Leaf architecture and direction of incident light influence mesophyll fluorescence profiles

Light propagation and distribution inside leaves have been recognized as important processes influencing photosynthesis. Monochromatic light absorption across the mesophyll was measured using chlorophyll fluorescence generated from illumination of the cut edge (epi-illumination), as well as the adaxial or abaxial surfaces of the leaf. Species were selected that had basic leaf types: laminar leaf with adaxial palisade layer (Rhododendron catawbiense), needle with palisade (Abies fraseri), and needle without palisade (Picea rubens). Fluorescence was more evenly distributed across the mesophyll for adaxially illuminated leaves with a palisade cell layer, as well as for the needles (cylindrical) without palisade, when compared to fluorescence generated by abaxial illumination. Moreover, fluorescence from green light illumination remained high across the mesophyll of adaxially illuminated R. catawbiense, indicating a possible influence of mesophyll structure on internal light distribution beyond that of chlorophyll levels. These data support the idea that light propagation within the mesophyll is associated with asymmetric mesophyll structure, in particular the presence of palisade cell layers. In addition, we propose that the evolution of a more cylindrical leaf form, such as found in conifer species, may be a structural solution to excessive sunlight that replaces the highly differentiated mesophyll found in most laminar-leaved species.     

Leaf structure and specific leaf mass: the alpine desert plants of the Eastern Pamirs, Tadjikistan

This study examines interrelationships between eight leaf attributes (specific leaf mass, area, dry mass, lamina thickness, mesophyll cell number per cm², mesophyll cell volume, chloroplast volume, and number of chloroplasts per mesophyll cell) in field-grown plants of 94 species from the Eastern Pamir Mountains, at elevations between 3800 and 4750 m. Unlike most other mountain areas, the Eastern Pamirs, Karakorum system, Tadjikistan provide localities where low temperatures and radiation combine with moisture stress at high altitudes. For all the attributes measured, significant differences were found between plants with different mesophyll types. Leaves with dorsiventral palisade structure (dorsal palisade, ventral spongy mesophyll cells) had thicker leaves with larger but fewer mesophyll cells, containing more and larger chloroplasts. These differences in mesophyll type are reflected in differences in the total surface of mesophyll cells per unit leaf area ( A _mes/ A ) or volume ( A _mes/ V ). Plants with isopalisade leaf structure (palisade cells under both dorsal and ventral surfaces) are more commonly xerophytes and their increased values of A _mes/ A and A _mes/ V decrease CO₂ mesophyll resistance, which is an important adaptation to drought. Path analysis shows the critical importance of mesophyll cell volume in leading to the covariance between the different leaf attributes and hence to specific leaf mass (SLM), even though mesophyll cell volume is not itself strongly correlated with SLM. This is because mesophyll cell volume increases SLM through its effects on leaf thickness and chloroplast number per cell, but decreases SLM through its negative effect on mesophyll cell density.     

Tropospheric ozone-induced structural changes in leaf mesophyll cell walls in grapevine plants

The structural changes in leaves of grapevine plants (Vitis vinifera L.) exposed to different ozone concentrations were investigated. Ozone fumigations were performed in open-top chambers at four different ozone levels (charcoal-filtered air (F), ambient air (N), ambient air + 25 mm³m⁻³ ozone (O-25) and ambient air + 50 mm³m⁻³ ozone (O-50)). The leaves of plants from chambers with increased ozone concentrations (O-25 and O-50) were significantly thicker than the controls (F), owing to increased thickness of the mesophyll layer. Observing O-50 leaves, it was found that the mesophyll cell wall displayed structural changes. In some places cell wall thickness increased up to 1 μm. We found callose deposits on the inner side of the cell walls of mesophyll cells. These data are in accord with the concept that the mesophyll cell wall acts as a barrier against the penetration of tropospheric ozone into the cells.     

