Morphological diversity of the velamen radicum in the genus Anthurium (Araceae).

Epiphytes develop anatomical features to improve efficiency of the uptake of water and nutrients, such as absorptive foliar scales or a velamen radicum. Despite substantial studies on the occurrence, morphology, development and phylogeny of the velamen, most of the available literature is focused on Orchidaceae, making current knowledge on velamen clearly biased. A recent publication firmly established that velamina are common in Anthurium species. Thus, this study provides further insights by describing velamen morphological characteristics of Anthurium species and classifying them into different velamen types. Furthermore, we investigate if the different velamen morphological traits are clade-specific and phylogenetically conserved within the genus. Using SEM, we performed a morphological study on 89 Anthurium species, describing six micromorphological traits of velamen and exodermis, following traits used to classify Orchidaceae velamen by Porembski & Barthlott (1988). We distinguished nine velamen types, including two that are unique to Anthurium and not similar to any type found in Orchidaceae. Comparing velamen morphology within the phylogenetic tree of Anthurium revealed clear phylogenetic signals. This study provides detailed morphological descriptions among 89 species of Anthurium from the Araceae, and substantially broadens our knowledge of this tissue. However, velamen function has been even less studied, with hardly anything known about functional significance of having secondary cell wall thickening and perforations on velamen cell walls. Therefore, a logical next step would be to connect these anatomical features to their functions.

Temperature dependence of germination and growth in Anthurium (Araceae).

By the year 2100, temperatures are predicted to increase by about 6 °C at higher latitudes and about 3 °C in the tropics. In spite of the smaller increase in the tropics, consequences may be more severe because the climatic niches of tropical species are generally assumed to be rather narrow due to a high degree of climate stability and higher niche specialisation. However, rigorous data to back up this notion are rare. We chose the megadiverse genus Anthurium (Araceae) for study. Considering that the regeneration niche of a species is crucial for overall niche breadth, we focused on the response of germination and early growth through a temperature range of 24 °C of 15 Anthurium species, and compared the thermal niche breadth (TNB) with the temperature conditions in their current range, modelled from occurrence records. Surprisingly, an increase of 3 °C would lead to a larger overlap of TNB of germination and modelled in situ temperature conditions, while the overlap of TNB of growth with in situ conditions under current and future conditions is statistically indistinguishable. We conclude that future temperatures tend to be closer to the thermal optima of most species. Whether this really leads to an increase in performance depends on other abiotic and biotic factors, most prominently potentially changing precipitation patterns.

Epiphytic bromeliads in a changing world: the effect of elevated CO2 and varying water supply on growth and nutrient relations.

Global climate change is likely to impact all plant life. Vascular epiphytes represent a life form that may be affected more than any other by possible changes in precipitation leading to water shortage, but negative effects of drought may be mitigated through increasing levels of atmospheric CO2 . We studied the response of three epiphytic Aechmea species to different CO2 and watering levels in a full-factorial climate chamber study over 100 days. All species use crassulacean acid metabolism (CAM). Response variables were relative growth rate (RGR), nocturnal acidification and foliar nutrient levels (N, P, K, Mg). Both elevated CO2 and increased water supply stimulated RGR, but the interaction of the two factors was not significant. Nocturnal acidification was not affected by these factors, indicating that the increase in growth in these CAM species was due to higher assimilation in the light. Mass-based foliar nutrient contents were consistently lower under elevated CO2 , but most differences disappeared when expressed on an area basis. Compared to previous studies with epiphytes, in which doubling of CO2 increased RGR, on average, by only 14%, these Aechmea species showed a relatively strong growth stimulation of up to +61%. Consistent with earlier findings with other bromeliads, elevated CO2 did not mitigate the effect of water shortage.

Latitudinal variation in the degree of crassulacean acid metabolism in Puya chilensis.

