LOCAL POLYPLOID VARIATION IN THE NATIVE PRAIRIE GRASS ANDROPOGON GERARDII

The microscale distribution of polyploid variants of the dominant grass big bluestem (Andropogon gerardii) in virgin tallgrass prairie was mapped using flow cytometry. The correlation between DNA content and polyploidy allows the use of flow cytometry for nondestructive determination of polyploidy in intact plants. At Konza Prairie, local plots contained from 0 to 100% hexaploid cytotypes but most showed fine-scale mixing of the polyploid variants. The relationship of cytotype frequency to moisture availability or burning history was nonsignificant.     

SURVIVORSHIP AND FECUNDITY OF THE POLYCARPIC PERENNIAL MENTZELIA NUDA (LOASACEAE) IN NEBRASKA SANDHILLS PRAIRIE

Survivorship and fecundity of the polycarpic perennial Mentzelia nuda were investigated to help understand its population dynamics. Between 1978 and 1984, all 685 individuals found in a Nebraska sandhills prairie plot were marked and their fate followed. Three plants lived seven years, but the average lifespan was less than two years. Most plants spent their first year as nonflowering rosettes: only 9% flowered the first summer after germination. Overall, first-year survivorship ranged from 0.20 to 0.50 between 1978 and 1984. Fifty-two percent of rosettes survived to flower. Survivorship of flowering individuals was about the same as the survivorship of rosettes, although actual survivorships varied significantly between years. Both rainfall and age affected survivorship and fecundity.     

VARIATION IN CHROMATOGRAPHIC PATTERNS IN THE TRAGOPOGON DUBIUS-PRATENSIS-PORRIFOLIUS COMPLEX (COMPOSITAE)

Natural populations of the diploid species, Tragopogon dubius, T. pratensis and T. porrifolius, the F₁ hybrids of these species, and the two amphidiploids, T. mirus and T. miscellus, were obtained from the same localities used for previous morphological and cytological studies of the evolutionary relationships of this complex. Inter- and intra-populational comparisons were made utilizing 2-dimensional chromatograms of these taxa. Members of the complex shared a group of 18 flavonoid-type pattern components, no one of which was completely species specific. Therefore, species patterns were expressed for a population rather than an individual with pattern components being expressed as a frequency. Patterns for F₁ hybrids and amphidiploids, expressed in the same manner, were relatable to the parental species. Diploid species and F₁ hybrid patterns were the most variable in areas of greatest sympatry, indicating considerable genetic interchange between species. The distribution of specific components in certain populations was interpreted as chemical introgression. Evidence was presented for separate origins of T. mirus in two localities. The results confirm previous interpretations of the evolutionary relationships in this complex, and when compared to those obtained for Baptisia, they implicate the different breeding systems as important factors in establishing the kind of variation observed.     

MIGRATION-SELECTION BALANCE AND LOCAL ADAPTATION OF MITOCHONDRIAL HAPLOTYPES IN RUFOUS-COLLARED SPARROWS (ZONOTRICHIA CAPENSIS) ALONG AN ELEVATIONAL GRADIENT

Variable selection pressures across heterogeneous landscapes can lead to local adaptation of populations. The extent of local adaptation depends on the interplay between natural selection and gene flow, but the nature of this relationship is complex. Gene flow can constrain local adaptation by eroding differentiation driven by natural selection, or local adaptation can itself constrain gene flow through selection against maladapted immigrants. Here we test for evidence that natural selection constrains gene flow among populations of a widespread passerine bird ( Zonotrichia capensis ) that are distributed along an elevational gradient in the Peruvian Andes. Using multilocus sequences and microsatellites screened in 142 individuals collected along a series of replicate transects, we found that mitochondrial gene flow was significantly reduced along elevational transects relative to latitudinal control transects. Nuclear gene flow, however, was not similarly reduced. Clines in mitochondrial haplotype frequency were strongly associated with transitions in environmental variables along the elevational transects, but this association was not observed for the nuclear markers. These results suggest that natural selection constrains mitochondrial gene flow along elevational gradients and that the mitonuclear discrepancy may be due to local adaptation of mitochondrial haplotypes.     

