L. A. Ivanov

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.

Changes in Leaf Characteristics as Indicator of the Alteration of Functional Types of Steppe Plants along the Aridity Gradient

Changes in some leaf characteristics: leaf mass area (LMA), content of photosynthetic pigments and nitrogen in the leaves, leaf mass ratio (LMR) and leaf area ratio (LAR) were investigated in steppe plants of the Volga land along the gradient of aridity. When drought stress became stronger, the content of chlorophylls in the leaves, LMR and LAR decreased, whereas LMA and the proportion of carotenoids in the leaves rose. In the North to South direction, the content of pigments and nitrogen per unit whole plant weight considerably decreased (4 and 2 times, respectively). The relationship between leaf indices (chlorophyll and nitrogen contents and LMA) differed along this gradient. It was concluded that adaptation of steppe plants to drought stress generally depended on predominant development of heterotrophic tissues in the leaf and the whole plant. During aridization, the stress-tolerant species became more numerous.

Changes in the specific density of leaves of Eurasian plants along the aridity gradient

The specific surficial density of a leaf (SSDL or dryweight of the leaf area unit) is a fundamental characteristic of a plant [1, 2] and is used in global models ofthe Earth’s vegetation [3]. The SSDL is related tomany aspects of plant growth and development [4] andcorrelates with such paramete rs as the level of net photosynthesis [5], content of nitrogen in leaves [6], thelifetime of leaves, and the relative growth rate [4].Generally, the SSDL is an integral parameter of a leafbecause it reflects the adaptation of the photosyntheticapparatus of plants to the complex of environmentalconditions. It is known that plants from differentbiomes differ in the density of the leaf blade [2].The SSDL of plants along global climatic gradients[6–9] are intensively investigate due to the necessity offorecasting the state of vegetation during climatechange. Earlier, it was shown that the SSDL of plantscorrelates with the average annual temperature,potential evapotranspiration, and the solar radiationlevel [6]. The plants of arid regions of Eurasia are stillinadequately known in this respect.In the present study, 495 species of steppe anddesert territories of Russia and Mongolia are investigated. It is shown that in arid regions of Eurasia, thechange in the SSDL of plants is nonlinear under conditions of linear change in the aridity index of the territory.Investigations were conducted during 1995–2007in Russia and Mongolia within 43.25° and 56.5° N(Table 1). Plants with typical flat leaves were studied,mainly belonging to dominants and edificatory plantsof the plant cover. Succulents and plants with aphylllous shoots were not studied. The SSDL (mg/dm

Changes in the chlorophyll and carotenoid contents in the leaves of steppe plants along a latitudinal gradient in South Ural

We studied the content of chlorophylls and carotenoids in the leaves of steppe plants of South Ural growing along a latitudinal gradient from southern steppe to forest-steppe. The content of chlorophylls (a + b) was 5–6 mg per 1 g of the leaf dry weight and did not depend on the latitude, whereas the content of carotenoids in the leaves increased northward from 1.0 to 1.5 mg/g dry wt. At the same time, the greatest changes occurred in the ratios between the forms of pigments: the chlorophyll a/b ratio increased from 1.8 to 2.8, and the chlorophyll/carotenoid ratio decreased from 5.6 to 3.5. The obtained results indicate that adaptation of the pigment apparatus of steppe plants growing along the latitudinal gradient occurs due to the transformation of the light-harvesting complex.

Construction costs and mesostructure of leaves in hydrophytes

Construction costs (CC) and parameters of leaf structure (specific leaf weight, dry matter content, volume of photosynthesizing cells, and the number of cells per leaf area unit) were determined for 19 species of aquatic higher plants. The CC of 1 g dry matter varied from 0.98 g glucose in Lemna gibba L. to 1.48 g glucose in Nuphar pumila (Timm) DC. and Potamogeton natans L. The CC of leaf area unit varied to a greater extent than the CC of 1 g dry wt (from 10 to 97 g glucose/m2) and depended on the type of mesophyll structure. In leaves of hydrophytes with dorsoventral mesophyll structure, the CC of 1 m2 leaf area was 3–9 times larger than in leaves with homogeneous structure. Variations in CC of 1 m2 leaf area in hydrophytes were affected insignificantly (by 2% only) by variations of CC per 1 g dry wt and were mainly determined (by 82%) by changes in specific leaf weight. Two-factor analysis of variance has shown that the CC of 1 g dry wt in hydrophytes depended on the attachment of plants to the sediment: the CC was 1.2 times larger in rooted hydrophytes than in free floating plants. The second factor (the extent of submergence) potentiated the effect of rooting on CC. Reliable differences were found between the leaf CC for hydrophytes belonging to four groups distinguished by the extent of their contact with water and sediment. In a group series: rooted hydrophytes with floating leaves → submerged rooted hydrophytes → free floating submerged hydrophytes → free floating surface inhabiting hydrophytes, the CC of 1 g dry wt decreased by 1.3 times. Path analysis has shown that this trend was due to the increase in photosynthesizing cell volume and to reduction in number of cells per leaf area unit, which caused the decrease in dry matter content. The decrease in the content of leaf dry matter was accompanied by changes in its chemical composition: the content of carbon and nitrogen decreased. This led to a consistent decrease in leaf CC expressed per 1 g dry wt upon the increase in extent of plant hydrophilicity.

