Veränderliche Markierungsmuster bei 14CO2-Fütterung von Bryophyllum tubiflorum zu verschiedenen Zeitpunkten der Hell-Dunkelperiode

Summary The distribution of radioactivity between the products of ¹⁴CO₂ light fixation in phyllodia of Bryophyllum tubiflorum could be influenced experimentally by manipulating the malic acid content of the cells. Accelerating the deacidification of the tissue during the light period by application of higher light intensities accelerated the increase of malate labelling and the decrease of the sucrose labelling after ¹⁴CO₂ light fixation under our standard conditions (10 min preillumination, 15 min ¹⁴CO₂ light fixation, 8000 lux).In other experiments different malate contents of the tissues were induced by treating the phyllodia with different temperatures during the night period. In the morning, phyllodia with low malate content transferred most of the label into malate, and phyllodia with high malate content incorporated most of the ¹⁴C radioactivity into sugars. However, this was true only after preillumination of 1 hour. When the phyllodia fixed ¹⁴CO₂ without preillumination, no differences in the labelling patterns between acidified and non-acidified phyllodia could be observed.In experiments using leaf tissue slices of Bryophyllum daigremontianum we could again observe that malate was labelled more heavily in the deacidified tissue than in the acidified controls, with less radioactivity being transferred into phosphate esters and sugars. The rates of ¹⁴CO₂ light fixation were identical in tissue slices with high and low malate content. However, the rates of CO₂ dark fixation in the acidified samples were clearly lower than those in the deacidified ones. The low rate of CO₂ dark fixation in acidified samples could not be inhibited by an inhibitor of PEP-carboxylase as the high CO₂ dark fixation rate of the deacidified tissue could be inhibited.The results are discussed in relation to the feed back inhibition of PEP-carboxylase in vivo by malate. Compartmentation also seemed to be involved in controlling the flow of carbon during CO₂ light fixation in succulent tissue.     

Gas Exchange Characteristics of the Submerged Aquatic Crassulacean Acid Metabolism Plant, Isoetes howellii

The submerged aquatic plant Isoetes howellii Engelmann possesses Crassulacean acid metabolism (CAM) comparable to that known from terrestrial CAM plants. Infrared gas analysis of submerged leaves showed Isoetes was capable of net CO₂ uptake in both light and dark. CO₂ uptake rates were a function of CO₂ levels in the medium. At 2,500 microliters CO₂ per liter (gas phase, equivalent to 1.79 milligrams per liter aqueous phase), Isoetes leaves showed continuous uptake in both the light and dark. At this CO₂ level, photosynthetic rates were light saturated at about 10% full sunlight and were about 3-fold greater than dark CO₂ uptake rates. In the dark, CO₂ uptake rates were also a function of length of time in the night period. Measurements of dark CO₂ uptake showed that, at both 2,500 and 500 microliters CO₂ per liter, rates declined during the night period. At the higher CO₂ level, dark CO₂ uptake rates at 0600 h were 75% less than at 1800 h. At 500 microliters CO₂ per liter, net CO₂ uptake in the dark at 1800 h was replaced by net CO₂ evolution in the dark at 0600 h. At both CO₂ levels, the overnight decline in net CO₂ uptake was marked by periodic bursts of accelerated CO₂ uptake. CO₂ uptake in the light was similar at 1% and 21% O₂, and this held for leaves intact as well as leaves split longitudinally. Estimating the contribution of light versus dark CO₂ uptake to the total carbon gain is complicated by the diurnal flux in CO₂ availability under field conditions.     

Diurnal fluctuation of light and dark CO2 fixation in peeled and unpeeled leaves of Bryophyllum daigremontiana

Diurnal fluctuation of light and dark CO₂ fixation in peeledand unpeeled leaves of Bryophyllum daigremontiana was examined.A distinct difference in light CO₂ fixation was observed inunpeeled leaves but not in peeled ones. No measurable differencein dark CO₂ fixation was observed in either type. These resultsindicate that the leaves of CAM plants have a high capacityfor CO₂ fixation in the daytime, but it is suppressed by theclosing of the stomata. Also, the rapid depression of CO₂ uptakewhen the illumination was directed at on dark acidified leavescould be prevented by peeling off the epidermis. The net photosyntheticCO₂ uptake in peeled leaves was 77 µmoles/mg chllrophyll/hrin the 3rd leaf and 62 in the 4th leaf. (Received August 7, 1978; )     

