13种栎属植物叶片的气孔特征及其相关性分析和分类学意义

以13种栎属(Quercus Linn.)植物〔包括产自辽宁省的槲栎组(Sect. Quercus)8种和麻栎组(Sect. Aegilops)2种以及引自北美洲的沼生栎组(Sect. Erythobalanus)2种和引自波兰的白栎组(Sect. Lepidabalanus)1种〕为试材,观察了这些种类叶片的气孔形态,并分析了气孔器和气孔的形态参数,还对各形态参数间的相关性进行了分析;在此基础上,对13种栎属植物进行了聚类分析。结果表明：栎属植物叶片的气孔仅分布于下表皮;气孔器排列属无规则型;气孔保卫细胞呈肾形,内壁加厚;气孔下陷,呈椭圆形或狭长椭圆形。供试种类的气孔和气孔器的形态参数差异明显,变异系数差异较大;气孔器的长轴长度、短轴长度和面积分别为18.24~27.46μm、14.63~23.18μm和221.56~501.70μm2,变异系数分别为7.73%~15.90%、7.10%~17.44%和14.13%~32.73%,气孔器指数为0.73~0.85;气孔的长轴长度、短轴长度、面积和密度分别为8.69~15.41μm、1.94~8.49μm、15.15~104.93μm2和462.32~984.44 mm-2,变异系数分别为12.03%~22.17%、13.65%~34.10%、27.95%~54.13%和8.10%~16.99%,气孔指数为0.22~0.57;总体上,按照多数气孔器和气孔参数从大至小的变化趋势依次排序为沼生栎组、白栎组、麻栎组、槲栎组。相关性分析结果表明：供试种类的气孔密度与气孔器和气孔的长轴及短轴长度均呈极显著(P<0.01)负相关;气孔器面积与气孔器指数和气孔指数分别呈极显著和显著(P<0.05)正相关;气孔面积与气孔器和气孔的长轴和短轴长度、气孔器指数和气孔指数呈极显著正相关,但与气孔密度呈显著负相关。聚类分析结果表明：在欧氏距离10处可将13种栎属植物分为3类,第Ⅰ类包含槲栎组的8种,第Ⅱ类包含麻栎组和白栎组的3种,第Ⅲ类包含沼生栎组的2种。研究结果显示：栎属植物叶片的气孔特征具有一定的稳定性,可作为栎属植物组间分类及亲缘关系分析的依据之一。     

润楠属植物叶片和枝条水力特征的协同关系

以润楠属(Machilus) 7种植物成年个体为材料,对其进行生理指标测定,并对它们的叶片水分供需关系以及木质部纹孔特征和导水效率之间的关联进行分析。结果显示,润楠属7种植物相比原始被子植物具有更高的叶脉密度(VD),叶脉密度为9.8～14.1 mm/mm~2;气孔密度(SD)与叶脉密度呈显著正相关,说明叶片水分供需存在协同关系;气孔密度与气孔大小(GLC)呈负相关;气孔越大的叶片其膨压丧失点(TLP)的绝对值越低。枝条边材比导率(Ks)较低,为0.13～1.87 kg·m~(-1)·s~(-1)·MPa~(-1),且种间差异较大。叶脉和气孔密度均与边材比导率呈正相关。边材比导率与纹孔膜面积、纹孔口面积以及纹孔口长短轴比例相关性不显著。研究结果表明润楠属植物虽然叶脉密度较高,且木质部水分供应和叶片结构具有协同关系,但木质部解剖结构较为原始,导管多具梯形穿孔板,导水效率低,只能适应比较湿润的生境。     

气孔在四季秋海棠营养器官和繁殖器官上的发育及相关性分析

主要观察了气孔在四季秋海棠营养器官和繁殖器官上的分布和发育情况,并分别对叶片和翅上气孔簇大小、气孔簇密度等指标的相关性进行了研究、结果表明:在叶片的下表皮、雌花和雄花的花被片、苞片、小苞片和翅上有气孔分布,而在茎、花梗上却未见气孔分布.叶片下表皮和翅上气孔通常成簇分布.在叶片的下表皮,气孔簇大小与气孔簇密度呈显著的负相关(P<0.05);气孔簇密度与叶片长度呈极显著的负相关(P<0.01).而翅上的气孔簇密度、气孔簇大小与子房长度无显著相关性(P>0.05).在四季秋海棠中,不同器官表皮的气孔簇大小是不同的,这可能与生理功能的不同有关.     

