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Epidemiology and genetics in the coevolution of parasites and hosts.

Recent studies suggest that parasites (interpreted broadly to include viruses, bacteria, protozoans and helminths) may influence the numerical magnitude or geographical distribution of their host populations; most of such studies focus on the population biology and epidemiology of the host-parasite association, taking no explicit account of the genetics. Other researchers have explored the possibility that the coevolution of hosts and parasites may be responsible for much of the genetic diversity found in natural populations, and may even be the main reason for sexual reproduction; such genetic studies rarely take accurate account of the density- and frequency-dependent effects associated with the transmission and maintenance of parasitic infections. This paper aims to combine epidemiology and genetics, reviewing the way in which earlier studies fit into a wider scheme and offering some new ideas about host-parasite coevolution. One central conclusion is that 'successful' parasites need not necessarily evolve to be harmless: both theory and some empirical evidence (particularly from the myxoma-rabbit system) indicate that many coevolutionary paths are possible, depending on the relation between virulence and transmissibility of the parasite or pathogen.

Animals↗

Conservation and divergence in multigene families: alternatives to selection and drift.

It is generally assumed that conservation and divergence of DNA signify function (selection) and no function (drift), respectively. This assumption is based on the view that a mutation is a unique event on a single chromosome, the fate of which depends on selection or drift. Knowledge of the rates, units and biases of widespread mechanisms of non-reciprocal DNA exchange, in particular within multigene families, provides alternative explanations for conservation and divergence, notwithstanding biological function. Such mechanisms of DNA turnover cause continual fluctuations in the copy-number of variant genes in an individual and, hence, promote the gradual and cohesive spread of a variant gene throughout a family (homogenization) and throughout a population (fixation). The dual processes (molecular drive) of homogenization and fixation are inextricably linked. Data are presented of the expected stages of transition in the spread of variant repeats by molecular drive in some non-genic families of DNA, seemingly not under the influence of selection. When a molecularly driven change in a given gene family is accompanied by the coevolution (mediated by selection) of other DNA, RNA or protein molecules that interact with the gene family then biological function is observed to be maintained despite sequence divergence. Conversely, the mechanics of DNA turnover and a turnover bias in favour of ancestral sequences can dramatically retard the rate of sequence change, in the absence of function. Examples of the maintenance of function by molecular coevolution and conservation of sequences in the absence of function, are drawn mainly from the rDNA multigene family.

Animals↗

The evolution of culture: from primate social learning to human culture.

Cultural transmission in our species works most of the time as a cumulative inheritance system allowing members of a group to incorporate behavioral features not only with a positive biological value but sometimes also with a neutral, or even negative, biological value. Most of models of dual inheritance theory and gene-culture coevolution suggest that an increase, either qualitative or quantitative, in the efficiency of imitation is the key factor to explain the transformation of primate social learning in a cumulative cultural system of inheritance as it happens during hominization. We contend that more efficient imitation is necessary but not enough for this transformation to occur and that the key factor enabling such a transformation is that some hominids developed the capacity to approve or disapprove their offspring's learned behavior. This capacity to approve or disapprove offspring's behavior makes learning both less costly and more accurate, and it transformed the hominid culture into a system of cumulative cultural inheritance similar to that of humans, although the system was still prelinguistic in nature.

Animals↗

How to be a fig wasp.

In the two decades since Janzen described how to be a fig, more than 200 papers have appeared on fig wasps (Agaonidae) and their host plants (Ficus spp., Moraceae). Fig pollination is now widely regarded as a model system for the study of coevolved mutualism, and earlier reviews have focused on the evolution of resource conflicts between pollinating fig wasps, their hosts, and their parasites. Fig wasps have also been a focus of research on sex ratio evolution, the evolution of virulence, coevolution, population genetics, host-parasitoid interactions, community ecology, historical biogeography, and conservation biology. This new synthesis of fig wasp research attempts to integrate recent contributions with the older literature and to promote research on diverse topics ranging from behavioral ecology to molecular evolution.

Animals↗

The evolution of pathogenic trypanosomes.

