Characteristics of leaf structure and photosynthetic apparatus within the crown of systematically shadedQuercus petraea andNothofagus procera seedlings

Four-year-old seedlings ofQuercus petraea (Matt.) Liebl. andNothofagus procera (Poepp. et Endl.) Querst were grown outdoors in pots while subjected to full, medium and low irradiances. Shading and decrease in height of leaf attachment generally increased specific leaf area, the diameters of chloroplasts and of palisade and spongy mesophyll cells, but decreased leaf thickness, number of palisade cell layers, length of palisade and spongy mesophyll cells, number of chloroplasts per mesophyll cell and epidermal cell and cuticle thickness, stomata and hair densities per unit leaf area, hair length, maximum hair breath and cell wall thickness in the two species. However, inN. procera grown under full irradiance, leaves at the upper and middle positions had hairs on both upper and lower epidermes, whereas those in other treatments and all leaves in all treatments inQ. petraea, had theirs only on the upper epidermis.     

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.     

A relationship between leaf structure and growth potential in apple

The number of palisade mesophyll cells per unit area leaf lamina was related to the growth potential of apple rootstocks covering a wide range of vigour. The number of these cells per unit leaf surface depends on cell diameter, the number of layers of palisade, and the amount of intercellular airspace. In composite, grafted trees rootstock influence was observed on the leaf structure of scions of low, or intermediate, growth potential.     

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.     

Shading-induced changes in the leaf mesophyll of plants of different functional types

Changes in the structural characteristics of mesophyll induced by shading were investigated in ten species of wild plants of diverse functional types. In all plant types, shading reduced leaf thickness and density by 30–50% and total surface of mesophyll, by 30–70%. The extent and mechanisms of mesophyll structural rearrangement depended on the plant functional type. In the ruderal plants, integral parameters of mesophyll, such as the surface of cells and chloroplasts and mesophyll resistance, changed threefold predominantly because of changes in the dimensions of the cells and chloroplasts. In these plants, shading reduced the volume of chloroplasts by 30%, and the chloroplast numbers per cell declined. The competitor plants showed a twofold increase in mesophyll resistance due to a decrease in the number of photosynthesizing cells per leaf area unit. Moreover, these plants maintained constant dimensions of mesophyll cells, ratios mesophyll surface/mesophyll volume and chloroplast surface/cell surface. In stress-tolerant plants, diffusion resistance of mesophyll remained the same irrespective of the growing conditions, and mesophyll rearrangement was associated with inversely proportional changes in the dimensions of the cells and cell volume per chloroplast. Noteworthy of these plants were relatively constant chloroplasts number per cell, per leaf area unit and total surface area of chloroplasts. The nature of relationship between the mesophyll diffusion resistance and structural parameters of leaf mesophyll differed in plants of diverse functional types.     

Variation in Mesophyll Cell Number and Size in Wheat Leaves

The numbers of mesophyll cells in wheat leaves were determinedin a variety of wheat species differing in ploidy level andin leaves from different positions on the wheat plant. Leafsize and mesophyll cell number are linearly related in bothcases but differences were observed in mesophyll cell numberper unit leaf area with changing leaf size. Where changes incell size are caused either by nuclear ploidy or leaf position,differences in mesophyll cell number per unit leaf are negativelycorrelated with mesophyll cell plan area. The decrease in cellsize with increasing leaf position also results in a greaternumber of chloroplasts per unit leaf area. These results arediscussed in relation to anatomical variation of the wheat leaf. Mesophyll cell, cell numbers, leaf size, Triticum     

Estimating near-infrared leaf reflectance from leaf structural characteristics

The relationship between near-infrared reflectance at 800 nm (NIRR) from leaves and characteristics of leaf structure known to affect photosynthesis was investigated in 48 species of alpine angiosperms. This wavelength was selected to discriminate the effects of leaf structure vs. chemical or water content on leaf reflectance. A quantitative model was first constructed correlating NIRR with leaf structural characteristics for six species, and then validated using all 48 species. Among the structural characteristics tested in the reflectance model were leaf trichome density, the presence or absence of both leaf bicoloration and a thick leaf cuticle (>1 μm), leaf thickness, the ratio of palisade mesophyll to spongy mesophyll thickness (PM/SM), the proportion of the mesophyll occupied by intercellular air spaces (%IAS), and the ratio of mesophyll cell surface area exposed to IAS (A(mes)) per unit leaf surface area (A), or A(mes)/A. Multiple regression analysis showed that measured NIRR was highly correlated with A(mes)/A, leaf bicoloration, and the presence of a thick leaf cuticle (r = 0.93). In contrast, correlations between NIRR and leaf trichome density, leaf thickness, the PM/SM ratio, or %IAS were relatively weak (r < 0.25). A model incorporating A(mes)/A, leaf bicoloration, and cuticle thickness predicted NIRR accurately for 48 species (r = 0.43; P < 0.01) and may be useful for linking remotely sensed data to plant structure and function.     

