从叶内生物量分配策略的角度理解叶大小的优化

叶大小的变化是许多因素综合作用的结果, 对叶大小优化机制的研究有助于我们更好地理解植物的适应进化和生活史策略。该研究通过对浙江省清凉峰常绿阔叶混交林中的19个常绿阔叶物种和30个落叶阔叶物种叶水平上的相关性状进行分析, 探讨叶内生物量分配策略对叶大小优化的限制性影响。研究结果显示: 无论叶大小用面积还是质量表示, 常绿物种和落叶物种均呈现出叶内生物量分配到支撑结构的比例随着叶大小的增加而增加的规律, 这主要是由叶柄大小与叶片大小之间显著的异速生长关系导致的。这种异速生长关系在常绿物种和落叶物种中普遍存在。然而, 由于常绿物种对叶柄具有较高的机械以及抵抗冰冻栓塞等不利环境的需求, 在某一给定的叶面积下, 常绿物种比落叶物种具有更高的叶柄生物量投资。这些结果表明: 作为整个植株支撑投资的一个重要组成部分, 叶内支撑投资所占的生物量比例对叶大小的优化具有一定的限制性影响。     

The scaling of leaf area and mass: the cost of light interception increases with leaf size

For leaves, the light-capturing surface area per unit dry mass investment (specific leaf area, SLA) is a key trait from physiological, ecological and biophysical perspectives. To address whether SLA declines with leaf size, as hypothesized due to increasing costs of support in larger leaves, we compiled data on intraspecific variation in leaf dry mass (LM) and leaf surface area (LA) for 6334 leaves of 157 species. We used the power function LM=alpha LAbeta to test whether, within each species, large leaves deploy less surface area per unit dry mass than small leaves. Comparing scaling exponents (beta) showed that more species had a statistically significant decrease in SLA as leaf size increased (61) than the opposite (7) and the average beta was significantly greater than 1 (betamean=1.10, 95% CI 1.08-1.13). However, scaling exponents varied markedly from the few species that decreased to the many that increased SLA disproportionately fast as leaf size increased. This variation was unrelated to growth form, ecosystem of origin or climate. The average within-species tendency found here (allometric decrease of SLA with leaf size, averaging 13%) is in accord with concurrent findings on global-scale trends among species, although the substantial scatter around the central tendency suggests that the leaf size dependency does not obligately shape SLA. Nonetheless, the generally greater mass per unit leaf area of larger than smaller leaves directly translates into a greater cost to build and maintain a unit of leaf area, which, all else being equal, should constrain the maximum leaf size displayed.     

亚热带常绿林不同冠层小枝叶面积-叶生物量关系研究

叶面积与叶生物量的关系对于理解植物叶片的碳收益和投资权衡策略具有重要意义。收益递减假说认为植物的叶面积与叶生物量成显著异速生长关系,其异速生长指数<1.0,但该假说是否适用于不同生活型(常绿与落叶)亚热带木本植物不同冠层高度(上下冠层)当年生小枝的叶片仍不清楚。以江西亚热带常绿阔叶林的69种常绿与落叶木本植物当年生小枝上的叶为研究对象,采用标准化主轴回归估计(standardized major axis estimation, SMA)方法检验不同冠层高度和生活型叶面积与叶生物量的异速生长关系。结果显示:(1)当年生小枝叶生物量在不同冠层高度和生活型的植物中无显著差异(P>0.05),叶面积在常绿和落叶植物中有显著差异(P<0.05),常绿和落叶植物的比叶重存在显著差异(P<0.05),而落叶植物的比叶重在不同冠层高度存在显著差异(P<0.05),同一冠层,常绿植物比叶重显著高于落叶植物(P<0.05);(2)69种植物的叶面积与叶生物量异速生长指数具有物种特异性,60.9%的物种叶面积与叶生物量呈等速生长关系;(3)不同冠层和生活型植物的叶面积与...     

