Mesophyll Cell Surface Area Research Articles

3-D reconstruction of rice leaf tissue for proper estimation of surface area of mesophyll cells and chloroplasts facing intercellular airspaces from 2-D section images.

The surface area of mesophyll cells (Smes) and chloroplasts (Sc) facing the intercellular airspace (IAS) are important parameters for estimating photosynthetic activity from leaf anatomy. Although Smes and Sc are estimated based on the shape assumption of mesophyll cells (MCs), it is questionable if the assumption is correct for rice MCs with concave-convex surfaces. Therefore, in this study, we establish a reconstruction method for the 3-D representation of the IAS in rice leaf tissue to calculate the actual Smes and Sc with 3-D images and to determine the correct shape assumption for the estimation of Smes and Sc based on 2-D section images. We used serial section light microscopy to reconstruct 3-D representations of the IAS, MCs and chloroplasts in rice leaf tissue. Actual Smes and Sc values obtained from the 3-D representation were compared with those estimated from the 2-D images to find the correct shape-specific assumption (oblate or prolate spheroid) in different orientations (longitudinal and transverse sections) using the same leaf sample. The 3-D representation method revealed that volumes of the IAS and MCs accounted for 30 and 70 % of rice leaf tissue excluding epidermis, respectively, and the volume of chloroplasts accounted for 44 % of MCs. The shape-specific assumption on the sectioning orientation affected the estimation of Smes and Sc using 2-D section images with discrepancies of 10-38 %. The 3-D representation of rice leaf tissue was successfully reconstructed using serial section light microscopy and suggested that estimation of Smes and Sc of the rice leaf is more accurate using longitudinal sections with MCs assumed as oblate spheroids than using transverse sections with MCs as prolate spheroids.

Cost-benefit analysis of mesophyll conductance: diversities of anatomical, biochemical and environmental determinants.

Plants invest photosynthates in construction and maintenance of their structures and functions. Such investments are considered costs. These costs are recovered by the CO2 assimilation rate (A) in the leaves, and thus A is regarded as the immediate, short-term benefit. In photosynthesizing leaves, CO2 diffusion from the air to the carboxylation site is hindered by several structural and biochemical barriers. CO2 diffusion from the intercellular air space to the chloroplast stroma is obstructed by the mesophyll resistance. The inverses is the mesophyll conductance (gm). Whether various plants realize an optimal gm, and how much investment is needed for a relevant gm, remain unsolved. This review examines relationships among leaf construction costs (CC), leaf maintenance costs (MC) and gm in various plants under diverse growth conditions. Through a literature survey, we demonstrate a strong linear relationship between leaf mass per area (LMA) and leaf CC. The overall correlation of CC vs. gm across plant phylogenetic groups is weak, but significant trends are evident within specific groups and/or environments. Investment in CC is necessary for an increase in LMA and mesophyll cell surface area (Smes). This allows the leaf to accommodate more chloroplasts, thus increasing A. However, increases in LMA and/or Smes often accompany other changes, such as cell wall thickening, which diminishes gm. Such factors that make the correlations of CC and gm elusive are identified. For evaluation of the contribution of gm to recover CC, leaf life span is the key factor. The estimation of MC in relation to gm, especially in terms of costs required to regulate aquaporins, could be essential for efficient control of gm over the short term. Over the long term, costs are mainly reflected in CC, while benefits also include ultimate fitness attributes in terms of integrated carbon gain over the life of a leaf, plant survival and reproductive output.

The α subunit of the heterotrimeric G protein regulates mesophyll CO2 conductance and drought tolerance in rice

Summary Mesophyll conductance g m determines CO2 diffusion rates from mesophyll intercellular air spaces to the chloroplasts and is an important factor limiting photosynthesis. Increasing g m in cultivated plants is a potential strategy to increase photosynthesis and intrinsic water use efficiency (WUEi). The anatomy of the leaf and metabolic factors such as aquaporins and carbonic anhydrases have been identified as important determinants of g m. However, genes involved in the regulation and modulation of g m remain largely unknown.In this work, we investigated the role of heterotrimeric G proteins in g m and drought tolerance in rice d1 mutants, which harbor a null mutation in the Gα subunit gene, RGA1. d1 mutants in both cv Nipponbare and cv Taichung 65 exhibited increased g m, fostering improvement in photosynthesis, WUEi, and drought tolerance compared with wild‐type. The increased surface area of mesophyll cells and chloroplasts exposed to intercellular airspaces and the reduced cell wall and chloroplast thickness in the d1 mutant are evident contributors to the increase in g m.Our results indicate that manipulation of heterotrimeric G protein signaling has the potential to improve crop WUEi and productivity under drought.