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.     

Structure of the Photosynthetic Apparatus in Leaves of Freshwater Hydrophytes: 2. Quantitative Characterization of Leaf Mesophyll and the Functional Activity of Leaves with Different Degrees of Submersion

The structure of leaf photosynthetic elements was investigated on 42 boreal plant species characterized by different degrees of submergence (helophytes, neustophytes, and hydatophytes). Six main types of mesophyll structures were identified. Quantitative characteristics for the mesostructure of the photosynthetic apparatus in these groups were determined, such as the size and abundance of cells and chloroplasts in the mesophyll and epidermis, the number of plastids per cell in each tissue, the total surface area of the mesophyll cells, epidermal cells, and chloroplasts per unit leaf area. Analysis showed that quantitative characteristics of the photosynthetic apparatus in hydrophytes are determined by two factors: (a) the degree of leaf submergence and (b) the type of mesophyll structure. With an increasing degree of immersion in water, the mesophyll types change in a sequence isopalisade dorsoventral homogeneous. The leaves become thinner, their weight per unit area diminishes, cells and chloroplasts become less numerous (on a per unit leaf area basis), but their dimensions become larger. Adaptation to aquatic medium is also manifested in the increasing contribution of the epidermal tissue to the overall photosynthesis: in submerged leaves, the epidermis accounts for more than 50% of the photosynthetic activity. The occurrence of six structural types of leaves contrasting in their characteristics was confirmed by discriminatory analysis according to the qualitative parameters of mesophyll.     

Associations between leaf structure, orientation, and sunlight exposure in five Western Australian communities

Five plant communities in Western Australia, as well as selected desert and Rocky Mountain species of the western USA, were surveyed to evaluate associations among leaf structure, orientational properties, and the sunlight exposure and precipitation characteristic of each community. Selected leaf structural features have been associated previously with photosynthetic function and included shape, thickness, the ratio of thickness to width, stomatal distribution, leaf surface coloration, and the number and distribution of palisade cell layers. Decreases in annual precipitation (<4 to over 15 cm/yr) and increases in total daily sunlight (4.2 to 29.2 mol photons/m1) corresponded strongly to an increase in the percentage of species in a given community with more inclined (more inclined than +/- 45 degrees from horizontal) or thicker leaf mesophyll (>0.4 mm) leaves. Also, the percentage of species with a leaf thickness to width ratio >0.1, which were amphistomatous, or which had palisade cell layers beneath both leaf surfaces, increased from >20% in the highest rainfall and lowest sunlight community to >80% in the community with least rainfall but greatest sunlight exposure. Over 70% of the species in the most mesic, shaded community had lighter abaxial than adaxial leaf surfaces (leaf bicoloration). All of the above structural features were positively associated with a more inclined leaf orientation (r1 = 0.79), except for leaf bicoloration, which was negatively associated (r1 = 0.75). The ratio of adaxial to abaxial light was more strongly associated with leaf bicoloration (r1 = 0.83) and the presence of multiple adaxial and isobilateral palisade cell layers(r1 = 0.80) than with total incident sunlight on just the adaxial leaf surface (r1 = 0.69 and 0.73, respectively). These results provide field evidence that leaf orientation and structure may have evolved in concert to produce a photosynthetic symmetry in leaf structure in response to the amount of sunlight and other limiting factors of the community. This structural symmetry may serve fundamentally to regulate the distribution of both light and CO2 levels inside the leaf and, thus, increase photosynthetic CO2 uptake per unit leaf biomass.     

