{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,25]],"date-time":"2026-09-25T19:30:30Z","timestamp":1790364630589,"version":"4.1.0"},"update-to":[{"DOI":"10.12688\/wellcomeopenres.11594.1","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2017,7,24]],"date-time":"2017-07-24T00:00:00Z","timestamp":1500854400000}}],"reference-count":35,"publisher":"F1000 Research Ltd","license":[{"start":{"date-parts":[[2017,7,24]],"date-time":"2017-07-24T00:00:00Z","timestamp":1500854400000},"content-version":"unspecified","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100007197","name":"U.S. Public Health Service","doi-asserted-by":"crossref","award":["AI054961"],"award-info":[{"award-number":["AI054961"]}],"id":[{"id":"10.13039\/100007197","id-type":"DOI","asserted-by":"crossref"}]},{"name":"Wellcome Trust","award":["103875"],"award-info":[{"award-number":["103875"]}]},{"name":"European Research Council","award":["ERC-2012-StG 309255-EndoTox"],"award-info":[{"award-number":["ERC-2012-StG 309255-EndoTox"]}]},{"name":"Wellcome Trust","award":["085349"],"award-info":[{"award-number":["085349"]}]},{"DOI":"10.13039\/100007197","name":"U.S. Public Health Service","doi-asserted-by":"crossref","award":["AI105191"],"award-info":[{"award-number":["AI105191"]}],"id":[{"id":"10.13039\/100007197","id-type":"DOI","asserted-by":"crossref"}]},{"name":"Wellcome Trust","award":["087582"],"award-info":[{"award-number":["087582"]}]}],"content-domain":{"domain":["wellcomeopenresearch.org"],"crossmark-restriction":false},"short-container-title":["Wellcome Open Res"],"abstract":"<ns4:p>\n                    <ns4:italic>Background:<\/ns4:italic>\n                    Micronemal proteins of the thrombospondin-related anonymous protein (TRAP) family are believed to play essential roles during gliding motility and host cell invasion by apicomplexan parasites, and currently represent major vaccine candidates against\n                    <ns4:italic>Plasmodium falciparum<\/ns4:italic>\n                    , the causative agent of malaria. However, recent evidence suggests that they play multiple and different roles than previously assumed. Here, we analyse a null mutant for MIC2, the TRAP homolog in\n                    <ns4:italic>Toxoplasma gondii<\/ns4:italic>\n                    .\n                    <ns4:italic>Methods:<\/ns4:italic>\n                    We performed a careful analysis of parasite motility in a 3D-environment, attachment under shear stress conditions, host cell invasion and\n                    <ns4:italic>in vivo<\/ns4:italic>\n                    virulence.\n                    <ns4:italic>Results:<\/ns4:italic>\n                    We verified the role of MIC2 in efficient surface attachment, but were unable to identify any direct function of MIC2 in sustaining gliding motility or host cell invasion once initiated. Furthermore, we find that deletion of\n                    <ns4:italic>mic2<\/ns4:italic>\n                    causes a slightly delayed infection\n                    <ns4:italic>in vivo,<\/ns4:italic>\n                    leading only to mild attenuation of virulence; like with wildtype parasites, inoculation with even low numbers of\n                    <ns4:italic>mic2<\/ns4:italic>\n                    KO parasites causes lethal disease in mice. However, deletion of\n                    <ns4:italic>mic2<\/ns4:italic>\n                    causes delayed host cell egress\n                    <ns4:italic>in vitro<\/ns4:italic>\n                    , possibly via disrupted signal transduction pathways.\n                    <ns4:italic>Conclusions:<\/ns4:italic>\n                    We confirm a critical role of MIC2 in parasite attachment to the surface, leading to reduced parasite motility and host cell invasion. However, MIC2 appears to not be critical for gliding motility or host cell invasion, since parasite speed during these processes is unaffected. Furthermore, deletion of MIC2 leads only to slight attenuation of the parasite.\n                  <\/ns4:p>","DOI":"10.12688\/wellcomeopenres.11594.2","type":"journal-article","created":{"date-parts":[[2017,7,24]],"date-time":"2017-07-24T11:30:12Z","timestamp":1500895812000},"page":"32","update-policy":"https:\/\/doi.org\/10.12688\/wellcomeopenres.crossmark-policy","source":"Crossref","is-referenced-by-count":35,"title":["Parasites lacking the micronemal protein MIC2 are deficient in surface attachment and host cell egress, but remain virulent in vivo"],"prefix":"10.12688","volume":"2","author":[{"given":"Simon","family":"Gras","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Allison","family":"Jackson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stuart","family":"Woods","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gurman","family":"Pall","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jamie","family":"Whitelaw","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jacqueline M.","family":"Leung","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gary E.","family":"Ward","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Craig W.","family":"Roberts","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4816-5221","authenticated-orcid":false,"given":"Markus","family":"Meissner","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"2560","published-online":{"date-parts":[[2017,7,24]]},"reference":[{"key":"ref-1","doi-asserted-by":"publisher","first-page":"125-127","DOI":"10.1038\/nmeth.2301","article-title":"Conditional genome engineering in Toxoplasma gondii uncovers alternative invasion mechanisms.","volume":"10","author":"N Andenmatten","year":"2013","journal-title":"Nat Methods."