The stomatal response to vapor pressure deficit drives the apparent temperature response of photosynthesis in tropical forests.

As temperature rises, net carbon uptake in tropical forests decreases, but the underlying mechanisms are not well understood. High temperatures can limit photosynthesis directly, for example by reducing biochemical capacity, or indirectly through rising vapor pressure deficit (VPD) causing stomatal closure. To explore the independent effects of temperature and VPD on photosynthesis we analyzed photosynthesis data from the upper canopies of two tropical forests in Panama with Generalized Additive Models. Stomatal conductance and photosynthesis consistently decreased with increasing VPD, and statistically accounting for VPD increased the optimum temperature of photosynthesis (Topt) of trees from a VPD-confounded apparent Topt of c. 30-31°C to a VPD-independent Topt of c. 33-36°C, while for lianas no VPD-independent Topt was reached within the measured temperature range. Trees and lianas exhibited similar temperature and VPD responses in both forests, despite 1500 mm difference in mean annual rainfall. Over ecologically relevant temperature ranges, photosynthesis in tropical forests is largely limited by indirect effects of warming, through changes in VPD, not by direct warming effects of photosynthetic biochemistry. Failing to account for VPD when determining Topt misattributes the underlying causal mechanism and thereby hinders the advancement of mechanistic understanding of global warming effects on tropical forest carbon dynamics.

A medida que aumenta la temperatura, disminuye la absorción neta de carbono en los bosques tropicales, sin embargo, aún no se conocen bien los mecanismos que la subyacen. Las altas temperaturas pueden limitar la fotosíntesis directamente, por ejemplo, reduciendo la eficiencia de los procesos bioquímicos, pero también de forma indirecta a través del aumento del déficit de presión de vapor (DPV) que resulta en el cierre estomático. Para explorar los efectos independientes de la temperatura y el DPV en la fotosíntesis, analizamos datos de la absorción neta de carbono del dosel de dos bosques tropicales en Panamá utilizando modelos aditivos generalizados. La conductancia estomática y la fotosíntesis disminuyó consistentemente con el aumento de DPV, y considerando el DPV en modelas estadísticas, la temperatura óptima de la fotosíntesis (Topt) aumentó, de un Topt aparente influida por la DVP de c. 30­31°C a un Topt independiente del DPV de c. 33­36°C. Los árboles y las lianas mostraron respuestas similares a la temperatura y a la DVP en ambos bosques, a pesar de la diferencia de 1500 mm en la precipitación media anual. La fotosíntesis en los bosques tropicales está limitada en gran medida por los efectos indirectos del aumento de la temperatura, a través de cambios en el DPV y no por los efectos directos en los procesos bioquímicos. Si no se tiene en cuenta el DPV al determinar el Topt, se atribuye erróneamente el mecanismo causal subyacente y, por lo tanto, se obstaculiza el avance en la comprensión de los efectos del calentamiento global en la dinámica del carbono.

Shifting photosynthesis between the fast and slow lane: Facultative CAM and water-deficit stress.

Five decades ago, the first report of a shift from C3 to CAM (crassulacean acid metabolism) photosynthesis following the imposition of stress was published in this journal. The annual, Mesembryanthemum crystallinum (Aizoaceae), was shown to be a C3 plant when grown under non-saline conditions, and a CAM plant when exposed to high soil salinity. This observation of environmentally triggered CAM eventually led to the introduction of the term facultative CAM, which categorises CAM that is induced or upregulated in response to water-deficit stress and is lost or downregulated when the stress is removed. Reversibility of C3-to-CAM shifts distinguishes stress-driven facultative-CAM responses from purely ontogenetic increases of CAM activity. We briefly review how the understanding of facultative CAM has developed, evaluate the current state of knowledge, and highlight questions of continuing interest. We demonstrate that the long-lived leaves of a perennial facultative-CAM arborescent species, Clusia pratensis, can repeatedly switch between C3 and CAM in response to multiple wet-dry-wet cycles. Undoubtedly, this is a dedicated response to environment, independent of ontogeny. We highlight the potential for engineering facultative CAM into C3 crops to provide a flexible capacity for drought tolerance.

The CAM lineages of planet Earth.

