Mesophyll cell properties for some C3 and C4 species with high photosynthetic rates

Leaves of twelve C₃ species and six C₄ species were examined to understand better the relationship between mesophyll cell properties and the generally high photosynthetic rates of these plants. The CO₂ diffusion conductance expressed per unit mesophyll cell surface area (g_CO2^cell) cell was determined using measurements of the net rate of CO₂ uptake, water vapor conductance, and the ratio of mesophyll cell surface area to leaf surface area (A^mes/A). A^mes/A averaged 31 for the C₃ species and 16 for the C₄ species. For the C₃ species g_CO2^cell ranged from 0.12 to 0.32 mm s^-1, and for the C₄ species it ranged from 0.55 to 1.5 mm s^-1, exceeding a previously predicted maximum of 0.5 mm s^-1. Although the C₃ species Cammissonia claviformis did not have the highest g_CO2^cell, the combination of the highest A^mes and highest stomatal conductance resulted in this species having the greatest maximum rate of CO₂ uptake in low oxygen, 93 μmol m^-2 s^-1 (147 mg dm^-2 h^-1). The high g_CO2^cell of the C₄ species Amaranthus retroflexus (1.5 mm s^-1) was in part attributable to its thin cell wall (72 nm thick).     

Influence of differences in leaf anatomy on net photosynthetic rates of some cultivars of Cassava

Gas exchange measurements and leaf anatomy of 10 cassava cultivars were conducted to study the interrelationship between the relatively high photosynthetic rates and the factors limiting internal CO₂ diffusion. The internal mesophyll surface area per unit leaf surface area (A^mes/A) and the intracellular components of CO₂ diffusion and fixation resistance (R^cellCO₂) were determined. Among the group of cultivars tested net CO₂ exchange rates were 26±2.5 mol CO₂ m^–2 s^–1 in normal air and intense light and A^mes/A ranged from 14 to 38. Estimated R^cellCO₂ ranged from 4300 to 13,000 s m^–1. The combined and compensating effects of A^mes/A and R^cellCO₂ accounted for both the high net photosynthetic rates (P_n) and the lack of large differences in P_n among cultivars.     

Relation between Mesophyll Surface Area, Photosynthetic Rate, and Illumination Level during Development for Leaves of Plectranthus parviflorus Henckel

The influence of illumination level during leaf development on the mesophyll cell surface area per unit leaf area (A^mes/A), CO₂ resistances, and the photosynthetic rate was determined for leaves of Plectranthus parviflorus Henckel. The relative importance of A^mes/A versus CO₂ resistances in accounting for observed changes in photosynthesis was quantitatively evaluated using equations based on analogies to electrical circuits.     

Internal Leaf Area and Cellular CO2 Resistance: Photosynthetic Implications of Variations with Growth Conditions and Plant Species

The influences of illumination, temperature, and soil water potential during development on leaf thickness, mesophyll cell wall area per unit leaf area (^Ames/A), and the cellular CO₂, resistance expressed on a mesophyll cell wall area basis (r_CO2^cell,) were examined for Plectranthus parviflorus Henckel. Although the ranges of all three growth conditions caused at least 9-fold variations in the leaf biomass produced in 4 weeks, only the illumination had a major effect on internal leaf morphology, e.g. the thickness went from 279 to 831 μm and A^mes/A from 10.5 to 34.8 as the photosynthetically active radiation was raised from 3 to 53 nEinsteins cm^?2 s^?1, while r_CO2^cell remained close to 154 s cm^?1. Variations in the growth temperature, soil water potential, and the nutritional status of the plant, affected photosynthesis mainly by changes in r_CO2^cell. To compare the influence of internal leaf area on photosynthesis for other plants, especially those with low A^mes/A values, the maximum rates of CO₂ uptake at light saturation and photosynthetically optimal temperatures were also determined for a moss, Mnium ciliare (C. Muell.) Schimp., and two ferns, Adiantum decorum Moore and Alsophila australe R. Br. As Ames/A went from 2.00 for the moss to 3.8, 7.5, 11.7, and 20.8 for the fens, the illumination at light saturation and the maximum rate of photosynthesis both progressively increased. The cellular CO₂ resistance, which theoretically might have a lower limit of 20 s cm^?1, ranged from 85 to 190 s cm^?1.     

