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.     

青藏高原矮火绒草(Leontopodium nanum)叶片性状对海拔高度变化的响应

植物叶片对环境变化十分敏感,能反映植物适应环境所形成的生存策略。为揭示高寒植物叶片性状对海拔高度变化的响应,对位于青藏高原东北部的冷龙岭3400—4200 m之间5个不同海拔高度的矮火绒草(Leontopodium nanum)叶片进行取样,采用常规石蜡制片技术和显微观察方法测定叶片外部形态、表皮气孔特征和解剖结构,探讨其叶片性状随海拔的变化,结果表明：（1）随海拔高度升高,叶面积呈减小的趋势,而比叶重和叶干物质含量增加;（2）叶片下表皮气孔密度随海拔升高呈先增加后下降的趋势,且气孔密度、气孔器面积、长度、宽度和潜在气孔导度指数等气孔特征之间存在显著相关性;（3）叶厚、栅栏组织和海绵组织厚度随海拔升高呈显著增厚的趋势;（4）叶片解剖结构可塑性和相关性分析显示,上、下角质层厚度的可塑性指数最大,而部分解剖结构指标间存在极显著的相关性。研究表明,矮火绒草为适应沿海拔上升温度降低的环境,主要采取叶片变小、变厚的对策,使植物趋于保温、保水和抗机械损伤的方向发展,并将资源最大化地投入到自身生长发育中。     

岷江上游干旱河谷海拔梯度上四川黄栌叶片特征及其与环境因子的关系

研究了岷江上游干旱河谷海拔梯度上四川黄栌（Cotinus coggygria）叶片特征及变化特点,以及它们与环境变化的相互关系。测定指标有叶片形念结构（叶长、叶宽、叶面积和叶长／叶宽）,表皮特征（气孔器密度和面积、孔径、表皮细胞密度和面积）以及生态特征（生物量、比叶最、饱和含水量）。结果表明。随着海拔升高叶长,叶宽、叶面积、生物量、饱和含水量以及表皮细胞面积呈增大趋势;而表皮细胞密度呈减小趋势;气孔器面积和比叶重在海拔1950m以下,随着海拔上升而增加。1950m的区域明显减小,气孔器密度的变化趋势与之相反;孔径和叶长／叶宽无明显变化。多元统计分析显示,叶面积、饱和含水世和气孔器面积主要与温和度有关,三者随着海拔升高和温和度降低呈增大趋势;随着水热综合因子的增加,气孔器密度减小而孔径和比叶重增大;随着年降水量的增加生物量和气孔器面积基增大趋势,面表皮细胞密度减小;表皮细胞面积随着生物温度降低而增加。综合分析表明,岷江上游干旱河谷缺水和生长季节高温是影响叶片特征变化的主要因素。     

15种蹄盖蕨科植物叶表皮形态特征的研究

利用光学显微镜对蹄盖蕨科15种植物的叶表皮形态特征进行观察和研究。结果表明:蹄盖蕨科15种植物的上、下表皮细胞均为无规则型,垂周壁为凹凸状或深波状;上表皮细胞长宽比在1.0~3.2之间,下表皮细胞长宽比在1.0~2.6之间;在这15种植物中共观察到6种气孔器类型,即极细胞型、腋下细胞型、聚合极细胞型、聚腋下细胞型、无规则四细胞型和无规则型,每种植物具有3~4种气孔器类型,气孔均为下生型,多呈椭圆形。气孔的长宽比在1.3~2.1之间;气孔密度在32~90个/mm2之间;气孔指数为17.7%~40.9%。依据上述叶表皮形态特征将15种蹄盖蕨科植物分为3类,即双盖蕨类、蹄盖蕨类和对囊蕨类。该研究在一定程度上支持Christenhusz分类系统对蹄盖蕨科的划分,为蹄盖蕨科植物的系统分类及演化研究提供基础资料。     

