Gas Exchange Analysis of the Fast Phase of Photosynthetic Induction in Alocasia macrorrhiza

When leaves of Alocasia macrorrhiza that had been preconditioned in 10 micromoles photons per square meter per second for at least 2 hours were suddenly exposed to 500 micromoles photons per square meter per second, there was an almost instantaneous increase in assimilation rate. After this initial increase, there was a secondary increase over the next minute. This secondary increase was more pronounced in high CO₂ (1400 microbars), where assimilation rate was assumed to be limited by the rate of regeneration of ribulose 1,5-bisphosphate (RuBP). It was absent in low CO₂ (75 microbars), where RuBP carboxylase/oxygenase (Rubisco) was assumed to be limiting. It was therefore concluded that it represented an increase in the capacity to regenerate RuBP. This fast-inducing component not only gained full induction rapidly, but also lost it rapidly in low photon flux density (PFD) with a half time of 150 to 200 seconds. It was concluded that in environments with fluctuating PFD, this fast-inducing component is an important factor in determining a leaf's potential for photosynthetic carbon gain. It is especially important during brief periods (<30 seconds) of high PFD that follow moderately long periods (1 to 10 minutes) of low PFD.     

Limitation of CO(2) Assimilation and Regulation of Benson-Calvin Cycle Activity in Barley Leaves in Response to Changes in Irradiance, Photoinhibition, and Recovery

The response of the Benson-Calvin cycle to changes in irradiance and photoinhibition was measured in low-light grown barley (Hordeum vulgare) leaves. Upon the transition from the growth irradiance (280 micromoles per square meter per second) to a high photoinhibitory irradiance (1400 micromoles per square meter per second), the CO₂ assimilation rate of the leaves doubled within minutes but high irradiance rapidly caused a reduction in quantum efficiency. Following exposure to high light the activities of NADP-malate dehydrogenase and fructose-1,6-bisphosphatase obtained near maximum values and the activation state of ribulose-1,5-bisphosphate carboxylase increased. The activity of the latter remained constant throughout the period of photoinhibitory irradiance, but the increase in the activities of fructose-1,6-bisphosphatase and NADP-malate dehydrogenase was transient decreasing once more to much lower values. This suggests that immediately following the transition to high light reduction and activation of redox-modulated enzymes occurred, but then the stroma became relatively oxidized as a result of photoinhibition. The leaf contents of glucose 6-phosphate and fructose 6-phosphate increased following exposure to high light but subsequently decreased, suggesting that following photoinhibition sucrose synthesis exceeded the rate of carbon assimilation. The ATP content attained a constant value much higher than that in low light. During photoinhibition the glycerate 3-phosphate content greatly increased while ribulose-1,5-bisphosphate decreased. The fructose-1,6-bisphosphate and triose phosphate contents increased initially and then remained constant. During photoinhibition CO₂ assimilation was not limited by ribulose-1,5-bisphosphate carboxylase activity but rather by the regeneration of the substrate, ribulose-1,5-bisphosphate, related to a restriction on the supply of reducing equivalents.     

Biphasic Activation of Ribulose Bisphosphate Carboxylase in Spinach Leaves as Determined from Nonsteady-State CO(2) Exchange

The activation kinetics of ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) following an increase in photon flux density (PFD) were studied by analyzing CO₂ assimilation time courses in spinach leaves (Spinacia oleracea). When leaves were exposed to 45 minutes of darkness before illumination at 690 micromoles per square meter per second, Rubisco activation followed apparent first-order kinetics with a relaxation time of about 3.8 minutes. But when leaves were illuminated for 45 minutes at 160 micromoles per square meter per second prior to illumination at 690 micromoles per square meter per second the relaxation time for Rubisco activation was only 2.1 minutes. The kinetics of this change in relaxation times were investigated by exposing dark-adapted leaves to 160 micromoles per square meter per second for different periods before increasing the PFD to 690 micromoles per square meter per second. It was found that the apparent relaxation time for Rubisco activation changed from 3.8 to 2.1 minutes slowly, requiring at least 8 minutes for completion. This result indicates that at least two sequential, slow processes are involved in light-mediated activation of Rubisco in spinach leaves and that the relaxation times characterizing these two processes are about 4 and 2 minutes, respectively. The kinetics of the first process in the reverse direction and the dependence of the relaxation time for the second process on the magnitude of the increase in PFD were also determined. Evidence that the first slow process is activation of the enzyme Rubisco activase and that the second slow process is the catalytic activation of Rubisco by activase is discussed.     

