This review describes the life cycle of Mesembryantheum crystallinum L. (the common ice plant, Aizoaceae, Caryophyllales), a halophyte with a developmentally programmed switch from C3 photosynthesis to Crassulacean acid metabolism (CAM) which is accelerated by salinity and drought. Since there has been controversy regarding the interplay between genes and environmental stimuli during the development of M. crystallinum, it is timely to summarize the life cycle for a defined set of conditions. We seek to establish the framework whereby five stages of development can be described in terms of morphology, physiology, and molecular biology. Stages 1 and 2, representing germination and growth of a juvenile form, show a determinate pattern of growth. Although specific genes for salt tolerance can be induced at these stages, stress early in development prevents progression to the mature form (stages 3–5) in which the plants advance to mature growth, flowering, and seed development. Growth in stage 3 is indeterminate in the absence of stress, but development and flowering are accelerated by environmental stresses, and CAM is constitutively expressed. Depending on the severity of the stress, plants start to flower (stage 4) and then die from the roots, ultimately with only seed capsules remaining viable, with salt sequestered into large epidermal bladder cells (stage 5). We highlight responses to salinity leading to compartmentation of ions and compatible solutes, turgor maintenance, and CAM. Finally, the molecular genetics of the ice plant are characterized, emphasizing selected genes and their products. We conclude with an analysis of the multiple stages of growth as an ecological adaptation to progressive stress. The initial determinate and inflexible juvenile phase provides a critical mass of plant material which supports the indeterminate, mature phase. Depending on the degree of stress, the mature form is then propelled towards flowering and seedset.

Changes in leaf water content, night-time accumulation of malic (Δ-malate) and citric acid (Δ-citrate) and phosphoenolpyruvate carboxylase (PEPC, EC 4 . 1 . 1 . 31) activity were followed for 60 d after germination in well watered and salt-stressed plants of the facultatively halophytic ephemeral Mesembryanthemum crystallinum L. To separate the effects of development, salt stress and water deficit on crassulacean acid metabolism (CAM) induction plants were stressed initially 10 d after germination and then successively at 1-wk intervals (five sets). Related to dry mass or organic matter (i.e. dry mass corrected for the mass of inorganic ions) water content started to decrease during the late embryonal phase of the life cycle. Water content on a dry mass basis was always lower in salt-stressed than in well watered individuals. However, on an organic matter basis no difference was detectable. This indicated that salt treatment did not reduce leaf water content but falsified the basis (dry mass). Increases in leaf succulence and in pressure potential prevented long-term water deficit in well watered and in salt-stressed plants. Instead, these changes displayed enhanced vacuolisation, which is an essential prerequisite for the development of CAM. The end of that differentiation process might allow the initiation of nocturnal malic acid accumulation in a threshold response. At the onset of each salt treatment, short-term water deficits occurred due to an incomplete osmotic adaptation independent of plant age. As Δ-malate only appeared when plants were c. 35 d old this water deficit was unlikely to be a decisive CAM-inducing factor. About 2 wk after germination water content began to decline during the light periods in plants of all treatments. This pattern disappeared again when CAM had been fully established. Daytime transpirational water loss is therefore unlikely to be the decisive factor because it failed to induce the metabolic shift in young plants. Environmental stress (e.g. salt or drought) can therefore only induce Δ-malate when leaf and plant differentiation has reached a certain stage.