High photosynthesis rate in two wild rice species is driven by leaf anatomy mediating high Rubisco activity and electron transport rate.

The importance of increasing photosynthetic efficiency for sustainable crop yield increases to feed the growing world population is well recognized. The natural genetic variation in leaf photosynthesis in crop plants is largely unexploited for increasing yield potential. The genus Oryza, including cultivated rice and wild relatives, offers tremendous genetic variability to explore photosynthetic differences and underlying biochemical, photochemical, and developmental traits. We quantified leaf photosynthesis and related physiological parameters for six cultivated and three wild rice genotypes, and identified photosynthetically efficient wild rice accessions. Fitting A/Ci curves and biochemical analyses showed that leaf photosynthesis in cultivated rice varieties IR 64 and Nipponbare was limited due to leaf nitrogen content, Rubisco activity, and electron transport rate compared with photosynthetically efficient wild rice accessions Oryza australiensis and Oryza latifolia. The selected wild rice accessions with high leaf photosynthesis per unit area had anatomical features such as larger mesophyll cells with more chloroplasts, fewer mesophyll cells between two adjacent veins, and higher mesophyll cell and chloroplast surface area exposed to intercellular space. Our results show the existence of desirable variations in Rubisco activity, electron transport rate, and leaf anatomical features that could be targeted for increasing the photosynthetic efficiency of cultivated rice varieties.

Ploidy influences wheat mesophyll cell geometry, packing and leaf function.

Leaf function is influenced by leaf structure, which is itself related not only to the spatial arrangement of constituent mesophyll cells, but also their size and shape. In this study, we used confocal microscopy to image leaves of Triticum genotypes varying in ploidy level to extract 3D information on individual mesophyll cell size and geometry. Combined with X‐ray Computed Tomography and gas exchange analysis, the effect of changes in wheat mesophyll cell geometry upon leaf structure and function were investigated. Mesophyll cell size and shape were found to have changed during the course of wheat evolution. An unexpected linear relationship between mesophyll cell surface area and volume was discovered, suggesting anisotropic scaling of mesophyll cell geometry with increasing ploidy. Altered mesophyll cell size and shape were demonstrated to be associated with changes in mesophyll tissue architecture. Under experimental growth conditions, CO2 assimilation did not vary with ploidy, but stomatal conductance was lower in hexaploid plants, conferring a greater instantaneous water‐use efficiency. We propose that as wheat mesophyll cells have become larger with increased ploidy, this has been accompanied by changes in cell geometry and packing which limit water loss while maintaining carbon assimilation.

High leaf mass per area Oryza genotypes invest more leaf mass to cell wall and show a low mesophyll conductance.

The intraspecific variations of leaf structure and anatomy in rice leaves and their impacts on gas diffusion are still unknown. Researches about the tradeoff between structural compositions and intracellular chemical components within rice leaves are still lacking. The objectives of the present study were to investigate the varietal differences in leaf structure and leaf chemical compositions, and the tradeoff between leaf structural tissues and intracellular chemical components in rice leaves. Leaf structure, leaf anatomy, leaf chemical composition concentrations and gas exchange parameters were measured on eight Oryza sativa L. genotypes to investigate the intraspecific variations in leaf structure and leaf anatomy and their impacts on gas exchange parameters, and to study the tradeoff between leaf structural compositions (cell wall compounds) and intracellular chemical components (non-structural carbohydrates, nitrogen, chlorophyll). Leaf thickness increased with leaf mass per area (LMA), while leaf density did not correlate with LMA. Mesophyll cell surface area exposed to intercellular airspace (IAS) per leaf area, the surface area of chloroplasts exposed to IAS and cell wall thickness increased with LMA. Cell wall compounds accounted for 71.5 % of leaf dry mass, while mass-based nitrogen and chlorophyll concentrations decreased with LMA. Mesophyll conductance was negatively correlated with LMA and cell wall thickness. High LMA rice genotypes invest more leaf mass to cell wall and possess a low mesophyll conductance.

