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PubMed1991

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↗

PubMed1984

Stasis: a coevolutionary model.

Traditional neo-Darwinian explanations for stasis involve limited variation, developmental constraints and stabilizing selection. Of these, stabilizing selection is regarded as the mechanism operating most widely. Arguments based on stabilizing selection, however, implicitly assume a one-way evolutionary relationship between organism and environment. In this paper, I suggest that stasis may arise in a number of different ways as a result of organism/environment coevolution. The chief causes of stasis may be the attainment of coadapted equilibria between organism and environment and periods of quiescence within and between arms races. I also suggest that many cases of stasis in the fossil records may be apparent rather than real due to a reliance on gross morphological trends and that apparently large environmental changes during which stasis persists may not reflect change in the coadapted components of the organisms' environment.

Adaptation, Physiological↗

PubMed2003

Runaway sexual selection with paternal transmission of the male trait and gene-culture determination of the female preference.

Sexual selection is modeled with a male viability-reducing trait and a female mating preference for that trait both of which are culturally transmitted. Both the male trait and the female preference are transmitted only between same-sex individuals, so that non-random association between the trait and the preference, which would give rise to a Fisherian runaway process, cannot arise. Inclusion of an autosomal gene that confers a female predisposition to acquire a certain preference is shown to allow the coevolution of the male trait and the female preference by a Fisherian process. This holds true even when the female preference has a slight viability cost, provided the male cultural transmission is not perfect. It is also suggested that a Fisherian process can be more easily initiated in these models than in the conventional genetic models. Furthermore, a Fisherian process may cause cultural transmission of female preference to evolve. Additionally, polymorphism can be maintained at the predisposition locus if heterozygous females have a stronger predisposition to acquire the preference than homozygotes. Our models may be applicable to the case when the male trait is a Y-linked genetic or environmentally determined trait.

Animals↗

PubMed2002

The ecology of fish parasites with particular reference to helminth parasites and their salmonid fish hosts in Welsh rivers: a review of some of the central questions.

Ecological studies carried out in Welsh rivers on the feeding behaviour of salmonid fish, their helminth parasites and intermediate hosts in the early 1950s and in 1998 have been used as a basis to review the literature dealing with the following questions. First, how are the helminth populations dispersed in space-time? Second, to what extent are the distributional patterns and the life history strategies of the parasites influenced by physicochemical factors? Third, to what extent are populations of helmith parasites in salmonid fish influenced by host characteristics including the genome, sex, age, size, social position and Feeding behaviour? Fourth, are the populations of parasites regulated in a density-dependent manner? Fifth, do the parasites influence the survival and wellbeing of their salmonid hosts and the evolution of sex? Sixth, to what extent is the parasite community influenced by environmental changes including those of an anthropogenic nature and can the parasites be used as bioindicators of pollution? As with most parasites the helminth species found were highly overdispersed thus making it necessary to undertake a log10 (1 + x) conversion for statistical analyses. Statistical analyses confirm that the genome, age and sex of salmonid fish hosts, the station and seasonal change in radiation levels were significant factors in predicting the number of parasites. The evidence given supports the hypothesis that the feeding behaviour and habitat selection by the host fish, their position in the social hierarchy and the overdispersed nature of the transmission sites are the key factors in causing differences in the parasitic fauna related to host species, age, size and sex. Differences in the helminth parasite community related to station can be explained on the basis of differences in water types, sediments and chemistry. Although the evidence presented is in accord with the consensus view that temperature is correlated with seasonal changes in the abundance of many species of helminth parasites, it is argued that it may not be the direct causative mechanism. It is postulated that the life history strategy that results in a decline in abundance of the more vulnerable adult parasites in the gut of the salmonid hosts during the summer has arisen as a result of evolutionary pressures. At this time, the gut environment is particularly inhospitable because of the temperature-related enhancement of the host's immune mechanism and the increased gut turnover rate. In contrast, the larval stages in the immunologically and metabolically more benign intermediate host would be under less intensive selective pressures. It is postulated therefore that evolutionary pressures have caused the parasites to leave the definitive host and concentrate their reproductive efforts in the intermediate hosts during the warmer months. Evidence is given in support of the hypothesis that the parasite populations are regulated in a density-dependent manner and that the regulatory mechanisms may involve the host's immune mechanisms and intraspecies competition and interspecies competition of an exploitative or interference nature. Quantitative studies using 'K' factor analysis and biochemical research to elucidate the nature of the interference mechanisms are required to test this hypothesis. The absence of age-related resistance indicates an old and stable relationship in which the immunosuppressive and immunoavoidance mechanisms of the parasites and hosts, respectively, are in balance. This indicates that the introduction of novel parasites or new genetic strains of host fish could result in harmful epidemics. Despite causing tissue damage, there was no evidence of parasite-induced mortality among the salmonids in the Teifi. This finding is in accord with the generally accepted view that most freshwaters are not troubled by parasite problems. although parasites are present in abundance. In fact, parasite abundance in the salmonid fish in the Teifi was positively correlated with the condition factor and the adipose index. Two testable hypotheses were advanced to explain these observations. First, the more dominant well-conditioned fish in the hierarchy are more likely to acquire parasites because they ingest more food items and spend more time in sheltered habitats with depositing sediments where transmission mainly occurs. Second, the parasites may release factors that stimulate the host's immune and endocrinological systems to produce factors that enhance somatic growth and inhibit reproduction of the host. This benign relationship is considered to be indicative of long-term coevolution. The sex of the fish had a significant influence on the abundance of the parasites in total and also on particular species with the bias in all cases being in favour of the female fish. This review shows that sex bias in parasitism is generally not strong and that male bias in parasitism is not a general rule. Taken as a whole, the results fail to support most of the predictions based on the Hamilton-Zuk and the immunocompetence hypotheses. Possible hypotheses to explain why parasitism tends to be higher in female than in male trout include testosterone immunosuppression, corticosteroid-based immune suppression and differences between the size and behaviour of the sexes. However, the latter two hypotheses have more credence, although testosterone levels are higher in female than male trout. Between the early 1950s and 1998 there has been a marked decline in the prevalence, abundance and diversity of the helminth parasite communities in salmonid fish as well as their intermediate hosts. Possible reasons for these declines include heavy metal pollution, increased acidity and habitat degradation linked to changes in land use. It is concluded that although helminth parasites can provide supplementary information on pollution. the use of biotic indices based on the Biological monitoring working party (BMWP) or River invertebrate prediction and classification system (RIVPACS) methods are preferable. However, as these methods were designed to measure the impact of organic pollution they lack the sensitivity for measuring metal pollution. It is advocated therefore that new biomonitoring methods should be developed to measure the impact of heavy metal pollution using biotic indices based on the sampling of the susceptible invertebrate communities inhabiting depositing sediments in the transmission sites of helminth parasites.