Sudden Collapse of Vacuoles in Saintpaulia sp. Palisade Cells Induced by a Rapid Temperature Decrease

It is well known that saintpaulia leaf is damaged by the rapid temperature decrease when cold water is irrigated onto the leaf surface. We investigated this temperature sensitivity and the mechanisms of leaf damage in saintpaulia (Saintpaulia sp. cv. ‘Iceberg’) and other Gesneriaceae plants. Saintpaulia leaves were damaged and discolored when subjected to a rapid decrease in temperature, but not when the temperature was decreased gradually. Sensitivity to rapid temperature decrease increased within 10 to 20 min during pre-incubation at higher temperature. Injury was restricted to the palisade mesophyll cells, where there was an obvious change in the color of the chloroplasts. During a rapid temperature decrease, chlorophyll fluorescence monitored by a pulse amplitude modulated fluorometer diminished and did not recover even after rewarming to the initial temperature. Isolated chloroplasts were not directly affected by the rapid temperature decrease. Intracellular pH was monitored with a pH-dependent fluorescent dye. In palisade mesophyll cells damaged by rapid temperature decrease, the cytosolic pH decreased and the vacuolar membrane collapsed soon after a temperature decrease. In isolated chloroplasts, chlorophyll fluorescence declined when the pH of the medium was lowered. These results suggest that a rapid temperature decrease directly or indirectly affects the vacuolar membrane, resulting in a pH change in the cytosol that subsequently affects the chloroplasts in palisade mesophyll cells. We further confirmed that the same physiological damage occurs in other Gesneriaceae plants. These results strongly suggested that the vacuoles of palisade mesophyll cells collapsed during the initial phase of leaf injury.     

青藏高原草地植物叶解剖特征

运用常规石蜡制片技术对我国青藏高原66种草地植物优势种的叶解剖特征进行研究,并分析了叶解剖特征与海拔、生长季降水及生长季均温之间的关系.结果表明:青藏高原草地植物叶片具有很多适应高寒环境的结构特征,如表皮层厚且表皮细胞大小差异显著,表皮毛等表皮附属物发达,异细胞丰富,通气组织普遍发达等;叶片各组成部分厚度的变异程度不同,其中海绵组织厚度变异最大,其次为上角质层、下表皮层、下角质层、上表皮层、栅栏组织,叶片厚度的变异最小;青藏高原草地植物叶片各组成部分的厚度存在协同进化,上下角质层厚度呈强烈正相关,海绵组织厚度与叶片厚度相关性最强;青藏高原草地植物叶片各组成部分的厚度与海拔、生长季降水、生长季均温3个重要环境变量呈较弱的相关性,总体表现为随海拔升高叶片各组成部分的厚度减小,而随生长季降水和生长季均温的增加叶片厚度增加.     

叶片叶肉结构对环境光强的适应及对光合作用的影响

本文利用Kubelka-Munk理论描述了平行光在叶片内的吸收和散射,同时利用叶片分层光合作用非直角双曲线光反应模型,给出了整张叶片光合作用计算式。最后利用优化理论阐明了叶片叶肉分化成光合特性具有明显差异的栅栏组织和海绵组织可能是对叶片内光梯度的一种适应;同时证明了叶片叶肉在一定环境光强下存在一个最佳的栅栏组织和海绵组织比例,并且这个比例随环境光强增大而增大,这最佳比例也受叶肉组织光合特性差异的影响。     

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.     

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.     

Ageing-related Anatomical and Ultrastructural Changes in Leaves of Birch (Betula pendula Roth.) Clones as Affected by Low Ozone Exposure

