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.     

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。     

Formation mechanisms of the alpine Erman’s birch (Betula ermanii) treeline on Changbai Mountain in Northeast China

Key message

The treeline on Changbai Mountain controlled by low temperature and water stress, has not reached the position most commonly expected.

Abstract

Treeline pattern is an important consideration in exploring the general mechanisms controlling the response of treelines to climatic change. However, most of the present conclusions were derived from evergreen and/or conifer treeline, it is still not clear about the deciduous treeline. This study analyzed concentrations of non-structural carbohydrates (NSC) and their components (total soluble sugars and starch) in tree tissues of the deciduous species Erman’s birch (Betula ermanii) at four points along an elevational gradient ranging from 1,908–2,058 m a.s.l at the end of the growing season on Changbai Mountain in Northeast China. The mean 10-cm soil temperature of 8.2 °C under trees across the 129-day growing season at the treeline in this region was higher than that of the average threshold temperature found at treeline positions in the global and China’s climate studies. However, altitudinal trends of NSC concentrations increased significantly in all tissue types along the altitudinal gradients, revealing no depletion of carbon reserves at the treeline on Changbai Mountain. At the same time, the pronounced variation of δ¹³C in leaves and aged branches suggested that low temperature and water stress may simultaneously be operating at high altitudes to restrict the growth and NSC accumulation in trees above the treeline. In light of the above, we conclude that treeline formation on Changbai Mountain is no carbon depletion at the end of growing season, and most likely the result of sink limitation reflecting the combined effects of low temperature and water stress that determined the actual position of the treeline.     

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.     

Carbon sink limitation and frost tolerance control performance of the tree <Emphasis Type="Italic">Kageneckia angustifolia</Emphasis> D. Don (Rosaceae) at the treeline in central Chile

The decrease in temperature with increasing elevation may determine the altitudinal tree distribution in different ways: affecting survival through freezing temperatures, by a negative carbon balance produced by lower photosynthetic rates, or by limiting growth activity. Here we assessed the relative importance of these direct and indirect effects of altitudinal decrease in temperature in determining the treeline in central Chile (33°S) dominated by Kageneckia angustifolia. We selected two altitudes (2000 and 2200 m a.s.l.) along the treeline ecotone. At each elevation, leaf non-structural carbohydrates (NSC) and gas exchange parameters were measured on five individuals during the growing season. We also determined the cold resistance of K.␣angustifolia, by measuring temperatures that cause 50% seedling mortality (LT₅₀) and ice nucleation (IN). No differences in net photosynthesis were found between altitudes. Although no differences were detected on NSC concentration on a dry matter basis between 2000 and 2200 m, when NSC concentration was expressed on a leaf area basis, higher contents were found at the higher elevation. Thus, carbon sink limitations may occur at the K. angustifolia’s upper altitudinal limit. For seedlings derived from seeds collected at the 2200 m, LT₅₀ of cold-acclimated and non-acclimated plants were −9.5 and −7 °C, respectively. However, temperatures as low as −10 °C can frequently occur at this altitude during the end of winter. Therefore, low temperature injury of seedlings seems also be involved in the treeline formation in this species. Hence, a confluence of global (carbon sink limitation) and regional (freezing tolerance) mechanisms explains the treeline formation in the Mediterranean-type climate zone of central Chile.     

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.     

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.     

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.     

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。     

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 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.     