THE VASCULAR SYSTEM OF THE INFLORESCENCE AXIS OF ANDROPOGON GERARDII (GRAMINEAE) AND ITS BEARING ON CONCEPTS OF MONOCOTYLEDON VASCULAR TISSUE

The vascular bundles in the inflorescence axis of Andropogon gerardii occur in inner and outer systems. The inner system is made up of large, early developing strands that, at earliest stages of development, are precocious (= the appendage they are to serve has not yet been initiated). The outer system consists of later developing smaller strands that are open ended in a proximal direction (= strands differentiate basipetally in the cortex below the appendage they serve). Bundles of both the inner and outer systems are not connected to other procambium early in their development but exist as isolated strands. The veins of the inner system of the inflorescence axis occur as sympodia. The presence of inner and outer systems in the vascular tissue is common to most monocotyledons. However, amongst monocotyledons, only certain grasses have been shown to have strands of the inner system that are isolated early in development. Many dicotyledons have large strands which are precocious and some have smaller, later developing strands which are open ended in a proximal direction, hence they occur as isolated strands. These smaller strands in dicotyledons occur between large strands. Certain dicotyledons have an inner and an outer system of veins. Of these, some have veins of the inner system that differ from the inner system bundles of monocotyledons in that they also form part of the outer system of veins, or develop at a different time. One other dicotyledon with an inner and outer system, Bougainvillea, differs from monocotyledons only in that the bundles of the outer system do not seem to be isolated early in their development and anastomoses are seen between the inner and outer systems. Thus, it appears that monocotyledons differ from dicotyledons only in the presence of independent inner and outer systems of vascular bundles in the former. Thus, the hypothesis of Zimmermann and Tomlinson that there are basic differences between monocotyledon and dicotyledon vascular systems is not substantiated. It is even suspected that monocotyledon and dicotyledon vascular systems will be demonstrated to be modifications of a basic plan consisting of large, acropetally differentiating and smaller, basipetally differentiating strands.     

A QUANTITATIVE STUDY OF VARIATION IN THE CHENOPODIUM ATROVIRENS-DESICCATUM-PRATERICOLA COMPLEX

Multivariate analyses were applied to 96 populations (OTUs) of plants, traditionally referred to Chenopodium atrovirens, C. desiccatum, and C. pratericola, in an attempt to evaluate the numerous and often contradictory taxonomic treatments of plants in this complex. The study consisted of two major parts. The first involved the use of cluster and principal components analyses using 14 morphological characters on the entire set of 96 OTUs to search for phenetically distinct groupings; these analyses were conducted without knowledge of traditional taxonomic designations of individual OTUs. Three reasonably well-defined groups emerged from these preliminary analyses. When traditional taxonomic designations were applied to member OTUs of each group, one group was composed primarily of C. atrovirens, another of C. pratericola and the third of C. desiccatum. The second part of the study utilized canonical analysis to: 1) confirm the integrity of the phenetic groups, 2) to classify OTUs difficult to identify to species using traditional methods, and 3) to provide an evaluation of characters important in the separation of these groups. This analysis confirmed the integrity of the groups and provided a classification to species of nearly all of the otherwise difficult OTUs. In addition, canonical analysis demonstrated that a combination of characters was important in the separation of the phenetic groups.     

A GRADIENT IN BENTHIC INTERTIDAL ALGAL ASSEMBLAGES ALONG THE SOUTHERN CALIFORNIA COAST1

Based on an analysis floristic data, a gradient appears to exist in the composition of intertidal algae along the 450 km of southern California coastline immediately south of Point Conception. Reciprocal averaging ordination of the algal flora at 51 sites in this area suggests that the gradient is not strictly latitudinal. Variation from a latitudinal gradient occurs ca. 60-80 km south of Point Conception. An ordination of frequently occurring species indicated that compositional changes are substantial when sites at the extreme north and south are compared, but that the changes are gradual and continuous. Possible explanations for the gradient are discussed, and these include: sea temperature, upwelling, sand movement, human disturbance (i.e., pollution, trampling), and wave action.     