Contrasting photosynthetic responses to ambient air pollution between the urban shrub Rhododendron × pulchrum and urban tall tree Ginkgo biloba in Kyoto city: stomatal and leaf mesophyll morpho-anatomies are key traits

Rhododendron × pulchrum avoided air pollution by adjusting its stomatal density, while Ginkgo biloba tolerated air pollution via a small stomatal density and high mesophyll thickness. Information on the physiological mechanisms underlying species-specific photosynthetic responses to ambient air pollution is essential for enhancing the multiple services provided by urban trees. We conducted an on-site investigation of Rhododendron × pulchrum and Ginkgo biloba, which are the most common shrub and tall roadside tree used in Japan, in order to clarify their photosynthetic responses at sites with different air pollution levels in Kyoto city, Japan. The shrub tree R. × pulchrum and tall tree G. biloba exhibited contrasting responses to air pollution mainly from automobile exhaust gas. R. × pulchrum had a lower photosynthetic rate and stomatal conductance at high-pollution sites than at low-pollution sites, while no reductions were observed at high-pollution sites for G. biloba. The stomatal density of R. × pulchrum negatively correlated with atmospheric nitrogen oxides (NO and NO2) concentrations. R. × pulchrum avoided the effects of air pollution by reducing stomatal density at high-pollution sites at the expense of reducing CO2 uptake, while G. biloba appeared to have the ability to tolerate high air pollution levels by reducing the pollution load per mesophyll cell surface area with a low stomatal density and large mesophyll thickness. In conclusion, R. × pulchrum and G. biloba both acclimate to urban environments through an avoidance or tolerance strategy for air pollution by regulating stomatal and/or mesophyll morphologies.

Mesophyll conductance in cotton bracts: anatomically determined internal CO2 diffusion constraints on photosynthesis.

Mesophyll conductance (gm) has been shown to affect photosynthetic capacity and thus the estimates of terrestrial carbon balance. While there have been some attempts to model gm at the leaf and larger scales, the potential contribution of gm to the photosynthesis of non-leaf green organs has not been studied. Here, we investigated the influence of gm on photosynthesis of cotton bracts and how it in turn is influenced by anatomical structures, by comparing leaf palisade and spongy mesophyll with bract tissue. Our results showed that photosynthetic capacity in bracts is much lower than in leaves, and that gm is a limiting factor for bract photosynthesis to a similar extent to stomatal conductance. Bract and the spongy tissue of leaves have lower mesophyll conductance than leaf palisade tissue due to the greater volume fraction of intercellular air spaces, smaller chloroplasts, lower surface area of mesophyll cells and chloroplasts exposed to leaf intercellular air spaces and, perhaps, lower membrane permeability. Comparing bracts with leaf spongy tissue, although bracts have a larger cell wall thickness, they have a similar gm estimated from anatomical characteristics, likely due to the cumulative compensatory effects of subtle differences in each subcellular component, especially chloroplast traits. These results provide the first evidence for anatomical constraints on gm and photosynthesis in non-leaf green organs.

STEREOLOGY, AN UNBIASED METHODOLOGICAL APPROACH TO STUDY PLANT ANATOMY AND CYTOLOGY: PAST, PRESENT AND FUTURE

This review presents an historical overview of stereological methods used for the quantitative evaluation of plant anatomical and cytological structures. It includes the origins of these methods up to the most recent developments such as the application of stereology based on 3D images. We focus especially on leaf, as the vast majority of studies of plant microscopic structure examine this organ. An overview of plant cell ultrastructure measurements as well as plant anatomical characteristics (e.g. plant tissue volume density, internal leaf surface area, number and mean size of mesophyll cells and chloroplast number), which were estimated by stereological methods most frequently, is presented. We emphasize the importance of proper sampling needed for unbiased measurements. Furthermore, we mention other methods used for plant morphometric studies and briefly discuss their relevance, precision, unbiasedness and efficiency in comparison with unbiased stereology. Finally, we discuss reasons for the sparse use of stereology in plant anatomy and consider the future of stereology in plant research.

