Flexible and reversible responses to different irradiance levels during photosynthetic acclimation of Cypripedium guttatum

Cypripedium guttatum can be found both in open and shady habitats. Photosynthetic acclimation of C. guttatum to different light availabilities was detected using measurements of chlorophyll fluorescence, photosynthesis and leaf traits. When growing under low light conditions, C. guttatum exhibited a greater efficiency in photochemical utilization of absorbed light energy, and a lower ability for non-photochemical dissipation of excess light energy, as compared to the plants growing under high light conditions. Under intermediate light conditions, C. guttatum exhibited higher photosynthetic capacity (A(max)) than those under both low light or high light conditions. The differences in A(max) among three light environments was linked to the differences in biochemical efficiency, leaf N content (LNC) and leaf dry mass per unit area (LMA), but not to the differences of chlorophyll content. However, there were no significant differences in the light compensation points (LCP) and light saturation points (LSP) for photosynthesis for the plants growing under the three light conditions. These results indicate that the photosynthetic capacity of C. guttatum leaves allows for flexible and reversible responses to different irradiance levels. Photosynthetic acclimation in C. guttatum was affected by biochemical changes, the changes in LMA and ratio of Chl a/b. Successful acclimation of C. guttatum to a broad range of light levels likely allows for its wide geographical distribution. A level of about 45% sunlight appears to be optimal for photosynthesis.     

Photosynthetic performance along a light gradient as related to leaf characteristics of a naturally occurring <Emphasis Type="Italic">Cypripedium flavum</Emphasis>

Photosynthesis, leaf structure, nitrogen content and nitrogen allocation in photosynthetic functions of Cypripedium flavum were studied in a naturally varying light regime. Light-saturated leaf net photosynthetic rate (A (max)) was strongly correlated with leaf dry mass per area (LMA), mesophyll conductance (g (m)) and area-based leaf nitrogen content (N(area)), with all variables increasing with increasing irradiance. Such coordinate variation of all these parameters illustrates the plastic response of leaf structure to high light (HL). Leaf N(area) was greater under HL than in low light (LL). The fractions of leaf nitrogen partitioning in carboxylation (P (R)) and bioenergetics (P (B)) were positively related to LMA. In contrast, P (R) and P (B) decreased with increasing mass-based leaf nitrogen content (N(mass)). However, no correlation was found between leaf nitrogen investment in light harvesting (P (L)) and either LMA or N(mass). Like maximum rate of carboxylation (V (cmax)) and electron transport (J (max)), the J (max)/V (cmax) ratio, which was strongly correlated to LMA, also increased significantly with irradiance. Under HL, leaf maximum photosynthetic nitrogen efficiency (ANUE) and intrinsic water use efficiency (WUE) were greater than in LL conditions, despite a small difference in WUE. This suggests that a functional balance in the photosynthetic machinery favors leaf photosynthetic plasticity of C. flavum in response to different light conditions. Given an ample soil nitrogen supply, C. flavum may offset its susceptibility to HL by efficient nitrogen use and higher stomatal and mesophyll conductance against photoinhibition so as to keep leaf photosynthesis positive.     

Seasonal changes in photosynthetic characteristics of Pachysandra terminalis (Buxaceae), an evergreen woodland chamaephyte,in the cool temperate regions of Japan

Summary Seasonal changes in photosynthetic capacity, and photosynthetic responses to intercellular CO₂ concentration and irradiance were investigated under laboratory conditions on intact leaves of Pachysandra terminalis. Photosynthetic capacity and stomatal conductance under saturating light intensity and constant water vapor pressure deficit showed almost the same seasonal trend. They increased from early June just after the expansion of leaves, reached the maximum in late-Septemer, and then decreased to winter. In over-wintering leaves they recovered and increased immediately after snow-melting, reached a first maximum in late April, and then decreased to early July in response to the reduction of light intensity on the forest floor. There-after, they increased from mid August, reached a second maximum in late September, and then decreased to winter. The parallel changes of photosynthesis and stomatal conductane indicate a more or less constant intercellular CO₂ concentration throughout the year. The calculated values of relative stomatal limitation of photosynthesis were nearly constant throughout the year, irrespective of leaf age. The results indicate that the seasonal changes in light-saturated photosynthetic capacity are not due to a change of stomatal conductance, but to a change in the photosynthetic capacity of mesophyll. Indeed, carboxylation efficiency assessed by the inital slope of the Ci-photosynthesis curve changed in proportion to seasonal changes of the photosynthetic capacity in both current-year and over-wintered leaves. High photosynthetic capacity in current-year leaves as compared with one-year-old leaves was also due to the high photosynthetic capacity of mesophyll. Nevertheless, stomatal conductance changed in proportion to photosynthetic capacity, indicating that stomatal conductance is regulated by the mesophyll photosynthetic capacity such that the intercellular CO₂ concentrations are maintained constant. The quantum yield also changed seasonally parallel with that in the photosynthetic capacity.Contribution No. 2893 from the Institute of Low Temperature Science     

