Dissecting succulence: Crassulacean acid metabolism and hydraulic capacitance are independent adaptations in Clusia leaves.

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 (CFT ). The effect of CAM and elevated CFT 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 CFT , across the genus Clusia. We also simulated the effects of CAM and CFT on growth and water conservation during drought using the Photo3 model. Within Clusia leaves, CAM and CFT are independent traits: CAM requires large palisade chlorenchyma cells, whereas hydrenchyma tissue governs interspecific differences in CFT . In addition, our model suggests that CAM supersedes CFT as a means to maximise CO2 assimilation and minimise transpiration during drought. Our study challenges the assumption that CAM and CFT are mutually dependent traits within succulent leaves.

Corrigendum to: Does the C4 plant Trianthema portulacastrum (Aizoaceae) exhibit weakly expressed crassulacean acid metabolism (CAM)?

We examined whether crassulacean acid metabolism (CAM) is present in Trianthema portulacastrum L. (Aizoaceae), a pantropical, salt-tolerant C4 annual herb with atriplicoid-type Kranz anatomy in leaves but not in stems. The leaves of T. portulacastrum are slightly succulent and the stems are fleshy, similar to some species of Portulaca, the only genus known in which C4 and CAM co-occur. Low- level nocturnal acidification typical of weakly expressed, predominantly constitutive CAM was measured in plants grown for their entire life-cycle in an outdoor raised garden box. Acidification was greater in stems than in leaves. Plants showed net CO2 uptake only during the light irrespective of soil water availability. However, nocturnal traces of CO2 exchange exhibited curved kinetics of reduced CO2 loss during the middle of the night consistent with low-level CAM. Trianthema becomes the second genus of vascular land plants in which C4 and features of CAM have been demonstrated to co-occur in the same plant and the first C4 plant with CAM-type acidification described for the Aizoaceae. Traditionally the stems of herbs are not sampled in screening studies. Small herbs with mildly succulent leaves and fleshy stems might be a numerically significant component of CAM biodiversity.

Leaf heat tolerance of 147 tropical forest species varies with elevation and leaf functional traits, but not with phylogeny.

Exceeding thermal thresholds causes irreversible damage and ultimately loss of leaves. The lowland tropics are among the warmest forested biomes, but little is known about heat tolerance of tropical forest plants. We surveyed leaf heat tolerance of sun-exposed leaves from 147 tropical lowland and pre-montane forest species by determining the temperatures at which potential photosystem II efficiency based on chlorophyll a fluorescence started to decrease (TCrit ) and had decreased by 50% (T50 ). TCrit averaged 46.7°C (5th-95th percentile: 43.5°C-49.7°C) and T50 averaged 49.9°C (47.8°C-52.5°C). Heat tolerance partially adjusted to site temperature; TCrit and T50 decreased with elevation by 0.40°C and 0.26°C per 100 m, respectively, while mean annual temperature decreased by 0.63°C per 100 m. The phylogenetic signal in heat tolerance was weak, suggesting that heat tolerance is more strongly controlled by environment than by evolutionary legacies. TCrit increased with the estimated thermal time constant of the leaves, indicating that species with thermally buffered leaves maintain higher heat tolerance. Among lowland species, T50 increased with leaf mass per area, suggesting that in species with structurally more costly leaves the risk of leaf loss during hot spells is reduced. These results provide insight in variation in heat tolerance at local and regional scales.

Low-level CAM photosynthesis in a succulent-leaved member of the Urticaceae, Pilea peperomioides.

Pilea peperomioides Diels (Urticaceae) is a semi-succulent herbaceous species native to south-western China that has become popular in cultivation as an ornamental plant. To investigate whether this species possesses the capacity for CAM photosynthesis, measurements were made of CO2 gas exchange and titratable acidity in plants under both well-watered and water-deficit conditions. Plants were found to assimilate CO2 almost exclusively in the light via C3 photosynthesis. However, distinct transient reductions in the rate of net nocturnal CO2 release were consistently observed during the course of the dark period, and under water-deficit conditions one plant exhibited a brief period of net nocturnal CO2 uptake, providing unequivocal evidence of CAM activity. Furthermore, nocturnal increases in titratable acidity in both leaf laminas and petioles were observed in all plants exposed to wet-dry-wet cycles. This is the first report of CAM in the family Urticaceae. The results are discussed in relation to the phylogenetic position of Pilea and the partially shaded montane habitats in which this species is typically found. An updated list of all plant families currently known to contain species with CAM is presented.

