David D. Ackerly

Interpreting phenotypic variation in plants

Plant ecologists and evolutionary biologists frequently examine patterns of phenotypic variation across variable environments or genetic identities. Too often, we ignore the fact that most phenotypic traits change throughout growth and development of individual plants, and that rates of growth and development are highly variable. Plants growing in different environments are likely to grow at different rates, and will be of different sizes and stages of development at a particular age. When we compare plants as a function of plant size or developmental stage, as well as a function of age, we broaden our understanding of phenotypic variation between plants.

Leaf dynamics, self-shading and carbon gain in seedlings of a tropical pioneer tree

We examined leaf dynamics and leaf age gradients of photosynthetic capacity and nitrogen concentration in seedlings of the tropical pioneer tree, Heliocarpus appendiculatus, grown in a factorial design under controlled conditions with two levels each of nutrients, ambient light (light levels incident above the canopy), and self-shading (the gradient of light levels from upper to lower leaves on the shoot). Correlations among these parameters were examined in order to determine the influence of self-shading, and the regulation of standing leaf numbers, on leaf longevity and its association with leaf photosynthetic capacity. Leaf longevity and the number of leaves on the main shoot were both reduced in high light, while in the low light environment, they were reduced in the steeper self-shading gradient. In high nutrients, leaf longevity was reduced whereas leaf number increased. Leaf initiation rates were higher in the high nutrient treatment but were not influenced by either light treatment. Maximum-light saturated photosynthetic rate, on an area basis, was greater in the high light and nutrient treatments, while the decline in photosynthetic capacity in realtion to leaf position on the shoot was more rapid in high light and in low nutrients. Leaf longevity was negatively correlated among treatments with initial photosynthetic capacity. The leaf position at which photosynthetic capacity was predicted to reach zero was positively correlated with the number of leaves on the shoot, supporting the hypothesis that leaf numbers are regulated by patterns of self-shading. The negative association of longevity and initial photosynthetic capacity apparently arises from different associations among gradients of photosynthetic capacity, leaf numbers and leaf initiation rates in relation to light and nutrient availability. The simultaneous consideration of age and position of leaves illuminates the role of self-shading as an important factor influencing leaf senescence and canopy structure and dynamics.

Self-shading, carbon gain and leaf dynamics: a test of alternative optimality models

Abstract A simple model of shoot-level carbon gain is presented addressing the optimal number and life span of leaves in relation to alternative optimality criteria: (1) maximizing carbon export from the shoot, or (2) maximizing the rate of leaf production at the shoot tip. Additionally, the processes that cause declining assimilation with leaf age are considered in relation to (1) leaf position on the shoot (e.g., self-shading) versus (2) leaf age per se. Using these alternative scenarios, only a model based on position-dependent assimilation and maximization of leaf production rates resulted in quantitative predictions for all aspects of leaf dynamics on the shoot (i.e., leaf number, life span, and birth rate), while other approaches predicted that one or more parameters would be infinite. This formulation of the model also predicted that leaves should be maintained on the shoot until the diurnal carbon balance declines to zero, in contrast with other scenarios which predict that leaves should be shed while maintaining a positive carbon balance. Predictions of the model were supported by the results of a field study of carbon gain and leaf dynamics in saplings of three species of tropical pioneer trees (Carica papaya, Cecropia obtusifolia, and Hampea nutricia) which differ in the number of leaves per shoot. The results illustrate that in these fast-growing plants, leaf production and height growth may be more appropriate measures of performance than net carbon export from the shoot, and suggest that leaf senescence is primarily a function of the position of a leaf within the canopy, rather than its chronological age.

