铁皮石斛叶片光合作用的碳代谢途径

 利用LI-6400光合测定系统测定了不同天气条件下铁皮石斛（Dendrobium officinale）叶片24h CO2吸收的动态以及CO2吸收对光强和温度的响应。晴天的白天和夜间铁皮石斛都能吸收CO2,中午CO2吸收速率为负值, CO2的交换方式具景天酸代谢途径(CAM)的特点。阴雨天,只有白天吸收CO2,夜间表现为暗呼吸,光合作用碳代谢的途径为C3途径。在多云的天气条件下,白天吸收CO2,并持续至日落后。夜间21∶00仍有CO2吸收,23∶00以后至次日凌晨处于暗呼吸状态。在500 μmol·m-2·s-1光照件下,20℃出现最大CO2吸收值。在夜间,25℃时CO2的吸收速率最高。有光和无光条件下,低温或高温引起CO2吸收速率下降均为非气孔因素所致。晴天上午,铁皮石斛叶片的表观量子产额为0.035,光合补偿点为2.9μmol·m-2·s-1,饱和光强为500μmol·m-2·s-1,强光下出现光抑制现象。叶片受到强光预先照射后,即使光照减弱光抑制效应仍保持一段时间,致使光合补偿点升高,表观量子产额下降,相同光强下的CO2吸收效率降低。结果表明：铁皮石斛为兼性CAM植物,随着环境条件的变化,其光合作用在景天酸代谢途径(CAM)与C3途径间变化。     

几种CAM植物PEP羧化酶同工酶的比较研究

比较研究几种兼性和专一性ＣＡＭ植物材料的ＰＥＰＣ同工酶表明：经自然干旱诱导,兼性ＣＡＭ植物露花（Ｍｅｓｅｍｂｒｙａｎｔｈｅｍｕｍｃｏｒｄｉｆｏｌｉｕｍ）、长药景天（Ｓｕｄｕｍｓｐｅｃｔａｂｉｌｅ）有新的ＰＥＰＣ同工酶的出现,诱导前后各同工酶的天然分子量变化不大;而土三七（Ｓｅｄｕｍａｉｚｏｏｎ）则没有新的ＰＥＰＣ同工酶出现,但诱导后其同工酶的天然分子量有所增大。以上几种兼性ＣＡＭ植物的ＰＥＰＣ同工酶酶谱无明显昼夜变化。专一性ＣＡＭ植物的ＰＥＰＣ酶谱和天然分子量均较一致,亦无昼夜差异。     

The "Kluge-Lüttge Kammer": a preliminary evaluation of an enclosed, Crassulacean acid metabolism (CAM) Mesocosm that allows separation of synchronized and desynchronized contributions of plants to whole system gas exchange

Crassulacean acid metabolism (CAM) is recognized as a photosynthetic adaptation of plants to arid habitats. This paper presents a proof-of-concept evaluation of partitioning net CO2 exchanges for soil and plants in an arid, exclusively CAM mesocosm, with soil depth and succulent plant biomass approximating that of natural Sonoran Desert ecosystems. We present the first evidence that an enclosed CAM-dominated soil and plant community exposed to a substantial day/night temperature difference (30/20 degrees C), exhibits a diel gas exchange pattern consisting of four consecutive phases with a distinct nocturnal CO2 uptake. These phases were modulated by plant assimilation and soil respiration processes. Day-time stomatal closure of the CAM cycle during phase III was used to eliminate aboveground photosynthetic assimilation and respiration and thereby to estimate belowground plant plus soil respiration. Rapid changes in temperature appeared to synchronize single plant gas exchange but individual plant gas exchange patterns were desynchronized at constant day/night temperatures (25 degrees C), masking the distinct mesocosm pattern. Overall, the mean carbon budget of this CAM model Sonoran Desert system was negative, releasing an average of 22.5 mmol CO2 m-2 d-1. The capacity for nocturnal CO2 assimilation in this exclusively CAM mesocosm was inadequate to recycle CO2 released by plant and soil respiration.     

