Crassulacean Acid Metabolism Expression Research Articles

Implications of leaf ontogeny on drought-induced gradients of CAM expression and ABA levels in rosettes of the epiphytic tank bromeliad Guzmania monostachia

Guzmania monostachia is an epiphytic heteroblastic bromeliad that exhibits rosette leaves forming water-holding tanks at maturity. Different portions along its leaf blades can display variable degrees of crassulacean acid metabolism (CAM) up-regulation under drought. Since abscisic acid (ABA) can act as an important long-distance signal, we conducted a joint investigation of ontogenetic and drought impacts on CAM intensity and ABA levels in different leaf groups within the G. monostachia rosette. For this, three groups of leaves were analysed according to their position within the mature-tank rosette (i.e., younger, intermediate, and older leaves) to characterize the general growth patterns and magnitude of drought-modulated CAM expression. CAM activity was evaluated by analysing key molecules in the biochemical machinery of this photosynthetic pathway, while endogenous ABA content was comparatively measured in different portions of each leaf group after seven days under well-watered (control) or drought treatment. The results revealed that G. monostachia shows more uniform morphological characteristics along the leaves when in the atmospheric stage. The drought treatment of mature-tank rosettes generally induced in older leaves a more severe water loss, followed by the lowest CAM activity and a higher increase in ABA levels, while younger leaves showed an opposite response. Therefore, leaf groups at distinct ontogenetic stages within the tank rosette of G. monostachia responded to drought with variable degrees of water loss and CAM expression. ABA seems to participate in this tissue-compartmented response as a long-distance signalling molecule, transmitting the drought-induced signals originated in older leaves towards the younger ones.

Effects of exogenously applied hydrogen peroxide on antioxidant and osmoprotectant profiles and the C3-CAM shift in the halophyte Mesembryanthemum crystallinum L.

Exogenously applied H2O2 (50, 100 and 200mM) to Mesembryanthemum crystallinum root medium induced a transition from C3 to Crassulacean Acid Metabolism (CAM), as evaluated by diurnal malate (Δmal) fluctuations. A very high concentration of H2O2 (400mM) reduced Δmal below the value measured in control plants. An increase of malate content during the night in 400mM H2O2-treated plants might suggest that malate decarboxylation is crucial for CAM functioning. We conclude that malate plays a dual role: i) a protective and signaling function before CAM expression, and ii) a storage form of CO2 in plants performing CAM. A slight stimulation of photosystem II (PSII) photochemistry and net photosynthesis observed during the C3-CAM shift indicated that neither photoinhibition nor reduction of the photosynthetic rate were prerequisites for CAM. Moreover, CAM induction corresponded to a decrease of catalase activity. In CAM-performing plants, α-tocopherol, polyamines (putrescine and spermidine) and proline showed daily alterations and the content of α-tocopherol and polyamines was lower at the end of the day. In contrast, the proline concentration correlated with the applied H2O2 concentration and was higher at the end of the day in treated plants. The dynamic changes of antioxidant and osmolyte levels suggest their active role in preventing oxidative damage, stress acclimation mechanisms and involvement in metabolic regulation and/or signal transduction cascades.

Plasticity of crassulacean acid metabolism at subtropical latitudes: a pineapple case study.

Plants with the crassulacean acid metabolism (CAM) express high-metabolic plasticity, to adjust to environmental stresses. This article hypothesizes that irradiance and nocturnal temperatures are the major limitations for CAM at higher latitudes such as the Azores (37°45'N). Circadian CAM expression in Ananas comosus L. Merr. (pineapple) was assessed by the diurnal pattern of leaf carbon fixation into l-malate at the solstices and equinoxes, and confirmed by determining maximal phosphoenolpyruvate carboxylase (PEPC) activity in plant material. Metabolic adjustments to environmental conditions were confirmed by gas exchange measurements, and integrated with environmental data to determine CAM's limiting factors: light and temperature. CAM plasticity was observed at the equinoxes, under similar photoperiods, but different environmental conditions. In spring, CAM expression was similar between vegetative and flowering plants, while in autumn, flowering (before anthesis) and fructifying (with fully developed fruit before ripening) plants accumulated more l-malate. Below 100 µmol m(-2) s(-1) , CAM phase I was extended, reducing CAM phase III during the day. Carbon fixation inhibition may occur by two major pathways: nocturnal temperature (<15°C) inhibiting PEPC activity and l-malate accumulation; and low irradiance influencing the interplay between CAM phase I and III, affecting carboxylation and decarboxylation. Both have important consequences for plant development in autumn and winter. Observations were confirmed by flowering time prediction using environmental data, emphasizing that CAM expression had a strong seasonal regulation due to a complex network response to light and temperature, allowing pineapple to survive in environments not suitable for high productivity.

