In the terrestrial bromeliad, Puya floccosa, a value of carbon isotopic composition (δ 13 C) of -22‰ has been previously reported, suggesting the operation of weak and/or intermediate (C 3 -CAM) crassulacean acid metabolism (CAM). In order to characterize the operation of CAM in P. floccosa and its possible induction by drought, plants were grown in Caracas and subjected to four independent drought cycles. Additionally, since plants of this species grow in Venezuela in a large range of elevations, leaf samples were collected at elevations ranging from 725 to 2,100 m a.s.l. in the Venezuelan Andes and the Coastal Range, in order to evaluate the effect of elevation on CAM performance. Even though nocturnal acid accumulation occurred in both watered and droughted plants, mean ΔH + was higher in droughted than watered plants [ΔH + = 60.17.5 and 22.9 ± 5.2 μmol g -1 (FM), respectively]. The majority of plants from all the natural populations sampled had low values of δ 13 C not differing significantly from those of C 3 plants collected as standards and δ 13 C did not change with elevation. We conclude that P. floccosa is capable of a weak CAM activity, with a large variability among populations and drought experiments probably due to local and temporal differences in microclimatic variables and drought stress; elevation bears no influence on values of δ 13 C in this species.Additional key words: Bromeliaceae, elevation, nocturnal acid accumulation.
Clusia minor L. is a C 3 -CAM species in which Crassulacean acid Metabolism (CAM) is induced, among other factors, by water deficit. We propose that CAM induction by natural drought in C. minor shifts the sap flow pattern from daytime to a night-time one, and that the decreased osmotic potential due to increased malate content in droughted plants aids in the increase in nocturnal sap flow. In order to test these hypotheses, we followed for 2 years the seasonal changes in parameters of water relationships and sap flow velocity in one single, freestanding tree growing in Caracas. Leaf water and osmotic potential were measured psychrometrically, nocturnal proton accumulation by titration of aqueous leaf extracts and sap flow density with thermal dissipation probes. Leaf water, osmotic and turgor potential remained relatively high throughout the seasons. Nocturnal proton accumulation was nil under extreme drought or after frequent and heavy rains, and high after moderate rainfall. Estimated malate and citrate concentrations contributed up to 80 and 60%, respectively, of the value of osmotic potential. The shape of the daily courses of sap flow velocity varied seasonally, from mostly diurnal during the dry season to mostly nocturnal after a short dry spell during the rainy season, when nocturnal acid accumulation attained high values. There was a strong positive relationship between the proportion of the integrated sap flow courses corresponding to the night and dawn [H + ] (r 2 = 0.88). Increased nocturnal sap flow in the CAM stage of the tree of C. minor may be explained by a lower osmotic potential due to an increased acid concentration, together with increased stomatal aperture, as suggested by increased nocturnal acid accumulation probably due to nocturnal CO 2 fixation.
Vegetative as well as reproductive phenology of tropical trees frequently occur synchronously at the end of the dry season. It has been repeatedly observed on two individuals of the tropical species Ficus obtusifolia growing in Caracas that they lose their entire foliage in a period of 48 h during the dry season and renew it within the next 72 h. In order to gain knowledge on the mechanisms that govern leaf exchange in F. obtusifolia, we followed in these two individuals the seasonal changes in leaf and fruit phenology, water relations and environmental variables, and in another 25, phenology only. Although foliar exchange was always observed before the rainy season, it also occurred near the end of the rainy season. An increase in leaf turgor potential of mature leaves and a low sap flow velocity coincided with leaf fall. Foliar exchange in all 27 individuals, growing in locations with different elevation and, possibly, access to underground water, was bi-modal and occurred in a period of approximately one month around both equinoxes. The presence of fruits was also bi-modal, occurring after leaf renewal. The sequence of phenological events repeated itself in all the individuals studied, although with inter-annual and individual variations in date of foliar exchange. Leaf fall was independent of leaf water status and significantly correlated with, in decreasing order, day length, minimum air temperature, rainfall, maximum air temperature and total radiation. We conclude that in F. obtusifolia high nocturnal temperature, increasing daytime temperature and radiation, and low cloud cover apparently drove foliar exchange. Day length showed a strong correlation with leaf renewal possibly to due to the significant relationships between day length and environmental variables. Direct effects of day length alone were not evidenced, as foliar exchange took place during a long time-span around the equinoxes.
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