CO2 ELEVATION, CANOPY PHOTOSYNTHESIS, AND OPTIMAL LEAF AREA INDEX

We studied the effects of CO2 elevation on leaf and canopy photosynthesis and optimal leaf area index (LAI) for stands of the annual species Abutilon theophrasti and Ambrosia artemisiifolia. Leaf photosynthesis was modeled as a function of photosyn- thetic photon flux density (PPFD) and nitrogen content per unit leaf area (NL). There was a curvilinear relationship between the light-saturated rates of leaf photosynthesis (Pmax) and NL. CO2 elevation significantly increased Pmax as a function of NL in both species. Dark respiration (Rd) was linearly correlated with NL. CO2 elevation slightly but significantly increased Rd in Abutilon, while it had no significant effect on Rd in Ambrosia. The initial slope of a light-response curve was determined from quantum yield (4gabs) multiplied by leaf absorptance and then calibrated against NL. Daily canopy photosynthesis, calculated by integration of leaf photosynthesis with the actual distribution of leaf area, leaf N, and PPFD within a canopy, showed fairly good agreement with the canopy photosynthesis estimated from growth analysis. CO2 elevation increased canopy photosynthesis by 30- 50%. Based on the leaf photosynthesis model for Abutilon, we calculated daily canopy photosynthesis for a given LAI and N availability, in which N was assumed to be distributed optimally within a leaf canopy to maximize daily canopy photosynthesis. An optimal LAI to maximize daily canopy photosynthesis was obtained for each level of N availability and this optimum increased with increasing N availability. Contrary to the often predicted increase in LAI with CO2 elevation, the optimum LAI did not increase at high CO2 when N availability was limited. Two factors were suggested to be involved in counteracting the increase in LAI in a high-CO2 world. One is the higher 4abs of plants grown in elevated C02, which makes leaves in the canopy more N limited, favors higher NL and thus lowers optimal LAI. The other is the higher Rd in elevated C02, which leads to higher light compensation points, and lowers optimal LAI.

CO2 ELEVATION, CANOPY PHOTOSYNTHESIS, ANDOPTIMAL LEAF AREA INDEX

We studied the effects of CO2 elevation on leaf and canopy photosynthesis and optimal leaf area index (LAI) for stands of the annual species Abutilon theophrasti and Ambrosia artemisiifolia. Leaf photosynthesis was modeled as a function of photosynthetic photon flux density (PPFD) and nitrogen content per unit leaf area (NL). There was a curvilinear relationship between the light-saturated rates of leaf photosynthesis (Pmax) and NL. CO2 elevation significantly increased Pmax as a function of NL in both species. Dark respiration (Rd) was linearly correlated with NL. CO2 elevation slightly but significantly increased Rd in Abutilon, while it had no significant effect on Rd in Ambrosia. The initial slope of a light-response curve was determined from quantum yield (ϕabs) multiplied by leaf absorptance and then calibrated against NL. Daily canopy photosynthesis, calculated by integration of leaf photosynthesis with the actual distribution of leaf area, leaf N, and PPFD within a canopy, showed fairly good agree...

Landscape and species-level distribution of morphological and life history traits in a temperate woodland flora

We developed a description of a central New Eng- land deciduous hardwood forest based on the distribution within the community of morphological and life history traits (N = 34) and environmental factors. Classification by TWINSPAN of 186 species based on morphological and life history traits identified six major functional groups of species largely corresponding to growth form. A data matrix of plots x traits was ordinated using PCA. Each of the resulting four PCA axes was associated with a major environmental gradi- ent: drainage, site exposure, disturbance due to past land use and degree of disturbance in the 1938 New England Hurricane (24.9, 19.8, 11.7 and 8.4% of the variation respectively). Two patterns suggested that a suite of potentially functional traits, rather than a few key characters (e.g. vital attributes), govern the distribution of species in this community: (1) each of the four axes was largely associated with a different group of traits and (2) each axis was associated with several traits that ap- peared to sort independently (i.e. not to co-occur within species). Evaluating one often-examined trait, there was no evidence that dispersal ability limited the colonization of species into secondary woodlands. We also found that landscape-scale abundance was associated with a small number of traits. Production of fleshy fruits and few diaspores per plant were positively associated with landscape-level abundance. Our results suggest that attempts to understand the overall struc- ture and function of this plant community based on a few key characters, such as dispersal ability, will meet with limited success. However, when focusing on one aspect of the com- munity, such as frequency across the landscape, relatively few characters may be important.

