基部被子植物气孔性状与叶脉密度的关联进化

植物叶片通过气孔的水分蒸腾散失和叶脉的水分供应达到水分平衡,而基部被子植物在进化过程中叶片水分供应和蒸腾散失是否达到平衡或关联进化还缺乏了解。本研究以11种基部被子植物为材料,测定了气孔密度、气孔长度、叶脉密度和叶片厚度4个叶片性状,并结合系统发育树,利用系统发育独立对比的方法分析这些性状之间的关联进化。结果显示：沿进化方向,气孔密度和叶脉密度逐渐增加,而气孔长度和叶片厚度有减小的趋势;无论是否考虑系统发育的影响,气孔密度都与叶脉密度呈显著正相关关系,说明二者之间存在关联进化,并证实了基部被子植物叶片水分平衡假说;气孔密度和长度、叶脉密度均与叶片厚度呈显著线性相关,但在去除系统发育的影响后这种线性相关关系不再显著,说明叶片厚度与其它三个叶片性状不存在关联进化。本研究结果还表明,叶片的水分供应和散失乃至CO2通透性的平衡主导着基部被子植物叶片结构和功能的进化。     

润楠属广布种和狭域种幼苗生理生态特征

润楠属植物属于基部被子植物类群,大部分物种局限分布于热带亚热带森林的潮湿生境,但也有些物种分布范围较广。本研究以润楠属11个物种的幼苗为材料,测定了一系列植物叶片水力学性状和解剖结构,包括:比叶重、叶片密度、气孔密度、叶脉密度、膨压丧失点水势、栅栏组织和海绵组织厚度等。结果表明:与其它分布在热带亚热带地区的被子植物相比,润楠属植物的叶脉密度较低,推测叶脉密度受强烈的进化限制;该属植物叶脉密度与气孔密度、栅栏和海绵组织的比值呈显著的正相关,表明该属植物能够维持叶片水平的水分供需平衡;广布种比狭域种具有更低的叶片膨压丧失点和更高的叶片密度,耐旱能力更强,但是比叶重差异不显著;叶片的膨压丧失点与物种的最大树高呈反比,即更高的物种叶片耐失水能力更强,说明植物叶片耐旱性与植物本身的遗传特性有关。本研究结果显示,叶片水力学性状可以较好地用于解释润楠属植物的地理分布。     

4种槭树叶功能性状及其关系对季节变化的响应

以牡丹江地区的白牛槭（Acer mandshuricum）、茶条槭（Acer tataricum subsp. ginnala.）、糖槭（Acer saccharum）、五角槭（Acer pictum subsp. mono）4种槭树为研究对象,分别于春季、夏季和秋季进行取样,测定叶片性状指标（叶厚度、气孔长度、气孔宽度、气孔密度、叶脉密度、比叶面积及色素质量分数）,分析叶片各性状的季节变化趋势,并探讨色素与叶性状间在不同季节下的关系。结果如下（：1）4种槭树均表现为在夏季具有较高的叶厚度、较低的比叶面积和气孔密度,在秋季具较高的比叶面积和叶脉密度、较低的叶厚度。（2）4种槭树均为在夏季有较高的叶绿素a、b,在秋季色素质量分数均降低,季节变化区间分别为叶绿素a 77.40%～98.80%,叶绿素b 85.60%～99.53%,类胡萝卜素4.29%～78.52%。（3）色素与叶性状关系密切,季节动态下色素与比叶面积、气孔密度、叶脉密度正相关,与叶厚度、气孔长度、气孔宽度负相关（P<0.05）,但不同季节相关性略有差异。4种彩叶植物的叶片在应对不同季节的气候条件时形成了不同的构建策略...     