Light-dependent changes in ribulose bisphosphate carboxylase activase activity in leaves

An assay for the activity of ribulose bisphosphate carboxylase (Rubisco) activase in crude leaf extracts was developed. The assay is based on a spectrophotometric assay of Rubisco, and activase activity (in nanomoles activated Rubisco per minute per milligram chlorophyll) was calculated from the rate of increase in Rubisco activity over time. Activase activity measurements were made using samples from spinach (Spinacia oleracea) leaves undergoing (a) steady-state photosynthesis at various photon flux density (PFD) values and (b) nonsteady-state photosynthesis following an increase from darkness to a high PFD. Analysis of these samples showed that steady-state Rubisco activase activity was relatively low in darkness, increased with PFD, and saturated below 300 micromoles per square meter per second. Rubisco activity (measured spectrophotometrically) was also found to be low in darkness and to increase with PFD, but it saturated at much higher PFD values (approximately 1000 micromoles per square meter per second) along with the rate of photosynthesis. Following an increase in PFD from darkness to 650 micromoles per square meter per second, activase activity increased more or less linearly over a period of 5 to 6 minutes, after which it was constant. Rubisco activity, however, increased more slowly. The light-dependence of Rubisco activase is consistent with previous gas-exchange data showing two interdependent processes in the activation of Rubisco following an increase in PFD.     

Regulation of Photosynthetic Rate of Two Sunflower Hybrids under Water Stress

The effect of short-term water stress on photosynthesis of two sunflower hybrids (Helianthus annuus L. cv Sungro-380 and cv SH-3622), differing in productivity under field conditions, was measured. The rate of CO₂ assimilation of young, mature leaves of SH-3622 under well-watered conditions was approximately 30% greater than that of Sungro-380 in bright light and elevated CO₂; the carboxylation efficiency was also larger. Growth at large photon flux increased assimilation rates of both hybrids. The changes in leaf composition, including cell numbers and sizes, chlorophyll content, and amounts of total soluble and ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) protein, and in Rubisco activity and amount of ribulose-1,5-bisphosphate (RuBP) were determined to assess the factors regulating the differences in assimilation of the hybrids at high and low water potentials. The amounts of chlorophyll, soluble protein, Rubisco protein and the initial activity of Rubisco and its activation state did not differ significantly between hybrids. However, unstressed leaves of SH-3622 had more, smaller cells per unit area and 60% more RuBP per unit leaf area than that of Sungro-380. Water stress developing over 4 days decreased the assimilation of both hybrids similarly. Changes in the amounts of chlorophyll, soluble and Rubisco protein, and Rubisco activity and activation state were small and were not sufficient to explain the decrease in photosynthesis; neither was decreased stomatal conductance (or stomatal “patchiness”). Reduction of photosynthesis per unit leaf area from 25 to 5 micromoles CO₂ per square meter per second in both hybrids was caused by a decrease in the amount of RuBP from approximately 130 to 40 micromoles per square meter in SH-3622 and from 80 to 40 micromoles per square meter in Sungro. Differences between hybrids and their response to water stress is discussed in relation to control of RuBP regeneration.     

Starch and Sucrose Synthesis in Phaseolus vulgaris as Affected by Light, CO(2), and Abscisic Acid

Phaseolus vulgaris L. leaves were subjected to various light, CO₂, and O₂ levels and abscisic acid, then given a 10 minute pulse of ¹⁴CO₂ followed by a 5 minute chase with unlabeled CO₂. After the chase period, very little label remained in the ionic fractions (presumed to be mostly carbon reduction and carbon oxidation cycle intermediates and amino acids) except at low CO₂ partial pressure. Most label was found in the neutral, alcohol soluble fraction (presumed sucrose) or in the insoluble fraction digestable by amyloglucosidase. Sucrose formation was linearly related to assimilation rate (slope = 0.35). Starch formation increased linearly with assimilation rate (slope = 0.56) but did not occur if the assimilation rate was below 4 micromoles per square meter per second. Neither abscisic acid, nor high CO₂ in combination with low O₂ (thought to disrupt control of carbon metabolism) caused significant perturbations of the sucrose/starch formation ratio. These studies indicate that the pathways for starch and sucrose synthesis both are controlled by the rate of net CO₂ assimilation, with sucrose the preferred product at very low assimilation rates.     

