Alan P. Smith

Growth and photosynthetic response of nine tropical species with long-term exposure to elevated carbon dioxide

SummarySeedlings of nine tropical species varying in growth and carbon metabolism were exposed to twice the current atmospheric level of CO2 for a 3 month period on Barro Colorado Island, Panama. A doubling of the CO2 concentration resulted in increases in photosynthesis and greater water use efficiency (WUE) for all species possessing C3 metabolism, when compared to the ambient condition. No desensitization of photosynthesis to increased CO2 was observed during the 3 month period. Significant increases in total plant dry weight were also noted for 4 out of the 5 C3 species tested and in one CAM species, Aechmea magdalenae at high CO2. In contrast, no significant increases in either photosynthesis or total plant dry weight were noted for the C4 grass, Paspallum conjugatum. Increases in the apparent quantum efficiency (AQE) for all C3 species suggest that elevated CO2 may increase photosynthetic rate relative to ambient CO2 over a wide range of light conditions. The response of CO2 assimilation to internal Ci suggested a reduction in either the RuBP and/or Pi regeneration limitation with long term exposure to elevated CO2. This experiment suggests that: (1) a global rise in CO2 may have significant effects on photosynthesis and productivity in a wide variety of tropical species, and (2) increases in productivity and photosynthesis may be related to physiological adaptation(s) to increased CO2.

Water potential gradients for gaps and slopes in a Panamanian tropical moist forest's dry season

Soil water potentials were measured weekly by psychrometers at 20 cm depth during the dry season in a tropical moist forest on Barro Colorado Island, Panama. There was a persistent gradient of decreasing soil moisture from gap centre to gap edge to adjacent understorey at both a large and a small gap. On both a north-south and an east-west transect, the soil was drier (water potentials were more negative) on an upland surface than on moderate slopes. This trend was reflected in the predawn, total water potentials of shallow-rooted Psychotria horizontalis and deep-rooted Trichilia tuberculata measured in the understorey during the last two months of the dry season. P. horizontalis from the wettest sites on the transects had higher osmotic potentials at full hydration and at zero turgor, indicating less drought resistance than for conspecifics from the driest sites.

Analysis of forest light environments part I. Computerized estimation of solar radiation from hemispherical canopy photographs

Abstract A microcomputer based system for the semi-automatic analysis of hemispherical canopy photos is described and evaluated. Images are transferred from film by a video camera to a digitizer for processing. Our program calculates diffuse and direct site factors (proportions of unobstructed radiation) according to Andersons methods. These factors are used to estimate absolute amounts of radiation on a horizontal, subcanopy surface from measured or estimated values in the open. Additional indices calculated include the proportion of open canopy in the entire image and in the solar band, the cardinal octants, and 20 concentric rings plus a frequency histogram of potential sunfleck duration. About 20 photos can be processed in an hour. The program was designed for large-scale studies correlating plant biology and distribution or animal behavior with radiation regimes where time scales of several weeks or longer are of interest. Consistency and accuracy of estimates of mean daily global radiation by different operators permit meaningful intersite comparisons.

Comparative physiology and demography of three Neotropical forest shrubs: alternative shade-adaptive character syndromes

A suite of functionally-related characters and demography of three species of Neotropical shadeadapted understory shrubs (Psychotria, Rubiaceae) were studied in the field over five years. Plants were growing in large-scale irrigated and control treatments in gaps and shade in old-growth moist forest at Barro Colorado Island, Panama. Irrigation demonstrated that dry-season drought limited stomatal conductance, light saturated photosynthesis, and leaf longevity in all three species. Drought increased mortality of P. furcata. In contrast, irrigation did not affect measures of photosynthetic capacity determined with an oxygen electrode or from photosynthesis-CO2 response curves in the field. Drought stress limited field photosynthesis and leaf and plant survivorship without affecting photosynthetic capacity during late dry season. Leaves grown in high light in naturally occurring treefall gaps had higher photosynthetic capacity, dark respiration and mass per unit area than leaves grown in the shaded understory. P. furcata had the lowest acclimation to high light for all of these characters, and plant mortality was greater in gaps than in shaded understory for this species. The higher photosynthetic capacity of gap-grown leaves was also apparent when photosynthetic capacity was calculated on a leaf mass basis. Acclimation to high light involved repackaging (higher mass per unit leaf area) as well as higher photosynthetic capacity per unit leaf mass in these species. The three species showed two distinct syndromes of functionally-related adaptations to low light. P. limonensis and P. marginata had high leaf longevity (∼3 years), high plant survivorship, low leaf nitrogen content, and high leaf mass per unit area. In contrast, P. furcata had low leaf survivorship (∼1 year), high plant mortality (77–96% in 39 months), low leaf mass per unit area, high leaf nitrogen content, and the highest leaf area to total plant mass; the lowest levels of shelf shading, dark respiration and light compensation; and the highest stem diameter growth rates. This suite of characters may permit higher whole-plant carbon gain and high leaf and population turnover in P. furcata. Growth in deep shade can be accomplished through alternative character syndromes, and leaf longevity may not be correlated with photosynthetic capacity in shade adapted plants.

