Leaf morphology of 40 evergreen and deciduous broadleaved subtropical tree species and relationships to functional ecophysiological traits

We explored potential of morphological and anatomical leaf traits for predicting ecophysiological key functions in subtropical trees. We asked whether the ecophysiological parameters stomatal conductance and xylem cavitation vulnerability could be predicted from microscopy leaf traits. We investigated 21 deciduous and 19 evergreen subtropical tree species, using individuals of the same age and from the same environment in the Biodiversity‐Ecosystem Functioning experiment at Jiangxi (BEF‐China). Information‐theoretic linear model selection was used to identify the best combination of morphological and anatomical predictors for ecophysiological functions. Leaf anatomy and morphology strongly depended on leaf habit. Evergreen species tended to have thicker leaves, thicker spongy and palisade mesophyll, more palisade mesophyll layers and a thicker subepidermis. Over 50% of all evergreen species had leaves with multi‐layered palisade parenchyma, while only one deciduous species (Koelreuteria bipinnata) had this. Interactions with leaf habit were also included in best multi‐predictor models for stomatal conductance (g_s) and xylem cavitation vulnerability. In addition, maximum g_s was positively related to log ratio of palisade to spongy mesophyll thickness. Vapour pressure deficit (vpd) for maximum g_s increased with the log ratio of palisade to spongy mesophyll thickness in species having leaves with papillae. In contrast, maximum specific hydraulic conductivity and xylem pressure at which 50% loss of maximum specific xylem hydraulic conductivity occurred (Ψ₅₀) were best predicted by leaf habit and density of spongy parenchyma. Evergreen species had lower Ψ₅₀ values and lower maximum xylem hydraulic conductivities. As hydraulic leaf and wood characteristics were reflected in structural leaf traits, there is high potential for identifying further linkages between morphological and anatomical leaf traits and ecophysiological responses.     

Photosynthetic and anatomic responses of peanut leaves to cadmium stress

Cadmium (Cd) treatments caused an inhibition in the net photosynthetic rate (P _N) of peanut (Arachis hypogaea) plants, due to the reduction of stomatal conductance (g _s) and photosynthetic pigment contents, as well as the alteration in leaf structure. The decrease of the transpiration rate and g _s might result from the Cd-induced xerophyte anatomic features of leaves (i.e. thick lamina, upper epidermis, palisade mesophyll, high palisade to spongy thickness ratio, as well as abundant and small stomata). The decline of P _N was independent of the impairment in photosystem 2.     

Variability of stomatal conductance, leaf anatomy, and seasonal leaf wettability of young and adult European beech leaves along a vertical canopy gradient

This study assessed the variation of leaf anatomy, chlorophyll content index (CCI), maximal stomatal conductance (g _s ^max ) and leaf wettability within the canopy of an adult European beech tree (Fagus sylvatica L.) and for beech saplings placed along the vertical gradient in the canopy. At the top canopy level (CL_28m) of the adult beech, CCI and leaf anatomy reflected higher light stress, while g _s ^max increased with height, reflecting the importance of gas exchange in the upper canopy layer. Leaf wettability, measured as drop contact angle, decreased from 85.5°?±?1.6° (summer) to 57.5°?±?2.8° (autumn) at CL_28m of the adult tree. At CL_22m, adult beech leaves seemed to be better optimized for photosynthesis than the CL_28m leaves because of a large leaf thickness with less protective and impregnated substances, and a higher CCI. The beech saplings, in contrast, did not adapt their stomatal characteristics and leaf anatomy according to the same strategy as the adult beech leaves. Consequently, care is needed when scaling up experimental results from seedlings to adult trees.     

Effects of reduced irradiance on leaf morphology,photosynthetic capacity,and fruit yield in olive (<Emphasis Type="Italic">Olea europaea</Emphasis> L.)