Animals↗

PubMed2003

Phylogenetic insights into the evolution of parasitism in hymenoptera.

The Hymenoptera are one of the four megadiverse orders of insects, with over 100000 described species and several times this number still waiting to be described. A major part of this diverse group is formed of large lineages of parasitoid wasps. Some of these lineages have in turn given rise to subgroups that have gone on to diversify into other lifestyles, such as gall-forming on, and pollination of, plants, as well as a broad array of food-collecting behaviors associated with social living in colonies. Thus, the Hymenoptera demonstrate the large evolutionary potential of parasitism as a lifestyle, in contrast to early assertions that parasitism tends to lead to evolutionary 'dead ends' driven by overspecialization. Phylogenetic approaches have already led to a number of important insights into the evolution of parasitism in Hymenoptera. A series of examples are discussed in this review, including the origin of parasitism in the order, the development of koinobiosis in some groups, coevolution with symbiotic viruses, and the evolution in some groups away from parasitism and into such habits as gall formation, pollination of figs, nest building and sociality. The potential for comparative analysis of hymenopteran habits is large, but progress is still in its early stages due to the paucity of available well-supported phylogenies, and the still limited accumulation of basic biological data for many taxa.

Animals↗

PubMed2003

Life cycle evolution in the digenea: a new perspective from phylogeny.

We use a new molecular phylogeny, developed from small and large subunit ribosomal RNA genes, to explore evolution of the digenean life cycle. Our approach is to map character states on the phylogeny and then use parsimony to infer how the character evolved. We conclude that, plesiomorphically, digenean miracidia hatched from eggs and penetrated gastropod first intermediate hosts externally. Fork-tailed cercariae were produced in rediae and emerged from the snail to be eaten directly by the teleost definitive host. These plesiomorphic characters are seen in extant Bivesiculidae. We infer that external encystment and the use of second intermediate hosts are derived from this behaviour and that second intermediate hosts have been adopted repeatedly. Tetrapod definitive hosts have also been adopted repeatedly. The new phylogeny proposes a basal dichotomy between 'Diplostomida' (Diplostomoidea, Schistosomatoidea and Brachylaimoidea) and 'Plagiorchiida' (all other digeneans). There is no evidence for coevolution between these clades and groups of gastropods. The most primitive life cycles are seen in basal Plagiorchiida. Basal Diplostomida have three-host life cycles and are associated with tetrapods. The blood flukes (Schistosomatoidea) are inferred to have derived their two-host life cycles by abbreviating three-host cycles. Diplostomida have no adult stages in fishes except by life cycle abbreviation. We present and test a radical hypothesis that the blood-fluke cycle is plesiomorphic within the Diplostomida.