What is the potential for dark CO2 fixation in the facultative crassulacean acid metabolism species Talinum triangulare?

In obligate Crassulacean acid metabolism (CAM) plants, dark CO2 fixation is almost the sole route of CO2 fixation and, under drought, continues for long periods. In contrast, in plants of the facultative CAM species Talinum triangulare under experimental drought, dark CO2 fixation provides a small proportion of the daily assimilation observed in watered plants and occurs only for a few days, after which almost nil CO2 fixation is observed. Under field conditions, with a practically unlimited substrate volume, drought-induced CAM might operate for a longer period and make a higher contribution to daily CO2 fixation. Greenhouse-grown plants of T. triangulare were subjected to low and nearly constant soil water content; the operation of CAM was assessed through the measurement of nocturnal proton accumulation and dark CO2 fixation. Dark CO2 fixation appeared 19d after the onset of drought; its contribution during three months of experiment to daily CO2 assimilation ranged from 0.5 to 30.7% with a mean of 13.5%. Twenty days after the beginning of treatment, nocturnal proton accumulation increased six times and remained high for over three months. In spite of low soil water content, leaves did not engage in dark CO2 fixation all the time but dark CO2 fixation was large enough to produce an increase in relative 13C composition of mature leaves compared to watered plants but not to the value in short-term drought experiments. Leaf anatomical characteristics may guarantee the achievement of higher rates of dark CO2 fixation but results evidence the occurrence of a limit to the expression of CAM that remains to be determined.

Multiple isoforms of phosphoenolpyruvate carboxylase in the Orchidaceae (subtribe Oncidiinae): implications for the evolution of crassulacean acid metabolism.

Phosphoenolpyruvate carboxylase (PEPC) catalyses the initial fixation of atmospheric CO2 into oxaloacetate and subsequently malate. Nocturnal accumulation of malic acid within the vacuole of photosynthetic cells is a typical feature of plants that perform crassulacean acid metabolism (CAM). PEPC is a ubiquitous plant enzyme encoded by a small gene family, and each member encodes an isoform with specialized function. CAM-specific PEPC isoforms probably evolved from ancestral non-photosynthetic isoforms by gene duplication events and subsequent acquisition of transcriptional control elements that mediate increased leaf-specific or photosynthetic-tissue-specific mRNA expression. To understand the patterns of functional diversification related to the expression of CAM, ppc gene families and photosynthetic patterns were characterized in 11 closely related orchid species from the subtribe Oncidiinae with a range of photosynthetic pathways from C3 photosynthesis (Oncidium cheirophorum, Oncidium maduroi, Rossioglossum krameri, and Oncidium sotoanum) to weak CAM (Oncidium panamense, Oncidium sphacelatum, Gomesa flexuosa and Rossioglossum insleayi) and strong CAM (Rossioglossum ampliatum, Trichocentrum nanum, and Trichocentrum carthagenense). Phylogenetic analysis revealed the existence of two main ppc lineages in flowering plants, two main ppc lineages within the eudicots, and three ppc lineages within the Orchidaceae. Our results indicate that ppc gene family expansion within the Orchidaceae is likely to be the result of gene duplication events followed by adaptive sequence divergence. CAM-associated PEPC isoforms in the Orchidaceae probably evolved from several independent origins.