Light-dependent damage to photosynthesis in olive leaves during chilling and high temperature stress

Abstract The leaves of olive are long lived and likely to experience both chilling and high temperature stress during their life. Changes in photosynthetic CO₂ assimilation resulting from chilling and high temperature stress, in both dim and high light, are investigated. The quantum yield (φ) of photosynthesis at limiting light levels was reduced following chilling (at 5°C for 12 h), in dim light by approximately 10%, and in high light by 75%; the difference being attributed to photoinhibition. Similar reductions were observed in the light-saturated rate of CO₂ uptake (A_max). Decrease in A_max correlated with a halving of the leaf internal CO₂ concentration (c_i), suggesting an increased limitation by stomata following photoinhibition. Leaves were apparently more susceptible to photoinhibitory damage if the whole plant, rather than the leaf alone, was chilled. On return to 26 °C, I he photosynthetic capacity recovered to pre-stress levels within a few hours if leaves had been chilled in high light for 8 h or less, but did not fully recover from longer periods of chilling when loss of chlorophyll occurred. Leaves which were recovering from chilling in high light showed far more damage on being chilled a second time in high light. Three hours in high light at 38 °C reduced φ by 80%, but φ recovered within 4h of return to 26 °C. Although leaves of Olive are apparently less susceptible to photoinhibitory damage during chilling stress than the short-lived leaves of chilling-sensitive annual? crops, the results nevertheless show that photoinhibition during temperature stress is potentially a major factor influencing the photosynthetic productivity of Olive in the field.     

Whole Leaf Carbon Exchange Characteristics of Phosphate Deficient Soybeans (Glycine max L.)

Low phosphate nutrition results in increased chlorophyll fluorescence, reduced photosynthetic rate, accumulation of starch and sucrose in leaves, and low crop yields. This study investigated physiological responses of soybean (Glycine max [L.] Merr.) leaves to low inorganic phosphate (Pi) conditions. Responses of photosynthesis to light and CO₂ were examined for leaves of soybean grown at high (0.50 millimolar) or low (0.05 millimolar) Pi. Leaves of low Pi plants exhibited paraheliotropic orientation on bright sunny days rather than the normal diaheliotropic orientation exhibited by leaves of high Pi soybeans. Leaves of plants grown at high Pi had significantly higher light saturation points (1000 versus 630 micromole photons [400-700 nanometers] per square meter per second) and higher apparent quantum efficiency (0.062 versus 0.044 mole CO₂ per mole photons) at ambient (34 pascals) CO₂ than did low Pi leaves, yet stomatal conductances were similar. High Pi leaves also had significantly higher carboxylation efficiency (2.90 versus 0.49 micromole CO₂ per square meter per second per pascal), a lower CO₂ compensation point (6.9 versus 11.9 pascals), and a higher photosynthetic rate at 34 pascals CO₂ (19.5 versus 6.7 micromoles CO₂ per square meter per second) than did low Pi leaves. Soluble protein (0.94 versus 0.73 milligram per square centimeter), ribulose-1,5-bisphosphate carboxylase/oxygenase content (0.33 versus 0.25 milligram per square centimeter), and ribulose-1,5-bisphosphate carboxylase/oxygenase specific activity (25.0 versus 16.7 micromoles per square meter per second) were significantly greater in leaves of plants in the high Pi treatment. The data indicate that Pi stress alters the plant's CO₂ reduction characteristics, which may in turn affect the plant's capacity to accommodate normal radiation loads.     

The Susceptibility of Photosynthesis to Photoinhibition and the Capacity of Recovery in High and Low Light Grown Cyanobacteria, Anacystis nidulans

The susceptibility of photosynthesis to photoinhibition and the rate of its recovery were studied in the cyanobacterium Anacystis nidulans grown at a low (10 micromoles per square meter per second) and a high (120 micromoles per square meter per second) photosynthetically active radiation. The rate of light limited photosynthetic O₂ evolution was measured to determine levels of photoinhibition and rates of recovery. Studies of photoinhibition and recovery with and without the translation inhibitor streptomycin demonstrated the importance of a recovery process for the susceptibility of photosynthesis to photoinhibition. We concluded that the approximately 3 times lower susceptibility to photoinhibition of high light than of low light grown cells, significantly depended on high light grown cells having an approximately 3 times higher recovery capacity than low light grown cells. It is suggested that these differences in susceptibility to photoinhibition and recovery depends on high light grown cells having a higher turnover rate of photosystem II protein(s) that is(are) the primary site(s) of photodamage, than have low light grown cells. Furthermore, we demonstrated that photoinhibition of A. nidulans may occur under physiological light conditions without visible harm to the growth of the cell culture. The results give support for the hypotheses that the net photoinhibitory damage of photosystem II results from the balance between the photoinhibitory process and the operation of a recovery process; the capacity of the latter determining significant differences in the susceptibility of photosynthesis to photoinhibition of high and low light grown A. nidulans.     