Animals↗

PubMed2001

The emergence of life on Earth.

Combined top-down and bottom-up research strategies and the principle of biological continuity were employed in an attempt to reconstruct a comprehensive origin of life theory, which is an extension of the coevolution theory (Lahav and Nir, Origins of Life Evol. Biosphere (1997) 27, 377-395). The resulting theory of emergence of templated-information and functionality (ETIF) addresses the emergence of living entities from inanimate matter, and that of the central mechanisms of their further evolution. It proposes the emergence of short organic catalysts (peptides and proto-ribozymes) and feedback-loop systems, plus their template-and-sequence-directed (TSD) reactions, encompassing catalyzed replication and translation of populations of molecules organized as chemical-informational feedback loop entities, in a fluctuating (wetting-drying) environment, functioning as simplified extant molecular-biological systems. The feedback loops with their TSD systems are chemically and functionally continuous with extant living organisms and their emergence in an inanimate environment may be defined as the beginning of life. The ETIF theory considers the emergence of bio-homochirality, a primordial genetic code, information and the incorporation of primordial metabolic cycles and compartmentation into the emerging living entities. This theory helps to establish a novel measure of biological information, which focuses on its physical effects rather than on the structure of the message, and makes it possible to estimate the time needed for the transition from the inanimate state to the closure of the first feedback-loop systems. Moreover, it forms the basis for novel laboratory experiments and computer modeling, encompassing catalytic activity of short peptides and proto-RNAs and the emergence of bio-homochirality and feedback-loop systems.

Biological Evolution↗

PubMed2004

Structural determinants of allosteric ligand activation in RXR heterodimers.

Allosteric communication underlies ligand-dependent transcriptional responses mediated by nuclear receptors. While studies have elucidated many of the components involved in this process, the energetic architecture within the receptor protein that mediates allostery remains unknown. Using a sequence-based method designed to detect coevolution of amino acids in a protein, termed the statistical coupling analysis (SCA), we identify a network of energetically coupled residues that link the functional surfaces of nuclear receptor ligand binding domains. Functional analysis of these predicted residues demonstrates their participation in an allosteric network that governs the ability of heterodimeric receptors to activate transcription in response to ligand binding by either partner. Interestingly, mutation of a single network residue can discriminate between receptor activation by endocrine, dietary, and synthetic agonists. These results reveal a structural network required for RXR heterodimer allosteric communication and suggest that the specificity of ligand response and permissivity coevolved to enable signal discrimination.

Allosteric Regulation↗

PubMed1991

Co-evolution of parasites and adaptive immune responses.

The interplay between evolving host populations and evolving parasite populations is dominated by two key genetically based elements, namely the virulence of parasites and the resistance of their hosts. Here, Graham Mitchell gives a personal overview of ideas on the coevolution of parasite-host relationships and the contributions to immunology that are likely to emerge from systematic studies on this relationship.

Adaptation, Biological↗

PubMed2000

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↗

PubMed2000

Are there pros as well as cons to being parasitized?

The diversity of ways in which parasites reduce the fitness of their hosts has been documented during the past decades, and clearly indicates that parasites can often be considered as direct agents of selection. In natural systems, however, the outcome of a host-parasite interaction might be strongly determined by other ecological factors. Parasites can be detrimental to host fitness in one environment, whereas they can be beneficial to it in another. From an evolutionary perspective, this phenomenon is of considerable importance for understanding the dynamics of coevolution among geographically structured populations evolving under different ecological pressures. Here, Frédéric Thomas and colleagues review several ecological situations in which parasitized individuals enjoy a selective advantage over unparasitized conspecifics.

Animals↗

PubMed2000

Persistence of selfish genetic elements: population structure and conflict.

