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(PDF) Coevolution of Parasite Virulence and Host Reproductive Effort
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Effort","grobid_abstract":"Epidemiological models generally explore the evolution of parasite life-history traits, namely, virulence and transmission, against a background of constant host life-history traits. However, life-history models have predicted the evolution of host traits in response to parasitism. The coevolution of host and parasite lifehistory traits remains largely unexplored. We present an epidemiological model, based on resource allocation theory, that provides an analysis of the coevolution between host reproductive effort and parasite virulence. This model allows for hosts with either a fixed (i.e., genetic) or conditional (i.e., a phenotypically plastic) response to parasitism. It also considers superinfections. We show that parasitism always favors increased allocation to host reproduction, but because of epidemiological feedbacks, the evolutionarily stable host reproductive effort does not always increase with parasite virulence. Superinfection drives the evolution of parasite virulence and acts on the evolution of the host through parasite evolution, generally leading to higher host reproductive effort. Coevolution, as opposed to cases where only one of the antagonists evolves, may generate correlations between host and parasite life-history traits across environmental gradients affecting the fecundity or the survival of the host. Our results provide a theoretical framework against which experimental coevolution outcomes or field observations can be contrasted.","publication_date":"2002,,","publication_name":"The American Naturalist","grobid_abstract_attachment_id":"78870273"},"document_type":"paper","pre_hit_view_count_baseline":null,"quality":"high","language":"en","title":"Coevolution between parasite virulence and host life‐history traits","broadcastable":false,"draft":null,"has_indexable_attachment":true,"indexable":true}}["work"]; window.loswp.workCoauthors = [48909677]; window.loswp.locale = "en"; window.loswp.countryCode = "SG"; window.loswp.cwvAbTestBucket = ""; window.loswp.designVariant = "ds_vanilla"; window.loswp.fullPageMobileSutdModalVariant = "full_page_mobile_sutd_modal"; window.loswp.useOptimizedScribd4genScript = false; window.loginModal = {}; window.loginModal.appleClientId = 'edu.academia.applesignon';</script><script defer="" src="https://accounts.google.com/gsi/client"></script><div class="ds-loswp-container"><div class="ds-work-card--grid-container"><div class="ds-work-card--container js-loswp-work-card"><div class="ds-work-card--cover"><div class="ds-work-cover--wrapper"><div class="ds-work-cover--container"><button class="ds-work-cover--clickable js-swp-download-button" data-signup-modal="{"location":"swp-splash-paper-cover","attachmentId":78870273,"attachmentType":"pdf"}"><img alt="First page of “Coevolution between parasite virulence and host life‐history traits”" class="ds-work-cover--cover-thumbnail" src="https://0.academia-photos.com/attachment_thumbnails/78870273/mini_magick20220116-32022-1a2zela.png?1642324712" /><img alt="PDF Icon" class="ds-work-cover--file-icon" src="//a.academia-assets.com/images/single_work_splash/adobe_icon.svg" /><div class="ds-work-cover--hover-container"><span class="material-symbols-outlined" style="font-size: 20px" translate="no">download</span><p>Download Free PDF</p></div><div class="ds-work-cover--ribbon-container">Download Free PDF</div><div class="ds-work-cover--ribbon-triangle"></div></button></div></div></div><div class="ds-work-card--work-information"><h1 class="ds-work-card--work-title">Coevolution between parasite virulence and host life‐history traits</h1><div class="ds-work-card--work-authors ds-work-card--detail"><a class="ds-work-card--author js-wsj-grid-card-author ds2-5-body-md ds2-5-body-link" data-author-id="48909677" href="https://independent.academia.edu/SylvainGandon"><img alt="Profile image of Sylvain Gandon" class="ds-work-card--author-avatar" src="//a.academia-assets.com/images/s65_no_pic.png" />Sylvain Gandon</a></div><div class="ds-work-card--detail"><p class="ds-work-card--detail ds2-5-body-sm">2002, The American Naturalist</p><div class="ds-work-card--work-metadata"><div class="ds-work-card--work-metadata__stat"><span class="material-symbols-outlined" style="font-size: 20px" translate="no">visibility</span><p class="ds2-5-body-sm" id="work-metadata-view-count">…</p></div><div class="ds-work-card--work-metadata__stat"><span class="material-symbols-outlined" style="font-size: 20px" translate="no">description</span><p class="ds2-5-body-sm">15 pages</p></div><div class="ds-work-card--work-metadata__stat"><span class="material-symbols-outlined" style="font-size: 20px" translate="no">link</span><p class="ds2-5-body-sm">1 file</p></div></div><script>(async () => { const workId = 68372555; 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However, life-history models have predicted the evolution