Changes in Morphology,Anatomy, and Photosynthetic Capacity of Needles of Japanese Larch (Larix kaempferi) Seedlings Grown in High CO2 Concentrations

Photosynthetic traits of two-year-old Japanese larch seedlings (Larix kaempferi Carr.) grown at elevated CO₂ concentrations were studied in relation to structural changes in the needles. Seedlings were grown at two CO₂ concentrations, 360 (AC) and 720 (EC) mol mol^–1 at high and low nutrient supply rates, high N (HN) and low N (LN). The photosynthetic capacity fell significantly in EC+LN, but increased significantly in EC+HN. Since the mesophyll surface area exposed to intercellular space per unit leaf area (A^mes/A) is correlated with the photosynthetic rate, we measured A^mes/A for larch needles growing in EC. Changes of A^mes/A in both EC+HN and EC+LN were very similar to the changes in photosynthetic capacity. This suggests that the changes of A^mes/A in EC probably caused the changes in the photosynthetic capacity. The changes of A^mes/A in EC were attributed to changes in the mesophyll cell size and mesophyll cell number. The photosynthetic capacity in EC can be explained by taking morphological and structural adaptations into account as well as biochemical factors.     

Determination of mesophyll diffusion resistance in Chamaerion angustifolium by the method of three-dimensional reconstruction of the leaf cell packing

A detailed quantitative analysis of the three-dimensional organization of the mesophyll was performed, and mesophyll diffusion resistance to CO₂ in the leaves of Chamaerion angustifolium formed under different irradiance was calculated using an original method of stereometric cellular packing. For each type of leaves (sun and shade), we determined structural components of gas exchange: the volume of mesophyll per unit leaf area (V _mes), the volume of the intercellular space in the mesophyll (V _is), the area of the total mesophyll surface (S), the area of the free mesophyll surface facing the intercellular spaces (S _mes), and the ratios of the total and the free mesophyll surfaces to its volume (S/V and S _mes/V). As compared with sun leaves, in the shade leaves of Ch. angustifolium, S and V _mes decreased twofold, tissue density was reduced twofold, and the share of the intercellular space in the mesophyll rose from 49 to 72%. In shade, the diffusion resistance of the mesophyll increased by 1.8 times because of changes in the leaf structure. At the same time, the ratio S _mes/V was found to increase by 1.4 times, which facilitated the diffusion of CO₂. In the shade leaves of Ch. angustifolium, the diffusion resistance of the intercellular air spaces was reduced twofold as a result of an increase in their share in the leaf mesophyll and simplification of their geometry. Thus, the method of three-dimensional reconstruction of sun and shade leaves of Ch. angustifolium showed a comprehensive rearrangement of the mesophyll spatial organization in shade and revealed the structural mechanisms of changes in the resistance to CO₂ diffusion within the leaf.     

Nutrient Influences on Leaf Photosynthesis: EFFECTS OF NITROGEN, PHOSPHORUS, AND POTASSIUM FOR GOSSYPIUM HIRSUTUM L

The net rate of CO₂ uptake for leaves of Gossypium hirsutum L. was reduced when the plants were grown at low concentrations of NO₃^-, PO₄^2-, or K⁺. The water vapor conductance was relatively constant for all nutrient levels, indicating little effect on stomatal response. Although leaves under nutrient stress tended to be lower in chlorophyll and thinner, the ratio of mesophyll surface area to leaf area did not change appreciably. Thus, the reduction in CO₂ uptake rate at low nutrient levels was due to a decrease in the CO₂ conductance expressed per unit mesophyll cell wall area (g^cell_CO2). The use of g^cell_CO2 and nutrient levels expressed per unit of mesophyll cell wall provides a new means of assessing nutrient effects on CO₂ uptake of leaves.     

