The influence of density on frequency-dependent food selection: a comparison of four experiments with wild birds

We compare the results of four experiments, conducted at different times and with different protocols, that explored the relationship between frequency-dependent selection and prey density in wild birds feeding on artificial populations of coloured baits. One (experiment 4) used pastry baits that differed only in the presence or absence of a red stripe, and this experiment provided no evidence for any kind of selective behaviour. The other three experiments used green and brown baits, and they all provided evidence for a trend towards increasing anti-apostatic selection with high densities (>100 baits m^–2). However, one of these (experiment 3) provided no evidence for frequency-dependent selection at low densities (0.5–20 baits m^–2), while the other two experiments concurred in suggesting a trend towards increasing apostatic selection with low densities (down to 2 baits m^–2). Together, these experiments both support and qualify the published findings of experiment 1 that frequency- dependent selection by wild birds on bait populations is modified by density. Experiment 4 indicates that frequency-dependent selection may break down entirely if bait types are too similar, while experiment 3 indicates that some details of this trend with density will depend either on the protocol used or on exogenous changes in the birds’ feeding behaviour. Received: 1 September 1999 / Accepted: 22 March 2000     

Running with the Red Queen: the role of biotic conflicts in evolution

What are the causes of natural selection? Over 40 years ago, Van Valen proposed the Red Queen hypothesis, which emphasized the primacy of biotic conflict over abiotic forces in driving selection. Species must continually evolve to survive in the face of their evolving enemies, yet on average their fitness remains unchanged. We define three modes of Red Queen coevolution to unify both fluctuating and directional selection within the Red Queen framework. Empirical evidence from natural interspecific antagonisms provides support for each of these modes of coevolution and suggests that they often operate simultaneously. We argue that understanding the evolutionary forces associated with interspecific interactions requires incorporation of a community framework, in which new interactions occur frequently. During their early phases, these newly established interactions are likely to drive fast evolution of both parties. We further argue that a more complete synthesis of Red Queen forces requires incorporation of the evolutionary conflicts within species that arise from sexual reproduction. Reciprocally, taking the Red Queen''s perspective advances our understanding of the evolution of these intraspecific conflicts.     

Differentiating the effects of origin and frequency in reciprocal transplant experiments used to test negative frequency-dependent selection hypotheses

Reciprocal transplant experiments have been used to estimate the probability that negative frequency-dependent selection by natural enemies has occurred in host populations by determining whether pest populations are less adapted to “foreign” (rare) hosts, which originate from a population with which the pests have not coevolved. However, these experiments usually confound the effects of frequency and origin: the rare genotypes are also genotypes that did not originate at a site. When clonal organisms are used, and the clones occur in more than one population, it is possible to separate the effects of origin and frequency. Here I present the results of an experiment in which Arabis clones of known frequency were reciprocally transplanted among sites. Contrary to expectations, clones at their site of origin had less disease, less herbivory, and higher fitness than foreign clones. However, variation within and among sites in herbivory and infection was large, suggesting that the number of sites and clones needed to thoroughly test the hypothesis of negative frequency-dependent selection in this system is very large: thus, these results are suggestive but not conclusive. Received: 20 October 1997 / Accepted: 8 February 1998     

Effects of genetic architecture on the evolution of assortative mating under frequency-dependent disruptive selection

We consider a model of sympatric speciation due to frequency-dependent competition, in which it was previously assumed that the evolving traits have a very simple genetic architecture. In the present study, we numerically analyze the consequences of relaxing this assumption. First, previous models assumed that assortative mating evolves in infinitesimal steps. Here, we show that the range of parameters for which speciation is possible increases when mutational steps are large. Second, it was assumed that the trait under frequency-dependent selection is determined by a single locus with two alleles and additive effects. As a consequence, the resultant intermediate phenotype is always heterozygous and can never breed true. To relax this assumption, here we add a second locus influencing the trait. We find three new possible evolutionary outcomes: evolution of three reproductively isolated species, a monomorphic equilibrium with only the intermediate phenotype, and a randomly mating population with a steep unimodal distribution of phenotypes. Both extensions of the original model thus increase the likelihood of competitive speciation.     

