Morphometric Characterization of Halophytic Plant Chloroplasts

Russian Journal of Plant Physiology - The data concerning morphometric characterization of chloroplasts belonging to two groups of halophytes distinguished by their salt...     

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.     

Structural and Functional Changes in the Leaves of Plants from Steppe Communities as Affected by Aridization of the Eurasian Climate

Morphological and physiological characteristics of leaves from plant species collected in steppe communities in the various climatic zones in Eurasia were compared. The changes in leaf structure correlated with the major climatic factors. The mean thickness of leaves increased with increasing mean temperature of July and decreasing mean precipitation, which corresponded to aridity increase. The increased leaf thickness correlated with an increase in the specific leaf weight. The content of chlorophylls (a + b) in leaves greatly varied with plant habitats, whereas the chlorophyll a/b ratio remained unchanged. The chlorophyll content in leaf tissues had a general tendency to decrease with increasing leaf thickness. The leaf chlorophyll content positively correlated (R ² = 0.77) with the proportion of chlorenchyma in leaf tissues. It is concluded that steppe plants adapt to climate aridization at the structural level by increasing the proportion of protective heterotrophic components of the leaf without changing the functional activity of photosynthetic tissues.     

Photosynthetic Metabolism and Activity of Carboxylating Enzymes in Emergent,Floating, and Submersed Leaves of Hydrophytes

Radioisotope techniques were used to compare photosynthetic CO₂ fixation, activities of carboxylating enzymes, and the composition of photosynthates in 42 species of aquatic plants (emergent, floating, and submersed hydrophytes) collected from rivers Sysert' and Iset' in Sverdlovsk oblast (Russia). The submersed leaves, in comparison with the emergent and floating leaves, featured lower rates of potential photosynthesis (by 2.2 mg CO²/(dm² h) on average), low content of the fraction I protein, and low activity of Rubisco and phosphoenolpyruvate carboxylase (PEPC). The averaged activities of Rubisco and PEPC were diminished in submersed leaves by 10 and 1 mg/(dm² h), respectively. Different hydrophyte groups showed similar composition of assimilates accumulated after 5-min photosynthesis and did not differ in this respect from terrestrial plants. However, the incorporation of ¹⁴C into sucrose and starch in submersed leaves (30 and 9% of total labeling, respectively) was lower than in emergent and floating leaves (45 and 15%, respectively). At the same time, the incorporation of ¹⁴C into C₄ acids (malate and aspartate) was 1.5 times higher in submersed leaves than in other leaf types. Analysis of leaf differentiation, the Rubisco/PEPC activity ratio, the PEPC activity, and the composition of primary photosynthates in the pulse–chase experiments revealed no evidence of the C₄ effect in the submersed hydrophytes examined. The adaptation of hydatophytes to specific conditions of an aquatic environment was structurally manifested in the reduction (by a factor of 3–5) in the number of chloroplasts per 1 cm² leaf area. This small number of chloroplasts was responsible for low photosynthetic rates in submersed leaves, although metabolic activities of individual chloroplasts were similar for all three hydrophyte groups.     

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.     

Structural and Functional Grounds for Ephedra sinica Expansion in Mongolian Steppe Ecosystems

Morphological and structural characteristics of photosynthetic organs, diurnal changes in photosynthetic and transpiration rates, and the efficiency of water use were studied in three plant species from mountain-steppe ecosystems in Mongolia, Ephedra sinica Stapf, Stipa glareosa P. Smirn., and Allium polyrhizum Furcz. ex Regel. The species studied differed in the structural and functional mechanisms for the adaptation of photosynthetic apparatus to arid conditions. E. sinica has thick, vertical assimilating shoots, which are characterized by a high density (620 mg/cm³) and a small proportion of photosynthetic tissues (13%). The proportion of meso-phyll in the leaves of A. polyrhizum and S. glareosa was two and three times higher, respectively. The low content of phototrophic tissues in E. sinica shoot was compensated for by a high photosynthetic activity of single chloroplasts (25 mg CO₂/(10⁹ chloroplast h)), which was six times higher, than in two other species. Daily course of photosynthesis and transpiration inE. sinica differed from those of A. polyrhizum and S. glareosa by the absence of the midday depression. E. sinica had the highest efficiency of water use (45 mg CO₂/g H₂O) due to a low transpiration rate (0.25 g/g fr wt h). It is concluded that, in E. sinica, the main strategy for adaptation to arid stress is to develop in the shoot a few photosynthesizing cells of high assimilation activity. Such structural organization of photosynthetic organs in ephedra contributes to its higher efficiency of water use and stability of physiological characteristics under changing environmental conditions. These specific features of the structure of assimilating organs and their functional features contribute to a greater expansion of E. sinica with increasing climate aridization in Mongolia.     

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.     

Over-expression of gsh1 in the cytosol affects the photosynthetic apparatus and improves the performance of transgenic poplars on heavy metal-contaminated soil

Recent studies of transgenic poplars over‐expressing the genes gsh1 and gsh2 encoding γ‐glutamylcysteine synthetase (γ‐ECS) and glutathione synthetase, respectively, provided detailed information on regulation of GSH synthesis, enzymes activities and mRNA expression. In this experiment, we studied quantitative parameters of leaves, assimilating tissues, cells and chloroplasts, mesophyll resistance for CO₂ diffusion, chlorophyll and carbohydrate content in wild‐type poplar and transgenic plants over‐expressing gsh1 in the cytosol after 3 years of growth in relatively clean (control) or heavy metal‐contaminated soil in the field. Over‐expression of gsh1 in the cytosol led to a twofold increase of intrafoliar GSH concentration and influenced the photosynthetic apparatus at different levels of organisation, i.e., leaves, photosynthetic cells and chloroplasts. At the control site, transgenic poplars had a twofold smaller total leaf area per plant and a 1.6‐fold leaf area per leaf compared to wild‐type controls. Annual aboveground biomass gain was reduced by 50% in the transgenic plants. The reduction of leaf area of the transformants was accompanied by a significant decline in total cell number per leaf, indicating suppression of cell division. Over‐expression of γ‐ECS in the cytosol also caused changes in mesophyll structure, i.e., a 20% decrease in cell and chloroplast number per leaf area, but also an enhanced volume share of chloroplasts and intercellular airspaces in the leaves. Transgenic and wild poplars did not exhibit differences in chlorophyll and carotenoid content of leaves, but transformants had 1.3‐fold fewer soluble carbohydrates. Cultivation on contaminated soil caused a reduction of palisade cell volume and chloroplast number, both per cell and leaf area, in wild‐type plants but not in transformants. Biomass accumulation of wild‐type poplars decreased in contaminated soil by more than 30‐fold, whereas transformants showed a twofold decrease compared to the control site. Thus, poplars over‐expressing γ‐ECS in the cytosol were more tolerant to heavy metal stress under field conditions than wild‐type plants according to the parameters analysed. Correlation analysis revealed strong dependence of cell number per leaf area unit, chloroplast parameters and mesophyll resistance with the GSH level in poplar leaves.