One-year-old olive trees (cv. Koroneiki) were grown in plastic containers of 50 000 cm³ under full daylight and 30, 60, and 90 % shade for two years. The effects of shade on leaf morphology and anatomy, including stomatal density and chloroplast structure, net photosynthetic rate (P _N), stomatal conductance (g _s), and fruit yield were studied. Shade reduced leaf thickness due to the presence of only 1–2 palisade layers and reduced the length of palisade cells and spongy parenchyma. The number of thylakoids in grana as well as in stroma increased as shade increased, while the number of plastoglobuli decreased in proportion to the reduced photosynthetically active radiation (PAR). The higher the level of shade, the lower the stomatal and trichome density, leaf mass per area (ALM), g _s, and P _N. Shade of 30, 60, and 90 % reduced stomatal density by 7, 16, and 27 %, respectively, while the corresponding reduction in P _N was 21, 35, and 67 %. In contrast, chlorophyll a+b per fresh mass, and leaf width, length, and particularly area increased under the same shade levels (by 16, 33, and 81 % in leaf area). P _N reduction was due both to a decrease in PAR and to the morphological changes in leaves. The effect of shade was more severe on fruit yield per tree (32, 67, and 84 %) than on P _N indicating an effect on bud differentiation and fruit set. The olive tree adapts well to shade compared with other fruit trees by a small reduction in stomatal and trichome density, palisade parenchyma, and a significant increase in leaf area.     

LEAF ANATOMY OF TISSUE-CULTURED LIQUIDAMBAR STYRACIFLUA (HAMAMELIDACEAE) DURING ACCLIMATIZATION

Structural changes accompanying the acclimation process were observed in leaves of sweetgum, Liquidambar styraciflua, using light and transmission electron microscopy (TEM). Comparisons were made of leaves obtained from tissue culture, plantlets acclimated after transfer from the in vitro environment to soil, and field grown trees. Leaves of cultured plantlets lacked a differentiated palisade parenchyma and had spongy parenchyma interspersed with large air spaces. Field grown leaves showed distinct palisade and spongy tissues and a high cell density. New leaves from acclimated plantlets showed an elongation of the upper mesophyll with fewer intercellular spaces than cultured plants. Cells from leaves from in vitro plantlets had large vacuoles, limited cytoplasmic content and flattened chloroplast with an irregularly arranged internal membrane system. Acclimated and field leaf cells had a greater cytoplasmic content than cultured leaves, with the former having more dominate vacuoles. Chloroplasts had evident grana. Acclimated and field leaves had a well developed cuticle unlike leaves from culture.     

枫香变红过程中叶片组织结构、光合特性及色素含量变化研究

枫香(Liquidambar formosana)因其叶片入秋后逐渐变红而极具观赏价值,是优良的景观生态树种。为了解枫香叶片结构变化与叶色的关系,该文通过连续监测枫香叶片变红过程中组织结构、光合特性及色素含量的变化,分析叶片结构与其光合特性和色素的关系。结果表明：(1)叶片变色过程中,表皮细胞均为椭圆形,紧密排列,未观察到明显的细胞变异,表面未附着绒毛和蜡质,且上表皮细胞与栅栏组织细胞间排列紧密,未出现较大的气室。(2)随着叶片逐渐变红,叶片结构变化显著,其中叶片、上表皮、栅栏组织和海绵组织厚度及气孔开度均逐渐减小,而气孔器长和宽、单个气孔器面积则逐渐增大。(3)随着叶片结构的变化,其叶绿素含量逐渐减少,致使净光合速率逐渐减小,在出现光破坏时,叶片通过在栅栏组织细胞液泡内合成花色苷来自我保护,而大量的花色苷致使叶片表面呈现红色。综上认为,叶绿素含量降低,花色素苷大量积累是导致枫香叶片变红的直接原因,而枫香叶色变红则是其一系列生理结构特征综合作用的结果。     

Leaf plasticity in peanut (Arachis hypogaea L.) in response to heavy metal stress