Constitutive and facultative crassulacean acid metabolism (CAM) in Cuban oregano, Coleus amboinicus (Lamiaceae).

Plants exhibiting the water-conserving crassulacean acid metabolism (CAM) photosynthetic pathway provide some of the most intriguing examples of photosynthetic diversity and plasticity. Here, a largely unnoticed facet of CAM-plant photosynthesis is highlighted: the co-occurrence of ontogenetically controlled constitutive and environmentally controlled facultative CAM in a species. Both forms of CAM are displayed in leaves of Coleus amboinicus Lour. (Lamiaceae), a semi-succulent perennial plant with oregano-like flavour that is native to southern and eastern Africa and naturalised elsewhere in the tropics. Under well-watered conditions, leaves assimilate CO2 predominantly by the C3 pathway. They also display low levels of CO2 uptake at night accompanied by small nocturnal increases in leaf tissue acidity. This indicates the presence of weakly expressed constitutive CAM. CAM expression is strongly enhanced in response to drought stress. The drought-enhanced component of CAM is reversible upon rewatering and thus considered to be facultative. In contrast to C. amboinicus, the thin-leaved closely related Coleus scutellarioides (L.) Benth. exhibits net CO2 fixation solely in the light via the C3 pathway, both under well-watered and drought conditions. However, low levels of nocturnal acidification detected in leaves and stems indicate that the CAM cycle is present. The highly speciose mint family, which contains few known CAM-exhibiting species and is composed predominantly of C3 species, appears to be an excellent group of plants for studying the evolutionary origins of CAM and for determining the position of facultative CAM along the C3-full CAM trajectory.

Does the C4 plant Trianthema portulacastrum (Aizoaceae) exhibit weakly expressed crassulacean acid metabolism (CAM)?

We examined whether crassulacean acid metabolism (CAM) is present in Trianthema portulacastrum L. (Aizoaceae), a pantropical, salt-tolerant C4 annual herb with atriplicoid-type Kranz anatomy in leaves but not in stems. The leaves of T. portulacastrum are slightly succulent and the stems are fleshy, similar to some species of Portulaca, the only genus known in which C4 and CAM co-occur. Low- level nocturnal acidification typical of weakly expressed, predominantly constitutive CAM was measured in plants grown for their entire life-cycle in an outdoor raised garden box. Acidification was greater in stems than in leaves. Plants showed net CO2 uptake only during the light irrespective of soil water availability. However, nocturnal traces of CO2 exchange exhibited curved kinetics of reduced CO2 loss during the middle of the night consistent with low-level CAM. Trianthema becomes the second genus of vascular land plants in which C4 and features of CAM have been demonstrated to co-occur in the same plant and the first C4 plant with CAM-type acidification described for the Aizoaceae. Traditionally the stems of herbs are not sampled in screening studies. Small herbs with mildly succulent leaves and fleshy stems might be a numerically significant component of CAM biodiversity.

Facultative crassulacean acid metabolism in a C3-C4 intermediate.