Seasonal changes in photosynthetic characteristics of Anemone raddeana,a spring-active geophyte,in the temperate region of Japan

Summary Seasonal changes in the photosynthetic characteristics of intact involucral leaves of Anemone raddeana were investigated under laboratory conditions. Net photosynthesis and constant water vapor pressure deficit showed almost the same seasonal trend. They increased rapidly from mid-April immediately after unfolding of the leaves and reached the maximum in late-April, before the maximum expansion of the leaves. They retained the maximum values until early-May and then decreased toward late-May with a progress of leaf senescence. The calculated values of intercellular CO₂ concentration and relative stomatal limitation of photosynthesis showed no significant change throughout the season. The carboxylation efficiency as assessed by the initial slope of Ci-photosynthesis curve and the net photosynthesis under a high Ci regime varied seasonally in parallel with the change of the light-saturated photosynthesis. The results indicate that the seasonal changes in light-saturated net photosynthesis are not due to a change of stomatal conductance, but to a change in the photosynthetic capacity of mesophyll. Nevertheless, leaf conductance changed concomitantly with photosynthetic capacity, indicating that the seasonal change in stomatal conductance is modulated by the mesophyll photosynthetic capacity such that the intercellular CO₂ concentrations is maintained constant. The shape of light-photosynthesis curve was similar to that of sun-leaf type. The quantum yield also changed simultaneously with the photosynthetic capacity throughout the season.Contribution No. 2965 from the Institute of Low Temperature Science     

Micropropagation of Cypripedium flavum through multiple shoots of seedlings derived from mature seeds

Cypripedium flavum, known as the rare lady’s slipper orchid, is one of the endemics with a yellow flower in China. Due to its conservation and commercial requirement, establishment of an efficient method for micropropogation is urgently needed. Multiple shoots were obtained by placing seedlings from seeds of C. flavum on Harvais media supplemented with two cytokinins (BAP or KIN) used alone or in addition to different concentration of potato homogenate. The effect of BAP was better than that of KIN on shoot multiplication. The Havais media supplemented with BAP (2.22 μM) and potato homogenate (20 g l⁻¹) was the most effective, providing high shoot multiplication frequencies (95%) associated with a high number of shoots per explant (2.55 shoots/plant). For root formation, high rooting and survival were achieved using 1/2 Harvais media supplemented with 0.6 g l⁻¹activated charcoals. High-level activated charcoal increased the number and the length of roots because the activated charcoal could absorb BAP in the media. This study demonstrated that C. flavum could be micropropagated by using multiple shoots of seedlings derived from mature seeds.     

Interactions among leaf photosynthetic rates,flowering and pod set in soybeans

The seasonal maximum in photosynthetic CO₂ exchange rate (CER) and the cessation of leaf expansion in soybeans (Glycine max (L.) Merr) accompany fruiting under normal agricultural conditions. To investigate whether these phenomena were obligatively tied together, we caused early flowering of long-season varieties by imposing artificial short-day treatments. Comparisons of CER and leaf area between vegetative (long-day treatment) and fruiting (short-day treatment) plants of long-season cultivar confirmed the relationship of these phenomena. The same comparisons made between a long-season and a short-season cultivar, both at the same daylength, also confirmed the relationship.     