Diurnaler Säurerhythmus bei Tillandsia usneoides: Untersuchungen über den Weg des Kohlenstoffs sowie die Abhängigkeit des CO2-Gaswechsels von Lichtintensität,Temperatur und Wassergehalt der Pflanze

Summary Tillandsia usneoides, in the common sense a non-succulent plant, exhibits CO₂ exchange characterized by net CO₂ dark fixation during the night and depression of CO₂ exchange during the day. Malate has been demonstrated to accumulate during CO₂ dark fixation and to be converted to carbohydrates in light. Thus, T. usneoides exhibits CAM like typical succulents.Net CO₂ uptake during the day is increased with net CO₂ output being suppressed in duration of time and extent when light intensity increases. Furthermore, a slight increase in CO₂ fixation during the following night can be observed if the plants were treated with high light intensity during the previous day.Curves of CO₂ exchange typical for CAM are obtained if T. usneoides is kept at 15°C and 20°C. Lower temperature tend to increase CO₂ uptake during the day and to inhibit CO₂ dark fixation. Temperatures higher than 20°C favour loss of CO₂ by respiration, which becomes apparent during the whole day and night at 30°C and higher temperatures. Thus, T. usneoides gains carbon only at temperatures well below 25°C.Net CO₂ uptake during the day occurs only in moist plant material and is inhibited in plants cept under water stress conditions. However, CO₂ uptake during the night is clearly favoured if the plants dry out. Therefore dry plants gain more carbon than moist ones.Curves of CO₂ exchange typical for CAM were also obtained with 13 other species of the genus Tillandsia.The exhibition of CAM by the non-succulent T. usneoides calls for a new definition of the term succulence if it is to remain useful in characterizing this metabolic pathway. Because CO₂-fixing cells of T. usneoides possess relatively large vacuoles and are relatively poor in chloroplasts, they resembles the assimilatory cells of typical CAM-exhibiting succulents. Therefore, if succulence only means the capacity of big vacuoles to store malate, the assimilatory cells in T. usneoides are succulent. It seems to be useful to investigate parameters which would allow a definition of the term succulence on the level of the cell rather than on the level of the whole plant or plant organs.     

The role of the epidermis in the generation of the circadian rhythm of carbon dioxide fixation in leaves of Bryophyllum fedtschenkoi

The role of the epidermis in the generation of the endogenous circadian rhythm of CO₂ exchange in leaves of Bryophyllum fedtschenkoi has been examined. At 25° C the rhythm of CO₂ output exhibited by whole leaves kept in continuous darkness and an initially CO₂-free air stream also occurs in isolated pieces of mesophyll. The sensitivity to light of the rhythms in whole leaves and in isolated mesophyll appears to be identical. At 15° C, however, no rhythm is observed in isolated mesophyll tissue, despite there being a conspicuous rhythm in intact leaves. The rhythm of net CO₂ assimilation in whole leaves kept in continuous light and a stream of normal air at either 25° C or at 15° C is abolished by removal of the epidermis, although at 15° C and under the higher of the two light levels used, there is an indication that rhythmicity may begin to reappear after the third day of the experiment. Thus, only under certain environmental conditions is the rhythm of CO₂ exchange in Bryophyllum leaves independent of the epidermis. The results indicate that the rhythm of carbon dioxide fixation in continuous darkness and CO₂-free air is generated primarily in the mesophyll cells, whereas the rhythm in continuous light and normal air is generated in the stomatal guard cells or in an interaction of these cells with the mesophyll cells.Abbreviation PEPCase phosphoenolpyruvate carboxylase     

Dependence of CO2 gas exchange and acid metabolism of the alpine CAM plant Sempervivum montanum on temperature and light

Summary The influence of light intensity and temperature on the diurnal course and magnitude of CO₂ gas exchange and on acid metabolism was studied in the laboratory with rooted rosettes of Sempervivum montanum collected at 2,200 m above sea level in the Central Alps. Under a temperature regime having a cool dark period and warm light period, S. montanum exhibited the time course of CO₂ gas exchange typical of a CAM plant; the response was very distinct even when the plants were well-watered. At day temperatures of less than 10° C and at night temperatures greater than 35° C, S. montanum behaved like a C₃ plant. Characteristic for S. montanum are a broad temperature optimum and a wide range of temperatures in which CO₂ uptake in light is possible (-2° to 45° C). Dark fixation of CO₂ is evident between-2° and 35° C, an apparent uptake of external CO₂, on the other hand, only as high as 20° C. Light saturation of CO₂ uptake is reached at 60–80 W m^-2 while the rate of deacidification is nearly maximal at 40 W m^-2. These results show that, due to their specific metabolism, CAM plants can be favored not only in xeric habitats, but also in heat stressed mountain habitats where the daily variation in temperature may be extreme.Dedicated with appreciation to Dr. K.F. Springer     