Characterisation of Triticum vavilovii-derived stripe rust resistance using genetic,cytogenetic and molecular analyses and its marker-assisted selection

Stripe rust resistance was identified in Triticum vavilovii (T. vavilovii Aus22498)-derived Russian wheat aphid (RWA)-resistant germplasm. Inheritance studies indicated monogenic control of resistance. The resistance gene was tentatively designated as Yrvav and was located on chromosome 1B by monosomic analysis. A close association (1.5±0.9% recombination) of Yrvav with a T. vavilovii-derived gliadin allele (Gli-B1vav) placed it in chromosome arm 1BS. Yrvav was allelic with Yr10. Tests with Yr10 avirulent and virulent pathotypes showed that Yrvav and Yr10 possess identical pathogenic specificity. Yrvav and Yr10 showed close genetic associations with alternate alleles at the Xpsp3000 (microsatellite marker), Gli-B1 and Rg1 loci. Based on these observations Yrvav was named as Yr10vav. The close association between Xpsp3000 and Gli-B1 was also confirmed. The Yr10vav-linked Xpsp3000 allele (285 bp) was not present in 65 Australian cultivars, whereas seven Australian wheats lacking Yr10 carried the same Xpsp3000 allele (260 bp) as Yr10 carrying wheat cultivar Moro. Xpsp3000 and/or Gli-B1 could be used in marker-assisted selection for pyramiding Yr10vav or Yr10 with other stripe rust resistance genes. Yr10vav was inherited independently of the T. vavilovii-derived RWA resistance. Received: 5 December 2000 / Accepted: 3 April 2001     

Path coefficient analysis of the effects of stripe rust and cultivar mixtures on yield and yield components of winter wheat

Four club wheat cultivars and three two-component cultivar mixtures, planted at five frequencies, were grown in three environments in both the presence and absence of stripe rust. The effect of stripe rust on wheat yield was through the yield components, with weight of individual seed being the component most affected by rust. In some cases, yield component compensation was indicated by the presence of negative correlations among the yield components. Path analysis of the yield components revealed that components with the highest correlations to yield also had the largest direct effects on yield. Of the yield components, number of heads per unit area exerted the largest direct influence on yield. The direct effects of number of seeds per head and weight of individual seed were similar, although number of seeds per head was more important in the absence of rust than in its presence. The pure stands and mixtures differed considerably with respect to correlation coefficients, but were very similar for direct effects of yield components on yield. Most of these discrepancies were due to opposing indirect effects, which were not evident from correlation coefficients alone.Paper No. 10,788 of the Journal Series of the Oregon Agricultural Experiment Station     

Perturbation expansions of multilocus fixation probabilities for frequency-dependent selection with applications to the Hill-Robertson effect and to the joint evolution of helping and punishment

Natural populations are of finite size and organisms carry multilocus genotypes. There are, nevertheless, few results on multilocus models when both random genetic drift and natural selection affect the evolutionary dynamics. In this paper we describe a formalism to calculate systematic perturbation expansions of moments of allelic states around neutrality in populations of constant size. This allows us to evaluate multilocus fixation probabilities (long-term limits of the moments) under arbitrary strength of selection and gene action. We show that such fixation probabilities can be expressed in terms of selection coefficients weighted by mean first passages times of ancestral gene lineages within a single ancestor. These passage times extend the coalescence times that weight selection coefficients in one-locus perturbation formulas for fixation probabilities. We then apply these results to investigate the Hill-Robertson effect and the coevolution of helping and punishment. Finally, we discuss limitations and strengths of the perturbation approach. In particular, it provides accurate approximations for fixation probabilities for weak selection regimes only (Ns?1), but it provides generally good prediction for the direction of selection under frequency-dependent selection.     