Changes in chemical composition of hydrophyte leaves during adaptation to aquatic environment

Leaf chemical composition of 19 hydrophytes was studied. The content of carbon, nitrogen, nonstructural carbohydrates, organic acids, minerals, and water was determined. Hydrophytes were shown to contain less carbon (below 410 mg/g dry wt in 60% species) than terrestrial plants. Hydrophytes and terrestrial plants did not differ in the nitrogen concentration in the leaves (33 and 29 mg/g dry wt, respectively). Hydrophytes were characterized by a low content of organic acids (40–90 mg/g dry wt in 60% species) and high content of mineral compounds (90–170 mg/g dry wt in 50% species). Total amount of nonstructural carbohydrates was similar in the leaves of hydrophytes and terrestrial plants (from 120 to 190 mg/g dry wt), but the proportions of various carbohydrate fractions differed substantially. In the hydrophyte leaves, the content of soluble carbohydrates was 2.4-fold lower, whereas the content of nonstructural polysaccharides 1.2-fold higher than in terrestrial plant leaves. Two groups of correlations between parameters of leaf chemical composition were distinguished: the contents of carbon, nitrogen, and soluble sugars were positively correlated, and the negative correlation was observed between these parameters and the amounts of mineral compounds, organic acids, water, and nonstructural polysaccharides. We concluded that hydrophyte leaf chemical composition reflects a specificity of plant adaptation to aquatic environment.

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.

Chemical composition of leaves and structure of photosynthetic apparatus in aquatic higher plants

Chemical composition of leaves (the content of carbon, nitrogen, nonstructural carbohydrates, organic acids, mineral substances, and water) and the structure of photosynthetic apparatus (specific leaf weight, cell volume, and the number of cells per unit leaf area) were investigated for 18 species of aquatic plants featuring various degrees of contact with aqueous environment and sediment. The rooted hydrophytes with floating leaves were characterized by comparatively high content of carbon and nitrogen (437 and 37 mg/g dry wt, respectively) and by low concentration of nonstructural carbohydrates, mineral substances, and organic acids (161, 54, and 60 mg/g dry wt, respectively). Unlike rooted plants, the free-floating nonrooted hydrophytes had characteristically higher content of nonstructural polysaccharides and mineral substances (by a factor of 1.3 and 1.6, respectively), while the leaf nitrogen content was 1.4 times lower, and the proportion of soluble carbohydrates in the total content of nonstructural carbohydrates was rather low (9%). The chemical composition of leaves in submerged rooted hydrophytes was intermediate between those for rooted hydrophytes with floating leaves and for nonrooted free plants. We found reliable positive correlations between the volume of photosynthesizing cells and the leaf content of organic acids (r = 0.69), as well as between specific leaf weight, the number of photosynthesizing cells per unit leaf area, and carbon content (r = 0.67 and r = 0.62, respectively). The content of nitrogen and nonstructural carbohydrates in hydrophytes was unrelated to structural characteristics of photosynthetic apparatus and depended on the absence or presence of plant attachment to the sediment. It is concluded that the structural traits of photosynthetic apparatus and the leaf chemical composition in hydrophytes featuring different degrees of plant contact with water and sediment reflect the specificity of plant adaptation to complex conditions of their habitats.

Mesostructure of the ginseng photosynthetic apparatus in relation to ecological strategy of the species

Mesostructure of the photosynthetic apparatus was investigated in the wild plants of ginseng (Panax ginseng C.A. Mey, Araliaceae) taken from various habitats. The revealed features of the structure of phototrophic tissues (large cells, small number of photosynthesizing cells and chloroplasts per leaf area unit, low values of membrane indices of the cells and chloroplasts) point to stress-tolerant type of ecological strategy ginseng pursues in Nature.

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 (Ames/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 Ames/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.