Crassulacean acid metabolism in the Basellaceae (Caryophyllales)

• C₄ and crassulacean acid metabolism (CAM) have evolved in the order Caryophyllales many times but neither C₄ nor CAM have been recorded for the Basellaceae, a small family in the CAM‐rich sub‐order Portulacineae.
• 24 h gas exchange and day–night changes in titratable acidity were measured in leaves of Anredera baselloides exposed to wet–dry–wet cycles.
• While net CO₂ uptake was restricted to the light period in well‐watered plants, net CO₂ fixation in the dark, accompanied by significant nocturnal increases in leaf acidity, developed in droughted plants. Plants reverted to solely C₃ photosynthesis upon rewatering.
• The reversible induction of nocturnal net CO₂ uptake by drought stress indicates that this species is able to exhibit CAM in a facultative manner. This is the first report of CAM in a member of the Basellaceae.

     

Diurnal Pattern of Transpiration,Water Uptake and Water Budget of Succulents with Different CO2 Fixation Pathways

The water fluxes and the CO₂ exchange of three leaf succulents, Othonna opima, Cotyledon orbiculata and Senecio medley-woodii, with different leaf anatomy, growth form and CO₂ fixation pathways (C₃, CAM) were monitored with a gas exchange cuvette which was combined with a potometric system to quantify water uptake. Measurements, which are primarily valid for plants with a sufficient water supply, were made during 6 to 10 consecutive days under constant experimental conditions. Water uptake for 24 h exceeded water loss by transpiration only for a S, medley-woodii plant with 10 expanding but only 7 mature leaves. In this case the gained water evidently is put into leaf expansion. All other plants showed balanced transpiration and water uptake rates. O. opima and C. orbiculata have a similar life form, similar water storage volumes and the same natural habitat but their diurnal water uptake patterns differ significantly. In the C₃ plant O. opima water uptake increased when the transpiration increased or transpiration rates were higher than uptake rates and vice versa. On the contrary the CAM plant C. orbiculata transpired during the dark period at constant or decreasing rates but showed steadily increasing uptake rates. Senecio medley-woodii- and C. orbiculata are CAM plants with similar diurnal water uptake patterns with its maximum in uptake during or towards the end of the CO₂ dark fixation period. Water uptake of C. orbiculata was at its minimum at the end of the light period despite transpiration being maximal. The results were discussed considering the different CO₂ fixation pathways. In the investigated CAM succulents, C. orbiculata and S. medley-woodii, the CAM influenced water uptake throughout the whole day and not only during the CO₂ dark fixation period.     

Light and dark CO2 fixation in Clusia uvitana and the effects of plant water status and CO2 availability

Summary In well-watered plants of Clusia uvitana, a species capable of carbon fixation by crassulacean acid metabolism (CAM), recently expanded leaves gained 5 to 13-fold more carbon during 12 h light than during 12 h dark periods. When water was withheld from the plants, daytime net CO₂ uptake strongly decreased over a period of several days, whereas there was a marked increase in nocturnal carbon gain. Photosynthetic rates in the chloroplasts were hardly affected by the water stress treatment, as demonstrated by measurements of chlorophyll a fluorescence of intact leaves, indicating efficient decarboxylation of organic acids and refixation of carbon in the light. Within a few days after rewatering, plants reverted to the original gas exchange pattern with net CO₂ uptake predominantly occurring during daytime. The reversible increase in dark CO₂ fixation was paralleled by a reversible increase in the content of phosphoenolpyruvate (PEP) carboxylase protein. In wellwatered plants, short-term changes in the degree of dark CO₂ fixation were induced by alterations in CO₂ partial pressure during light periods: a decrease from 350 to 170 bar CO₂ caused nocturnal carbon gain, measured in normal air (350 bar), to increase, whereas an increase to 700 bar CO₂, during the day, caused net dark CO₂ fixation to cease. The increased CAM activity in response to water shortage may, at least to some extent, be directly related to the reduced carbon gain during daytime.     