BACKGROUND AND SCOPE: The growth of experimental studies of crassulacean acid metabolism (CAM) in diverse plant clades, coupled with recent advances in molecular systematics, presents an opportunity to re-assess the phylogenetic distribution and diversity of species capable of CAM. It has been more than two decades since the last comprehensive lists of CAM taxa were published, and an updated survey of the occurrence and distribution of CAM taxa is needed to facilitate and guide future CAM research. We aimed to survey the phylogenetic distribution of these taxa, their diverse morphology, physiology and ecology, and the likely number of evolutionary origins of CAM based on currently known lineages. RESULTS AND CONCLUSIONS: We found direct evidence (in the form of experimental or field observations of gas exchange, day-night fluctuations in organic acids, carbon isotope ratios and enzymatic activity) for CAM in 370 genera of vascular plants, representing 38 families. Further assumptions about the frequency of CAM species in CAM clades and the distribution of CAM in the Cactaceae and Crassulaceae bring the currently estimated number of CAM-capable species to nearly 7 % of all vascular plants. The phylogenetic distribution of these taxa suggests a minimum of 66 independent origins of CAM in vascular plants, possibly with dozens more. To achieve further insight into CAM origins, there is a need for more extensive and systematic surveys of previously unstudied lineages, particularly in living material to identify low-level CAM activity, and for denser sampling to increase phylogenetic resolution in CAM-evolving clades. This should allow further progress in understanding the functional significance of this pathway by integration with studies on the evolution and genomics of CAM in its many forms.

Forty years of research into crassulacean acid metabolism in the genus Clusia: anatomy, ecophysiology and evolution.

Clusia is the only genus containing dicotyledonous trees with a capacity to perform crassulacean acid metabolism (CAM). Since the discovery of CAM in Clusia 40 years ago, several studies have highlighted the extraordinary plasticity and diversity of life forms, morphology and photosynthetic physiology of this genus. In this review, we revisit aspects of CAM photosynthesis in Clusia and hypothesize about the timing, the environmental conditions and potential anatomical characteristics that led to the evolution of CAM in the group. We discuss the role of physiological plasticity in influencing species distribution and ecological amplitude in the group. We also explore patterns of allometry of leaf anatomical traits and their correlations with CAM activity. Finally, we identify opportunities for further research on CAM in Clusia, such as the role of elevated nocturnal accumulation of citric acid, and gene expression in C3-CAM intermediate phenotypes.

Brief reflections on 50 years as a plant ecophysiologist.

SCOPE: This paper is a short biographical sketch of my life as a plant ecophysiologist in which serendipity and outstanding collaborators have been key allies.

Plant water use theory should incorporate hypotheses about extreme environments, population ecology, and community ecology.

Plant water use theory has largely been developed within a plant-performance paradigm that conceptualizes water use in terms of value for carbon gain and that sits within a neoclassical economic framework. This theory works very well in many contexts but does not consider other values of water to plants that could impact their fitness. Here, we survey a range of alternative hypotheses for drivers of water use and stomatal regulation. These hypotheses are organized around relevance to extreme environments, population ecology, and community ecology. Most of these hypotheses are not yet empirically tested and some are controversial (e.g. requiring more agency and behavior than is commonly believed possible for plants). Some hypotheses, especially those focused around using water to avoid thermal stress, using water to promote reproduction instead of growth, and using water to hoard it, may be useful to incorporate into theory or to implement in Earth System Models.

Leaf vein density correlates with crassulacean acid metabolism, but not hydraulic capacitance, in the genus Clusia.

BACKGROUND AND AIMS: Many succulent species are characterized by the presence of Crassulacean acid metabolism (CAM) and/or elevated bulk hydraulic capacitance (CFT). Both CAM and elevated CFT substantially reduce the rate at which water moves through transpiring leaves. However, little is known about how these physiological adaptations are coordinated with leaf vascular architecture. METHODS: The genus Clusia contains species spanning the entire C3-CAM continuum, and also is known to have >5-fold interspecific variation in CFT. We used this highly diverse genus to explore how interspecific variation in leaf vein density is coordinated with CAM and CFT. KEY RESULTS: We found that constitutive CAM phenotypes were associated with lower vein length per leaf area (VLA) and vein termini density (VTD), compared to C3 or facultative CAM species. However, when vein densities were standardized by leaf thickness, this value was higher in CAM than C3 species, which is probably an adaptation to overcome apoplastic hydraulic resistance in deep chlorenchyma tissue. In contrast, CFT did not correlate with any xylem anatomical trait measured, suggesting CAM has a greater impact on leaf transpiration rates than CFT. CONCLUSIONS: Our findings strongly suggest that CAM photosynthesis is coordinated with leaf vein densities. The link between CAM and vascular anatomy will be important to consider when attempting to bioengineer CAM into C3 crops.