Diel leaf growth cycles in Clusia spp. are related to changes between C3 photosynthesis and crassulacean acid metabolism during development and during water stress

This study reports evidence that the timing of leaf growth responds to developmental and environmental constraints in Clusia spp. We monitored diel patterns of leaf growth in the facultative C₃-crassulacean acid metabolism (CAM) species Clusia minor and in the supposedly obligate CAM species Clusia alata using imaging methods and followed diel patterns of CO₂ exchange and acidification. Developing leaves of well-watered C. minor showed a C₃-like diel pattern of gas exchange and growth, with maximum relative growth rate (RGR) in the early night period. Growth slowed when water was withheld, accompanied by nocturnal CO₂ exchange and the diel acid change characteristic of CAM. Maximum leaf RGR shifted from early night to early in the day when water was withheld. In well-watered C. alata , similar changes in the diel pattern of leaf growth occurred with the development of CAM during leaf ontogeny. We hypothesize that the shift in leaf growth cycle that accompanies the switch from C₃ photosynthesis to CAM is mainly caused by the primary demand of CAM for substrates for nocturnal CO₂ fixation and acid synthesis, thus reducing the availability of carbohydrates for leaf growth at night. Although the shift to leaf growth early in the light is presumably associated with the availability of carbohydrates, source–sink relationships and sustained diurnal acid levels in young leaves of Clusia spp. need further evaluation in relation to growth processes.     

阴天和晴天对黄河三角洲芦苇湿地净生态系统CO2交换的影响

云量以及大气气溶胶含量变化引起的阴天和晴天会对局地的微气候环境产生综合效应, 影响地面接收的太阳辐射强度, 同时引起环境因子的变化, 最终对净生态系统CO₂交换(NEE)产生影响。该文通过涡度相关系统以及微气象梯度观测系统, 对黄河三角洲芦苇(Phragmites australis)湿地NEE以及环境要素进行了观测。在自然条件下选择12对相邻阴天和晴天数据, 在生物要素(生物量、叶面积指数)、土壤水分以及养分特征保持不变的前提下, 揭示了阴天和晴天变化对湿地生态系统NEE的光响应和温度响应的影响。结果表明: 12对阴天和晴天生态系统NEE的日平均动态均呈“U”型曲线, 但阴天NEE的变幅较小。晴天条件下湿地生态系统NEE的日均值显著高于阴天(p < 0.01)。阴天和晴天湿地生态系统NEE与光合有效辐射(PAR)之间均呈直角双曲线关系, 但晴天条件下, 最大光合速率(A_max)显著大于阴天(p < 0.01), 同时白天生态系统呼吸(R_eco,daytime)也显著大于阴天(p < 0.01)。不论阴天还是晴天, R_eco,daytime与气温均呈显著的指数关系。晴天湿地生态系统呼吸的温度敏感系数Q₁₀ (5.5)远大于阴天(1.9)。阴天和晴天昼间PAR差值以及气温差值对NEE差值的协同影响达到63%。     

CO(2)-concentrating: consequences in crassulacean acid metabolism

The consequences of CO(2)-concentrating in leaf air-spaces of CAM plants during daytime organic acid decarboxylation in Phase III of CAM (crassulacean acid metabolism) are explored. There are mechanistic consequences of internal CO(2) partial pressures, p(i)(CO(2)). These are (i) effects on stomata, i.e. high p(i)(CO(2)) eliciting stomatal closure in Phase III, (ii) regulation of malic acid remobilization from the vacuole, malate decarboxylation and refixation of CO(2) via Rubisco (ribulose bisphosphate carboxylase/oxygenase), and (iii) internal signalling functions during the transitions between Phases II and III and III and IV, respectively, in the natural day/night cycle and in synchronizing the circadian clocks of individual leaf cells or leaf patches in the free-running endogenous rhythmicity of CAM. There are ecophysiological consequences. Obvious beneficial ecophysiological consequences are (i) CO(2)-acquisition, (ii) increased water-use- efficiency, (iii) suppressed photorespiration, and (iv) reduced oxidative stress by over-energization of the photosynthetic apparatus. However, the general potency of these beneficial effects may be questioned. There are also adverse ecophysiological consequences. These are (i) energetics, (ii) pH effects and (iii) Phase III oxidative stress. A major consequence of CO(2)-concentrating in Phase III is O(2)-concentrating, increased p(i)(CO(2)) is accompanied by increased p(i)(O(2)). Do reversible shifts of C(3)/CAM-intermediate plants between the C(3)-CAM-C(3) modes of photosynthesis indicate that C(3)-photosynthesis provides better protection from irradiance stress? There are many open questions and CAM remains a curiosity.     