Crassulacean acid metabolism (CAM) is a photosynthetic pathway found in many plant species from arid and semiarid environments. Few studies aiming to characterise plant species as CAM or C3 account for inter-population differences in photosynthetic pathway, often relying on samples taken from herbarium material and/or a single plant or population. This may be especially problematic for species growing under contrasting climate conditions, as is the case for species with a wide geographic range. We used Puya chilensis, a species previously reported as CAM and C3, to study among-population variation in expression of the CAM pathway within its distribution range, which spans a significant climate gradient. We carried out a wide sampling scheme, including five populations and a combination of analytical methods (quantification of nocturnal acidification and stable isotope measurements). The study populations of P. chilensis encompass the entire latitudinal distribution range, from semi-arid to temperate oceanic climates. Our results indicate that CAM decreased with latitude. However, even in the southern (wetter) populations, where δ13C values were indicative of C3 metabolism, we found some nocturnal acidification. We stress the value of using two methods along with the use of samples from different populations, as this allows more reliable conclusions on the photosynthetic pathway for 'probable' CAM species that face varying climate conditions within their distribution ranges.

The temperature acclimation potential of tropical bryophytes.

Bryophyte biomass and diversity in tropical moist forests decrease dramatically from higher altitudes towards the lowlands. High respiratory carbon losses at high temperatures may partly explain this pattern, if montane species are unable to acclimatise their metabolic rates to lowland temperatures. We transplanted ten bryophyte species from two altitudes (1200 and 500 m a.s.l.) to lower (warmer) altitudes (500 m and sea level) in Panama. We studied short-term temperature acclimation of CO2 exchange for 2.5 months, and survival and growth for 21 months following transplantation. Short-term acclimation did not occur, and on a longer time scale mortality was highest and growth lowest in the transplanted samples. A few transplanted samples of most species, however, survived the whole experiment and finished with growth rates similar to controls. This recovery of growth rate suggests temperature acclimation, in spite of no measurable metabolic changes in smaller random samples. This acclimation even compensated for shorter periods of CO2 uptake due to more rapid drying. Nevertheless, these species are not abundant in lowland forests, perhaps due to dispersal or establishment limitation. The apparent heterogeneity of the acclimation potential within species may allow populations to adapt locally and avoid being forced uphill under climatic warming.

Phytic acid in green leaves.

Phytic acid or phytate, the free-acid form of myo-inositolhexakiphosphate, is abundant in many seeds and fruits, where it represents the major storage form of phosphorus. Although also known from other plant tissues, available reports on the occurrence of phytic acid, e.g. in leaves, have never been compiled, nor have they been critically reviewed. We found 45 published studies with information on phytic acid content in leaves. Phytic acid was almost always detected when studies specifically tried to detect it, and accounted for up to 98% of total P. However, we argue that such extreme values, which rival findings from storage organs, are dubious and probably result from measurement errors. Excluding these high values from further quantitative analysis, foliar phytic acid-P averaged 2.3 mg·g(-1) , and represented, on average, 7.6% of total P. Remarkably, the ratio of phytic acid-P to total P did not increase with total P, we even detected a negative correlation of the two variables within one species, Manihot esculenta. This enigmatic finding warrants further attention.

What is the proximate cause for size-dependent ecophysiological differences in vascular epiphytes?

Size-related variation in physiological parameters as diverse as photosynthetic capacity, abscisic acid relationships or the relative water deficit at stomatal closure have been reported for a large number of vascular epiphytes, but the proximate mechanism behind these observations has not been identified. We test four possible reasons for size-related changes in photosynthetic capacity, leaf-N content and specific leaf area: (i) plant size itself, (ii) plant age or developmental stage, (iii) previous nutrition, or (iv) previous water regime. A suite of study species and approaches were used: a 'natural experiment' with the orchid Polystachya foliosa; an experimental field study with another orchid, Dimerandra emarginata; and a study under controlled conditions with the tank bromeliad, Vriesea sanguinolenta. Neither size, age nor differences in water supply caused differences in leaf N and photosynthetic capacity, while low supply of nutrients yielded, and high supply with nutrients completely removed, size-related trends. The observed size-related trends are thus a consequence of in situ differences in nutrient acquisition. Arguably, the improved nutrient status of larger plants under natural conditions results from larger tanks, holding moisture for increasingly longer intervals, which allows longer periods of decomposition of detritus and of nutrient uptake.

Growth in epiphytic bromeliads: response to the relative supply of phosphorus and nitrogen.

