Frequency dependent natural selection during character displacement in sticklebacks

Abstract We know little about how natural selection on a species is altered when a closely related species consuming similar resources appears in its environment. In a pond experiment with threespine sticklebacks I tested the prediction that divergent natural selection between competitors is frequency-dependent, changing with the distribution of phe-notypes in the environment. Differential growth and survival of phenotypes in a target stickleback population were contrasted between two treatments. In one treatment an offshore zooplankton feeder (the limnetic stickleback species) was added to the same pond as the target. In the other treatment I added the benthic stickleback instead, a species adapted to feeding on invertebrates from sediments and inshore vegetation. The target population was ecologically and morphologically intermediate with phenotypic variance artificially inflated by hybridization. Growth rates of phenotypes within the target population differed between treatments as predicted by character displacement. The impact of adding a second species always fell most heavily on those phenotypes in the target population resembling the added species most closely. However, those individuals in the target population that most resembled the added species did not experience reduced survival. Instead, consistent survival differences between populations suggested the presence of an inshore –offshore gradient in mortality risk. These results provide further support for the hypothesis of character displacement in sympatric sticklebacks. They suggest that displacement along the resource gradient also led to divergence in vulnerability to agents of mortality, probably including predation.     

Divergent selection drives the adaptive radiation of crossbills

Abstract Knowledge of how phenotype influences fitness is necessary if we are to understand the basis of natural selection and how natural selection contributes to adaptive radiations. Here I quantify selection on a wild population of red crossbills ( Loxia curvirostra complex) in the South Hills, Idaho. Bill depth is the target of selection and selection on bill depth is stabilizing. I then show how fitness is related to both bill depth and performance. I use these and previously published relationships to estimate a fitness surface for five species of red crossbills that are part of an ongoing adaptive radiation in western North America. The fitness surface for crossbills has distinct peaks and valleys, with each crossbill species residing on or very near the summits. This work strongly supports a key tenet of the ecological theory of adaptive radiations; namely, divergent selection for utilizing alternative resources is the ultimate cause of adaptive radiations.     

Adaptive evolutionary conservation: towards a unified concept for defining conservation units

Recent years have seen a debate over various methods that could objectively prioritize conservation value below the species level. Most prominent among these has been the evolutionarily significant unit (ESU). We reviewed ESU concepts with the aim of proposing a more unified concept that would reconcile opposing views. Like species concepts, conflicting ESU concepts are all essentially aiming to define the same thing: segments of species whose divergence can be measured or evaluated by putting differential emphasis on the role of evolutionary forces at varied temporal scales. Thus, differences between ESU concepts lie more in the criteria used to define the ESUs themselves rather than in their fundamental essence. We provide a context-based framework for delineating ESUs which circumvents much of this situation. Rather than embroil in a befuddled debate over an optimal criterion, the key to a solution is accepting that differing criteria will work more dynamically than others and can be used alone or in combination depending on the situation. These assertions constitute the impetus behind adaptive evolutionary conservation.     

Interspecific Differences of Leaf Gas Exchange and Water Relations of Three Evergreen Mediterranean Shrub Species

Leaf gas exchange and plant water relations of three co-occurring evergreen Mediterranean shrubs species, Quercus ilex L. and Phillyrea latifolia L. (typical evergreen sclerophyllous shrubs) and Cistus incanus L. (a drought semi-deciduous shrub), were investigated in order to evaluate possible differences in their adaptive strategies, in particular with respect to drought stress. C. incanus showed the highest annual rate of net photosynthetic rate (P _N) and stomatal conductance (g _s) decreasing by 67 and 69 %, respectively, in summer. P. latifolia and Q. ilex showed lower annual maximum P _N and g _s, although P _N was less lowered in summer (40 and 37 %, respectively). P. latifolia reached the lowest midday leaf water potential (₁) during the drought period (–3.54±0.36 MPa), 11 % lower than in C. incanus and 19 % lower than in Q. ilex. Leaf relative water content (RWC) showed the same trend as ₁. C. incanus showed the lowest RWC values during the drought period (60 %) while they were never below 76 % in P. latifolia and Q. ilex; moreover C. incanus showed the lowest recovery of ₁ at sunset. Hence the studied species are well adapted to the prevailing environment in Mediterranean climate areas, but they show different adaptive strategies that may be useful for their co-occurrence in the same habitat. However, Q. ilex and P. latifolia by their water use strategy seem to be less sensitive to drought stress than C. incanus.     