The evolution of autotrophy in relation to phosphorus requirement

The evolution of autotrophy is considered in relation to the availability of phosphorus (P), the ultimate elemental resource limiting biological productivity through Earth's history. Work on microbes and plants is emphasized, dealing in turn with the main uses for P in cells, namely nucleic acids, phospholipids, and water-soluble low molecular mass phosphate esters plus metabolically active inorganic orthophosphate. There is a greater minimum gene number and minimum DNA content in autotrophic than in osmochemoorganotrophic archaea and bacteria, as well as a lower rate of biomass increase per unit P (P-use efficiency) in autotrophs than in osmochemoorganotrophs, in eukaryotes as well as bacteria. This may be due to the diversion of rRNA from producing proteins common to all organisms to producing highly expressed proteins specific to autotrophs. The P requirement for phospholipids is decreased in oxygenic photolithotrophs, and some anoxygenic photolithotrophs, by substituting galactolipids and sulpholipids for phospholipids in the photosynthetic, and some other, membranes. The six different autotrophic inorganic carbon assimilation pathways have varying requirements for low molecular mass water-soluble phosphate esters. In oxygenic photolithotrophs, there is no clear evidence of a different P requirement for growth in the absence (diffusive CO2 entry) relative to the presence of CO2-concentrating mechanisms (CCMs). P limitation increases the expression of crassulacean acid metabolism (CAM) in facultative CAM plants, decreases the extent of inorganic carbon accumulation in algae with CCMs, and (usually) their inorganic carbon affinity and the water-use efficiency of growth of terrestrial plants, and the light-use efficiency of photolithotrophs.

A system dynamics model integrating physiology and biochemical regulation predicts extent of crassulacean acid metabolism (CAM) phases

A system dynamics (SD) approach was taken to model crassulacean acid metabolism (CAM) expression from measured biochemical and physiological constants. SD emphasizes state-dependent feedback interaction to describe the emergent properties of a complex system. These mechanisms maintain biological systems with homeostatic limits on a temporal basis. Previous empirical studies on CAM have correlated biological constants (e.g. enzyme kinetic parameters) with expression over the CAM diel cycle. The SD model integrates these constants within the architecture of the CAM 'system'. This allowed quantitative causal connections to be established between biological inputs and the four distinct phases of CAM delineated by gas exchange and malic acid accumulation traits. Regulation at flow junctions (e.g. stomatal and mesophyll conductance, and malic acid transport across the tonoplast) that are subject to feedback control (e.g. stomatal aperture, malic acid inhibition of phosphoenolpyruvate carboxylase, and enzyme kinetics) was simulated. Simulated expression for the leaf-succulent Kalanchoë daigremontiana and more succulent tissues of Agave tequilana showed strong correlation with measured gas exchange and malic acid accumulation (R(2) =0.912 and 0.937, respectively, for K. daigremontiana and R(2) =0.928 and 0.942, respectively, for A. tequilana). Sensitivity analyses were conducted to quantitatively identify determinants of diel CO2 uptake. The transition in CAM expression from low to high volume/area tissues (elimination of phase II-IV carbon-uptake signatures) was achieved largely by the manipulation three input parameters.

Spatial patterns of photosynthesis in thin- and thick-leaved epiphytic orchids: unravelling C3-CAM plasticity in an organ-compartmented way.

A positive correlation between tissue thickness and crassulacean acid metabolism (CAM) expression has been frequently suggested. Therefore, this study addressed the question of whether water availability modulates photosynthetic plasticity in different organs of two epiphytic orchids with distinct leaf thickness. Tissue morphology and photosynthetic mode (C3 and/or CAM) were examined in leaves, pseudobulbs and roots of a thick-leaved (Cattleya walkeriana) and a thin-leaved (Oncidium 'Aloha') epiphytic orchid. Morphological features were studied comparing the drought-induced physiological responses observed in each organ after 30 d of either drought or well-watered treatments. Cattleya walkeriana, which is considered a constitutive CAM orchid, displayed a clear drought-induced up-regulation of CAM in its thick leaves but not in its non-leaf organs (pseudobulbs and roots). The set of morphological traits of Cattleya leaves suggested the drought-inducible CAM up-regulation as a possible mechanism of increasing water-use efficiency and carbon economy. Conversely, although belonging to an orchid genus classically considered as performing C3 photosynthesis, Oncidium 'Aloha' under drought seemed to express facultative CAM in its roots and pseudobulbs but not in its leaves, indicating that such photosynthetic responses might compensate for the lack of capacity to perform CAM in its thin leaves. Morphological features of Oncidium leaves also indicated lower efficiency in preventing water and CO2 losses, while aerenchyma ducts connecting pseudobulbs and leaves suggested a compartmentalized mechanism of nighttime carboxylation via phosphoenolpyruvate carboxylase (PEPC) (pseudobulbs) and daytime carboxylation via Rubisco (leaves) in drought-exposed Oncidium plants. Water availability modulated CAM expression in an organ-compartmented manner in both orchids studied. As distinct regions of the same orchid could perform different photosynthetic pathways and variable degrees of CAM expression depending on the water availability, more attention should be addressed to this in future studies concerning the abundance of CAM plants.