Selfish genetic elements are vertically transmitted factors that spread by obtaining a transmission advantage relative to the rest of the genome of their host organism, often with a cost to overall host fitness. In many cases, conventional population genetics theory predicts them spreading through populations, reaching fixation and becoming undetectable or sometimes driving the population extinct. However, in several well studied systems, these genetic elements are known to persist at relatively low, stable frequencies. Recent research suggests that several processes might explain these observations, including population structure, intragenomic conflict and coevolution.

Journal Article↗

PubMed2000

Polymorphism of attack and defense.

Coevolution of attack and defense occurs in host-parasite systems and various forms of genomic conflict. Attackers benefit when their specific molecules allow entry past host defenses. Defenders gain when their matching biochemical specificities aid recognition. Selection continually favors new attack specificities that avoid matching defense and, in turn, new defense specificities that match novel attackers. The introduction of novel specificities strongly influences the spatial and temporal dynamics of conflict. Lack of reciprocally matching diversity in a particular system suggests biochemical constraints that prevent diversification. New work on cytoplasmic male sterility, B chromosomes and meiotic drive suggests that varying biochemical constraints on recognition cause varying patterns of diversity and spatiotemporal dynamics

Journal Article↗

PubMed1991

Biology of ticks.

Ticks are an ancient group of obligate bloodsucking ectoparasites that has evolved over millions of years. Two general types of ticks are evident today: argasid or soft ticks, and ixodid or hard ticks. Each lineage exhibits distinct patterns of host coevolution and preference. However, about 10% of the approximately 850 species are of medical importance because of their indiscriminate host selection and catholic feeding behavior. As a result, a number of diseases have begun to emerge in the temperate zones, including Lyme borreliosis and several others putatively associated with ticks. Ticks may serve as both pathogens and disease vectors. Because of the unique physiology of the salivary glands and the contents in tick saliva of toxins, feeding alone may cause disease. Ticks also transmit a number of different types of pathogens (viruses, rickettsiae, spirochetes and bacteria, fungi, protozoa, filarial nematodes) and even exceed mosquitoes in this regard. Abatement and control of ticks emphasizes a broad approach because of the differing types of habitats in which pest species may be found. The use of repellents and acaricides as well as cultural and management practices are of primary importance. Other approaches (ivermectin) may be beneficial; with the advent of molecular genetics and its usefulness in immunology, the development of tick vaccines for common pest species appears promising.

Animals↗

PubMed1996

The HLA-DRB9 gene and the origin of HLA-DR haplotypes.

HLA-DRB9 is a gene fragment consisting of exon 2 and flanking intron sequences. It is located at the extreme end of the DRB subregion, whose other end is demarcated by the DRB1 locus. We sequenced approximately 1400 base pairs of the segment encompassing the DRB9 locus from eight human haplotypes (DR1, DR10, DR2, DR3, DR5, DR6, DR8, and DR9, the DR4 and DR7 having been sequenced by others earlier), as well as two chimpanzee, five gorillas, one orangutan and one macaque haplotype. The analysis of these sequences indicates that the DRB9 locus, which we estimate to be more than 58 million years (my) old, has been coevolving with the DRB1 locus for the last 4.2 my. As a consequence of this coevolution, the human DRB9 alleles fall into groups that correlate with the DRB1 allelic groups and with the gene organization of the human haplotypes. This observation implies that the present-day HLA-DR haplotype groups (DR1, DR51, DR52, DR8, and DR53) were founded more than 4 my ago and have remained intact (barring minor internal rearrangements that did not recombine the DRB1 and DRB9 genes) for this period of time. The haplotypes have been transmitted during speciations from ancestral to emerging species just like allelic lineages at the DRB1 locus. Thus not only allelic but also haplotype polymorphism evolves trans-specifically.

Animals↗

PubMed2002

Full-length sequence of VERL, the egg vitelline envelope receptor for abalone sperm lysin.