Mathematical modeling of the onset of capillary formation initiating angiogenesis

It is well accepted that neo-vascular formation can be divided into three main stages (which may be overlapping): (1) changes within the existing vessel, (2) formation of a new channel, (3) maturation of the new vessel. In this paper we present a new approach to angiogenesis, based on the theory of reinforced random walks, coupled with a Michaelis-Menten type mechanism which views the endothelial cell receptors as the catalyst for transforming angiogenic factor into proteolytic enzyme in order to model the first stage. In this model, a single layer of endothelial cells is separated by a vascular wall from an extracellular tissue matrix. A coupled system of ordinary and partial differential equations is derived which, in the presence of an angiogenic agent, predicts the aggregation of the endothelial cells and the collapse of the vascular lamina, opening a passage into the extracellular matrix. We refer to this as the onset of vascular sprouting. Some biological evidence for the correctness of our model is indicated by the formation of teats in utero. Further evidence for the correctness of the model is given by its prediction that endothelial cells will line the nascent capillary at the onset of capillary angiogenesis. Received: 27 May 1999 / Revised version: 28 December 1999 / Published online: 16 February 2001     

Statistical methods for detecting molecular adaptation

The past few years have seen the development of powerful statistical methods for detecting adaptive molecular evolution. These methods compare synonymous and nonsynonymous substitution rates in protein-coding genes, and regard a nonsynonymous rate elevated above the synonymous rate as evidence for darwinian selection. Numerous cases of molecular adaptation are being identified in various systems from viruses to humans. Although previous analyses averaging rates over sites and time have little power, recent methods designed to detect positive selection at individual sites and lineages have been successful. Here, we summarize recent statistical methods for detecting molecular adaptation, and discuss their limitations and possible improvements.     

Some problems of crustacean taxonomy related to the phenomenon of Horohalinicum

If gene regulatory processes form the basis of phenotypic plasticity that evidences itself as discrete adaptive norms (sensu developmental conversion), it seems evident that the forms capable of such adaptations would possess a correspondingly richer genome than those lacking such adaptations. We examine evidence for the existence of genetic triads which are comprised of one form with wide niches due to phenotypic plasticity and two forms, each which is genetically fixed and exhibits a phenotype at one of the extremes of the plastic form. The invasion of organisms into new environments may occur by acquisition of discrete adaptive norms (DAN), which enable a transition between the old and new environments. The horohalinicum (=critical salinity) is defined as a salinity zone which always corresponds to 5–8 ppm and divides freshwater and marine faunas and floras, as well as many other physico-chemical characteristics. The current paper explores the existence of discrete adaptive norms and genetic triads with respect to salinity in some euryhaline crustaceans, particularly the arctic copepods, and discusses how such adaptations might inform us about species boundaries that affect the taxonomic status of these forms.     

Patterns of parapatric speciation

Abstract. Geographic variation may ultimately lead to the splitting of a subdivided population into reproductively isolated units in spite of migration. Here, we consider how the waiting time until the first split and its location depend on different evolutionary factors including mutation, migration, random genetic drift, genetic architecture, and the geometric structure of the habitat. We perform large-scale, individual-based simulations using a simple model of reproductive isolation based on a classical view that reproductive isolation evolves as a by-product of genetic divergence. We show that rapid parapatric speciation on the time scale of a few hundred to a few thousand generations is plausible even when neighboring subpopulations exchange several individuals each generation. Divergent selection for local adaptation is not required for rapid speciation. Our results substantiates the claims that species with smaller range sizes (which are characterized by smaller local densities and reduced dispersal ability) should have higher speciation rates. If mutation rate is small, local abundances are low, or substantial genetic changes are required for reproductive isolation, then central populations should be the place where most splits take place. With high mutation rates, high local densities, or with moderate genetic changes sufficient for reproductive isolation, speciation events are expected to involve mainly peripheral populations.     

Limits to adaptive plasticity: temperature and photoperiod influence shade-avoidance responses

In plants, the ratio of red to far-red wavelengths (R:FR) reliably indicates neighbor proximity and influences stem elongation. Enhanced elongation increases light interception and fitness under crowded conditions. However, many environmental factors vary simultaneously such that responses to R:FR may be affected by abiotic conditions or maternal environmental conditions. This study examines the effects of temperature, photoperiod, and maternal environment on stem-elongation responses to R:FR. Four populations of Abutilon theophrasti (two from disturbed, weedy areas and two from cornfields) were used in factorial common-garden experiments of temperature × R:FR × population and photoperiod × R:FR × population. Seedling growth of greenhouse- and field-derived seed was compared to evaluate maternal effects. Maternal environment did not alter seedling elongation. Higher temperatures resulted in both a twofold increase in average elongation and increased responsiveness to R:FR. Significant three-way interactions in both experiments demonstrate that population responses to R:FR differ depending on temperature and photoperiod conditions. These results indicate that elongation responses to R:FR are more variable than previously realized. The observed variability in elongation also suggests that the outcome of competitive interactions in the natural environment will depend on ambient temperature, photoperiod length, and population origin.