},{"key":"ref-2","doi-asserted-by":"publisher","first-page":"e1004273","DOI":"10.1371\/journal.ppat.1004273","article-title":"Host cell invasion by apicomplexan parasites: the junction conundrum.","volume":"10","author":"D Bargieri","year":"2014","journal-title":"PLoS Pathog."},{"key":"ref-3","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms3552","article-title":"Apical membrane antigen 1 mediates apicomplexan parasite attachment but is dispensable for host cell invasion.","volume":"4","author":"D Bargieri","year":"2013","journal-title":"Nat Commun."},{"key":"ref-4","doi-asserted-by":"publisher","first-page":"618-630","DOI":"10.1016\/j.chom.2016.10.015","article-title":"Plasmodium Merozoite TRAP Family Protein Is Essential for Vacuole Membrane Disruption and Gamete Egress from Erythrocytes.","volume":"20","author":"D Bargieri","year":"2016","journal-title":"Cell Host Microbe."},{"key":"ref-5","doi-asserted-by":"publisher","first-page":"e1003031","DOI":"10.1371\/journal.ppat.1003031","article-title":"Semaphorin-7A is an erythrocyte receptor for P. falciparum merozoite-specific TRAP homolog, MTRAP.","volume":"8","author":"S Bartholdson","year":"2012","journal-title":"PLoS Pathog."},{"key":"ref-6","doi-asserted-by":"publisher","first-page":"5197-5208","DOI":"10.1074\/jbc.M509807200","article-title":"A conserved molecular motor drives cell invasion and gliding motility across malaria life cycle stages and other apicomplexan parasites.","volume":"281","author":"J Baum","year":"2006","journal-title":"J Biol Chem."},{"key":"ref-7","doi-asserted-by":"publisher","first-page":"9399-9408","DOI":"10.1128\/MCB.20.24.9399-9408.2000","article-title":"Ionophore-resistant mutants of Toxoplasma gondii reveal host cell permeabilization as an early event in egress.","volume":"20","author":"M Black","year":"2000","journal-title":"Mol Cell Biol."},{"key":"ref-9","doi-asserted-by":"publisher","first-page":"6221-6227","DOI":"10.1093\/emboj\/18.22.6221","article-title":"CTRP is essential for mosquito infection by malaria ookinetes.","volume":"18","author":"J Dessens","year":"1999","journal-title":"EMBO J."},{"key":"ref-36","doi-asserted-by":"publisher","first-page":"14010-14019","DOI":"10.1074\/jbc.271.24.14010","article-title":"Insertional tagging, cloning, and expression of the Toxoplasma gondii hypoxanthine-xanthine-guanine phosphoribosyltransferase gene. 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actomyosin system of Toxoplasma gondii is important for efficient gliding motility and invasion.","volume":"15","author":"J Whitelaw","year":"2017","journal-title":"BMC Biol."}],"container-title":["Wellcome Open Research"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/wellcomeopenresearch.org\/articles\/2-32\/v2\/xml","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/wellcomeopenresearch.org\/articles\/2-32\/v2\/pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/wellcomeopenresearch.org\/articles\/2-32\/v2\/iparadigms","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2020,5,17]],"date-time":"2020-05-17T23:10:01Z","timestamp":1589757001000},"score":1,"resource":{"primary":{"URL":"https:\/\/wellcomeopenresearch.org\/articles\/2-32\/v2"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2017,7,24]]},"references-count":35,"URL":"https:\/\/doi.org\/10.12688\/wellcomeopenres.11594.2","relation":{"has-review":[{"id-type":"doi","id":"10.21956\/wellcomeopenres.13129.r24449","asserted-by":"subject"},{"id-type":"doi","id":"10.21956\/wellcomeopenres.13129.r24448","asserted-by":"subject"},{"id-type":"doi","id":"10.21956\/wellcomeopenres.13129.r24449","asserted-by":"object"},{"id-type":"doi","id":"10.21956\/wellcomeopenres.13129.r24448","asserted-by":"object"}]},"ISSN":["2398-502X"],"issn-type":[{"value":"2398-502X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2017,7,24]]},"assertion":[{"value":"Indexed","URL":"https:\/\/wellcomeopenresearch.org\/articles\/2-32\/v2#article-reports","order":0,"name":"referee-status","label":"Referee status","group":{"name":"current-referee-status","label":"Current Referee Status"}},{"value":"10.21956\/wellcomeopenres.13129.r24449, Vernon Carruthers, My-Hang Huynh, Department of Microbiology and Immunology, University of Michigan School of Medicine, Ann Arbor, MI, 48109, USA, 02 Aug 