Regulation of Ribulose-1,5-Bisphosphate Carboxylase Activity in Alocasia macrorrhiza in Response to Step Changes in Irradiance

The regulation of ribulose-1,5-bisphosphate (RuBP) carboxylase (Rubisco) activity and pool sizes of RuBP and P-glycerate were examined in the tropical understory species Alocasia macrorrhiza following step changes in photon flux density (PFD). Previous gas exchange analysis of this species following a step increase in PFD from 10 to 500 micromoles quanta per square meter per second suggested that the increase in photosynthetic rate was limited by the rate of increase of Rubisco activity for the first 5 to 10 minutes. We demonstrate here that the increase in photosynthetic rate was correlated with an increase in both the activation state of Rubisco and the total k_cat (fully activated specific activity) of the enzyme. Evidence presented here suggests that a change in the pool size of the naturally occurring tight binding inhibitor of Rubisco activity, 2-carboxyarabinitol 1-phosphate, was responsible for the PFD-dependent change in the total k_cat of the enzyme. RuBP pool size transiently increased after the increase in PFD, indicating that photosynthesis was limited by the capacity for carboxylation. After 5 to 10 minutes, RuBP pool size was again similar to the pool size at low PFD, presumably because of the increased activity of Rubisco. Following a step decrease in PFD from 500 to 10 micromoles quanta per square meter per second, Rubisco activity declined but at a much slower rate than it had increased in response to a step increase in PFD. This slower rate of activity decline than increase was apparently due to the slower rate of 2-carboxyarabinitol 1-phosphate synthesis than degradation and, to a lesser degree, to slower deactivation than activation. RuBP pool size initially declined following the decrease in PFD, indicating that RuBP regeneration was limiting photosynthesis. As Rubisco activity decreased, RuBP slowly increased to its original level at high PFD. The slow rate of activity loss by Rubisco in this species suggests a biochemical basis for the increased efficiency for CO₂ assimilation of successive lightfleck use by species such as A. macrorrhiza.     

Photosynthetic and Respiratory Activity of Fruiting Forms within the Cotton Canopy

The supply of photosynthates by leaves for reproductive development in cotton (Gossypium hirsutum L.) has been extensively studied. However, the contribution of assimilates derived from the fruiting forms themselves is inconclusive. Field experiments were conducted to document the photosynthetic and respiratory activity of cotton leaves, bracts, and capsule walls from anthesis to fruit maturity. Bracts achieved peak photosynthetic rates of 2.1 micromoles per square meter per second compared with 16.5 micromoles per square meter per second for the subtending leaf. However, unlike the subtending leaf, the bracts did not show a dramatic decline in photosynthesis with increased age, nor was their photosynthesis as sensitive as leaves to low light and water-deficit stress. The capsule wall was only a minor site of ¹⁴CO₂ fixation from the ambient atmosphere. Dark respiration by the developing fruit averaged −18.7 micromoles per square meter per second for 6 days after anthesis and declined to −2.7 micromoles per square meter per second after 40 days. Respiratory loss of CO₂ was maximal at −158 micromoles CO₂ per fruit per hour at 20 days anthesis. Diurnal patterns of dark respiration for the fruit were age dependent and closely correlated with stomatal conductance of the capsule wall. Stomata on the capsule wall of young fruit were functional, but lost this capacity with increasing age. Labeled ¹⁴CO₂ injected into the fruit interior was rapidly assimilated by the capsule wall in the light but not in the dark, while fiber and seed together fixed significant amounts of ¹⁴CO₂ in both the light and dark. These data suggest that cotton fruiting forms, although sites of significant respiratory CO₂ loss, do serve a vital role in the recycling of internal CO₂ and therein, function as important sources of assimilate for reproductive development.     