Spatial division of phosphoenolpyruvate carboxylase and nitrate reductase activity and its regulation by cytokinins in CAM-induced leaves of Guzmania monostachia (Bromeliaceae)

Crassulacean acid metabolism (CAM) is a physiological adaptation of plants that live in stress environment conditions. A good model of CAM modulation is the epiphytic bromeliad, Guzmania monostachia, which switches between two photosynthetic pathways (C3–CAM) in response to different environmental conditions, such as light stress and water availability. Along the leaf length a gradient of acidity can be observed when G. monostachia plants are kept under water deficiency. Previous studies showed that the apical portions of the leaves present higher expression of CAM, while the basal regions exhibit lower expression of this photosynthetic pathway. The present study has demonstrated that it is possible to induce the CAM pathway in detached leaves of G. monostachia kept under water deficit for 7 d. Also, it was evaluated whether CAM expression can be modulated in detached leaves of Guzmania and whether some spatial separation between NO3− reduction and CO2 fixation occurs in basal and apical portions of the leaf. In addition, we analyzed the involvement of endogenous cytokinins (free and ribosylated forms) as possible signal modulating both NO3− reduction and CO2 fixation along the leaf blade of this bromeliad. Besides demonstrating a clear spatial and functional separation of carbon and nitrogen metabolism along G. monostachia leaves, the results obtained also indicated a probable negative correlation between endogenous free cytokinins – zeatin (Z) and isopentenyladenine (iP) – concentration and PEPC activity in the apical portions of G. monostachia leaves kept under water deficit. On the other hand, a possible positive correlation between endogenous Z and iP levels and NR activity in basal portions of drought-exposed and control leaves was verified. Together with the observations presented above, results obtained with exogenous cytokinins treatments, strongly suggest that free cytokinins might act as a stimulatory signal involved in NR activity regulation and as a negative regulator of PEPC activity in CAM-induced leaves of G. monostachia during a diel cycle.

Abscisic acid and nitric oxide signaling in two different portions of detached leaves of Guzmania monostachia with CAM up-regulated by drought

Guzmania monostachia is an epiphyte tank bromeliad capable of up-regulating crassulacean acid metabolism (CAM) in response to several environmental stimuli, including drought and light stress. In other plant species, abscisic acid (ABA) and nitric oxide (NO) seem to be involved in CAM induction. Because the leaves of tank bromeliads perform different functions along their length, this study attempted to investigate whether ABA and NO are involved in regulation of CAM expression in this species by quantifying these compounds in apical and basal portions of the leaf, and whether there would be differences in this event for each leaf portion. Detached leaves exposed to a 30% polyethylene glycol solution showed a significant upregulation of CAM on the seventh day of treatment only in the apical portion, as indicated by nocturnal acid accumulation and phosphoenolpyruvate carboxylase (PEPC) activity. On the three days prior to CAM induction, ABA, NO and H2O2 were quantified. The amounts of ABA were higher in PEG-exposed leaves, along their entire length. NO, however, was higher only in the apical portion, precisely where CAM was up-regulated. H2O2 was higher only in the basal portion of PEG-exposed leaves. Our results suggest that ABA might be a systemic signal to drought, occurring in the entire leaf. NO and H2O2, however, may be signals restricted only to the apical or basal portions, respectively.

Induction and reversal of crassulacean acid metabolism in Calandrinia polyandra: effects of soil moisture and nutrients.

Calandrinia polyandra Benth. (Montiaceae), an annual succulent herb endemic to Australia, is an exemplary facultative crassulacean acid metabolism (CAM) plant as demonstrated by continuous whole-plant lifetime CO2 exchange measurements under controlled conditions in the laboratory. Reduced soil water availability induced a shift from solely daytime CO2 fixation to dark CO2 fixation. The shift from C3 photosynthesis to CAM was reversible either upon rewatering alone, or upon a combination of rewatering and addition of nutrients. These observations highlight the role of edaphic conditions in controlling CAM expression in a plant that has the option of fixing CO2 either during the day or during the night, providing further evidence that this extreme form of photosynthetic plasticity is primarily controlled by the environment rather than plant ontogeny. The stimulating effect of soil nutrients on CO2 fixation in the light and its negative effect on dark CO2 fixation have not been described previously and deserve further attention. In the most widely used CAM model system, the halophytic Mesembryanthemum crystallinum L., CAM is typically induced by high salinity, and some metabolic responses may be CAM-unrelated and related to salt stress per se. C. polyandra could be an excellent complementary system for studying the biochemical and molecular foundations of CAM because drought stress elicits a complete C3 to CAM transition.