Phenotypic plasticity in morphological, anatomical and physiological traits of peanut (Arachis hypogaea L.) leaves was tested at four different concentrations of Cd, Cu and Zn under greenhouse conditions. Among 18 characteristics tested, nine were found to be the most sensitive and demonstrate the greatest phenotypic plasticity. These were: the leaf area (LA), the leaf mass per area (LMA), chlorophyll a concentration (Chl a), chlorophyll b concentration (Chl b), total chlorophyll concentration (Chl t), the effective quantum yield of photosystem II (ΦPS II), stomatal density of upper epidermis (SDU), palisade thickness (PT), and palisade to spongy thickness ratio (P/S). The plasticity of chlorophyll concentration and fluorescence parameters may be maladaptive and reflects metal toxicity to leaves, whereas the anatomical plasticity is adaptive, indicative of a tradeoff between the physiological and anatomic plasticity. The anatomical plasticity resulted in a xerophyte feature of leaves (i.e. small leaflets, thick lamina, upper epidermis, palisade mesophyll, as well as abundant and small stomata), which enhanced the capacity to resist drought caused by heavy metals via a decrease in root growth.     

Stomatal activity within the crowns of tall deciduous trees under forest conditions

The variation in stomatal activity within the crowns ofAcer campestre, Carpinus betulus andQuercus cerris was measured by vapour exchange porometer on several summer days in an oak-hornbeam forest, in SW Slovakia, Czechoslovakia. Variation resulted from crown position in the forest stand and from leaf position within the canopy. The highest stomatal conductance was in sunlit sun leaves in the upper part of the canopy. Stomatal conductance decreased with increasing depth in the canopy. The steepest decrease was in the upper canopy, in the intermediate zone between fully sunlit and fully shaded leaves, and was caused by the decline in leaf irradiance and in stomatal density. In codominant trees, the conductance in shade leaves at the base of the crown was significantly lower than in the sun leaves at the top of the crown. In a dominant tree,Q. cerris, the differences in stomatal conductance were small and most frequently insignificant. Variation in incident light also determined the diurnal variation of stomatal conductance with respect to crown aspect. Differences between sun leaves on the east and west facing aspects of the overstory crown ofQ. cerris were demonstrated for several days.     

Developmental and physiological correlates of leaf size in Hyeronima alchorneoides (Euphorbiaceae)

The tropical emergent tree Hyeronima alchorneoides has large decreases in leaf size with tree age: 1200 cm(2) at 1 yr, 900 cm(2) at 3 yr, 200 cm(2) at 11 yr, and 80 cm(2) in old (>30 yr) individuals. We tracked leaf growth and physiological attributes on trees of three different ages (1, 3, and 11 yr) to determine the developmental basis and functional consequences of this variation. Leaves on young trees grew faster and sustained maximum rates of leaf expansion longer than leaves on older trees. Leaf mass per area (LMA) did not differ among age classes. Maximum photosynthetic rates reflected differences in leaf nitrogen concentration, in which leaves from the lower crown of younger trees outperformed those at a comparable crown position in older trees. One-year-old trees had the lowest stomatal conductance and the greatest instantaneous water use efficiency. Ontogenetic plasticity in mature leaf size, structure, and physiology may be a balance between the advantages conferred by rapid height growth when trees are young and the benefits derived from producing branches that increase light harvesting ability as trees reach the canopy.     

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。     

Elevated carbon dioxide does not affect average canopy stomatal conductance of Pinus taeda L.

