in deciduous–evergreen mixed species forests across elevational gradients, with the hypothesis that deciduous treeline species show a different carbon storage trend from that shown by evergreen species across elevations.

Methods

Tree growth and concentrations of non-structural carbohydrates (NSCs) in foliage, branch sapwood and stem sapwood tissues were measured at four elevations in six deciduous–evergreen treeline ecotones (including treeline) in the southern Andes of Chile (40°S, Nothofagus pumilio and Nothofagus betuloides; 46°S, Nothofagus pumilio and Pinus sylvestris) and in the Swiss Alps (46°N, Larix decidua and Pinus cembra).

Key Results

Tree growth (basal area increment) decreased with elevation for all species. Regardless of foliar habit, NSCs did not deplete across elevations, indicating no shortage of carbon storage in any of the investigated tissues. Rather, NSCs increased significantly with elevation in leaves (P < 0·001) and branch sapwood (P = 0·012) tissues. Deciduous species showed significantly higher NSCs than evergreens for all tissues; on average, the former had 11 % (leaves), 158 % (branch) and 103 % (sapwood) significantly (P < 0·001) higher NSCs than the latter. Finally, deciduous species had higher NSC (particularly starch) increases with elevation than evergreens for stem sapwood, but the opposite was true for leaves and branch sapwood.

Conclusions

Considering the observed decrease in tree growth and increase in NSCs with elevation, it is concluded that both deciduous and evergreen treeline species are sink limited when faced with decreasing temperatures. Despite the overall higher requirements of deciduous tree species for carbon storage, no indication was found of carbon limitation in deciduous species in the alpine treeline ecotone.     

Carbon dynamics in the deciduous broadleaf tree Erman's birch (Betula ermanii) at the subalpine treeline on Changbai Mountain,Northeast China

Premise of the Study

The growth limitation hypothesis (GLH) and carbon limitation hypothesis (CLH) are two dominant explanations for treeline formation. The GLH proposes that low temperature drives the treeline through constraining C sinks more than C sources, and it predicts that non‐structural carbohydrate (NSC) levels are static or increase with elevation. Although the GLH has received strong support globally for evergreen treelines, there is still no consensus for deciduous treelines, which experience great asynchrony between supply and demand throughout the year.

Methods

We investigated growth and the growing‐season C dynamics in a common deciduous species, Erman's birch (Betula ermanii), along an elevational gradient from the closed forest to the treeline on Changbai Mountain, Northeast China. Samples were collected from developing organs (leaves and twigs) and main storage organs (stems and roots) for NSC analysis.

Key Results

Tree growth decreased with increasing elevation, and NSC concentrations differed significantly among elevations, organs, and sampling times. In particular, NSC levels varied slightly during the growing season in leaves, peaked in the middle of the growing season in twigs and stems, and increased continuously throughout the growing season in roots. NSCs also tended to increase or vary slightly in developing organs but decreased significantly in mature organs with increasing elevation.

Conclusions

The decrease in NSCs with elevation in main storage organs indicates support for the CLH, while the increasing or static trends in new developing organs indicate support for the GLH. Our results suggest that the growth limitation theory may be less applicable to deciduous species' growth than to that of evergreen species.     

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。     

Insights into intraspecific wood density variation and its relationship to growth,height and elevation in a treeline species

• The wood economics spectrum provides a general framework for interspecific trait–trait coordination across wide environmental gradients. Whether global patterns are mirrored within species constitutes a poorly explored subject. In this study, I first determined whether wood density co‐varies together with elevation, tree growth and height at the within‐species level. Second, I determined the variation of wood density in different stem parts (trunk, branch and twigs).
• In situ trunk sapwood, trunk heartwood, branch and twig densities, in addition to stem growth rates and tree height were determined in adult trees of Nothofagus pumilio at four elevations in five locations spanning 18° of latitude. Mixed effects models were fitted to test relationships among variables.
• The variation in wood density reported in this study was narrow (ca. 0.4–0.6 g cm^?3) relative to global density variation (ca. 0.3–1.0 g cm^?3). There was no significant relationship between stem growth rates and wood density. Furthermore, the elevation gradient did not alter the wood density of any stem part. Trunk sapwood density was negatively related to tree height. Twig density was higher than branch and trunk densities. Trunk heartwood density was always significantly higher than sapwood density.
• Negative across‐species trends found in the growth–wood density relationship may not emerge as the aggregate of parallel intraspecific patterns. Actually, trees with contrasting growth rates show similar wood density values. Tree height, which is tightly related to elevation, showed a negative relationship with sapwood density.