Phospholipid synthesis and exchange in castor bean endosperm homogenates

Leaves on a bush of Hyptis emoryi Torr. varied in length from less than 1 cm when development occurred in full sunlight (e.g. 40 Mjoules m⁻²) to over 7 cm when the total daily solar irradiance was less than 3 Mjoules m⁻². The 1-cm sun leaves were 3-fold higher than the 7-cm shade leaves in chlorophyll per unit area, mesophyll thickness, and the internal to external leaf area ratio (A^mes/A). The higher A^mes/A caused a 1.2-cm leaf to have a 3-fold lower CO₂ liquid phase resistance than did a 7.1-cm leaf. Large thin shade leaves captured photosynthetically active radiation effectively (less than 7% passed through), but were not adapted to full sunlight. Specifically, when a 6.9-cm leaf was placed at 910 w m⁻² for 30 min, its temperature exceeded that of the air by nearly 8 C. For the common daytime air temperatures above 30 C for this desert shrub, large shade leaves would have temperatures far in excess of that optimum for photosynthesis for H. emoryi, 29 to 32 C.     

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.     

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.     

Resistance Analysis of Nocturnal Carbon Dioxide Uptake by a Crassulacean Acid Metabolism Succulent, Agave deserti

Nocturnal CO₂ uptake by a Crassulacean acid metabolism succulent, Agave deserti Engelm. (Agavaceae), was measured so that the resistance properties of the mesophyll chlorenchyma cells and their CO₂ concentrations could be determined. Two equivalents of acidity were produced at night per mole of CO₂ taken up. The nocturnal CO₂ uptake became light-saturated at 3.5 mEinsteins cm⁻² of photosynthetically active radiation (400-700 nm) incident during the preceding day; at least 46 Einsteins were required per mole of CO₂ fixed. Variations in the daytime leaf temperature between 20 and 37 C had little effect on nocturnal CO₂ uptake. After the first few hours in the dark, the leaf liquid phase CO₂ resistance (r^liq_CO2) and the CO₂ concentration in the chlorenchyma cells (cⁱ_CO2) both increased, the latter usually reaching the ambient external CO₂ level at the end of the dark period. Increasing the leaf surface temperature above 15 C at night markedly increased the stomatal resistance, r^liq_CO2, and cⁱ_CO2.

The minimum r^liq_CO2 at night was about 1.6 seconds cm⁻¹. Based on the ratio of chlorenchyma surface area to total leaf surface area of 82, this r^liq_CO2 corresponded to a minimum cellular resistance of approximately 130 seconds cm⁻¹, comparable to values for mesophyll cells of C₃ plants. The contribution of the carboxylation reaction and/or other biochemical steps to r^liq_CO2 may increase appreciably as the nighttime temperature shifts a few degrees from the optimum or after a few hours in the dark, both of which caused large increases in r^liq_CO2. This necessitates a large internal leaf area for CO₂ diffusion into the chlorenchyma to support moderate nocturnal CO₂ uptake rates by these succulent leaves.

     