of host traits in response to parasitism. The coevolution of host and parasite lifehistory traits remains largely unexplored. We present an epidemiological model, based on resource allocation theory, that provides an analysis of the coevolution between host reproductive effort and parasite virulence. This model allows for hosts with either a fixed (i.e., genetic) or conditional (i.e., a phenotypically plastic) response to parasitism. It also considers superinfections. We show that parasitism always favors increased allocation to host reproduction, but because of epidemiological feedbacks, the evolutionarily stable host reproductive effort does not always increase with parasite virulence. Superinfection drives the evolution of parasite virulence and acts on the evolution of the host through parasite evolution, generally leading to higher host reproductive effort. Coevolution, as opposed to cases where only one of the antagonists evolves, may generate correlations between host and parasite life-history traits across environmental gradients affecting the fecundity or the survival of the host. Our results provide a theoretical framework against which experimental coevolution outcomes or field observations can be contrasted.</p><div class="ds-work-card--button-container"><button class="ds2-5-button js-swp-download-button" data-signup-modal="{"location":"continue-reading-button--work-card","attachmentId":78870273,"attachmentType":"pdf","workUrl":"https://www.academia.edu/68372555/Coevolution_between_parasite_virulence_and_host_life_history_traits"}">See full PDF</button><button class="ds2-5-button ds2-5-button--secondary js-swp-download-button" data-signup-modal="{"location":"download-pdf-button--work-card","attachmentId":78870273,"attachmentType":"pdf","workUrl":"https://www.academia.edu/68372555/Coevolution_between_parasite_virulence_and_host_life_history_traits"}"><span class="material-symbols-outlined" style="font-size: 20px" translate="no">download</span>Download PDF</button></div></div></div></div><div data-auto_select="false" data-client_id="331998490334-rsn3chp12mbkiqhl6e7lu2q0mlbu0f1b" data-doc_id="78870273" data-landing_url="https://www.academia.edu/68372555/Coevolution_between_parasite_virulence_and_host_life_history_traits" data-login_uri="https://www.academia.edu/registrations/google_one_tap" data-moment_callback="onGoogleOneTapEvent" id="g_id_onload"></div><div class="ds-top-related-works--grid-container"><div class="ds-related-content--container ds-top-related-works--container"><h2 class="ds-related-content--heading">Related papers</h2><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="0" data-entity-id="87309488" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/87309488/The_Evolution_of_Parasite_Virulence_Superinfection_and_Host_Resistance">The Evolution of Parasite Virulence, Superinfection, and Host Resistance</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="48909677" href="https://independent.academia.edu/SylvainGandon">Sylvain Gandon</a></div><p class="ds-related-work--metadata ds2-5-body-xs">The American Naturalist, 2002</p><p class="ds-related-work--abstract ds2-5-body-sm">We analyze the evolutionary consequences of host resistance (the ability to decrease the probability of being infected by parasites) for the evolution of parasite virulence (the deleterious effect of a parasite on its host). When only single infections occur, host resistance does not affect the evolution of parasite virulence. However, when superinfections occur, resistance tends to decrease the evolutionarily stable (ES) level of parasite virulence. We first study a simple model in which the host does not coevolve with the parasite (i.e., the frequency of resistant hosts is independent of the parasite). We show that a higher proportion of resistant host decreases the ES level of parasite virulence. Higher levels of the efficiency of host resistance, however, do not always decrease the ES parasite virulence. The implications of these results for virulence management (evolutionary consequences of public health policies) are discussed. Second, we analyze the case where host resistance is allowed to coevolve with parasite virulence using the classical gene-for-gene (GFG) model of host-parasite interaction. It is shown that GFG coevolution leads to lower parasite virulence (in comparison with a fully susceptible host population). The model clarifies and relates the different components of the cost of parasitism: infectivity (ability to infect the host) and virulence (deleterious effect) in an evolutionary perspective.