The Portulacaceae enable the study of the evolutionary relationship between C4 and crassulacean acid metabolism (CAM) photosynthesis. Shoots of well-watered plants of the C3-C4 intermediate species Portulaca cryptopetala Speg. exhibit net uptake of CO2 solely during the light. CO2 fixation is primarily via the C3 pathway as indicated by a strong stimulation of CO2 uptake when shoots were provided with air containing 2% O2. When plants were subjected to water stress, daytime CO2 uptake was reduced and CAM-type net CO2 uptake in the dark occurred. This was accompanied by nocturnal accumulation of acid in both leaves and stems, also a defining characteristic of CAM. Following rewatering, net CO2 uptake in the dark ceased in shoots, as did nocturnal acidification of the leaves and stems. With this unequivocal demonstration of stress-related reversible, i.e. facultative, induction of CAM, P. cryptopetala becomes the first C3-C4 intermediate species reported to exhibit CAM. Portulaca molokiniensis Hobdy, a C4 species, also exhibited CAM only when subjected to water stress. Facultative CAM has now been demonstrated in all investigated species of Portulaca, which are well sampled from across the phylogeny. This strongly suggests that in Portulaca, a lineage in which species engage predominately in C4 photosynthesis, facultative CAM is ancestral to C4. In a broader context, it has now been demonstrated that CAM can co-exist in leaves that exhibit any of the other types of photosynthesis known in terrestrial plants: C3, C4 and C3-C4 intermediate.

Operating at the very low end of the crassulacean acid metabolism spectrum: Sesuvium portulacastrum (Aizoaceae).

Demonstration of crassulacean acid metabolism (CAM) in species with low usage of this system relative to C3-photosynthetic CO2 assimilation can be challenging experimentally but provides crucial information on the early steps of CAM evolution. Here, weakly expressed CAM was detected in the well-known pantropical coastal, leaf-succulent herb Sesuvium portulacastrum, demonstrating that CAM is present in the Sesuvioideae, the only sub-family of the Aizoaceae in which it had not yet been shown conclusively. In outdoor plots in Panama, leaves and stems of S. portulacastrum consistently exhibited a small degree of nocturnal acidification which, in leaves, increased during the dry season. In potted plants, nocturnal acidification was mainly facultative, as levels of acidification increased in a reversible manner following the imposition of short-term water-stress. In drought-stressed plants, nocturnal net CO2 exchange approached the CO2-compensation point, consistent with low rates of CO2 dark fixation sufficient to eliminate respiratory carbon loss. Detection of low-level CAM in S. portulacastrum adds to the growing number of species that cannot be considered C3 plants sensu stricto, although they obtain CO2 principally via the C3 pathway. Knowledge about the presence/absence of low-level CAM is critical when assessing trajectories of CAM evolution in lineages. The genus Sesuvium is of particular interest because it also contains C4 species.

Two tropical conifers show strong growth and water-use efficiency responses to altered CO2 concentration.

BACKGROUND AND AIMS: Conifers dominated wet lowland tropical forests 100 million years ago (MYA). With a few exceptions in the Podocarpaceae and Araucariaceae, conifers are now absent from this biome. This shift to angiosperm dominance also coincided with a large decline in atmospheric CO2 concentration (ca). We compared growth and physiological performance of two lowland tropical angiosperms and conifers at ca levels representing pre-industrial (280 ppm), ambient (400 ppm) and Eocene (800 ppm) conditions to explore how differences in ca affect the growth and water-use efficiency (WUE) of seedlings from these groups. METHODS: Two conifers (Araucaria heterophylla and Podocarpus guatemalensis) and two angiosperm trees (Tabebuia rosea and Chrysophyllum cainito) were grown in climate-controlled glasshouses in Panama. Growth, photosynthetic rates, nutrient uptake, and nutrient use and water-use efficiencies were measured. KEY RESULTS: Podocarpus seedlings showed a stronger (66 %) increase in relative growth rate with increasing ca relative to Araucaria (19 %) and the angiosperms (no growth enhancement). The response of Podocarpus is consistent with expectations for species with conservative growth traits and low mesophyll diffusion conductance. While previous work has shown limited stomatal response of conifers to ca, we found that the two conifers had significantly greater increases in leaf and whole-plant WUE than the angiosperms, reflecting increased photosynthetic rate and reduced stomatal conductance. Foliar nitrogen isotope ratios (δ15N) and soil nitrate concentrations indicated a preference in Podocarpus for ammonium over nitrate, which may impact nitrogen uptake relative to nitrate assimilators under high ca SIGNIFICANCE: Podocarps colonized tropical forests after angiosperms achieved dominance and are now restricted to infertile soils. Although limited to a single species, our data suggest that higher ca may have been favourable for podocarp colonization of tropical South America 60 MYA, while plasticity in photosynthetic capacity and WUE may help account for their continued persistence under large changes in ca since the Eocene.