Veränderliche Markierungsmuster bei14CO2-Fütterung vonBryophyllum tubiflorum zu verschiedenen Zeitpunkten der Hell/Dunkelperiode

Summary Detached phyllodia ofBryophyllum tubiflorum were fed under illumination with¹⁴CO₂ at different times during the light/dark period (12:12 hours). After photosynthesis in presence of¹⁴CO₂ during the intrinsic dark period the greatest part of soluble radioactivity was found in malate. When the same experiment was repeated during the light period, radioactivity was incorporated mainly into sucrose in the first hours while malate was labelled rather weakly. In the late afternoon (last third of the light period), malate became most heavily labelled again during photosynthesis with¹⁴CO₂.Our results indicate that the synthesis of malate by PEP-carboxylase/malate dehydrogenase is inhibited at certain times during the night/day period by end product inhibition of PEP-carboxylase, as was demonstrated byQueiroz (1967, 1968) andTing (1968) in vitro.During inhibition of the PEP-carboxylase there is no competition between the synthesis of malate and CO₂-fixation by the Calvin cycle. Thus radioactivity can flow into sucrose via the Calvin cycle during this time. When the malate content of the phyllodia is low, CO₂-fixation by PEP-carboxylase is not inhibited. Now this pathway dominates over photosynthesis via the Calvin cycle, for PEP-carboxylase has a higher affinity for CO₂ than carboxydismutase. Therefore malate now becomes more labelled than sucrose.     

Einfluß der Temperatur auf die Lichtatmung der Blaualge Anacystis nidulans

Summary CO₂ exchange, ¹⁴CO₂ fixation and ¹⁴C-products of Anacystis nidulans (strain L 1402-1) were studied during the induction period at temperatures of +15°C and+35°C. At+15°C the stationary rates of CO₂ uptake and respiration were reached directly. At+35°C a maximum of CO₂ uptake could be observed at the beginning of the illumination period followed by a lower steady rate of photosynthesis. In the following dark period a CO₂ gush appeared at+35°C. The magnitude of the CO₂ outburst is relatively independent of the photosyntbetic period. The autoradiographic studies showed that the Calvin cycle is the main carboxylation pathway in Anucystis. At a temperature of +35°C serine was labelled after 20 sec of photosynthesis. At+15°C, on the other hand, a low radio-activity appeared in serine after 5 min of photosynthesis. The results show that photorespiration of Anacystis is stimulated by high temperatures.     

The role of temperature in the regulation of the circadian rhythm of CO2 fixation in Bryophyllum fedtschenkoi

Detached leaves of Bryophyllum fedtschenkoi Hamet et Perrier kept in normal air show a single period of net CO₂ fixation on transfer to constant darkness at temperatures in the range 0–25 °C. The duration of this initial fixation period is largely independent of temperature in the range 5–20 °C, but lengthens very markedly at temperatures below 4 °C, and is reduced at temperatures above 25 °C. The onset of net fixation of CO₂ on transfer of leaves to constant darkness is immediate at low temperatures, but is delayed as the temperature is increased. The ambient temperature also determines whether or not a circadian rhythm of CO₂ exchange occurs. The rhythm begins to appear at about 20 °C, is most evident at 30 °C and becomes less distinct at 35 °C. The occurrence of a distinct circadian rhythm in CO₂ output at 30° C in the absence of a detectable rhythm in PEPCase kinase activity shows that the kinase rhythm is not a mandatory requirement for the rhythm of PEPCase activity. However, when it occurs, the kinase rhythm undoubtedly amplifies the PEPCase rhythm.Abbreviation PEPCase phosphoenolpyruvate carboxylase We thank the Agricultural and Food Research Council for financial support for this work.     