Dissecting succulence: Crassulacean acid metabolism and hydraulic capacitance are independent adaptations in Clusia leaves.

Succulence is found across the world as an adaptation to water-limited niches. The fleshy organs of succulent plants develop via enlarged photosynthetic chlorenchyma and/or achlorophyllous water storage hydrenchyma cells. The precise mechanism by which anatomical traits contribute to drought tolerance is unclear, as the effect of succulence is multifaceted. Large cells are believed to provide space for nocturnal storage of malic acid fixed by crassulacean acid metabolism (CAM), whilst also buffering water potentials by elevating hydraulic capacitance (CFT ). The effect of CAM and elevated CFT on growth and water conservation have not been compared, despite the assumption that these adaptations often occur together. We assessed the relationship between succulent anatomical adaptations, CAM, and CFT , across the genus Clusia. We also simulated the effects of CAM and CFT on growth and water conservation during drought using the Photo3 model. Within Clusia leaves, CAM and CFT are independent traits: CAM requires large palisade chlorenchyma cells, whereas hydrenchyma tissue governs interspecific differences in CFT . In addition, our model suggests that CAM supersedes CFT as a means to maximise CO2 assimilation and minimise transpiration during drought. Our study challenges the assumption that CAM and CFT are mutually dependent traits within succulent leaves.

Evidence for phylogenetic signal and correlated evolution in plant-water relation traits.

Evolutionary relationships are likely to play a significant role in shaping plant physiological and structural traits observed in contemporary taxa. We review research on phylogenetic signal and correlated evolution in plant-water relation traits, which play important roles in allowing plants to acquire, use, and conserve water. We found more evidence for a phylogenetic signal in structural traits (e.g. stomatal length and stomatal density) than in physiological traits (e.g. stomatal conductance and water potential at turgor loss). Although water potential at turgor loss is the most-studied plant-water relation trait in an evolutionary context, it is the only trait consistently found to not have a phylogenetic signal. Correlated evolution was common among traits related to water movement efficiency and hydraulic safety in both leaves and stems. We conclude that evidence for phylogenetic signal varies depending on: the methodology used for its determination, that is, model-based approaches to determine phylogenetic signal such as Blomberg's K or Pagel's λ vs statistical approaches such as ANOVAs with taxonomic classification as a factor; on the number of taxa studied (size of the phylogeny); and the setting in which plants grow (field vs common garden). More explicitly and consistently considering the role of evolutionary relationships in shaping plant ecophysiology could improve our understanding of how traits compare among species, how traits are coordinated with one another, and how traits vary with the environment.

Twelve-month efficacy of second-generation cryoballoon ablation for atrial fibrillation performed at community hospitals: results of the German register on cryoballoon ablation in local hospitals (regional).

BACKGROUND: The 12-month follow-up (F/U) efficacy of CBA PVI performed at community hospitals for treatment of symptomatic paroxysmal and persistent atrial fibrillation (AF) is unknown. This study determined the 12-month efficacy of pulmonary vein isolation (PVI) using cryoballoon ablation (CBA) performed at community hospitals with limited annual case numbers. METHODS: This registry study included 983 consecutive patients (pts) from 19 hospitals, each with an annual procedural volume of < 100 PVI procedures/year. Pts underwent CBA PVI for paroxysmal AF (n = 520), persistent AF (n = 423), or redo PVI (n = 40). The primary endpoint was frequency of documented recurrent AF, the occurrence of atrial flutter or tachycardia following a 90-day period after the index ablation and up to 12 months. The frequency of repeat ablation was determined. RESULTS: Isolation of all PVs was documented in 98% of pts at the end of the procedure. Twelve-month F/U data could be obtained in 916 pts. A 24-h ECG registration was performed in 641 pts (70.0%); in 107 pts (16.7%) of them, recurrent AF was documented. The primary endpoint was met in 193 F/U pts (21.1%). It occurred in 80/486 F/U pts with paroxysmal AF (16.4%), and in 107/390 F/U pts with persistent AF (27.4%). Redo PVI was performed in 71 pts (7.8%), and atrial flutter ablation was performed in 12 pts (1.4%). CONCLUSIONS: CBA PVI for paroxysmal or persistent AF can be performed at community hospitals with adequate rates of 12-month symptom freedom and arrhythmia recurrence. The study was registered at the German register of clinical studies (DRKS00016504).