Protection by light against heat stress in leaves of tropical crassulacean acid metabolism plants containing high acid levels.

Heat tolerance of plants exhibiting crassulacean acid metabolism (CAM) was determined by exposing leaf sections to a range of temperatures both in the dark and the light, followed by measuring chlorophyll a fluorescence (Fv/Fm and F0) and assessing visible tissue damage. Three CAM species, Clusia rosea Jacq., Clusia pratensis Seem. and Agave angustifolia Haw., were studied. In acidified tissues sampled at the end of the night and exposed to elevated temperatures in the dark, the temperature that caused a 50% decline of Fv/Fm (T50), was remarkably low (40-43°C in leaves of C. rosea). Conversion of chlorophyll to pheophytin indicated irreversible tissue damage caused by malic acid released from the vacuoles. By contrast, when acidified leaves were illuminated during heat treatments, T50 was up to 50-51°C. In de-acidified samples taken at the end of the light period, T50 reached ∼54°C, irrespective of whether temperature treatments were done in the dark or light. Acclimation of A. angustifolia to elevated daytime temperatures resulted in a rise of T50 from ∼54° to ∼57°C. In the field, high tissue temperatures always occur during sun exposure. Measurements of the heat tolerance of CAM plants that use heat treatments of acidified tissue in the dark do not provide relevant information on heat tolerance in an ecological context. However, in the physiological context, such studies may provide important clues on vacuolar properties during the CAM cycle (i.e. on the temperature relationships of malic acid storage and malic acid release).

Light-stimulated heat tolerance in leaves of two neotropical tree species, Ficus insipida and Calophyllum longifolium.

Previous heat tolerance tests of higher plants have been mostly performed with darkened leaves. However, under natural conditions, high leaf temperatures usually occur during periods of high solar radiation. In this study, we demonstrate small but significant increases in the heat tolerance of illuminated leaves. Leaf disks of mature sun leaves from two neotropical tree species, Ficus insipida Willd. and Calophyllum longifolium Willd., were subjected to 15min of heat treatment in the light (500µmol photons m-2s-1) and in the dark. Tissue temperatures were controlled by floating the disks on the surface of a water bath. PSII activity was determined 24h and 48h after heating using chlorophyll a fluorescence. Permanent tissue damage was assessed visually during long-term storage of leaf sections under dim light. In comparison to heat treatments in the dark, the critical temperature (T50) causing a 50% decline of the fluorescence ratio Fv/Fm was increased by ~1°C (from ~52.5°C to ~53.5°C) in the light. Moreover, illumination reduced the decline of Fv/Fm as temperatures approached T50. Visible tissue damage was reduced following heat treatment in the light. Experiments with attached leaves of seedlings exposed to increasing temperatures in a gas exchange cuvette also showed a positive effect of light on heat tolerance.

Thermal tolerance, net CO2 exchange and growth of a tropical tree species, Ficus insipida, cultivated at elevated daytime and nighttime temperatures.

Global warming and associated increases in the frequency and amplitude of extreme weather events, such as heat waves, may adversely affect tropical rainforest plants via significantly increased tissue temperatures. In this study, the response to two temperature regimes was assessed in seedlings of the neotropical pioneer tree species, Ficus insipida. Plants were cultivated in growth chambers at strongly elevated daytime temperature (39°C), combined with either close to natural (22°C) or elevated (32°C) nighttime temperatures. Under both growth regimes, the critical temperature for irreversible leaf damage, determined by changes in chlorophyll a fluorescence, was approximately 51°C. This is comparable to values found in F. insipida growing under natural ambient conditions and indicates a limited potential for heat tolerance acclimation of this tropical forest tree species. Yet, under high nighttime temperature, growth was strongly enhanced, accompanied by increased rates of net photosynthetic CO2 uptake and diminished temperature dependence of leaf-level dark respiration, consistent with thermal acclimation of these key physiological parameters.

