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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Contribution of Murine Models to the Study of Malaria During Pregnancy</title><author><name sortKey="Barateiro, Andre" sort="Barateiro, Andre" uniqKey="Barateiro A" first="André" last="Barateiro">André Barateiro</name><affiliation><nlm:aff id="aff1"><institution>Department of Parasitology, Institute of Biomedical Sciences, University of São Paulo</institution>,<addr-line>São Paulo</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Pereira, Marcelo L M" sort="Pereira, Marcelo L M" uniqKey="Pereira M" first="Marcelo L. M." last="Pereira">Marcelo L. M. Pereira</name><affiliation><nlm:aff id="aff1"><institution>Department of Parasitology, Institute of Biomedical Sciences, University of São Paulo</institution>,<addr-line>São Paulo</addr-line>,<country>Brazil</country></nlm:aff></affiliation><affiliation><nlm:aff id="aff2"><institution>Institute of Biosystems and Integrative Sciences, Faculty of Sciences, University of Lisbon</institution>,<addr-line>Lisbon</addr-line>,<country>Portugal</country></nlm:aff></affiliation></author><author><name sortKey="Epiphanio, Sabrina" sort="Epiphanio, Sabrina" uniqKey="Epiphanio S" first="Sabrina" last="Epiphanio">Sabrina Epiphanio</name><affiliation><nlm:aff id="aff3"><institution>Department of Clinical Analysis and Toxicology, Faculty of Pharmaceutical Sciences, University of São Paulo</institution>,<addr-line>São Paulo</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Marinho, Claudio R F" sort="Marinho, Claudio R F" uniqKey="Marinho C" first="Claudio R. F." last="Marinho">Claudio R. F. Marinho</name><affiliation><nlm:aff id="aff1"><institution>Department of Parasitology, Institute of Biomedical Sciences, University of São Paulo</institution>,<addr-line>São Paulo</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">31275284</idno><idno type="pmc">6594417</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6594417</idno><idno type="RBID">PMC:6594417</idno><idno type="doi">10.3389/fmicb.2019.01369</idno><date when="2019">2019</date><idno type="wicri:Area/Pmc/Corpus">000976</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000976</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Contribution of Murine Models to the Study of Malaria During Pregnancy</title><author><name sortKey="Barateiro, Andre" sort="Barateiro, Andre" uniqKey="Barateiro A" first="André" last="Barateiro">André Barateiro</name><affiliation><nlm:aff id="aff1"><institution>Department of Parasitology, Institute of Biomedical Sciences, University of São Paulo</institution>,<addr-line>São Paulo</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Pereira, Marcelo L M" sort="Pereira, Marcelo L M" uniqKey="Pereira M" first="Marcelo L. M." last="Pereira">Marcelo L. M. Pereira</name><affiliation><nlm:aff id="aff1"><institution>Department of Parasitology, Institute of Biomedical Sciences, University of São Paulo</institution>,<addr-line>São Paulo</addr-line>,<country>Brazil</country></nlm:aff></affiliation><affiliation><nlm:aff id="aff2"><institution>Institute of Biosystems and Integrative Sciences, Faculty of Sciences, University of Lisbon</institution>,<addr-line>Lisbon</addr-line>,<country>Portugal</country></nlm:aff></affiliation></author><author><name sortKey="Epiphanio, Sabrina" sort="Epiphanio, Sabrina" uniqKey="Epiphanio S" first="Sabrina" last="Epiphanio">Sabrina Epiphanio</name><affiliation><nlm:aff id="aff3"><institution>Department of Clinical Analysis and Toxicology, Faculty of Pharmaceutical Sciences, University of São Paulo</institution>,<addr-line>São Paulo</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Marinho, Claudio R F" sort="Marinho, Claudio R F" uniqKey="Marinho C" first="Claudio R. F." last="Marinho">Claudio R. F. Marinho</name><affiliation><nlm:aff id="aff1"><institution>Department of Parasitology, Institute of Biomedical Sciences, University of São Paulo</institution>,<addr-line>São Paulo</addr-line>,<country>Brazil</country></nlm:aff></affiliation></author></analytic><series><title level="j">Frontiers in Microbiology</title><idno type="eISSN">1664-302X</idno><imprint><date when="2019">2019</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Annually, many pregnancies occur in areas of <italic>Plasmodium</italic> spp. transmission, particularly in underdeveloped countries with widespread