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。     

青藏高原高寒草地和内蒙古高原温带草地主要双子叶植物叶片解剖特征的比较研究

青藏高原高寒草地和内蒙古高原温带草地同属中国地带性的草地类型.然而,主导它们的环境因子有所不同.前者主要受生长季低温的影响,后者主要受干旱的影响.发育在这两类草地上的植物,在解剖特征上有何共性和差异,目前少有报道,但却是理解植物适应环境的关键所在.本文通过对青藏高原高寒草地和内蒙古高原温带草地71个样地65种双子叶植物叶片解剖特征的研究,对比分析了不同科、属及不同生活型植物叶片对干旱环境以及高寒环境的适应特征,探讨了叶片总厚度、上表皮和下表皮厚度、叶肉组织厚度、叶肉细胞密度、叶肉细胞表面积和体积等多个解剖特征之间的关系以及气候因子(生长季温度、降水)对它们的影响.结果表明:(1)总体上看,青藏高原与内蒙古高原草地植物的叶片解剖特征存在显著差异,除叶肉细胞密度外,青藏高原高寒草地植物的叶片各组成部分厚度、叶肉细胞表面积和体积均大于内蒙古高原温带草地植物;(2)同一科属植物的叶片解剖特征在两地存在显著差异,并显示出与所有物种总体分析一致的规律;(3)在调查范围内,两地草本植物除叶肉细胞密度外,叶片各组成部分厚度以及叶肉细胞表面积、体积均显著大于木本植物;(4)叶片的各解剖指标间存在显著的协同变化:叶片总厚度,上、下表皮厚度,叶肉细胞厚度,叶肉细胞表面积和体积之间均呈显著正相关,而叶肉细胞密度与这些指标显著负相关;(5)总体上看,青藏高原和内蒙古高原草地植物叶片的解剖特征与生长季温度的关系比与生长季降水密切.因此,尽管青藏高原高寒草地和内蒙古高原温带草地在外貌上有趋同性,但叶片解剖结构有着显著差异,这可能是由于植物对环境的适应进化及(或)环境的筛选作用共同作用的结果.     

内蒙古高原植物气孔性状的空间变异及其适应机制

气孔是植物与大气进行水气交换的阀门,调节着植物的蒸腾与光合作用。因此,气孔性状的变异影响着植物功能,并能够反映植物对环境的适应。目前,内蒙古高原草地植物气孔性状的空间变异及其对环境的响应尚未被探究。于2018年在内蒙古高原草原区沿降水梯度设置了10个受人为干扰较少的天然草地样地,采集每个样地及周围的植物,并对每个物种叶片上、下表皮的气孔性状进行了测量和计算。结果表明：(1)气孔密度(SD)、气孔大小(SS)和气孔相对面积(SRA)的均值分别为262.29个/mm²、378.84μm²和8.10%;下表皮与上表皮的气孔密度比(R.SD)、大小比(R.SS)和相对面积比(R.SRA)的均值分别为1.54、1.04和1.61。(2)由于草地生境开阔,气孔形态性状在不同生活型(草本与木本植物、一二年生与多年生草本)间没有显著差异(P>0.05)。(3)从草甸草原到荒漠草原,叶片SD和SRA减小,SS增大,气孔性状变异的主要驱动因素是年降水。(4)气孔综合指标SRA主要受SD驱动,降低叶片SD是内蒙古高原草地植物提高耐旱性的重要机制之一。(5)植...     

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。     

植物气孔扩散传导率的研究

本文对生长在可控环境温室中的花生气孔扩散传导率进行了实验研究,揭示了单个植株之间、上下表面之间、叶片不同部位以及冠层垂直方向上气孔扩散传导率的变异性。同时以气孔扩散传导率与环境条件的测定值为基础,对传导率对环境因子的反应进行了分析,植株顶部叶片气孔扩散传导率与太阳总辐射和空气饱和差有关系;冠层传导率与冠层截留辐射和空气饱和差有相关关系。     

羊草叶片气孔导度对环境因子的响应模拟

准确定量描述植物气孔对环境的响应是了解植物光合作用机理、预测植物生产力及其大气－植被－土壤系统中水分和热量交换的关键。利用松嫩平原盐碱化草地羊草光合生理特征的野外观测数据,分析了羊草叶片气孔导度对环境因子的反应,结果表明：羊草叶片气孔导度对环境因子变化敏感,尤其对瞬时光合有效辐射（ＰＡＲ）、叶片与空气间的水汽压亏损（ＶＰＤ）和空气温度（Ｔａ）反应十分明显。依据野外实测资料对国际上两类代表性气孔导度     