Allometric scaling relationship between above‐ and below‐ground biomass within and across five woody seedlings

Allometric biomass allocation theory predicts that leaf biomass (M_L) scaled isometrically with stem (M_S) and root (M_R) biomass, and thus above‐ground biomass (leaf and stem) (M_A) and root (M_R) scaled nearly isometrically with below‐ground biomass (root) for tree seedlings across a wide diversity of taxa. Furthermore, prior studies also imply that scaling constant should vary with species. However, litter is known about whether such invariant isometric scaling exponents hold for intraspecific biomass allocation, and how variation in scaling constants influences the interspecific scaling relationship between above‐ and below‐ground biomass. Biomass data of seedlings from five evergreen species were examined to test scaling relationships among biomass components across and within species. Model Type II regression was used to compare the numerical values of scaling exponents and constants among leaf, stem, root, and above‐ to below‐ground biomass. The results indicated that M_L and M_S scaled in an isometric or a nearly isometric manner with M_R, as well as M_A to M_R for five woody species. Significant variation was observed in the Y‐intercepts of the biomass scaling curves, resulting in the divergence for intraspecific scaling and interspecific scaling relationships for M_L versus M_S and M_L versus M_R, but not for M_S versus M_R and M_A versus M_R. We conclude, therefore, that a nearly isometric scaling relationship of M_A versus M_R holds true within each of the studied woody species and across them irrespective the negative scaling relationship between leaf and stem.     

Plant leaf traits,height and biomass partitioning in typical ephemerals under different levels of snow cover thickness in an alpine meadow

Aims In the cold life zones, snow cover is a comprehensive environmental factor that directly influences soil temperature, soil water content, light and nutrient availability. Plants in these zones develop a series of unique mechanisms involving phenological characteristics, reproductive strategies, physiology and morphology to adapt to environmental changes. This paper is focused on the responses of plant leaf traits, height and biomass partitioning to variations in snow cover thickness, in order to better understand the responses of plant functional traits and specific adaptation strategies under global climate change scenarios. Methods Three transects were established along a gradient of snow cover in an alpine meadow of Mt. Kaka, in the eastern Qinghai-Xizang Plateau. Primula purdomii, Pedicularis kansuensis and Ranunculus tanguticus, which are three widely distributed and dominant ephemerals, were sampled and studied, particularly at their blooming stages. Plant height, specific leaf area (SLA) and biomass partitioning were measured accordingly. Important findings The values of SLA in Pedicularis kansuensis and R. tanguticus were relatively greater under better soil conditions; it was smaller in Primula purdomii with thick snow cover. The relationship between aboveground biomass and belowground biomass in Primula purdomii was allometric at sites with both thick and thin snow cover. No significant relationships were found between aboveground biomass and belowground biomass in Pedicularis kansuensis and R. tanguticus at some individual sites. However, when samples of the three species were pooled, the relationships between aboveground biomass and belowground biomass were allometric at all sites, which did not support isometric scaling hypothesis. In addition, on sites with either thick or thin snow cover, aboveground biomass had greater rate of accumulation than belowground biomass; whereas on sites with medium snow cover, the rate of biomass accumulation was greater for belowground component than aboveground component. Functional traits and biomass variables were better correlated in Primula purdomii and Pedicularis kansuensis than in R. tanguticus.     