近70年黄土高原3种植物叶片气孔特征参数比较

以黄土高原地区3种典型植物辽东栎(Quercus liaotungensis Koidz.)、虎榛子(Ostryopsis davidiana Decne.)和酸枣[Ziziphusjujuba Mill.var.spinosa(Bunge)Hu ex H.F.Chow]标本为材料,利用数码图像显微处理系统,研究了从20世纪30年代至2002年近70 a中植物叶片气孔长度、宽度、面积与密度的变化状况及其相关性.结果表明,气候变化对3种植物气孔性状并无一致的影响.辽东栎叶片气孔长度、宽度、面积和密度变化总体上均呈上升趋势,升高率分别为11.62%、3.17%、18.01%和1.32%.酸枣叶片气孔长度、宽度和面积呈上升趋势,升高率分别为21.90%、13.60%和35.61%;而气孔密度下降,降低率为-27.86%.虎榛子叶片气孔长度、面积和密度均呈下降趋势,降低率分别为-7.91%、-1.43%和-9.73%;而气孔宽度升高率为2.56%.酸枣叶片气孔4个特征参数的变化率均明显大于辽东栎和虎榛子,虎榛子叶片气孔性状的3个参数(除密度外)变化率最小.3种植物叶片气孔长度的变幅均大于气孔宽度,证实气孔宽度是相对比较稳定的性状.     

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.     

15种铁角蕨科植物叶表皮形态特征的研究

利用光学显微镜对铁角蕨科15种植物的叶表皮形态特征进行观察。研究结果表明：（1）15种铁角蕨科植物的叶上、下表皮细胞形状为不规则型,垂周壁为深波状、波状或浅波状;上表皮细胞长宽比1.3~2.6,下表皮细胞长宽比1.3~4.1;（2）在15种铁角蕨科植物中共观察到7种气孔器类型,分别为腋下细胞型、不等细胞型、无规则四细胞型、无规则型、极细胞型、聚合极细胞型和聚腋下细胞型,每种植物具有2~5种气孔器类型,气孔均为下生型,多为椭圆形;气孔的长宽比1.12~2.81,气孔密度16.4~105.1个·mm^-2,气孔指数为5.7%~21.1%;（3）铁角蕨科植物叶表皮形态特征中的上表皮细胞形状、垂周壁形状、上表皮细胞长宽比、主要气孔器类型及衍生类型等具有一定的属内稳定性,可作为铁角蕨科属间分类的依据之一;（4）该研究在一定程度上支持秦仁昌和吴兆洪对铁角蕨科的划分以及铁角蕨属内分组和组内分系,并为铁角蕨科植物的分类鉴定及系统演化研究提供基础资料。     

田野菟丝子寄生对薇甘菊气孔行为的影响

田野菟丝子寄生已成为当前控制薇甘菊入侵蔓延的最佳生物防治手段之一。应用扫描电子显微镜技术,观察并统计分析了田野菟丝子寄生对薇甘菊叶片气孔形态参数的影响,为了解寄生对薇甘菊生理特征的影响提供一定的形态学依据。结果表明,田野菟丝子寄生对薇甘菊叶片的气孔长宽比和气孔面积产生显著影响:对照薇甘菊叶片的气孔长宽比和气孔面积表现为双峰日变化动态;而被寄生的薇甘菊的叶片气孔长宽比和气孔面积则表现为单峰日变化规律。对照薇甘菊气孔的长宽比分别在12:00和10:00,气孔面积分别在10:00和14:00达到日变化中的最大值与最小值;在15:00到16:00气孔长宽比和气孔面积再一次达到一个峰值,使气孔长宽比和气孔面积的日波动形成一个双峰曲线。被寄生的薇甘菊的叶片气孔长宽比和气孔面积则表现出单峰曲线,峰值分别出现在12:00和14:00。田野菟丝子寄生对薇甘菊的气孔密度没有产生显著影响。     

兰州北山不同坡向刺槐叶脉密度与气孔性状的关联性分析

植物叶脉和气孔性状的关系反映了叶片的水力特性, 对认识它们与植物水分利用有关的生理功能间的关系及其调控作用具有重要意义。该文利用GIS (geographic information system)与实验生态学相结合的方法, 采用标准化主轴估计方法, 研究了兰州市北山不同坡向人工林刺槐(Robinia pseudoacacia)叶脉密度与气孔密度、气孔大小的关系。结果表明: 随着坡向由南坡向东坡、西坡和北坡转变, 植被群落的郁闭度、高度和土壤含水量呈逐渐增加的趋势, 刺槐的净光合速率(P_n)、蒸腾速率(T_r)、光合有效辐射(PAR)、叶脉密度和气孔密度呈逐渐减小的趋势, 气孔与叶面积呈逐渐增大的趋势; 各个坡向的刺槐叶脉密度与气孔密度呈显著正相关关系, 与气孔大小呈显著负相关关系, 且在南坡达到极显著相关关系。生长在南坡的刺槐具有高的叶脉密度和密而小的气孔, 生长在北坡的刺槐具有低的叶脉密度和疏而大的气孔。不同坡向刺槐叶脉密度与气孔特征间的资源分配模式, 反映了植物在异质性生境中根据其功能需求在自身性状之间进行投资权衡机制的优化。     