</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"The Evolution of Parasite Virulence, Superinfection, and Host Resistance","attachmentId":91555238,"attachmentType":"pdf","work_url":"https://www.academia.edu/87309488/The_Evolution_of_Parasite_Virulence_Superinfection_and_Host_Resistance","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/87309488/The_Evolution_of_Parasite_Virulence_Superinfection_and_Host_Resistance"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="1" data-entity-id="53631624" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/53631624/Coevolution_of_parasite_virulence_and_host_life_history">Coevolution of parasite virulence and host life history</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="3764553" href="https://independent.academia.edu/JacobKoella">Jacob Koella</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Ecology Letters, 2001</p><p class="ds-related-work--abstract ds2-5-body-sm">Most models about the evolutionary interactions between a parasite's virulence and its host's life history neglect two potentially important aspects: epidemiological and coevolutionary feedback. We emphasize their importance by presenting models that describe the coevolution of a semelparous host's age at reproduction and a parasite's virulence in different environmental conditions. In particular, we first show that an epidemiological feedback will lead to a nonmonotonic response of the host's age at reproduction as virulence increases. We then show that the coevolutionary pressure on virulence can lead to complex associations between the host's life history and the parasite's virulence, which would not be expected with more traditional models of host or parasite evolution. Thus, for example, a high mortality rate of the host favours avirulent parasites and late reproduction of the host when the environmental conditions allow the host to grow rapidly, but early reproduction and high virulence when growth is slow.</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Coevolution of parasite virulence and host life history","attachmentId":70386329,"attachmentType":"pdf","work_url":"https://www.academia.edu/53631624/Coevolution_of_parasite_virulence_and_host_life_history","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/53631624/Coevolution_of_parasite_virulence_and_host_life_history"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="2" data-entity-id="53631807" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/53631807/Shared_Control_of_Epidemiological_Traits_in_a_Coevolutionary_Model_of_Host_Parasite_Interactions">Shared Control of Epidemiological Traits in a Coevolutionary Model of Host‐Parasite Interactions</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="3764553" href="https://independent.academia.edu/JacobKoella">Jacob Koella</a></div><p class="ds-related-work--metadata ds2-5-body-xs">The American Naturalist, 2003</p><p class="ds-related-work--abstract ds2-5-body-sm">Most models concerning the evolution of a parasite's virulence and its host's resistance assume that each component of the relationship (transmission, virulence, recovery, etc.) is controlled by either the host or the parasite but not by both. We present a model that describes the coevolution of host and parasite, assuming that the rate of transmission or the virulence depends on both genotypes. The evolution of these traits is constrained by trade-offs that account for costs of defense and attack strategies, in line with previous studies on the separate evolution of the host and the parasite. Considering shared control by the host and the parasite in determining the traits of the relationship leads to several novel predictions. First, the host should evolve maximal investment in defense against parasites with an intermediate replication rate. Second, the evolution of the parasite strongly depends on the way the host's defense is described. Third, the coevolutionary process may lead to decreasing the parasite's virulence as a response to a rise in the host's background mortality, contrary to classical predictions.</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Shared Control of Epidemiological Traits in a Coevolutionary Model of Host‐Parasite Interactions","attachmentId":70386377,"attachmentType":"pdf","work_url":"https://www.academia.edu/53631807/Shared_Control_of_Epidemiological_Traits_in_a_Coevolutionary_Model_of_Host_Parasite_Interactions","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/53631807/Shared_Control_of_Epidemiological_Traits_in_a_Coevolutionary_Model_of_Host_Parasite_Interactions"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="3" data-entity-id="68372554" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/68372554/Host_life_history_and_the_evolution_of_parasite_virulence">Host life history and the evolution of parasite virulence</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="48909677" href="https://independent.academia.edu/SylvainGandon">Sylvain Gandon</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Evolution, 2001</p><p class="ds-related-work--abstract ds2-5-body-sm">We present a general epidemiological model of host-parasite interactions that includes various forms of superinfection. We use this model to study the effects of different host life-history traits on the evolution of parasite virulence. In particular, we analyze the effects of natural host death rate on the evolutionarily stable parasite virulence. We show that, contrary to classical predictions, an increase in the natural host death rate may select for lower parasite virulence if some form of superinfection occurs. This result is in agreement with the experimental results and the verbal argument presented by Ebert and Mangin (1997). This experiment is discussed in the light of the present model. We also point out the importance of superinfections for the effect of nonspecific immunity on the evolution of virulence. In a broader perspective, this model demonstrates that the occurrence of multiple infections may qualitatively alter classical predictions concerning the effects of various host life-history traits on the evolution of parasite virulence.