The maintenance of sex: host–parasite coevolution with density‐dependent virulence

Why don’t asexual females replace sexual females in most natural populations of eukaryotes? One promising explanation is that parasites could counter the reproductive advantages of asexual reproduction by exerting frequency‐dependent selection against common clones (the Red Queen hypothesis). One apparent limitation of the Red Queen theory, however, is that parasites would seem to be required by theory to be highly virulent. In the present study, I present a population‐dynamic view of competition between sexual females and asexual females that interact with co‐evolving parasites. The results show that asexual populations have higher carrying capacities, and more unstable population dynamics, than sexual populations. The results also suggest that the spread of a clone into a sexual population could increase the effective parasite virulence as population density increases. This combination of parasite‐mediated frequency‐dependent selection, and density‐dependent virulence, could lead to the coexistence of sexual and asexual reproductive strategies and the long‐term persistence of sex.     

Parasitism and breeding system variation in North American populations of Daphnia pulex

The Red Queen hypothesis proposes that frequency-dependent selection by parasites may be responsible for the evolutionary maintenance of sexual reproduction. We sought to determine whether parasites could be responsible for variation in the occurrence of sexual reproduction in 21 populations of Daphnia pulex (Crustacea; Cladocera) that previous studies have shown to consist of either cyclical parthenogens, obligate parthenogens, or a mixture of both. We measured parasite prevalence over a four-week period (which essentially encompasses an entire season for the temporary snow-melt habitats we sampled) and regressed three different measures of sexuality against mean levels of parasite prevalence. Levels of parasitism were low and we found no relationship between levels of sexuality and mean parasite prevalence. Genetic variation with infection level was detected in 2 of the 21 populations, with several different clones showing signs of overparasitism or underparasitism. Overall, however, our results suggest that parasites are not a major source of selection in these populations and it thus seems unlikely they are responsible for maintaining breeding system variation across the study region.     

Frequency-dependent selection by pollinators: mechanisms and consequences with regard to behaviour of bumblebees Bombus terrestris (L.) (Hymenoptera: Apidae)

Bombus terrestris , a typical pollinating insect species, was offered artificial flowers of two different corolla colours with the same sucrose solution reward in an array. Common colours were significantly preferred, and the strength of the frequency-dependent response increased as a result of learning. There were also frequency-independent biases towards blue flowers, probably because blue flowers appeared more conspicuous to bumblebees than yellow flowers, and the degree of preference for blue was greater when flowers had low nectar rewards. Flower-to-flower movements by individual bumblebees between flowers were non-random, were biased to movements within the same flower colour, and were also dependent on morph frequency. The mechanisms governing flower selection in bumblebees are discussed. Pollinators foraging similarly in a natural situation would induce positive frequency-dependent selection, assortative mating, and directional selection on different corolla colour morphs of the plant population being visited, resulting in stabilizing selection for a single flower colour.     

Resistance of wild wheat to stripe rust: Predictive method by ecology and allozyme genotypes

From 114 accessions of wild emmer wheat from 11 sites in Israel, known for their allozymic variation (Nevo & al. 1982), individual genotypes were tested for resistance to one isolate of stripe rust both in the seedling stage in a growth chamber and in the adult plant stage in the field. The results indicate that resistance to stripe rust in seedlings and adults are significantly correlated (r_s = 0.40, p < 0.001). Genetic polymorphisms of resistance to stripe rust vary geographically and are predictable by climatic, as well as allozymic markers. Three variable combinations of rainfall, evaporation, and temperature explain significantly 0.40–0.53 of the spatial variance in disease resistance to stripe rust, suggesting the operation of natural selection. Several allozyme genotypes are significantly associated with disease resistance. We conclude that natural populations of wild emmer wheat in Israel contain large amounts of disease resistance genes. These populations could be effectively screened and then utilized by the phytopathologist for identifying resistant genotypes and producing new resistant cultivars.Patterns of Resistance of Wild Wheat to Pathogens in Israel II.     