Leaf anatomical and photosynthetic acclimation to cool temperature and high light in two winter versus two summer annuals

Acclimation of foliar features to cool temperature and high light was characterized in winter (Spinacia oleracea L. cv. Giant Nobel; Arabidopsis thaliana (L.) Heynhold Col‐0 and ecotypes from Sweden and Italy) versus summer (Helianthus annuus L. cv. Soraya; Cucurbita pepo L. cv. Italian Zucchini Romanesco) annuals. Significant relationships existed among leaf dry mass per area, photosynthesis, leaf thickness and palisade mesophyll thickness. While the acclimatory response of the summer annuals to cool temperature and/or high light levels was limited, the winter annuals increased the number of palisade cell layers, ranging from two layers under moderate light and warm temperature to between four and five layers under cool temperature and high light. A significant relationship was also found between palisade tissue thickness and either cross‐sectional area or number of phloem cells (each normalized by vein density) in minor veins among all four species and growth regimes. The two winter annuals, but not the summer annuals, thus exhibited acclimatory adjustments of minor vein phloem to cool temperature and/or high light, with more numerous and larger phloem cells and a higher maximal photosynthesis rate. The upregulation of photosynthesis in winter annuals in response to low growth temperature may thus depend on not only (1) a greater volume of photosynthesizing palisade tissue but also (2) leaf veins containing additional phloem cells and presumably capable of exporting a greater volume of sugars from the leaves to the rest of the plant.     

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.     

Leaf acclimation to experimental climate warming in meadow plants of different functional types

The use of open-top chambers (OTCs) installed in natural plant cover is one of the approaches to study plant responses to climate change. Three OTCs made from polyethylene film were installed on a herbgrass meadow in the subzone of the southern taiga before the beginning of the growing season. A significant increase in the average daily temperature values (by 0.5°C) and the relative humidity (by 10%) compared to control conditions was observed inside the chambers. Plant height, leaf parameters, and the pigments content were studied for six species of meadow plants during the growing season in two variants—inside the chamber and outside the chamber (control); more than 20 quantitative parameters of the mesophyll were studied for four of the species. It was found that the differences in microclimatic conditions had no effect on plant height and leaf area. A slight decrease in the thickness and density of the leaves and an increased water content were noted inside the OTCs. In contrast to weak changes in external leaf parameters, the internal leaf structure and the content of photosynthetic pigments varied considerably. Warming caused the reduction of the content of chlorophyll and carotenoids per unit leaf area in the majority of studied species, except for Veronica chamaedrys L., but the ratio of pigment forms did not change. Changes in the pigments content in the leaf were associated with some structural rearrangements in the mesophyll, whose mechanism depended on the functional properties of the species. Increased size of palisade cells and the number of chloroplasts per cell was noted in the ruderal species (R/CSR-strategist) Taraxacum officinale Wigg. s. l.; the reduction of chlorophyll content per leaf area occurred due to the decrease in chlorophyll content per a single chloroplast. Decreased number of cells and chloroplasts per leaf unit area without any changes in their size was marked for the species with S/CSR strategy Alchemilla vulgaris L. s. 1. and V. chamaedrys L. in a chamber, but the content of chlorophyll per a chloroplast increased. An increase in the number of cells and a simultaneous decrease in their size was observed in CR-strategist Cirsium setosum (Willd.) Bess. inside the OTC; the chlorophyll content per chloroplast did not change. It was concluded that the acclimation of plants to short-term climate warming was associated with the restructuring of leaf mesophyll, whose mechanism depended on the functional properties of the species.     

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。     

Ontogenetic differences in mesophyll structure and chlorophyll distribution in Eucalyptus globulus ssp. globulus

Mesophyll structure has been associated with the photosynthetic performance of leaves via the regulation of internal light and CO(2) profiles. Differences in mesophyll structure and chlorophyll distribution within three ontogenetically different leaf types of Eucalyptus globulus ssp. globulus were investigated. Juvenile leaves are blue-grey in color, dorsiventral (adaxial palisade layer only), hypostomatous, and approximately horizontal in orientation. In contrast, adult leaves are dark green in color, isobilateral (adaxial and abaxial palisade), amphistomatous, and nearly vertical in orientation. The transitional leaf type has structural features that appear intermediate between the juvenile and adult leaves. The ratio of mesophyll cell surface area per unit leaf surface area (A(mes)/A) of juvenile leaves was maximum at the base of a single, adaxial palisade layer and declined through the spongy mesophyll. Chlorophyll a + b content showed a coincident pattern, while the chlorophyll a:b ratio declined linearly from the adaxial to abaxial epidermis. In comparison, the mesophyll of adult leaves had a bimodal distribution of A(mes)/A, with maxima occurring beneath both the adaxial and abaxial surfaces within the first layer of multiple palisade layers. The distribution of chlorophyll a + b content had a similar pattern, although the maximum ratio of chlorophyll a:b occurred immediately beneath the adaxial and abaxial epidermis. The matching distributions of A(mes)/A and chlorophyll provide further evidence that mesophyll structure may act to influence photosynthetic performance. These changes in internal leaf structure at different life stages of E. globulus may be an adaptation for increased xeromorphy under increasing light exposure experienced from the seedling to adult tree, similar to the characteristics reported for different species according to sunlight exposure and water availability within their native habitats.     