</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Host life history and the evolution of parasite virulence","attachmentId":78870345,"attachmentType":"pdf","work_url":"https://www.academia.edu/68372554/Host_life_history_and_the_evolution_of_parasite_virulence","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/68372554/Host_life_history_and_the_evolution_of_parasite_virulence"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="4" data-entity-id="61378880" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/61378880/Host_parasite_dynamics_and_the_evolution_of_host_immunity_and_parasite_fecundity_strategies">Host-parasite dynamics and the evolution of host immunity and parasite fecundity strategies</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="33928342" href="https://independent.academia.edu/VeijoKaitala">Veijo Kaitala</a></div><p class="ds-related-work--metadata ds2-5-body-xs">1997</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Host-parasite dynamics and the evolution of host immunity and parasite fecundity strategies","attachmentId":74427206,"attachmentType":"pdf","work_url":"https://www.academia.edu/61378880/Host_parasite_dynamics_and_the_evolution_of_host_immunity_and_parasite_fecundity_strategies","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/61378880/Host_parasite_dynamics_and_the_evolution_of_host_immunity_and_parasite_fecundity_strategies"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="5" data-entity-id="53631657" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/53631657/A_correlated_response_of_a_parasite_s_virulence_and_life_cycle_to_selection_on_its_host_s_life_history">A correlated response of a parasite’s virulence and life cycle to selection on its host’s life history</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="3764553" href="https://independent.academia.edu/JacobKoella">Jacob Koella</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Journal of Evolutionary Biology, 1999</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"A correlated response of a parasite’s virulence and life cycle to selection on its host’s life history","attachmentId":70386356,"attachmentType":"pdf","work_url":"https://www.academia.edu/53631657/A_correlated_response_of_a_parasite_s_virulence_and_life_cycle_to_selection_on_its_host_s_life_history","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/53631657/A_correlated_response_of_a_parasite_s_virulence_and_life_cycle_to_selection_on_its_host_s_life_history"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="6" data-entity-id="22323876" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/22323876/Host_Availability_and_the_Evolution_of_Parasite_Life_History_Strategies">Host Availability and the Evolution of Parasite Life-History Strategies</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="43641886" href="https://independent.academia.edu/AndyFenton">Andy Fenton</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Evolution, 2007</p><p class="ds-related-work--abstract ds2-5-body-sm">Parasites exploit an inherently patchy resource, their hosts, which are discrete entities that may only be available for infection within a relatively short time window. However, there has been little consideration of how heterogeneities in host availability may affect the phenotypic or genotypic composition of parasite populations or how parasites may evolve to cope with them. Here we conduct a selection experiment involving an entomopathogenic nematode (Steinernema feltiae) and show for the first time that the infection rate of a parasite can evolve rapidly to maximize the chances of infecting within an environment characterized by the rate of host availability. Furthermore, we show that the parasite's infection rate trades off with other fitness traits, such as fecundity and survival. Crucially, the outcome of competition between strains with different infection strategies depends on the rate of host availability; frequently available hosts favor "fast" infecting nematodes, whereas infrequently available hosts favor "slow" infecting nematodes. A simple evolutionarily stable strategy (ESS) analysis based on classic epidemiological models fails to capture this behavior, predicting instead that the fastest infecting phenotype should always dominate. However, a novel model incorporating more realistic, discrete bouts of host availability shows that strain coexistence is highly likely. Our results demonstrate that heterogeneities in host availability play a key role in the evolution of parasite life-history traits and in the maintenance of phenotypic variability. Parasite life-history strategies are likely to evolve rapidly in response to changes in host availability induced by disease management programs or by natural dynamics in host abundance. Incorporating parasite evolution in response to host availability would therefore enhance the predictive ability of current epidemiological models of infectious disease.