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In order to assess the response of cocoa trees to drought, changes in water status, gas exchange, leaf carbon isotopic ratio (δ13C), photochemical activity, and leaf N and chlorophyll content during the rainy and dry season were measured in 31 Venezuelan cocoa clones (17 Trinitarios, 6 Criollos, and 8 Modern Criollos) grown in a common garden. Drought caused a 40% decrease in water potential (ψ) in all but the Modern Criollos, and a reduction in net photosynthetic rate (A) and stomatal conductance (gs) without an increase in instantaneous water use efficiency (WUE) in 93% of clones, and an increase inδ13C (long-term WUE) in 74% of clones; these responses suggest differences in tolerance to drought among clones. A positive correlation betweenAand bothgsand leaf N content was found for all genotypes. Leaf N content, chlorophyll content, and photochemical activity were reduced during drought, suggesting that metabolism was also inhibited. The best performance during drought was shown by Modern Criollos with the highest WUE, while five Trinitario clones seemed to be less sensitive to drought, since neither chlorophyll, N, total soluble protein concentration, norgschanged with drought, indicating that those Trinitario clones, with lowerA, have a conservative water use. Modern Criollos showed no reductions in eitherψorgs;Aremained unchanged, as did WUE, which was the highest, suggesting that these clones would be more successful in environments with low water availability. Our results indicate large variation in physiological response to drought over a range of parameters, suggesting possible differences in tolerance among clones.

We evaluated the effects of an elevated [CO2] on photosynthesis and growth of cassava plants grown in open-top chambers with an adequate supply of water and N and a sufficient rooting volume. Cassava plants (Manihot esculenta Crantz. cv. Motilona) showed higher photosynthetic rates (Pn) when grown and measured at elevated [CO2] (680 µmol mol-1) than when grown and measured at ambient [CO2] (480 µmol mol-1). No downregulation of photosynthesis due to elevated [CO2] was found, since carboxylation efficiency increased after 220 d in spite of a decrease in leaf soluble protein, Rubisco, and leaf N content. Soluble sugar and starch contents decreased with time under elevated [CO2], the decrease in starch content coinciding with the beginning of the increase in root mass. Canopy Pn by leaf area decreased with time under elevated [CO2] but, when canopy Pn was expressed by ground area, higher and constant rates were observed, suggesting a higher productivity in plants grown at elevated [CO2]. The absence of differences between growth [CO2] in root : shoot ratio observed suggests that elevated [CO2], while causing increases in the shoot as well as the root, did not affect the pattern of biomass allocation. Acclimation responses of gas exchange parameters changed during the experiment. The absence of downregulation of photosynthesis was associated with a decrease in leaf sugar and starch contents of plants grown at elevated [CO2], which suggests a favourable source/sink relationship.

BACKGROUND AND AIMS: Crassulacean acid metabolism (CAM) can be induced by salinity, thus conferring the plant higher water-use efficiency. Talinum triangulare does not frequently encounter salt in its natural habitat but is cultivated in soils that may become salinized. Here we examined whether plants of T. triangulare can grow in saline soils and show salt-induced CAM. METHODS: Leaf gas exchange, carbon isotopic ratio (δ(13)C), nocturnal acid accumulation (ΔH(+)), water relations, photosynthetic pigment and mineral contents, leaf anatomy and growth were determined in greenhouse in plants irrigated with 0, 150, 300 and 400 mm NaCl. KEY RESULTS: Salinity reduced gas exchange and induced CAM, ΔH(+) reaching 50.2 μmol H(+) g(−1) fresh mass under 300 mm NaCl. No nocturnal CO(2) uptake, but compensation, was observed. Values of δ(13)C were lowest under 0 and 400 mm NaCl, and highest under 150 and 300 mm. The difference in osmotic potential (ψ(s)) between control and treated plants averaged 0.45 MPa for the three [NaCl] values, the decrease in ψ(s) being accounted for by up to 63 % by Na(+) and K(+). Pigment contents were unaffected by treatment, suggesting lack of damage to the photosynthetic machinery. Changes in stomatal index with unchanged stomatal density in newly expanded leaves suggested inhibited differentiation of epidermal cells into stomata. Whole-leaf and parenchymata thickness increased under 150 and 300 mm NaCl. Only plants irrigated with 400 mm NaCl showed reductions in biomass (stems, 41 %; reproductive structures, 78 %). The K/Na molar ratio decreased with [NaCl] from 2.0 to 0.4. CONCLUSIONS: The operation of CAM in the recycling mode was evidenced by increased ΔH(+) with no nocturnal CO(2) uptake. Talinum triangulare can be classified as a halo-tolerant species based on its low K/Na molar ratio under salinity and the relatively small reduction in growth only at the highest [NaCl].