Anatomical and diffusional determinants inside leaves explain the difference in photosynthetic capacity between Cypripedium and Paphiopedilum, Orchidaceae.

Comparing with other angiosperms, most members within the family Orchidaceae have lower photosynthetic capacities. However, the underlying mechanisms remain unclear. Cypripedium and Paphiopedilum are closely related phylogenetically in Orchidaceae, but their photosynthetic performances are different. We explored the roles of internal anatomy and diffusional conductance in determining photosynthesis in three Cypripedium and three Paphiopedilum species, and quantitatively analyzed their diffusional and biochemical limitations to photosynthesis. Paphiopedilum species showed lower light-saturated photosynthetic rate (A N), stomatal conductance (g s), and mesophyll conductance (g m) than Cypripedium species. A N was positively correlated with g s and g m. And yet, in both species A N was more strongly limited by g m than by biochemical factors or g s. The greater g s of Cypripedium was mainly affected by larger stomatal apparatus area and smaller pore depth, while the less g m of Paphiopedilum was determined by the reduced surface area of mesophyll cells and chloroplasts exposed to intercellular airspace per unit of leaf area, and much thicker cell wall thickness. These results suggest that leaf anatomical structure is the key factor affecting g m, which is largely responsible for the difference in photosynthetic capacity between those two genera. Our findings provide new insight into the photosynthetic physiology and functional diversification of orchids.

Quantitative mesophyll parameters rather than whole-leaf traits predict response of C3 steppe plants to aridity.

We studied the impact of aridity on leaf and mesophyll traits in dominant and very abundant plant species of Eurasian steppe plant communities. We covered a 500-km latitudinal gradient across three vegetation zones in the Volga region of southern European Russia. Whole-leaf traits, volumetric fractions of leaf tissues, quantitative parameters of photosynthetic cells and chloroplasts, and chlorophyll, carbon (C) and nitrogen (N) contents were analyzed and related to plant functional type (PFT), type of mesophyll anatomy, phylogeny and climate aridity. The proportions of prevailing PFTs in the communities, such as C3 monocots, C3 dicots with dorsiventral and isopalisade anatomy and C4 dicots, changed with increasing aridity which influenced the whole-leaf parameters and tissue composition in the leaf. Leaf mass per unit area and leaf thickness slightly increased along the aridity gradient, but the most significant changes were observed in the mesophyll. Mesophyll cell surface area, chloroplast number and chloroplast surface area per unit leaf area were higher in C3 plants growing in the desert steppe compared with those of the forest steppe, while chlorophyll content per single chloroplast and per unit N content as well chlorophyll a/b ratio decreased. Our results identify a suite of mesophyll traits as a typical 'syndrome' of increasingly drought-adapted steppe plants.

Leaf photosynthetic rate and mesophyll cell anatomy changes during ontogenesis in backcrossed indica × japonica rice inbred lines.

The high-yielding indica rice variety, 'Takanari', has the high rate of leaf photosynthesis compared with the commercial japonica varieties. Among backcrossed inbred lines from a cross between 'Takanari' and a japonica variety, 'Koshihikari', two lines, BTK-a and BTK-b, showed approximately 20% higher photosynthetic rate than that of 'Takanari' for a flag leaf at full heading. This is a highest recorded rate of rice leaf photosynthesis. Here, the timing and cause of the increased leaf photosynthesis in the BTK lines were investigated by examining the photosynthesis and related parameters, as well as mesophyll cell anatomy during ontogenesis. Their photosynthetic rate was greater than that of 'Takanari' in the 13th leaf, as well as the flag leaf, but there were no differences in the 7th and 10th leaves. There were no consistent differences in the stomatal conductance, or the leaf nitrogen and Rubisco contents in the 13th and flag leaves. The total surface area of mesophyll cells per leaf area (TAmes) in the 13th and flag leaves increased significantly in the BTK lines due to the increased number and developed lobes of mesophyll cells compared with in 'Takanari'. The mesophyll conductance (g m) became greater in the BTK lines compared with 'Takanari' in the flag leaves but not in the 10th leaves. A close correlation was observed between TAmes and g m. We concluded that the increased mesophyll conductance through the development of mesophyll cells during the reproductive period is a probable cause of the greater photosynthetic rate in the BTK lines.