山东银莲花叶片形态结构对异质生境和海拔变化的响应

山东银莲花为一分布极其狭域的稀有物种,对海拔600 m以上的针阔混交林和山顶灌丛两种异质的生境都具有较高的适应性。为探索其适应策略,选择两种异质生境中的5个海拔梯度样带,采用常规石蜡切片法和显微观察技术,对叶片进行观察、分析和测量,通过比较叶片外部形态特征参数和内部解剖结构的差异,推测其叶片适应海拔和异质生境的响应策略。结果表明:为适应阴暗、潮湿的针阔混交林和干旱、强光照的山顶灌丛两种不同环境,山东银莲花分别表现出不同的适应策略。针阔混交林下,叶片的背腹表皮毛密度、比叶面积和气孔相对开度较山顶灌丛的大,而气孔密度、叶片厚度、栅栏组织和海绵组织的厚度较山顶灌丛的小;山顶灌丛植株叶片栅栏组织细胞排列较林下更加整齐紧密。两种生境中叶片腹面表皮毛的长度、气孔相对开度都随海拔的升高而减小,且差异明显;而叶片厚度、比叶面积、气孔指数等对600 m以上海拔变化未表现出明显的规律性。本研究将为山东银莲花的保护和利用提供理论基础及依据,为其他植物的相关研究提供参考。     

盾叶薯蓣四倍体与二倍体叶表皮及气孔器显微结构的比较

对盾叶薯蓣四倍体和二倍体阴叶和阳叶表皮及气孔器的显微形态结构进行了比较。结果显示:叶片表皮细胞及气孔器的形态结构因染色体倍性及光照的不同而存在明显差异。表皮细胞有规则多边形与不规则细胞壁内褶2种类型;气孔器为典型的双子叶植物无规则型气孔器,仅分布在下表皮。四倍体表皮细胞密度、气孔器密度及气孔指数平均值分别为476.28±6.87个.mm-2、78.22±3.1个.mm-2、14.11,较二倍体的分别小4.60%、17.95%和11.98%。前者气孔器长、宽及保卫细胞宽度平均值分别为32.78±2.09μm、26.07±1.55μm、9.63±1.14μm,较后者的分别大10.73%、3.90%和18.01%,差异极显著。前者阴叶的较其阳叶的分别大5.77%、6.00%、8.72%;后者的则分别大1.72%、1.74%、2.41%。前者叶片保卫细胞中的叶绿体数目为23.93±3.19个,较后者的多74.00%。表皮细胞及气孔器密度、气孔指数,气孔器大小,保卫细胞叶绿体数目、尤其是保卫细胞宽度,可作为倍性鉴定的参考依据。四倍体高产有其良好的叶片结构基础。     

太白山栎属树种气孔特征沿海拔梯度的变化规律

气孔是植物与外界环境进行水分和气体交换的主要通道,调节植物碳同化和水分散失的平衡关系,在一定程度上反映植物对外界环境变化的适应。沿太白山北坡1100—2300 m海拔,测定4种栎属树种的气孔性状,分析气孔性状沿海拔的变化规律和其对环境因子的响应。结果表明:(1)气孔密度与气孔长度间的负相关在4个树种间均显著存在(P0.05);除栓皮栎(Quercus variabilis)外,气孔密度与潜在气孔导度指数的正相关关系均达显著水平;而气孔宽度与气孔长度之间只在栓皮栎和锐齿栎(Q. aliena var. acuteserrata)达到显著水平。(2)栓皮栎和槲栎(Q. aliena)的气孔长度和宽度随海拔升高而下降,气孔密度、潜在气孔导度指数增加,辽东栎(Q. wutaishansea)变化形式则相反;锐齿栎气孔宽度减小,其余性状沿海拔呈单峰变化,在约1600 m处气孔长度达到最小值,气孔密度和潜在气孔导度指数达到最大值。(3)与土壤因子相比,气孔性状主要受气候因素的影响。潜在气孔导度指数与大气温度、空气湿度成极显著正相关(P0.01),与降水量显著负相关(P0.05)。其中,空气相对湿度是影响潜在气孔导度指数的主要因素,能够解释气孔变异的22.9%。本研究结果对于深入认识秦岭太白山地区栎属树种对环境变化的响应和适应提供理论证据。     