EVOLUTION OF LEAF ARCHITECTURE IN THE CHAPARRAL SPECIES FREMONTODENDRON CALIFORNICUM SSP. CALIFORNICUM (STERCULIACEAE)

The chaparral species Fremontodendron californicum ssp. californicum and the cloud forest species Cheirostemon platanoides arose from a common mesophytic stock. The leaf of C. platanoides is proposed as a model for the ancestral leaf and provides an architectural baseline allowing identification of the types and relative magnitudes of morphological and anatomical modifications which have occurred in the leaf of F. californicum. The leaf of F. californicum reflects intense selection for reduced surface area; however, both deletion and addition of lobes has occurred. Vein density has increased due to a greater number of more highly branched terminal veinlets. Mechanical structures, i.e., sclerenchyma, cuticle, show a quantitative reduction; whereas, water storage tissue, i.e., mucilage cells, has been augmented at the expense of spongy mesophyll. The leaf has become weakly isolateral and the number of palisade strata has increased. No increase in the volume of palisade tissue per unit volume of mesophyll is apparent due to a reduction in palisade cell length and palisade packing density. Total leaf thickness has not increased. The indumentum shows selection for non-living, pedestalled stellate trichomes which form a transpiration-limiting canopy above the stomata. Stomatal density has decreased while guard cell dimensions have become larger. Some structural modifications in F. californicum appear to have evolved directly in response to long-term water stress, others seem more directly related to leaf diminution. Xeroplastic changes in foliar structure are in some cases unreliable indicators of ultimate xeromorphic modifications arising in an evolutionary context.     

Variation of leaf traits and pigment content in three species of steppe plants depending on the climate aridity

Mesophyll structure and content of photosynthetic pigments in the leaves of three species of steppe plants, Centaurea scabiosa L., Euphorbia virgata Waldst. et Kit., Helichrysum arenarium (L.) Moench, were investigated in four geographical sites of the Volga region and the Urals located in the forest-steppe and steppe zones. Variations of the studied parameters between geographical points depended both on the species and on the structural organization of the leaf. The highest level of variation was observed for leaf area and pigment content per unit leaf area, the size and the number of chloroplasts in the cell changed to a lesser extent. The leaf thickness, leaf area and mesophyll cell sizes mostly depended on the plant species. C. scabiosa had large leaves (40–50 cm²) with large thickness (280–290 μm) and large mesophyll cells (up to 15000 μm³). The leaves of H. arenarium and E. virgata were ten times smaller and characterized by 1.5 times smaller thickness and 2?3 times smaller cell size. Geographical location and climate of the region affected leaf density, proportion of partial tissue volume, and the ratio of the photosynthetic pigments. In the southern point of Volga region with the highest climate aridity, all studied species were characterized by maximum values of volumetric leaf density (LD), due to the high proportion of sclerenchyma and vascular bundles, and specificity of the mesophyll structure. With the decline in latitude, chlorophyll (Chl) and carotenoid (Car) contents in leaf area were reduced, the ratio Chl/Car was increased, and the ratio Chl a/b was declined. The reduction of the pigment content in the leaf in all species was associated with a reduction in the amount of Chl per chloroplast, and for C. scabiosa and H. arenarium it was associated also with the reduction of chloroplast amount in the leaf area. In turn, chloroplast number per leaf area and the total cell area (A_mes/A) depended on the ratio of the number and size of mesophyll cells inherent to this plant species. At the same time, we found a similar mechanism of spatial organization of leaf restructuring for all studied species—decrease in A_mes/A was accompanied by increasing in the proportion of intercellular air spaces in the leaf. It is concluded that variations in structural and functional parameters of the photosynthetic apparatus of steppe plants were associated with plant adaptation to climate features. General direction of the changes of leaf parameters of the studied species with aridity was the increase of LD and the decrease of pigment content per leaf area however the cellular mechanisms of changes in the pigment content and integral parameters of mesophyll were determined by the plant species properties.     