CAM photosynthesis: the acid test.

There is currently considerable interest in the prospects for bioengineering crassulacean acid metabolism (CAM) photosynthesis - or key elements associated with it, such as increased water-use efficiency - into C3 plants. Resolving how CAM photosynthesis evolved from the ancestral C3 pathway could provide valuable insights into the targets for such bioengineering efforts. It has been proposed that the ability to accumulate organic acids at night may be common among C3 plants, and that the transition to CAM might simply require enhancement of pre-existing fluxes, without the need for changes in circadian or diurnal regulation. We show, in a survey encompassing 40 families of vascular plants, that nocturnal acidification is a feature entirely restricted to CAM species. Although many C3 species can synthesize malate during the light period, we argue that the switch to night-time malic acid accumulation requires a fundamental metabolic reprogramming that couples glycolytic breakdown of storage carbohydrate to the process of net dark CO2 fixation. This central element of the CAM pathway, even when expressed at a low level, represents a biochemical capability not seen in C3 plants, and so is better regarded as a discrete evolutionary innovation than as part of a metabolic continuum between C3 and CAM.

Corrigendum to: Does the C4 plant Trianthema portulacastrum (Aizoaceae) exhibit weakly expressed crassulacean acid metabolism (CAM)?

We examined whether crassulacean acid metabolism (CAM) is present in Trianthema portulacastrum L. (Aizoaceae), a pantropical, salt-tolerant C4 annual herb with atriplicoid-type Kranz anatomy in leaves but not in stems. The leaves of T. portulacastrum are slightly succulent and the stems are fleshy, similar to some species of Portulaca, the only genus known in which C4 and CAM co-occur. Low- level nocturnal acidification typical of weakly expressed, predominantly constitutive CAM was measured in plants grown for their entire life-cycle in an outdoor raised garden box. Acidification was greater in stems than in leaves. Plants showed net CO2 uptake only during the light irrespective of soil water availability. However, nocturnal traces of CO2 exchange exhibited curved kinetics of reduced CO2 loss during the middle of the night consistent with low-level CAM. Trianthema becomes the second genus of vascular land plants in which C4 and features of CAM have been demonstrated to co-occur in the same plant and the first C4 plant with CAM-type acidification described for the Aizoaceae. Traditionally the stems of herbs are not sampled in screening studies. Small herbs with mildly succulent leaves and fleshy stems might be a numerically significant component of CAM biodiversity.

Safety and acute efficacy of cryoballoon ablation for atrial fibrillation at community hospitals.

AIMS: Pulmonary vein isolation (PVI) using cryoballoon ablation (CBA) is an established procedure for treating symptomatic paroxysmal and persistent atrial fibrillation (AF). The safety and efficacy of PVI performed at community hospitals are unknown. We aimed to determine the safety and acute efficacy of PVI using CBA performed at community hospitals with limited annual case numbers. METHODS AND RESULTS: This registry study included 1004 consecutive patients who had PVI performed for symptomatic paroxysmal (n = 563) or persistent AF (n = 441) from January 2019 to September 2020 at 20 hospitals. Each hospital performed fewer than 100 CBA-PVI procedures/year according to local standards. Procedural data, efficacy, and complication rates were determined. The mean number of CBA procedures performed/year at each centre was 59 ± 25. The average procedure time was 90.1 ± 31.6 min and the average fluoroscopy time was 19.2 ± 11.4 min. Isolation of all pulmonary veins was documented in 97.9% of patients. The most frequent reason for not achieving complete isolation was development of phrenic nerve palsy. No hospital deaths were observed. Two patients (0.2%) suffered a clinical stroke. Pericardial effusion occurred in six patients (0.6%), two of whom (0.2%) required pericardial drainage. Vascular complications occurred in 24 patients (2.4%), two of whom (0.2%) required vascular surgery. Phrenic nerve palsy occurred in 48 patients (4.8%) and persisted up to hospital discharge in six patients (0.6%). CONCLUSION: Pulmonary vein isolation procedures for paroxysmal or persistent AF using CBA can be performed at community hospitals with high acute efficacy and low complication rates.

Large differences in leaf cuticle conductance and its temperature response among 24 tropical tree species from across a rainfall gradient.