不同海拔高度泡沙参叶片形态研究

对不同海拔高度的泡沙参叶表形态、气孔特征进行了观察,比较了不同海拔高度对泡沙参叶片生态适应的影响。结果表明：泡沙参叶表皮角质层较厚,被有大量表皮毛和蜡质颗粒,海拔越高越明显;泡沙参气孔复体类型均为不规则型,气孔保卫细胞角质层较厚,气孔口较小,气孔密度与气孔大小成反比,并且气孔大小随海拔升高而减小,气孔密度随海拔升高而增大,是一种抗旱适应的表现。     

五个猕猴桃种的皮孔、气孔器和叶片下表皮微观特征

为了探讨不同种类猕猴桃皮孔、气孔器和叶片下表皮特征的差异及其分类学意义,该文利用光学显微镜观察五个猕猴桃种共计9份样品材料的一年生枝条韧皮部上皮孔的形态,结果发现供试猕猴桃皮孔呈长椭圆形或长梭形,不同种皮孔的长×宽、皮孔密度、皮孔面积和皮孔面积的百分比值存在差异但与种类划分无明显规律,皮孔的宽可明显区分所选4个中华猕猴...     

Correlation analysis between vein density and stomatal traits of Robinia pseudoacacia in different aspects of Beishan Mountain in Lanzhou

AimsThe relationship between leaf venation and stomatal traits reflects leaf water potential of plants, and is fundamental to probe the relationship between physiological functions and water use. Our objective was to study the changes in the relationship between venation density and stomatal traits (stomatal density and stomatal length) in Robinia pseudoacacia with the changes in slope aspects.MethodsIn Beishan Mountain of Lanzhou in Gansu Province, China, 20 transects were laid out horizontally along the contour at intervals of 50 m from an elevation of 1550-1750 m in four different slope aspects, and 12 plots were set up along each transect at intervals of 5 m. A handed GPS (global positioning system) was used to measure latitude, longitude and altitude at each plot. Community characteristics, including crown density, tree height and soil moisture, were investigated. Robinia pseudoacacia within all plots were sampled and used for measurements of individual leaf area, venation density, stomatal density and stomatal length in laboratory in each plot. The 240 plots were categorized into groups of southern, eastern, western and northern aspects, and the standardized major axis (SMA) estimation method was then used to examine the relationship between the stomatal density, stomatal length and vein density.Important findings The results showed that with a change of the aspect from south to east, west, and north, the crown density, average tree height and soil moisture of the plant community increased, while leaf net photosynthetic rate (P_n), transpiration rate (T_r), photosynthetically active radiation (PAR), vein density and stomatal density decreased. In addition, stomatal length and individual leaf area increased. Venation density of each aspect of R. pseudoacacia was positively correlated with stomatal density (p < 0.05) and negatively with stomatal length (p < 0.05). These correlations were strongest in south slope (p < 0.01).     

Leaf Factors Affecting Light-Saturated Photosynthesis in Ecotypes of Solidago virgaurea from Exposed and Shaded Habitats

Plants of Solidago virgaurea L. from exposed and shaded habitats differ with respect to the response of the photosynthetic apparatus to the level of irradiance during growth. An analysis was carried out on leaf characteristies which might be responsible for the differences established in the rates of Hght-saturated CO₂ uptake. The clones were grown in controlled environment chambers at high and low levels of irradiance. Light-saturated rates of photosynthesis and transpiration were measured at natural and lower ambient CO₂ concentrations. A low temperature dependence of light-saturated CO₂ uptake at natural CO₂ concentrations, and a strong response to changes in stomatal width, suggested that the rate of CO₂ transfer from ambient air towards reaetion sites in chloroplasts was mainly limiting the pholosynthetic rate. Resistances to transfer of CO₂ for different parts of the pathway were calculated. There was a weak but significant correlation between stomatal conductance and the product stomatal frequency ± pore length. Mesopbyll conductance and dry weight per unit area were highly correlated in leaves not damaged by high irradiance. This suggests that mesophyll conductance increases with increasing cross sectional area (per unit leaf area) of the pathways of CO₂ transfer in the mesophyll from cell surfaces to reaction sites. The higher light-saturated photosynthesis in clones from exposed habitats when grown at high irradiance than when grown at low irradiance was attributable mainly to a lower mesophyll resistance. In shade clones the effect upon CO₂ uptake of the increase in leaf thickness when grown at high irradiance was counteracted by the associated inactivation of the photosynthetic apparatus. The difference in CO₂ uptake present between clones from exposed and shaded habitats when preconditioned to high irradiance resulted from differences in both mesophyll and stomatal resistances. A few hybrid clones of an F₁-population from a cross between a clone from an exposed habitat and a clone from a shaded habitat reacted, on the whole, in the same way as the exposed habitat parent. When grown at high irradiance, the hybrid clones showed higher photosynthetic rates than either parent; this was largely attributable to the unusually low stomatal resistance of the hybrid leaves.     