金星蕨科16种植物叶表皮特征的研究

利用光学显微镜对金星蕨科(Thelypteridaceae)8属16种植物的叶表皮形态进行了观察和研究,为金星蕨科植物的系统演化及分类提供科学依据。结果表明:(1)16种金星蕨科植物的叶片表皮细胞为不规则型,表皮细胞垂周壁大多呈深波状。(2)气孔为下生型,多呈椭圆形;共观察到6种气孔器类型(极细胞型、腋下细胞型、聚合极细胞型、聚腋下细胞型、不规则型和聚围绕细胞型),每种植物具2~5种不同类型的气孔器。(3)金星蕨科8属植物在表皮细胞大小、垂周壁形状、气孔大小和形状、气孔密度、气孔指数和气孔器类型上存在一定的属间差异。     

杉木衰退个体截干后叶片气孔器特征及其对3个因素的响应

本研究以广西全州县咸水林场杉木种子园中的衰退个体为对象进行截干试验,截干1年和4年后,采集当年抽梢的半木质化枝条进行叶片气孔特征观察,从不同无性系、不同截干时间和不同枝条方向等3个方面探讨其叶片气孔器特征的差异性。结果表明:杉木叶片的气孔,中等密度占多数,总体近乎正态分布;气孔器长度与宽度呈极显著正相关,与气孔密度则呈极显著负相关;对于气孔形态,不同无性系的影响最为显著,截干因素次之,枝条方向的影响最小。本试验揭示了截干措施对杉木叶片生理结构的影响,可为进一步探究杉木种子园衰退母树个体的复壮机制,提高经营水平、延长使用期限提供依据。     

西北干旱区葡萄净光合速率变化及其影响因素

葡萄作为西北干旱区主要经济作物之一,认识其光合生产过程对种植栽培至关重要。为探究大田自然条件下葡萄光合生理特征及影响葡萄光合作用的主要影响因子,该研究于2019年6—9月测定葡萄(品种：无核白)叶片光合作用及其生理生态因子日变化,采用通径分析方法分析各因子对叶片净光合速率的直接和间接作用,确定其主要影响因子,同时在全天分时段模式下进一步分析葡萄叶片净光合速率对各生理生态因子的响应。结果表明：(1)葡萄叶片净光合速率日变化总体呈现先升高、后下降的单峰型曲线变化特征。(2)葡萄叶片净光合速率与光合有效辐射、饱和水汽压差、空气温度、气孔导度和蒸腾速率呈极显著正相关,与相对湿度和胞间CO₂浓度呈极显著负相关。(3)各月影响葡萄叶片净光合速率变化的主要决定因子6月、8月和9月为蒸腾速率,而7月为气孔导度。(4)6—9月的葡萄叶片净光合速率与空气温度、光合有效辐射、饱和水汽压差的响应均呈“迟滞回环”关系,与蒸腾速率、气孔导度呈良好的线性关系(R²>0.85),与胞间CO₂浓度呈指数函数关系(R²=0.53...     

落羽杉属(杉科)叶表皮结构及气孔参数

落羽杉属Taxodium Rich.现生3种植物——落羽杉T. distichum (L.) Rich.、池杉T. ascendens Brongn.和墨西哥落羽杉T. mucronatum Tenore.的条形叶为双面气孔型或单面气孔型。叶片远轴面气孔分布于中脉两侧,每侧各有4-8列气孔。叶片中部气孔数量稳定,顶部和基部气孔数量比中部略少。近轴面气孔在中脉两侧各有1-4行,有时仅少数几个气孔或没有气孔分布。非气孔分布区内,表皮细胞长方形,细胞壁直或稍微呈波状,细胞长轴与叶片长轴一致。气孔分布区内的表皮细胞有时为多边形。气孔器椭圆形,长轴与叶片长轴垂直或成一定的角度。保卫细胞壁加厚明显,极端联合形成极层结构。落羽杉属3种现生植物的气孔密度和气孔指数差异显著,不同采集地的落羽杉气孔密度和气孔指数差异不显著。这3种植物的气孔指数的变异系数均小于气孔密度的变异系数,用气孔指数指示大气CO2浓度比用气孔密度可靠。     

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。     

青藏高原草地双子叶植物叶片的气孔特征研究

利用光学显微镜对青藏高原29种草地双子叶植物叶片的气孔形态与数量特征进行观察及差异显著性分析,为揭示青藏高原草地双子叶植物对高原环境的适应机制及探索气孔作为辅助分类的依据奠定理论基础。结果表明:(1)青藏高原草地双子叶植物大多数种类在叶片上、下表皮均分布有气孔,气孔随机排列,气孔器多为无规则型。(2)气孔长度(SL)较小,上、下表皮的气孔平均长度分别为26.20μm与25.56μm,且气孔密度(SD)与气孔指数(SI)相对较大。(3)不同科、属、种间叶片上、下表皮的SL、SD、SI差异均极显著。(4)叶片上、下表皮的6个气孔数量特征之间具有显著相关关系。(5)上表皮的SL、SD与不同科、属、种间显著相关,下表皮除SI与物种间相关不显著外,其他指标与科、属、种间显著相关。研究认为,青藏高原草地双子叶植物独特的气孔形态与数量特征是对高寒极端环境长期适应的结果,且气孔数量特征对植物辅助分类具有重要价值。     