Effects of ozone on the leaf anatomy and ultrastructure of fivebirch (Betula pendula Roth.) clones were studied during onegrowing season in open-field conditions. Cumulative ozone exposurewas 1·5 times higher than ambient. Ozone exposure decreasedtotal leaf thickness in one, ozone sensitive, clone. The effecton palisade spongy mesophyll thickness was clone-specific, whilethe amount of palisade intercellular space was reduced in allclones. A second effect was a change in the relative amountsof adaxial and abaxial epidermis. In palisade and spongy parenchymacells of all clones, ozone increased the number of irregularand spherical shaped chloroplasts, the electron density of chloroplaststroma, swelling and curling of thylakoids, translucency ofthe mitochondrial matrix and also the amount of cytoplasmiclipids. In the sensitive clone shorter chloroplasts and reducedamount of starch were observed in ozone-exposed plants, whilst,in the tolerant clone, the size of chloroplasts and the amountof starch were unaffected. Ozone effects on number, size andelectron density of plastoglobuli and vacuolar tannin were clone-dependent.At the ultrastructural level, the normal leaf ageing processprogressed at different rates in the birch clones. Ozone acceleratedsenescence-related structural changes, in accordance with earlierobservations of deciduous species.Copyright 1995, 1999 AcademicPress Betula pendula Roth., birch, clones, ageing, ozone, leaf anatomy, ultrastructure     

Fluorescence and Delayed Light Emission from Mesophyll and Bundle Sheath Cells in Leaves of Normal and Salt-Treated Panicum miliaceum

A modified fluorescence microscope system was used to measure chlorophyll fluorescence and delayed light emission from mesophyll and bundle sheath cells in situ in fresh-cut sections from leaves of Panicum miliaceum L. The fluorescence rise in 3-(3,4-dichlorophenyl)-1, 1-dimethylurea (DCMU)-treated leaves and the slow fluorescence kinetics in untreated leaves show that mesophyll chloroplasts have larger photosystem II unit sizes than do bundle sheath chloroplasts. The larger photosystem II units imply more efficient noncyclic electron transport in mesophyll chloroplasts. Quenching of slow fluorescence also differs between the cell types with mesophyll chloroplasts showing complex kinetics and bundle sheath chloroplasts showing a relatively simple decline. Properties of the photosynthetic system were also investigated in leaves from plants grown in soil containing elevated NaCl levels. As judged by changes in both fluorescence kinetics in DCMU-treated leaves and delayed light emission in leaves not exposed to DCMU, salinity altered photosystem II in bundle sheath cells but not in mesophyll cells. This result may indicate different ionic distributions in the two cell types or, alternatively, different responses of the two chloroplast types to environmental change.     

Ozone tolerance in snap bean is associated with elevated ascorbic acid in the leaf apoplast

Ascorbic acid (AA) in the leaf apoplast has the potential to limit ozone injury by participating in reactions that detoxify ozone and reactive oxygen intermediates and thus prevent plasma membrane damage. Genotypes of snap bean ( Phaseolus vulgaris L) were compared in controlled environments and in open-top field chambers to assess the relationship between extracellular AA content and ozone tolerance. Vacuum infiltration methods were employed to separate leaf AA into extracellular and intracellular fractions. For plants grown in controlled environments at low ozone concentration (4 nmol mol⁻¹ ozone), leaf apoplast AA was significantly higher in tolerant genotypes (300–400 nmol g⁻¹ FW) compared with sensitive genotypes (approximately 50 nmol g⁻¹ FW), evidence that ozone tolerance is associated with elevated extracellular AA. For the open top chamber study, plants were grown in pots under charcoal-filtered air (CF) conditions and then either maintained under CF conditions (29 nmol mol⁻¹ ozone) or exposed to elevated ozone (67 nmol mol⁻¹ ozone). Following an 8-day treatment period, leaf apoplast AA was in the range of 100–190 nmol g⁻¹ FW for all genotypes, but no relationship was observed between apoplast AA content and ozone tolerance. The contrasting results in the two studies demonstrated a potential limitation in the interpretation of extracellular AA data. Apoplast AA levels presumably reflect the steady-state condition between supply from the cytoplasm and utilization within the cell wall. The capacity to detoxify ozone in the extracellular space may be underestimated under elevated ozone conditions where the dynamics of AA supply and utilization are not adequately represented by a steady-state measurement.     