Sensitivity of Altitudinal Ecotypes of the Wild Tomato Lycopersicon hirsutum to Chilling Injury

The transition temperature of the leaf polar lipids and the critical temperature for chill-induced inhibition of photosynthesis was determined for three altitudinal ecotypes of the wild tomato Lycopersicon hirsutum. Photosynthesis was measured as CO₂-dependent O₂ evolution at 25°C after leaf slices were exposed to chilling temperatures for 2 hours at a moderate photon flux density of 450 micromoles per square meter per second. The transition temperature of the leaf polar lipids was detected from the change in the temperature coefficient of the fluorescence intensity of trans-parinaric acid. Chill-induced photoinhibition was evident in the three tomato ecotypes when they were chilled below a critical temperature of 10°, 11°, and 13°C, respectively, for the high (LA1777), mid (LA1625), and low (LA1361) altitudinal ecotypes. The temperature differential, below the critical temperature, required to produce a 50% inhibition was also similar for the three ecotypes. A transition was detected in the leaf polar lipids of these plants at a temperature similar to that of the critical temperature for photoinhibition. The results show that the three tomato ecotypes are similar with respect to their critical temperature for chilling-induced photoinhibition and the rate of their response to the chilling stress. They are, thus, similarly sensitive to chilling.     

Photosynthetic dynamics in chrysanthemum in response to single step increases and decreases in photon flux density

The time-course of CO₂ assimilation rate and stomatal conductance to step changes in photosynthetic photon flux density (PPFD) was observed in Chrysanthemum × morifolium Ramat. `Fiesta'. When PPFD was increased from 200 to 600 micromoles per square meter per second, the rate of photosynthetic CO<sub>2</sub> assimilation showed an initial rapid increase over the first minute followed by a slower increase over the next 12 to 38 minutes, with a faster response in low-light-grown plants. Leaves exposed to small step increases (100 micromoles per square meter per second) reached the new steady-state assimilation rate within a minute. Both stomatal and biochemical limitations played a role during photosynthetic induction, but carboxylation limitations seemed to predominate during the first 5 to 10 minutes. Stomatal control during the slow phase of induction was less important in low-light compared to high-light-grown plants. In response to step decreases in PPFD, photosynthetic rate decreased rapidly and a depression in CO<sub>2</sub> assimilation prior to steady-state was observed. This CO<sub>2</sub> assimilation `dip' was considerably larger for the large step (400 micromoles per square meter per second) than for the small step. The rapid photosynthetic response seems to be controlled by biochemical processes. High- and low-light-grown plants did not differ in their photosynthetic response to PPFD step decreases.     

Inhibition of photosynthesis by carbohydrates in wheat leaves

The rate of net CO₂ assimilation of mature wheat (Triticum aestivum L.) leaves in ambient air (21% O₂, 340 microbars CO₂) declined with time of illumination at temperatures lower than 25°C, but not at higher temperatures, and the rate of decline increased when maintained in air with higher CO₂ concentration (700-825 microbars). In this latter case, the decline in the rate of net CO₂ assimilation also occurred at high temperatures. Stomatal conductance also declined with time in some cases and stomata became more sensitive to CO₂, but this was not the primary cause of the decrease in CO₂ assimilation because internal partial pressure of CO₂ remained constant. Treatments which reduced the rate of translocation (e.g. lower temperatures, chilling the base of the leaf) produced a marked decline in CO₂ assimilation of leaves in atmospheric and high CO₂ concentrations. The decreased net CO₂ assimilation was correlated with carbohydrate accumulation in each case, suggesting end product inhibition of photosynthesis. Analysis of CO₂ assimilation in high carbohydrate leaves as a function of intercellular CO₂ partial pressure showed reduction in the upper part of the curve. The initial slope of this curve, however, was not affected. Photosynthetic rates in the upper part of this curve generally recovered after a short period in darkness in which carbohydrates were removed from the leaf. The stimulation of net CO₂ assimilation by 2% O₂ (Warburg effect), and the apparent quantum yield, decreased after several hours of light.     