poverty. Estimations have suggested that several million women are at risk of developing malaria during pregnancy. In particular cases, systemic infection caused by <italic>Plasmodium</italic> spp. may extend to the placenta, dysregulating local homeostasis and promoting the onset of placental malaria; these processes are often associated with increased maternal and fetal mortality, intrauterine growth restriction, preterm delivery, and reduced birth weight. The endeavor to understand and characterize the mechanisms underlying disease onset and placental pathology face several ethical and logistical obstacles due to explicit difficulties in assessing human gestation and biological material. Consequently, the advent of murine experimental models for the study of malaria during pregnancy has substantially contributed to our understanding of this complex pathology. Herein, we summarize research conducted during recent decades using murine models of malaria during pregnancy and highlight the most relevant findings, as well as discuss similarities to humans and the translational capacity of achieved results.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Abrams, E T" uniqKey="Abrams E">E. T. Abrams</name></author><author><name sortKey="Brown, H" uniqKey="Brown H">H. Brown</name></author><author><name sortKey="Chensue, S W" uniqKey="Chensue S">S. W. Chensue</name></author><author><name sortKey="Turner, G D H" uniqKey="Turner G">G. D. H. Turner</name></author><author><name sortKey="Molyneux, M E" uniqKey="Molyneux M">M. E. Molyneux</name></author><author><name sortKey="Meshnick, S R" uniqKey="Meshnick S">S. R. Meshnick</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Adachi, M" uniqKey="Adachi M">M. 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<front><journal-meta><journal-id journal-id-type="nlm-ta">Front Microbiol</journal-id><journal-id journal-id-type="iso-abbrev">Front Microbiol</journal-id><journal-id journal-id-type="publisher-id">Front. Microbiol.</journal-id><journal-title-group><journal-title>Frontiers in Microbiology</journal-title></journal-title-group><issn pub-type="epub">1664-302X</issn><publisher><publisher-name>Frontiers Media S.A.</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">31275284</article-id><article-id pub-id-type="pmc">6594417</article-id><article-id pub-id-type="doi">10.3389/fmicb.2019.01369</article-id><article-categories><subj-group subj-group-type="heading"><subject>Microbiology</subject><subj-group><subject>Review</subject></subj-group></subj-group></article-categories><title-group><article-title>Contribution of Murine Models to the Study of Malaria During Pregnancy</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Barateiro</surname><given-names>André</given-names></name><xref ref-type="aff" rid="aff1"><sup>1</sup></xref><uri xlink:type="simple" xlink:href="http://loop.frontiersin.org/people/615315/overview"></uri></contrib><contrib contrib-type="author"><name><surname>Pereira</surname><given-names>Marcelo L. M.</given-names></name><xref ref-type="aff" rid="aff1"><sup>1</sup></xref><xref ref-type="aff" rid="aff2"><sup>2</sup></xref><uri xlink:type="simple" xlink:href="http://loop.frontiersin.org/people/492373/overview"></uri></contrib><contrib contrib-type="author"><name><surname>Epiphanio</surname><given-names>Sabrina</given-names></name><xref ref-type="aff" rid="aff3"><sup>3</sup></xref><uri xlink:type="simple" xlink:href="http://loop.frontiersin.org/people/113848/overview"></uri></contrib><contrib contrib-type="author"><name><surname>Marinho</surname><given-names>Claudio R. F.</given-names></name><xref ref-type="aff" rid="aff1"><sup>1</sup></xref><xref ref-type="corresp" rid="c001"><sup>*</sup></xref><uri xlink:type="simple" xlink:href="http://loop.frontiersin.org/people/615145/overview"></uri></contrib></contrib-group><aff id="aff1"><sup>1</sup><institution>Department of Parasitology, Institute of Biomedical Sciences, University of São Paulo</institution>,<addr-line>São Paulo</addr-line>,<country>Brazil</country></aff><aff id="aff2"><sup>2</sup><institution>Institute of Biosystems and Integrative Sciences, Faculty of Sciences, University of Lisbon</institution>,<addr-line>Lisbon</addr-line>,<country>Portugal</country></aff><aff id="aff3"><sup>3</sup><institution>Department of Clinical Analysis and Toxicology, Faculty of Pharmaceutical Sciences, University of São Paulo</institution>,<addr-line>São