In the absence of a fossil record, the evolution of protozoa has until recently largely remained a matter for speculation. However, advances in molecular methods and phylogenetic analysis are now allowing interpretation of the "history written in the genes". This review focuses on recent progress in reconstruction of trypanosome phylogeny based on molecular data from ribosomal RNA, the miniexon and protein-coding genes. Sufficient data have now been gathered to demonstrate unequivocally that trypanosomes are monophyletic; the phylogenetic trees derived can serve as a framework to reinterpret the biology, taxonomy and present day distribution of trypanosome species, providing insights into the coevolution of trypanosomes with their vertebrate hosts and vectors. Different methods of dating the divergence of trypanosome lineages give rise to radically different evolutionary scenarios and these are reviewed. In particular, the use of one such biogeographically based approach provides new insights into the coevolution of the pathogens, Trypanosoma brucei and Trypanosoma cruzi, with their human hosts and the history of the diseases with which they are associated.

Africa↗

Impact of parasite sympatry on the geographic mosaic of coevolution.

Slave-making ants are specialized social parasites that steal the young from colonies of their host species to augment their slave supply. The degree of parasite-host specialization has been shown to shape the trajectory along which parasites and hosts coevolve and is a prime contributor to the geographic mosaic of coevolution. However, virtually nothing is known about extrinsic influences on parasite-host dynamics, although the simple addition of a competing slave-maker may significantly alter selection pressures. Here we report the effect of two sympatric slave-makers on a single host. We measured temporal and spatial changes in colonies of the primary host Temnothorax curvispinosus that had been placed in field enclosures along with a single colony of either one or both species of the North American slave-making ants Protomognathus americanus and Temnothorax duloticus. Each slave-maker species alone had a negative impact on its hosts, although one slave-maker species more frequently decimated its host assemblage and then went extinct. Nevertheless, the combined effect in mixed-parasite enclosures was, surprisingly, greatly attenuated. Virulent slave-maker growth and prudent slave-maker decay in these shared enclosures, together with field data showing an inverse proportional relationship between the two slave-makers in natural populations, suggest that their checkered distribution is a consequence of direct asymmetrical antagonism between parasites. Thus, our results imply a tripartite coevolutionary arms race, whereby intraguild interactions among social parasites strongly affect the realized selection pressures on hosts and contribute to the geographic mosaic of coevolution.

Animals↗

[Are symbolic behaviour and neuroplasticity an example of gene-culture coevolution?].

INTRODUCTION AND DEVELOPMENT: The brain size in the Homo genus not only has not increased during the last 150,000 years but has also experienced a slight reduction in the last 35,000 years. This reduction coincides with the generalization of the symbolic culture that was most likely established during the Upper Palaeolithic. Therefore, the cognitive capacities characteristic in the Homo sapiens could be due to structural and functional changes during the brain evolution, rather than an increase of the brain size. Dependence of symbolic culture probably required an increase of the learning and memory skills, thus demanding, at the same time, an improvement of neuroplasticity. CONCLUSIONS: The epsilon3 and epsilon2 alleles of the apolipoprotein E seem to contribute to a better synaptic repairing, in relation to the ancestral epsilon4 allele. Mutation leading to the epsilon3 allele occurred between 220,000 and 150,000 years ago. Its selection and expansion may have continued until a relatively recent period that coincides with the emergence and expansion of the complex symbolic culture. Other factors favouring neuroplasticity, such as certain polymorphisms and the expression increase of certain proteins as reelin, could also have been selected. Emergence of the symbolic behaviour and increase of its deriving technical and social complexity could have made an intense selective pressure leading to a selection of genes that induced an improvement in neuroplasticity. This would constitute an example of gene-culture coevolution.

Alleles↗

Identification of selective sources: partitioning selection based on interactions.

Interspecific interactions are an inescapable reality in nature. The evolution of a species is largely determined by the environment, abiotic or biotic, in which selection occurs. Quantifying the magnitude of selection is crucial to understanding which aspects of the environment are important to the evolution of a species. Such knowledge is particularly important to fields such as conservation biology, which attempts to maintain a suitable environment for the prosperity of a species, or coevolution, where dynamics are determined by the strength of reciprocal selection between species. I present a general method by which selection due to interspecific interactions may be quantified. This technique is based on past quantitative genetic models of selection and can be used with other methodologies that build on these standard models. The approach may be expanded to account for n-species interactions (e.g., a plant with two pollinators). Simulation studies conducted using this method indicate that the magnitude of selection between two species is strongly correlated with the presence of nonrandom interactions.