While photosynthetic responses of C₃ plants to elevated CO₂ are fairly well documented, whole-plant water use under such conditions has been less intensively studied. Woody species, in particular, have exhibited highly variable stomatal responses to high CO₂ as determined by leaf-level measurements. In this study, sap flux of Pinus taeda L. saplings was periodically monitored during the 4th year of an open-top chamber CO₂ fumigation experiment. Water use per unit sapwood area did not differ between treatments. Furthermore, the ratio of leaf area to sapwood area did not change under high CO₂, so that average canopy stomatal conductance (on a unit leaf area basis) remained unaffected by the CO₂ treatment. Thus, the only effect of high CO₂ was to increase whole-plant water use by increasing sapling leaf area and associated conducting sapwood area. Such an effect may not directly translate to forest-level responses as the feedback effects of higher leaf area at the canopy scale cannot be incorporated in a chamber study. These feedbacks include the potential effect of higher leaf area index on rainfall and light interception, both of which may reduce average stomatal conductance in intact forest canopies. Received: 13 March 1998 / Accepted: 8 July 1998     

Structural Adaptation of the Leaf Mesophyll to Shading

Structural characteristics of the mesophyll were studied in five boreal grass species experiencing a wide range of light and water supply conditions. Quantitative indices of the palisade and spongy mesophyll tissues (cell and chloroplast sizes, the number of chloroplasts per cell, the total cell and chloroplast surface area per unit leaf surface area) were determined in leaves of each of the species. The cell surface area and the cell volume in spongy mesophyll were determined with a novel method based on stereological analysis of cell projections. An important role of spongy parenchyma in the photosynthetic apparatus was demonstrated. In leaves of the species studied, the spongy parenchyma constituted about 50% of the total volume and 40% of the total surface area of mesophyll cells. The proportion of the palisade to spongy mesophyll tissues varied with plant species and growth conditions. In a xerophyte Genista tinctoria, the total cell volume, cell abundance, and the total surface area of cells and chloroplasts were 30–40% larger in the palisade than in the spongy mesophyll. In contrast, in a shade-loving species Veronica chamaedris, the spongy mesophyll was 1.5–2 times more developed than the palisade mesophyll. In mesophyte species grown under high light conditions, the cell abundance and the total cell surface area were 10–20% greater in the palisade mesophyll than in the spongy parenchyma. In shaded habitats, these indices were similar in the palisade and spongy mesophyll or were 10–20% lower in the palisade mesophyll. In mesophytes, CO₂ conductance of the spongy mesophyll accounted for about 50% of the total mesophyll conductance, as calculated from the structural characteristics, with the mesophyll CO₂ conductance increasing with leaf shading.     

青藏高原不同海拔3种菊科植物叶片结构变化及其生态适应性

采用常规石蜡制片技术和显微观察方法,对分布于青藏高原祁连山东部山地冷龙岭3600—4400 m不同海拔高度的蒲公英(Taraxacum mongolicum Hand-Mazz)、火绒草(Leontopodium leontopodioides(Willd.)Beauv)和美丽风毛菊(Saussurea superba Anthony)的叶片形态解剖结构进行了研究,并探讨了其对海拔高度的响应及生态适应性。结果表明:随海拔高度的升高,3种植物气孔器外拱盖内缘、角质层纹饰、气孔与表皮细胞的位置关系以及上、下表皮气孔器内缘呈现不同的变化趋势;火绒草上、下表皮气孔密度随海拔升高而增加,而蒲公英和美丽风毛菊的气孔密度则降低;3种植物上、下表皮气孔指数随海拔高度的增加均未出现规律性变化;3种植物叶片厚度、上下表皮厚度、上下角质层厚度、栅栏细胞系数均随海拔升高而增加;解剖学指标之间大多呈明显的协同进化;叶片结构,尤其是气孔密度对海拔高度变化表现出较大的可塑性。研究表明3种植物采取不同的响应机制来适应海拔高度的变化,植物对高原环境变化的适应具有多样性。     

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。     

Leaf responses to iron nutrition and low cadmium in peanut: anatomical properties in relation to gas exchange

Aims

This study evaluated how iron nutrition affect leaf anatomical and photosynthetic responses to low cadmium and its accumulation in peanut plants.

Methods

Seedlings were treated with Cd (0 and 0.2 μM CdCl₂) and Fe (0, 10, 25, 50 or 100 μM EDTA-Na₂Fe) in hydroponic culture.

