Rapid mixing between old and new C pools in the canopy of mature forest trees

Stable C isotope signals in plant tissues became a key tool in explaining growth responses to the environment. The technique is based on the fundamental assumption that the isotopic composition of a given unit of tissue (e.g. a tree ring) reflects the specific C uptake conditions in the leaf at a given time. Beyond the methodological implications of any deviation from this assumption, it is of physiological interest whether new C is transferred directly from sources (a photosynthesizing leaf) to structural sinks (e.g. adjacent stem tissue), or inherently passes through existing (mobile) C pools, which may be of variable (older) age. Here, we explore the fate of (13)C-labelled photosynthates in the crowns of a 30-35 m tall, mixed forest using a canopy crane. In all nine study species labelled C reached woody tissue within 2-9 h after labelling. Four months later, very small signals were left in branch wood of Tilia suggesting that low mixing of new, labelled C with old C had taken place. In contrast, signals in Fagus and Quercus had increased, indicating more intense mixing. This species-specific mixing of new with old C pools is likely to mask year- or season-specific linkages between tree ring formation and climate and has considerable implications for climate reconstruction using stable isotopes as proxies for past climatic conditions.     

Fruit-bearing branchlets are carbon autonomous in mature broad-leaved temperate forest trees

In order to evaluate the degree of carbon autonomy for fruit development, the carbon source-sink relationship in fruit-bearing branchlets of mature deciduous forest trees was manipulated in situ. The tests included half and complete defoliation, girdling or the combination of both treatments, which were applied on fruiting branchlets by using a canopy crane. Concentrations of non-structural carbohydrates (NSC) were analysed in different branchlet tissues and fruits, to identify situations of carbon imbalances induced by the treatments. NSC concentrations of branchlets were generally lower under treatments resulting in decreased fruit growth. All three investigated species (Carpinus betulus, Fagus sylvatica and Tilia platyphyllos) exhibited complete carbon autonomy of fruiting at the level of whole, undisturbed branchlets, since neither a decrease of total infructescence biomass, nor of individual fruit mass occurred on girdled, un-defoliated branchlets. On girdled, 100% defoliated branchlets, fruit biomass relative to controls decreased by approximately 50% in Carpinus and Tilia, but by almost 80% in Fagus, which can be explained by different proportions of photosynthetically active infructescence tissues among the species. In contrast to the other two species, Tilia branchlets did not import carbon to compensate for assimilate loss after defoliation.     

Drought and shade deplete nonstructural carbohydrate reserves in seedlings of five temperate tree species

Plants that store nonstructural carbohydrates (NSC) may rely on carbon reserves to survive carbon‐limiting stress, assuming that reserves can be mobilized. We asked whether carbon reserves decrease in resource stressed seedlings, and if NSC allocation is related to species' relative stress tolerances. We tested the effects of stress (shade, drought, and defoliation) on NSC in seedlings of five temperate tree species (Acer rubrum Marsh., Betula papyrifera Marsh., Fraxinus americana L., Quercus rubra L., and Quercus velutina Lam.). In a greenhouse experiment, seedlings were subjected to combinations of shade, drought, and defoliation. We harvested seedlings over 32–97 days and measured biomass and NSC concentrations in stems and roots to estimate depletion rates. For all species and treatments, except for defoliation, seedling growth and NSC accumulation ceased. Shade and drought combined caused total NSC decreases in all species. For shade or drought alone, only some species experienced decreases. Starch followed similar patterns as total NSC, but soluble sugars increased under drought for drought‐tolerant species. These results provide evidence that species deplete stored carbon in response to carbon limiting stress and that species differences in NSC response may be important for understanding carbon depletion as a buffer against shade‐ and drought‐induced mortality.     

Leaf phenology and carbon dynamics in six leguminous trees

I documented photosynthetic rates and seasonal stem total nonstructural carbohydrates (TNC) in six leguminous tree species Burkea africana, Baikiaea plurijuga, Erythrophleum africanum, Guibourtia coleosperma, Julbernardia globiflora and Pterocarpus angolensis exhibiting a range of leaf phenological patterns. My goal was to (i) measure photosynthetic characteristics and levels of stored stem carbohydrates in species with varying patterns of leaf phenology and (ii) determine seasonal patterns of stem carbohydrate storage. Despite significant differences in the timing of bud break and leaf cover between the six species, there were no significant differences in maximum photosynthetic rate, quantum efficiency or light saturation point between species. Similarly, there was no significant difference in seasonal mean stem TNC levels despite significant differences in the timing of bud break and leaf cover both between species and within a single species. However, while the average amount of TNC does not seem to be related to leaf phenology, the patterns of carbohydrate use and storage do seem to be related to leaf phenology.     