冬小麦品种(系)旗叶叶脉性状及其与气孔性状间的关联性

植物叶脉结构决定着叶水分运输能力,而气孔特性则与水分散失有关,研究二者间的关联性对认识与植物水分利用有关的生理功能间关系及其调控有重要理论价值。本研究以冬小麦为例,系统研究了15个小麦品种(系)旗叶的叶脉性状及其与气孔性状间的关联性。结果表明:在小麦3种类型叶脉中,小纵向脉的密度占叶脉密度的65.8%;小纵向脉间距与大纵向脉间距及横向脉间距均呈显著正相关,大、小纵向脉密度与其间距呈极显著负相关;供试小麦品种(系)叶片上、下表面的气孔大小相差不大,但上表面气孔密度显著高于下表面,上、下表面的气孔长、气孔宽与气孔密度存在显著正相关,但气孔长与气孔密度之间存在显著负相关;3种类型叶脉中,大纵向脉与气孔特征间并无显著相关关系,而小纵向脉、横向脉和气孔特征间存在一定的相关性,小麦品种(系)叶脉密度与下表面气孔密度和总气孔密度呈显著正相关,但与上表面气孔长呈显著负相关,从而揭示了小麦品种(系)水分供给和需求存在协调性的结构基础。     

叶脉网络功能性状及其生态学意义

叶脉网络结构是叶脉系统在叶片里的分布和排列样式。早期叶脉网络结构研究主要集中在其分类学意义上; 近年来叶脉网络功能性状及其在植物水分利用上的意义已成为植物生态学研究的热点。该文介绍了叶脉网络功能性状的指标体系(包括叶脉密度、叶脉直径、叶脉之间的距离、叶脉闭合度等), 综述了叶脉网络功能性状与叶脉系统功能(包括水分、养分和光合产物等物质运输、机械支撑和虫害防御等)的关系, 叶脉网络功能性状与叶片其他功能性状(包括比叶重、叶寿命、光合速率、叶片大小、气孔密度等)的协同变异和权衡关系, 以及叶脉网络功能性状随环境因子(包括水分、温度、光照等)的变化规律等方面的最新研究进展。此外, 叶脉网络功能性状的研究成果也被应用于古环境重建、城市交通规划、流域规划及全球变化研究中。由于叶脉网络功能性状是环境因子与系统发育共同作用的结果, 未来开展分子—叶片—植物—生态系统等多尺度的叶脉网络功能性状研究, 理清叶脉网络功能性状与气孔失水—茎干导水—根系吸水等植物水分利用的关系, 将为预测植物及生态系统对全球变化的响应提供新的启示。     

Plant ontogenetic changes in vein and stomatal traits and their relationship with economic traits in leaves of three Mediterranean oaks

3种地中海橡树在叶脉和气孔性状上的个体发育性变化及其与叶片经济性状的关系 我们比较了3个地中海栎属(Quercus)树种幼苗和成龄树的叶脉和气孔性状。之前的研究表明,幼苗的气体交换率往往比成龄树的要高。我们的研究目标是确认叶脉和气孔的性状在整个植物个体发育过程中是否会随着气体交换率的变化而同步改变。我们提出了以下两个备选假设：幼苗比成龄树有着更高的叶脉和气孔密度(假设1);幼苗在维管组织上的投资更少,从而降低了建设成本(假设2)。本研究是在西班牙中西部进行的,我们对每个树种不同生长阶段的植株随机采集了10个样本。我们测量了气孔和叶脉的平均性状(单位叶面积里的气孔大小和数量、叶脉密度、叶脉体积、叶脉到表皮的距离),比叶质量和叶片厚度。研究结果表明,细脉密度和单位面积叶脉体积随着树龄的增加而增加,这似乎与气体交换率方面的个体发育趋势不一致。这种差异支持了我们的假设2,说明植株在幼苗阶段减少对维管组织的投资以最大限度地提高生长速度是其当务之急。幼苗叶片中较大的叶脉间距可以由叶脉到表皮的较短距离所补偿。因此,薄叶片可能是幼苗的一个必要性状,它可以让水分沿较短路径输送到蒸发点,而无需对高成本的维管组织进行大量投资。     