Photochemical efficiency of photosystem II,photon yield of O2 evolution,photosynthetic capacity,and carotenoid composition during the midday depression of net CO2 uptake in Arbutus unedo growing in Portugal

During the midday depression of net CO₂ exchange in the mediterranean sclerophyllous shrub Arbutus unedo, examined in the field in Portugal during August of 1987, several parameters indicative of photosynthetic competence were strongly and reversibly affected. These were the photochemical efficiency of photosystem (PS) II, measured as the ratio of variable to maximum chlorophyll fluorescence, as well as the photon yield and the capacity of photosynthetic O₂ evolution at 10% CO₂, of which the apparent photon yield of O₂ evolution was most depressed. Furthermore, there was a strong and reversible increase in the content of the carotenoid zeaxanthin in the leaves that occurred at the expense of both violaxanthin and -carotene. Diurnal changes in fluorescence characteristics were interpreted to indicate three concurrent effects on the photochemical system. First, an increase in the rate of radiationless energy dissipation in the antenna chlorophyll, reflected by changes in 77K fluorescence of PSII and PSI as well as in chlorophyll a fluorescence at ambient temperature. Second, a state shift characterized by an increase in the proportion of energy distributed to PSI as reflected by changes in PSI fluorescence. Third, an effect lowering the photon yield of O₂ evolution and PSII fluorescence at ambient temperature without affecting PSII fluorescence at 77K which would be expected from a decrease in the activity of the water splitting enzyme system, i.e. a donor side limitation.Abbreviations and symbols c_i concentration of CO₂ within the leaf - F_o instantaneous fluorescence emission - F_M maximum fluorescence emission - F_v variable fluorescence emission - PFD photon flux density (400–700 nm) - PSI, II photosystem I, II - T_L leaf temperature     

Partitioning of photosynthetic electron flow between CO2 and O2 reduction in a C3 leaf (Phaseolus vulgaris L.) at different CO2 concentrations and during drought stress

Photosystem II chlorophyll fluorescence and leaf net gas exchanges (CO₂ and H₂O) were measured simultaneously on bean leaves (Phaseolus vulgaris L.) submitted either to different ambient CO₂ concentrations or to a drought stress. When leaves are under photorespiratory conditions, a simple fluorescence parameter F/ F_m (B. Genty et al. 1989, Biochem. Biophys. Acta 990, 87–92; F = difference between maximum, F_m, and steady-state fluorescence emissions) allows the calculation of the total rate of photosynthetic electron-transport and the rate of electron transport to O₂. These rates are in agreement with the measurements of leaf O₂ absorption using ¹⁸O₂ and the kinetic properties of ribulose-1,5bisphosphate carboxylase/oxygenase. The fluorescence parameter, F/F_m, showed that the allocation of photosynthetic electrons to O₂ was increased during the desiccation of a leaf. Decreasing leaf net CO₂ uptake, either by decreasing the ambient CO₂ concentration or by dehydrating a leaf, had the same effect on the partitioning of photosynthetic electrons between CO₂ and O₂ reduction. It is concluded that the decline of net CO₂ uptake of a leaf under drought stress is only due, at least for a mild reversible stress (causing at most a leaf water deficit of 35%), to stomatal closure which leads to a decrease in leaf internal CO₂ concentration. Since, during the dehydration of a leaf, the calculated internal CO₂ concentration remained constant or even increased we conclude that this calculation is misleading under such conditions.Abbreviations Ca, Ci ambient, leaf internal CO₂ concentrations - F_m, F_o, F_s maximum, minimal, steady-state fluorescence emission - F_v variable fluorescence emission - PPFD photosynthetic photon flux density - q_p, q_N photochemical, non-photochemical fluorescence quenching - Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase     

Light,leaf age,and leaf nitrogen concentration in a tropical vine

Summary Tropical vines in the Araceae family commonly exhibit alternating periods of upward and downward growth, decoupling the usual relationship between decreasing light environment with increasing age among the leaves on a shoot. In this study I examined patterns of light, leaf specific mass, and leaf nitrogen concentration in relation to leaf position, a measure of developmental age, in field collected shoots of Syngonium podophyllum. These data were analyzed to test the hypothesis that nitrogen allocation parallels within-shoot gradients of light availability, regardless of the relationship between light and leaf age. I found that leaf nitrogen concentration, on a mass basis, was weakly correlated with leaf level light environment. However, leaf specific mass, and consequently nitrogen per unit leaf area, were positively correlated with gradients of light within the shoot, and either increased or decreased with leaf age, providing support for the hypothesis that nitrogen allocation parallels gradients of light availability.     