POPULATION STRUCTURE ALONG A STEEP ENVIRONMENTAL GRADIENT: CONSEQUENCES OF FLOWERING TIME AND HABITAT VARIATION IN THE SNOW BUTTERCUP,RANUNCULUS ADONEUS

Few studies have determined how gene flow and selection interact to generate population genetic structure in heterogeneous environments. One way to identify the potential role played by natural selection is to compare patterns of spatial genetic structure between different life cycle stages and among microenvironments. We examined patterns of spatial structure in a population of the snow buttercup (Ranunculus adoneus), using both adult plants and newly emerged seedlings. The study population spans a steep environmental gradient caused by gradual melting of snow within a permanent snowbed. Early-melting sites are characterized by denser vegetation, more fertile soils, and a longer growing season than late-melting sites tens of meters away. The flowering time of R. adoneus is controlled entirely by time of snowmelt, so the contiguous population is phenologically substructured into a series of successively flowering cohorts, reducing the opportunity for direct pollen transfer between early- and late-melting sites. For four highly polymorphic enzyme loci in this tetraploid species, there was subtle, but statistically significant, genetic differentiation between early, middle, and late-melting cohorts; adults usually showed greater differentiation among snowmelt zones than did seedlings. At two loci in adults and one locus in seedlings, homozygotes were more common than predicted at Hardy-Weinberg equilibrium, even when assuming maximum levels of double reduction during meiosis. This pattern suggests the occurrence of self-fertilization and/or population substructure. To determine how spatial isolation and phenological separation each contribute to genetic substructure, we used bivariate regression models to predict the numbers of allele differences between randomly paired individuals as a function of meters separation in space and days separation in flowering time. For newly emerged seedlings, we found that spatial separation was positively associated with genetic difference, but that the additional contribution of phenological separation to genetic difference was not significant. This implies that seeds and/or pollen move effectively across the snowmelt gradient, despite differences in flowering time. As was true for seedlings, spatial separation between paired adults contributed to greater genetic difference, but for a given spatial separation, the genetic difference between adult plants was reduced by phenological separation. This result implies that postemergence selection is favoring at least some seeds that migrate across the snowmelt gradient. Directional gene flow across the snowmelt gradient probably results from a genetic source-sink interaction, that is, the colonization of ecologically marginal late-melting sites by high quality seeds produced by the larger subpopulation in early-melting sites. Effective gene flow from high to low quality microenvironments is likely to impede adaptation to late-melting locations.     

THE LEVEL OF AGAMOSPERMY IN A NEBRASKA POPULATION OF SPIRANTHES CERNUA (ORCHIDACEAE)

This study investigated reproduction in a prairie population of Spiranthes cernua in eastern Nebraska. Spiranthes cernua reproduces both sexually and asexually, through adventitious embryony. Agamospermous seeds are monoembryonic, polyembryonic, have extruded or free embryos, or lack embryos. These kinds of seeds are extremely rare in Spiranthes vernalis, a close relative that relies exclusively on sexual reproduction. Sexually derived seeds are monoembryonic or lack embryos. One objective was to quantify the amount of sexual reproduction in a natural population. In 1985 and 1986, maximum estimates of sexual seed production per capsule were 19.2% and 33.6%, respectively. Thus, every capsule examined contained a high proportion of agamospermous seeds. In addition, attempts were made to manipulate the reproductive system with experimental crosses performed in the field. The treatments examined agamospermy, autogamy, and outcrossing. In all treatments the majority of seeds in a capsule were agamospermous.     

RESPONSE OF ANDROPOGON GERARDII (POACEAE) TO FIRE-INDUCED HIGH VS. LOW IRRADIANCE ENVIRONMENTS IN TALLGRASS PRAIRIE: LEAF STRUCTURE AND PHOTOSYNTHETIC PIGMENTS

Photosynthetic pigments and several structural characteristics were measured in leaves of Andropogon gerardii from tallgrass prairie populations in an unburned, low-irradiance site and a burned, high-irradiance site to determine if these species displayed sun/shade differences similar to those documented in forest species. Early in the growing season, leaves of A. gerardii in the low-irradiance, unburned site had significantly lower stomatal density, pore length, and conductance, as well as specific leaf mass and thickness than leaves from the high-irradiance, burned site. Moreover, the chlorophyll a:b ratio, carotenoid content, and bundle sheath-vascular complex area were significantly lower, but chlorophyll content (mass/mass) was greater in leaves in unburned vs. burned sites. These differences are consistent with sun/shade adaptations reported for forest understory plants and may contribute to the low productivity of A. gerardii in unburned tallgrass prairie.     