植物气孔扩散传导率的研究(以花生为例)

本文对生长在可控环境温室中的花生气孔扩散传导率进行了实验研究,揭示了单个植株之间、上下表面之间、叶片不同部位以及冠层垂直方向上气孔扩散传导率的变异性。同时以气孔扩散传导率与环境条件的测定值为基础,对传导率对环境因子的反应进行了分析,植株顶部叶片气孔扩散传导率与太阳总辐射和空气饱和差有关系;冠层传导率与冠层截留辐射和空气饱和差有相关关系。     

兰州市主要绿化植物气孔性状特征

气孔是植物叶片与外界环境进行水汽交换的门户, 利用气孔特征反映植物对环境变化的响应与适应有助于了解干旱区绿化植物的适应策略。于2019年7月通过气孔印迹法对兰州市40种主要绿化植物气孔性状进行观察与测定。采用标准化主轴估计和系统独立比较分析建立气孔性状间的相关关系; 通过计算Blomgerg’sK值以检验系统发育信号; 利用聚类分析和主成分分析对气孔性状特征进行功能群划分。结果表明: (1)在所研究的植物中, 气孔性状特征在个体间差异显著, 植物生长型(乔木、小乔木、灌木和草本)会显著影响气孔长度、宽度、开度和密度, 叶习性(落叶和常绿)仅对气孔开度有显著影响; (2)气孔长度与宽度、气孔开度与面积间均为显著的异速生长关系, 气孔密度与面积和开度间为负异速生长关系; (3)系统发育会对气孔性状的相关关系产生显著影响, 虽然气孔性状的系统发育保守性不强(K < 1), 但气孔开度和气孔开张比具有显著的系统发育信号; 4)依据气孔性状特征可以将绿化植物划分为: 低密度-大面积、高密度-小面积和中等密度-适中面积共3种功能群。结合系统发育和异速生长理论能更好地解释气孔性状变异及适应策略。     

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。     

羊草叶片气孔导度特征及数值模拟

对松嫩平原草地羊草叶片气孔导特征及与环境因子关系的研究结果表明,羊草叶片气孔导度日变化与环境因子密切相关,晴天表现为双峰曲线,阴天为单峰曲线,同时叶片气孔导度（gs)对瞬时光合有效辐射（PAR）,叶片与空气间的水汽压亏损（VPD）,空气温度（Ta）反应十分明显,依据野外实测资料,在对国际上两类代表性气孔导度模型验证比较的基础上,建立了适用于羊草草原的羊草叶片气孔导度对环境因子的响应模型gs=PAR(2.01Ta^2 147.74Ta-2321.11)/(444.62 PAR)(-538.04 VPD).     

Large-scale patterns of stomatal traits in Tibetan and Mongolian grassland species

We aimed to disentangle the influence of environmental variables on the spatial patterns in stomatal occurrence and stomatal traits. We surveyed the stomatal occurrence and the stomatal length (SL), density (SD) and index (SI) of 99 species in 150 grassland sites on the Tibetan and Inner Mongolian Plateau to explore their spatial patterns and evaluate the influence of climate. Of all species studied, two thirds were amphistomatous. The species from the Tibetan Plateau had larger but fewer stomata than those from Inner Mongolia. Among the climate factors examined, temperature and insolation affected SD, SI and SL, whereas the CO₂ partial pressure and the relative humidity affected SD. The climatic variables showed similar effects on the spatial variation of the adaxial and abaxial stomatal traits. We conclude that the low temperature and high insolation at high altitudes may be responsible for the larger and fewer stomata in plants on the Tibetan Plateau. The stomatal parameters that are presented here can be used to model the gas exchanges at the ecosystem scale.     

Abaxial and adaxial stomata from Pima cotton (Gossypium barbadense L.) differ in their pigment content and sensitivity to light quality