Ecophysiological studies on the shrub Vaccinium myrtillus L. taken from a wide altitudinal range

Summary Observations have been made on the gas exchange and morphology of Vaccinium myrtillus taken from altitudes of 200 m, 610 m and 1,100 m along an altitudinal gradient in central Scotland. Under saturating irradiance, optimum temperatures and a range of vapour pressure deficits, photosynthetic rate and stomatal conductance increased with the altitude of origin of the populations. Correlated with these increases was an increase in the adaxial stomatal density with altitude. This response to altitude could be simulated in controlled conditions, by growing plants in a CO₂ concentration below ambient, similar to that expected at altitude.Plant height decreased with altitude, a feature which was maintained in cultivation. Stem rigidity declined with altitude, in a manner which is predicted to limit the reproductive capacity of the population from 1,100 m in high wind speeds.Total leaf nitrogen increased with altitude. The nitrogen economy of the shoot is discussed in terms of nitrogen availability for stems and leaves and its control over maximum rates of photosynthesis, competitive ability and reproductive capacity.     

Morphological and stomatal responses of Norway spruce foliage to irradiance within a canopy depending on shoot age

Morphological and stomatal responses of Norway spruce (Picea abies) foliage to light availability were studied in respect to shoot age. Needle minor diameter (D(1), anatomical width), major diameter (D(2), anatomical thickness), dry weight (M), and tissue density index (I(D)) increased, and needle flatness (Fl) and specific leaf area (SLA) decreased with foliage age, while shade foliage demonstrated higher morphological plasticity as compared to sun foliage. Needle minor diameter, dry weight, and the ratio of total to projected leaf area increased, and needle flatness and specific leaf area decreased with daily average photosynthetic photon flux density (Q(D)). The current-year foliage exhibited the highest variation with irradiance, while the morphological plasticity decreased with needle ageing. The morphological characteristics of needles were independent of irradiance if Q(D) was above 300 μmol m(-2) s(-1). D(1) was the only linear needle characteristic which significantly changed with light availability within a canopy, and thus determined needle flatness, SLA, as well as the ratio of total to projected leaf area (TLA/PLA). Needle flatness was a characteristic responding most sensitively to the photosynthetic photon flux density, R(2) was 0.68, 0.44, and 0.49 for the current-year, 1-year-old, and 2-year-old foliage, respectively. TLA/PLA ranged from 2.2 to 4.0 depending on D(1). Variation in SLA in response to light availability can be attributed to changes both in needle shape and tissue density. Stomatal responses to photosynthetic photon flux density (Q(P)) depended on foliage type (sun or shade) and age. Sun needles demonstrated higher daily maximum leaf conductances to water vapour compared to shade needles. The shade needles responded more sensitively to changes in Q(P) at dawn and sunset than the sun needles, while older needles of both foliage types exhibited faster stomatal responses. The light-saturation of leaf conductance (g(L)) was achieved by 20 μmol m(-2) s(-1) for shade foliage, and approximately by 50 μmol m(-2) s(-1) for sun foliage. As a rule, g(L) changed in response to irradiance faster in the evening, i.e. at decreasing irradiance. Stomata were not usually completely closed in the dark before sunrise and after sunset, the phenomenon being more pronounced in older shoots and sun needles. Nightly water losses from spruce foliage are attributable primarily to older shoots, and are related to age-dependent changes in stomatal responsiveness.     

Stomatal Morphology during Acclimatization of Tobacco Plantlets to ex vitro Conditions

Image analysis was used in studying stomatal morphology during acclimatization of tobacco plantlets to ex vitro conditions, 45 d after transfer leaf area was 15 times, and total number of stomata per leaf four times increased. During acclimatization stomatal density was decreased considerably on both leaf sides, and was compensated by an increase in stomatal sizes, e.g., in stomatal length and in stomatal area (both guard cells and pore). Elongation of stomata was increased indicating that the originally circular stomata of in vitro plantlets were changed into elliptical ones in ex vitro acclimatized plants. This revised version was published online in June 2006 with corrections to the Cover Date.     

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.