Demographic consequences of drought in the herbaceous perennial Cryptantha flava: effects of density,associations with shrubs,and plant size

The demographic consequences of a severe drought year were examined for two experimental plantings of the herbaceous desert perennial Cryptantha flava(Boraginaceae) in northeastern Utah, United States. A total of 6680 nutlets were planted individually or in clusters of four both under shrubs and in open microhabitats within two natural populations. Survival, growth, and flowering as a function of density and microhabitat were followed for 7 years, including 1 year when precipitation just before and during the growing season was 74.5% below normal. The design permitted assessment of how intraspecific density and shrub cover affect demographic response to drought. Mortality increased and flowering decreased dramatically during drought but neither varied with density or between shrub and open microhabitats. For plants growing under shrubs, survival (at Site 1) and growth (at Site 2) varied with shrub species. Average aboveground plant size also decreased during drought. Population size hierarchies were rearranged because larger plants lost leaf rosettes while many smaller plants grew. Density and microhabitat affected plant performance in non-drought years but more often at Site 1 than at Site 2. Individuals growing alone often were more likely to flower and/or produced more inflorescences when they did flower than did individuals growing with at least one other plant. However, for 2 years, survival rates at Site 1 were higher for plants growing in clumps than for single individuals. Shrubs also had mixed effects on plant performance. In some years, survival was higher under shrubs, but at Site 1 plants in the open often were more likely to flower and/or produced more inflorescences. Thus despite severe demographic consequences of drought, the study provided no evidence that intraspecific competition, interference by shrubs, or facilitation by shrubs increases under limited soil water.     

Non-destructive estimation of lateral root distribution in an aridland perennial

Estimation of root distributions in natural systems remains challenging due to the difficulties in excavation and easy breakage of fine roots. Identifying lateral fine root distribution is necessary to determine the potential exploitation of spatially and temporally variable nutrient supplies that characterize most arid ecosystems. We estimated this potential by taking field measurements of lateral root distribution of the small herbaceous perennial Cryptantha flava (A. Nels.) Payson using ¹⁵N-enriched nutrient solutions wicked into the soil at various distances from study plants. Leaves were subsequently harvested from these plants and analyzed for N isotopic ratios. C. flava plants were capable of N uptake at distances of greater than 1.0 m from the outer edge of their aboveground canopy. The considerable lateral root neighborhood area of C. flava increases the amount of spatially variable N that is exploitable in these low-N soils. The ability to acquire spatially variable N and rapidly translate N uptake into photosynthetic carbon gain are traits that aid C. flava in maintaining its position as a successful subordinate competitor in a community dominated by larger, woody perennials.     

Cleistogamy in the rare high Andean perennial herb Cryptantha capituliflora (Boraginaceae)

 Axial and apical flowers of Cryptantha capituliflora were analyzed with regard to morphology and pollen tube growth to assess the occurrence of cleistogamy. Although intermediate floral forms do occur, cleistogamous flowers were significantly smaller than chasmogamous flowers, had fewer anthers, and showed a distinctive stigmatic surface. Chasmogamous flowers can be cross-pollinated. Nevertheless, the growth of self-pollen tubes in few chasmogamous buds jointly with flower characters suggests that these flowers can probably produce fruits through autonomous selfing. The mean seed number per fruit did not differ between fruits from chasmogamous and cleistogamous flowers. Cleistogamous flowers were only observed in axial inflorescences, which are completely covered by the leaf. Other species of section Cryptantha also show the same trend, with cleistogamous flowers located in the lower half of the stems. This pattern is discussed in relation to dissimilarities in the outcrossing opportunities between flower types within the plant. Received May 22, 2002; accepted November 14, 2002 Published online: March 20, 2003     

Variation in carbon isotope discrimination in relation to plant performance in a natural population of <Emphasis Type="Italic">Cryptantha flava</Emphasis>

Few studies of phenotypic selection have focused on physiological traits, especially in natural populations. The adaptive significance of plant water-use efficiency, the ratio of photosynthesis to water loss through transpiration, has rarely been examined. In this study, carbon isotopic discrimination, Δ, an integrated measure of water-use efficiency, was repeatedly measured in juveniles and adults in a natural population of the herbaceous desert perennial Cryptantha flava over a 4-year period and examined for plasticity in Δ, consistency between years in values of Δ, and evidence for selection on Δ phenotypes. There was significant concordance in Δ values among the 4 years for adult plants and significant correlations in Δ values measured in different years for juveniles and adults combined. The wettest year of the study, 1998, proved an exception because Δ values that year were not correlated with Δ values in any other year of the study. Consistency in Δ measured on the same plants in different years could indicate genotypic variation and/or consistency in the water status of the microhabitats the plants occupied. Two forms of plasticity in Δ were also evident; mean seasonal values were correlated with precipitation the preceding autumn, and Δ values also declined with plant size, indicating increasing water-use efficiency. Phenotypic selection was evident because in the first year of the study juvenile plants that would survive until year five averaged lower Δ values than did those that failed to survive. During the driest year, 2000, Δ was significantly negatively correlated with adult plant size, measured as the number of leaf rosettes, but the negative relationship between Δ and the number of flowering stalks, a more direct measure of fitness, was not significant. These results suggest that the direction of phenotypic selection on Δ changes as plants grow.     

Two ATPases

In this article, I reflect on research on two ATPases. The first is F(1)F(0)-ATPase, also known as ATP synthase. It is the terminal enzyme in oxidative phosphorylation and famous as a nanomotor. Early work on mitochondrial enzyme involved purification in large amount, followed by deduction of subunit composition and stoichiometry and determination of molecular sizes of holoenzyme and individual subunits. Later work on Escherichia coli enzyme utilized mutagenesis and optical probes to reveal the molecular mechanism of ATP hydrolysis and detailed facets of catalysis. The second ATPase is P-glycoprotein, which confers multidrug resistance, notably to anticancer drugs, in mammalian cells. Purification of the protein in large quantity allowed detailed characterization of catalysis, formulation of an alternating sites mechanism, and recently, advances in structural characterization.     

Two neo-Darwinisms

There are two extant theories of evolution, each of which deserves the honourific "neo-Darwinism": Modern Synthesis Replicator theory and a theory I shall call Developmental Darwinism. The principal difference concerns the canonical unit of biological organization. Modern Synthesis replicator theory explains the process of evolution by appeal to the activities of genes or replicators. Developmental Darwinism explains the process of evolution by appeal to the capacities of organisms. In particular, it is the plasticity of organisms, manifested most distinctly during development, that causes adaptive evolution. Despite the fact that each, in its own way, traces its origin to the theory outlined by Darwin, they are radically different. The objectives of this essay are twofold: to underscore the differences between these theories, and to argue that Developmental Darwinism, though nascent, is a viable alternative to Modern Synthesis replicator theory.