Drought-stress-induced up-regulation of CAM in seedlings of a tropical cactus, Opuntia elatior, operating predominantly in the C3 mode

Immediately after unfolding, cotyledons of the tropical platyopuntoid cactus, Opuntia elatior Mill., exhibited a C(3)-type diel CO(2) exchange pattern characterized by net CO(2) uptake in the light. Significant nocturnal increases in titratable acidity typical of crassulacean acid metabolism (CAM) were not detected at this early developmental stage. As cotyledons matured and the first cladode (flattened stem) developed, features of CAM were observed and the magnitude of CAM increased. Nonetheless, in well-watered seedlings up to 10 cm tall, C(3) photosynthetic CO(2) fixation in the light remained the major pathway of carbon fixation. Reduced soil water availability led to an up-regulation of net dark CO(2) fixation and greater nocturnal increases in tissue acidity, consistent with facultative CAM. These observations demonstrate that C(3) photosynthesis, drought-stress-related facultative CAM, and developmentally controlled constitutive CAM can all contribute to the early growth of O. elatior. The strong C(3) component and facultative CAM features expressed in young O. elatior contrast with mature plants in which obligate CAM is the major pathway of carbon acquisition.     

Seasonal Shifts of Photosynthesis in Portulacaria afra (L.) Jacq

Portulacaria afra (L.) Jacq., a perennial facultative Crassulacean acid metabolism (CAM) species, was studied under natural photoperiods and temperatures in San Diego, California. The plants were irrigated every fourth day throughout the study period. Measurements of ¹⁴CO₂ uptake, stomatal resistance, and titratable acidity were made periodically from July 1981 through May 1982. P. afra maintained C₃ photosynthesis during the winter and the spring. Diurnal acid fluctuations were low and maximal ¹⁴CO₂ uptake occurred during the day. The day/night ratio of carbon uptake varied from 5 to 10 and indicated little nocturnal CO₂ uptake. CAM photosynthesis occurred during the summer and a mixture of both C₃ and CAM during the fall. Large acid fluctuations of 100 to 200 microequivalents per gram fresh weight were observed and maximal ¹⁴CO₂ uptake shifted to the late night and early morning hours. Daytime stomatal closure was evident. A reduction in the day/night ratio of carbon uptake to 2 indicated a significant contribution of nocturnal CO₂ uptake to the overall carbon gain of the plant. The seasonal shift from C₃ to CAM was facilitated by increasing daytime temperature and accompanied by reduced daytime CO₂ uptake despite irrigation.     

Correlation between CAM-Cycling and Photosynthetic Gas Exchange in Five Species of Talinum (Portulacaceae)

Photosynthetic gas exchange and malic acid fluctuations were monitored in 69 well-watered plants from five morphologically similar species of Talinum in an investigation of the ecophysiological significance of the Crassulacean acid metabolism (CAM)-cycling mode of photosynthesis. Unlike CAM, atmospheric CO₂ uptake in CAM-cycling occurs exclusively during the day; at night, the stomata are closed and respiratory CO₂ is recaptured to form malic acid. All species showed similar patterns of day-night gas exchange and overnight malic acid accumulation, confirming the presence of CAM-cycling. Species averages for gas exchange parameters and malic acid fluctuation were significantly different such that the species with the highest daytime gas exchange had the lowest malic acid accumulation and vice versa. Also, daytime CO₂ exchange and transpiration were negatively correlated with overnight malic acid fluctuation for all individuals examined together, as well as within one species. This suggests that malic acid may effect reductions in both atmospheric CO₂ uptake and transpiration during the day. No significant correlation between malic acid fluctuation and water-use efficiency was found, although a nonsignificant trend of increasing water-use efficiency with increasing malic acid fluctuation was observed among species averages. This study provides evidence that CO₂ recycling via malic acid is negatively correlated with daytime transpirational water losses in well-watered plants. Thus, CAM-cycling could be important for survival in the thin, frequently desiccated soils of rock outcrops on which these plants occur.     