</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Host Availability and the Evolution of Parasite Life-History Strategies","attachmentId":42961459,"attachmentType":"pdf","work_url":"https://www.academia.edu/22323876/Host_Availability_and_the_Evolution_of_Parasite_Life_History_Strategies","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/22323876/Host_Availability_and_the_Evolution_of_Parasite_Life_History_Strategies"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="7" data-entity-id="11017127" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/11017127/The_Implications_of_Coevolutionary_Dynamics_to_Host_Parasite_Interactions">The Implications of Coevolutionary Dynamics to Host‐Parasite Interactions</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="26667611" href="https://independent.academia.edu/AndyWhite11">Andy White</a></div><p class="ds-related-work--metadata ds2-5-body-xs">American Naturalist, 2009</p><p class="ds-related-work--abstract ds2-5-body-sm">Due to the importance of infectious disease, there is a large body of theory on the evolution of either hosts or, more commonly, parasites. Here we present a fully coevolutionary model of a host-parasite system that includes ecological dynamics that feed back into the coevolutionary outcome, and we show that highly virulent parasites may evolve due to the coevolutionary process. Parasite evolution is very sensitive to evolution in the host, and virulence fluctuates substantially when mutation rates vary between host and parasite. Evolutionary branching in the host leads to parasites increasing their virulence, and small changes in host resistance drive large changes in parasite virulence. Evolutionary branching in one species does not cause branching in the other. Our work emphasizes the importance of considering coevolutionary dynamics and shows that certain highly virulent parasites may result from responses to host evolution.</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"The Implications of Coevolutionary Dynamics to Host‐Parasite Interactions","attachmentId":36741132,"attachmentType":"pdf","work_url":"https://www.academia.edu/11017127/The_Implications_of_Coevolutionary_Dynamics_to_Host_Parasite_Interactions","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/11017127/The_Implications_of_Coevolutionary_Dynamics_to_Host_Parasite_Interactions"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="8" data-entity-id="10682025" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/10682025/Ecological_conditions_that_favor_the_evolution_of_intermediate_virulence_in_an_environmentally_transmitted_parasite">Ecological conditions that favor the evolution of intermediate-virulence in an environmentally transmitted parasite</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="26074682" href="https://independent.academia.edu/ErnestBarany">Ernest Barany</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Journal of Mathematical Biology, 2005</p><p class="ds-related-work--abstract ds2-5-body-sm">In this paper we develop and analyze several population-dynamic models of an environmentally transmitted symbiotic parasite infecting an isolated population of susceptible hosts. In our most basic model infection acts only to decrease the average lifetime of the infected host, parasites are only transmitted to uninfected hosts, there is no recovery from infection, and the rate of parasite transmission is an increasing function of the level of parasite virulence. It is shown that invasion of the parasite-free equilibrium cannot occur for virulence levels that are either too high or too low. We then incorporate a number of modifications to the model, among them the possibility that host fertility is reduced by infection, and that transmission rate depends additionally on susceptible host density. It is shown that the essential nature of the conditions for invasion are preserved. Thus, natural selection for intermediate virulence is a generic property of a broad class of population models.