不同水分处理下喀斯特土层厚度异质性对两种草本叶片解剖结构和光合特性的影响

为了探究不同水分处理下草本植物对喀斯特土层厚度变化的叶片形态建成和光合生理响应,以黑麦草(Lolium perenne L.)和苇状羊茅(Festuca arundinacea Schreb.)为研究对象,通过盆栽水分受控试验,研究了3种水分处理[正常供水(W_(ck)),减水1组(D1)和减水2组(D2)]下3种土层厚度[浅土组(S_S)、对照组(S_(CK))和深土组(S_D)]对两种草本叶片解剖结构和光合特性的影响。结果表明:(1)正常供水下(W_(ck)),黑麦草和苇状羊茅在浅土组(S_S)的气孔密度和气孔限制值(Ls)均显著高于对照组(S_(CK)),净光合速率(Pn)、胞间CO_2浓度(Ci)和蒸腾速率(Tr)降低;在深土组(S_D),两种植物的气孔密度都有所下降,黑麦草的叶脉密度、Pn和Tr均低于对照组,而苇状羊茅的叶脉密度和Pn表现出增加;(2)D1水分条件下,黑麦草在浅土组的气孔密度较对照组增加,叶脉密度、Pn和Tr均降低,而苇状羊茅的气孔密度有所降低,叶脉密度、Pn和Tr未受到显著影响;在深土组中,黑麦草的气孔密度不变,叶脉密度增加,而Pn和Tr均降低;苇状羊茅的气孔密度降低,但叶脉密度、Pn和Tr均升高;(3)D2水分条件下,两种植物在浅土组的叶脉密度较对照组均增加,气孔密度、Pn和Tr均受到抑制;在深土组,黑麦草的远轴面气孔密度较对照组下降,两种植物的其他指标未受到明显影响。可见,在不同水分条件下,植物的叶片解剖结构和光合特性对不同土层厚度的响应不一,且不同物种间也有差异。总体上随着水分减少,土层厚度降低对植物的光合抑制作用增强,而厚度增加对深根植物的光合促进作用和对浅根植物的光合抑制作用先增强后减弱。植物气孔和叶脉性状特征随水分条件的变化在一定程度上与叶面积和叶片宽度的变化有关。     

气孔特征与叶片功能性状之间关联性沿海拔梯度的变化规律——以长白山为例

气孔是陆生植物与外界环境进行水分和气体交换的主要通道,控制着植物的光合作用和蒸腾过程。植物往往通过多种性状的组合来适应变化的环境,叶片功能性状之间的紧密关系已经在不同尺度得到证实。然而,植物气孔特征与叶片其它功能性状是否存在关联性以及这种关联性是否会受到环境变化梯度的影响仍鲜少报道。沿长白山北坡6个海拔梯度,测定了150种植物的气孔特征和叶片功能性状。结果发现,气孔密度(SD)与比叶面积(SLA)负相关,与单位面积的叶氮含量(N_(area))正相关;除了SLA和N_(area)外,气孔长度(SL)与SLA、叶片厚度(LT)和单位质量的叶氮含量(N_(mass))均存在显著的相关性(P0.05)。然而,气孔特征与叶片功能性状的相关性只在部分海拔梯度存在。此外,发现SD与SL之间存在稳定一致的负相关关系。这些结果表明,植物气孔特征与叶片形态和化学特征对环境变化的适应存在一定的协同变异性,但这种关系不具有普适性,主要与气孔特征和叶片功能性状的选择压力存在差异以及物种分布范围相关。未来仍需要在更多物种和不同区域内来验证气孔特征与植物功能性状之间的关联关系。     