Evaluation of diel patterns of relative changes in cell turgor of tomato plants using leaf patch clamp pressure probes

Relative changes in cell turgor of leaves of well‐watered tomato plants were evaluated using the leaf patch clamp pressure probe (LPCP) under dynamic greenhouse climate conditions. LPCP changes, a measure for relative changes in cell turgor, were monitored at three different heights of transpiring and non‐transpiring leaves of tomato plants on sunny and cloudy days simultaneously with whole plant water uptake. Clear diel patterns were observed for relative changes of cell turgor of both transpiring and non‐transpiring leaves, which were stronger on sunny days than on cloudy days. A clear effect of canopy height was also observed. Non‐transpiring leaves showed relative changes in cell turgor that closely followed plant water uptake throughout the day. However, in the afternoon the relative changes of cell turgor of the transpiring leaves displayed a delayed response in comparison to plant water uptake. Subsequent recovery of cell turgor loss of transpiring leaves during the following night appeared insufficient, as the pre‐dawn turgescent state similar to the previous night was not attained.     

Crassulacean acid metabolism in the epiphytic ferns Drymoglossum piloselloides and Pyrrosia longifolia: studies on responses to environmental signals

Abstract The paper reports the results of the comprehensive study of crassulacean acid metabolism in two epiphytic tropical ferns, Drymoglossum piloselloides and Pyrrosia longifolia. The plants were investigated under different light, temperature and water status. It was found that both species are obligate CAM plants. The diurnal acidity rhythm is due to the fluctuation in malic acid concentration, which accounts for the change in titratable acidity. Besides malic acid, shikimate and oxalate are found to be present, but not contributing to the CAM acid rhythm. The diurnal rhythm of malic acid content results in a corresponding rhythm in leaf water relations. Both Φ_Φ and Φ_total, were lowest at the end of the night, i.e. when the level of malic acid was highest. The effects of temperature on CO₂ exchange were inverse to those observed in other CAM plants. In both ferns studied, dark CO₂ fixation increased when the night temperature was increased. Increase in day temperature reduced CO₂ uptake during phase IV and during the following night. The observed responses of the ferns to temperature changes suggest that the in situ environmental conditions are optimal for their CAM performance. In weak light, the plants showed net CO₂ output during the midday deacidification period. Increases in light intensity reduced such CO₂ output. Under drought conditions, the CO₂ exchange in the ferns was reduced to zero within 5–6 d, indicating that the ferns studied are more susceptible to water deficiency than other CAM plants. This could be due to a higher cuticular conductance for water. The results are discussed, in particular, in relation to CAM performance of epiphytes growing in the wet tropics.     

Cool-night temperature induces spike emergence and affects photosynthetic efficiency and metabolizable carbohydrate and organic acid pools in <Emphasis Type="Italic">Phalaenopsis aphrodite</Emphasis>

After being acclimated to constant warm (28 degrees C day/28 degrees C night) and cool-night temperature (28 degrees C day/20 degrees C night) regimes in growth chambers for 2 weeks, the two groups of mature Phalaenopsis aphrodite subsp. formosana plants both clearly exhibited a diurnal oscillation of stomatal conductance, net CO(2) uptake rate, malate and starch levels, and the phosphoenolpyruvate carboxylase (EC 4.1.1.31) and NAD(+)-malic enzyme (EC 1.1.1.39) activities. Hence, P. aphrodite is an obligate crassulacean acid metabolism plant. Nevertheless, different night temperature greatly affected both the stomatal conductance and the contribution of ambient and respiratory CO(2) to the nocturnal accumulation of malate. However, the amounts of nocturnal accumulated malate and daily deposited starch appeared to have no significant difference between the two groups. These results demonstrate that P. ahrodite is congruent with the characteristics of CAM plants having great flexibility and plasticity in response to changes in environmental conditions. In addition, the formation of reproductive stem, viz. spike, was noticeably inhibited by a constant warm temperature, but induced by a fluctuating warm day and cool night condition. The relationship between the metabolic pool variation and spike induction of Phalaenopsis is also discussed.     