Abalone sperm use 16 kDa lysin to create a hole in the egg vitelline envelope (VE) by a species-specific, nonenzymatic mechanism. To create the hole, lysin binds tightly to VERL (the VE receptor for lysin), a giant, unbranched glycoprotein comprising 30% of the VE. Binding of lysin to VERL causes the VERL molecules to lose cohesion and splay apart creating the hole. Lysin and VERL represent a cognate pair of gamete recognition proteins, one male the other female, which mediate fertilization. The coevolution of such cognate pairs may underlie the establishment of species-specific fertilization which could be a component of the mechanism to achieve reproductive isolation and hence new species. Here we present the full-length cDNA sequence (11,166 bp) of VERL from the red abalone (Haliotis rufescens). There are 42 amino acids from the start Met residue to the beginning of the first 'VERL repeat'. Most of VERL (9981 bp; 89.4%) consists of 22 tandem repeats of a approximately 153 amino acid sequence that is predicted to be beta-sheet. The last VERL repeat is followed by 353 non-repeat amino acid residues containing a furin cleavage site (RTRR), a ZP domain and a hydrophobic COOH-terminus with a 3' UTR of only 10 nucleotides. VERL repeats 3-22 have been subjected to concerted evolution and consequently have almost identical sequences. Curiously, comparisons of repeats from other species shows that repeats 1 and 2 of red abalone VERL have not been subjected to concerted evolution since the divergence of the red species from the other six California species.

Amino Acid Sequence↗

PubMed2003

A novel gene family encoding leucine-rich repeat transmembrane proteins differentially expressed in the nervous system.

Leucine-rich repeat containing proteins are involved in protein-protein interactions and they regulate numerous cellular events during nervous system development and disease. Here we have isolated and characterized a new four-membered family of genes from human and mouse, named LRRTMs, that encode putative leucine-rich repeat transmembrane proteins. Human and mouse LRRTMs are highly conserved, and orthologous genes exist in other vertebrates but not in invertebrates. All LRRTMs, except LRRTM4, are located in the introns of different alpha-catenin genes, suggesting coevolution of these two gene families. We show by in situ hybridization and RT-PCR that LRRTM mRNAs are predominantly expressed in the nervous system and that each LRRTM possesses a specific, partially nonoverlapping expression pattern. The structure and expression profile of LRRTM mRNAs suggest that they may have a role in the development and maintenance of the vertebrate nervous system.

Amino Acid Sequence↗

PubMed1997

Biochemical, molecular, and phylogenetic analysis of pyruvate carboxylase in the yellow fever mosquito, Aedes aegypti.

Pyruvate carboxylase (PC, pyruvate: carbon dioxide ligase [ADP-forming], EC 6.4.1.1) was purified from the yellow fever mosquito, Aedes aegypti. The purified PC showed two polypeptides of similar M(r) (133 and 128 k). The N-terminal sequences of both polypeptides were shown to be very similar, if not identical. A polyclonal antiserum against the 133 kDa polypeptide cross-reacted strongly with the 128 kDa polypeptide. PC was found in all tissues examined. Using a semi-quantitative Western blot assay, PC was shown to be concentrated in the indirect flight muscles and fat body preparations. The ratios of the 133 to 128 kDa polypeptides were shown to differ in various tissues and an Aedes albopictus cell line. The indirect flight muscle was the only tissue in which the 128 kDa polypeptide was more abundant, while both the midgut and the cell line showed almost exclusively the 133 kDa polypeptide. Both peptides were present in varying amounts in brain, malpighian tubule, ovary and fat body preparation. The two isoforms of PC could play different roles in the flight muscle and other tissues. Clones covering a complete cDNA of PC of A. aegypti were obtained using a directional approach. The 3952 bp nucleotide sequence, including a 3585 bp coding region, was determined from these cDNA clones. The deduced 1195 amino acid sequence has a calculated M(r) of 132,200. A putative mitochondrial targeting sequence was determined by comparing the deduced amino acid sequence to the N-terminal sequences of the mature protein. The presence of a mitochondrial targeting sequence indicates that the mosquito PC encoded by the cloned cDNA may be localized in the mitochondria. After the targeting sequence, three functional domains were identified in the following order; biotin carboxylase (BC), carboxyltransferase (CT) and biotin carboxyl carrier protein (BCCP). The mosquito PC showed very high similarity to PCs from other sources (55.1-75.2% identity). Genomic Southern analysis indicated that there could be two similar PC genes or a single PC gene with allelic polymorphism in the A. aegypti genome. The evolutionary relationship of PCs among different organisms was consistent with the accepted evolutionary relationship of their host organisms. The evolution of the domain structures of the biotin-dependent carboxylases including PC was also investigated. This analysis indicates that biotin-dependent carboxylases evolved from a common origin. The analysis also provides evidence for early gene duplication events that shaped the family of biotin-dependent carboxylases. Clear evidence for the coevolution of BC and BCCP domains is presented, although they are associated with very different CT domains and the relative position of the three functional domains varies between members of the biotin-dependent carboxylases.

Aedes↗