叶脉结构与功能及其对叶片经济谱的影响

叶脉由贯穿于叶肉内部的维管组织及其外围机械组织构成,多样化的脉序及网络结构使叶脉系统发生变异和功能分化。该文综述了叶脉系统结构与功能的最新研究进展。通过聚焦叶脉分级系统的结构与功能及其在叶片经济谱(LES)中的重要性,解释叶脉性状与其它叶片功能性状之间的关系及机制。不同等级叶脉在机械支撑与水分运输方面存在功能分化,其中1–3级粗脉在维持叶片形状和叶表面积以及物理支撑方面发挥重要作用,有利于维持叶片最大受光面积;4级及以上细脉具有水分调节功能,它们与气孔相互协调,影响叶片水分运输、蒸腾散热和光合作用速率。叶片生长过程与叶脉发育的动态变化模式决定叶脉密度,并影响叶脉密度与叶片大小之间的关系:叶面积与粗脉密度呈显著负相关,与粗脉直径呈显著正相关,而与细脉密度无关。与叶脉性状相关的叶片经济谱框架模型预测,叶脉密度较高的叶片寿命短、比叶重较小,叶片最大碳同化速率、代谢速率以及资源获取策略潜力较高。     

高粱、紫苏叶脉密度与光合特性的关系

叶脉是植物叶片光合作用水分输送的重要结构。为阐述叶脉与光合特性之间的关系,以C4植物高粱(Sorghum bicolor)、C3植物紫苏(Perilla frutescens)为实验材料研究了叶脉密度和光合特性之间的关系。结果表明,与紫苏相比,高粱叶片叶脉密度大,导水能力强,蒸腾速率高,但气孔密度小。进一步分析表明,高粱叶片近轴侧气孔密度占总气孔的比例明显高于紫苏。叶脉密度大的高粱具有较高的净光合速率;而紫苏叶脉密度小,净光合速率也较低。由此表明,较高的叶脉密度有利于支持较高的光合速率,但研究表明叶脉密度和气孔密度可能不存在严格的协同变异关系。研究结果对理解植物光合作用适应有重要意义。     

Leaf Photosynthetic Rate of Tropical Ferns Is Evolutionarily Linked to Water Transport Capacity

Ferns usually have relatively lower photosynthetic potential than angiosperms. However, it is unclear whether low photosynthetic potential of ferns is linked to leaf water supply. We hypothesized that there is an evolutionary association of leaf water transport capacity with photosynthesis and stomatal density in ferns. In the present study, a series of functional traits relating to leaf anatomy, hydraulics and physiology were assessed in 19 terrestrial and 11 epiphytic ferns in a common garden, and analyzed by a comparative phylogenetics method. Compared with epiphytic ferns, terrestrial ferns had higher vein density (D_vein), stomatal density (SD), stomatal conductance (g_s), and photosynthetic capacity (A_max), but lower values for lower epidermal thickness (LET) and leaf thickness (LT). Across species, all traits varied significantly, but only stomatal length (SL) showed strong phylogenetic conservatism. A_max was positively correlated with D_vein and g_s with and without phylogenetic corrections. SD correlated positively with A_max, D_vein and g_s, with the correlation between SD and D_vein being significant after phylogenetic correction. Leaf water content showed significant correlations with LET, LT, and mesophyll thickness. Our results provide evidence that A_max of the studied ferns is linked to leaf water transport capacity, and there was an evolutionary association between water supply and demand in ferns. These findings add new insights into the evolutionary correlations among traits involving carbon and water economy in ferns.     

Variation in leaf traits at different altitudes reflects the adaptive strategy of plants to environmental changes

Leaf anatomical traits play key roles in plant functions and display evolutionary adaptive changes to suit the surrounding environment. To reveal the adaptive mode and mechanisms of plants in response to global warming, we analyzed leaf morphology and anatomical structures in three different species, Epilobium amurense Hausskn., Pedicularis densispica Franch., and Potentilla fulgens Wall. ex Hook., growing along an elevational gradient (3,000–4,600 m) in the Yulong Mountains. The results showed leaf length and width decreased, whereas leaf thickness increased with increasing altitude in all three species. Thickness of leaf upper epidermis, lower epidermis, palisade and spongy mesophyll, and main vein increased with rising altitude. Stomatal density in each species increased with rising elevation. These results illustrate that plants can adapt to the environmental changes that accompany high altitudes by decreasing leaf area and increasing leaf thickness, mesophyll tissue thickness, and stomatal density. Such morphological and anatomical plasticity would lead to lower transpiration rates, enhanced internal temperature and water status, and improved photosynthetic capability.     