Effects of Simulated Acid Rain on Anatomy of Primary Leaves of Phaseolus Vulgaris

Ten-days-old bean plants (Phaseolus vulgaris L., cv. Cheren Starozagorski) were treated with simulated acid rain (pH 2.4, 2.2, 2.0 and 1.8). Anatomical changes in the primary leaves were studied 3, 48 and 168 h after a single treatment. This treatment induced: 1) change in the shape of palisade cells, contraction of their contact surfaces and expansion of spongy cells (pH 1.8, 3 h after treatment); 2) reduction of symplast connections among palisade cells and of apoplast in the spongy mesophyll (pH 1.8, 48 h after treatment); 3) destruction of adaxial epidermis and portions of palisade mesophyll, plasmolysis of spongy cells (pH 1.8, 168 h after treatment); 4) full destruction of mesophyll (pH 2.4, 2.2, 2.0 and 1.8, 168 h after treatment). The structure of abaxial epidermis was more stable than that of the adaxial one. With respect to anatomical parameters the studied species could be considered as comparatively resistant to acid rain.     

Ozone effects on the ultrastructure of peatland plants: Sphagnum mosses, Vaccinium oxycoccus, Andromeda polifolia and Eriophorum vaginatum

BACKGROUND AND AIMS: Ozone effects on peatland vegetation are poorly understood. Since stress responses are often first visible in cell ultrastructure, electron microscopy was used to assess the sensitivity of common peatland plants to elevated ozone concentrations. METHODS: Three moss species (Sphagnum angustifolium, S. magellanicum and S. papillosum), a graminoid (Eriophorum vaginatum) and two dwarf shrubs (Vaccinium oxycoccus and Andromeda polifolia), all growing within an intact canopy on peat monoliths, were exposed to a concentration of 0, 50, 100 or 150 ppb ozone in two separate growth chamber experiments simulating either summer or autumn conditions in central Finland. After a 4- or 5-week-long exposure, samples were photographed in a transmission electron microscope and analysed quantitatively using image processing software. KEY RESULTS: In the chlorophyllose cells of the Sphagnum moss leaves from the capitulum, ozone exposure led to a decrease in chloroplast area and in granum stack thickness and various changes in plastoglobuli and cell wall thickness, depending on the species and the experiment. In E. vaginatum, ozone exposure significantly reduced chloroplast cross-sectional areas and the amount of starch, whereas there were no clear changes in the plastoglobuli. In the dwarf shrubs, ozone induced thickening of the cell wall and an increase in the size of plastoglobuli under summer conditions. In contrast, under autumn conditions the cell wall thickness remained unchanged but ozone exposure led to a transient increase in the chloroplast and starch areas, and in the number and size of plastoglobuli. CONCLUSIONS: Ozone responses in the Sphagnum mosses were comparable to typical ozone stress symptoms of higher plants, and indicated sensitivity especially in S. angustifolium. The responses in the dwarf shrubs suggest stimulation of photosynthesis by low ozone concentrations and ozone sensitivity only under cool autumn conditions.     

The paraveinal mesophyll of soybean leaves in relation to assimilate transfer and compartmentation