Paulo</addr-line>,<country>Brazil</country></aff><author-notes><fn fn-type="edited-by"><p>Edited by: Demba Sarr, University of Georgia, United States</p></fn><fn fn-type="edited-by"><p>Reviewed by: Justin Yai Alamou Doritchamou, National Institutes of Health (NIH), United States; Adrian John Frederick Luty, Institut de Recherche pour le Développement (IRD), France; Julie M. Moore, University of Florida, United States</p></fn><corresp id="c001">*Correspondence: Claudio R. F. Marinho, <email>marinho@usp.br</email></corresp><fn fn-type="other" id="fn004"><p>This article was submitted to Infectious Diseases, a section of the journal Frontiers in Microbiology</p></fn></author-notes><pub-date pub-type="epub"><day>19</day><month>6</month><year>2019</year></pub-date><pub-date pub-type="collection"><year>2019</year></pub-date><volume>10</volume><elocation-id>1369</elocation-id><history><date date-type="received"><day>02</day><month>4</month><year>2019</year></date><date date-type="accepted"><day>31</day><month>5</month><year>2019</year></date></history><permissions><copyright-statement>Copyright © 2019 Barateiro, Pereira, Epiphanio and Marinho.</copyright-statement><copyright-year>2019</copyright-year><copyright-holder>Barateiro, Pereira, Epiphanio and Marinho</copyright-holder><license xlink:href="http://creativecommons.org/licenses/by/4.0/"><license-p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</license-p></license></permissions><abstract><p>Annually, many pregnancies occur in areas of <italic>Plasmodium</italic> spp. transmission, particularly in underdeveloped countries with widespread poverty. Estimations have suggested that several million women are at risk of developing malaria during pregnancy. In particular cases, systemic infection caused by <italic>Plasmodium</italic> spp. may extend to the placenta, dysregulating local homeostasis and promoting the onset of placental malaria; these processes are often associated with increased maternal and fetal mortality, intrauterine growth restriction, preterm delivery, and reduced birth weight. The endeavor to understand and characterize the mechanisms underlying disease onset and placental pathology face several ethical and logistical obstacles due to explicit difficulties in assessing human gestation and biological material. Consequently, the advent of murine experimental models for the study of malaria during pregnancy has substantially contributed to our understanding of this complex pathology. Herein, we summarize research conducted during recent decades using murine models of malaria during pregnancy and highlight the most relevant findings, as well as discuss similarities to humans and the translational capacity of achieved results.</p></abstract><kwd-group><kwd>murine models</kwd><kwd><italic>Plasmodium</italic> spp.</kwd><kwd>malaria</kwd><kwd>pregnancy</kwd><kwd>placental malaria</kwd></kwd-group><funding-group><award-group><funding-source id="cn001">Fundação de Amparo à Pesquisa do Estado de São Paulo<named-content content-type="fundref-id">10.13039/501100001807</named-content></funding-source><award-id rid="cn001">2018/20468-0</award-id><award-id rid="cn001">2017/05782-8</award-id><award-id rid="cn001">2017/03939-7</award-id><award-id rid="cn001">2012/10081-5</award-id></award-group></funding-group><counts><fig-count count="3"></fig-count><table-count count="2"></table-count><equation-count count="0"></equation-count><ref-count count="104"></ref-count><page-count count="18"></page-count><word-count count="0"></word-count></counts></article-meta></front><body><sec id="S1"><title>Malaria in Human Pregnancy</title><p>Malaria still represents a serious public health issue for several communities distributed worldwide. Estimations from 2017 have shown that at least 219 million cases occurred in areas of <italic>Plasmodium</italic> spp. transmission, in which approximately half a million victims died from the disease (<xref rid="B104" ref-type="bibr">World Health Organization, 2018</xref>). These estimations encompass pregnant women, who are particularly more susceptible to developing severe clinical manifestations resulting from malaria in pregnancy (MiP) induced by <italic>Plasmodium</italic> spp. Although outdated, estimations performed by <xref rid="B20" ref-type="bibr">Dellicour et al. (2010)</xref> noted 125.2 million pregnancies occurring in malaria-endemic areas. This study came as a follow-up of previous estimations suggesting that 25 million pregnancies occur in areas of <italic>P. falciparum</italic> transmission alone, distributed across sub-Saharan Africa (<xref rid="B21" ref-type="bibr">Desai et al., 2007</xref>). Nevertheless, current epidemiologic knowledge is inaccurate and imprecise and might support outdated and underestimated predictions, hiding a much more alarming reality.