Growth of autotrophic plants is directly and dramatically influenced by the intensity of light — the driving force of photosynthesis — which provides nearly all of the carbon and chemical energy needed for plant growth. Moreover, light intensity (quantum flux density) is perhaps the most conspicuous environmental variable with which plants must cope. Contrasting terrestrial habitats may differ by at least two orders of magnitude in the daily quantum flux available for photosynthesis. In any habitat the quantum flux density varies seasonally, diurnally, and spatially (such as within a canopy of a given plant stand).

Responses to Different Quantum Flux Densities

The photosynthetic CO2-fixation rates, chlorophyll content, chloroplast ultrastructure and other leaf characteristics (e.g. variable fluorescence, stomata density, soluble carbohydrate content) were studied in a comparative way in sun and shade leaves of beech (Fagus sylvatica) and in high-light and low-light seedlings. 1. Sun leaves of the beech possess a smaller leaf area, higher dry weight, lower water content, higher stomata density, higher chlorophyll a/b ratios and are thicker than the shade leaves. Sun leaves on the average contain more chlorophyll in a leaf area unit; the shade leaf exhibits more chlorophyll on a dry weight basis. Sun leaves show higher rates for dark respiration and a higher light saturation of photosynthetic CO2-fixation. Above 2000 lux they are more efficient in photosynthetic quantum conversion than the shade leaves. 2. The development of HL-radish plants proceeds much faster than that of LL-plants. The cotyledons of HL-plants show a higher dry weight, lower water content, a higher ratio of chlorophyll a/b and a higher gross photosynthesis rate than the cotyledons of the LL-plants, which possess a higher chlorophyll content per dry weight basis. The large area of the HL-cotyledon on the one hand, as well as the higher stomata density and the higher respiration rate in the LL-cotyledon on the other hand, are not in agreement with the characteristics of sun and shade leaves respectively. 3. The development, growth and wilting of wheat leaves and the appearance of the following leaves (leaf succession) is much faster at high quanta fluence rates than in weak light. The chlorophyll content is higher in the HL-leaf per unit leaf area and in the LL-leaf per g dry weight. There are no differences in the stomata density and leaf area between the HL- and LL-leaf. There are fewer differences between HL- and LL-leaves than in beech or radish leaves. 4. The chloroplast ultrastructure of shade-type chloroplasts (shade leaves, LL-leaves) is not only characterized by a much higher number of thylakoids per granum and a higher stacking degree of thylakoids, but also by broader grana than in sun-type chloroplasts (sun leaves, HL-leaves). The chloroplasts of sun leaves and of HL-leaves exhibit large starch grains. 5. Shade leaves and LL-leaves exhibit a higher maximum chlorophyll fluorescence and it takes more time for the fluorescence to decline to the steady state than in sun and HL-leaves. The variable fluorescence VF (ratio of fluorescence decrease to steady state fluorescence) is always higher in the sun and HL-leaf of the same physiological stage (maximum chlorophyll content of the leaf) than in the shade and LL-leaf. The fluorescence emission spectra of sun and HL-leaves show a higher proportion of chlorophyli fluorescence in the second emission maximum F2 than shade and LL-leaves. 6. The level of soluble carbohydrates (reducing sugars) is significantly higher in sun and HL-leaves than in shade and LL-leaves and even reflects changes in the amounts of the daily incident light. 7. Some but not all characteristics of mature sun and shade leaves are found in HL- and LL-leaves of seedlings. Leaf thickness, dry weight, chlorophyll content, soluble carbohydrate level, photosynthetic CO2-fixation, height and width of grana stacks and starch content, are good parameters to describe the differences between LL- and HL-leaves; with some reservations concerning age and physiological stage of leaf, a/b ratios, chlorophyll content per leaf area unit and the variable fluorescence are also suitable.

Photosynthetic activity, chloroplast ultrastructure, and leaf characteristics of high-light and low-light plants and of sun and shade leaves.