落叶收集法测定叶面积指数的快速取样方法

叶面积指数(LAI)是植被冠层结构的一个重要参数,它不仅是许多生态和气候模型的重要输入变量,而且是生态系统动态变化监测的一个重要指标。LAI可通过各种间接和直接的手段来观测,而间接观测的LAI值常常需要直接的观测数据来校验。落叶收集法是一种广泛使用的直接观测LAI的方法,过去的研究还未发现有涉及落叶收集的取样技术及其观测精度的内容。对长白山和北京地区落叶阔叶林的落叶进行了3a的观测,每年一次性收集落叶样品分析,研究结果表明:①不同层次落叶的含水量差异巨大,且落叶含水量的日变化明显。上下层落叶的含水量绝对值差异高达10%以上,日变化绝对值差异高达20%以上。因此,在野外收集落叶样本时,为减小落叶含水量变化所导致的LAI观测误差,应从上到下直到地面进行取样,且尽可能多地收集落叶样本。②在落叶阔叶人工林和天然林里,无论样地的大小(1hm2或30m×30m样地),无论取样单元的大小(1m2或25m2分辨率),林内的LAI分布很不均匀,LAI介于0-15.5(1m2分辨率的1hm2样地)或者2.6-9.1(25m2分辨率的30m×30m样地)。③要准确测定落叶林的LAI,收集落叶的样地面积越大越好,且尽量选择地势平坦的样地。对于1hm2或者30m×30m大小的样地,可随机布设一个10m×10m的小样地来观测,精度分别可达85%、80%。④10m×10m小样地的LAI观测,可将其分为4个相邻的5m×5m小样进行取样。对每个5m×5m小样,快速的取样方法是:Ⅰ.随机布设6个1m2取样,这样取样可以保证在99%概率水平上,100m2、30m×30m和1hm2样地的LAI观测精度分别为90%、75%、70%左右。Ⅱ.随机布设11个1m2取样。可以保证在99%概率水平上,100m2、30m×30m和1hm2样地的LAI观测精度分别为94%、80%、75%左右。     

不同雪被厚度下典型高山草地早春植物叶片性状、株高及生物量分配的研究

在高寒生命带, 雪被作为重要的综合环境因子, 影响着植物的生理生态特征、种群动态及群落演替等过程, 进而作用于生态系统的功能与服务。通过在青藏高原东缘高寒草甸设置厚雪、中雪和浅雪3个地段, 选取早春开花的常见种紫罗兰报春(Primula purdomii)、甘肃马先蒿(Pedicularis kansuensis)、高原毛茛(Ranunculus tanguticus), 研究了三种植物株高、叶片性状和生物量分配随雪被厚度的变化规律, 以及三者之间的关系。结果表明: 甘肃马先蒿和高原毛茛在生境状况较好的地段比叶面积相对较高, 紫罗兰报春由于具有根状茎和肉质根, 在厚雪地段比叶面积相对较小; 关于单个物种的地上-地下生物量的关系, 紫罗兰报春在厚雪和浅雪地段为完全一致的异速生长关系, 而甘肃马先蒿和高原毛茛在个别地段并无显著相关关系。总体而言, 三种植物整体样本的地上-地下生物量在不同雪被地段均为异速生长关系, 不支持等速生长假说, 并且地上生物量能够很好地解释地下生物量的变异; 功能性状和生物量指标间的相关性, 在紫罗兰报春和高原毛茛表现较好, 而在甘肃马先蒿表现较弱。植物对环境变化的适应具有一定的规律, 同时也表现出物种特异性。今后的研究中, 应注重构建植物适应环境变化的功能性状谱, 以更好地理解全球变化背景下植物功能性状的响应及其适应策略。     

森林自然更新过程中地上氮贮量与生物量异速生长的关系

研究植物氮贮量与生物量(M)之间的异速生长关系对于开展生态系统碳收支和氮循环研究具有重要意义。目前大量对氮贮量与生物量关系的研究主要集中在个体水平,对于群落水平的氮贮量和生物量之间的异速生长关系,以及群落自然更新过程如何影响该关系仍有待深入研究。利用3种森林类型皆伐后20多年自然更新过程中氮贮量和生物量的数据,采用简化主轴回归方法(reduced major axis,RMA)对不同自然更新阶段的森林氮贮量和生物量之间的异速生长指数和常数进行比较。结果表明:在不同的更新阶段,3种森林类型的植物氮贮量和生物量之间的异速生长指数均接近于1.0(即N∝M0.91-1.07)。异速生长常数随更新时间的增加而逐渐降低,导致3种森林类型整体上氮贮量正比于生物量的0.85次幂。异速生长常数的降低可能是由于在更新过程中叶生物量占整体生物量的比例逐渐下降,导致其对N吸收的生态化学计量制约所造成。     