Changes in photosynthetic capacity,carboxylation efficiency,and CO2 compensation point associated with midday stomatal closure and midday depression of net CO2 exchange of leaves of Quercus suber

The carbon-dioxide response of photosynthesis of leaves of Quercus suber, a sclerophyllous species of the European Mediterranean region, was studied as a function of time of day at the end of the summer dry season in the natural habitat. To examine the response experimentally, a standard time course for temperature and humidity, which resembled natural conditions, was imposed on the leaves, and the CO₂ pressure external to the leaves on subsequent days was varied. The particular temperature and humidity conditions chosen were those which elicited a strong stomatal closure at midday and the simultaneous depression of net CO₂ uptake. Midday depression of CO₂ uptake is the result of i) a decrease in CO₂-saturated photosynthetic capacity after light saturation is reached in the early morning, ii) a decrease in the initial slope of the CO₂ response curve (carboxylation efficiency), and iii) a substantial increase in the CO₂ compensation point caused by an increase in leaf temperature and a decrease in humidity. As a consequence of the changes in photosynthesis, the internal leaf CO₂ pressure remained essentially constant despite stomatal closure. The effects on capacity, slope, and compensation point were reversed by lowering the temperature and increasing the humidity in the afternoon. Constant internal CO₂ may aid in minimizing photoinhibition during stomatal closure at midday. The results are discussed in terms of possible temperature, humidity, and hormonal effects on photosynthesis.Abbreviations and symbols CE carboxylation efficiency - NP net photosynthesis rate - PAR photosynthetically active radiation - P_i leaf internal CO₂ partial pressure - W water vapor mole fraction difference between leaf and air - T CO₂ compensation pressure Dedicated to Professor Dr. Hubert Ziegler on the occasion of his 60th birthday     

Effect of CO2 enrichment and nitrogen availability on resource acquisition and resource allocation in a grass,Bromus mollis

Summary The effects of CO₂ enrichment on the growth, biomass partitioning, photosynthetic rates, and leaf nitrogen concentration of a grass, Bromus mollis (C₃), were investigated at a favorable and a low level of nitrogen availability. Despite increases in root: shoot ratios, leaf nitrogen concentrations were decreased under CO₂ enrichment at both nitrogen levels. For the low-nitrogen treatment, this resulted in lower photosynthetic rates measured at 650 l/l for the CO₂-enriched plants, compared to photosynthetic rates measured at 350 l/l for the non-enriched plants. At higher nitrogen availability, photosynthetic rates of plants grown and measured at 650 l/l were greater than photosynthetic rates of the non-enriched plants measured at 350 l/l. Water use efficiency, however, was increased in enriched plants at both nitrogen levels. CO₂ enrichment stimulated vegetative growth at both high and low nitrogen during most of the vegetative growth phase but, at the end of the experiment, total biomass of the high and low CO₂ treatments did not differ for plants grown at low nitrogen availability. While not statistically significant, CO₂ tended to stimulate seed production at high nitrogen and to decrease it at low nitrogen.     

CO2 environment,microclimate and photosynthetic characteristics of the moss Hylocomium splendens in a subarctic habitat

Summary In order to document the natural CO₂ environment of the moss Hylocomium splendens, and ascertain whether or not the moss was adapted to this, and its interactions with other microenvironmental factors, two studies were carried out. Firstly, the seasonal variations of CO₂ concentration, photosynthetically active radiation (PAR), tissue water content and temperature were measured in the natural microenvironment of H. splendens in a subarctic forest during the summer period (July–September). Secondly, the photosynthetic responses of the species to controlled CO₂ concentrations, PAR, temperature, and hydration were measured in the laboratory. CO₂ concentrations around the upper parts of the plant, when PAR was above the compensation point (30 mol m^–2 s^–1), were mostly between 400 and 450 ppm. They occasionally increased up to 1143 ppm for short periods. PAR flux densities below saturating light levels for photosynthesis (100 mol m^–2 s^–1), occurred during 65% (July), 76% (August) and 96% (September) of the hours of the summer period. The temperature optimum of photosynthesis was 20° C: this temperature coincided with PAR above the compensation point during 5%, 6% and 0% of the time in July, August and September, respectively. Optimal hydration of tissues was infrequent. Hence PAR, temperature and water limit CO₂ uptake for most of the growing season. Our data suggest that the higher than normal ambient CO₂ concentration in the immediate environment of the plant counteracts some of the limitations in PAR supply that it experiences in its habitat. This species already experiences concentrations of atmospheric CO₂ predicted to occur over the next 50 years.     