We aimed at determining whether C 4 and CAM species had a higher reproductive efficiency than C 3 species because of their higher water- and nitrogen-use efficiency. The resource-limitation hypothesis was also examined in the context of photosynthetic pathway. Reproductive efficiency was evaluated by measuring fruit set (fruit/flower ratio), seed set (seed/ovule ratio), and relative fecundity (the proportion of well-developed seed formed per ovule per infrutescence), along with seed abortion, and flower, fruit and seed biomass were evaluated. Photosynthetic pathway was determined according to literature, taxonomy, leaf anatomy and carbon isotopic composition (δ 13 C). Additionally, carbon and nitrogen content and nitrogen isotopic composition (δ 15 N) were determined. Plant life form and breeding system were considered. The traits of δ 13 C and δ 15 N were found to be positively correlated to fruit set, seed set and relative fecundity. The C 4 species exhibited the highest values of fruit and seed set associated to the lowest biomass of flower, fruit, seed, and seed/fruit. The CAM species exhibited the highest values of flower and seed/fruit biomass and C 3 species had the highest values of fruit and seed biomass. Reproductive efficiency was negatively related to the cost of reproductive structures in C 3 species. The highest values of maximum relative fecundity were reached in CAM and C 4 species under contrasting costs of reproductive structures: high values of fruits and seeds biomass in CAM species and low values of fruit and seed biomass in C 4 species, suggesting that CAM species may reach high values of maximum fecundity irrespectively of fruit and seed cost. Patterns found in reproductive efficiency may be slightly modified according to life form and breeding system: fruit set was higher for annual than perennial herbs, which may be initially associated to the self-compatibility and autogamy of annual herbs. Fruit set increased from C 3 to CAM species for annual herbs and decreased from C 3 to CAM species for perennial herbs, indicating that C 4 and CAM photosynthetic pathways could be associated with the increment of reproductive efficiency in annual species. Besides the traditionally acknowledged relationship with resistance to stressful environments, reproductive strategies of C 4 and CAM plants could be also related to an increment in plant reproduction.

Cocoa is sensitive to both water deficit (WD) and waterlogging (WL), especially during the juvenile stage. The aim of this study was to compare in four cocoa clones the water status, gas exchange and photochemical activity of PSII of adult trees subjected to natural drought in the field, and young individuals in the greenhouse subjected to either WD or WL. In the greenhouse, pots in which eight-month-old saplings were growing were placed in 8-L plastic buckets and either weekly watered (control, C), left without watering (WD), or filled with tap water up to 1 cm above soil surface (WL). In the field, drought affected clones differently in their water relations and photosynthetic responses. In the greenhouse, after 35 days of treatment, the four clones showed decreases in water potential which were stronger under WD than WL. Photosynthetic rate ( P N ) decreased on average 86% due to WD and 60% due to WL, clone 415 being the most susceptible to both stresses. On average, P N reached 100% of control values ​​7 d after re-watering but only 74% after drainage. Stomatal closure with maintenance of leaf water was the general response to WD. Clone 439, with highest relative lenticel area, was the one in which P N recovered most from WL. Lenticel hypertrophy, particularly in clone 439, apparently was one of the mechanisms of acclimation to WL. Chronic photoinhibition occurred in no case. We conclude that all clones are adequate for growth in the field, although 415 is the one with lower P N during the rainy season. Saplings of all clones are suitable for cultivation in drought-prone areas for dry periods of at least a month, and clone 439 is the one best suited for soils under short-term WL .

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.