4种城市绿化树种叶片PAHs含量特征与叶面结构的关系

用气质联用仪测定了长沙市樟树(Cinnamomu camphora)、广玉兰(Magnolia grandiflora)、桂花(Opsmanthus fragrans)和红檵木(Redrlowered loropetalum)4个主要绿化树种叶片中PAHs含量,同时测定了叶片的气孔密度、气孔长宽比、叶片的宽长比和叶面积等叶面结构特征值,探讨了叶面结构与叶片中PAHs含量的关系。结果表明:红檵木叶片的PAHs含量最高,为11.13mg·kg-1,16种PAHs在4树种叶片中均有不同程度的检出,其中以3环和4环为主,菲的浓度最高。除桂花外,在气温较低的秋冬季节,其余3种植物叶片气孔密度大PAHs含量高。叶面宽长比、气孔长宽比均与叶片PAHs含量呈极显著正相关,而叶面积与PAHs含量呈极显著负相关。表明叶面结构是影响叶片PAHs含量的重要因素。研究结果可为城市绿化树种合理选择与配置提供科学依据。     

七种热带雨林树苗叶片气孔特征及其可塑性对不同光照强度的响应

对生长在荫棚3种不同光照条件下和全自然光下的热带雨林4个冠层种(望天树、绒毛番龙眼、团花、红厚壳)和3个中层种(玉蕊、藤黄、滇南风吹楠)树苗叶片气孔特征以及它们的可塑性进行了研究、结果表明,这些植物的气孔全部着生在远轴面．7种植物中,玉蕊和绒毛番龙眼的气孔密度较大,滇南红厚壳和团花的保卫细胞最长．随光强的增大,气孔密度和气孔指数增大,单位叶气孔数在低光强下较大．除团花外,其它植物叶片气孔导度在50％光强处最大,而光强对保卫细胞的长度影响不显著．相关分析表明,气孔密度与植物单位叶的面积呈负相关。而与气孔导度的相关性不显著、尽管两种不同生活型植物气孔指数和单位叶气孔数在不同光强下的可塑性差异较小,但冠层树种气孔密度和气孔导度的可塑性显著高于中层树种．     

Allometric co-variation of xylem and stomata across diverse woody seedlings

Leaf stomatal density is known to co-vary with leaf vein density. However, the functional underpinning of this relation, and how it scales to whole-plant water transport anatomy, is still unresolved. We hypothesized that the balance of water exchange between the vapour phase (in stomata) and liquid phase (in vessels) depends on the consistent scaling between the summed stomatal areas and xylem cross-sectional areas, both at the whole-plant and single-leaf level. This predicted size co-variation should be driven by the co-variation of numbers of stomata and terminal vessels. We examined the relationships of stomatal traits and xylem anatomical traits from the entire plant to individual leaves across seedlings of 53 European woody angiosperm species. There was strong and convergent scaling between total stomatal area and stem xylem area per plant and between leaf total stomatal area and midvein xylem area per leaf across all the species, irrespective of variation in leaf habit, growth-form or relative growth rate. Moreover, strong scaling was found between stomatal number and terminal vessel number, whereas not in their respective average areas. Our findings have broad implications for integrating xylem architecture and stomatal distribution and deepen our understanding of the design rules of plants' water transport network.     

Stomatal response to air humidity and its relation to stomatal density in a wide range of warm climate species

The gas exchange of 19 widely different warm climate species was observed at different leaf to air vapour pressure deficits (VPD). In all species stomata tended to close as VPD increased resulting in a decrease in net photosynthesis. The absolute reduction in leaf conductance per unit increase in VPD was greatest in those species which had a large leaf conductance at low VPDs. This would be expected even if stomata of all species were equally sensitive. However the percentage reduction in net photosynthesis (used as a measure of the relative sensitivity of stomata of the different species) was also closely related to the maximal conductance at low VPD. Similarily the relative sensitivity of stomata to changes in VPD was closely related to the weighted stomatal density or crowding index.The hypothesis is presented that stomatal closure at different VPDs is related to peristomatal evaporation coupled with a high resistance between the epidermis and the mesophyll and low resistance between the stomatal apparatus and the epidermal cells. This hypothesis is consistent with the greater relative sensitivity of stomata on leaves with a high crowding index.The results and the hypothesis are discussed in the light of selection, for optimal productivity under differing conditions of relative humidity and soil water availablility, by observation of stomatal density and distribution on the two sides of the leaf.Visiting scientist, plant physiologist and research assitant of the Cassava Program     