Testing the pluralist approach to sex: the influence of environment on synergistic interactions between mutation load and parasitism in Daphnia magna

Both deleterious mutations and parasites have been acknowledged as potential selective forces responsible for the evolutionary maintenance of sexual reproduction. The pluralist approach to sex proposes that these two factors may have to interact synergistically in order to stabilize sex, and one of the simplest ways this could occur is if parasites are capable of causing synergistic epistasis between mutations in their hosts. However, the effects of both deleterious mutations and parasitism are known to be influenced by a range of environmental factors, so the nature of the interaction may depend upon the organisms' environment. Using chemically mutated Daphnia magna lines, we examined the effects of mutation and parasitism under a range of temperature and food regimes. We found that although parasites were capable of causing synergistic epistasis between mutations in their hosts, these effects were dependent upon an interaction between parasite genotype and temperature.     

A robust KASP marker for selection of four pairs of linked leaf rust and stripe rust resistance genes introgressed on chromosome arm 5DS from different wheat genomes

Molecular Biology Reports - Stripe rust and leaf rust are among the most devastating diseases of wheat, limiting its production globally. Wheat wild relatives harbour genetic diversity for new...     

Impact of negative frequency-dependent selection on mating pattern and genetic structure: a comparative analysis of the S-locus and nuclear SSR loci in Prunus lannesiana var. speciosa

Mating processes of local demes and spatial genetic structure of island populations at the self-incompatibility (S-) locus under negative frequency-dependent selection (NFDS) were evaluated in Prunus lannesiana var. speciosa in comparison with nuclear simple sequence repeat (SSR) loci that seemed to be evolutionarily neutral. Our observations of local mating patterns indicated that male-female pair fecundity was influenced by not only self-incompatibility, but also various factors, such as kinship, pollen production and flowering synchrony. In spite of the mating bias caused by these factors, the NFDS effect on changes in allele frequencies from potential mates to mating pollen was detected at the S-locus but not at the SSR loci, although the changes from adult to juvenile cohorts were not apparent at any loci. Genetic differentiation and isolation-by-distance over various spatial scales were smaller at the S-locus than at the SSR loci, as expected under the NFDS. Allele-sharing distributions among the populations also had a unimodal pattern at the S-locus, indicating the NFDS effect except for alleles unique to individual populations probably due to isolation among islands, although this pattern was not exhibited by the SSR loci. Our results suggest that the NFDS at the S-locus has an impact on both the mating patterns and the genetic structure in the P. lannesiana populations studied.     

The distribution and prevalence of helminths,coccidia and blood parasites in two competing species of gecko: implications for apparent competition

Across the Pacific the invading gecko species Hemidactylus frenatus has competitively displaced the resident gecko species Lepidodactylus lugubris in urban/surburban habitats. Do parasites enhance, inhibit, orhave no effect on this invasion? Parasites can confer an advantage to an invading species when the invader (1) introduces a new parasite to a resident species that has a greater detrimental effect on the resident than the invader, (2) is less susceptible to endemic parasites than the resident, and/or (3) increases the susceptibility of the resident to parasites. Conversely, parasites may protect a resident against invasion when endemic parasites have a greater impact on the invader than the resident. We screened more than one thousand H. frenatus and L. lugubris in areas of sympatry and allopatry from 28 islands and 5 sites on mainland Asia for a broad array of blood parasites, coccidia and helminths in order to evaluate the potential for parasites to affect their interaction. We found that 1) There were no parasites which appear to protect L. lugubris against invasion by H. frenatus. 2) H. frenatus does not introduce the same parasite to L. lugubris in every location where the two come in to contact, but probably has introduced different parasites in different locations. L. lugubris also seems to have introduced at least one parasite to H. frenatus. 3) The prevalence of parasite species shared by the two hosts is generally higher in H. frenatus; however, prevalence is determined by many factors and cannot be directly translated as susceptibility. We discuss the implications of this difference in prevalence for the Red Queen hypothesis. 4) The prevalence of the cestode Cylindrotaenia sp. is significantly higher in L. lugubris that are sympatric with H. frenatus than those which are allopatric.     