</p><p>This concerning epidemiological scenario highlights the importance of conducting preventive measures to control MiP, which might drastically evolve to placental malaria (PM), a pathology frequently associated with the occurrence of poor outcomes during pregnancy, such as maternal and fetal mortality, intrauterine growth restriction (IUGR), preterm birth and reduced birth weight (<xref rid="B21" ref-type="bibr">Desai et al., 2007</xref>; <xref rid="B95" ref-type="bibr">Umbers et al., 2011</xref>; <xref rid="B77" ref-type="bibr">Rogerson et al., 2018</xref>). These deleterious consequences affecting both the mother and the growing fetus are reflections of histological and physiological changes occurring within the placenta [recently summarized elsewhere (<xref rid="B88" ref-type="bibr">Sharma and Shukla, 2017</xref>)]. One of the key features of <italic>P. falciparum</italic> PM is the accumulation of <italic>P. falciparum</italic>-infected erythrocytes (IE) in the placenta (<xref rid="B8" ref-type="bibr">Beeson et al., 2002</xref>; <xref rid="B57" ref-type="bibr">Muthusamy et al., 2004</xref>). Upregulation of the VAR2CSA protein, a variant of the highly polymorphic adhesion peptide <italic>P. falciparum</italic> erythrocyte membrane protein 1 (PfEMP1) (<xref rid="B82" ref-type="bibr">Salanti et al., 2003</xref>), is responsible for parasite sequestration upon preferential binding to chondroitin sulfate A (CSA), which is abundantly expressed in the placenta (<xref rid="B26" ref-type="bibr">Fried et al., 2006</xref>; <xref rid="B58" ref-type="bibr">Muthusamy et al., 2007</xref>). Consequently, a severe local inflammatory process is triggered, characterized by the infiltration of monocytes and leukocytes in the placenta and eventual inflammation in response to parasite accumulation (<xref rid="B34" ref-type="bibr">Ismail et al., 2000</xref>; <xref rid="B67" ref-type="bibr">Parekh et al., 2010</xref>; <xref rid="B44" ref-type="bibr">Lucchi et al., 2011</xref>; <xref rid="B91" ref-type="bibr">Souza et al., 2013</xref>). This process has been frequently associated with placental histological alterations during MiP, such as dysregulation of placental architecture, formation of syncytial knots, fibrin deposition, necrosis, and placental barrier thickening (<xref rid="B102" ref-type="bibr">Walter et al., 1982</xref>; <xref rid="B34" ref-type="bibr">Ismail et al., 2000</xref>; <xref rid="B91" ref-type="bibr">Souza et al., 2013</xref>), and has been reported to occur during IUGR and preterm delivery, as well as in cases of reduced birth weight (<xref rid="B55" ref-type="bibr">Moormann et al., 1999</xref>; <xref rid="B52" ref-type="bibr">Menendez et al., 2000</xref>; <xref rid="B79" ref-type="bibr">Rogerson et al., 2003</xref>; <xref rid="B95" ref-type="bibr">Umbers et al., 2011</xref>).</p><p>Although we have obtained substantial knowledge in the field, studies on the epidemiology and pathology of MiP are frequently challenging due to related ethical and logistic difficulties. Long gestational periods, uncontrollable experimental planning, difficult access to biological and placental samples, and critical constraints associated with human experiments represent significant barriers that slow research progress and the understanding of this severe and complex disease. Therefore, alternatives have emerged with the advent of murine models to study MiP that have brought fundamental knowledge to the field. Herein, we have compiled research conducted for the past four decades using experimental rodent models, highlighting the most relevant findings, similarities to humans and, consequently, the translational capacity of achieved results.