In order to determine how flooding affects sap flow and hydraulic conductivity of the tolerant species, Campsiandra laurifolia, trees growing in a tropical seasonally flooded forest in Venezuela were studied. We hypothesized that trees respond to rising-waters with a decrease in root-water absorption, caused by hypoxia, and stomatal conductance, and that this is reverted later on through a process of acclimation that involves improvement in water absorption. We followed the seasonal changes, of trees with the whole or part of the canopy exposed to air, in sap flow density, leaf stomatal conductance, leaf transpiration rate and xylem water potential. The highest daytime sap flow density occurred at noon and its proportion relative to the yearly maximum (drainage at falling-waters) was 41 (dry season), 15 (flooding by rising-waters for 2 weeks), 54 (2 months of flooding) and 41% (6 months of flooding). Since at rising-waters dawn xylem water potential remained high, it became apparent that the initial stages of flooding imposed a restriction to sap flow unrelated to water deficit. The decrease at rising-waters in highest daytime sap flow density was due to reduced leaf-specific hydraulic conductivity, whereas the recovery observed 1.5 months later was correlated to an increase in hydraulic conductivity, and attributed to acclimation. Sap flow density was highly and positively correlated with radiation at all seasons but rising-waters; also, the relationship with air water vapor saturation deficit was high and significant on dates other than at rising-waters. Results suggest that early flooding inhibited water absorption by roots and that this inhibition was overcome later on at a higher water column through an acclimation process involving the improvement of internal aeration by adventitious roots.

Clusia minor L. is a C3-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 CO2 fixation.

Seasonal changes in water relations, net photosynthetic rate (P(N)), and fluorescence of chlorophyll (Chl) a of two perennial C(3) deciduous shrubs, Ipomoea carnea and Jatropha gossypifolia, growing in a thorn scrub in Venezuela were studied in order to establish the possible occurrence of photoinhibition during dry season and determine whether changes in photochemical activity of photosystem 2 (PS2) may explain variations of PN in these species. Leaf water potential (psi) decreased from -0.2 to -2.1 MPa during drought in both species. The PN decreased with y in T carnea and J. gossypifolia by 64 and 74%, respectively. Carboxylation efficiency (CE) decreased by more than 50 and 70% in I. carnea and J gossypifolia, respectively. In I. carnea, relative stomatal limitation (L(s)) increased by 17% and mesophyll limitation (L(m)) by 65% during drought, while in J. gossypifolia Ls decreased by 27% and L. increased by 51%. Drought caused a reduction in quantum yield of PS2 (theta(PS2)) in both species. Drought affected the capacity of energy dissipation of leaves, judging from the changes in the photochemical (q(P)) and non-photochemical quenching (NPQ) coefficients. Photo-inhibition during drought in I. carnea and J gossypifolia was evidenced in the field by a drop in the maximum quantum yield of PS2 (F(v)/F(m)) below 0.8 and also by non-coordinated changes in phi(PS2) and quantum yield of non-photochemical excitation quenching (Y(n)). Total soluble protein content on an area basis increased with psi but the ribulose-1,5-bisphosphate carboxylase/oxygenase content remained unchanged. A reduction of total Chl content with drought was observed. Hence in the species studied photoinhibition occurred, which imposed an important limitation on carbon assimilation during drought.

Water relations and photosynthetic characteristics of plants of Lycium nodosum grown under increasing water deficit (WD), saline spray (SS) or saline irrigation (SI) were studied. Plants of this perennial, deciduous shrub growing in the coastal thorn scrubs of Venezuela show succulent leaves which persist for approx. 1 month after the beginning of the dry season; leaf succulence is higher in populations closer to the sea. These observations suggested that L. nodosum is tolerant both to WD and salinity. In the glasshouse, WD caused a marked decrease in the xylem water potential (y), leaf osmotic potential (ys) and relative water content (RWC) after 21 d; additionally, photosynthetic rate (A), carboxylation efficiency (CE) and stomatal conductance (gs) decreased by more than 90 %. In contrast, in plants treated for 21 d with a foliar spray with 35 ‰ NaCl or irrigation with a 10 % NaCl solution, y and RWC remained nearly constant, while ys decreased by 30 %, and A, CE and gs decreased by more than 80 %. An osmotic adjustment of 0·60 (SS) and 0·94 MPa (SI) was measured. Relative stomatal and mesophyll limitations to A increased with both WD and SS, but were not determined for SI-treated plants. No evidence of chronic photoinhibition due to any treatment was observed, since maximum quantum yield of PSII, Fv/Fm, did not change with either drought in the field or water or salinity stress in the glasshouse. Nevertheless, WD and SI treatments caused a decrease in the photochemical (qP) and an increase in the nonphotochemical (qN) quenching coefficients relative to controls; qN was unaffected by the SS treatment. The occurrence of co-limitation of A by stomatal and non-stomatal factors in plants of L. nodosum may be associated with the extended leaf duration under water or saline stress. Additionally, osmotic adjustment may partly explain the relative maintenance of A and gs in the SS and SI treatments and the tolerance to salinity of plants of this species in coastal habitats. a 2003 Annals of Botany Company