三种锦鸡儿属植物水力结构特征及其干旱适应策略

水分胁迫是干旱半干旱区限制植物生长的主要因素。以干旱半干旱区的3种锦鸡儿属植物为研究对象,从生态适应策略角度来分析3种锦鸡儿植物产生生态分离的原因。对三种锦鸡儿属植物茎干叶片的显微结构、生理功能(导水率、光合速率以及水分利用效率)进行测定,并统计了3种锦鸡儿植株的形态特征,如一、二级枝的直径、长度、末端叶面积。结果表明:三种锦鸡儿属植物都能形成较小的导管直径来适应旱生环境,但是在导水结构上又表现出一定的差异性。中间锦鸡儿的导管直径最小,次脉密度和最大净光合速率最大;柠条锦鸡儿的导管直径、叶片厚度和比叶重(LMA)最大。小叶锦鸡儿在导水率下降50%时的水势(P_(50))最大,水分胁迫时极易发生栓塞,但正是由于导管的栓塞降低了水分运输效率,使其在旱生环境中能够通过减少水分的供应来降低水分的丧失,从而保证自身生长的水分需求;而中间锦鸡儿则主要通过减小导管直径来适应旱生环境;柠条锦鸡儿的水分利用效率最高,抗栓塞能力最强,抗旱性最好,同时柠条锦鸡儿可以通过减少蒸腾面积来减少水分的丧失。植物的导管直径大小、叶片厚度、LMA、叶脉密度对植物导水速率、光合速率等生理功能都有一定的影响。     

Linking vein properties to leaf biomechanics across 58 woody species from a subtropical forest

• Leaf venations have elements with relatively lower elasticity than other leaf tissue components, which are thought to contribute to leaf biomechanics. A better mechanistic understanding of relationships between vein traits and leaf mechanical properties is essential for ecologically relevant interpretation of leaf structural variations.
• We investigated 13 major (first to third order) and minor (>third order) vein traits, six leaf mechanical properties and other structural traits across 58 woody species from a subtropical forest to elucidate how vein traits contribute to leaf biomechanics.
• Across species, vein dry mass density (ρ_v), total vein dry mass per leaf area (VMA) and minor vein diameter (VD_min), but not the lower‐order vein density (VLA_1?2), were positively correlated with leaf force to punch (F_p) and force to tear (F_t). Structural equation models showed that ρ_v and VD_min not only contribute to leaf mechanical properties directly (direct pathway), but also had impacts on leaf biomechanics by influencing leaf thickness and leaf dry mass per area (indirect pathway).
• Our study demonstrated that vein dry mass density and minor vein diameter are the key vein properties for leaf biomechanics. We also suggest that the mechanical characteristics of venations are potential factors influencing leaf mechanical resistance, structure and leaf economics spectrum.

     

濒危种观光木小枝生物量分配与功能性状的纬度变异规律及其影响因素

为了探讨观光木当年生枝条生物量的分配规律和叶片功能性状的变化规律及其影响因素,该研究以濒危物种观光木(Michelia odora)为对象,测定了广西地区5个不同纬度上观光木当年生小枝及叶片功能性状.结果表明:(1)随着纬度的增加小枝总重和总叶重总体呈异速生长关系,其生物量更多趋于对小枝的构建.(2)观光木叶功能性状呈...     

Transpiration and stomatal control: a cross-species study of leaf traits in 39 evergreen and deciduous broadleaved subtropical tree species

Key message

Using an extensive dataset for 39 subtropical broadleaved tree species, we found traits of the leaf economics spectrum to be linked to mean stomatal conductance but not to stomatal regulation.