On the nature of facultative and constitutive CAM: environmental and developmental control of CAM expression during early growth of Clusia, Kalanchöe, and Opuntia

The capacity to induce crassulacean acid metabolism developmentally (constitutive CAM) and to up-regulate CAM expression in response to drought stress (facultative CAM) was studied in whole shoots of seven species by measuring net CO(2) gas exchange for up to 120 day-night cycles during early growth. In Clusia rosea, CAM was largely induced developmentally. Well-watered seedlings began their life cycle as C(3) plants and developed net dark CO(2) fixation indicative of CAM after the initiation of the fourth leaf pair following the cotyledons. Thereafter, CAM activity increased progressively and drought stress led to only small additional, reversible increases in dark CO(2) fixation. In contrast, CAM expression was overwhelmingly under environmental control in seedlings and mature plants of Clusia pratensis. C(3)-type CO(2) exchange was maintained under well-watered conditions, but upon drought stress, CO(2) exchange shifted, in a fully reversible manner, to a CAM-type pattern. Clusia minor showed CO(2) exchange reponses intermediate to those of C. rosea and C. pratensis. Clusia cretosa operated in the C(3) mode at all times. Notably, reversible stress-induced increases of dark CO(2) fixation were also observed during the developmental progression to pronounced CAM in young Kalancho? daigremontiana and Kalancho? pinnata, two species considered constitutive CAM species. Drought-induced up-regulation of CAM was even detected in young cladodes of a cactus, Opuntia ficus-indica, an archetypal constitutive CAM species. Evidently, the defining characteristics of constitutive and facultative CAM are shared, to variable degrees, by all CAM species.     

Crassulacean acid metabolism and fitness under water deficit stress: if not for carbon gain,what is facultative CAM good for?

Background

In obligate Crassulacean acid metabolism (CAM), up to 99 % of CO₂ assimilation occurs during the night, therefore supporting the hypothesis that CAM is adaptive because it allows CO₂ fixation during the part of the day with lower evaporative demand, making life in water-limited environments possible. By comparison, in facultative CAM (inducible CAM, C₃-CAM) and CAM-cycling plants drought-induced dark CO₂ fixation may only be, with few exceptions, a small proportion of C₃ CO₂ assimilation in watered plants and occur during a few days. From the viewpoint of survival the adaptive advantages, i.e. increased fitness, of facultative CAM and CAM-cycling are not obvious. Therefore, it is hypothesized that, if it is to increase fitness, CAM must aid in reproduction.

Scope

An examination of published reports of 23 facultative CAM and CAM-cycling species finds that, in 19 species, drought-induced dark CO₂ fixation represents on average 11 % of C₃ CO₂ assimilation of watered plants. Evidence is discussed on the impact of the operation of CAM in facultative and CAM-cycling plants on their survival – carbon balance, water conservation, water absorption, photo-protection of the photosynthetic apparatus – and reproductive effort. It is concluded that in some species, but not all, facultative and cycling CAM contribute, rather than to increase carbon balance, to increase water-use efficiency, water absorption, prevention of photoinhibition and reproductive output.Key words: Facultative CAM, CAM-cycling, water, crassulacean acid metabolism, deficit     

Operation of the xanthophyll cycle and degradation of D1 protein in the inducible CAM plant,Talinum triangulare,under water deficit