In obligate Crassulacean acid metabolism (CAM) plants, dark CO2 fixation is almost the sole route of CO2 fixation and, under drought, continues for long periods. In contrast, in plants of the facultative CAM species Talinum triangulare under experimental drought, dark CO2 fixation provides a small proportion of the daily assimilation observed in watered plants and occurs only for a few days, after which almost nil CO2 fixation is observed. Under field conditions, with a practically unlimited substrate volume, drought-induced CAM might operate for a longer period and make a higher contribution to daily CO2 fixation. Greenhouse-grown plants of T. triangulare were subjected to low and nearly constant soil water content; the operation of CAM was assessed through the measurement of nocturnal proton accumulation and dark CO2 fixation. Dark CO2 fixation appeared 19d after the onset of drought; its contribution during three months of experiment to daily CO2 assimilation ranged from 0.5 to 30.7% with a mean of 13.5%. Twenty days after the beginning of treatment, nocturnal proton accumulation increased six times and remained high for over three months. In spite of low soil water content, leaves did not engage in dark CO2 fixation all the time but dark CO2 fixation was large enough to produce an increase in relative (13)C composition of mature leaves compared to watered plants but not to the value in short-term drought experiments. Leaf anatomical characteristics may guarantee the achievement of higher rates of dark CO2 fixation but results evidence the occurrence of a limit to the expression of CAM that remains to be determined.

The mechanisms underlying the drought tolerance of Peperomia carnevalii Steyermark (Piperaceae), a succulent herb growing in the understorey of seasonally dry forests, were examined. Crassulacean acid metabolism (CAM) was studied in the field and laboratory, and measurements of water status were made in plants subjected to drought in the greenhouse. Nocturnal acid accumulation and day and night-time CO 2 assimilation rates were greatest in watered plants and decreased in drought. The proportion of CO 2 recycled through CAM in droughted plants, with nocturnal CO 2 uptake close to zero, was higher than in watered plants. Maximum quantum yield of chlorophyll fluorescence remained unchanged during drought, but the PSII quantum yield at the photosynthetic photon flux density at which the plants were grown was significantly decreased. Leaf anatomy consists of a chlorophyll-less hydrenchyma located beneath the upper epidermis, and a two-layered mesophyll. Leaves nearer to the apex are thinner than those nearer to the base of the shoot. Drought caused a reduction in leaf thickness due to shrinkage of the hydrenchyma, but not of the mesophyll. This was associated with the occurrence of a gradient of osmotic potential between these tissues. Comparison of water loss from thin leaves of watered and droughted plants, either partly defoliated at the lower nodes or intact, suggested that water moved from the thick to the thin leaves. This process was related to the occurrence of a gradient of water potential between the thick and the thin leaves. Drought tolerance in P. carnevalii is achieved by the operation of CAM and the occurrence of water movement within and between leaves.

We studied the responses of leaf gas exchange and growth to an increase in atmospheric CO2 concentration in four tropical deciduous species differing in carbon fixation metabolism: Alternanthera crucis, C3-C4; Ipomoea carnea, C3; Jatropha gossypifolia, C3; and Talinum triangulare, inducible-CAM. In the first stage, plants were grown in one open-top chamber at a CO2 concentration of 560±40 μmol mol-1 (EC), one ambient CO2 concentration chamber (AC), and one unenclosed plot (U). In the second stage, plants were grown in five EC chambers (CO2 concentration = 680±30 μmol mol-1), five AC chambers, and five unenclosed plots. During the first weeks under EC in the first stage, plants of all the species had a very marked increase in their maximal net photosynthetic rates (Pmax) of 3.5 times on average; this stimulatory effect was maintained for 11-15 weeks, rates dampening afterward to values still higher than controls for 37 weeks. After a suspension of CO2 enrichment for 6 weeks, an increase in Pmax of EC plants over the controls was found in plants of all the species until week 82 of the experiment. Stomatal conductance (g) showed no response to EC. Carboxylation efficiency decreased in all the species under EC