Photosynthesis, photoprotection, and growth of shade-tolerant tropical tree seedlings under full sunlight.

High solar radiation in the tropics is known to cause transient reduction in photosystem II (PSII) efficiency and CO(2) assimilation in sun-exposed leaves, but little is known how these responses affect the actual growth performance of tropical plants. The present study addresses this question. Seedlings of five woody neotropical forest species were cultivated under full sunlight and shaded conditions. In full sunlight, strong photoinhibition of PSII at midday was documented for the late-successional tree species Ormosia macrocalyx and Tetragastris panamensis and the understory/forest gap species, Piper reticulatum. In leaves of O. macrocalyx, PSII inhibition was accompanied by substantial midday depression of net CO(2) assimilation. Leaves of all species had increased pools of violaxanthin-cycle pigments. Other features of photoacclimation, such as increased Chl a/b ratio and contents of lutein, ß-carotene and tocopherol varied. High light caused strong increase of tocopherol in leaves of T. panamensis and another late-successional species, Virola surinamensis. O. macrocalyx had low contents of tocopherol and UV-absorbing substances. Under full sunlight, biomass accumulation was not reduced in seedlings of T. panamensis, P. reticulatum, and V. surinamensis, but O. macrocalyx exhibited substantial growth inhibition. In the highly shade-tolerant understory species Psychotria marginata, full sunlight caused strongly reduced growth of most individuals. However, some plants showed relatively high growth rates under full sun approaching those of seedlings at 40 % ambient irradiance. It is concluded that shade-tolerant tropical tree seedlings can achieve efficient photoacclimation and high growth rates in full sunlight.

Transpiration efficiency over an annual cycle, leaf gas exchange and wood carbon isotope ratio of three tropical tree species.

Variation in transpiration efficiency (TE) and its relationship with the stable carbon isotope ratio of wood was investigated in the saplings of three tropical tree species. Five individuals each of Platymiscium pinnatum (Jacq.) Dugand, Swietenia macrophylla King and Tectona grandis Linn. f. were grown individually in large (760 l) pots over 16 months in the Republic of Panama. Cumulative transpiration was determined by repeatedly weighing the pots with a pallet truck scale. Dry matter production was determined by destructive harvest. The TE, expressed as experiment-long dry matter production divided by cumulative water use, averaged 4.1, 4.3 and 2.9 g dry matter kg(-1) water for P. pinnatum, S. macrophylla and T. grandis, respectively. The TE of T. grandis was significantly lower than that of the other two species. Instantaneous measurements of the ratio of intercellular to ambient CO(2) partial pressures (c(i)/c(a)), taken near the end of the experiment, explained 66% of variation in TE. Stomatal conductance was lower in S. macrophylla than in T. grandis, whereas P. pinnatum had similar stomatal conductance to T. grandis, but with a higher photosynthetic rate. Thus, c(i)/c(a) and TE appeared to vary in response to both stomatal conductance and photosynthetic capacity. Stem-wood delta(13)C varied over a relatively narrow range of just 2.2 per thousand, but still explained 28% of variation in TE. The results suggest that leaf-level processes largely determined variation among the three tropical tree species in whole-plant water-use efficiency integrated over a full annual cycle.

Sun-shade patterns of leaf carotenoid composition in 86 species of neotropical forest plants.

A survey of photosynthetic pigments, including 86 species from 64 families, was conducted for leaves of neotropical vascular plants to study sun-shade patterns in carotenoid biosynthesis and occurrence of α-carotene (α-Car) and lutein epoxide (Lx). Under low light, leaves invested less in structural components and more in light harvesting, as manifested by low leaf dry mass per area (LMA) and enhanced mass-based accumulation of chlorophyll (Chl) and carotenoids, especially lutein and neoxanthin. Under high irradiance, LMA was greater and ß-carotene (ß-Car) and violaxanthin-cycle pool increased on a leaf area or Chl basis. The majority of plants contained α-Car in leaves, but the α- to ß-Car ratio was always low in the sun, suggesting preference for ß-Car in strong light. Shade and sun leaves had similar ß,ε-carotenoid contents per unit Chl, whereas sun leaves had more ß,ß-carotenoids than shade leaves. Accumulation of Lx in leaves was found to be widely distributed among taxa: >5 mmol mol Chl-1 in 20% of all species examined and >10 mmol mol Chl-1 in 10% of woody species. In Virola elongata (Benth.) Warb, having substantial Lx in both leaf types, the Lx cycle was operating on a daily basis although Lx restoration in the dark was delayed compared with violaxanthin restoration.