ALLOZYMIC VARIATION IN LOCAL APOMICTIC POPULATIONS OF LEMNA MINOR (LEMNACEAE)

Flowering occurrence and allozymic variation were studied in eight local populations of Lemna minor in eastern Ontario, Canada. After 2 years of survey, not a single flower was observed. This absence of flowering suggests the possibility of loss of sexual reproduction. This may have had no net adverse effect on fitness given the simple life history and prolific vegetative propagation in L. minor. However, the absence of sexual reproduction may limit genotypic diversity. The allozymic analysis detected 18 loci from 13 enzyme systems. Large deviations from Hardy-Weinberg equilibrium were common because of an excess of homozygotes for several enzyme systems. The genotypic diversity within these eight populations had a mean D value of 0.973 with an average number of genotypes per population of 19.6. These results suggest that genotypic diversity within these populations is not severely limited by the rarity of sexual reproduction. The mean genotypic distance index (D₁₄ = 0.801) suggests a high degree of differentiation between populations. The mean number of populations per genotype was 1.78. Using a Mantel test, the genotypic distance matrix was not significantly related to the population-to-population distance matrix (t = -0.161, P = 0.413). Although rare events of sexual reproduction may help maintain genetic variation, somatic mutations and multiple origins of clones may be important factors maintaining genetic diversity both within and between populations of L. minor.     

VARIATION AND EVOLUTION OF BREEDING SYSTEMS IN THE TURNERA ULMIFOLIA L. COMPLEX (TURNERACEAE)

The evolutionary and functional relationships among breeding systems and floral morphology were investigated in the Turnera ulmifolia complex. Predictions of a model of breeding system evolution among distylous and homostylous varieties were tested. Chromosome counts of 73 accessions revealed an association between breeding system and chromosome number. Diploid and tetraploid populations of five taxonomic varieties are distylous and self-incompatible, whereas hexaploid populations of three varieties are homostylous and self-compatible. The latter occur at different margins of the geographical range of the complex. Crossing studies and analyses of pollen and ovule fertility in F₁'s revealed that the three homostylous varieties are intersterile. To test the prediction that, homostylous varieties are long homostyles that have originated by crossing over within the distyly supergene, a crossing program was undertaken among distylous and homostylous plants. Residual incompatibility was observed in styles and pollen of each homostylous variety with patterns consistent with predictions of the cross-over model. The intersterility of hexaploid varieties suggests that long homostyly has arisen on at least three occasions in the complex by recombination within the supergene controlling distyly. Deviation from expected compatibility behavior occurs in populations of var. angustifolia that have the longest styles. These phenotypes displayed the greatest separation between anthers and stigmas (herkogamy) and set little seed in crosses with long- or short-styled plants. This suggests that they are derived from long homostyles with shorter length styles. It is proposed that selection for increased outcrossing has favored the evolution of herkogamy in long homostyles. Estimates of outcrossing rate in a distylous population using allozyme markers confirmed that dimorphic incompatibility enforces complete outcrossing. Significant genetic variation for floral traits likely to influence the mating system, such as stigma-anther separation, occurs within and among homostylous populations of var. angustifolia on Jamaica. Estimates of the mating system of families from a population with varying degrees of stigma-anther separation, using five isozyme loci, were heterogeneous and ranged from t = 0.04–0.79. Families exhibiting the largest mean stigma-anther separation have higher outcrossing rates than those with little separation.     