RELATIONSHIPS BETWEEN PHOTOSYNTHETIC CAPACITY AND LEAF STRUCTURE IN SEVERAL SHADE PLANTS

Leaf structure, photosynthetic characteristics and related physiological parameters have been studied in three ornamental shade species: Fatsia japonica, Cissus rhombifolia (relatively light-tolerant plants), and Philodendron scandens (obligate shade plant). Species were grown in a shadehouse. Maximum photosynthetic photon flux density was 470 μmol m^-2 s^-1. Net rate of CO₂ uptake at light saturation (maximum Pn) in Fatsia was 6.90 ± 1.27 μmol m^-2 s^-1. In Cissus and Philodendron values were about 30% and 63% less respectively, than those measured in Fatsia. The nitrogen content, relative dry wt, specific leaf dry wt (SLDW), chlorophyll a/b ratio, and nitrogen to chlorophyll ratio were lower in Philodendron. However, leaf thickness in Philodendron (296 ± 17 μm) was about 54% and 160% higher, respectively, than in Fatsia and Cissus, and the ratio between mesophyll cell area and leaf surface area (A^mes/A) was nearly similar in the three species. However Philodendron exhibited a percentage of palisade parenchyma about three times lower than that observed in the two other species. The chloroplast number per mm of cell wall in transverse sections (chloroplast density) in the palisade parenchyma was fairly constant (about 65), irrespective of species. The “chloroplast density” in the spongy parenchyma of Philodendron was about 53% and 63%, respectively, of Fatsia and Cissus values. In Fatsia and Cissus chloroplast ultrastructure seems to change gradually and continuously from sun to shade type with the depth from the adaxial to abaxial surface. Special emphasis was given in order to determine the structural parameters best correlated with maximum Pn between the different species. In this way chloroplast number in transverse sections (chloroplast number) and the ratio between chloroplast area and leaf surface area (A^chl/A) were the parameters best correlated with maximum Pn, and stomatal frequency was also a good determinant of maximum Pn. However, leaf thickness, SLDW, and even A^mes/A ratio were weakly correlated with maximum Pn.     

Mesostructure and activity of photosynthetic apparatus for three crassulacean species grown in cold climate

A comparative study of leaf anatomy and morphology and of CO₂ exchange was conducted with Rhodiola rosea L., Hylotelephium triphyllum (Haw.) Holub., and Sedum acre L. as representative Crassulacean species occurring in the northeast European Russia. The leaf mesophyll in R. rosea was clearly differentiated into the palisade and spongy tissues, whereas the mesophyll of stonecrops (H. triphyllum and S. acre) was composed of round-shaped cells. The leaves of S. acre featured the largest volume of mesophyll cells and possessed water-retaining cells located around conducting bundles. The chloroplast volume in S. acre (50 μm³) was three times smaller and the number of chloroplasts per cell (170 cell^?1) was three times higher than in R. rosea and H. triphyllum (50–55 cell^?1). The content of chlorophylls (5–7 mg/g dry wt) and carotenoids (1.5–2.0 mg/g dry wt) in R. rosea leaves was 2–3 times higher than in leaves of stonecrops. The rate of CO₂ net uptake in Crassulacean species depended on mesostructure and correlated with the content of pigments and soluble carbohydrates. The photosynthetic rate in R. rosea under optimal irradiance and temperature attained the value of 40 mg/(g dry wt), which is 3 and 8 times higher than in H. triphyllum and S. acre, respectively. The temperature optimum for photosynthesis of R. rosea was observed at 8–18°C, while the optimum for stonecrops was shifted towards higher temperatures by 3–5°C. At chilling temperatures (5–7°C), the leaves of R. rosea retained 50% of their maximal photosynthetic rate, while photosynthetic rates in H. triphyllum and S. acre leaves lowered to 25–30% of the maximal rate. The increase in temperature to 25–30°C led to depression of CO₂ net uptake in leaves of Crassulacean species. In R. rosea and H. triphyllum, the rate of photosynthetic electron flow was depressed at high irradiances and temperatures that were supraoptimal for net photosynthesis. It is concluded that the photosynthetic apparatus of Crassulacean species is well adapted to moderate and chilling temperatures, which adjusts the plant metabolism to “life strategies” under conditions of cold climate.     

Correlation of Stomatal Conductance with Photosynthetic Capacity of Cotton Only in a CO2-Enriched Atmosphere: Mediation by Abscisic Acid?