Lutein epoxide cycle, light harvesting and photoprotection in species of the tropical tree genus Inga.

Dynamics and possible function of the lutein epoxide (Lx) cycle, that is, the reversible conversion of Lx to lutein (L) in the light-harvesting antennae, were investigated in leaves of tropical tree species. Photosynthetic pigments were quantified in nine Inga species and species from three other genera. In Inga, Lx levels were high in shade leaves (mostly above 20 mmol mol(-1) chlorophyll) and low in sun leaves. In Virola surinamensis, both sun and shade leaves exhibited very high Lx contents (about 60 mmol mol(-1) chlorophyll). In Inga marginata grown under high irradiance, Lx slowly accumulated within several days upon transfer to deep shade. When shade leaves of I. marginata were briefly exposed to the sunlight, both violaxanthin and Lx were quickly de-epoxidized. Subsequently, overnight recovery occurred only for violaxanthin, not for Lx. In such leaves, containing reduced levels of Lx and increased levels of L, chlorophyll fluorescence induction showed significantly slower reduction of the photosystem II electron acceptor, Q(A), and faster formation as well as a higher level of non-photochemical quenching. The results indicate that slow Lx accumulation in Inga leaves may improve light harvesting under limiting light, while quick de-epoxidation of Lx to L in response to excess light may enhance photoprotection.

Photoprotection, photosynthesis and growth of tropical tree seedlings under near-ambient and strongly reduced solar ultraviolet-B radiation.

Seedlings of two late-successional tropical rainforest tree species, Tetragastris panamensis (Engler) O. Kuntze and Calophyllum longifolium (Willd.), were field grown for 3-4 months at an open site near Panama City (9 degrees N), Panama, under plastic films that either transmitted or excluded most solar UV-B radiation. Experiments were designed to test whether leaves developing under bright sunlight with strongly reduced UV-B are capable of acclimating to near-ambient UV-B conditions. Leaves of T. panamensis that developed under near-ambient UV-B contained higher amounts of UV-absorbing substances than leaves of seedlings grown under reduced UV-B. Photosynthetic pigment composition, content of alpha-tocopherol, CO(2) assimilation, potential photosystem II (PSII) efficiency (evaluated by F(v)/F(m) ratios) and growth of T. panamensis and C. longifolium did not differ between seedlings developed under near-ambient and reduced solar UV-B. When seedlings were transferred from the reduced UV-B treatment to the near-ambient UV-B treatment, a pronounced inhibition of photosynthetic capacity was observed initially in both species. UV-B-mediated inhibition of photosynthetic capacity nearly fully recovered within 1 week of the transfer in C. longifolium, whereas in T. panamensis an about 35% reduced capacity of CO(2) uptake was maintained. A marked increase in UV-absorbing substances was observed in foliage of transferred T. panamensis seedlings. Both species exhibited enhanced mid-day photoinhibition of PSII immediately after being transferred from the reduced UV-B to the near-ambient UV-B treatment. This effect was fully reversible within 1d in T. panamensis and within a few days in C. longifolium. The data show that leaves of these tropical tree seedlings, when developing in full-spectrum sunlight, are effectively protected against high solar UV-B radiation. In contrast, leaves developing under conditions of low UV-B lacked sufficient UV protection. They experienced a decline in photosynthetic competence when suddenly exposed to near-ambient UV-B levels, but exhibited pronounced acclimative responses.

Research note: Large gene family of phosphoenolpyruvate carboxylase in the crassulacean acid metabolism plant Kalanchoe pinnata (Crassulaceae) characterised by partial cDNA sequence analysis.