Changes in photochemical activity induced by water deficit were investigated in Talinum triangulare, an inducible CAM plant. The aim was to analyse the interactions between C3 photosynthesis, induction and activity of CAM, photosynthetic energy regulation and the mechanisms responsible for photoprotection and photoinhibition under water stress. Gas exchange, chlorophyll a fluorescence, titratable acidity, carotenoid composition and relative contents of the PSII reaction centre protein (D1) were measured. A decrease in xylem tension (psi) from -0.14 to -0.2 MPa substantially decreased daytime net CO2 assimilation and daily carbon gain, and induced CAM, as shown by CO2 assimilation during the night and changes in titratable acidity; a further decrease in psi decreased nocturnal acid accumulation by 60%. Non-photochemical quenching of chlorophyll a fluorescence (NPQ) increased with water deficit, but decreased with a more severe drought (psi below -0.2 MPa), when CAM activity was low. NPQ was lower at 0900 h (during maximum decarboxylation rates) than at 1400 h, when malate pools were depleted. Down-regulation of PSII activity related to the rise in NPQ was indicated by a smaller quantum yield of PSII photochemistry (phiPSII) in droughted compared with watered plants. However, phiPSII was larger at 0900 h than at 1400 h. The de-epoxidation state of the xanthophyll cycle increased with drought and was linearly related to NPQ. Intrinsic quantum yield of PSII (FV/FM) measured at dusk was also lower in severely stressed plants than in controls. Under maximum photosynthetic photon flux and high decarboxylation rates of organic acids, the D1 content in leaves of droughted plants showing maximal CAM activity was identical to the controls; increased drought decreased D1 content by more than 30%. Predawn samples had D1 contents similar to leaves sampled at peak irradiance, with no signs of recovery after 12 h of darkness. It is concluded that under mild water stress, early induction of CAM, together with an increased energy dissipation by the xanthophyll cycle, prevents net degradation of D1 protein; when water deficit is more severe, CAM and xanthophyll cycle capacities for energy dissipation decline, and net degradation of D1 proceeds.     

Discrimination Processes and Shifts in Carboxylation during the Phases of Crassulacean Acid Metabolism

The magnitude and extent of Crassulacean acid metabolism (CAM) activity in two Clusia species was manipulated to investigate the regulation of the distinct CAM phases. First, in response to leaf-air vapor pressure deficit at night, changes in leaf conductance altered on-line carbon-isotope discrimination throughout the theoretical range for dark CO2 uptake during CAM. These ranged from the limit set by phosphoenolpyruvate carboxylase (PEPc) (-6[per mille (thousand) sign], [delta]13C equivalent of -2[per mille (thousand) sign]) to that imposed by diffusion limitation (+4[per mille (thousand) sign], [delta]13C equivalent of -12[per mille (thousand) sign]), but the lowest carbon-isotope discrimination occurred when P[square root]pa was only 0.7. Second, when the availability of external or internal sources of CO2 was reduced for both field- and greenhouse-grown plants, CO2 uptake by day via PEPc during phase II largely compensated. Third, by reducing the dark period, plants accumulated low levels of acidity, and CO2 uptake occurred throughout the subsequent light period. Discrimination switched from being dominated by PEPc (phase II) to ribulose 1,5-bisphosphate carboxylase/oxygenase (phase III), with both enzymes active during phase IV. Under natural conditions, photochemical stability is maintained by extended PEPc activity in phase II, which enhances acid accumulation and delays decarboxylation until temperature and light stress are maximal at midday.     

Crassulacean acid metabolism photosynthesis: `working the night shift'