Environmental regulation of carbon isotope composition and crassulacean acid metabolism in three plant communities along a water availability gradient.

Expression of crassulacean acid metabolism (CAM) is characterized by extreme variability within and between taxa and its sensitivity to environmental variation. In this study, we determined seasonal fluctuations in CAM photosynthesis with measurements of nocturnal tissue acidification and carbon isotopic composition (δ13C) of bulk tissue and extracted sugars in three plant communities along a precipitation gradient (500, 700, and 1,000 mm year−1) on the Yucatan Peninsula. We also related the degree of CAM to light habitat and relative abundance of species in the three sites. For all species, the greatest tissue acid accumulation occurred during the rainy season. In the 500 mm site, tissue acidification was greater for the species growing at 30% of daily total photon flux density (PFD) than species growing at 80% PFD. Whereas in the two wetter sites, the species growing at 80% total PFD had greater tissue acidification. All species had values of bulk tissue δ13C less negative than −20‰, indicating strong CAM activity. The bulk tissue δ13C values in plants from the 500 mm site were 2‰ less negative than in plants from the wetter sites, and the only species growing in the three communities, Acanthocereus tetragonus (Cactaceae), showed a significant negative relationship between both bulk tissue and sugar δ13C values and annual rainfall, consistent with greater CO2 assimilation through the CAM pathway with decreasing water availability. Overall, variation in the use of CAM photosynthesis was related to water and light availability and CAM appeared to be more ecologically important in the tropical dry forests than in the coastal dune.

Nitric oxide mediates the hormonal control of Crassulacean acid metabolism expression in young pineapple plants.

Genotypic, developmental, and environmental factors converge to determine the degree of Crassulacean acid metabolism (CAM) expression. To characterize the signaling events controlling CAM expression in young pineapple (Ananas comosus) plants, this photosynthetic pathway was modulated through manipulations in water availability. Rapid, intense, and completely reversible up-regulation in CAM expression was triggered by water deficit, as indicated by the rise in nocturnal malate accumulation and in the expression and activity of important CAM enzymes. During both up- and down-regulation of CAM, the degree of CAM expression was positively and negatively correlated with the endogenous levels of abscisic acid (ABA) and cytokinins, respectively. When exogenously applied, ABA stimulated and cytokinins repressed the expression of CAM. However, inhibition of water deficit-induced ABA accumulation did not block the up-regulation of CAM, suggesting that a parallel, non-ABA-dependent signaling route was also operating. Moreover, strong evidence revealed that nitric oxide (NO) may fulfill an important role during CAM signaling. Up-regulation of CAM was clearly observed in NO-treated plants, and a conspicuous temporal and spatial correlation was also evident between NO production and CAM expression. Removal of NO from the tissues either by adding NO scavenger or by inhibiting NO production significantly impaired ABA-induced up-regulation of CAM, indicating that NO likely acts as a key downstream component in the ABA-dependent signaling pathway. Finally, tungstate or glutamine inhibition of the NO-generating enzyme nitrate reductase completely blocked NO production during ABA-induced up-regulation of CAM, characterizing this enzyme as responsible for NO synthesis during CAM signaling in pineapple plants.

Canopy CO2 exchange of two neotropical tree species exhibiting constitutive and facultative CAM photosynthesis, Clusia rosea and Clusia cylindrica.