Coordination of Leaf Photosynthesis, Transpiration, and Structural Traits in Rice and Wild Relatives (Genus Oryza)

The genus Oryza, which includes rice (Oryza sativa and Oryza glaberrima) and wild relatives, is a useful genus to study leaf properties in order to identify structural features that control CO(2) access to chloroplasts, photosynthesis, water use efficiency, and drought tolerance. Traits, 26 structural and 17 functional, associated with photosynthesis and transpiration were quantified on 24 accessions (representatives of 17 species and eight genomes). Hypotheses of associations within, and between, structure, photosynthesis, and transpiration were tested. Two main clusters of positively interrelated leaf traits were identified: in the first cluster were structural features, leaf thickness (Thick(leaf)), mesophyll (M) cell surface area exposed to intercellular air space per unit of leaf surface area (S(mes)), and M cell size; a second group included functional traits, net photosynthetic rate, transpiration rate, M conductance to CO(2) diffusion (g(m)), stomatal conductance to gas diffusion (g(s)), and the g(m)/g(s) ratio.While net photosynthetic rate was positively correlated with gm, neither was significantly linked with any individual structural traits. The results suggest that changes in gm depend on covariations of multiple leaf (S(mes)) and M cell (including cell wall thickness) structural traits. There was an inverse relationship between Thick(leaf) and transpiration rate and a significant positive association between Thick(leaf) and leaf transpiration efficiency. Interestingly, high g(m) together with high g(m)/g(s) and a low S(mes)/g(m) ratio (M resistance to CO(2) diffusion per unit of cell surface area exposed to intercellular air space) appear to be ideal for supporting leaf photosynthesis while preserving water; in addition, thick M cell walls may be beneficial for plant drought tolerance.

Comparative analyses of leaf anatomy of dicotyledonous species in Tibetan and Inner Mongolian grasslands

Knowledge of the leaf anatomy of grassland plants is crucial for understanding how these plants adapt to the environment. Tibetan alpine grasslands and Inner Mongolian temperate grasslands are two major grassland types in northern China. Tibetan alpine grasslands occur in high-altitude regions where the low temperatures limit plant growth. Inner Mongolian temperate grasslands are found in arid regions where moisture is the limiting factor. Few comparative studies concerning the leaf anatomy of grassland plants of the Tibetan Plateau and Inner Mongolian Plateau have been conducted. We examined leaf characteristics at 71 sites and among 65 species, across the alpine grasslands of the Tibetan Plateau and the temperate grasslands of the Inner Mongolian Plateau. We compared the leaf structures of plants with different life forms and taxonomies, and their adaptation to arid or cold environments. We explored relationships among leaf features and the effects of climatic factors (i.e., growing season temperature and precipitation) on leaf characteristics. Our results showed that (i) there were significant differences in leaf anatomy between Tibetan alpine and Inner Mongolian temperate grasslands. Except for mesophyll cell density, the values obtained for thickness of leaf tissue, surface area and volume of mesophyll cells were larger on the Tibetan Plateau than on the Inner Mongolian Plateau. (ii) Within the same family or genus, leaf anatomy showed significant differences between two regions, and trends were consistent with those of whole species. (iii) Leaf anatomy of woody and herbaceous plants also showed significant differences between the regions. Except for mesophyll cell density, the values obtained for the thickness of leaf tissue, and the surface area and volume of mesophyll cells were larger in herbaceous than in woody plants. (iv) Leaf anatomical traits changed accordingly. Total leaf thickness, thicknesses of lower and upper epidermal cells, and surface area and volume of mesophyll cells were positively correlated, while mesophyll cell density was negatively associated with those traits. (v) Growing season temperature had stronger effects on leaf anatomy than growing season precipitation. Although the communities in Tibetan and Inner Mongolian grasslands were similar in appearance, leaf anatomy differed; this was probably due to the combined effects of evolutionary adaptation of plants to environment and environmental stress induced by climatic factors.