The effect of a very high CO2 mole fraction (27000–35000 μmol mol−1) on photosynthesis and water relations was studied during the dry and the rainy season in plants of Spatiphylum cannifolium (Dryand.) Schott and Bauhinia multinervia (H.B.K.) DC. growing near natural cold CO2 springs. Xylem water potential in plants of both species was lowered by drought, high CO2 growth-concentration decreasing it further in S. cannifolium. In plants of both species growing under high CO2 concentration photosynthetic rates measured at a CO2 mole fraction of 1000 μmol mol−1 were higher than in plants growing at ambient CO2 mole fraction and measured at 350 μmol mol−1. The response was the result of a direct effect of CO2 on the photosynthetic machinery. Changes in carboxylation efficiency in response to high CO2 were found during the rainy season, with an increase in S. cannifolium and a decrease in B. multinervia; a significant interaction between growth CO2 concentration and season in B. multinervia resulted from significant effects of both factors. An increase in intrinsic water-use efficiency due to high CO2 was determined in both species by an increase in photosynthetic rate as well as a decrease in leaf conductance. In high-CO2 plants of S. cannifolium a 71% decrease in stomatal density and 73% in stomatal index suggested that CO2 affected stomatal initiation, whereas in B. multinervia an 85% decrease in stomatal index and a 72% decrease in stomatal density indicated that CO2 influenced stomatal initiation as well as epidermal cell expansion. Our results indicate that very high CO2 concentrations did not inhibit photosynthesis in these species, and that growth under high CO2 allowed plants to attain carbon balances higher than those of plants growing under low CO2. This was particularly so during the dry season, since the photosynthetic rates at the corresponding ambient concentration were higher in plants nearer the springs, and carboxylation efficiency and some stomatal characteristics of both species apparently acclimated to high CO2, but patterns were not consistent and bore no obvious relationship to photosynthetic capacity.

To investigate the possible induction of Crassulacean acid metabolism (CAM) by drought in Talinum paniculatum ([Jacq.] Gaertn.), a deciduous herb with succulent leaves and lignified stems, nocttimal acid accumulation and COj-exchange were studied in watered and droughted greenhouse-grown plants. Watered plants had a t>'pica] C3 pattern of COj-exchange. When plants were subjected to drought, noetumal acid accumulation increased significantly from 0.9 to 13.4 pmol H*cm after 21 days. Water deficit provoked a rapid reduction of daytime CO2 assimilation of as mtich as 92% and a slower increase in night-time fixation. A maximum of 24% of the diel carbon gain was contributed by daik fixation in droughted plants. After 34 days of drought, only CO2 compensation and a small accumulation of acid (idling) was detected during the night. Relative recycling of respiratory C02 was approximately 100% for most of the water deficit treatment, the amount of CO2 recycled showing a high positive correlation with nocturnal acid accumulation. A low rate of nocturnal loss of CO3 in watered plants did not explain the amount recycled nightly in droughted plants, implying that respiration increased with drought. Leaf lamina area was reduced by 49% during drought due to rolling. Leaf biomass remained unchanged during the water-deficit treatment Neither apparent quantum yield nor light-saturated photosynthetic rate differed significantly between control and 14-day water-stressed plants rewatered for 20 h. ChlorophyE content did not change with drought. These results confirm that CAM is induced by drought in T. paniculatum; the carbon acquired through this pathway only contributes to maintain, but not to inciease, leaf biomass; also, CAM is responsible for a high recycling of respiratory CO2 during the night. Recycling through CAM, plus the reduction of exposed isif area during drought,, may help explain the maintenance of chlorophyll, quantum yield and saturated photosynthetic rates in water-stressed plants of T. paniculatum.