Drought acclimation among tropical forest shrubs (Psychotria, Rubiaceae)

SummaryMechanisms of dry-season drought resistance were evaluated for five evergreen shrubs (Psychotria, Rubiaceae) which occur syntopically in tropical moist forest in central Panama. Rooting depths, leaf conductance, tissue osmotic potentials and elasticity, and the timing of leaf production were evaluated. From wet to dry season, tissue osmotic potentials declined and moduli of elasticity increased in four and five species, respectively. Irrigation only affected osmotic adjustment by P. furcata. The other seasonal changes in leaf tissue properties represented ontogenetic change. Nevertheless, they made an important contribution to dry-season turgor maintenance. Small between-year differences in dry season rainfall had large effects on plant water status. In 1986, 51 mm of rain fell between 1 January and 31 March, and pre-dawn turgor potentials averaged <0.1 MPa for all five Psychotria species in March (Wright 1991). In 1989, 111 mm of rain fell in the same period, pre-dawn turgor potentials averaged from 0.75 to 1.0 MPa for three of the species in April, and only P. chagrensis lost turgor. The relation between leaf production and drought differed among species. P. limonensis was buffered against drought by the lowest dry-season conductances and the deepest roots (averaging 244% deeper than its congeners) and was the only species to produce large numbers of leaves in the dry season. P. chagrensis was most susceptible to drought, and leaf production ceased as turgor loss developed. For the other species, water stress during severe dry seasons may select against dry-season leaf production.

Ecology of a leaf color polymorphism in a tropical forest species: habitat segregation and herbivory

SummaryByttneria aculeata (Sterculiaceae), a subcanopy liane with a shrubby juvenile form has two distinct leaf color morphs in juvenile plants- a given juvenile has plain green leaves or leaves with whitish variegation. Both forms occur together in the forest and in clearings; however, the variegated morph is more common in open sites, and the plain morph predominates in the forest. Percent variegation per leaf for variegated plants increased from closed to open sites. Measurements of growth support the idea that variegation is favored in open habitats. Within a given habitat, rate of herbivory by leaf miners on a given morph increases with increasing relative frequency of that morph; however, at a given relative frequency, the variegated morph is less heavily attacked than is the plain morph when it occurs elsewhere at that same frequency.

Spatial autocorrelation of solar radiation in a tropical moist forest understory.

Abstract Several indices of solar radiation were estimated from hemispherical canopy photographs of mature forest on Barro Colorado Island, Panama. The photographs were taken during two wet seasons at 400 uniformly spaced stations on two 1-km transects. In an El Nino year, mean daily global radiation was positively autocorrelated from 2.5 to 12.5–22.5 m when light penetration was abnormally high due to drought-induced leaf fall in the previous dry season. In a more typical year, weak positive spatial autocorrelation occurred only between adjacent sites 2.5 m apart. Other radiation indices, including the proportion of unobstructed sky in the canopy on the solar path, showed correlograms nearly identical to that of global radiation in the El Nino year. Spectral analysis revealed a period of 125 m in mean daily global radiation along a transect in the El Nino, but not in the normal year.

Growth and photosynthesis of Aechmea magdalenae , a terrestrial CAM plant in a tropical moist forest, Panama

Aechmea magdalenae is a terrestrial bromeliad that dominates areas of forest understorey on Barro Colorado Island, Panama. Nocturnal CO2 uptake via crassulacean acid metabolism was the primary means of carbon gain under well-watered conditions and all light regimes. The ability to maintain a positive carbon balance under conditions of very low light was demonstrated by laboratory measurements of photosynthesis and forest measurements of growth. Low-light-grown juvenile rosettes had the same daily net assimilation whether tested at photon flux densities of 15 or 300 ,umol m-2 s-1. Growth rates of rosettes in treefall gaps were similar to those of plants in closed canopy forest. Growth rates of forest plants were increas- ingly correlated with canopy openness as the wet season progressed due to increased growth by gap plants, suggesting that water availability rather than light may limit growth during the annual dry season.

The cost of reproduction in Senecio keniodendron, a giant rosette species of Mt. Kenya

SummaryA three year study of Senecio keniodendron (Compositae), a giant rosette species of the alpine zone of Mt. Kenya, demonstrated that individuals which reproduce are more likely to die, and less likely to reproduce in the future if they do survive, than are vegetative individuals of the same size. However, if an individual reproduces, survives and reproduces again, then it produces more seeds during the second reproductive episode than does a plant of the same height reproducing for the first time, because reproduction is followed by production of lateral rosettes, increasing the number of potentially-reproductive rosettes per plant.Slow-growing rosettes are less likely to reproduce than fast-growing rosettes. For rosettes which do reproduce, rosette size and rate of leaf production, measured before reproduction begins, are good predictors of fecundity.

Morphological and physiological correlates of niche breadth in two species of Espeletia (Compositae) in the Venezuelan Andes

SummaryWe compared adaptive strategies in two plants of Venezuelan páramos (alpine areas): the widely distributed, caulescent, and pubescent Espeletia schultzii Wedd. with the acaulescent, nearly glabrous E. atropurpurea A.C. Smith which is restricted to mesic sites just above treeline. Both species occur together at 3,450 m, near treeline.Physiologically, E. schultzii was more drought resistant than E. atropurpurea, and was better adapted for carbon dioxide fixation under low temperatures. The densely pubescent leaves of E. schultzii are highly reflective; this increases the intensity of light needed for photosynthetic saturation and influences leaf temperature. Leaf pubescence may reduce the level of insect predation.Measurements of leaf productivity indicate higher values for E. atropurpurea during the rainy season and higher values for E. schultzii during the dry season. However, annual values of leaf productivity are similar for both species.Benefits of specialization in E. atropurpurea include reduced costs for stem and leaf hair production, higher growth rates during the rainy season and the ability to grow beneath canopies of some larger arborescent species. Costs of specialization include lower growth rates during the dry season, great susceptibility to insect predation and restriction to low elevation, mesic sites.