Insufficient nitrogen (N) and phosphorus (P) frequently limit primary production. Although most nutrient studies on vascular epiphytes have focused on N uptake, circumstantial evidence suggests that P rather than N is the most limiting element for growth in this plant group. We directly tested this by subjecting a total of 162 small individuals of three bromeliad species (Guzmania monostachia, Tillandsia elongata, Werauhia sanguinolenta) to three N and three P levels using a full-factorial experimental design, and determined relative growth rates (RGR) and nutrient acquisition over a period of 11 weeks. Both N and P supply had a significant effect on RGR, but only tissue P concentrations were correlated with growth. Uptake rates of N and P, in contrast, were not correlated with RGR. Increased nutrient supply led to an up to sevenfold increase in tissue P concentration compared to natural conditions, while concentrations of N hardly changed or even decreased. All treatment combinations, even at the lowest experimental P supply, led to decreased N:P ratios. We conclude that P is at least as limiting as N for vegetative function under natural conditions in these epiphytic bromeliads. This conclusion is in line with the general notion of the prevalence of P limitation for the functioning of terrestrial vegetation in the tropics.

A hierarchical framework for investigating epiphyte assemblages: networks, meta-communities, and scale.

Epiphytes are an important component of many forested ecosystems, yet our understanding of epiphyte communities lags far behind that of terrestrial-based plant communities. This discrepancy is exacerbated by the lack of a theoretical context to assess patterns in epiphyte community structure. We attempt to fill this gap by developing an analytical framework to investigate epiphyte assemblages, which we then apply to a data set on epiphyte distributions in a Panamanian rain forest. On a coarse scale, interactions between epiphyte species and host tree species can be viewed as bipartite networks, similar to pollination and seed dispersal networks. On a finer scale, epiphyte communities on individual host trees can be viewed as meta-communities, or suites of local epiphyte communities connected by dispersal. Similar analytical tools are typically employed to investigate species interaction networks and meta-communities, thus providing a unified analytical framework to investigate coarse-scale (network) and fine-scale (meta-community) patterns in epiphyte distributions. Coarse-scale analysis of the Panamanian data set showed that most epiphyte species interacted with fewer host species than expected by chance. Fine-scale analyses showed that epiphyte species richness on individual trees was lower than null model expectations. Therefore, epiphyte distributions were clumped at both scales, perhaps as a result of dispersal limitations. Scale-dependent patterns in epiphyte species composition were observed. Epiphyte-host networks showed evidence of negative co-occurrence patterns, which could arise from adaptations among epiphyte species to avoid competition for host species, while most epiphyte meta-communities were distributed at random. Application of our "meta-network" analytical framework in other locales may help to identify general patterns in the structure of epiphyte assemblages and their variation in space and time.

No down-regulation of leaf photosynthesis in mature forest trees after three years of exposure to elevated CO2.

The photosynthetic responses of six species of mature forest trees to long-term exposure to elevated CO2 (ca. 530 ppm) were determined at the Swiss Canopy Crane (SCC) site near Basel, Switzerland. In the third year of growth in elevated CO2, using web-FACE technology, net photosynthesis (As) in fully sunlit, upper canopy foliage was stimulated by ca. 40% compared to ambient controls. This enhancement did not differ from the instantaneous increase in As found in ambient-grown leaves that were temporarily measured at elevated CO2. A complete lack of down-regulation of photosynthesis was found in all species and in both the early and the late growing season. Neither was leaf nitrogen content significantly affected by long-term exposure to elevated CO2. Our results document a persistent enhancement in leaf level photosynthesis in response to elevated CO2 in mature forest trees over a period of three years. Circumstantial evidence suggests that the additional assimilates feed into large sinks other than stem and shoot growth.

Effects of natural intensities of visible and ultraviolet radiation on epidermal ultraviolet screening and photosynthesis in grape leaves.