The subdominant status of Echinocereus viridiflorus and Mammillaria vivipara in the shortgrass prairie: The role of temperature and water effects on gas exchange

Summary The subdominant CAM species, Echinocereus viridiflorus and Mammillaria vivipara, collected from the shortgrass prairie in northeastern Colorado were pretreated and analyzed for gas exchange under cool temperatures (20/15°C) and warm temperatures (35/15°C). Well watered plants of both species under a 35/15°C thermoperiod fixed atmospheric CO₂ during the night and early moring. Echinocereus viridiflorus grown and analyzed at 20/15°C fixed CO₂ during the night, early morning and late afternoon but total carbon gain over a 24 h period is less than when grown and analyzed under the 35/15°C thermoperiod. Mammillaria vivipara grown and analyzed at 20/15°C assimilates CO₂ at low rates during all parts of a 24 h period with the greatest CO₂ fixation rates occuring from midday to late afternoon. The total carbon gain under the 20/15°C thermoperiod is less than that for this species under the 35/15°C thermoperiod. Decreasing the night temperature of plants grown under the warm conditions to 10°C or 5°C results in a depression of the night CO₂ fixation in both species. E. viridiflorus from the cool growth conditions showed an enhancement of the CO₂ uptake during the night, early morning and late afternoon when subjected to the cooler night temperatures (10°C and 5°C). The CO₂ uptake of M. vivipara grown at 20/15°C shows an enhancement during the night and early morning while the CO₂ fixation during midday and late afternoon is slightly depressed under cool night temperatures (10° and 5°C). Under the 35/15°C thermoperiod both species exhibit depressed rates of CO₂ fixation during the night and early morning when water stressed. Plants of both species grown under the 20/15°C thermoperiod exhibit no net CO₂ fixation following five weeks of water deprivation. Upon rewatering, E. viridiflorus begins to recover its capacity for CO₂ fixation within 24 h under both the warm and cool temperature regimes. However, M. vivipara did not show recovery within 48 h following rewatering under the warm or cool temperature regime. Contrasting the patterns of gas exchange of the subdominant species, E. viridiflorus and M. vivipara, with a dominant CAM species of the shortgrass prairie, Opuntia polyacantha reveals significant differences that may well dictate the role of these species in this ecosystem. E. viridiflorus and M. vivipara have a lower capacity of carbon gain and recovery from water stress than O. polyacantha mainly due to their lack of late afternoon CO₂ uptake. This study suggests that carbon gain plays an important role in limiting E. viridiflorus and M. vivipara in the shortgrass prairie ecosystem.     

Responses of CAM species to increasing atmospheric CO2 concentrations

Crassulacean acid metabolism (CAM) species show an average increase in biomass productivity of 35% in response to a doubled atmospheric CO₂ concentration. Daily net CO₂ uptake is similarly enhanced, reflecting in part an increase in chlorenchyma thickness and accompanied by an even greater increase in water‐use efficiency. The responses of net CO₂ uptake in CAM species to increasing atmospheric CO₂ concentrations are similar to those for C₃ species and much greater than those for C₄ species. Increases in net daily CO₂ uptake by CAM plants under elevated atmospheric CO₂ concentrations reflect increases in both Rubisco‐mediated daytime CO₂ uptake and phosphoenolpyruvate carboxylase (PEPCase)‐mediated night‐time CO₂ uptake, the latter resulting in increased nocturnal malate accumulation. Chlorophyll contents and the activities of Rubisco and PEPCase decrease under elevated atmospheric CO₂, but the activated percentage for Rubisco increases and the K_M(HCO₃ ^? ) for PEPCase decreases, resulting in more efficient photosynthesis. Increases in root:shoot ratios and the formation of additional photosynthetic organs, together with increases in sucrose‐Pi synthase and starch synthase activity in these organs under elevated atmospheric CO₂ concentrations, decrease the potential feedback inhibition of photosynthesis. Longer‐term studies for several CAM species show no downward acclimatization of photosynthesis in response to elevated atmospheric CO₂ concentrations. With increasing temperature and drought duration, the percentage enhancement of daily net CO₂ uptake caused by elevated atmospheric CO₂ concentrations increases. Thus net CO₂ uptake, productivity, and the potential area for cultivation of CAM species will be enhanced by the increasing atmospheric CO₂ concentrations and the increasing temperatures associated with global climate change.     