The joint effects of kin, multilevel selection and indirect genetic effects on response to genetic selection

Kin and levels-of-selection models are common approaches for modelling social evolution. Indirect genetic effect (IGE) models represent a different approach, specifying social effects on trait values rather than fitness. We investigate the joint effect of relatedness, multilevel selection and IGEs on response to selection. We present a measure for the degree of multilevel selection, which is the natural partner of relatedness in expressions for response. Response depends on both relatedness and the degree of multilevel selection, rather than only one or the other factor. Moreover, response is symmetric in relatedness and the degree of multilevel selection, indicating that both factors have exactly the same effect. Without IGEs, the key parameter is the product of relatedness and the degree of multilevel selection. With IGEs, however, multilevel selection without relatedness can explain evolution of social traits. Thus, next to relatedness and multilevel selection, IGEs are a key element in the genetical theory of social evolution.     

Stable genetic polymorphism in heterogeneous environments: balance between asymmetrical dispersal and selection in the acorn barnacle

Elucidating the processes responsible for maintaining polymorphism at ecologically relevant genes is intimately related to understanding the interplay between selection imposed by habitat heterogeneity and a species' capacity for dispersal in the face of environmental constraints. In this paper, we used a model-based approach to solve equilibria of balanced polymorphism, given values of fitness and larval dispersal among different habitats in the acorn barnacle Semibalanus balanoides from the Gulf of St Lawrence. Our results showed that allele frequencies observed at both MPI* and GPI* loci represented stable equilibria, given empirical estimates of fitness values, and that considerably more larvae dispersed from one region (north) to the other (south) than vice versa. Dispersal conditions were predicted to be similar for the maintenance of polymorphism at both loci. Moreover, the values of asymmetrical dispersal required by the model to reach stable equilibria were compatible with empirical estimates of larval dispersal and oceanic circulation documented in this system. Overall, this study illustrated the usefulness of a modified and computable version of Bulmer's model (1972) in order to test hypotheses of balanced polymorphism resulting from interactions between spatial selection and asymmetrical dispersal.     

In vitro selection of barley and wheat for resistance against Helminthosporium sativum

Summary Calli derived from immature embryos of barley and wheat genotypes were screened for their resistance to purified culture filtrate produced by the fungus Helminthosporium sativum P.K. and B. Two selection methods were used: a continuous method in which four cycles of selection were performed one after another on toxic medium and a discontinuous method in which a pause on non-toxic medium was given after the second or third cycle of selection. The latter was superior as it allowed the calli to regain their regeneration ability. About 3,000 calli from two barley genotypes and 2,000 from two wheat genotypes were used for selection. The selection with the pathotoxins resulted in 6% to 17% surviving calli. Toxin tolerant callus lines of barley were characterised by protein isozymes. Zymograms showed one more isozyme than with the unselected sensitive callus. Barley and wheat plants have been regenerated from callus lines surviving the toxin treatment and in vivo testing against pathogen revealed that the majority of these plants were less sensitive.     

Daisyworld inhabited with daisies incorporating a seed size/number trade-off: the mechanism of negative feedback on selection from a standpoint of the competition theory

We reexamined a Daisyworld model from the traditional view of competition theory. Unlike the original model, white and black daisies in our model incorporate a seeding/germination trade-off against bare ground area without assuming the local temperature reward. As a result, the planetary temperature is automatically regulated by two species if the following conditions are met: (i) the species react equally to an environmental condition, but one can alter the environmental condition in the opposite direction to the other. (ii) that one of the two cannot have both a higher maximal growth rate (mu(max)) and lower half-saturation constant (K) than those of the other. In other words, a pair of phenotypes incorporates a trade-off between quality and number of seeds. We found that the homeostatic regulation can also be reconciled with the adaptive evolution of optimal temperature. The results of simulation imply that biotic environmental feedback can also be maintained when the emergence of polymorphisms (black and white daisies) is closely linked to such a trade-off.