Winter photosynthesis by saplings of evergreen broad-leaved trees in a deciduous temperate forest

* Here we investigated photosynthetic traits of evergreen species under a deciduous canopy in a temperate forest and revealed the importance of CO2 assimilation during winter for annual CO2 assimilation. * Saplings were shaded by the canopy trees from spring through to autumn, but were less shaded during the winter months. Photosynthetic rates at light saturation (Aarea) were lower during winter than during the growing season. Aarea was higher in Camellia, Ilex and Photinia than in Castanopsis, Cleyera and Quercus during the winter, but differed little during summer and autumn. * Estimated daily CO2 assimilation (Aday) was higher during the winter than during the growing season in Camellia, Ilex and Photinia but was higher than that during the growing season only at the beginning and end of winter in Castanopsis, Cleyera and Quercus. Aday was higher in Camellia, Ilex and Photinia than in Castanopsis, Cleyera and Quercus but differed little among them during the growing season. * These results reveal the importance of winter CO2 assimilation for the growth of Camellia, Ilex and Photinia. Furthermore, differences in annual CO2 assimilation among species are strongly modified by species-specific photosynthetic traits during the winter under deciduous canopy trees.     

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.     

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.     

广西光皮桦林的分类和演替

光皮桦林是常绿阔叶林或山地常绿落叶阔叶混交林遭受砍伐后,在保护较好的情况下自然恢复起来的一种演替系列群落,属亚热带落叶阔叶林的一种类型。本文通过群落分类研究,划分出３个群丛纲和１１个群丛。从这些群丛的外貌、结构和种类成分的变化,可确定其演替的趋向和所处阶段,是合理经营管理它的科学依据。     

中国森林凋落物不同组分异速比例关系

森林凋落物(litterfall)是森林植物在其生长发育过程中新陈代谢的产物, 在物质循环和能量流动方面起着重要作用。该文利用已发表的我国主要森林凋落物的研究数据, 分析了不同组分(叶、枝和繁殖器官)凋落物量之间及其与总凋落物量之间的异速比例关系。结果表明: 我国森林叶、枝和繁殖器官的平均凋落物量分别为3 810.34、1 019.07和767.95 kg·hm^-2·a^-1; 温度、降水量、林龄对森林凋落物量均有一定程度的影响, 其中温度对各组分凋落物量的影响最大。叶凋落物量(L_L)与总凋落物量(L_T)之间呈等速生长关系(L_L ∝ L_T^0.96), 繁殖器官和枝的凋落物量(分别为L_P和L_B)与L_T之间呈异速比例关系, 分别为L_P ∝ L_T^1.84和L_B ∝ L_T^1.61。不同组分凋落物量之间具有显著的异速比例关系, 其异速指数均小于1.0。不同林型(常绿林和落叶林)各组分凋落物量之间的异速比例关系无显著差异。了解不同组分凋落物量与总凋落物量之间的异速比例关系可以为更加精确地估算森林生产力提供理论依据。     

Effect of nitrate on acetylene reduction activity and carbohydrate composition of Phaseolus vulgaris nodules

When Phaseolus vulgaris L. cv. Kentucky Wonder plants were supplied with various levels of nitrate for 34 days, nodule weight (plant)⁻¹, acetylene reduction activity (g nodule)⁻¹, and sugar concentration in nodules were depressed >60% (7.5 m M nitrate vs nil nitrate). Starch concentration in nodules was more than double the sugar concentration and declined only slightly in response to nitrate level. At the highest level of nitrate, sugar concentration in nodules was 50% greater than that in roots and nodule starch was about 6-fold greater than root starch on a fresh weight basis. When plants were grown with 1 m M nitrate and then supplied with 12 m M nitrate for 7 days, the rapid decline in acetylene reduction activity coincided with a decline in sucrose concentration. However, glucose and fructose concentrations declined only after the largest decrease in acetylene reduction had occurred, and the quantitative decrease in glucose and fructose in nodules was small relative to sucrose. Other results showed that the magnitude of the effect of nitrate on some nodule carbohydrate compounds depends on Rhizobium phaseoli strain and on whether plants were grown with or without nitrate prior to experimental treatments. Some of the results are consistent with the carbohydrate-deprivation hypothesis for inhibition of legume nodules by nitrate. However, there are several complications involved in the interpretation of results of this type, and other possible explanations for the results are suggested.     

