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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., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

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., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

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., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Annals of Botany Company.)

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 (C FT ). The effect of CAM and elevated C FT 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 C FT , across the genus Clusia. We also simulated the effects of CAM and C FT on growth and water conservation during drought using the Photo3 model. Within Clusia leaves, CAM and C FT are independent traits: CAM requires large palisade chlorenchyma cells, whereas hydrenchyma tissue governs interspecific differences in C FT . In addition, our model suggests that CAM supersedes C FT as a means to maximise CO 2 assimilation and minimise transpiration during drought. Our study challenges the assumption that CAM and C FT are mutually dependent traits within succulent leaves., (© 2023 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

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 C 3 plants. Resolving how CAM photosynthesis evolved from the ancestral C 3 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 C 3 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 C 3 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 CO 2 fixation. This central element of the CAM pathway, even when expressed at a low level, represents a biochemical capability not seen in C 3 plants, and so is better regarded as a discrete evolutionary innovation than as part of a metabolic continuum between C 3 and CAM., (© 2021 The Authors. New Phytologist © 2021 New Phytologist Foundation This article has been contributed to by US Government employees and their work is in the public domain in the USA.)

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.

Clusia is a widely distributed neotropical genus with 321 currently described species. This remarkable genus is the only one known to contain trees sensu stricto with CAM photosynthesis. To survey the occurrence of CAM in Clusia species from Colombia, we determined the leaf stable carbon isotope composition (d13C) of 568 specimens from 114 species deposited in 12 Colombian herbaria. In the vast majority of specimens, d13C values indicated that C3 photosynthesis was the principal contributor to carbon gain. d13C values typical of strong CAM (less negative than -20‰) were observed in only five species, in four of them for the first time. All samples with CAM-type isotopic signatures were collected below 1,000 m a.s.l., whereas species with predominantly C3 occurred from sea level to 3,500 m a.s.l. Together with information already available in the literature, we conclude that CAM is present in 22% (35/156) of the species of Clusia investigated thus far. [ABSTRACT FROM AUTHOR]

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

Abstract Crassulacean acid metabolism (CAM), an advanced photosynthetic pathway conferring water conservation to plants in arid habitats, has enigmatically been reported in some species restricted to extremely wet tropical forests. Of these, epiphytic Bromeliaceae may possess absorbent foliar trichomes that hinder gas-exchange when wetted, imposing further limitations on carbon dioxide (CO2 ) uptake. The hypothesis that the metabolic plasticity inherent to CAM confers an ecological advantage over conventional C3 plants, when constant rainfall and mist might inhibit gas-exchange was investigated. Gas-exchange, fluorometry and organic acid and mineral nutrient contents were compared for the bromeliads Aechmea dactylina (CAM) and Werauhia capitata (C3 ) in situ at the Cerro Jefe cloud forest, Panama (annual rainfall > 4 m). Daily carbon gain and photosynthetic nutrient use efficiencies were consistently higher for A. dactylina , due to a greater CO2 uptake period, recycling of CO2 from respiration and a dynamic response of CO2 uptake to wetting of leaf surfaces. During the dry season CAM also had water conserving and photoprotective roles. A paucity of CAM species at Cerro Jefe suggests a recent radiation of this photosynthetic pathway into the wet cloud forest, with CAM extending diversity in form and function for epiphytes. [ABSTRACT FROM AUTHOR]