Stomatal frequency of Quercus myrsinaefolia grown under different irradiances

Stomatal and epidermal cell frequencies and leaf area were measured in leaves of Quercus myrsinaefolia grown in the field under different relative photon flux density (PFD), which was the ratio of integrated PFD at the leaf surface to that at an open site. Leaf area showed a linear relationship with the relative PFD. Stomatal and epidermal cell frequencies increased with increasing relative PFD. Numbers of stomata and epidermal cells per leaf, and stomatal index (ratio of stomatal number to epidermal cell number) increased with increasing relative PFD.     

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。     

不同种源刨花楠林下幼苗叶功能性状与地理环境的关系

研究9个种源地天然刨花楠林下幼苗主要叶功能性状差异及其与地理环境的关系,分析刨花楠林下幼苗对地理环境变化的响应与适应机制.结果表明: 不同种源间刨花楠林下幼苗主要叶功能性状种内变异系数较大(8.8%～28.2%),其中种源间比叶面积、叶相对含水率、叶组织密度和叶厚差异显著,表明刨花楠林下幼苗具有较强的叶片形态可塑性.叶组织密度与叶干物质含量、叶相对含水率均呈显著正相关,与比叶面积、叶厚则呈显著负相关;比叶面积与叶干物质含量、叶面积均呈显著负相关,反映刨花楠林下幼苗可通过叶片性状组合的调整和平衡以响应地理环境变化.影响刨花楠林下幼苗叶功能性状可塑性的主要环境因子为经度、纬度、>10 ℃年积温和年均温.叶厚随着经度的增加而降低,叶干物质含量和叶相对含水率则随着经度的增加而增加;叶组织密度与经度和年均温呈显著正相关,且经度对其影响大于年均温;叶面积与>10 ℃年积温和经度呈显著正相关,且前者对其影响大于后者.     

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.     

Stomatal frequency of Quercus myrsinaefolia grown under different irradiances

Stomatal and epidermal cell frequencies and leaf area were measured in leaves of Quercus myrsinaefolia grown in the field under different relative photon flux density (PFD), which was the ratio of integrated PFD at the leaf surface to that at an open site. Leaf area showed a linear relationship with the relative PFD. Stomatal and epidermal cell frequencies increased with increasing relative PFD. Numbers of stomata and epidermal cells per leaf, and stomatal index (ratio of stomatal number to epidermal cell number) increased with increasing relative PFD. This revised version was published online in June 2006 with corrections to the Cover Date.     

Distribution of stomata on the second leaf ofZea mays following root hypoxia

The changes in leaf dimensions, transverse and longitudinal gradients in stomatal density and the total number of stomata under the influence of root hypoxia were followed. In spite of considerably reduced leaf area following hypoxia the total number of stomata per leaf was not changed significantly. The resulting increase in stomatal density was not uniform being the most prominent in the basal part of the leaf where the distances between stomata and between rows of stomata became shorter.     

灰木莲种源幼苗叶片性状表型多样性分析

以越南收集的12个灰木莲（Manglietia conifera Dandy）种源为材料,对叶片形态性状（叶长、叶宽、叶面积、叶周长、长宽比、叶柄）以及微形态特征（气孔器密度、气孔器长、气孔器宽、气孔器面积、长宽比）进行测定,采用方差分析、变异系数、相关分析和主成分分析等方法进行分析。结果表明：灰木莲不同种源间的叶表型性状存在显著差异;种源中,LC2的叶形态性状的平均变异系数最大（22.09%）,TQ2的变异系数最小（12.76%）;叶表型性状中,叶面积的变异系数最大（28.60%）,气孔器宽的变异系数最小（5.19%）;相关性分析结果表明叶表型性状间存在显著或极显著的相关关系,而地理因素中经度与叶周长显著相关,经纬度与叶长宽比呈显著相关,海拔与叶表型性状间的相关性不显著（P<0.05）;主成分分析表明前3个主成分的累计贡献率达到了89.29%,基本代表原始性状的全部信息。灰木莲12个种源经聚类分析在欧式距离5阈值处可分为4类。灰木莲种源间的叶表型性状存在着丰富的变异,纬度对灰木莲叶形态特征有明显影响,灰木莲种源间和性状间的变异程度存在着差异,本研究为灰木莲遗传改良提供理论依据。