Community composition and diversity of riparian forests regulate decomposition of leaf litter in stream ecosystems

Riparian forest plantings are a well‐established restoration technique commonly used to stabilize banks and intercept nutrient flow from adjacent agricultural fields. Tree species planted for these efforts may not reflect mature forest communities within the same region. Given contemporary research on links between biodiversity and ecosystem functioning, we conducted a leaf‐litter decomposition study to investigate how mixing of detrital resources that reflect forest community composition would regulate in‐stream leaf litter. Leaf litter bags containing material from a mature forest (Liriodendron tulipifera, Acer rubrum, Quercus rubra, full factorial treatments = 7) and a restored riparian forest (Cornus sericea, Fraxinus pennsylvanica, Platanus occidentalis, full factorial treatments = 7) were deployed in a stream reach that experienced riparian reforestation in 2004. Litter from the restored riparian community had less mass remaining (45.28 ± 2.27%) than that from the mature riparian community (54.95 ± 2.19%) after 5 weeks. In addition, mixed litter treatments in the restored riparian community had less mass remaining (40.54 ± 2.37%) than single‐species treatments (51.80 ± 4.05%), a pattern not observed in the mature forest community. Results highlight the importance of planting mixed‐species assemblages as this structure may regulate processes such as decomposition and food‐web structure, processes often not targeted in the restoration plans.     

桂西北喀斯特区原生林与次生林鲜叶和凋落叶化学计量特征

研究喀斯特生态脆弱区植物新鲜叶片与凋落叶的元素化学计量学性状,对该地区森林生态系统的恢复与重建具有重要指导意义。在桂西北喀斯特区分别选取了3个原生林群落与3个次生林群落,研究其建群种植物新鲜叶片和凋落叶的C、N、P元素含量及其生态化学计量特征。结果发现,6个群落建群种新鲜叶片C、N、P含量(其平均含量分别为404.3、22.5、1.75 mg/g)均大于凋落叶(平均含量分别为376.5、19.0、1.35 mg/g),鲜叶C:N、C:P、N:P比值(均值分别为17.8、244.9、13.8)均小于凋落叶(均值分别为19.3、315.3、16.3)。6种植物新鲜叶片N、P含量大于凋落叶,而N:P比小于凋落叶,表明喀斯特区植物对N的再吸收率大于P。3个原生林群落建群种鲜叶与凋落叶的平均C、N含量均大于次生林,而P含量则略小于次生林;原生林鲜叶与凋落叶的C:N比均小于次生林,C:P、N:P则大于次生林,推测次生林相对于原生林有更快的生长速率。原生林鲜叶N:P比为13—15之间,次生林鲜叶N:P比为11—12之间,次生林鲜叶与凋落叶的N:P比均小于原生林,说明原生林凋落物分解相对较慢,原生林能相对多的保留养分以供植物吸收,更能适应喀斯特石生环境。植物鲜叶和凋落叶的C:N与N:P比值均呈极显著正相关,说明叶片养分元素间具有共变的特性;叶片N、P含量呈正相关关系,表明植物N:P比具有相对的稳定性,这是高等陆生植物C-N-P元素计量的普遍规律,体现了植物群落对环境的适应。     

三种相思人工林和木荷林凋落叶分解的动态分析

对3种相思（Acacia Mill．）人工林和木荷（Schima superba Gardn．et．Champ．）林凋落叶的分解速率及磷和钾的动态变化进行了研究。结果表明,在分解过程中,卷荚相思（A．cincinnata F．Muell．）林、黑木相思（A．melanoxylon R．Br．）林、马占相思（A．magium Willd．）林和木荷林凋落叶干质量残留率分别为24．37％、13．51％、12．49％和41．86％,其中马占相思林的年分解系数最大,木荷林最小。凋落叶的磷含量大多呈上升趋势,其中黑木相思林和马占相思林表现为磷的净释放;卷荚相思林则在分解过程的前180d内表现为净固持,180d后表现为净释放;木荷林基本表现为磷的净固持。凋落叶的钾含量均为单调下降,且各林分均呈现出钾的净释放,分解末期各林分的钾释放率达90．14％～98．72％。与木荷林相比,3种相思人工林对土壤养分含量,特别是磷含量的维持非常有益。     