Sensitivity to light quality and pigment composition were analysed and compared in abaxial and adaxial stomata of Gossypium barbadense L. (Pima cotton). In most plants, abaxial (lower) stomatal conductances are higher than adaxial (upper) ones, and stomatal opening is more sensitive to blue light than to red. In greenhouse-grown Pima cotton, abaxial stomatal conductances were two to three times higher than adaxial ones. In contrast, adaxial stomatal conductances were 1·5 to two times higher than abaxial ones in leaves from growth chamber-grown plants. To establish whether light quality was a factor in the regulation of the relationship between abaxial and adaxial stomatal conductances, growth-chamber-grown plants were exposed to solar radiation outdoors and to increased red light in the growth chamber. In both cases, the ratios of adaxial to abaxial stomatal conductance reverted to those typical of greenhouse plants. We investigated the hypothesis that adaxial stomata are more sensitive to blue light and abaxial stomata are more sensitive to red light. Measurements of stomatal apertures in mechanically isolated epidermal peels from growth chamber and greenhouse plants showed that adaxial stomata opened more under blue light than under red light, while abaxial stomata had the opposite response. Using HPLC, we quantified the chlorophylls and carotenoids extracted from isolated adaxial and abaxial guard cells. All pigments analysed were more abundant in the adaxial than in the abaxial guard cells. Antheraxanthin and β-carotene contents were 2·3 times higher in adaxial than in abaxial guard cells, comparing with ad/ab ratios of 1·5–1·9 for the other pigments. We conclude that adaxial and abaxial stomata from Pima cotton have a differential sensitivity to light quality and their distinct responses are correlated with different pigment content.     

14种木犀榄族植物叶表皮微形态的研究

利用光学显微镜和扫描电子显微镜对木犀榄族（Oleeae）5属14种植物的叶表皮微形态特征进行了观察,结果表明：（1）叶表皮细胞形状有无规则形和多边形两种;下表皮腺点的数目远远大于上表皮,流苏树上表皮细胞没有腺点;叶下表皮有气孔器,形状为圆形、椭圆形;气孔器均为不规则型;气孔器外围角质层有放射状、条状、环状、颗粒状等多种类型;（2）气孔器和叶表皮细胞许多特征具有明显的种间差异,可以作为种间鉴定的重要依据;（3）叶表皮微形态结构上的许多共同特征,表明木犀榄族植物是单独一自然类群。上述特征可为木犀榄族植物属及物种的划分与鉴定、系统演化关系的探讨等提供参考。     

Leaf epidermal micromorphology of Cercis (Fabaceae: Caesalpinioideae)

The micromorphology of leaf epidermal cells and stomata of all eight species and one form (11 samples) of Cercis were observed by scanning electron microscopy and light microscopy. Both the adaxial and abaxial epidermal cells are polygonal or irregular in shape; the anticlinal walls are straight and arched or undulate. Two types of stomata, which occur only on the abaxial surface of the leaves, are found in the genus. The atypical paracytic type is present in only one species, Cercis chingii, and the anomocytic type is present in all other species. Interspecific differences are minor in the genus with regard to leaf epidermal characters, except for C. chingii, which is characterized by atypical paracytic‐type stomata, a two‐lipped outer stomatal rim, the highest stomatal density and undulate and densely pitted anticlinal walls in the adaxial epidermis. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 539–547.     

地黄叶和茎的解剖学及组织化学研究

采用解剖学和组织化学方法对地黄叶和茎的显微结构以及梓醇、多糖的分布进行观察研究,以明确梓醇和多糖在地黄叶和茎中的分布特征。结果显示:(1)地黄叶的上、下表皮均分布有腺毛和非腺毛,腺毛都属于头状腺毛,包括长柄和短柄的头状腺毛,两类腺毛的分泌物化学成分主要是黄酮和多糖;叶的上、下表皮上都分布有无规则型气孔,下表皮的气孔密度比上表皮的大,但气孔指数相差不大;栅栏组织由2～3层薄壁细胞构成,排列紧密,海绵组织薄壁细胞形状无规则,细胞间隙大。(2)组织化学研究表明,海绵组织中黄斑样的薄壁细胞是梓醇和多糖的贮存场所,这类薄壁细胞在叶片边缘的齿末端处最为集中,茎的皮层、韧皮部和木质部的薄壁细胞也都是梓醇和多糖的贮存场所。     

Relationship between stomatal frequencies,yield components and morphological characters in collections of winter wheat cultivars

Genotypes on the Belgian national list as well as about 120 genotypes of winter wheat from the world collection were examined for the relationships of their stomatal frequencies in the flag leaf with yield, yield components and other morphological characters. Stomata were in rows, with a higher density in adaxial leaf surface. Cultivars differed significantly in both stomatal density and number of rows of stomata per unit of leaf width. Yield and morphological characters in general showed no significant correlation with stomatal frequencies. Densities in all series examined ranged from 46 to 78 stomata per mm² in the adaxial leaf epidermis and from 32 to 93 in the abaxial one.     