Grape (Vitis vinifera cv Silvaner) vine plants were cultivated under shaded conditions in the absence of ultraviolet (UV) radiation in a greenhouse, and subsequently placed outdoors under three different light regimes for 7 d. Different light regimes were produced by filters transmitting natural radiation, or screening out the UV-B (280-315 nm), or screening out the UV-A (315-400 nm) and the UV-B spectral range. During exposure, synthesis of UV-screening phenolics in leaves was quantified using HPLC: All treatments increased concentrations of hydroxycinnamic acids but the rise was highest, reaching 230% of the initial value, when UV radiation was absent. In contrast, UV-B radiation specifically increased flavonoid concentrations resulting in more than a 10-fold increase. Transmittance in the UV of all extracted phenolics was lower than epidermal UV transmittance determined fluorimetrically, and the two parameters were curvilinearly related. It is suggested that curvilinearity results from different absorption properties of the homogeneously dissolved phenolics in extracts and of the non-homogeneous distribution of phenolics in the epidermis. UV-B-dependent inhibition of maximum photochemical yield of photosystem II (PSII), measured as variable fluorescence of dark-adapted leaves, recovered in parallel to the buildup of epidermal screening for UV-B radiation, suggesting that PSII is protected against UV-B damage by epidermal screening. However, UV-B inhibition of CO(2) assimilation rates was not diminished by efficient UV-B screening. We propose that protection of UV-B inactivation of PSII is observed because preceding damage is efficiently repaired while those factors determining UV-B inhibition of CO(2) assimilation recover more slowly.

The physiological ecology of vascular epiphytes: current knowledge, open questions.

The current knowledge of the physiological ecology of vascular epiphytes is reviewed here with an emphasis on the most recent literature. It is argued that by far the most relevant abiotic constraint for growth and vegetative function of vascular epiphytes is water shortage, while other factors such as nutrient availability or irradiation, are generally of inferior importance. However, it is shown that the present understanding of epiphyte biology is still highly biased, both taxonomically and ecologically, and it is concluded that any generalizations are still preliminary. Future studies should include a much wider range of taxa and growing sites within the canopy to reach a better understanding how abiotic factors are limiting epiphyte growth and survival which, in turn, should affect epiphyte community composition. Finally, a more integrative approach to epiphyte biology is encouraged: physiological investigations should be balanced by studies of other possible constraints, for example, substrate instability, dispersal limitation, competition or herbivory.

Small plants, large plants: the importance of plant size for the physiological ecology of vascular epiphytes.

Recently, a number of publications have reported that many physiological properties of vascular epiphytes are a function of plant size. This short review will summarize what is known to date about this phenomenon, describe the possible mechanism and will discuss the consequences for the present understanding of epiphyte biology. Size-related changes are also known from other plant groups and it is argued that close attention should be paid to the size of the organisms under study in order to understand the performance and survival of a species in the field. In the light of these findings, the results of many earlier studies on epiphyte ecophysiology are now difficult to interpret because essential information on the size of the specimens used is missing.

Ecophysiological consequences of differences in plant size: abscisic acid relationships in the epiphytic orchid Dimerandra emarginata.

Changes in abscisic acid (ABA) content and several water relations parameters were studied in the epiphytic orchid Dimerandra emarginata in the tropical lowlands of Barro Colorado Island, Panama. Similar to previous observations of size dependency of physiological parameters in vascular epiphytes, we found significant differences in ABA accumulation (both in terms of temporal patterns and amount) in small and large plants under drought stress. The highest values of ABA content in leaves, stems, and roots were found in the smaller specimens, reaching the levels of poikilohydric resurrection plants. Substantial accumulation of ABA occurred after stomatal closure, indicating other functions of ABA than stomatal regulation in this bark epiphyte.

6-Hydroxyluteolin-7-O-(1''-alpha-rhamnoside) from Vriesea sanguinolenta Cogn. and Marchal (Bromeliaceae).

The isolation of 6-hydroxyluteolin-7-O-(1"-alpha-rhamnoside) from the Central American epiphyte Vriesea sanguinolenta Cogn. and Marchal (Bromeliaceae) is described here. Its stereostructure was established by spectroscopic methods and an X-ray structure analysis of its hepta-O-acetyl derivative. This flavonoid glycoside had previously been reported from some Teucrium species (Labiatae), yet without sufficient physical data and spectroscopic evidence.

Hydraulic architecture and water relations of a flood-tolerant tropical tree, Annona glabra.