Some evidence indicates that photosynthetic rate (A) and stomatal conductance (g) of leaves are correlated across diverse environments. The correlation between A and g has led to the postulation of a “messenger” from the mesophyll that directs stomatal behavior. Because A is a function of intercellular CO₂ concentration (c_i), which is in turn a function of g, such a correlation may be partially mediated by c_i if g is to some degree an independent variable. Among individual sunlit leaves in a cotton (Gossypium hirsutum L.) canopy in the field, A was significantly correlated with g (r² = 0.41, n = 63). The relative photosynthetic capacity of each leaf was calculated as a measure of mesophyll properties independent of c_i. This approach revealed that, in the absence of c_i effects, mesophyll photosynthetic capacity was unrelated to g (r² = 0.06). When plants were grown in an atmosphere enriched to about 650 microliters per liter of CO₂, however, photosynthetic capacity remained strongly correlated with g even though the procedure discounted any effect of variable c_i. This “residual” correlation implies the existence of a messenger in CO₂-enriched plants. Enriched CO₂ also greatly increased stomatal response to abscisic acid (ABA) injected into intact leaves. The data provide no evidence for a messenger to coordinate g with A at ambient levels of CO₂. In a CO₂-enriched atmosphere, though, ABA may function as such a messenger because the sensitivity of the system to ABA is enhanced.     

Leaf expansion and carbon assimilation in cotton leaves grown at two photosynthetic photon flux densities

Increasing photosynthetic photon flux density (PPFD) received during development from 5.5 to 31.2 mol m^-2 d^-1 resulted in greater leaf and mesophyll cell surface areas in cotton (Gossypium hirsutum L.). The relationships between the amounts of these surface areas and potential CO₂ assimilation by these leaves were evaluated. Leaf area (epidermal surface area of one side of a leaf), mesophyll cell surface area, and net rate of CO₂ uptake (P_n) were measured from the time leaves first unfolded until P., was substantially reduced. At the higher PPFD, leaf and mesophyll surface areas increased more rapidly during expansion, and P_n per unit leaf area was greater than at the lower PPFD. Although leaves at the higher PPFD reached the maximum P., per unit mesophyll cell surface area 4 to 5 days earlier than leaves at the lower PPFD, the maxima for these P., were similar. Leaves grown at the higher PPFD had the potential to assimilate 2.2, 3.5, or 5.8 times the amount of CO₂ as leaves from the lower PPFD when P., was expressed per unit mesophyll surface, per unit leaf surface, or per whole leaf, respectively. Greater and earlier development of both P., and mesophyll cell surface area at higher PPFD apparently had a compounding effect on the potential for carbon assimilation by a leaf.     

Estimation of Mesophyll Conductance to CO(2) Flux by Three Different Methods

The resistance to diffusion of CO₂ from the intercellular airspaces within the leaf through the mesophyll to the sites of carboxylation during photosynthesis was measured using three different techniques. The three techniques include a method based on discrimination against the heavy stable isotope of carbon, ¹³C, and two modeling methods. The methods rely upon different assumptions, but the estimates of mesophyll conductance were similar with all three methods. The mesophyll conductance of leaves from a number of species was about 1.4 times the stomatal conductance for CO₂ diffusion determined in unstressed plants at high light. The relatively low CO₂ partial pressure inside chloroplasts of plants with a low mesophyll conductance did not lead to enhanced O₂ sensitivity of photosynthesis because the low conductance caused a significant drop in the chloroplast CO₂ partial pressure upon switching to low O₂. We found no correlation between mesophyll conductance and the ratio of internal leaf area to leaf surface area and only a weak correlation between mesophyll conductance and the proportion of leaf volume occupied by air. Mesophyll conductance was independent of CO₂ and O₂ partial pressure during the measurement, indicating that a true physical parameter, independent of biochemical effects, was being measured. No evidence for CO₂-accumulating mechanisms was found. Some plants, notably Citrus aurantium and Simmondsia chinensis, had very low conductances that limit the rate of photosynthesis these plants can attain at atmospheric CO₂ level.     

