John M. Emlen

Antisymmetry, directional asymmetry, and dynamic morphogenesis

Fluctuating asymmetry is the most commonly used measure of developmental instability. Some authors have claimed that antisymmetry and directional asymmetry may have a significant genetic basis, thereby rendering these forms of asymmetry useless for studies of developmental instability. Using a modified Rashevsky-Turing reaction-diffusion model of morphogenesis, we show that both antisymmetry and directional asymmetry can arise from symmetry-breaking phase transitions. Concentrations of morphogen on right and left sides can be induced to undergo transitions from phase-locked periodicity, to phase-lagged periodicity, to chaos, by simply changing the levels of feedback and inhibition in the model. The chaotic attractor has two basins of attraction-right sidedominance and left side dominance. With minor disturbance, a developmental trajectory settles into one basin or the other. With increasing disturbance, the trajectory can jump from basin to basin. The changes that lead to phase transitions and chaos are those expected to occur with either genetic change or stress. If we assume that the morphogen influences the behavior of cell populations, then a transition from phase-locked periodicity to chaos in the morphogen produces a corresponding transition from fluctuating asymmetry to antisymmetry in both morphogen concentrations and cell populations. Directional asymmetry is easily modeled by introducing a bias in the conditions of the simulation. We discuss the implications of this model for researchers using fluctuating asymmetry as an indicator of stress.

Developmental stability in plants: symmetries, stress and epigenesis

Plant developmental stability has received little attention in the past three or four decades. Here we review differences in plant and animal development, and discuss the advantages of using plants as experimental subjects in exploring developmental stability. We argue that any type of developmental invariant may be used to assess developmental stability and review the use of fluctuating asymmetry in studies of plant developmental stability. We also examine the use of deviations from translatory, radial, and self-symmetry as measures of developmental instability. The role of nonlinear dynamics and epigenesis in the production of the phenotype is also discussed.

Differences in native soil ecology associated with invasion of the exotic annual chenopod, Halogeton glomeratus

Various biotic and abiotic components of soil ecology differed significantly across an area where Halogeton glomeratus is invading a native winterfat, [ Krascheninnikovia (= Ceratoides) lanata] community. Nutrient levels were significantly different among the native, ecotone, and exotic-derived soils. NO3, P, K, and Na all increased as the cover of halogeton increased. Only Ca was highest in the winterfat area. A principal components analysis, conducted separately for water-soluble and exchangeable cations, revealed clear separation between halogeton- and winterfat-derived soils. The diversity of soil bacteria was highest in the exotic, intermediate in the ecotone, and lowest in the native community. Although further studies are necessary, our results offer evidence that invasion by halogeton alters soil chemistry and soil ecology, possibly creating conditions that favor halogeton over native plants.

Nonlinear growth dynamics and the origin of fluctuating asymmetry

The nonlinear, complex nature of biosynthesis magnifies the impacts of small, random perturbations on organism growth, leading to distortions in adaptive allometries and, in particular, to fluctuating asymmetry. These distortions can be partly checked by cell-cell and inter-body part feedback during growth and development, though the latter mechanism also may lead to complex patterns in right-left asymmetry. Stress can be expected to increase the degree to which random growth perturbations are magnified and may also result in disruption of the check mechanisms, thus exaggerating fluctuating asymmetry.The processes described not only provide one explanation for the existence of fluctuating asymmetry and its augmentation under stress, but suggest additional effects of stress as well. Specifically, stress is predicted to lead to decreased fractal dimension of bone sutures and branching structures in animals, and in increased dimension of growth trace patterns such as those found in mollusc shells and fish otoliths and scales.A basic yet broad primer on fractals and chaos is provided as background for the theoretical development in this manuscript.

Heterosis and outbreeding depression: A multi-locus model and an application to salmon production

Abstract Both artificial propagation and efforts to preserve or augment natural populations sometimes involve, wittingly or unwittingly, the mixing of different gene pools. The advantages of such mixing vis-a-vis the alleviation of inbreeding depression are well known. Acknowledged, but less well understood, are the complications posed by outbreeding depression. This paper derives a simple model of outbreeding depression and demonstrates that it is reasonably possible to predict the generation-to-generation fitness course of hybrids derived from parents from different origins. Genetic difference, or distance between parental types, is defined by the drop in fitness experienced by one type reared at the site to which the other is locally adapted. For situations where decisions involving stock mixing must be made in the absence of complete information, a sensitivity analysis-based conflict resolution method (the Good-Bad-Ugly model) is described.