Photoinhibition of photosynthesis in intact kiwifruit (Actinidia deliciosa) leaves: effect of growth temperature on photoinhibition and recovery

Intact leaves of kiwifruit (Actinidia deliciosa (A. Chev.) C.F. Liang et A.R. Ferguson) from plants grown in a range of controlled temperatures from 15/10 to 30/25°C were exposed to a photon flux density (PFD) of 1500 μmol·m⁻²·s⁻¹ at leaf temperatures between 10 and 25°C. Photoinhibition and recovery were followed at the same temperatures and at a PFD of 20 μmol·m⁻²·s⁻¹, by measuring chlorophyll fluorescence at 77 K and 692 nm, by measuring the photon yield of photosynthetic O₂ evolution and light-saturated net photosynthetic CO₂ uptake. The growth of plants at low temperatures resulted in chronic photoinhibition as evident from reduced fluorescence and photon yields. However, low-temperature-grown plants apparently had a higher capacity to dissipate excess excitation energy than leaves from plants grown at high temperatures. Induced photoinhibition, from exposure to a PFD above that during growth, was less severe in low-temperature-grown plants, particularly at high exposure temperatures. Net changes in the instantaneous fluorescence,F ₀, indicated that little or no photoinhibition occurred when low-temperature-grown plants were exposed to high-light at high temperatures. In contrast, high-temperature-grown plants were highly susceptible to photoinhibitory damage at all exposure temperatures. These data indicate acclimation in photosynthesis and changes in the capacity to dissipate excess excitation energy occurred in kiwifruit leaves with changes in growth temperature. Both processes contributed to changes in susceptibility to photoinhibition at the different growth temperatures. However, growth temperature also affected the capacity for recovery, with leaves from plants grown at low temperatures having moderate rates of recovery at low temperatures compared with leaves from plants grown at high temperatures which had negligible recovery. This also contributed to the reduced susceptibility to photoinhibition in low-temperature-grown plants. However, extreme photoinhibition resulted in severe reductions in the efficiency and capacity for photosynthesis.     

Photoinhibition at low temperature in chilling-sensitive and -resistant plants

Photoinhibition resulting from exposure at 7°C to a moderate photon flux density (300 micromoles per square meter per second, 400-700 nanometers) for 20 hours was measured in leaves of annual crops differing widely in chilling tolerance. The incidence of photoinhibition, determined as the decrease in the ratio of induced to total chlorophyll fluorescence emission at 693 nanometers (F_v/F_max) measured at 77 Kelvin, was not confined to chilling-sensitive species. The extent of photoinhibition in leaves of all chilling-resistant plants tested (barley [Hordeum vulgare L.], broad bean [Vicia faba L.], pea [Pisum sativum L.], and wheat [Triticum aestivum L.]) was about half of that measured in chilling-sensitive plants (bean [Phaseolus vulgaris L.], cucumber [Cucumis sativus L.], lablab [Lablab purpureus L.], maize [Zea mays L.], pearl millet [Pennisetum typhoides (Burm. f.) Stapf & Hubbard], pigeon pea [Cajanus cajun (L.) Millsp.], sesame [Sesamum indicum L.], sorghum [Sorghum bicolor L. Moench], and tomato [Lycopersicon esculentum Mill.]). Rice (Oryza sativa L.) leaves of the indica type were more susceptible to photoinhibition at 7°C than leaves of the japonica type. Photoinhibition was dependent both on temperature and light, increasing nonlinearly with decreasing temperature and linearly with increasing light intensity. In contrast to photoinhibition during chilling, large differences, up to 166-fold, were found in the relative susceptibility of the different species to chilling injury in the dark. It was concluded that chilling temperatures increased the likelihood of photoinhibition in leaves of both chilling-sensitive and -resistant plants. Further, while the photoinhibition during chilling generally occurred more rapidly in chilling-sensitive plants, this was not related directly to chilling sensitivity.     

Gas Exchange Analysis of the Relative Importance of Stomatal and Biochemical Factors in Photosynthetic Induction in Alocasia macrorrhiza

When leaves of Alocasia macrorrhiza adapted to 10 micromole quanta per square meter per second were transferred to 500 micromole quanta per square meter per second, the rate of photosynthetic CO₂ assimilation increased for over 45 minutes. For the first 10 to 15 minutes, increases in both stomatal conductance and the leaf's photosynthetic capacity were responsible for the increase in assimilation rate. Thereafter, continuing increases in stomatal conductance were almost entirely responsible for further increases in assimilation rate. When conductances were initially high, assimilation rates 1 minute after the increase in photon flux density could be more than six times as high as for similar leaves with initially low conductance. Further increases in assimilation rate in these leaves with high conductance were predominantly due to increases in the induction state at the biochemical level and followed an exponential time course. When stomatal conductances were initially low, then increases in conductance were predominantly responsible for the increases in assimilation rate, with both following a sigmoidal time course. In these leaves, it was important to also consider the effect of cuticular water loss on the calculation of the intracellular partial pressure of CO₂, and an assessment of the relative importance of stomatal conductance differed considerably from one that did not include cuticular water loss.     