Nitrogen and carbohydrate assimilates were temporally and spatially compartmented among various cell types in soybean (Glycine max L., Merr.) leaves during seed filling. The paraveinal mesophyll (PVM), a unique cell layer found in soybean, was demonstrated to function in the synthesis, compartmentation and remobilization of nitrogen reserves prior to and during the seed-filling stages. At anthesis, the PVM vacuoles contain substantial protein which completely disappears by two weeks into the seed filling. Distinct changes in the PVM cytoplasm, tonoplast and organelles were correlated with the presence or absence of the vacuolar material. Microautoradiography following the accumulation of several radiolabeled sugars and amino acids demonstrated the glycoprotein nature of the vacuolar material. Incorporation of methionine, leucine, glucose, and glucosamine resulted in heavy labelling of the PVM vacuole, in contrast to galactose, proline, and mannose which resulted in a much reduced labelling pattern. In addition, starch is unequally compartmented and degraded among the various leaf cells during seed filling. At the end of the photoperiod at the flowering stage, the highest starch accumulation was in the second palisade layer followed by the spongy mesophyll and the first (uppermost) palisade layer. Starch in the first palisade layer was completely degraded during the dark whereas the starch in the second palisade and spongy mesophyll was not remobilized to any appreciable extent. By mid-podfilling (approximately five weeks postanthesis) starch was absent in the first palisade layer at the end of the photoperiod while the second palisade and spongy mesophyll layers contained substantial starch. Starch was remobilized from these latter cells during the remainder of seed filling when current photosynthetic production is low. Structural changes associated with cell senescence first appear in the upper palisade layer and then progress (excluding the PVM) to the second palisade and spongy mesophyll layer. The PVM and phloem appear to retain their structural integrity into the leaf yellowing stage. Reducing sink capacity by pod removal resulted in a continued accumulation of vacuolar protein, an increase in cytoplasmic volume, and fragmentation of the vacuole in the PVM. Pod removal also resulted in an increased amount of accumulated starch (which did not turn over) in all mesophyll layers, and an increase in cell size and cell-wall thickness.     

Nutritional quality of specific leaf tissues and selective feeding by a specialist leafminer

Summary Many folivorous insects are selective feeders which consume specific leaf tissues. For specialist herbivores feeding on plants of overall low nutritional quality, selective feeding may allow consumption of a high quality resource. Selective feeding may also allow insects to avoid structural or allelochemical defenses. We examined the structure and chemistry of leaves of American holly, Ilexopaca Aiton, and the feeding site of its principal insect herbivore, the native holly leafminer, Phytomyza ilicicola Loew (Diptera: Agromyzidae), to test the hypothesis that the leafminer consumes tissues which are of greater nutritional quality than the leaf as a whole. Holly leaves have a continuous layer of palisade mesophyll, uninterrupted by fibers or vascular bundles. The leafminer feeds entirely within this layer. The palisade mesophyll contained 196 mg/g dry wt extractable protein, more than twice as much as the leaf as a whole, and 375 mg/g dry wt saponins, more than 9 times that of the leaf as a whole. The water content of the palisade mesophyll was 66% higher than that of the leaf as a whole. The palisade mesophyll is 3–4 cell layers thick in leaves grown in full sun, but only 2 layers thick in shaded leaves. Crystals, probably of calcium oxalate, are abundant in the abaxial cell layer. These may impose mechanical constraints on larval feeding in shade leaves, which are thinner than sun leaves. Selective feeding on the middle palisade mesophyll of sun leaves allows the leafminer to consume a resource which is lacking in mechanical barriers and is rich in protein and water, but which contains large amounts of saponins.The investigation reported in this paper (No. 86-8-7-117) is in connection with a project of the Kentucky Agricultural Experiment Station and is published with the approval of the Director     

黄土高原不同植被带内草地植物叶片解剖性状的变异规律

叶片作为植物与外界进行物质交换的桥梁,其解剖性状能够相互协调以应对外界环境对植物生长造成的不利影响,从而反映出植物对环境变化所采取的适应策略。通过对黄土高原不同植被带(森林草原带、典型草原带、荒漠草原带)草地群落中常见115种植物(包括单子叶植物,双子叶植物,木本植物和草本植物四种功能型植物)叶片进行取样,并运用石蜡制片技术和光学显微技术获得叶片解剖性状(包括表皮厚度、栅栏组织厚度、海绵组织厚度、叶肉厚度和叶片厚度),旨在研究不同植被带内草地植物叶片解剖性状的变异规律及其与群落内物种相对优势度之间的关系,为黄土高原植被恢复和生态环境改善提供理论依据。结果表明：(1)沿着干旱梯度,从森林草原带、典型草原带到荒漠草原带,除叶肉厚度外,植物各叶片解剖性状值均呈现增大趋势,表明干旱地区叶片的旱生结构特征明显。(2)不同功能型植物叶片解剖性状与环境因子的关系各异。木本植物和草本植物的栅栏组织厚度和栅海比均与降水和土壤养分呈显著负相关关系(P<0.05)。同时,木本植物的叶片厚度与水分呈显著负相关关系(P<0.05),而草本植物表皮厚度仅与土壤养分呈显著负相关关系(P<0.05)...     