Long-term drought modifies the fundamental relationships between light exposure, leaf nitrogen content and photosynthetic capacity in leaves of the lychee tree (Litchi chinensis)

Drought has dramatic negative effects on plants' growth and crop productivity. Although some of the responses and underlying mechanisms are still poorly understood, there is increasing evidence that drought may have a negative effect on photosynthetic capacity. Biochemical models of leaf photosynthesis coupled with models of radiation transfer have been widely used in ecophysiological studies, and, more recently, in global change modeling. They are based on two fundamental relationships at the scale of the leaf: (i) nitrogen content-light exposure and (ii) photosynthetic capacity-nitrogen content. Although drought is expected to increase in many places across the world, such models are not adapted to drought conditions. More specifically, the effects of drought on the two fundamental relationships are not well documented. The objective of our study was to investigate the effects of a long-term drought imposed slowly on the nitrogen content and photosynthetic capacity of leaves similarly exposed to light, from 3-year-old lychee trees cv. Kwa? Mi. Leaf nitrogen and non-structural carbohydrate concentrations were measured along with gas exchanges and the light-saturated rate of photosynthetic electron transport (J(max)) after a 5.5-month-long period of drought. Leaf nitrogen content on a mass basis remained stable, while the leaf mass-to-area ratio (LMA) increased with increasing water stress. Consequently, the leaf nitrogen content on an area basis (N(a)) increased in a non-linear fashion. The starch content decreased, while the soluble sugar content increased. Stomata closed and net assimilation decreased to zero, while J(max) and the ratio J(max)/N(a) decreased with increasing water stress. The drought-associated decrease in photosynthetic capacity can be attributed to downregulation of photosynthetic electron transport and to reallocation of leaf nitrogen content. It is concluded that modeling photosynthesis in drought conditions will require, first, the modeling of the effect of drought on LMA and J(max).     

Decline of photosynthetic capacity with leaf age in relation to leaf longevities for five tropical canopy tree species

The effect of leaf aging on photosynthetic capacities was examined for upper canopy leaves of five tropical tree species in a seasonally dry forest in Panama. These species varied in mean leaf longevity between 174 and 315 d, and in maximum leaf life span between 304 and 679 d. The light-saturated CO2 exchange rates of leaves produced during the primary annual leaf flush measured at 7-8 mo of age were 33-65% of the rates measured at 1-2 mo of age for species with leaf life span of < 1 yr. The negative regression slopes of photosynthetic capacity against leaf age were steeper for species with shorter maximum leaf longevity. In all species, regression slopes were less steep than the slopes predicted by assuming a linear decline toward the maximum leaf age (20-80% of the predicted decline rate). Maximum oxygen evolution rates and leaf nitrogen content declined faster with age for species with shorter leaf life spans. Statistical significance of regression slopes of oxygen evolution rates against leaf age was strongest on a leaf mass basis (r = 0.49-0.87), followed by leaf nitrogen basis (r = 0.48-0.77), and weakest on a leaf area basis (r = 0.35-0.70).     

Secondary fluorescence kinetics of spinach leaves in relation to the onset of photosynthetic carbon assimilation

When spinach leaves are re-illuminated, after dark periods of 90 s or less, an initial fluorescence peak is observed which rapidly gives way to a much lower terminal value. After 2 min or more in the dark, however, there is a secondary rise, at about 50–70 s, which then gives way, more slowly, to approximately the same low terminal value as before. The secondary rise is eliminated or disguised by feeding D,L-glyceraldehyde (a specific inhibitor of photosynthetic carbon assimilation) and by manose, 2-deoxyglucose and glucosamine, all of which are believed to sequester cytoplasmic orthophosphate. This secondary rise in fluorescence is discussed in relation to photosynthetic induction and the manner in which these compounds may modulate fluorescence by their effect on the availability of orthophosphate and their consequent impact on the adenylate status of the stroma.Abbreviations DCMU 3(3,4-dichlorophenyl)-1,1-dimethylurea - CCCP carbonylcyanidchlorophenylhydrazon     

Light- and seasonal-induced plasticity in leaf morphology, N partitioning and photosynthetic capacity of two temperate deciduous species