Why are Sun Leaves Thicker than Shade Leaves? — Consideration based on Analyses of CO2 Diffusion in the Leaf

A ) depend not only on photosynthetic biochemistry but also on mesophyll structure. Because resistance to CO₂ diffusion from the substomatal cavity to the stroma is substantial, it is likely that mesophyll structure affects A through affecting diffusion of CO₂ in the leaf. To evaluate effects of various aspects of mesophyll structure on photosynthesis, we constructed a one-dimensional model of CO₂ diffusion in the leaf. When mesophyll thickness of the leaf is changed with the Rubisco content per unit leaf area kept constant, the maximum A occurs at an almost identical mesophyll thickness irrespective of the Rubisco contents per leaf area. On the other hand, with an increase in Rubisco content per leaf area, the mesophyll thickness that realizes a given photosynthetic gain per mesophyll thickness (or per leaf cost) increases. This probably explains the strong relationship between A and mesephyll thickness. In these simulations, an increase in mesophyll thickness simultaneously means an increase in the diffusional resistance in the intercellular spaces (R _ias), an increase in the total surface area of chloroplasts facing the intercellular spaces per unit leaf area (S _c ), and an increase in construction and maintenance cost of the leaf. Leaves can increase S _c and decrease R _ias also by decreasing cell size. Leaves with smaller cells are mechanically stronger. However, actual leaves do not have very small cells. This could be because actual leaves exhibiting considerable rates of leaf area expansion, adequate heat capacitance, high efficiency of N and/or P use, etc, are favoured. Relationships between leaf longevity and mesophyll structure are also discussed. Received 20 September 2000/ Accepted in revised form 4 January 2001     

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 (S_mes), surface area of chloroplasts facing intercellular spaces (S_c), 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.     

INFLUENCE OF IRRADIATION,SOIL WATER POTENTIAL,AND LEAF TEMPERATURE ON LEAF MORPHOLOGY OF A DESERT BROADLEAF,ENCELIA FARINOSA GRAY (COMPOSITAE)

Laboratory experiments were performed to evaluate observed seasonal changes in leaf morphology of the desert perennial shrub, Encelia farinosa Gray. Plants were grown under low or high conditions of photosynthetically active irradiation, soil water potential (Ψ^soil), and leaf temperature (8 different experimental regimes). The relative growth rate, leaf water vapor conductance, leaf water potential, and leaf length were all greater for the high Ψ^soil regimes, the largest leaves occurring at low irradiation. High irradiation during growth led to thicker leaves with a higher internal to external leaf area ratio (A^mes/A); low Ψ^soil tended to increase A^mes/A somewhat. High irradiation also led to decreased absorptance to solar irradiation caused by increased pubescence. High leaf temperature during development resulted in slightly smaller, thicker leaves with higher A^mes/A. Thus, irradiation appeared to have its major influence on leaf thickness, A^mes/A, and absorptance, with a secondary effect on leaf length; Ψ^soil affected primarily leaf length, growth rate, and water status, and secondarily A^mes/A. Results are discussed with regard to recent ecophysiological studies on the observed seasonal changes in leaf morphology of E. farinosa.     

Light effects on leaf development and photosynthetic capacity of Hydrocotyle bonariensis Lam.

Rates of net CO₂ uptake were examined in developing leaves of Hydrocotyle bonariensis. Leaves that developed under high photosynthetically active radiation (48 mol m^-2 day^-1 PAR) were smaller, thicker, and reached maximum size sooner than did leaves that developed under low PAR (4.8 mol m^-2 day^-1). Maximum net CO₂ uptake rates were reached after 5 to 6 days expansion for both the low and the high PAR leaves. Leaves grown at high PAR had higher maximum photosynthetic rates and a higher PAR required for light saturation but showed a more rapid decline in rate with age than did low PAR leaves. To assess the basis for the difference observed in photosynthetic rates, CO₂ diffusion conductances and the mesophyll surface available for CO₂ absorption were examined for mature leaves. Stomatal conductance was the largest conductance in all treatments and did not vary appreciably with growth PAR. Mesophyll conductance progressively increased with growth PAR (up to 48 mol m^-2 day^-1) as did the mesophyll surface area per unit leaf area, but the cellular conductance exhibited most of its increase at low PAR (up to 4.8 mol m^-2 day^-1).