Comparative ecophysiology of CAM and C3 bromeliads. II. Field measurements of gas exchange of CAM bromeliads in the humid tropics

Abstract The results described represent the first detailed measurements of gas exchange of epiphytic plants with crassulacean acid metabolism (CAM) in the humid tropics. A portable steady-state CO₂ and H₂O porometer was used to measure net exchange rates of CO₂ and H₂O vapour (J_CO2, J_H2O), leaf temperature (T₁), air temperature (T_A), air relative humidity (RH) and photosynthetically active radiation (PAR) for bromeliads in the field during the dry season in February and March 1983 on the tropical island of Trinidad. Different lengths of tubing (up to 25 m) were used so that the gas exchange could be measured of bromeliads in situ in their epiphytic habitats. Derived parameters such as leaf-air water-vapour-concentration difference (Δ_w), water-vapour conductance of leaves (g) and internal CO₂ partial pressure (pⁱ_CO2) could be calculated. The particular problems of making such measurements in the humid tropics due to high relative humidities and high dew-point temperatures are discussed. The long and often broad, strap-like leaves of bromeliads are well suited for measurements with the steady-state porometer. It is shown that CAM activity varies along the length of individual leaves, and variability between different leaves is also demonstrated. The major phases of CAM, i.e. nocturnal stomalal opening, CO₂ uptake and dark fixation as malic acid (Phase I), daytime stomatal closure and light-dependent assimilation of CO₂ derived from decarboxylation of the malic acid (Phase III), and late-afternoon stomatal opening with direct light-dependent assimilation of atmospheric CO₂ (Phase IV) were all clearly shown by CAM bromeliads in situ. Their expression and magnitude depended on the environmental conditions. An early-morning peak of CO₂ uptake as is characteristic of Phase II of CAM was not detected during the night-day transition. A bromeliad intermediate between C₃ and CAM, Guzmania monostachia, showed substantial net CO₂ uptake in the early morning but no net uptake integrated over the whole of the night.     

Differential usage of storage carbohydrates in the CAM bromeliad Aechmea 'Maya' during acclimation to drought and recovery from dehydration

CAM requires a substantial investment of resources into storage carbohydrates to account for nocturnal CO₂ uptake, thereby restricting carbohydrate partitioning to other metabolic activities, including dark respiration, growth and acclimation to abiotic stress. Flexible modulation of carbon flow to the different competing sinks under changing environmental conditions is considered a key determinant for the growth, productivity and ecological success of the CAM pathway. The aim of the present study was to examine how shifts in carbohydrate partitioning could assure maintenance of photosynthetic integrity and a positive carbon balance under conditions of increasing water deprivation in CAM species. Measurements of gas exchange, leaf water relations, malate, starch and soluble sugar (glucose, fructose and sucrose) contents were made in leaves of the CAM bromeliad Aechmea ‘Maya’ over a 6‐month period of drought and subsequently over a 2‐month period of recovery from drought. Results indicated that short‐term influences of water stress were minimized by elevating the level of respiratory recycling, and carbohydrate pools were maintained at the expense of export for growth while providing a comparable nocturnal carbon gain to that in well‐watered control plants. Longer term drought resulted in a disproportionate depletion of key carbohydrate reserves. Sucrose, which was of minor importance for providing substrate for the dark reactions under well‐watered conditions, became the major source of carbohydrate for nocturnal carboxylation as drought progressed. Flexibility in terms of the major carbohydrate source used to sustain dark CO₂ uptake is therefore considered a crucial factor in meeting the carbon and energy demands under limiting environmental conditions. Recovery from CAM‐idling was found to be dependent on the restoration of the starch pool, which was used predominantly for provision of substrate for nocturnal carboxylation, while net carbon export was limited. The conservation of starch for the nocturnal reactions might be adaptive with regard to responding efficiently to a return of water stress.     

Persistent circadian rhythms in the phosphorylation state of phosphoenolpyruvate carboxylase from Bryophyllum fedtschenkoi leaves and in its sensitivity to inhibition by malate

Phosphoenolpyruvate carboxylase (EC 4.1.1.31; PEPCase) from Bryophyllum fedtschenkoi leaves has previously been shown to exist in two forms in vivo. During the night the enzyme is phosphorylated and relatively insensitive to feedback inhibition by malate whereas during the day the enzyme is dephosphorylated and more sensitive to inhibition by malate. These properties of PEPCase have now been investigated in leaves maintained under constant conditions of temperature and lighting. When leaves were maintained in continuous darkness and CO₂-free air at 15°C, PEPCase exhibited a persistent circadian rhythm of interconversion between the two forms. There was a good correlation between periods during which the leaves were fixing respiratory CO₂ and periods during which PEPCase was in the form normally observed at night. When leaves were maintained in continuous light and normal air at 15°C, starting at the end of a night or the end of a day, a circadian rhythm of net uptake of CO₂ was observed. Only when these constant conditions were applied at the end of a day was a circadian rhythm of interconversions between the two forms of PEPCase observed and the rhythms of enzyme interconversion and CO₂ uptake did not correlate in phase or period.Abbreviations CAM Crassulacean acid metabolism - FW fresh weight - PEPCase phosphoenolpyruvate carboxylase - RuBPCase ribulose-1,5-bisphosphate carboxylase To whom correspondence should be addressed.     