Results

Cadmium accumulation is highest in Fe-deficient plants, and dramatically decreased with increasing Fe supply. The biomass, gas exchange, and reflectance indices were highest at 25 μM Fe²⁺ treatments, indicating the concentration is favorable for the growth of peanut plants. Both Fe deficiency and Cd exposure impair photosynthesis and reduce reflectance indices. However, they show different effects on leaf anatomical traits. Fe deficiency induces more and smaller stomata in the leaf surface, but does not affect the inner structure. Low Cd results in a thicker lamina with smaller stomata, thicker palisade and spongy tissues, and lower palisade to spongy thickness ratio. The stomatal length and length/width ratio in the upper epidermis, spongy tissue thickness, and palisade to spongy thickness ratio were closely correlated with net photosynthetic rate, stomatal conductance, and transpiration rate.

Conclusions

Cd accumulation rather than Fe deficiency alters leaf anatomy that may increase water use efficiency but inhibit photosynthesis.     

Photosynthesis in an Australian rainforest tree,Argyrodendron peralatum,during the rapid development and relief of water deficits in the dry season

Summary Rates of apparent photosynthesis were measured in situ at five positions between the upper crown and a lower branch of a 34 m tall Argyrodendron peralatum (F.M. Bailey) H.L. Edlin ex I.H. Boas tree, and on an understorey sapling of the same species growing in a northern Australian rainforest. At the end of the dry season, rapid reductions in photosynthetic rates occurred in the upper crown within three days after a rain event, but changes in the lower crown and the sapling were less marked. Complete recovery of photosynthesis followed a second rain event. At high photon flux densities, stomatal conductance to water vapour decreased in a curvilinear fashion as the vapour pressure difference between leaf and air increased. Apparent photosynthesis was linearly related to stomatal conductance on the first clear day after each rain event, but there was no relationship between these parameters at the end of a brief natural drying cycle. Under conditions of adequate water supply, stomatal conductances of both upper crown and understorey leaves increased linearly with increasing photon flux density up to about 300 mol m^-2 s^-1. During water deficits, stomatal conductances in leaves from the understorey increased much more rapidly at very low photon flux densities than did conductances in leaves from the upper canopy.     

遮荫对南美蟛蜞菊、蟛蜞菊及其自然杂交种叶片显微结构及叶绿体超微结构的影响

利用JSM-6360LV型扫描电镜和JEM-1010型透射电镜,观察了南美蟛蜞菊、蟛蜞菊及其自然杂交种新近成熟和老熟叶片的解剖结构及叶绿体超微结构。结果表明:遮荫后该杂交种与其亲本新近成熟叶片均表现为上下表皮气孔密度、叶片总厚度及上下表皮厚度、栅栏组织、海绵组织厚度减小,叶绿体肿胀变形,基粒片层垛叠程度增加,淀粉粒增多变大;遮荫后杂交种老熟叶片总厚度及上表皮、栅栏组织、海绵组织厚度增加,入侵种的下表皮厚度及本地种的上表皮厚度增加,叶绿体超微结构在遮荫后均出现严重损伤,基粒片层类囊体结构边缘溶解等。说明三种蟛蜞菊属物种及各物种不同叶龄叶片对弱光条件的响应存在差异;杂交种叶片显微及超微结构在不同光照下的变化介于亲本之间,对遮荫环境能较好适应。     

Changes in stomatal frequency, stomatal conductance and cuticle thickness during leaf expansion in the broad-leaved evergreen species, Eucalyptus regnans

Aspects of leaf anatomical and physiological development were investigated in the broad-leaved evergreen species, Eucalyptus regnans F.Muell. Newly emergent leaves were tagged in the field and measured for stomatal conductance while a subset was collected every 14 days for the measurement of stomata and cuticle over a 113-day period. Cuticle thickness increased during leaf expansion, the increase following a sigmoid curve. Stomatal frequency (no. mm⁻²) decreased from 56 to 113 days after leaf emergence. The frequency of both immature and intermediate developmental stages of stomata also decreased over this time, but the total number of stomata per leaf remained relatively constant. Stomatal conductance (g _s) of young expanding leaves increased during expansion, and was significantly linearly correlated with stomatal frequency (excluding immature stomata), and with cuticle thickness. The progressive increase in g _s in young developing leaves was contrary to the observed changes in structural characteristics (increased cuticle thickness and decreased stomatal frequency). This increase in g _s with development may be related to the progressive increase in number of mature stomata with larger apertures and, therefore, a higher total pore area in fully expanded leaves.     