MICROGEOGRAPHIC VARIATION IN MITOCHONDRIAL DNA OF MEADOW VOLES (MICROTUS PENNSYLVANICUS) IN RELATION TO POPULATION DENSITY

We examined mitochondrial-DNA (mtDNA) sequence heterogeneity on four adjacent trapping grids in an island population of meadow voles (Microtus pennsylvanicus) at two different population densities. Four restriction endonucleases revealed 20 different mtDNA composite phenotypes in samples totaling 198 meadow voles. There were significant heterogeneities in the distribution of four common mtDNA composite phenotypes among the four trapping grids, suggesting that there is population subdivision on a fine scale. Genetic distances between grids, mtDNA diversity within grids, and G_ST also varied during the study period. We found a decrease in genetic distance and an increase in diversity when the population density was high and vice versa when the population density was low. When population density was high, the coefficient of gene differentiation was smaller than the same coefficient observed when the population density was low. These changes in population subdivision and diversity are consistent with theoretical expectations of population structure in which effective female population size and dispersal are the critical variables. The data also support the hypothesis of maintenance of mtDNA diversity by population subdivision, rapid population growth rate, and dispersal.     

VARIATION IN ENVIRONMENTAL AND GENOTYPIC SEX-DETERMINING MECHANISMS ACROSS A LATITUDINAL GRADIENT IN THE FISH,MENIDIA MENIDIA

Models of environmental sex determination (ESD) usually assume that genetic influences on sex are polygenic, but the validity of this (or any other) form of genotype-environment interaction is virtually unknown. In the Atlantic silverside, Menidia menidia, sex is determined by an interaction between temperature and genotype and the response of sex ratio to temperature differs among populations from different latitudes. We examined the genetic basis of this pattern by measuring among family variation in the proportion of females, F/(F + M), within and among high (21°C) and low (15°C) temperatures for two populations: one from Nova Scotia (NS) where the level of ESD is low, and another from South Carolina (SC) where the level of ESD is high. In NS fish, temperature had a significant influence on sex ratio in only 1 of 23 families. The distribution of the fraction of females within temperatures for families from NS was highly heterogeneous and tended to fall into distinct classes (0.0, 0.25, 0.5, 1.0) like that expected from Mendelian segregation of a major sex factor(s). In contrast, temperature had a highly significant influence on sex ratio in all SC families examined (N = 24). Family sex ratios within temperatures were highly heterogeneous and, at least at 15°C, did not conform to simple Mendelian ratios. At 21°C, the proportion of females in most SC families was near zero and so the underlying sex tendencies of different families could not be discerned. Based on a previous study, mid-latitude fish appear to have an intermediate form of sex determination: simple Mendelian sex-ratio patterns exist and there is a moderate thermal influence on sex ratio in most but not all families. We suggest that sex determination in M. menidia is controlled by an interaction between major genetic factors, polygenic factors, and temperature and that the relative importance of each component differs with latitude. High latitude populations appear to have evolved a major sex-determining factor(s) that overrides the effect of temperature, and this factor(s) is lacking in low latitude populations.     

GENETIC ADAPTATION AND ACCLIMATION OF PHYTOPLANKTON ALONG A STRESS GRADIENT IN THE EXTREME WATERS OF THE AGRIO RIVER–CAVIAHUE LAKE (ARGENTINA)1

We tested if different adaptation strategies were linked to a stress gradient in phytoplankton cells. For this purpose, we studied the adaptation and acclimation of Dictyosphaerium chlorelloides (Naumann) Komárek et Perman (Chlorophyta) and Microcystis aeruginosa (Kütz.) Kütz. (Cyanobacteria) to different water samples (from extremely acid, metal‐rich water to moderate stressful conditions) of the Agrio River–Caviahue Lake system (Neuquén, Argentina). Both experimental strains were isolated from pristine, slightly alkaline waters. To distinguish between physiological acclimation and genetic adaptation (an adaptive evolution event), a modified Luria‐Delbrück fluctuation analysis was carried out with both species by using as selective agent sample waters from different points along the stress gradient. M. aeruginosa did not acclimate to any of the waters tested from different points along the stress gradient nor did D. chlorelloides to the two most acidic and metal‐rich waters. However, D. chlorelloides proliferated by rapid genetic adaptation, as the consequence of a single mutation (5.4 × 10^?7 resistant mutants per cell per division) at one locus, in less extreme water and also by acclimation in the least extreme water. It is hypothesized that the stress gradient resulted in different strategies of adaptation in phytoplankton cells from nonextreme waters. Thus, very extreme conditions were lethal for both organisms, but as stressful conditions decreased, adaptation of D. chlorelloides cells was possible by the selection of resistant mutants, and in less extreme conditions, by acclimation.