Nitric oxide production in tobacco leaf cells: a generalized stress response?

The function of nitric oxide (NO), a gaseous free radical emitted by many plants, is incompletely understood. In the present study the hypothesis that NO generation, like that of the reactive oxygen species, occurs as a general response to different environmental cues was tested. Leaf peels and mesophyll cell suspensions of Nicotiana tabacum cv. Xanthi were loaded with the NO‐specific fluorophore, diaminofluorescein, and subjected to an abiotic stressor. Light stress and mechanical injury had no apparent effect on NO production. In contrast, high temperatures, hyperosmotic stress, salinity and epi‐illumination in a microscope all led to rapid surges in NO‐induced fluorescence. The fluorescence originated from cells of the palisade mesophyll and across all epidermal cell types, including guard cells, subsidiary cells, and long and short trichomes. Fluorescence was evident first in the plastids, then in the nucleus and finally throughout the cytosol. Nicotiana plumbaginifolia cell suspensions expressing the calcium reporter aequorin provided evidence that, under hyperosmotic stress, NO participates in the elevation of free Ca²⁺ in the cytoplasm. The physiological significance of NO production in response to abiotic stressors is discussed.     

Basic Types of Structural Changes in the Leaf Mesophyll during Adaptation of Eastern Pamir Plants to Mountain Conditions

Quantitative characteristics of mesophyll structure were compared in leaves of eleven alpine plant species grown under natural conditions in the Eastern Pamirs at various altitudes, from 3800 to 4750 m. Basic types of changes in mesophyll structure, associated with plant adaptation to mountain conditions, were characterized. These changes manifested themselves in different numbers of cell layers and cell sizes in the palisade tissue and, as a consequence, in changed leaf thickness and cell number per unit of leaf area. Three plant groups were identified by the changes in the leaf structural characteristics depending on the type of mesophyll structure, ecological group of plant species, and altitude of plant habitat. The first group comprised alpine xerophytes with an isopalisade structure, in which the volume of palisade cells decreased and their number per unit of leaf area increased with the altitude of plant habitat. The number of mesophyll layers and leaf thickness decreased or did not change with altitude. The second group comprised subalpine plant species with a dorsoventral structure of mesophyll; these species occur below the line of continuous night frost. In these plant species, the number of mesophyll layers, leaf thickness, and cell number per unit of leaf area increased with altitude. The third group comprised mesophyte plants with a dorsoventral and homogenous mesophyll structure, which are encountered in a wide range of habitats, including the nival belt (from 4700 to 5000 m). In this group, cell volume increased and cell number per unit of leaf area decreased with altitude. We present a general scheme of leaf structural changes, which explains the changes in the quantitative characteristics of mesophyll as a function of altitude and highland environmental conditions.     

Observation of Apparently Unchanging Mesophyll Cell Diameters Throughout Leaf Ontogeny in Woody Species

The expansion of plant leaves usually lasts 3–6 weeks and it is widely believed that most cell types (epidermal and mesophyll) continue to expand in unison over a similar time period. The evidence supporting this account was derived from studies of herb leaves. We observed in woody species, however, that the diameter of mesophyll cells (spongy and palisade) changed little during leaf expansion from about 5 to 100 % maximum size. To keep pace with epidermal cell enlargement and leaf area expansion, mesophyll cells divided but palisade cell length expanded as leaves grew thicker. The prolonged division of mesophyll and apparently unchanging mesophyll cell diameters constitute a novel pattern of leaf cell development, different from that previously described for herbs. Possible mechanisms that attribute the varied expansion direction and speed to the different cellulose distributions in woody and herbaceous species are suggested. This finding could contribute to an enhanced understanding of the overall mechanism of leaf development.