Biological Evolution↗

Adventures with poxviruses of vertebrates.

Because they were the largest of all viruses and could be visualised with a light microscope, the poxviruses were the first viruses to be intensively studied in the laboratory. It was clear from an early date that they caused important diseases of humans and their domestic animals, such as smallpox, cowpox, camelpox, sheeppox, fowlpox and goatpox. This essay recounts some of the early history of their recognition and classification and then expands on aspects of research on poxviruses in which the author has been involved. Studies on the best-known genus, Orthopoxvirus, relate to the use of infectious ectromelia of mice as a model for smallpox, embracing both experimental epidemiology and pathogenesis, studies on the genetics of vaccinia virus and the problem of non-genetic reactivation (previously termed 'transformation') and the campaign for the global eradication of smallpox. The other group of poxviruses described here, the genus Leporipoxvirus, came to prominence when the myxoma virus was used for the biological control of Australian wild rabbits. This provided a unique natural experiment on the coevolution of a virus and its host. Future research will include further studies of the many immunomodulatory genes found in all poxviruses of vertebrates, since these provide clues about the workings of the immune system and how viruses have evolved to evade it. Some of the many recombinant poxvirus constructs currently being studied may come into use as vaccines or for immunocontraception. A field that warrants study but will probably remain neglected is the natural history of skunkpox, raccoonpox, taterapox, yabapox, tanapox and other little-known poxviruses. A dismal prospect is the possible use of smallpox virus for bioterrorism.

Animals↗

The dynamical theory of coevolution: a derivation from stochastic ecological processes.

In this paper we develop a dynamical theory of coevolution in ecological communities. The derivation explicitly accounts for the stochastic components of evolutionary change and is based on ecological processes at the level of the individual. We show that the coevolutionary dynamic can be envisaged as a directed random walk in the community's trait space. A quantitative description of this stochastic process in terms of a master equation is derived. By determining the first jump moment of this process we abstract the dynamic of the mean evolutionary path. To first order the resulting equation coincides with a dynamic that has frequently been assumed in evolutionary game theory. Apart from recovering this canonical equation we systematically establish the underlying assumptions. We provide higher order corrections and show that these can give rise to new, unexpected evolutionary effects including shifting evolutionary isoclines and evolutionary slowing down of mean paths as they approach evolutionary equilibria. Extensions of the derivation to more general ecological settings are discussed. In particular we allow for multi-trait coevolution and analyze coevolution under nonequilibrium population dynamics.

Animals↗

Are elicitins cryptograms in plant-Oomycete communications?

Stimulation of plant natural defenses is an important challenge in phytoprotection prospects. In that context, elicitins, which are small proteins secreted by Phytophthora and Pythium species, have been shown to induce a hypersensitive-like reaction in tobacco plants. Moreover, these plants become resistant to their pathogens, and thus this interaction constitutes an excellent model to investigate the signaling pathways leading to plant resistance. However, most plants are not reactive to elicitins, although they possess the functional signaling pathways involved in tobacco responses to elicitin. The understanding of factors involved in this reactivity is needed to develop agronomic applications. In this review, it is proposed that elicitins could interact with regulating cell wall proteins before they reach the plasma membrane. Consequently, the plant reactivity or nonreactivity status could result from the equilibrium reached during this interaction. The possibility of overexpressing the elicitins directly from genomic DNA in Pichia pastoris allows site-directed mutagenesis experiments and structure/function studies. The recent discovery of the sterol carrier activity of elicitins brings a new insight on their molecular activity. This constitutes a crucial property, since the formation of a sterol-elicitin complex is required to trigger the biological responses of tobacco cells and plants. Only the elicitins loaded with a sterol are able to bind to their plasmalemma receptor, which is assumed to be an allosteric calcium channel. Moreover, Phytophthora and Pythium do not synthesize the sterols required for their growth and their fructification, and elicitins may act as shuttles trapping the sterols from the host plants. Sequence analysis of elicitin genes from several Phytophthora species sheds unexpected light on the phylogenetic relationships among the genus, and suggests that the expression of elicitins is under tight regulatory control. Finally, general involvement of these lipid transfer proteins in the biology of Pythiaceae, and in plant defense responses, is discussed. A possible scheme for the coevolution between Phytophthora and tobacco plants is approached.

Algal Proteins↗

Coevolution in temporally variable environments.