Characteristics of free air carbon dioxide enrichment of a northern temperate mature forest

In 2017, the Birmingham Institute of Forest Research (BIFoR) began to conduct Free Air Carbon Dioxide Enrichment (FACE) within a mature broadleaf deciduous forest situated in the United Kingdom. BIFoR FACE employs large‐scale infrastructure, in the form of lattice towers, forming ‘arrays’ which encircle a forest plot of ~30 m diameter. BIFoR FACE consists of three treatment arrays to elevate local CO₂ concentrations (e[CO₂]) by +150 µmol/mol. In practice, acceptable operational enrichment (ambient [CO₂] + e[CO₂]) is ±20% of the set point 1‐min average target. There are a further three arrays that replicate the infrastructure and deliver ambient air as paired controls for the treatment arrays. For the first growing season with e[CO₂] (April to November 2017), [CO₂] measurements in treatment and control arrays show that the target concentration was successfully delivered, that is: +147 ± 21 µmol/mol (mean ± SD) or 98 ± 14% of set point enrichment target. e[CO₂] treatment was accomplished for 97.7% of the scheduled operation time, with the remaining time lost due to engineering faults (0.6% of the time), CO₂ supply issues (0.6%) or adverse weather conditions (1.1%). CO₂ demand in the facility was driven predominantly by wind speed and the formation of the deciduous canopy. Deviations greater than 10% from the ambient baseline CO₂ occurred <1% of the time in control arrays. Incidences of cross‐contamination >80 µmol/mol (i.e. >53% of the treatment increment) into control arrays accounted for <0.1% of the enrichment period. The median [CO₂] values in reconstructed three‐dimensional [CO₂] fields show enrichment somewhat lower than the target but still well above ambient. The data presented here provide confidence in the facility setup and can be used to guide future next‐generation forest FACE facilities built into tall and complex forest stands.     

峨眉山常绿落叶阔叶混交林的生物多样性及植物区系初探

对峨眉山常绿落叶阔叶混交林的群落结构﹑组成﹑生物多样性及植物区系等几方面进行了研究分析。结果显示峨眉山阔叶混交林由226种维管束植物组成,其中被子植物73科137属207种, 占总种数的91.6%,是峨眉山阔叶混交林的重要组成部分;群落分层现象明显,为乔木层﹑灌木层﹑草本层和层外植物4层,其中草本层发育情况差;科分布型是以热带—亚热带﹑热带—温带为主,各占22.7%, 在属的水平上则以温带分布占绝对优势(52.9%),揭示了峨眉山阔叶混交林的区系性质是以温带为主的亚热带类型;生物多样性指数处于较低的水平,在经过人为干扰后,群落处于稳定的恢复阶段。     

Weather and climate controls over the seasonal carbon isotope dynamics of sugars from subalpine forest trees

We examined the environmental variables that influence the δ ¹³C value of needle and phloem sugars in trees in a subalpine forest. We collected sugars from Pinus contorta , Picea engelmannii and Abies lasiocarpa from 2006 to 2008. Phloem and needle sugars were enriched in ¹³C during the autumn, winter and early spring, but depleted during the growing season. We hypothesized that the late-winter and early-spring ¹³C enrichment was due to the mobilization of carbon assimilated the previous autumn; however, needle starch concentrations were completely exhausted by autumn, and we observed evidence of new starch production during episodic warm weather events during the winter and early-spring. Instead, we found that ¹³C enrichment was best explained by the occurrence of cold night-time temperatures. We also observed seasonal decoupling in the ¹³C/¹²C ratios of needle and phloem sugars. We hypothesized that this was due to seasonally-changing source-sink patterns, which drove carbon translocation from the needles towards the roots early in the season, before bud break, but from the roots towards the needles later in the season, after bud break. Overall, our results demonstrate that the ¹³C/¹²C ratio of recently-assimilated sugars can provide a sensitive record of the short-term coupling between climate and tree physiology.     

油松幼苗非结构性碳水化合物对干旱胁迫的阶段性响应

为探讨干旱胁迫对油松非结构性碳水化合物（NSC）的影响及其响应机制,以2年生油松幼苗为对象,设置适宜水分、轻度、中度和重度干旱胁迫处理（即80%、60%、40%和20%田间持水量）,比较幼苗在不同胁迫时间（15、30、45和60 d）各器官NSC含量的变化规律。结果表明：不同干旱胁迫处理程度与时间对油松幼苗当年生叶、1年生叶、茎、粗根和细根的可溶性糖、淀粉及总NSC含量均具有显著交互作用。干旱胁迫15 d时,当年生叶NSC含量在干旱胁迫处理显著低于适宜水分处理,细根NSC含量在轻度和重度胁迫处理显著高于适宜水分处理。干旱胁迫45 d时,随着胁迫程度增加,当年生叶、1年生叶、茎、粗根和细根淀粉含量均呈增加趋势。干旱胁迫60 d时,随着胁迫程度增加,细根淀粉含量呈下降趋势,细根可溶性糖及NSC含量在胁迫处理显著低于适宜水分处理。综上所述,油松幼苗胁迫初期,NSC优先供给细根以促进吸收水分,而长期胁迫,将导致NSC向下运输受阻,根系生长和吸收功能下降。这种NSC对干旱胁迫阶段性响应策略的揭示为东北地区油松幼苗造林最佳时间选择及水分管理提供数据支持。     