</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Ecological conditions that favor the evolution of intermediate-virulence in an environmentally transmitted parasite","attachmentId":47207889,"attachmentType":"pdf","work_url":"https://www.academia.edu/10682025/Ecological_conditions_that_favor_the_evolution_of_intermediate_virulence_in_an_environmentally_transmitted_parasite","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/10682025/Ecological_conditions_that_favor_the_evolution_of_intermediate_virulence_in_an_environmentally_transmitted_parasite"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="9" data-entity-id="12701674" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/12701674/Evolutionary_implications_for_interactions_between_multiple_strains_of_host_and_parasite">Evolutionary implications for interactions between multiple strains of host and parasite</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="31770419" href="https://independent.academia.edu/ZhilanFeng">Zhilan Feng</a><span>, </span><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="31721515" href="https://independent.academia.edu/GregorySandland">Gregory Sandland</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Journal of Theoretical Biology, 2007</p><p class="ds-related-work--abstract ds2-5-body-sm">The interaction between multiple parasite strains within different host types may influence the evolutionary trajectories of parasites. In this article, we formulate a deterministic model with two strains of parasites and two host types in order to investigate how heterogeneities in parasite virulence and host life-history may affect the persistence and spread of diseases in natural systems. We compute the reproductive number of strain i (R i ) independently, as well as the (conditional) ''invasion'' reproductive number for strains i (R j i , jai) when strain j is at a positive equilibrium. We show that the disease-free equilibrium is locally asymptotically stable if R i o1 for both strains and is unstable if R i 41 for one stain. We establish the criterion R j i 41 for strain i to invade strain j. Subthreshold coexistence driven by coinfection is possible even when R i of one strain is below 1. We identify conditions that determine the evolution of parasite specialism or generalism based on the life-history strategies employed by hosts, and investigate how host strains may influence parasite persistence. r</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Evolutionary implications for interactions between multiple strains of host and parasite","attachmentId":45996709,"attachmentType":"pdf","work_url":"https://www.academia.edu/12701674/Evolutionary_implications_for_interactions_between_multiple_strains_of_host_and_parasite","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/12701674/Evolutionary_implications_for_interactions_between_multiple_strains_of_host_and_parasite"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div></div></div><div class="ds-sticky-ctas--wrapper js-loswp-sticky-ctas hidden"><div class="ds-sticky-ctas--grid-container"><div class="ds-sticky-ctas--container"><button class="ds2-5-button js-swp-download-button" data-signup-modal="{"location":"continue-reading-button--sticky-ctas","attachmentId":78870273,"attachmentType":"pdf","workUrl":null}">See full PDF</button><button class="ds2-5-button ds2-5-button--secondary js-swp-download-button" data-signup-modal="{"location":"download-pdf-button--sticky-ctas","attachmentId":78870273,"attachmentType":"pdf","workUrl":null}"><span class="material-symbols-outlined" style="font-size: 20px" translate="no">download</span>Download PDF</button></div></div></div><div class="ds-below-fold--grid-container"><div class="ds-work--container js-loswp-embedded-document"><div class="attachment_preview" data-attachment="Attachment_78870273" style="display: none"><div class="js-scribd-document-container"><div class="scribd--document-loading js-scribd-document-loader" style="display: block;"><img alt="Loading..." src="//a.academia-assets.com/images/loaders/paper-load.gif" /><p>Loading Preview</p></div></div><div style="text-align: center;"><div class="scribd--no-preview-alert js-preview-unavailable"><p>Sorry, preview is currently unavailable. You can download the paper by clicking the button above.</p></div></div></div></div><div class="ds-sidebar--container js-work-sidebar"><div class="ds-related-content--container"><h2 class="ds-related-content--heading">Related papers</h2><div class="ds-related-work--container js-related-work-sidebar-card" data-collection-position="0" data-entity-id="12507848" data-sort-order="default"><a class="ds-related-work--title js-related-work-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/12507848/The_evolution_of_host_parasite_range">The evolution of host-parasite range</a><div class="ds-related-work--metadata"><a class="js-related-work-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="31377409" href="https://independent.academia.edu/EKisdi">E. 