Ontogenetic changes in stomatal size and conductance of sunflowers

The ontogenetic changes in stomatal size, frequency and conductance (g_s) on abaxial and adaxial leaf surfaces of sunflower plants (Helianthus annuus L. Russian Mammoth) were examined under controlled environmental conditions. The stomatal frequency on the adaxial and abaxial leaf surfaces decreased with leaf ontogeny and insertion level. The ratio of adaxial to abaxial stomatal frequency did not change with leaf ontogeny and insertion level, and 42–44% of total stomata was apportioned to the adaxial surface. Ontogenetic changes in stomatal pore length were detected and increased with ontogenesis. The stomatal length of both leaf surfaces had linear relationships with leaf area. Ontogenetic changes in g_s were similar between the two surfaces. However the adaxial g_s was lower than abaxial g_s in leaves of higher insertion levels. Conductance had a linear relationship with width x frequency but not with pore area.     

Stomata and Structure of Tetraploid Apple Leaves cultured in Vitro

Leaves of anther-derived tetraploid apple (Malus pumila Mill.)shoots were examined by low-temperature scanning electron microscopy(LT-SEM). Leaves were serrate and wide with an undulating adaxialsurface due to convex epidermal cells, apparently without crystallineepicuticular wax. Stomata were absent from the adaxial surface,except for the marginal teeth which exhibited 40-60 stomataper leaf; they probably originated from residual mitotic activity.One third of abaxial stomata was occluded by the residual cuticleof the mother guard cell across the stomatal pore which rupturedwhen the stomata became functional. The stomatal index was 7·2(± 1·6) with 60-75 stomata mm^-2, i.e. abaxialstomata of tetraploid leaves expanded in vitro were less frequentthan those in triploid leaves either cultured in vitro (475-575stomata mm^-2) or grown on the tree (320-390 stomata mm^-2) wherethe stomatal index was 21 (± 4). Freeze-fracture transsectionsshowed that the tetraploid in vitro leaves were composed ofa layer of adaxial epidermal cells, followed by a single layerof palisade cells and four to five layers of spongy mesophyllcells and the abaxial layer of epidermal cells, in contrastto juvenile seedling-grown apple leaves in which the two layersof palisade cells comprised the majority (52-60%) of the leafvolume. The same morphological features, such as wide and lesspointed leaves, reduced stomatal density and stomatal index,and increased stomatal size that were previously reported fortree-grown tetraploid leaves were also expressed in vitro. Thus,causes of the stomatal deformation in tissue-cultured Rosaceaeare interpreted to be in part genetic and not purely environmental.Copyright1994, 1999 Academic Press Malus pumila Mill., apple, biotechnology, breeding, cryo-preservation, CO₂, juvenile, low temperature-scanning electron microscopy (LT-SEM), micropropagation, ploidy, stomata, tissue-culture, transpiration     

27种木犀属植物叶表皮微形态特征的研究

利用光学显微镜和扫描电子显微镜观察了27种木犀属植物的叶表皮,测量并统计了气孔器类型、气孔大小、气孔密度、气孔指数及腺点密度等指标,以明确各种的叶表皮细胞形状及垂周壁式样、表皮角质膜、蜡质纹饰及气孔外拱盖的具体特征。结果显示:木犀属植物叶下表皮有气孔器,形状为圆形、椭圆形;气孔器类型多为不规则型,只有总状桂花和狭叶木犀为环列型;气孔器外围角质层有放射状、条状、环状、颗粒状等多种类型;叶表皮细胞形状有无规则形和多边形2种;下表皮腺点的数目远远大于上表皮。研究表明,木犀属植物气孔器和叶表皮细胞特征在种之间差异比较明显,可以作为种间鉴定的重要依据,具有重要的分类学意义。     

施肥对旱地冬小麦近轴和远轴叶面气孔敏感性的影响

 施肥降低旱地冬小麦的叶片水势。当作物体内出现水分胁迫时,冬小麦叶片两面气孔对施肥的反应有明显差异。远轴叶面气孔对施肥的反应比近轴叶面气孔敏感。旱地施肥以后,冬小麦远轴叶面气孔首先收缩,且收缩的程度比近轴叶面大,从而使远轴叶面气孔阻力与近轴叶面气孔阻力的比值(Rab／Rad)增大。旱地施肥以后,远轴和近轴叶面气孔阻力均急剧增大,并且随肥力水平的提高(施肥量增加)而缓慢增大,二者呈直线关系发展趋势。旱地施肥对土壤水势有影响,但不论是提高还是降低土壤水势,均增大Rab／Rad。说明施肥确有增强旱地冬小麦远轴叶面气孔对环境因素变化敏感性的作用。