Global patterns of mobile carbon stores in trees at the high‐elevation tree line

Aim Across all latitudes, high‐elevation tree lines represent a drastic change in the dominant plant life‐form, from upright trees to low‐stature alpine plants. Although associated with low temperatures, the physiological mechanisms controlling this boundary are still not clear. The growth‐limitation hypothesis assumes a direct low‐temperature restriction of tissue formation at otherwise sufficient photoassimilation. In order to test this hypothesis, we present a global synthesis of previously published and new data on tree carbon supply status at high‐elevation tree lines. Location Global; 13 regions between 68° N and 45° S. Methods Late‐season concentrations of non‐structural carbohydrates (NSC) in foliage and branch wood were measured at three elevations across the tree line ecotones, from upper tall forests (timber line) to the edges of aborescent tree growth (tree line). Year‐round records of ?10 cm soil temperatures were taken at the tree line. Results Despite large differences in elevation and season length, the mean growing season temperature at the tree line was similar (approximately 6.6 °C) between all sites. NSC concentrations were not depleted at any of the elevation gradients between timber line and tree line, indicating no shortage of C at the uppermost tree stands. Tested across all sites, NSC concentrations in fact significantly increased with elevation. On average, NSC increased by 18% in leaves and 26% in branch wood from the lowest to the highest stands, primarily due to higher starch concentrations rather than free sugars. Hence, these responses do not reflect osmotic adjustments to lower temperatures at the end of the growing season. Main conclusions This global data set contributes to a mechanistic understanding of tree line formation based on biological principles across climatic zones and tree genera. No evidence of C shortage was found at the high‐elevation tree line in either seasonal and non‐seasonal regions. The increasing trend of NSC concentrations with elevation is in line with the growth‐limitation hypothesis.     

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.     

Asymmetric changes of growth and reproductive investment herald altitudinal and latitudinal range shifts of two woody species

Ongoing changes in global climate are altering ecological conditions for many species. The consequences of such changes are typically most evident at the edge of the geographical distribution of a species, where range expansions or contractions may occur. Current demographical status at geographical range limits can help us to predict population trends and their implications for the future distribution of the species. Thus, understanding the comparability of demographical patterns occurring along both altitudinal and latitudinal gradients would be highly informative. In this study, we analyse the differences in the demography of two woody species through altitudinal gradients at their southernmost distribution limit and the consistency of demographical patterns at the treeline across a latitudinal gradient covering the complete distribution range. We focus on Pinus sylvestris and Juniperus communis, assessing their demographical structure (density, age and mortality rate), growth, reproduction investment and damage from herbivory on 53 populations covering the upper, central and lower altitudes as well as the treeline at central latitude and northernmost and southernmost latitudinal distribution limits. For both species, populations at the lowermost altitude presented older age structure, higher mortality, decreased growth and lower reproduction when compared to the upper limit, indicating higher fitness at the treeline. This trend at the treeline was generally maintained through the latitudinal gradient, but with a decreased growth at the northern edge for both species and lower reproduction for P. sylvestris. However, altitudinal and latitudinal transects are not directly comparable as factors other than climate, including herbivore pressure or human management, must be taken into account if we are to understand how to infer latitudinal processes from altitudinal data.     

Intrinsic morphology and spatial distribution of non-structural carbohydrates contribute to drought resistance of two mulberry cultivars

• Drought is one of the most adverse environmental stresses limiting plant growth and productivity. However, the underlying mechanisms regarding metabolism of non-structural carbohydrates (NSC) in source and sink organs are still not fully elucidated in woody trees.
• Saplings of mulberry cv Zhongshen1 and Wubu were subjected to a 15-day progressive drought stress. NSC levels and gene expression involved in NSC metabolism were investigated in roots and leaves. Growth performance and photosynthesis, leaf stomatal morphology, and other physiological parameters were also analysed.
• Under well-watered conditions, Wubu had a higher R/S, with higher NSC in leaves than in roots; Zhongshen1 had a lower R/S with higher NSC in roots than leaves. Under drought stress, Zhongshen1 showed decreased productivity and increased proline, abscisic acid, ROS content and activity of antioxidant enzymes, while Wubu sustained comparable productivity and photosynthesis. Interestingly, drought resulted in decreased starch and slightly increased soluble sugars in leaves of Wubu, accompanied by notable downregulation of starch-synthesizing genes and upregulation of starch-degrading genes. Similar patterns in NSC levels and relevant gene expression were also observed in roots of Zhongshen1. Concurrently, soluble sugars decreased and starch was unchanged in roots of Wubu and leaves of Zhongshen1. However, gene expression of starch metabolism in roots of Wubu was unaltered, but in leaves of Zhongshen1 starch metabolism was more activated.
• These findings revealed that intrinsic R/S and spatial distribution of NSC in roots and leaves concomitantly contribute to drought resistance in mulberry.