Crassulacean acid metabolism (CAM) can be traced from Roman times through persons who noted a morning acid taste of some common house plants. From India in 1815, Benjamin-Heyne described a `daily acid taste cycle' with some succulent garden plants. Recent work has shown that the nocturnally formed acid is decarboxylated during the day to become the CO₂ for photosynthesis. Thus, CAM photosynthesis extends over a 24-hour day using several daily interlocking cycles. To understand CAM photosynthesis, several landmark discoveries were made at the following times: daily reciprocal acid and carbohydrate cycles were found during 1870 to 1887; their precise identification, as malic acid and starch, and accurate quantification occurred from 1940 to 1954; diffusive gas resistance methods were introduced in the early 1960s that led to understanding the powerful stomatal control of daily gas exchanges; C₄ photosynthesis in two different types of cells was discovered from 1965 to ∼1974 and the resultant information was used to elucidate the day and night portions of CAM photosynthesis in one cell; and exceptionally high internal green tissue CO₂ levels, 0.2 to 2.5%, upon the daytime decarboxylation of malic acid, were discovered in 1979. These discoveries then were combined with related information from C₃ and C₄ photosynthesis, carbon biochemistry, cellular anatomy, and ecological physiology. Therefore by ∼1980, CAM photosynthesis finally was rigorously outlined. In a nutshell, 24-hour CAM occurs by phosphoenol pyruvate (PEP) carboxylase fixing CO₂(HCO₃ ⁻) over the night to form malic acid that is stored in plant cell vacuoles. While stomata are tightly closed the following day, malic acid is decarboxylated releasing CO₂ for C₃ photosynthesis via ribulose bisphosphate carboxylase oxygenase (Rubisco). The CO₂ acceptor, PEP, is formed via glycolysis at night from starch or other stored carbohydrates and after decarboxylation the three carbons are restored each day. In mid to late afternoon the stomata can open and mostly C₃ photosynthesis occurs until darkness. CAM photo-synthesis can be both inducible and constitutive and is known in 33 families with an estimated 15 to 20 000 species. CAM plants express the most plastic and tenacious photosynthesis known in that they can switch photosynthesis pathways and they can live and conduct photosynthesis for years even in the virtual absence of external H₂O and CO₂, i.e., CAM tenaciously protects its photosynthesis from both H₂O and CO₂ stresses. This revised version was published online in August 2006 with corrections to the Cover Date.     

Crassulacean acid metabolism in the context of other carbon-concentrating mechanisms in freshwater plants: a review

Inorganic carbon can be in short supply in freshwater relative to that needed by freshwater plants for photosynthesis because of a large external transport limitation coupled with frequent depleted concentrations of CO(2) and elevated concentrations of O(2). Freshwater plants have evolved a host of avoidance, exploitation and amelioration strategies to cope with the low and variable supply of inorganic carbon in water. Avoidance strategies rely on the spatial variation in CO(2) concentrations within and among lakes. Exploitation strategies involve anatomical and morphological features that take advantage of sources of CO(2) outside of the water column such as the atmosphere or sediment. Amelioration strategies involve carbon-concentrating mechanisms based on uptake of bicarbonate, which is widespread, C(4)-fixation, which is infrequent, and crassulacean acid metabolism (CAM), which is of intermediate frequency. CAM enables aquatic plants to take up inorganic carbon in the night. Furthermore, daytime inorganic carbon uptake is generally not inhibited and therefore CAM is considered to be a carbon-conserving mechanism. CAM in aquatic plants is a plastic mechanism regulated by environmental variables and is generally downregulated when inorganic carbon does not limit photosynthesis. CAM is regulated in the long term (acclimation during growth), but is also affected by environmental conditions in the short term (response on a daily basis). In aquatic plants, CAM appears to be an ecologically important mechanism for increasing inorganic carbon uptake, because the in situ contribution from CAM to the C-budget generally is high (18-55%).     

Competing carboxylases: circadian and metabolic regulation of Rubisco in C3 and CAM Mesembryanthemum crystallinum L

The temporal co-ordination of ribulose 1·5-bisphosphate carboxylase/oxygenase (Rubisco) and phosphoenolpyruvate carboxylase (PEPc) activities by Mesembryanthemum crystallinum L. in C(3) and crassulacean acid metabolism (CAM) modes was investigated under conventional light-dark (LD) and continuous light (LL) conditions. When C(3) , net CO(2) assimilation rate increased during each subjective night under LL with maximum carboxylation unrelated to Rubisco activation state. The CAM circadian rhythm of CO(2) uptake was more pronounced, with CO(2) assimilation rate maximal towards the end of each subjective night. In vivo and in vitro techniques were integrated to map carboxylase enzyme regulation to the framework provided by CAM LL gas exchange activity. Rubisco was activated in vitro throughout each subjective dark period and consistently deactivated at each subjective dawn, similar to that observed at true dawn in constitutive CAM species. Instantaneous carbon isotope discrimination showed in vivo carboxylase co-dominance during the CAM subjective night, initially by Rubisco and latterly C(4) (PEPc), despite both enzymes seemingly activated in vitro. The circadian rhythm in titratable acidity accumulation was progressively damped over successive subjective nights, but maintenance of PEPc carboxylation capacity ensures that CAM plants do not become progressively more 'C(3) -like' with time under LL.