Increased chilling tolerance and altered carbon metabolism in tomato leaves following application of mechanical stress

We investigated the effects of brushing on the chilling tolerance and metabolism of nonstructural carbohydrates (soluble sugars and starch) in tomato leaves before, during and after a chilling stress. Tomato plants ( Lycopersicon esculentum Mill. cv. Caruso) were cultivated either without mechanical stress application (control plants) or with daily brushing treatments for 15 days (brushed plants), prior to a 7-day chilling treatment (8/5°C day/night). Brushing resulted in shorter plants with a 34% reduction in leaf dry weight per area and a 59% reduction of soluble sugars and starch, on a dry weight basis. The sugar to starch ratio was not affected by brushing. A greater chilling tolerance in the brushed plants was demonstrated by the maintenance of a significantly higher PSII efficiency in brushed plants (42%) compared to that of the control plants (30%) after 7 days of chilling treatment, less visible damage to the leaf tissue, and a more rapid resumption of growth during 3 days of recovery as compared to control plants. During the chilling treatment levels of soluble sugars per leaf dry weight increased 15-fold in the brushed plants and 5-fold in control plants. In the present study we have demonstrated that brushing can increase chilling tolerance in tomato plants. The observed differences in chilling tolerance and concentration of soluble sugars in the leaves may indicate an involvement of soluble sugar levels in acclimation to chilling.     

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。     

基于地形因子的天童地区常绿树种和落叶树种共存机制研究

位于亚热带的浙江天童和古田山常绿阔叶林大样地分布有较高比例的落叶树种,那么它们与常绿树种的共存机制是什么?常绿树种和落叶树种生态习性差异较大,二者对生境的选择应有所不同,我们推测生境分化可能是两类植物实现共存的主要机制。为检验该假设,我们以天童20ha动态样地调查数据为依托,选择个体数≥20的55个常绿树种和42个落叶树种作为分析对象,用典范对应分析(CCA)研究了地形因子对二者分布的影响差异,用torus转换检验来分析常绿树种和落叶树种与各类地形生境的关联。结果如下:(1)CCA分析表明地形因子对常绿树种分布的解释量为19.2%,对落叶树种分布的解释量为7.0%。(2)torus转换检验结果表明,与沟谷成正关联的常绿树种和落叶树种的比例分别为16.4%和28.6%,成负关联的比例分别为40%和7%;与山脊成正关联的常绿树种和落叶树种的比例分别为41.8%和4.8%,成负关联的比例分别为10.9%和47.6%;与受干扰生境成正关联的常绿树种和落叶树种的比例分别为16.4%和42.9%。上述结果说明地形对常绿树种分布的影响大于落叶树种;两个植物类群对生境的选择多呈现相反格局,尤其是在沟谷生境和山脊生境,这进一步表明生境分化是常绿树种和落叶树种共存的重要机制之一,生态位理论在一定程度上能较好地解释亚热带常绿阔叶林物种多样性的维持。     

Ecological importance of large-diameter trees in a temperate mixed-conifer forest

Large-diameter trees dominate the structure, dynamics and function of many temperate and tropical forests. Although both scaling theory and competition theory make predictions about the relative composition and spatial patterns of large-diameter trees compared to smaller diameter trees, these predictions are rarely tested. We established a 25.6 ha permanent plot within which we tagged and mapped all trees ≥1 cm dbh, all snags ≥10 cm dbh, and all shrub patches ≥2 m(2). We sampled downed woody debris, litter, and duff with line intercept transects. Aboveground live biomass of the 23 woody species was 507.9 Mg/ha, of which 503.8 Mg/ha was trees (SD?=?114.3 Mg/ha) and 4.1 Mg/ha was shrubs. Aboveground live and dead biomass was 652.0 Mg/ha. Large-diameter trees comprised 1.4% of individuals but 49.4% of biomass, with biomass dominated by Abies concolor and Pinus lambertiana (93.0% of tree biomass). The large-diameter component dominated the biomass of snags (59.5%) and contributed significantly to that of woody debris (36.6%). Traditional scaling theory was not a good model for either the relationship between tree radii and tree abundance or tree biomass. Spatial patterning of large-diameter trees of the three most abundant species differed from that of small-diameter conspecifics. For A. concolor and P. lambertiana, as well as all trees pooled, large-diameter and small-diameter trees were spatially segregated through inter-tree distances <10 m. Competition alone was insufficient to explain the spatial patterns of large-diameter trees and spatial relationships between large-diameter and small-diameter trees. Long-term observations may reveal regulation of forest biomass and spatial structure by fire, wind, pathogens, and insects in Sierra Nevada mixed-conifer forests. Sustaining ecosystem functions such as carbon storage or provision of specialist species habitat will likely require different management strategies when the functions are performed primarily by a few large trees as opposed to many smaller trees.