</p></sec><sec id="S2"><title>Human and Murine Gestation: Comparison and Translational Challenges for the Study of MiP</title><sec id="S2.SS1"><title>Comparative Gestation Development</title><p>In parallel with some other features, gestation length and development are somewhat different between human and murine mammals. Human gestation lasts for approximately 38 weeks (three trimesters) in contrast to rodents, in which gestation takes place over a 3-week period (<xref rid="B56" ref-type="bibr">Murray et al., 2010</xref>). In addition, there are also slight differences between species regarding the implantation period. Murine implantation takes place between the fourth and fifth days of gestation, somewhat sooner than in humans, in which this phenomenon occurs between the fifth and sixth days (<xref rid="B81" ref-type="bibr">Rossant and Tam, 2017</xref>). After implantation, gestation will progress toward a shift in fetal nutritional means. During early gestation in both humans and rodents, the growing fetus will survive through means of histiotrophic nutrition, in which nutrients are acquired by the uptake of substances secreted from the uterine glands present in the endometrium (<xref rid="B15" ref-type="bibr">Burton et al., 2002</xref>; <xref rid="B27" ref-type="bibr">Georgiades et al., 2002</xref>). However, the nutritional strategy changes around mid-gestation when the maternal blood supply to the placenta is completely established. Accordingly, the nutrient and gas exchanges between the mother and the fetus become functional, marking the onset of placental hemotrophic nutrition (<xref rid="B14" ref-type="bibr">Burton et al., 2001</xref>; <xref rid="B27" ref-type="bibr">Georgiades et al., 2002</xref>). In humans, this phenomenon occurs between the end of the first and the beginning of the second trimester, while in mice and rats, the same scenario occurs specifically at mid-gestation (between the twelfth and thirteenth days) (<xref rid="B27" ref-type="bibr">Georgiades et al., 2002</xref>). This shift in nutritional strategy occurs with the onset of organogenesis and fetal development, after which growth will continue until gestational term is reached.</p></sec><sec id="S2.SS2"><title>Comparative Placental Function, Structure, and Histology</title><p>Human and murine placentas have a considerable degree of similarity and are nearly identical from physiological and functional perspectives (<xref rid="B80" ref-type="bibr">Rossant and Cross, 2001</xref>; <xref rid="B27" ref-type="bibr">Georgiades et al., 2002</xref>). In both mammalian species, this transient organ ensures nutrient and gas exchange between the mother and the growing fetus (<xref rid="B39" ref-type="bibr">Lager and Powell, 2012</xref>), maintains tolerance to the maternal immune system (<xref rid="B37" ref-type="bibr">Kanellopoulos-Langevin et al., 2003</xref>), and works as a physical and immunological barrier against endogenous pathogens (<xref rid="B74" ref-type="bibr">Robbins and Bakardjiev, 2012</xref>).</p><p>The placentas in both species are discoid organs in which maternal blood coming from the endometrium circulates and is in direct contact with trophoblasts, the fetal-derived cells responsible for regulating most placental physiological functions. In both species, the placenta can be structurally divided into three main areas: (1) an outer layer, consisting of uterine decidual cells and maternal blood vessels; (2) a middle layer, considered the implantation site where the placenta attaches to the uterus; and (3) an inner layer, in which the maternal blood interacts with trophoblasts to promote vital metabolic exchanges [reviewed elsewhere (<xref rid="B27" ref-type="bibr">Georgiades et al., 2002</xref>)]. Accordingly, the outer area can be mainly described as being composed by the myometrium and decidua basalis, which are extensively infiltrated by maternal arteries. This area is partially invaded by fetal-derived trophoblasts during the implantation process, which is considerably similar between both mammalian species. Moving toward the inner area of the human placenta, it is possible to distinguish a structural layer with no counterpart in the murine placenta known as the basal plate. This implantation site contains several distinct trophoblast subpopulations, such as extravillous (EVT) and cytotrophoblast (CT) cells. Nevertheless, an analogous area defined as the junctional zone occurs in murine animals, which is populated by specific types of cells such as trophoblast giant cells (TGC) and spongiotrophoblasts (SPG). Inside the placenta, more pronounced differences are observed regarding morphology and structure. The central area, which is commonly