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 C(3)-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.

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 Kalanchoë daigremontiana and Kalanchoë 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.

Androecia in two Clusia species: development, structure and resin secretion.

Clusia fluminensis and C. lanceolata are dioecious shrubs having resiniferous flowers with strongly distinct androecia. The aim of this study was to investigate the development and anatomy of their androecia and the ultrastructure, histochemistry and secretory process of their androecium resin glands, examining whether the cellular aspects of resin secretion differed between these two morphologically distinct androecia. Stamens differ, being free in C. fluminensis and clustered in a synandrium in C. lanceolata. Staminode sterility is due to the undifferentiated nature of the anthers in C. lanceolata and degeneration of meiocytes and anther indehiscence in C. fluminensis. Resin is produced in subepidermal cavities and canals with wide lumens. In the secretory stage, epithelial cells present sinuous walls, voluminous nuclei, polymorphic plastids associated with periplastidial reticulum, mitochondria, oil bodies, multivesicular bodies, endoplasmic reticulum and dictyosomes. The resin is released through rupture points on the distal surface of stamens and staminodes, associated with disrupted cavities and canals. Our results show morphological diversity associated with functional similarity. Also, a secretion pattern shared by the two species includes initiation of the secretory process in young floral buds, compartmentalisation of the secretion in pre-anthesis buds and release of secretions at anthesis. Cellular aspects of resin secretion in these species are quite similar, as are the chemical identities of the main components of the floral resins of the genus.

Aspects of reproductive ecology of Clusia valerioi Standl. and Clusia peninsulae Hammel (sp. nov.), two Central American species of Clusiaceae with resin flowers.

Clusia valerioi Standl. and C. peninsulae Hammel, sp. nov. (formally established in the Appendix), two Central American species of Clusiaceae offering resin as a floral reward, were studied in the Piedras Blancas National Park, SW Costa Rica. Basic data on phenology, flower structure, course of anthesis, flower visitors, flower visitation and pollination processes, fruit development and seed dispersal are communicated. Resin collection and pollen release are reported and documented in greater detail. The significance of stingless bees for pollination is confirmed, but honeybees were also observed to visit the flowers and to collect resin. The seeds of C. valerioi, exhibiting a clearly ornithochorous character syndrome, were observed to be eaten and dispersed by four species of the tanager family (Thraupidae) and one species of finch (Fringillidae). Ants were also observed to carry away the seeds.