Photosynthetic and stomatal responses of spinach leaves to salt stress

The gas exchange of spinach plants, salt-stressed by adding NaCl to the nutrient solution in increments of 25 millimolar per day to a final concentration of 200 millimolar, was studied 3 weeks after starting NaCl treatment. Photosynthesis became light saturated at 1100 to 1400 micromoles per square meter per second in salt-treated plants and at approximately 2000 micromoles per square meter per second in control plants. Photosynthetic capacity of the mesophyll measured as a function of intercellular partial pressure of CO₂ at the light intensity prevailing during growth and at light saturation were both decreased in the salttreated plants. The CO₂ compensation points and relative enhancements of photosynthesis at low O₂ were not affected by salinity. The lower photosynthetic rates in salt-treated leaves at 450 micromoles per square meter per second were associated with a 70% reduction in stomatal conductance and low intercellular CO₂ (219 microbars; cf. 285 microbars for controls). Increasing photon flux density to light saturation extended the linear portions of the CO₂ response curves, increased stomatal conductances, increased intercellular CO₂ in the salt-treated plants, but lowered it in controls, and accentuated differences in photosynthetic rate (area basis) between the treatments.

Leaves from salt-treated plants were thicker but contained about 73% of the chlorophyll per unit area of control plants. When photosynthetic rates were expressed on a chlorophyll basis there was no difference in initial slope of assimilation versus intercellular CO₂ between treatments. Photosynthetic rates (chlorophyll basis) at light saturation differed only by 20% which was also observed earlier with isolated, intact chloroplasts (Robinson et al. 1983 Plant Physiol 73: 238-242).

Measurement of carbon isotope ratio revealed less discrimination against ¹³C with salt treatment and confirmed the persistence of low intercellular partial pressures of CO₂ during plant growth. The development of a thicker leaf with less chlorophyll per unit area during salt treatment permitted stomatal conductance and intercellular partial pressure of CO₂ to decline without restricting photosynthesis and had the benefit of greatly increasing water use efficiency.

     

Leaf photosynthetic parameters of two maize (Zea mays) cultivars in response to various patterns of chilling temperatures and photon flux densities

Chilling‐induced photosynthetic impairment was examined in leaves of maize (Zea mays L.) seedlings of two cultivars, one adapted to western Europe and one adapted to Mexican highlands. Three experiments were performed in a controlled environment. The effects of chilling night temperatures, of chilling at high light intensity and of variable chilling day temperatures on photosynthetic parameters, were evaluated. Chilling in the dark period resulted in stomatal limitation of net photosynthesis. Chilling at moderate to high light intensities caused chilling‐dependent photoinhibition of CO₂ uptake. Photobleached maize leaves did not resume normal photosynthetic function. Maize cv. Batan 8686 from the highlands of Mexico was less susceptible to photosynthetic damage than maize cv. Bastion adapted for cultivation in W. Europe, when exposed to chilling night temperatures, or to mild chilling photoinhibitory conditions.     

Photosynthetic Responses to Dynamic Light Environments by Hawaiian Trees : Time Course of CO(2) Uptake and Carbon Gain during Sunflecks

Gas exchange responses to rapid changes in light were studied in a C₃ tree, Claoxylon sandwicense Muell-Arg and a C₄ tree, Euphorbia forbesii Sherff that are native to the understory of a mesic Hawaiian forest. When light was increased to 500 micromoles per meter per second following a 2 hour preexposure at 22 micromoles per meter per second, net CO₂ uptake rates and stomatal conductance gradually increased for over 1 hour in C. sandwicense but reached maximum values within 30 minutes in E. forbesii. Calculation of the intercellular CO₂ pressures indicated that the primary limitation to CO₂ uptake during this induction was nonstomatal in both species. The photosynthetic response to simulated sunflecks (lightflecks) was strongly dependent on the induction state of the leaf. Total CO₂ uptake during a lightfleck was greater and the response was faster after exposure of the leaf to high light than when the leaf had been exposed only to low light for the previous 2 hours. During a series of lightflecks, induction resulted in increased CO₂ uptake in successive lightflecks. Significant postillumination CO₂ fixation was evident and contributed substantially to the total carbon gain, especially for lightflecks of 5 to 20 seconds' duration.