Abstract

The aim of our study was to establish links between stomatal control and functional leaf traits. We hypothesized that mean and maximum stomatal conductance (g _s) varies with the traits described by the leaf economics spectrum, such as specific leaf area and leaf dry matter content, and that high g _s values correspond to species with tender leaves and high photosynthetic capacity. In addition, we hypothesized that species with leaves of low stomata density have more limited stomatal closure than those with high stomata density. In order to account for confounding site condition effects, we made use of a common garden situation in which 39 deciduous and evergreen species of the same age were grown in a biodiversity ecosystem functioning experiment in Jiangxi (China). Daily courses of g _s were measured with porometry, and the species-specific g _s~vpd relationships were modeled. Our results show that mean stomatal conductance can be predicted from leaf traits that represent the leaf economics spectrum, with a positive relationship being related to leaf nitrogen content and a negative relationship with the leaf carbon: nitrogen ratio. In contrast, parameters of stomatal control were related to traits unassociated with the leaf economics spectrum. The maximum of the conductance~vpd curve was positively related to leaf carbon content and vein length. The vpd at the point of inflection of the conductance~vpd curve was lower for species with higher stomata density and higher for species with a high leaf carbon content. Overall, stomata size and density as well as vein length were more effective at explaining stomatal regulation than traits used in the leaf economics spectrum.     

不同光环境下枫香幼苗的叶片解剖结构

落叶阔叶树种枫香是常绿阔叶林的优势种之一。对比研究了重庆丰都世坪林场不同光环境下枫香幼苗叶片的解剖特征,结果表明:1)旷地枫香叶片的厚度大,气孔个体较大且排列紧密,栅栏组织发达,具有明显的旱生叶特征;而林下的枫香叶片明显变薄,气孔小且排列稀疏,海绵组织发达,具有一定的阴生叶特征;2)不同光环境下,气孔密度、栅栏组织的厚度和层数、栅栏细胞长度以及栅栏组织与海绵组织厚度比(P/S)等具有较高的可塑性,表明这些因素对于枫香适应不同光环境具有较大的作用;3)从叶片解剖特征与功能的统一来看,阳性树种枫香幼苗更适合生长在旷地等高光生境中,而不易生长在郁闭的林下;4)落叶阔叶树种在中亚热带常绿阔叶林群落恢复与演替中具有重要作用。     

Stem growth habit affects leaf morphology and gas exchange traits in soybean

Backgrounds and Aims

The stem growth habit, determinate or indeterminate, of soybean, Glycine max, varieties affects various plant morphological and developmental traits. The objective of this study is to identify the effect of stem growth habit in soybean on the stomatal conductance of single leaves in relation to their leaf morphology in order to better understand the ecological and agronomic significance of this plant trait.

Methods

The stomatal conductance of leaves on the main stem was measured periodically under favourable field conditions to evaluate g_max, defined as the maximum stomatal conductance at full leaf expansion, for four varieties of soybean and their respective determinate or indeterminate near isogenic lines (NILs). Leaf morphological traits including stomatal density, guard cell length and vein density were also measured.

Key Results

The value of g_max ranged from 0·383 to 0·754 mol H₂O m⁻² s⁻¹ across all the genotypes for both years. For the four pairs of varieties, the indeterminate lines exhibited significantly greater g_max, stomatal density, numbers of epidermal cells per unit area and total vein length per unit area than their respective determinate NILs in both years. The guard cell length, leaf mass per area and single leaf size all tended to be greater in the determinate types. The variation of g_max across genotypes and years was well explained by the product of stomatal density and guard cell length (r = 0·86, P < 0·01).

Conclusions

The indeterminate stem growth habit resulted in a greater maximum stomatal conductance for soybean than the determinate habit, and this was attributed to the differences in leaf structure. This raises the further hypothesis that the difference in stem growth habit results in different water use characteristics of soybean plants in the field. Stomatal conductance under favourable conditions can be modified by leaf morphological traits.Key words: Soybean, Glycine max, stem growth habit, stomatal conductance, stomatal density, guard cell length, near isogenic lines