Photon flux density (PFD) and water availability, the daily and seasonal factors that vary most in tropical environments, were examined to see how they influenced expression of crassulacean acid metabolism (CAM) in 3-year-old Clusia shrubs native to Panama. Instead of the commonly used single-leaf approach, diel CO2 exchange was measured for whole individual canopies of plants in large soil containers inside a naturally illuminated 8.8 m3 chamber. In well-watered C. rosea, a mainly constitutive CAM species, nocturnally fixed CO2 contributed about 50% to 24 h carbon gain on sunny days but the contribution decreased to zero following overcast days. Nonetheless, CO2 fixation in the light responded in such a way that 24 h carbon gain was largely conserved across the range of daily PFDs. The response of C. rosea to drought was similarly buffered. A facultative component of CAM expression led to reversible increases in nocturnal carbon gain that offset drought-induced reductions of CO2 fixation in the light. Clusia cylindrica was a C3 plant when well-watered but exhibited CAM when subjected to water stress. The induction of CAM was fully reversible upon rewatering. C. cylindrica joins C. pratensis as the most unambiguous facultative CAM species reported in the genus Clusia.

Carbon isotope composition and mode of photosynthesis in Clusia species from Mexico

The neotropical genus Clusia comprises arborescent species exhibiting Crassulacean Acid Metabolism (CAM) as was first reported for a Mexican species, Clusia lundellii. Here, the occurrence of CAM photosynthesis was studied in 20 species of Clusia, 18 from Mexico, and 2 from Guatemala, using leaf carbon isotopic composition. In most species, samples from individuals collected in different locations were analyzed. CAM was present in at least 11 species, eight of which contained specimens with δ13C values less negative than -20.0 ‰, indicating strong CAM (C. chanekiana, C. flava, C. lundellii, C. mexicana, C. quadrangula, C. rosea, C. suborbicularis, and C. tetra-trianthera). δ13C was highly variable in some species, but CAM expression was not correlated to life form (epiphytic, hemiepiphytic, terrestrial) or habitat. CAM specimens were not collected at altitudes above 1 700 m a.s.l.

Impact of developmental stage on CAM expression and growth in an Aechmea hybrid under greenhouse conditions

SummaryBesides environmental factors, expression of crassulacean acid metabolism (CAM) and the related production of malic acid may depend on plant developmental stage. This topic has not yet been investigated under commercial greenhouse conditions. In CAM plant cultivation, malic acid is a central determinant of plant growth, but can also cause physiological leaf damage problems. Here, we present data on diurnal leaf malic acid contents, relative water content dynamics, and measurements of growth at four stages of the plant growth cycle (i.e., ex vitro-acclimated plantlets, 6-month, 12-month, and 18-month-old plants) of Aechmea ‘Maya’. The results obtained showed that Aechmea ‘Maya’ was an obligate CAM plant at all developmental stages during cultivation. Nevertheless, there was some developmental control on the expression of CAM. Under the same environmental conditions, 6-month-old plants accumulated significantly higher amounts of malic acid in their leaves, thus supporting higher growth rates, than in the other growth stages. This larger malic acid pool implied a higher risk of physiological leaf damage, but since remobilisation of malic acid in the early morning remained unaffected during growth, physiological leaf damage might occur throughout vegetative growth.

The occurrence of crassulacean acid metabolism in Cymbidium (Orchidaceae) and its ecological and evolutionary implications

Crassulacean acid metabolism (CAM) is one of the photosynthetic pathways regarded as adaptations to water stress in land plants. Little is known about correlations among the level of CAM activity, environment of habitat, life form, and phylogenetic relationship of a plant group from an evolutionary perspective. We examined these relationships in 18 species of Cymbidium (Orchidaceae) because the genus shows distinctive diversification of habitats and life forms. The photosynthetic type was classed into three categories, strong CAM, weak CAM, and C(3) on the basis of CAM activity. CAM expression in Cymbidium was confined to the epiphytic and lithophytic species. Especially, all of these species from tropical to subtropical rainforest exhibited CAM activity. On the other hand, the terrestrial species always exhibited C(3) metabolism irrespective of their varied habitats. Regarding the evolution of photosynthetic characters, weak CAM was the ancestral state in Cymbidium and strong CAM and C(3) metabolism occurred subsequently. The evolution of strong CAM likely enabled Cymbidium to extend to exposed sites in tropical lowland where marked water stress exists. Further, different levels of CAM activity characterized each species and such potential plasticity of CAM may realize the radiation of Cymbidium into sites with different environmental conditions.