     

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.     

GIS‐analysis of tree‐line elevation in the Swiss Alps suggests no exposure effect

Abstract. Counter intuition, an analysis of tree‐line position across the Swiss Alps based on a geographical information system (GIS) with a spatial resolution of 100 m (2.5 million points) revealed no difference in climatic tree‐line altitude with slope exposure. Through step wise discrimination procedures our analysis accounts for anthropogenic tree‐line depression. Any land cover bias affects the frequency of GIS‐points corresponding to tree‐line forests rather than the mean elevation of such points, captured by our analysis. We explain this phenomenon (1) by the absence of significant drought effects in the Alps (no disadvantages for southwest slopes), (2) by the fact that tree tops, unlike low stature vegetation, do not profit from greater radiation warming on south slopes during the growing season but are thermally coupled to free air circulation, and (3) by preliminary data for root zone temperatures during the growing season, which do not differ between south and north slopes, as long as the soil is screened by a closed forest canopy. The overall difference in season length and snow cover, often seen between south and north slopes, does not seem to affect tree‐line position but explains greater natural forest fragmentation on north slopes. It is this greater fragmentation and patchiness (avalanche tracks, snow beds etc.) which seem to have nourished the idea of a generally lower limit of tree growth and tree lines at northern slopes. These results are in line with a recently developed theory, which suggests that tree‐line elevations in humid climates correspond to similar isotherms, irrespective of latitude and thus, season length.     

No evidence for depletion of carbohydrate pools in Scots pine (Pinus sylvestris L.) under drought stress

The physiological mechanisms leading to Scots pine (Pinus sylvestris L.) decline in the dry inner alpine valleys are still unknown. Testing the carbon starvation hypothesis, we analysed the seasonal course of mobile carbohydrate pools (NSC) of Scots pine growing at a xeric and a dry-mesic site within an inner alpine dry valley (750 m a.s.l., Tyrol, Austria) during 2009, which was characterised by exceptional soil dryness. Although, soil moisture content dropped to ca. 10% at both sites during the growing season, NSC concentrations rose in all tissues (branch, stem, root) until the end of July, except in needles, where maxima were reached around bud break. NSC concentrations were not significantly different in the analysed tissues at the xeric and the dry-mesic site. At the dry-mesic site, NSC concentrations in the aboveground tree biomass were significantly higher during the period of radial growth. An accumulation of NSC in roots at the end of July indicates a change in carbon allocation after an early cessation in aboveground growth, possibly due to elevated belowground carbon demand. In conclusion, our results revealed that extensive soil dryness during the growing season did not lead to carbon depletion. However, even though carbon reserves were not exhausted, sequestration of carbohydrate pools during drought periods might lead to deficits in carbon supply that weaken tree vigour and drive tree mortality.     

Uneven winter snow influence on tree growth across temperate China

Winter snow is an important driver of tree growth in regions where growing‐season precipitation is limited. However, observational evidence of this influence at larger spatial scales and across diverse bioclimatic regions is lacking. Here, we investigated the interannual effects of winter (here defined as previous October to current February) snow depth on tree growth across temperate China over the period of 1961–2015, using a regional network of tree ring records, in situ daily snow depth observations, and gridded climate data. We report uneven effects of winter snow depth on subsequent growing‐season tree growth across temperate China. There shows little effect on tree growth in drier regions that we attribute mainly to limited snow accumulation during winter. By contrast, winter snow exerts important positive influence on tree growth in stands with high winter snow accumulation (e.g., in parts of cold arid regions). The magnitude of this effect depends on the proportion of winter snow to pre‐growing‐season (previous October to current April) precipitation. We further observed that tree growth in drier regions tends to be increasingly limited by warmer growing‐season temperature and early growing‐season water availability. No compensatory effect of winter snow on the intensifying drought limitation of tree growth was observed across temperate China. Our findings point toward an increase in drought vulnerability of temperate forests in a warming climate.     

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.     