2017, version 2, indexed","URL":"https:\/\/wellcomeopenresearch.org\/articles\/2-32\/v2#referee-response-24449","order":0,"name":"referee-response-24449","label":"Referee Report","group":{"name":"article-reports","label":"Article Reports"}},{"value":"10.21956\/wellcomeopenres.13129.r24448, Sebastian Lourido, Whitehead Institute for Biomedical Research, Cambridge, MA, USA, 02 Aug 2017, version 2, indexed","URL":"https:\/\/wellcomeopenresearch.org\/articles\/2-32\/v2#referee-response-24448","order":1,"name":"referee-response-24448","label":"Referee Report","group":{"name":"article-reports","label":"Article Reports"}},{"value":"10.21956\/wellcomeopenres.12524.r22938, Tim W. Gilberger, Paul-Chrisian Burda, Bernhard Nocht Institute for Tropical Medicine, Hamburg, Germany, 12 Jun 2017, version 1, indexed","URL":"https:\/\/wellcomeopenresearch.org\/articles\/2-32\/v1#referee-response-22938","order":2,"name":"referee-response-22938","label":"Referee Report","group":{"name":"article-reports","label":"Article Reports"}},{"value":"<b>Markus Meissner<\/b>; \n<i>Posted: 10 Jul 2017<\/i>; We thank the reviewers for their valuable comments, which helped us to improve our manuscript. We addressed all comments in our revision. See our response to the reviewers comments below (in bold)&nbsp;Figure 3B: Please show SD also for the 0.5 dyn\/cm\n<sup>2<\/sup> values\n<b>The value at 0.5 dyn\/cm2 value have been set for each experiment to 100% to be our starting point and allow us to visualise the variation under stronger flow. By doing so, no SD can be calculated for this point<\/b>Figure 4A+C, Figure 6D: Please include scale bar information in the figure legend.\n<b>Information now included for all 3 panels<\/b>Fig. 4: Did the authors used a Students T-test (as described in the figure legend) or a&nbsp;two-way-Anova with&nbsp;Sidak\u2019s multiple comparisons test&nbsp;as indicated in the Materials &amp; Methods?\n<b>A two-way-Anova with&nbsp;Sidak\u2019s multiple comparisons test&nbsp;as indicated in the Materials &amp; Methods was used. The is now corrected in the text.<\/b>Figure 5C: This figure is confusing and not well embedded in the result section. It would be helpful to include co-localization of AMA1 and RON2 in the \n<i>mic<\/i>2 knock out cell line. Additionally, they could provide the information of the YFP expression in WT \n<i>vs<\/i> KO.\n<b>RON2 was used as a marker for the tight junction and was used ofr both<i> WT <\/i>and <i>mic2 KO <\/i>parasites to demonstrate normal junction formation. Ama-1 is only shown for WT to outline the parasite. In case of <i>mic2KO <\/i>parasites YFP expression is activation upon induction of DiCre, requiring no additional stain<\/b>.Figure 5D: The &quot;ns&quot;&nbsp;on top of the graphs is hardly readable\n<b>We changed this in the revision.<\/b>Figure S3: The Y-Axis description is missing for the second and third data set\n<b>We changed this in the revision.<\/b>The supplementary movie file cannot be opened\n<b>We have checked the files and they could be opened on our system.<\/b>Additional remarks:Abstract \u2013 \u201cMIC2 appears to not be critical for gliding motility \u2026\u201d. This is correct, but in the light of Fig. 4A, showing a 60% reduction in gliding motility, a re-phrasing of this sentence might be good to avoid confusion.\n<b>The sentence now reads \u201cWe verified the role of MIC2 in efficient surface attachment, but were unable to identify any direct function of MIC2 in sustaining gliding motility or host cell invasion once initiated.\u201d<\/b>&nbsp;Page 5: The up-regulation of various invasion-related genes in both the \n<i>mic<\/i>2 KO and the AMA1 KO could be -as suggested- connected to multifactorial compensation, although this has to analyze in detail and compared to additional KO strains lacking other invasion-related proteins. An alternative explanation could be that the \u201cgeneral\u201d up-regulation is \u201cnoise\u201d due to the perturbation of the functional protein network.\n<b>We agree with the reviews that a more thorough analysis of transcriptional changes in different null mutants for components of the invasion machinery is worthwhile for future investigations.<\/b>","URL":"https:\/\/wellcomeopenresearch.org\/articles\/2-32\/v2#referee-comment-2865","order":3,"name":"referee-comment-2865","label":"Referee Comment","group":{"name":"article-reports","label":"Article Reports"}},{"value":"This work was supported the Wellcome Trust [087582], [103875] and [085349]; an European Research Council-Starting grant [ERC-2012-StG 309255-EndoTox] to MM; US Public Health Service [AI105191] and [AI054961] to GW. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.","order":4,"name":"grant-information","label":"Grant Information"},{"value":"This is an open access article distributed under the terms of the Creative Commons Attribution Licence, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.","order":0,"name":"copyright-info","label":"Copyright"}]}}