Many potentially mutualistic interactions are conditional, with selection that varies between mutualism and antagonism over space and time. We develop a genetic model of temporally variable coevolution that incorporates stochastic fluctuations between mutualism and antagonism. We use this model to determine conditions necessary for the coevolution of matching traits between a host and a conditional mutualist. Using an analytical approximation, we show that matching traits will coevolve when the geometric mean interaction is mutualistic. When this condition does not hold, polymorphism and trait mismatching are maintained, and coevolutionary cycles may result. Numerical simulations verify this prediction and suggest that it remains robust in the presence of temporal autocorrelation. These results are compared with those from spatial models with unrestricted movement. The comparisons demonstrate that gene flow is unnecessary for generating empirical patterns predicted by the geographic mosaic theory of coevolution.

Animals↗

Detecting sexually antagonistic coevolution with population crosses.

The result of population crosses on traits such as mating rate, oviposition rate and survivorship are increasingly used to distinguish between modes of coevolution between the sexes. Two key hypotheses, erected from a verbal theory of sexually antagonistic coevolution, have been the subject of several recent tests. First, statistical interactions arising in population crosses are suggested to be indicative of a complex signal/receiver system. In the case of oviposition rates, an interaction between populations (x, y and z) would be indicated by the rank order of female oviposition rates achieved by x, y and z males changing depending upon the female (x, y or z) with which they mated. Second, under sexually antagonistic coevolution females will do 'best' when mated with their own males, where best is defined by the weakest response to the signal and the highest fitness. We test these hypotheses by crossing strains generated from a formal model of sexually antagonistic coevolution. Strains differ in the strength of natural selection acting on male and female traits. In our model, we assume sexually antagonistic coevolution of a single male signal and female receptor. The female receptor is treated as a preference function where both the slope and intercept of the function can evolve. Our results suggest that neither prediction is consistently supported. Interactions are not diagnostic of complex signal-receiver systems, and even under sexually antagonistic coevolution, females may do better mating with males of strains other than their own. These results suggest a reinterpretation of several recent experiments and have important implications for developing theories of speciation when sexually antagonistic coevolution is involved.

Animals↗

A simple model of host-parasite evolutionary relationships. Parasitism: compromise or conflict?

The evolutionary biology of host-parasite relationships are considered here using a simple game-theory model in which hosts play against parasite and vice versa. In this model, the players can choose between two strategies (aggressive or not aggressive) and the utility of the game is envisaged in terms of fitness and selective costs. The game solutions suggest that the two types of confrontation are encountered in symbiotic relationships and thus constitute two Evolutionary Stable Strategies (ESS). These observations lead us to discuss: (i) the status of different kinds of symbiotic relationships (i.e. parasitoidism; parasitism, commensalism and mutualism) related to selective costs and (ii) the position of coevolution in this game theory context.

Animals↗

Bacteroides thetaiotaomicron: a dynamic, niche-adapted human symbiont.

The coevolution of humans with their intestinal microflora has resulted in cooperative relationships that have shaped the biology and the genomes of these symbiotic partners. Bacteroides thetaiotaomicron is one such bacterial symbiont that is a dominant member of the intestinal microbiota of humans and other mammals. The recent report of the genome sequence of B. thetaiotaomicron is the first reported for an abundant Gram-negative organism of the human colonic microbiota and, as such, provides the first glimpse on a genomic scale of the genetic arsenal used by a Gram-negative symbiont to dominate in this ecosystem. The genome has revealed large expansions of many paralogous groups of genes that encode products essential to the organism's ability to successfully compete in this environment. Most noteable is the organism's abundant machinery for utilizing a large variety of complex polysaccharides as a source of carbon and energy. The proteome also reveals the organism's extensive ability to adapt and regulate expression of its genes in response to the changing ecosystem. These factors, as well as others highlighted below, suggest an incredibly flexible and adaptable organism that is exquisitely equipped to dominate in its challenging and competitive niche.

Bacteroides↗

Coevolution: mathematical analysis of host-parasite interactions.

An S----I----S epidemic transmitted by two similar strains of parasite acting on a host population of three genotypes which differ in their reaction to the disease is modelled and analyzed. Singular perturbation techniques are used to reduce the original system of nine differential equations to a coupled system of two equations describing the slowtime coevolution of gene frequency and parasite strain frequency.

Animals↗