The analysis of the causes of variability of the relationship between leaf dry mass and area in plants

The lamina dry mass: area ratio (LMA - Leaf Mass per Area) is a quite variable trait. Leaf dry mass consists of symplast mass (a set of all leaf protoplasts) and apoplast mass (a set of all cell walls in a leaf). The ratio between symplast and apoplast masses is positively related to any functional trait of leaf calculated per unit of dry mass. The value of this ratio is defined by cells size and their number per unit of leaf area, number of mesophyll cells layers and their differentiation between palisade and spongy ones, and also by density of cells packing. The LMA value is defined by leaf thickness and density. The extent and direction of variability in both leaf traits define the extent and direction of variability in LMA. Negative correlation between leaf thickness and density reduces the level of LMA variability. As a consequence of this correlation the following pattern emerges: the thinner a leaf, the denser it is. Changes in the traits that define the LMA value take place both within a species under the influence of environmental factors and between species that differ in leaf structure and functions. Light is the most powerful environmental factor that influences the LMA, increase in illumination leading to increase in LMA. This effect occurs during leaf growth at the expense of structural changes associated with the reduction of symplast/apoplast mass ratio. Under conditions of intense illumination, LMA may increase due to accumulation of starch. With regard to the majority of leaf functions, the mass of starch may be ascribed to apoplast. Starch accumulation in leaves is observed also under conditions of elevated CO2 concentration in the air. Under high illumination, however, LMA increases also due to increased apoplast contribution to leaf dry mass. Scarce mineral nutrition leads to LMA increase due to lowering of growth zones demands for phothosyntates and, therefore, to increase in starch content of leaves. High level of mineral nutrition during leaf growth period leads to LMA increase at the expense of mesophyll thickening where components of photosynthesis system are located. When additional environmental factors are involved, starch accumulation may be partly responsible for increase in LMA. LMA increase at the expense of starch accumulation, unlike that at the expense of mesophyll thickening, is accompanied by increased leaf density. Under conditions of water deficiency LMA increases, which in mature leaf may be caused by starch accumulation. LMA increase during leaf growth period under conditions of water deficiency is associated with decrease in the symplast/apoplast mass ratio.     

Genome-Dependent Factors in the Development of Leaf Phototrophic Tissues in Diploid and Alloploid Wheat Species

Growth and mesostructure of the photosynthetic apparatus were studied in leaves of ten Triticum L. species. Plants with the A^u genome were shown to develop larger leaf assimilation areas due to expanding areas of individual leaves and an increase in the absolute growth rate. Leaf and mesophyll thickness and mesophyll cell size decreased in the G-genome species. Leaf compactness, which depended on cell size and number per unit leaf area and leaf folding, determined the specific patterns of internal leaf organization in wheat species with diverse genotypes. These patterns did not affect cell plastid-to-cytoplasm ratio as shown by the stable indices of cell surface area/cell volume, cell surface area per chloroplast, and cell volume per chloroplast. The structural indices of leaf phototrophic tissues, mesophyll density, and mesophyll CO₂ conductance in alloploids, as compared to diploid species, depended on both ploidy and genome constitution.     

SOME EFFECTS OF NITROGEN NUTRITION ON THE MORPHOLOGY AND ANATOMY OF MARROW-STEM KALE

Marrow-stem kale plants grown on plots receiving frequent additions of sulphate of ammonia showed a 40% increase in length of internode and a 25% increase in number of nodes per plant, and the leaf size was increased by between 50 and 70% over plants in plots receiving no N fertilizer. Leaves of kale continue to increase in area until they turn yellow, and the high N leaves showed a greater rate of increase in area at every stage in the life of the leaf.
Various features of leaf structure, such as stomatal index, and thickness of palisade and mesophyll, were unaffected by N treatment. The size of the epidermal cells of the leaves was very variable, and although the high N leaves showed a 12% increase in area per epidermal cell over the low N leaves, this difference is not statistically significant. The increased area of the high N leaves can therefore be attributed mainly to increased cell division during the life of the leaf. Only a very slight increase in rate of cell division is necessary to produce the observed effect.
The greater leaf area of the high N plants can be attributed mainly to increased size of individual leaves, but there was also a significantly greater number of living functional leaves per plant on the high N plants; at 23 weeks from sowing the high N plants had an average of 13.4 living leaves, while the low N plants had only 11.7 living leaves per plant.
There was an appreciable degree of N succulence in the high N kale leaves, which showed a 2% greater moisture content than the low N leaves.
A seasonal drift in epidermal cell size, palisade thickness, and total leaf thickness, is shown to be fully significant, statistically. Marked variations in stomatal frequency are barely significant at the 5% level.