Processes involved in leaf photosynthetic acclimation to light and throughout the growing season were investigated in two hardwood species (Acer saccharum and Betula alleghaniensis), which differed in their level of shade-tolerance. For both species, variation in traits related to (i) leaf morphology (LMA, leaf mass:area ratio), (ii) leaf N content (N_A, leaf nitrogen content on an area basis and N_M, N concentration in leaf dry mass), (iii) leaf N partitioning among photosynthetic functions (P_r, N allocated to Rubisco, and P_b, N allocated to bioenergetics), and (iv) leaf photosynthetic capacity (V_cmax, maximal carboxylation rates, and J_max, maximal light-driven electron flow) were assessed at three different times during the growing season (early, mid- and late summer) and under four contrasting light regimes (40, 17, 6 and 2% of full sunlight). For both species, light-driven variation in most traits was greater than their seasonally driven variation. Furthermore, results showed for both species the pre-eminence of LMA changes in the light-driven acclimation of N_A. Importance of N_M to variation in N_A was restricted to seasonal acclimation, especially for the less shade-tolerant species, B. alleghaniensis. Similarly, for both species, light-driven acclimation of leaf photosynthetic capacities was tightly related to variation in N_A, which was related to LMA changes. However, variation in P_r and P_b better explained seasonally driven variation in V_cmax and J_max, specifically under lower light levels, where N_A was low. Thus, the great variability observed for leaf activity in response to contrasting light environments was related to efficient morphological adjustments, regardless of species level of shade-tolerance. Finally, physiological adjustments were mainly involved in fine-scale changes observed during seasonally driven acclimation of leaves, when LMA was constrained to a slight range of variation.     

Long term effects of naturally elevated CO2 on mediterranean grassland and forest trees

We investigated the carbon supply status in species-rich mediterranean plant communities growing in a bowl-shaped 1-ha CO₂ spring area near Sienna, Italy. A geothermic lime-kiln has provided these communities, for as long as historical records are available, with pure CO₂ that mixes with ambient air at canopy level to daytime means of 500–1000 ppm CO₂. Immediately outside the spring area similar plant communities are growing on similar substrate, and in the same climate, but under ca. 355 ppm CO₂. We found no evidence that plants in the CO₂ spring area grow faster, flower earlier or become larger. However, we found very large differences in tissue quality among the 40 species studied inside and outside the spring area. Depending on weather conditions, the mean concentration of total non-structural carbohydrates (TNC, sugars and starch) in leaves of herbaceous plants was 38–47% higher in the spring area. Fast growing ruderals growing on garden soil inside and outside the spring area show the same response. Among trees, leaves of the deciduousQuercus pubscens contain twice as much TNC inside as outside the vent area, whereas evergreenQ. ilex leaves show no significant difference. TNC levels in branch wood paralleled leaf values. TNC in shade leaves was also higher. Elevated CO₂ had no effect on the sugar fraction, therefore differences in TNC are due to starch accumulation. Leaf nitrogen concentration decreases under elevated CO₂. These observations suggest that the commonly reported TNC accumulation and N depletion in leaves growing under elevated CO₂ are not restricted to the artificial conditions of short-term CO₂ enrichment experiments but persist over very long periods. Such an alteration of tissue composition can be expected to occur in other plant communities also if atmospheric CO₂ levels continue to rise. Effects on food webs and nutrient cycling are likely.     

Gas exchange and photosynthetic performance of the tropical tree <Emphasis Type="Italic">Acacia nigrescens</Emphasis> when grown in different CO<Subscript>2</Subscript> concentrations

The photosynthetic responses of the tropical tree species Acacia nigrescens Oliv. grown at different atmospheric CO₂ concentrations—from sub-ambient to super-ambient—have been studied. Light-saturated rates of net photosynthesis (A _sat) in A. nigrescens, measured after 120 days exposure, increased significantly from sub-ambient (196 μL L⁻¹) to current ambient (386 μL L⁻¹) CO₂ growth conditions but did not increase any further as [CO₂] became super-ambient (597 μL L⁻¹). Examination of photosynthetic CO₂ response curves, leaf nitrogen content, and leaf thickness showed that this acclimation was most likely caused by reduction in Rubisco activity and a shift towards ribulose-1,5-bisphosphate regeneration-limited photosynthesis, but not a consequence of changes in mesophyll conductance. Also, measurements of the maximum efficiency of PSII and the carotenoid to chlorophyll ratio of leaves indicated that it was unlikely that the pattern of A _sat seen was a consequence of growth [CO₂] induced stress. Many of the photosynthetic responses examined were not linear with respect to the concentration of CO₂ but could be explained by current models of photosynthesis.     