More frequent droughts and rising temperatures pose serious threats to tropical forests. When stomata are closed under dry and hot conditions, plants lose water through leaf cuticles, but little is known about cuticle conductance (gmin ) of tropical trees, how it varies among species and environments, and how it is affected by temperature. We determined gmin in relation to temperature for 24 tropical tree species across a steep rainfall gradient in Panama, by recording leaf drying curves at different temperatures in the laboratory. In contrast with our hypotheses, gmin did not differ systematically across the rainfall gradient; species differences did not reflect phylogenetic patterns; and in most species gmin did not significantly increase between 25 and 50°C. gmin was higher in deciduous than in evergreen species, in species with leaf trichomes than in species without, in sun leaves than in shade leaves, and tended to decrease with increasing leaf mass per area across species. There was no relationship between stomatal and cuticle conductance. Large species differences in gmin and its temperature response suggest that more frequent hot droughts may lead to differential survival among tropical tree species, regardless of species' position on the rainfall gradient.

Diversity of CAM plant photosynthesis (crassulacean acid metabolism): a tribute to Barry Osmond.

This special issue is a tribute to the Australian plant biologist Professor Charles Barry Osmond - Fellow of the Australian Academy of Sciences, the Royal Society of London, and Leopoldina, the German National Academy of Sciences - and his many contributions to our understanding of the biochemistry and physiological ecology of CAM (crassulacean acid metabolism) photosynthesis. This water-conserving photosynthetic pathway is characterised by nocturnal uptake of atmospheric CO2 and typically enables succulent plants to perform and survive in warm semiarid terrestrial and epiphytic habitats. The idea for this issue is to mark the occasion of Barry's 80th birthday in 2019. The foreword highlights some of his outstanding contributions and introduces the research papers of the special issue.

Leaf water δ18O reflects water vapour exchange and uptake by C3 and CAM epiphytic bromeliads in Panama.

The distributions of CAM and C3 epiphytic bromeliads across an altitudinal gradient in western Panama were identified from carbon isotope (δ13C) signals, and epiphyte water balance was investigated via oxygen isotopes (δ18O) across wet and dry seasons. There were significant seasonal differences in leaf water (δ18Olw), precipitation, stored 'tank' water and water vapour. Values of δ18Olw were evaporatively enriched at low altitude in the dry season for the C3 epiphytes, associated with low relative humidity (RH) during the day. Crassulacean acid metabolism (CAM) δ18Olw values were relatively depleted, consistent with water vapour uptake during gas exchange under high RH at night. At high altitude, cloudforest locations, C3 δ18Olw also reflected water vapour uptake by day. A mesocosm experiment with Tillandsia fasciculata (CAM) and Werauhia sanguinolenta (C3) was combined with simulations using a non-steady-state oxygen isotope leaf water model. For both C3 and CAM bromeliads, δ18Olw became progressively depleted under saturating water vapour by day and night, although evaporative enrichment was restored in the C3 W. sanguinolenta under low humidity by day. Source water in the overlapping leaf base 'tank' was also modified by evaporative δ18O exchanges. The results demonstrate how stable isotopes in leaf water provide insights for atmospheric water vapour exchanges for both C3 and CAM systems.

Evolution of crassulacean acid metabolism (CAM) as an escape from ecological niche conservatism in Malagasy Bulbophyllum (Orchidaceae).

Despite growing evidence that niche shifts are more common in flowering plants than previously thought, little is known of whether such shifts are promoted by changes in photosynthetic pathways. Here we combine the most complete phylogeny for epiphytic Malagasy Bulbophyllum orchids (c. 210 spp.) with climatic niche and carbon isotope ratios to infer the group's spatial-temporal history, and the role of strongly expressed crassulacean acid metabolism (CAM) in facilitating niche shifts and diversification. We find that most extant species still retain niche (Central Highland) and photosynthesis (C3 ) states as present in the single mid-Miocene (c. 12.70 million yr ago (Ma)) ancestor colonizing Madagascar. However, we also infer a major transition to CAM, linked to a late Miocene (c. 7.36 Ma) invasion of species from the sub-humid highland first into the island's humid eastern coastal, and then into the seasonally dry 'Northwest Sambirano' rainforests, yet without significant effect on diversification rates. These findings indicate that CAM in tropical epiphytes may be selectively advantageous even in high rainfall habitats, rather than presenting a mere adaptation to dry environments or epiphytism per se. Overall, our study qualifies CAM as an evolutionary 'gateway' trait that considerably widened the spatial-ecological amplitude of Madagascar's most species-rich orchid genus.