Trees in the flooded forest of the Mapire River in Venezuela suffer a decrease in photosynthetic rate (A) when flood begins, which is reverted at maximum flood. Changes in A are accompanied by similar changes in stomatal conductance (g(s)), and the possibility of changes in photosynthetic capacity is not ruled out. In order to understand how relative stomatal and non-stomatal limitations of photosynthesis are affected by flooding, we studied the seasonal changes in A and its response to intercellular CO(2) concentration in trees of Campsiandra laurifolia, Symmeria paniculata, Acosmium nitens and Eschweilera tenuifolia. Flooding caused in trees of C. laurifolia and S. paniculata a reduction in A, g(s), carboxylation efficiency and total soluble protein (TSP), whereas gas exchange in A. nitens and E. tenuifolia was more sensitive to drought. Under flooding, relative stomatal limitation (L(s)) was on average half the highest, and relative non-stomatal limitation (L(ns)) increased from the dry season to flooding. Under full flood, A, g(s) and TSP regained high values. A was positively correlated to light-saturated electron transport rate, suggesting that part of the decrease in A under flooding was due to impairment of photosynthetic capacity. Under flooding, not only stomatal closure but also increased L(ns) causes a reduction in photosynthesis of all four species, and a process of acclimation as flooding progresses allows gas exchange and related variables to regain high values.

Changes in photochemical activity induced by water deficit were investigated in Talinum triangulare, an inducible CAM plant. The aim was to analyse the interactions between C3 photosynthesis, induction and activity of CAM, photosynthetic energy regulation and the mechanisms responsible for photoprotection and photoinhibition under water stress. Gas exchange, chlorophyll a fluorescence, titratable acidity, carotenoid composition and relative contents of the PSII reaction centre protein (D1) were measured. A decrease in xylem tension (psi) from -0.14 to -0.2 MPa substantially decreased daytime net CO2 assimilation and daily carbon gain, and induced CAM, as shown by CO2 assimilation during the night and changes in titratable acidity; a further decrease in psi decreased nocturnal acid accumulation by 60%. Non-photochemical quenching of chlorophyll a fluorescence (NPQ) increased with water deficit, but decreased with a more severe drought (psi below -0.2 MPa), when CAM activity was low. NPQ was lower at 0900 h (during maximum decarboxylation rates) than at 1400 h, when malate pools were depleted. Down-regulation of PSII activity related to the rise in NPQ was indicated by a smaller quantum yield of PSII photochemistry (phiPSII) in droughted compared with watered plants. However, phiPSII was larger at 0900 h than at 1400 h. The de-epoxidation state of the xanthophyll cycle increased with drought and was linearly related to NPQ. Intrinsic quantum yield of PSII (FV/FM) measured at dusk was also lower in severely stressed plants than in controls. Under maximum photosynthetic photon flux and high decarboxylation rates of organic acids, the D1 content in leaves of droughted plants showing maximal CAM activity was identical to the controls; increased drought decreased D1 content by more than 30%. Predawn samples had D1 contents similar to leaves sampled at peak irradiance, with no signs of recovery after 12 h of darkness. It is concluded that under mild water stress, early induction of CAM, together with an increased energy dissipation by the xanthophyll cycle, prevents net degradation of D1 protein; when water deficit is more severe, CAM and xanthophyll cycle capacities for energy dissipation decline, and net degradation of D1 proceeds.

We studied the flood tolerance of five tree species growing in the flooded forest adjacent to the Mapire river, in SW Venezuela. Mean photosynthetic rate and leaf conductance were 11mgr;mol m(-2) s(-1) and 700 mmol m(-2) s(-1), respectively. Xylem water potential ranged from -0.08 to -1.15 MPa. Based on leaf gas exchange as a criterion of tolerance to flooding, two response patterns were identified: (1) decreasing photosynthetic rate with increasing flooding and leaf conductance (Psidium ovatifolium Berg. ex Desc., Campsiandra laurifolia Benth., Symmeria paniculata Benth. and Acosmium nitens (Vog.) Benth); and (2) independence of photosynthesis and leaf conductance from flooding (Eschweilera tenuifolia (Berg.) Miers.). In the first response pattern, declining photosynthetic rate with flooding may be interpreted as a sign of reduced flood tolerance, whereas the second response pattern may indicate increased flood tolerance. An increase in xylem water potential with depth of water column was found for all species (with the possible exception of P. ovatifolium), indicating that flooding does not cause water stress in these trees. Submerged leaves that had been under water for between four days and four months generally had photosynthetic rates and leaf conductances similar to those of aerial leaves, indicating maintenance of photosynthetic capacity under water. Daily positive oscillations in glucan content in submerged leaves of P. ovatifolium and C. laurifolia suggest that submerged leaves do not represent a sink for photosynthates produced by aerial leaves.