Regulation of water flux through trunks, branches, and leaves in trees of a lowland tropical forest

We studied regulation of whole-tree water use in individuals of five diverse canopy tree species growing in a Panamanian seasonal forest. A construction crane equipped with a gondola was used to access the upper crowns and points along the branches and trunks of the study trees for making concurrent measurements of sap flow at the whole-tree and branch levels, and vapor phase conductances and water status at the leaf level. These measurements were integrated to assess physiological regulation of water use from the whole-tree to the single-leaf scale. Whole-tree water use ranged from 379 kg day⁻¹ in a 35 m-tall Anacardium excelsum tree to 46 kg day⁻¹ in an 18 m-tall Cecropia longipes tree. The dependence of whole-tree and branch sap velocity and sap flow on sapwood area was essentially identical in the five trees studied. However, large differences in transpiration per unit leaf area (E) among individuals and among branches on the same individual were observed. These differences were substantially reduced when E was normalized by the corresponding branch leaf area:sapwood area ratio (LA/SA). Variation in stomatal conductance (g _s) and crown conductance (g _c), a total vapor phase conductance that includes stomatal and boundary layer components, was closely associated with variation in the leaf area-specific total hydraulic conductance of the soil/leaf pathway (G _t). Vapor phase conductance in all five trees responded similarly to variation in G _t. Large diurnal variations in G _t were associated with diurnal variation in exchange of water between the transpiration stream and internal stem storage compartments. Differences in stomatal regulation of transpiration on a leaf area basis appeared to be governed largely by tree size and hydraulic architectural features rather than physiological differences in the responsiveness of stomata. We suggest that reliance on measurements gathered at a single scale or inadequate range of scale may result in misleading conclusions concerning physiological differences in regulation of transpiration. Received: 1 October 1997 / Accepted: 6 March 1998     

岷江上游干旱河谷海拔梯度上白刺花叶片生态解剖特征研究

对岷江上游干旱河谷海拔梯度上（1 650～1 950 m）白刺花(Sophora davidii)叶片进行生态解剖学研究.观测指标包括叶片形态特征（叶长宽比、叶面积、叶片厚度）、解剖结构（表皮厚度、栅栏组织厚度(P)、海绵组织厚度(S)、P/S比值、表皮角质膜厚度）及叶表皮特征（气孔器密度和面积、表皮细胞密度和面积、表皮毛密度和长度）.结果表明,白刺花叶片面积为0.144～0.208 cm²,叶总厚度为171.58～195.83 μm;叶肉组织分化明显,栅栏组织厚度与海绵组织厚度分别为69.83～82.42和62.00～ 80.67 μm,P/S的比值为1.14～1.01,上下表皮厚度分别为14.03～15.33和13.88～16.17 μm,上下角质膜厚度分别为2.66～4.56和2.76～2.02 μm;气孔密度为13.71～15.02个·mm^-2,其面积为249.86～280.43 μm²;表皮细胞密度为160.54～178.43个·mm^-2,其面积为557.43～626.85 μm²;表皮毛长度为186.51～260.99 μm,其密度为18.29～32.27个·mm^-2.随海拔升高叶面积、叶厚度、栅栏组织和海绵组织的厚度、气孔器面积、表皮细胞面积以及表皮毛密度呈增加趋势,而角质膜厚度、表皮细胞密度和表皮毛长度则呈减小趋势;叶长宽比、P/S的比值、表皮厚度与气孔器密度无明显差异.