Variation in mesophyll anatomy and photosynthetic capacity during leaf development in a deciduous mesophyte fruit tree (Prunus persica) and an evergreen sclerophyllous Mediterranean shrub (Olea europaea)

The relative importance that biomechanical and biochemical leaf traits have on photosynthetic capacity would depend on a complex interaction of internal architecture and physiological differences. Changes in photosynthetic capacity on a leaf area basis and anatomical properties during leaf development were studied in a deciduous tree, Prunus persica, and an evergreen shrub, Olea europaea. Photosynthetic capacity increased as leaves approached full expansion. Internal CO2 transfer conductance (g i) correlated with photosynthetic capacity, although, differences between species were only partially explained through structural and anatomical traits of leaves. Expanding leaves preserved a close functional balance in the allocation of resources of photosynthetic component processes. Stomata developed more rapidly in olive than in peach. Mesophyll thickness doubled from initial through final stages of development when it was twice as thick in olive as in peach. The surface area of mesophyll cells exposed to intercellular air spaces per unit leaf area tended to decrease with increasing leaf expansion, whereas, the fraction of mesophyll volume occupied by the intercellular air spaces increased strongly. In the sclerophyllous olive, structural protection of mesophyll cells had priority over efficiency of photochemical mechanisms with respect to the broad-leaved peach. The photosynthetic capacity of these woody plants during leaf development relied greatly on mesophyll properties, more than on leaf mass per area ratio (LMA) or nitrogen (N) allocation. Age-dependent changes in diffusion conductance and photosynthetic capacity affected photosynthetic relationships of peach versus olive foliage, evergreen leaves maturing functionally and structurally a bit earlier than deciduous leaves in the course of adaptation for xeromorphy.

Carbon dioxide uptake, water relations and drought survival for Dudleya saxosa, the ‘rock live‐forever’, growing in small soil volumes

Summary Although many plants grow in rock crevices and other regions of small soil volume, including over 20 000 epiphytic and hemi‐epiphytic species, analyses of the actual soil volume occupied, the water availability in that soil, the water‐storage capacity in the shoots and underground organs, and the photosynthetic pathway utilized have rarely been combined. Dudleya saxosa (M.F. Jones) Britton and Rose (Crassulaceae), growing in the Sonoran Desert, has very shallow roots that occupied soil volumes averaging only 43 × 10−6 m3 per medium‐sized plant. This volume of soil can hold about the same amount of water (10 g) as can be stored in the leaves, corm and roots combined (11 g), but at a sufficiently high water potential for transfer to the plant for less than 1 week after a substantial rainfall. About 80% of the net carbon dioxide uptake by D. saxosa over a 24‐h period occurred during the daytime (C3) under wet conditions, the daily total decreasing by 34% and the pattern shifting to nocturnal net CO2 uptake (CAM) after 46 days’ drought. Seventy‐seven days’ drought eliminated its daily net CO2 uptake. Stomatal frequency was only 67 mm−2 on the adaxial (upper) surface and twofold lower on the abaxial surface. The cuticle was thick, 34 µm for the adaxial surface. Leaves had 24 mesophyll cell layers, leading to a high mesophyll cell surface area per unit leaf area of 142. The three leaf anatomical features plus utilization of CAM increased net CO2 uptake per unit of water transpired, and helped D. saxosa thrive in a small soil volume, with the underground corm being a major supplier of water to the succulent leaves during 2.5 months of drought. The maximum water‐holding capacity of the soil explored by the roots closely matched the maximum water‐holding capacity of the plant, reflecting the conservative strategy used by D. saxosa in a stressful semi‐arid environment.

Morpho-anatomical and physiological leaf traits of two alpine herbs, Podophyllum hexandrum and Rheum emodi in the Western Himalaya under different irradiances

Morpho-anatomical leaf traits and photosynthetic activity of two alpine herbs, Podophyllum hexandrum (shade-tolerant) and Rheum emodi (light-requiring), were studied under field (PAR>2 000 µmol m−2 s−1) and greenhouse (PAR 500 µmol m−2 s−1) conditions. Mesophyll thickness, surface area of mesophyll cells facing intercellular spaces (Smes), surface area of chloroplasts facing intercellular spaces (Sc), intercellular spaces of mesophyll cells (porosity), photon-saturated rate of photosynthesis per unit leaf area (P Nmax), and ribulose-1,5-bisphosphate carboxylase/oxygenase activity decreased in the greenhouse with respect to the field and the decreases were significantly higher in R. emodi than in P. hexandrum. P. hexandrum had lower intercellular CO2 concentration than R. emodi under both irradiances. The differences in acclimation of the two alpine herbs to low irradiance were due to their highly unlikely changes in leaf morphology, anatomy, and P Nmax which indicated that the difference in radiant energy requirement related to leaf acclimation had greater impact under low than high irradiance.