From individual leaf elongation to whole shoot leaf area expansion: a comparison of three Aegilops and two Triticum species

BACKGROUND AND AIMS: Rapid leaf area expansion is a desirable trait in the early growth stages of cereal crops grown in low-rainfall areas. In this study, the traits associated with inherent variation in early leaf area expansion rates have been investigated in two wheat species (Triticum aestivum and T. durum) and three of its wild relatives (Aegilops umbellulata, A. caudata and A. tauschii) to find out whether the Aegilops species have a faster leaf area expansion in their early developmental stage than some of the current wheat species. METHODS: Growth of individual leaves, biomass allocation, and gas exchange were measured on hydroponically grown plants for 4 weeks. KEY RESULTS: Leaf elongation rate (LER) was strongly and positively correlated with leaf width but not with leaf elongation duration (LED). The species with more rapidly elongating leaves showed a faster increase with leaf position in LER, leaf width and leaf area, higher relative leaf area expansion rates, and more biomass allocation to leaf sheaths and less to roots. No differences in leaf appearance rate were found amongst the species. CONCLUSIONS: Aegilops tauschii was the only wild species with rapid leaf expansion rates similar to those of wheat, and it achieved the highest photosynthetic rates, making it an interesting species for further study.     

Linking canopy leaf area and light environments with tree size distributions to explain Amazon forest demography

Forest biophysical structure – the arrangement and frequency of leaves and stems – emerges from growth, mortality and space filling dynamics, and may also influence those dynamics by structuring light environments. To investigate this interaction, we developed models that could use LiDAR remote sensing to link leaf area profiles with tree size distributions, comparing models which did not (metabolic scaling theory) and did allow light to influence this link. We found that a light environment‐to‐structure link was necessary to accurately simulate tree size distributions and canopy structure in two contrasting Amazon forests. Partitioning leaf area profiles into size‐class components, we found that demographic rates were related to variation in light absorption, with mortality increasing relative to growth in higher light, consistent with a light environment feedback to size distributions. Combining LiDAR with models linking forest structure and demography offers a high‐throughput approach to advance theory and investigate climate‐relevant tropical forest change.     

呼伦贝尔草原植物叶片大小与数量的权衡关系及其内在机制

叶片大小与数量的权衡关系是植物生长策略的基础,研究叶片大小与数量权衡关系的内在机制对于深刻理解植物生长策略具有重要意义.本研究以单个叶片干重表示叶大小,以单位茎上的叶片数量表示出叶强度,采用标准主轴回归分析方法研究呼伦贝尔草原植物叶片大小与数量的权衡关系以及内在机制.结果表明:陈旗嵯岗(典型草原)和陈旗八一(草甸草原)...     

Relationships among the Stem, Aboveground and Total Biomass across Chinese Forests

Forest biomass plays a key role in the global carbon cycle. In the present study, a general allometric model was derived to predict the relationships among the stem biomass Ms, aboveground biomass MA and total biomass MT, based on previously developed scaling relationships for leaf, stem and root standing biomass. The model predicted complex scaling exponents for MT and/or MA with respect to Ms. Because annual biomass accumulation in the stem, root and branch far exceeded the annual increase in standing leaf biomass, we can predict that MT ∝MA ∝ Ms as a simple result of the model. Although slight variations existed in different phyletic affiliations （i.e. conifers versus angiosperms）, empirical results using Model Type Ⅱ （reduced major axis） regression supported the model＇s predictions. The predictive formulas among stem, aboveground and total biomass were obtained using Model Type I （ordinary least squares） regression to estimate forest biomass. Given the low mean percentage prediction errors for aboveground （and total biomass） based on the stem biomass, the results provided a reasonable method to estimate the biomass of forests at the individual level, which was insensitive to the variation in local environmental conditions （e.g. precipitation, temperature, etc.）.