The midday depression of CO2 assimilation in leaves of Arbutus unedo L.: diurnal changes in photosynthetic capacity related to changes in temperature and humidity

Parts of attached leaves of the sclerophyllous shrub Arbutus unedo were subjected to simulated mediterranean days. Gas exchange was recorded in order to recognize the causes of the midday depression in CO₂ assimilation. Depressions could be induced in part of a leaf: they were local responses. The CO₂-saturation curves of photosynthesis, determined during the morning and afternoon maxima of CO₂ assimilation and during the minimum at midday, established that depressions in CO₂ assimilation were in one-half of the investigated cases totally caused by reversible reductions in the photosynthetic capacity of the leaves, and in the other half almost totally caused by such reductions. An analysis of 37 daily courses showed that morning reductions and afternoon recoveries of stomatal conductance and rate of photosynthesis occurred simultaneously and in proportion to each other, with the result that the partial pressure of CO₂ in the intercellular spaces remained more or less constant. Midday depressions occurred also in detached leaves standing in water. The initiation of a midday depression was not caused by a circadian rhythm, nor was high quantum flux or high temperature a requirement. There was no correlation between the rate of water loss from the leaves, or the amount of water lost, with the degree of reduction of the photosynthetic capacity. However, depressions occurred if an apparent threshold in the water-vapor pressure difference between leaf and air was exceeded. This critical value varied between about 20 and 30 mbar, depending on the leaf investigated. The dominating role of humidity in the induction of the midday depression was further demonstrated when leaf temperature was held constant and the vapor-pressure difference was made to follow the pattern of the mediterranean day: depressions occurred. Depressions however were hardly noticeable when the water-vapor pressure difference was held constant and leaf temperature was allowed to vary. In another set of experiments, leaves were subjected to variations in temperature and humidity independent of the time of the day, under otherwise constant conditions. Photosynthetic capacity and stomatal conductance proved to be almost insensitive to changes in temperature (in a range extending from 20 to 37° C) as long as the water vapor-pressure difference was held constant. If it was not, the rate of photosynthesis began to decline with increasing temperature after a threshold water-vapor pressure difference was exceeded. The position of the resulting apparent temperature optimum of photosynthesis depended on the humidity of the air. We suggest that the ability of A. unedo to respond to a dry atmosphere with a reversible reduction of its photosynthetic capacity (by a still unknown mechanism) is the result of a co-evolution with the development of a strong stomatal sensitivity to changes in humidity.     

Photosynthetic carbon metabolism during leaf ontogeny in Zea mays L.: Enzyme studies

The activities of several enzymes, including ribulose-1,5-diphosphate (RuDP) carboxylase (EC 4.1.1.39) and phosphoenolpyruvate (PEP) carboxylase (EC 4.1.1.31) were measured as a function of leaf age in Z. mays. Mature leaf tissue had a RuDP-carboxylase activity of 296.7 mol CO₂ g^-1 fresh weight h^-1 and a PEP-carboxylase activity of 660.6 mol CO₂ g^-1 fresh weight h^-1. In young corn leaves the activity of the two enzymes was 11 and 29%, respectively, of the mature leaves. In senescent leaf tissue, RuDP carboxylase activity declined more rapidly than that of any of the other enzymes assayed. On a relative basis the activities of NADP malic enzyme (EC 1.1.1.40), aspartate (EC 2.6.1.1) and alanine aminotransferase (EC 2.6.1.2), and NAD malate dehydrogenase (EC 1.1.1.37) exceeded those of both PEP and RuDP carboxylase in young and senescent leaf tissue. Pulse-chase labeling experiments with mature and senescent leaf tissue show that the predominant C₄ acid differs between the two leaf ages. Labeling of alanine in senescent tissue never exceeded 4% of the total ¹⁴C remaining during the chase period, while in mature leaf tissue alanine accounted for 20% of the total after 60 s in ¹²CO₂. The activity of RuDP carboxylase during leaf ontogeny in Z. mays parallels the development of the activity of this enzyme in C₃ plants.Abbreviations RuDP ribulose-1,5-diphosphate - PEP phosphoenol pyruvate - PGA 3-phosphoglycerate