Stomatal responses to humidity in Opuntia inermis in relation to control of CO2 and H2O exchange patterns

Summary At constant cladode temperature the stomatal resistance of O. inermis increased when the cladode-air vapor pressure difference was increased and stomatal resistance decreased when the cladode-air vapor pressure difference was lowered. Net CO₂ fixation in the dark was very responsive to these humidity dependent changes in stomatal resistance. Net CO₂ fixation and stomatal resistance in the light did not respond to changes in cladode-air vapor pressure differences in the light under the conditions tested. When temperature response functions for dark CO₂ fixation were examined at constant ambient humidity, the reduction in dark CO₂ fixation at higher temperatures was largely due to stomatal closure in response to the increased vapor pressure difference. The water requirement for net CO₂ fixation in the dark at typical nocturnal vapor pressure differences was about 10 times lower than that of net CO₂ fixation in the light at vapor pressure differences typical of the late afternoon. The role of the stomatal responses to humidity in determining the patterns and rates of net CO₂ exchange in the light or dark, and its possible ecological significance is discussed.     

Effects of temperature on the activity of phosphoenolpyruvate carboxylase and on the control of CO2 fixation in Bryophyllum fedtschenkoi

The phosphorylation state and the malate sensitivity of phosphoenolpyruvate carboxylase (PEPCase, EC 4.1.1.31) in Bryophyllum fedtschenkoi Hamet et Perrier are altered by changes in the ambient temperature. These effects, in turn alter the in-vivo activity of the enzyme. Low temperature (3 °C or less), stabilizes the phosphorylated form of the enzyme, while high temperature (30 °C) promotes its dephosphorylation. The catalytic activity of the phosphorylated and dephosphorylated forms of PEPCase increases with temperature, but the apparent K _i values for malate of both forms of the enzyme decrease. Results of experiments with detached leaves maintained in darkness in normal air indicate that the changes in malate sensitivity and phosphorylation state of PEPCase with temperature are of physiological significance. When the phosphorylated form of PEPCase is stabilized by reducing the temperature of leaves 9 h after transfer to constant darkness at 15 °C, a prolonged period of CO₂ fixation follows. When leaves are maintained in constant darkness at 15 °C until CO₂ output reaches a low steady-state level and the PEPCase is dephosphorylated, reducing the temperature to 3 °C results in a further period of CO₂ fixation even though the phosphorylation state of PEPCase does not change.Abbreviations CAM Crassulacean acid metabolism - PEP phosphoenolpyruvate - PEPCase phosphoenolpyruvate carboxylase We thank the Agricultural and Food Research Council for financial support for this work.     

The effect of different night conditions on the CO2 fixation in a lichen Xanthoria parietina

CO₂ fixation was studied in a lichen, Xanthoria parietina, kept in continuous light, and with cyclic changes in light intensity, dark period or temperature. The diurnal and seasonal courses of CO₂ exchange were followed. The rate of net photosynthesis was observed to fall from morning to evening, and this decline was more pronounced in winter than in summer. The maximal net photosynthetic rate, 223 ng CO₂g^-1dws^-1, occured in winter and the minimum, 94 ng CO₂g^-1dws^-1, late in spring. The light compensation point in summer was four times as high as in winter. In continuous light (180 or 90 mol photons m^-2s^-1, 15°C) net photosynthesis decreased noticeably during one week, falling below the level maintained in a 12 h light: 12 h dark cycle. Photosynthetic activity did not decrease, however, in lichens held in continuous light (90 mol photons m^-2s^-1) with cyclic changes of temperature (12 h 20 °C: 12 h 5 °C). Active photosynthesis was also maintained in light of cyclically changing intensity (12 h: 12 h, 15 °C) when night-time light was at least 75% lower than illumination by day. A dark period of 4 hours in a 24-h light:dark cycle was sufficient to keep CO₂ fixation at the control level. It seems that plants need an unproductive period during the day to survive and this can be induced by fluctuations in light and/or temperature.