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Life history traits of hosts and parasites interact to influence the fitness payoffs of alternative anti-parasite strategies</a><div class="ds-related-work--metadata"><a class="js-related-work-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="34537691" href="https://unc.academia.edu/MariaServedio">Maria Servedio</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Journal of Evolutionary Biology, 2006</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"To eject or to abandon? Life history traits of hosts and parasites interact to influence the fitness payoffs of alternative anti-parasite strategies","attachmentId":51798464,"attachmentType":"pdf","work_url":"https://www.academia.edu/31436488/To_eject_or_to_abandon_Life_history_traits_of_hosts_and_parasites_interact_to_influence_the_fitness_payoffs_of_alternative_anti_parasite_strategies","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-related-work-grid-card-view-pdf" href="https://www.academia.edu/31436488/To_eject_or_to_abandon_Life_history_traits_of_hosts_and_parasites_interact_to_influence_the_fitness_payoffs_of_alternative_anti_parasite_strategies"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-related-work-sidebar-card" data-collection-position="3" data-entity-id="2691272" data-sort-order="default"><a class="ds-related-work--title js-related-work-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/2691272/Virulence_and_age_at_reproduction_new_insights_into_host_parasite_coevolution">Virulence and age at reproduction: new insights into host–parasite coevolution</a><div class="ds-related-work--metadata"><a class="js-related-work-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="305405" href="https://cnrs.academia.edu/MichaelHochberg">Michael Hochberg</a></div><p class="ds-related-work--metadata ds2-5-body-xs">2001</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Virulence and age at reproduction: new insights into host–parasite coevolution","attachmentId":30684250,"attachmentType":"pdf","work_url":"https://www.academia.edu/2691272/Virulence_and_age_at_reproduction_new_insights_into_host_parasite_coevolution","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-related-work-grid-card-view-pdf" href="https://www.academia.edu/2691272/Virulence_and_age_at_reproduction_new_insights_into_host_parasite_coevolution"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-related-work-sidebar-card" data-collection-position="4" data-entity-id="19998332" data-sort-order="default"><a class="ds-related-work--title js-related-work-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/19998332/The_unavoidable_costs_and_unexpected_benefits_of_parasitism_Population_and_metapopulation_models_of_parasite_mediated_competition">The unavoidable costs and unexpected benefits of parasitism: Population and metapopulation models of parasite-mediated competition</a><div class="ds-related-work--metadata"><a class="js-related-work-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="40899475" href="https://independent.academia.edu/ChihhorngKuo">Chih-horng Kuo</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Journal of Theoretical Biology, 2008</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"The unavoidable costs and unexpected benefits of parasitism: Population and metapopulation models of parasite-mediated competition","attachmentId":40950885,"attachmentType":"pdf","work_url":"https://www.academia.edu/19998332/The_unavoidable_costs_and_unexpected_benefits_of_parasitism_Population_and_metapopulation_models_of_parasite_mediated_competition","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-related-work-grid-card-view-pdf" href="https://www.academia.edu/19998332/The_unavoidable_costs_and_unexpected_benefits_of_parasitism_Population_and_metapopulation_models_of_parasite_mediated_competition"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-related-work-sidebar-card" data-collection-position="5" data-entity-id="117622831" data-sort-order="default"><a class="ds-related-work--title js-related-work-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/117622831/Direct_and_Correlated_Responses_to_Selection_in_a_Host_Parasite_System_Testing_for_the_Emergence_of_Genotype_Specificity">Direct and Correlated Responses to Selection in a Host-Parasite System: Testing for the Emergence of Genotype Specificity</a><div class="ds-related-work--metadata"><a class="js-related-work-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="291404642" href="https://independent.academia.edu/PhookanArundhati">Arundhati Phookan</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Evolution, 2007</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Direct and Correlated Responses to Selection in a Host-Parasite System: Testing for the Emergence of 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data-author-id="45028606" href="https://independent.academia.edu/AKarter">A Karter</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Trends in Ecology & Evolution, 1990</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Parasite-host coevolution","attachmentId":68163862,"attachmentType":"pdf","work_url":"https://www.academia.edu/50025425/Parasite_host_coevolution","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-related-work-grid-card-view-pdf" href="https://www.academia.edu/50025425/Parasite_host_coevolution"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" 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