Ecological Effects of Ranching: A Six-Point Critique

Abstract Ranching is the dominant land use in much of the American West. Although a copious literature has examined the effects of various grazing practices on native ecosystems, we present here the idea that ranching has important impacts on the land independent of those caused by grazing itself. If biological conservation is to be successful on the western grasslands and shrublands, ranchers must be central to any plan. Focusing on the Great Plains of the United States, and on Wyoming in particular, we raise six points of concern that must be addressed before we can hope to restore or maintain native ecosystems on the range.

Biased Sex Ratios in Plants: Theory and Trends

This paper examines the history of sex ratio theory and the effects of multiple variables on individual and population sex ratios. It also provides examples where plants have been used to test major predictions of sex ratio theory. Then, using over 200 studies from the literature, dioecious plant species are categorized based on their life form, pollination agent, fruit dispersal agent, and sex ratio. A loglinear analysis is used to look at possible correlations between the sex ratio of a population and other life history characteristics. These data are used to examine the predictions made by De Jong et al. (Journal of Evolutionary Biology 15:7, 2002), that relative pollen and seed dispersal distances can be used to predict sex ratio bias. Despite the limited sample size, strong relationships are still observed. 93% of insect pollinated dioecious vines that have biotically dispersed fruit have male-biased sex ratios. Conversely, 61% of shrubs that are wind pollinated and have abiotic fruit dispersal have female-biased sex ratios.

Developmental stability and its applications in ecotoxicology

Developmental stability refers to the ability of a developing organism to produce a consistent phenotype in a given environment. It provides a simple, reliable method of detecting stressed populations and monitoring their recovery. The most common measure of developmental instability, fluctuating asymmetry, assesses minor deviations from perfect bilateral symmetry in traits that are normally symmetrical. Measures of developmental instability are based upon the concept of developmental invariance. The biotest approach consists of the simultaneous analysis of developmental instability (and related physiological instability) in a variety of species.

Linking habitat quality with trophic performance of steelhead along forest gradients in the South Fork Trinity River Watershed, California.

Abstract We examined invertebrate prey, fish diet, and energy assimilation in relation to habitat variation for steelhead Oncorhynchus mykiss (anadromous rainbow trout) and rainbow trout in nine low-order tributaries of the South Fork Trinity River, northern California. These streams spanned a range of environmental conditions, which allowed us to use bioenergetics modeling to determine the relative effects of forest cover, stream temperature, season, and fish age on food consumption and growth efficiency. Evidence of seasonal shifts in reliance on aquatic versus terrestrial food sources was detected among forest cover categories and fish ages, although these categories were not robust indicators of O. mykiss condition and growth efficiency. Consumption estimates were generally less than 20% of maximum consumption, and fish lost weight in some streams during summer low-flow conditions when stream temperatures exceeded 15°C. Current 100-year climate change projections for California threaten to exacerbate ...

Animal population dynamics: Identification of critical components

Abstract There is a growing interest in the use of population dynamics models in environmental risk assessment and the promulgation of environmental regulatory policies. Unfortunately, because of species and areal differences in the physical and biotic influences on population dynamics, such models must almost inevitably be both complex and species- or site-specific. Given the emormous variety of species and sites of potential concern, this fact presents a problem; it simply is not possible to construct models for all species and circumstances. Therefore, it is useful, before building predictive population models, to discover what input parameters are of critical importance to the desired output. This information should enable the construction of simpler and more generalizable models. As a first step, it is useful to consider population models as composed to two, partly separable classes, one comprising the purely mechanical descriptors of dynamics from given demographic parameter values, and the other describing the modulation of the demographic parameters by environmental factors (changes in physical environment, species interactions, pathogens, xenobiotic chemicals). This division permits sensitivity analyses to be run on the first of these classes, providing guidance for subsequent model simplification. We here apply such a sensitivity analysis to network models of mammalian and avian population dynamics.