Light, leaf age, and leaf nitrogen concentration in a tropical vine

SummaryTropical vines in the Araceae family commonly exhibit alternating periods of upward and downward growth, decoupling the usual relationship between decreasing light environment with increasing age among the leaves on a shoot. In this study I examined patterns of light, leaf specific mass, and leaf nitrogen concentration in relation to leaf position, a measure of developmental age, in field collected shoots of Syngonium podophyllum. These data were analyzed to test the hypothesis that nitrogen allocation parallels within-shoot gradients of light availability, regardless of the relationship between light and leaf age. I found that leaf nitrogen concentration, on a mass basis, was weakly correlated with leaf level light environment. However, leaf specific mass, and consequently nitrogen per unit leaf area, were positively correlated with gradients of light within the shoot, and either increased or decreased with leaf age, providing support for the hypothesis that nitrogen allocation parallels gradients of light availability.

Size-dependent variation of gender in high density stands of the monoecious annual, Ambrosia artemisiifolia (Asteraceae).

SummaryWe examined the relationship between size variability and the distribution of functional gender in stands of the monoecious, wind-pollinated annual Ambrosia artemisiifolia. Populations of 60 individuals were grown in the greenhouse at a density of 372 m−2 and at two nutrient levels. Among the surviving plants, after self-thinning, variability in above-ground biomass and gender was higher in the high nutrient treatment. Among individuals there was a significant positive correlation between maleness and both height and biomass. Fecundity was also positively correlated with both measures of size. Based on the pattern of distribution of male and female flowers within the plant, it appears that the increase in maleness in larger plants is due to increased branching and axis elongation. These results demonstrate that competitive interactions, which lead to increased variability in biomass and fecundity, can also generate variability in gender within populations.

The forest-cerrado transition zone in southern Amazonia: Results of the 1985 Projeto Flora Amazônica expedition to Mato Grosso

An account of the history of collecting in the Mato Grosso is followed by a discussion of the current views on the nature of the vegetation along the transition from Amazonian forest to cerrado. The purpose of the 1985 Projeto Flora Amazônica expedition was to sample the vegetation in the transition region in northern Mato Grosso. In order to characterize the vegetation at the various localities, the identified collections were assigned to one of five broad distributional categories: Planaltine, Transitional, Amazonian, Wide-spread, or Other. In most cases, as expected, the collecting sites in the north and the west were characterized by Amazonian forest species while those in the south and east comprised mainly cerrado elements. Our collections show that the transition zone is a complex mosaic of Amazonian forest and cerrado formations.

Effects of CO2 elevation on canopy development in the stands of two co-occurring annuals

Elevated CO2 may increase dry mass production of canopies directly through increasing net assimilation rate of leaves and also indirectly through increasing leaf area index (LAI). We studied the effects of CO2 elevation on canopy productivity and development in monospecific and mixed (1:1) stands of two co-occurring C3 annual species, Abutilon theophrasti, and Ambrosia artemisiifolia. The stands were established in the glasshouse with two CO2 levels (360 and 700 μl/l) under natural light conditions. The planting density was 100 per m2 and LAI increased up to 2.6 in 53 days of growth. Root competition was excluded by growing each plant in an individual pot. However, interference was apparent in the amount of photons absorbed by the plants and in photon absorption per unit leaf area. Greater photon absorption by Abutilon in the mixed stand was due to different canopy structures: Abutilon distributed leaves in the upper layers in the canopy while Ambrosia distributed leaves more to the lower layers. CO2 elevation did not affect the relative performance and light interception of the two species in mixed stands. Total aboveground dry mass was significantly increased with CO2 elevation, while no significant effects on leaf area development were observed. CO2 elevation increased dry mass production by 30–50%, which was mediated by 35–38% increase in the net assimilation rate (NAR) and 37–60% increase in the nitrogen use efficiency (NUE, net assimilation rate per unit leaf nitrogen). Since there was a strong overall correlation between LAI and aboveground nitrogen and no significant difference was found in the regression of LAI against aboveground nitrogen between the two CO2 levels, we hypothesized that leaf area development was controlled by the amount of nitrogen taken up from the soil. This hypothesis suggests that the increased LAI with CO2 elevation observed by several authors might be due to increased uptake of nitrogen with increased root growth.