     

贡嘎山地区不同海拔冷杉比叶质量和非结构性碳水化合物含量变化

以贡嘎山地区磨西山谷东北坡和康定山谷西南坡为样地,研究了高山林线和低海拔冷杉休眠期和生长旺盛期针叶比叶质量和组织非结构性碳水化合物含量.结果表明:生长在温暖湿润的磨西山谷东北坡的冷杉针叶比叶质量和组织非结构性碳水化合物含量均高于炎热干燥的康定山谷西南坡;高山林线冷杉比叶质量大于低海拔冷杉;高山林线冷杉组织非结构性碳水化合物含量总体高于低海拔冷杉,且生长期比休眠期更为显著.研究结果不支持"碳供应不足导致高山林线形成"的假说.     

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_other显著正相关,说明叶片P_other和NSC含量的累积共同用于提高大叶醉鱼草在高海拔恶劣环境下的适应性。     

Carbon allocation in shoots of alpine treeline conifers in a CO<Subscript>2</Subscript> enriched environment

With a new approach we assessed the relative contribution of stored and current carbon compounds to new shoot growth in alpine treeline conifers. Within a free air CO₂ enrichment experiment at the alpine treeline in Switzerland, ¹³C-depleted fossil CO₂ was used to trace new carbon in the two tree species Larix decidua L. and Pinus uncinata Ramond over two subsequent years. The deciduous L. decidua was found to supply new shoot growth (structural woody part) by 46% from storage. Surprisingly, the evergreen P. uncinata, assumed to use current-year photosynthates, also utilized a considerable fraction of storage (42%) for new wood growth. In contrast, the needles of P. uncinata were built up almost completely from current-year photosynthates. The isotopic composition of different wood carbon fractions revealed a similar relative allocation of current and stored assimilates to various carbon fractions. Elevated CO₂ influenced the composition of woody tissue in a species-specific way, e.g. the water soluble fraction decreased in pine in 2001 but increased in larch in 2002 compared to ambient CO₂. Heavy defoliation applied as an additional treatment factor in the second year of the experiment decreased the lipophilic fraction in current-year wood in both species compared to undefoliated trees. We conclude that storage may play an important role for new shoot growth in these treeline conifers and that altered carbon availability (elevated CO₂, defoliation) results in significant changes in the relative amount of mobile carbon fractions in woody tissue. In particular, stored carbon seems to be of greater importance in the evergreen P. uncinata than has been previously thought.     

Bicarbonate stimulates non‐structural carbohydrate pools of Camptotheca acuminata

The role of root‐derived dissolved inorganic carbon (DIC) has been emphasized lately, as it can provide an alternative source of carbon for photosynthesis. The fate of newly fixed DIC and its effect on non‐structural carbohydrate (NSC) pools has not been thoroughly elucidated to date. To this end, we used ¹³C (NaHCO₃) as a substrate tracer to investigate the incorporation of newly fixed bicarbonate into the plant organs and NSC compounds of Camptotheca acuminata seedlings for 24 and 72 h. NSC levels across the organs were all markedly increased within 24 h of labeling treatment and afterward only decreased in stems at 72 h. The variation range of NSC concentrations in roots was considerably smaller than in the stem and leaves. As time passed, the δ¹³C in NSC compounds was significantly affected by ¹³C labeling and was more positive in the roots than in the stem and leaves. Starch was more ¹³C‐enriched than was soluble carbohydrate, and the δ¹³C of root starch was as high as ?4.70‰. Bicarbonate incorporation into newly formed NSC compounds contributed up to 0.24% of the root starch within 72 h. These data provided strong evidence that bicarbonate not only acted as a C source that contributed slightly to the NSC pools but also stimulated the increase in NSC pools. The present study expands our understanding of the rapid change of NSC pools across the organs in response to bicarbonate.