referred to as the human fetal placenta, is constituted by villi in an extremely ramified tree-like structure, which increases the contact surface with the maternal blood freely circulating within the intervillous space (IVS) (<xref ref-type="fig" rid="F1">Figure 1A</xref>). On the other hand, a similar structure known as the labyrinth develops in mice and rats (<xref ref-type="fig" rid="F2">Figures 2A,C</xref>). This impacts maternal blood circulation inside the placenta, where blood stays confined to the tortuous and sinusoidal channels (<xref rid="B80" ref-type="bibr">Rossant and Cross, 2001</xref>). Additionally, the surface area contacting the maternal blood is somewhat distinguished between humans and rodents from both cytological and structural perspectives. In the human placenta, maternal blood is separated from fetal capillaries by a single layer of syncytiotrophoblasts (ST), beyond which CT cells are present together with the basal lamina and fetal endothelium (schematically represented in <xref ref-type="fig" rid="F3">Figure 3</xref>). According to these characteristics, the human placenta is classified as hemomonochorial (<xref rid="B93" ref-type="bibr">Takata et al., 1997</xref>; <xref rid="B27" ref-type="bibr">Georgiades et al., 2002</xref>). In opposition, three layers compose the murine labyrinthine wall: one composed of mononuclear trophoblasts dispersed throughout the surface contacting maternal blood and two ST layers definitively separating the maternal and fetal compartments (illustrated in <xref ref-type="fig" rid="F3">Figure 3</xref>). Accordingly, murine placentas are classified as hemotrichorial (<xref rid="B93" ref-type="bibr">Takata et al., 1997</xref>).</p><fig id="F1" position="float"><label>FIGURE 1</label><caption><p>Histologic events associated with placental pathology during human malaria in pregnancy. Placental histologic sections of non-infected <bold>(A)</bold> and <italic>P. falciparum</italic>-infected <bold>(B–F)</bold> women stained with Hematoxylin-Eosin. <bold>(A)</bold> Healthy human placenta without tissue damage and normal architecture in which intervillous spaces (IVS), usually bathed in maternal blood are in contact with placental villi (V), feeding the growing fetus through the fetal vessels (FV). <bold>(B)</bold><italic>P. falciparum</italic>-infected placentas often characterized by extensive necrotic areas (Ne) and tissue disorganization. <bold>(C,D)</bold> Mononuclear cells infiltrate (arrow) is frequently patent due to the accumulation of parasitized erythrocytes (arrowhead). <bold>(E)</bold> Massive sequestration of <italic>P. falciparum</italic>-infected erythrocytes in the placenta is accompanied by <bold>(F)</bold> the accumulation of hemozoin in the IVS, which can be detected by polarized light microscopy. Scale bars represent <bold>(A,B,E,F)</bold> 100 μm, <bold>(C)</bold> 5 μm, and <bold>(D)</bold> 15 μm. Microscopy pictures were taken by Rodrigo Medeiros de Souza.</p></caption><graphic xlink:href="fmicb-10-01369-g001"></graphic></fig><fig id="F2" position="float"><label>FIGURE 2</label><caption><p>Histologic events associated with placental pathology during murine malaria in pregnancy. Placental histology of non-infected <bold>(A,C)</bold> and <italic>P. berghei</italic>-infected <bold>(B,C,E,F)</bold> mice stained with Hematoxylin-Eosin. <bold>(A)</bold> Healthy placenta with normal histologic structure characterized by normal distribution of layer-specific cell such decidual cells (DC), trophoblastic cells (Cy), and labyrinthic cells (La), having <bold>(C)</bold> a normal labyrinth organization. <bold>(B)</bold> Fibrinoid necrosis (Ne) is depicted in section of <italic>P. berghei</italic>-infected placentas with <bold>(D)</bold> massive tissue disorganization and trophoblast membrane thickening. <bold>(E)</bold> Mononuclear cell infiltrate is visible (arrow), which occurs in response to <bold>(D)</bold> parasite accumulation (arrow) and <bold>(F)</bold> hemozoin deposition observed using polarized light microscopy. Scale bars represent <bold>(A,B)</bold> 100 μm, <bold>(C–E)</bold> 15 μm, and <bold>(F)</bold> 30 μm. Figure was modified from <xref rid="B59" ref-type="bibr">Neres et al. (2008)</xref> and <xref rid="B47" ref-type="bibr">Marinho et al. (2009)</xref> upon permission granted on behalf of journal’s editorial board.