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。     

Wood vessel diameter is related to elevation and genotype in the Hawaiian tree Metrosideros polymorpha (Myrtaceae)

We tested the hypothesis that trees growing at high elevations with occasional freezing temperatures have smaller diameter xylem vessels than trees of the same species growing at lower and warmer elevations. The young branch wood of the wide-ranging Hawaiian tree species Metrosideros polymorpha (Myrtaceae) was examined in three natural field populations (high, middle, and low elevations: 2469, 1280, and 107 m a.s.l., respectively) and contrasted with seedlings from these populations that were grown in a common garden at middle elevation (1190 m). Previous studies showed that these populations have some genetic differences and have distinctive leaf structure and ecophysiological traits. Vessel diameter was significantly smaller in the high elevation field and common garden plants than in middle elevation plants. However, high elevation vessels were wider in common garden plants compared to field plants, indicating that vessel diameter is determined both by genotype (parental populations) and environment (growing conditions different from those of parents). Reduced vessel diameter has implications for resistance to cavitation induced by freezing and/or drought in plants growing near tree line in Hawaii.     

温带不同材性树种树干非结构性碳水化合物的径向分配差异

温带森林不同树种具有不同的非结构性碳水化合物(NSC)存储和利用策略, 树干是成年树木NSC主体储存库。但树干NSC径向变异和种间差异仍不清楚, 无孔材(裸子植物)、散孔材和环孔材(被子植物)所代表的木材孔性功能群对树干NSC浓度的影响尚缺乏定论。为探索温带森林主要树种树干NSC浓度随树木木材孔性和组织的变化特征, 该研究在黑龙江省穆棱市的东北典型阔叶红松(Pinus koraiensis)林中选择32个树种, 采集胸高位置树皮、边材和心材3种组织, 分析NSC浓度随木材孔性和组织的变化特征。结果表明: (1)树种、组织和木材孔性均显著影响树干的NSC浓度。3种组织可溶性糖、淀粉、总NSC浓度和糖/淀粉的种间变异较大, 变异系数最低为37% (树皮总NSC浓度), 最高达到101% (心材淀粉浓度), 树干组织、树种及其交互作用均显著影响NSC浓度。(2)总体上可溶性糖、淀粉和总NSC浓度均随径向深度增加而降低。无孔材树皮的可溶性糖浓度和糖/淀粉显著高于散孔材和环孔材, 而边材中的淀粉和总NSC浓度为环孔材>散孔材>无孔材。(3)无孔材可溶性糖、淀粉和总NSC浓度边材和心材比均在1左右, 显著低于散孔材和环孔材, 而且无孔材边材和心材之间淀粉浓度相关较紧密, 表明被子植物的边材、心材功能分化较裸子植物更为明显。研究结果表明木材孔性影响了温带树种树干NSC存储策略, 研究整树NSC以及树木生理生态学功能需要区分树干组织。     

Elevation effects on the carbon budget of tropical mountain forests (S Ecuador): the role of the belowground compartment

Carbon storage and sequestration in tropical mountain forests and their dependence on elevation and temperature are not well understood. In an altitudinal transect study in the South Ecuadorian Andes, we tested the hypotheses that (i) aboveground net primary production (ANPP) decreases continuously with elevation due to decreasing temperatures, whereas (ii) belowground productivity (BNPP) remains constant or even increases with elevation due to a shift from light to nutrient limitation of tree growth. In five tropical mountain forests between 1050 and 3060 m a.s.l., we investigated all major above‐ and belowground biomass and productivity components, and the stocks of soil organic carbon (SOC). Leaf biomass, stemwood mass and total aboveground biomass (AGB) decreased by 50% to 70%, ANPP by about 70% between 1050 and 3060 m, while stem wood production decreased 20‐fold. Coarse and large root biomass increased slightly, fine root biomass fourfold, while fine root production (minirhizotron study) roughly doubled between 1050 and 3060 m. The total tree biomass (above‐ and belowground) decreased from about 320 to 175 Mg dry mass ha^?1, total NPP from ca. 13.0 to 8.2 Mg ha^?1 yr^?1. The belowground/aboveground ratio of biomass and productivity increased with elevation indicating a shift from light to nutrient limitation of tree growth. We propose that, with increasing elevation, an increasing nitrogen limitation combined with decreasing temperatures causes a large reduction in stand leaf area resulting in a substantial reduction of canopy carbon gain toward the alpine tree line. We conclude that the marked decrease in tree height, AGB and ANPP with elevation in these mountain forests is caused by both a belowground shift of C allocation and a reduction in C source strength, while a temperature‐induced reduction in C sink strength (lowered meristematic activity) seems to be of secondary importance.