Leaf anatomy as a constraint for photosynthetic acclimation: differential responses in leaf anatomy to increasing growth irradiance among three deciduous trees

ABSTRACTInterspecific variation in the response to transfer from low to high growth irradiance with respect to anatomical and photosynthetic characteristics was studied in mature leaves of three tree species, Betula ermanii Cham., Acer rufinerve Sieb. et Zucc. and Fagus crenata Blume, which occur in different successional stages in temperate deciduous forests. Transfer from low to high irradiance increased the light‐saturated rate of photosynthesis per unit leaf area (Pmax) significantly in B. ermanii and A. rufinerve, but not in F. crenata. Leaves of B. ermanii grown at low irradiance were relatively thick and had vacant spaces along the mesophyll cell surfaces which was not occupied by chloroplasts or other organelles. After transfer to high irradiance, chloroplasts enlarged to fill the space along with Pmax without an increase in leaf thickness. Leaves of A. rufinerve were plastic in mesophyll cell surface area and in leaf thickness, both of which increased after the transfer to high irradiance, along with an increase in the amount of chloroplasts and in Pmax. On the other hand, F. crenata had little mesophyll cell surface unoccupied by chloroplasts and leaf anatomy was not changed after the transfer. In all species, Pmax was strongly correlated with chloroplast surface area adjacent to the exposed mesophyll surface across different growth irradiances. An increase in Pmax was observed only when chloroplast volume also increased. We conclude that light acclimation potential is primarily determined by the availability of unoccupied cell surface into which chloroplasts expand, as well as by the plasticity of the mesophyll that allows an increase in its surface area.

Interspecific variation of photosynthesis and leaf characteristics in canopy trees of five species of Dipterocarpaceae in a tropical rain forest.

Photosynthetic rate, nitrogen concentration and morphological properties of canopy leaves were studied in 18 trees, comprising five dipterocarp species, in a tropical rain forest in Sarawak, Malaysia. Photosynthetic rate at light saturation (Pmax) differed significantly across species, varying from 7 to 18 micro mol m(-2) s(-1). Leaf nitrogen concentration and morphological properties, such as leaf blade and palisade layer thickness, leaf mass per area (LMA) and surface area of mesophyll cells per unit leaf area (Ames/A), also varied significantly across species. Among the relationships with leaf characteristics, Pmax had the strongest correlation with leaf mesophyll parameters, such as palisade cell layer thickness (r2 = 0.76, P < 0.001) and Ames/A (r2 = 0.73, P < 0.001). Leaf nitrogen concentration and Pmax per unit area also had a significant but weaker correlation (r2 = 0.46, P < 0.01), whereas Pmax had no correlation, or only weakly significant correlations, with leaf blade thickness and LMA. Shorea beccariana Burck, which had the highest P(max) of the species studied, also had the thickest palisade layer, with up to five or more layers. We conclude that interspecific variation in photosynthetic capacity in tropical rain forest canopies is influenced more by leaf mesophyll structure than by leaf thickness, LMA or leaf nitrogen concentration.

Development and structure of drought-tolerant leaves of the Mediterranean shrub Capparis spinosa L.

Capparis spinosa (caper), a winter-deciduous perennial shrub, is a consistent floristic element of Mediterranean ecosystems, growing from May to October, i.e. entirely during the prolonged summer drought. The internal architecture of young and fully expanded leaves was studied, along with certain physiological characteristics. Capparis spinosa possesses thick, amphistomatic and homobaric leaves with a multilayered mesophyll. The latter possesses an increased number of photosynthesizing cells per unit leaf surface, a large surface area of mesophyll cells facing intercellular spaces (Smes) and a low percentage of intercellular space per tissue volume. Smes and chlorophyll content attain their maximum values synchronously, slightly before full leaf expansion. Nitrogen investment is also completed before full leaf expansion. The structural features, in combination with the water status, could contribute to enhanced rates of transpiration and photosynthesis under field water shortage conditions.