Diurnal patterns of carbon dioxide, water vapour, and energy fluxes in pineapple [Ananas comosus (L.) Merr. cv. Red Spanish] field using eddy covariance

We analyzed the eddy covariance measurements of momentum, mass, and energy taken daily throughout five consecutive seasonal courses (i.e. 840 d after planting) of a pineapple [Ananas comosus (L.) Merr. cv. Red Spanish] field growing in the Orinoco lowlands. This field provides an opportunity for micrometeorological studies because of the flat and windy site; the seasonal weather including ENSO effects and the Crassulacean Acid Metabolism (CAM) physiology of the crop were additional attributes. Soil CO2 flux was quantified and added to the net ecosystem exchange in order to obtain the canopy flux (FC). The canopy CO2 flux partially followed the four phases of CAM sensu Osmond (1978). The daily pattern of gaseous exchange in pineapple showed a continuum spectrum in which a major proportion of CO2 uptake occurring during the daytime was common and in which the CAM expression was related to day and nocturnal CO2 uptake. However, the benefits of CO2 uptake at low water cost were constrained by the limited nocturnal CO2 uptake. Seasonal and ontogenetic changes affected the energy exchange as well as the partitioning of available energy into sensible (QH) and latent (QLE) heat. When the hourly net radiation (QRn) reached its maximum value, latent heat flux (QLE) to available energy throughout the vegetative and reproductive stages was 0.65, 0.05, 0.30, 0.11, and 0.33 for the 1997 wet season, 1997/98 dry season, 1998 wet season, 1998/99 dry season, and 1999 wet season, respectively. Throughout the growth period, we found the pivotal role of surface conductance (g S) in both QLE and FC. Furthermore, the canopy responded to environmental changes. During the wet seasons the g S was strongly influenced by humidity mole fraction deficit and was usually lower than aerodynamic conductance, whereas during the dry seasons, soil water deficit limited evapotranspiration and production rates. For the fully canopy cover, the hourly trend of marginal water cost of pineapple carbon gain in the dry seasons indicated that g S became sufficiently efficient to reduce the amount of water transported per unit of carbon gain. In the wet season, the coupling of CO2 uptake and stomatal conductance was more effective in maintaining a higher proportionality between QLE and g S.

Functional constraints of CAM leaf anatomy: tight cell packing is associated with increased CAM function across a gradient of CAM expression

Increased cell size, increased leaf succulence, reduced intercellular air space (IAS), and reduced surface of mesophyll exposed to IAS (L(mes)/area) are traits associated with the Crassulacean acid metabolism (CAM) photosynthetic pathway. An examination was carried out to determine whether these anatomical and structural traits are related to the degree of CAM function in eight CAM species, as measured by CO(2) assimilation during the CAM and C(3) phases. Increased cell size and leaf succulence were not closely related to the degree of CAM function, indicating that succulence beyond a certain threshold does not enhance CAM function. Reduced IAS and L(mes)/area were positively related to CAM function, and negatively related to C(3) function. These results indicate that reduced IAS and L(mes)/area are beneficial for CAM function through the reduction of CO(2) efflux and the improvement of carbon economy. However, reduced IAS and L(mes)/area limit C(3) photosynthesis, potentially mediating a bimodal distribution of weak and strong CAM species with high and low IAS and L(mes)/area values, respectively.

Morpho-anatomical changes and photosynthetic metabolism of Stenocereus beneckei seedlings under soil water deficit

Characteristics developed by Cactaceae for adaptation to climates where water is limited include crassulacean acid metabolism (CAM), a thick cuticle, and spines and trichomes that intercept a proportion of solar radiation. A few studies consider morpho-anatomical and physiological characteristics of Cactaceae seedlings, which may help understand their establishment, growth, and eventual reproduction. In this study, photosynthetic metabolism (titratable protons) and morpho-anatomical features of Stenocereus beneckei seedlings were examined under limiting water conditions. Soil moisture treatments consisted of -0.03, -0.5, -1.5, and -3.0 MPa, and seedling samples were taken at 3 h intervals on one day at 7 and 9 months of age with three replicates per treatment. The results show irregular fluctuations in acidity concentrations during the first 6 and 7 months of age; at 9 months, an increase in titratable proton values was observed during the night, and it seems that soil moisture does not determine CAM expression. Seedlings from smaller seeds are less tolerant to water stress, they had poor growth in all treatments, and at -3.0 MPa after 3 months of drought none survived. Anatomical observations show collapsed cells associated with a high accumulation of calcium oxalate crystals and starch grains, as a response to water deficit. Titratable acidity concentration increased with seedling age, and CAM expression did not accelerate with soil water deficit.