Clones coding for a 1100-bp cDNA sequence of phosphoenolpyruvate carboxylase (PEPC) of the constitutive crassulacean acid metabolism (CAM) plant Kalanchoe pinnata (Lam.) Pers., were isolated by reverse transcription-polymerase chain reaction (RT-PCR) and characterised by restriction fragment length polymorphism analysis and DNA sequencing. Seven distinct PEPC isogenes were recovered, four in leaves and three in roots (EMBL accession numbers: AJ344052-AJ344058). Sequence similarity comparisons and distance neighbour-joining calculations separate the seven PEPC isoforms into two clades, one of which contains the three PEPCs found in roots. The second clade contains the four isoforms found in leaves and is divided into two branches, one of which contains two PEPCs most similar with described previously CAM isoforms. Of these two isoforms, however, only one exhibited abundant expression in CAM-performing leaves, but not in very young leaves, which do not exhibit CAM, suggesting this isoform encodes a CAM-specific PEPC. Protein sequence calculations suggest that all isogenes are likely derived from a common ancestor gene, presumably by serial gene duplication events. To our knowledge, this is the most comprehensive identification of a PEPC gene family from a CAM plant, and the greatest number of PEPC isogenes reported for any vascular plant to date.

Growth irradiance effects on photosynthesis and growth in two co-occurring shade-tolerant neotropical perennials of contrasting photosynthetic pathways.

Dieffenbachia longispatha (C3) and Aechmea magdalenae (Crassulacean acid metabolism, CAM) are syntopic, neotropical forest perennials in central Panama that are restricted to shaded habitats. This is of particular interest for A. magdalenae because, like other understory CAM bromeliad species, it appears functionally and structurally to be better suited to life in full sun. Growth irradiance (GI) effects on photosynthesis and growth in both species were explored in the context of sun/shade trade-off concepts largely derived from studies of C3 plants. Potted plants were grown outdoors in 1, 55, and 100% full sun for 5 mo under well-watered conditions. While both species grew faster in high compared to low light, maximum relative growth rates (RGR) in full sun were still extremely slow with A. magdalenae showing a RGR approximately half that of D. longispatha. Photosynthetic capacity increased with GI in D. longispatha but not in A. magdalenae. Aechmea magdalenae responded to GI with shifts in the activity of the different CAM phases. Both species were photoinhibited in full sun, but more so in A. magdalenae. Despite possessing many traits considered adaptive in high light, these results suggest that A. magdalenae is unlikely to attain sufficient growth rates to thrive in productive, high-light habitats.

Sudden exposure to solar UV-B radiation reduces net CO(2) uptake and photosystem I efficiency in shade-acclimated tropical tree seedlings.

Tree seedlings developing in the understory of the tropical forest have to endure short periods of high-light stress when tree-fall gaps are formed, and direct solar radiation, including substantial UV light, reaches the leaves. In experiments simulating the opening of a tree-fall gap, the response of photosynthesis in leaves of shade-acclimated seedlings (Anacardium excelsum, Virola surinamensis, and Calophyllum longifolium) to exposure to direct sunlight (for 20-50 min) was investigated in Panama (9 degrees N). To assess the effects of solar UV-B radiation (280-320 nm), the sunlight was filtered through plastic films that selectively absorbed UV-B or transmitted the complete spectrum. The results document a strong inhibition of CO(2) assimilation by sun exposure. Light-limited and light-saturated rates of photosynthetic CO(2) uptake by the leaves were affected, which apparently occurred independently of a simultaneous inhibition of potential photosystem (PS) II efficiency. The ambient UV-B light substantially contributed to these effects. The photochemical capacity of PSI, measured as absorbance change at 810 nm in saturating far-red light, was not significantly affected by sun exposure of the seedlings. However, a decrease in the efficiency of P700 photooxidation by far-red light was observed, which was strongly promoted by solar UV-B radiation. The decrease in PSI efficiency may result from enhanced charge recombination in the reaction center, which might represent an incipient inactivation of PSI, but contributes to thermal dissipation of excessive light energy and thereby to photoprotection.