Patient delay and benefit of timely reperfusion in ST-segment elevation myocardial infarction.

BACKGROUND: In patients with ST-segment elevation myocardial infarction (STEMI), it is unknown how patient delay modulates the beneficial effects of timely reperfusion. AIMS: To assess the prognostic significance of a contact-to-balloon time of less than 90 min on in-hospital mortality in different categories of symptom-onset-to-first-medical-contact (S2C) times. METHODS: A total of 20 005 consecutive patients from the Feedback Intervention and Treatment Times in ST-segment Elevation Myocardial Infarction (FITT-STEMI) programme treated with primary percutaneous coronary intervention (PCI) were included. RESULTS: There were 1554 deaths (7.8%) with a J-shaped relationship between mortality and S2C time. Mortality was 10.0% in patients presenting within 1 hour, and 4.9%, 6.0% and 7.3% in patient groups with longer S2C intervals of 1-2 hours, 2-6 hours and 6-24 hours, respectively. Patients with a short S2C interval of less than 1 hour (S2C<60 min) had the highest survival benefit from timely reperfusion with PCI within 90 min (OR 0.27, 95% CI 0.23 to 0.31, p<0.0001) as compared with the three groups with longer S2C intervals of 1 hour

Leaf heat tolerance of 147 tropical forest species varies with elevation and leaf functional traits, but not with phylogeny.

Exceeding thermal thresholds causes irreversible damage and ultimately loss of leaves. The lowland tropics are among the warmest forested biomes, but little is known about heat tolerance of tropical forest plants. We surveyed leaf heat tolerance of sun-exposed leaves from 147 tropical lowland and pre-montane forest species by determining the temperatures at which potential photosystem II efficiency based on chlorophyll a fluorescence started to decrease (TCrit ) and had decreased by 50% (T50 ). TCrit averaged 46.7°C (5th-95th percentile: 43.5°C-49.7°C) and T50 averaged 49.9°C (47.8°C-52.5°C). Heat tolerance partially adjusted to site temperature; TCrit and T50 decreased with elevation by 0.40°C and 0.26°C per 100 m, respectively, while mean annual temperature decreased by 0.63°C per 100 m. The phylogenetic signal in heat tolerance was weak, suggesting that heat tolerance is more strongly controlled by environment than by evolutionary legacies. TCrit increased with the estimated thermal time constant of the leaves, indicating that species with thermally buffered leaves maintain higher heat tolerance. Among lowland species, T50 increased with leaf mass per area, suggesting that in species with structurally more costly leaves the risk of leaf loss during hot spells is reduced. These results provide insight in variation in heat tolerance at local and regional scales.

CAM photosynthesis in desert blooming Cistanthe of the Atacama, Chile.

When plants of the Atacama desert undergo episodic blooms, among the most prominent are succulent-leaved Cistanthe (Montiaceae). We demonstrate that two Cistanthe species, the perennial Cistanthe sp. aff. crassifolia and the annual/biannual Cistanthe sp. aff. longiscapa, can exhibit net CO2 uptake and leaf acidification patterns typical of crassulacean acid metabolism (CAM). In C. sp. aff. crassifolia leaves, CAM expression was facultative. CAM-type nocturnal net CO2 uptake and acid accumulation occurred in drought-stressed but not in well-watered plants. By contrast, CAM expression in C. sp. aff. longiscapa was largely constitutive. Nocturnal acid accumulation was present in leaves of well-watered and in droughted plants. Following water-deficit stress, net nocturnal CO2 uptake was induced and the level of acid accumulated increased. Neither nocturnal CO2 uptake nor acid accumulation was reduced when the plants were re-watered. δ13C values of a further nine field-collected Cistanthe species are consistent with a contribution of CAM to their carbon pools. In the Portulacinae, a suborder with eight CAM-containing families, Cistanthe becomes the sixth genus with CAM within the family Montiaceae, and it is likely that the ancestor of all Portulacineae also possessed CAM photosynthesis. In the stochastic rainfall landscape of the Atacama, carbon uptake in the dark is a water-use efficient mechanism that increases the carbon pool available for seed production or dormancy. The next rain event may be years away.