Hydraulic architecture parameters, water relation parameters and wood anatomy were studied in roots and shoots of the flood-tolerant tree Annona glabra L. on Barro Colorado Island, Panama. Hydraulic conductivity, leaf specific conductivity, and Huber value were similar to the corresponding values for tree species living in non-flooded habitats. The vulnerability of stems to loss of hydraulic conductivity resulting from embolism was low (50% loss of conductivity at -3.3 MPa). The lowest leaf water potential measured in the field was about -1.0 MPa, indicating that A. glabra has a large margin of safety from embolism, which may provide protection against rare drought events, or may be an adaptation to brackish mangrove habitats. Low absolute conductivity of roots was compensated for by an increase in the number of roots. More than two-thirds of whole-plant resistance to water flow was located in the roots.

The occurrence of crassulacean acid metabolism among vascular epiphytes from Central Panama.

The occurrence of crassulacean acid metabolism (CAM) among the epiphyte flora of the lowland forest on Barro Colorado Island (BCJ), Panama, was investigated. A total of 116 species was included, i.e. about 2/3 of the known epiphyte taxa. As judged from the carbon isotope ratios and the absence of Kranz anatomy, indications of CAM were found in 29 species of three families, Orchidaceae (20), Bromeliaceae (7), and Cactaceae (2). We estimate that about 25% of the epiphyte flora of BCI are CAM plants. CAM was most prevalent in exposed sites, but even in the understorey two epiphyte species engage in CAM.

Photosynthesis of a tropical canopy tree, Ceiba pentandra, in a lowland forest in Panama.

Diel (24 h) courses of CO(2) and water-vapor exchange of Ceiba pentandra (L.) Gaertn. (Bombacaceae) were studied under natural tropical conditions in the semi-evergreen moist forest of Barro Colorado Island, Panama. Measurements were conducted from early February 1991 (dry season), shortly after new leaves emerged, until mid-October 1991 (wet season), when leaves were shed. Rates of net CO(2) uptake were significantly higher in the dry season than in the wet season, and showed a linear decrease with leaf age. Leaf nitrogen concentrations and contents also decreased with age. Our estimate of annual carbon gain (2640 g CO(2) m(-2) year(-1) or 21 g CO(2) g(DW) (-1) year(-1)) is considerably higher than estimates available for temperate forest trees.

Short-Term Regulation of Crassulacean Acid Metabolism Activity in a Tropical Hemiepiphyte, Clusia uvitana.

Diel courses of net CO2 exchange of leaves were studied in Clusia uvitana (Clusiaceae), a tropical Crassulacean acid metabolism (CAM) hemiepiphyte, growing in the crown of a 47-m tall kapok tree on Barro Colorado Island, Panama. Measurements on days without precipitation showed that net uptake of atmospheric CO2 occurred at night, a feature of CAM, as well as in the early morning and late afternoon. During 36 h of almost continuous rainfall, nocturnal net CO2 uptake was abolished and the diel pattern of net CO2 exchange became similar to that of a C3 plant. Exposing well-watered, potted plants of Clusia in the laboratory to temperatures and photosynthetic photon flux densities similar to those during the tropical rainstorm also abolished nocturnal net CO2 uptake. In contrast, Kalanchoe pinnata (Crassulaceae), an obligate CAM plant, still showed net CO2 dark fixation following the same low-light and moderate-temperature conditions, albeit at decreased rates. During these 12-h photoperiods, titratable acidity in Clusia increased slightly above its high level measured at the end of the previous dark period, whereas in Kalanchoe, the acid content decreased by about 40%. A survey among outer canopy leaves of Clusia on Barro Colorado Island showed that leaves that exhibited little or no nocturnal acidification maintained high levels of H+ at dawn and dusk. Progressively lower levels of H+ at dusk were accompanied by progressively higher nocturnal increases in H+. The data suggest that in C. uvitana the rapid switching between CAM- and C3-type carbon fixation that may occur within 24 h in response to environmental changes is controlled by the acidity status of the leaves in the light. Nocturnal CO2 fixation is enhanced by conditions that decrease the organic acid content during the light period.