Leaves and herbaceous leaf canopies photosynthesize efficiently although the distribution of light, the ultimate resource of photosynthesis, is very biased in these systems. As has been suggested in theoretical studies, if a photosynthetic system is organized such that every photosynthetic apparatus photosynthesizes in concert, the system as a whole has the sharpest light response curve and is most adaptive. This condition can be approached by (i) homogenization of the light environment and (ii) acclimation of the photosynthetic properties of leaves or chloroplasts to their local light environments. This review examines these two factors in the herbaceous leaf canopy and in the leaf. Changes in the inclination of leaves in the canopy and differentiation of mesophyll into palisade and spongy tissue contribute to the moderation of the light gradient. Leaf and chloroplast movements in the upper parts of these systems under high irradiances also moderate light gradients. Moreover, acclimation of leaves and chloroplasts to the local light environment is substantial. These factors increase the efficiency of photosynthesis considerably. However, the systems appear to be less efficient than the theoretical optimum. When the systems are optically dense, the light gradients may be too great for leaves or chloroplasts to acclimate. The loss of photosynthetic production attributed to the imperfect adjustment of photosynthetic apparatus to the local light environment is most apparent when the photosynthesis of the system is in the transition between the light-limited and light-saturated phases. Although acclimation of the photosynthetic apparatus and moderation of light gradients are imperfect, these markedly raise the efficiency of photosynthesis. Thus more mechanistic studies on these adaptive attributes are needed. The causes and consequences of imperfect adjustment should also be investigated.

Comparative ecophysiology of leaf and canopy photosynthesis

Physiological processes in plant ecology

Fragaria vesca, the woodland strawberry, was grown under a series of controlled environments including variations in light intensity, average temperatures, and temperature amplitude around a constant mean. Observations on CO2 exchange capacities, leaf anatomy, and cell ultrastructure were made for each treatment to determine relationships between these variables. With increasing light intensity, leaf thickness, leaf density, and mesophyll cell surface area and volume per leaf surface area increased. Net photosynthesis (NPS) per leaf weight decreased with increasing light pretreatment while NPS per area increased from low to medium intensity, then decreased at the highest intensity. Depression of photosynthesis at the highest light pretreatment may have been due to massive starch accumulation in the chloroplasts associated with the sodium vapor lamps used. Correlation of all anatomical variables was highly significant with dark respiration and NPS per dry weight but insignificant for NPS per leaf area. In the variable temperature treatments, photosynthetic acclimation occurred with a shift in optimum temperature for NPS in the direction of prevailing growth temperature. Absolute rates were highest at moderate pretreatment temperatures and were reduced by extreme growth temperatures. Thick leaves with low density mesophyll became thinner and more dense with increasing growth temperature corresponding to an increase in maximum net photosynthetic rates. Leaves became thicker and more dense at the highest temperatures, but with an increase in cell damage and indications of changes in metabolic pathways. Highest correlations for gas exchange rates were with specific leaf weight (weight per area). Correlation with other anatomical variables were scattered or insignificant. It was concluded that adaptation to a range of environmental conditions cannot be consistently attributed to changes in mesophyll cell volume or surface area.

Effects of light and temperature on leaf anatomy and photosynthesis in Fragaria vesca.

A B S T R A C T Soybean plants grown in controlled environment cabinets under light intensities of 220 w/m2 or 90 w/m2 (400-700 nm) and day to night temperatures of 27.5-22.5 C or 20.0-12.5 C in all combinations, exhibited differences in growth rate, leaf anatomy, chloroplast ultrastructure, and leaf starch, chlorophyll, and chloroplast lipid contents. Leaves grown under the lower light intensity at both temperatures had palisade mesophyll chloroplasts containing well-formed grana. The corresponding leaves developed under the higher light intensity had very rudimentary grana. Chloroplasts from high temperature and high light had grana consisting of two or three appressed thylakoids, while grana from the low temperature were confined to occasional thylakoid overlap. Spongy mesophyll chloroplasts were less sensitive to growth conditions. Transfer experiments showed that the ultrastructure of chloroplasts from mature leaves could be modified by changing the conditions, though the effect was less marked than when the leaf was growing. LITTLE IS known concerning the influence of light intensities and temperatures characteristic of field conditions on the chloroplast ultrastructure of developing and mature leaves of normal plants. Bjorkman and Holmgren (1963) using the light microscope showed that their higher light intensity destroyed the chloroplasts of shade ecotypes of Solidago virgaurea, whereas chloroplasts from exposed ecotypes were unaffected. The effect of light intensity on the ultrastructure of chloroplasts from pigment-deficient mutants was investigated by Walles (1965), Schmid, Price, and Gaffron (1966), Clewell and Schmid (1969); the effect of temperature and light on the greening of detached, etiolated leaves by Klein (1960), Eilam and Klein (1962); and the eff ect of transference of dark-grown seedlings to light by \'IcWilliam and Naylor (1967). This study concerned differences in growth rate, leaf anatomy, chloroplast ultrastructure, and chemical composition in soybean plants grown in cabinets under combinations of temperature and light similar to those occurring naturally. Comparisons were made with plants grown outside and in the glasshouse. The plants used were well beyond the seedling stage.

https://www.jstor.org/stable/pdfplus/2441353.pdf

B. J. Forde

Papers

The effect of light intensity and temperature on plant growth and chloroplast ultrastructure in soybean