</p></caption><graphic xlink:href="fmicb-10-01369-g002"></graphic></fig><fig id="F3" position="float"><label>FIGURE 3</label><caption><p>Schematic representation of possible histologic events in human and murine placentas during malaria in pregnancy. Human and murine fetuses are represented with corresponding placentas. Magnification of transversal slices of each placental core illustrate histologic architecture, which widely differs between human placental villi and murine labyrinthine structure. During malaria in pregnancy, both human and murine placentas experience histopathologic manifestations that impair local homeostasis and lead to poor pregnancy outcomes. A comparison between both infected placentas and their non-infected counterparts is depicted. The presence of monocyte infiltrate is visible, which occurs in response to parasite and hemozoin accumulation. Histologic modifications such as fibrin deposition and syncytial knots occur due to the onset of severe local inflammation. Legend of picture components is vertically displayed beside the scheme.</p></caption><graphic xlink:href="fmicb-10-01369-g003"></graphic></fig><p>Nevertheless, the structural divergences of human and murine placentas have occurred in parallel with an extensive variety of resident cell populations that are independent of their analogs but not similar in development and characteristics and are equally responsible for the same physiological and functional processes in human, mouse and rat placentas (<xref ref-type="fig" rid="F3">Figure 3</xref>).</p></sec><sec id="S2.SS3"><title>Using Murine Models to Study Malaria in Human Pregnancy</title><p>It is required that a suitable experimental model includes a group of features that lead to valid and translatable findings. As such, it is pivotal that rodent physiology and genetics, as well as pathologic manifestations during disease, resemble those in humans to a certain extent, ultimately validating them as models for human research (<xref rid="B36" ref-type="bibr">Justice and Dhillon, 2016</xref>). Accordingly, after several models were reviewed, mice and rats were validated by Desowitz as suitable to study MiP (<xref rid="B24" ref-type="bibr">Duffy and Fried, 2001</xref>) under the assumption that physiological and pathological similarities observed between rodents and humans were sufficient to consider them appropriate to study this disease. Shared placental characteristics, such as hemochorial and discoid structure (<xref rid="B27" ref-type="bibr">Georgiades et al., 2002</xref>; <xref rid="B103" ref-type="bibr">Wildman et al., 2006</xref>), hemotrophic nutrition (<xref rid="B14" ref-type="bibr">Burton et al., 2001</xref>; <xref rid="B27" ref-type="bibr">Georgiades et al., 2002</xref>) and analogous placental cell populations (<xref rid="B80" ref-type="bibr">Rossant and Cross, 2001</xref>; <xref rid="B27" ref-type="bibr">Georgiades et al., 2002</xref>), encourage their usage as models of MiP. These similarities are supported by molecular phylogenetic analysis, which clusters both rodents and humans into the same evolutionary clade (<xref rid="B103" ref-type="bibr">Wildman et al., 2006</xref>). The molecular similarities can be extended to immunity, an important aspect to consider when studying a disease in which poor outcomes are predominantly dependent on the host’s immune response to malaria (<xref rid="B55" ref-type="bibr">Moormann et al., 1999</xref>; <xref rid="B79" ref-type="bibr">Rogerson et al., 2003</xref>). Hence, despite controversial discussion, it was shown that patterns of gene expression and biological pathways altered in mouse models of inflammatory diseases were significantly correlated with those from corresponding human conditions (e.g., sepsis) (<xref rid="B92" ref-type="bibr">Takao and Miyakawa, 2015</xref>), further supporting the usage of these animals for studying inflammatory diseases such as MiP. Together with the features mentioned above, the short gestational period, capacity for frequent and successive pregnancies, as well as large litter sizes, allow the rapid gathering of a considerable number of biological samples, which makes the rodents a powerful model for studying any pregnancy-related disease.</p><p>In addition to host characteristics, parasite intrinsic features are essential to validate experimental models of MiP. Above all, the most relevant features to consider should be the similarities between human and murine parasites cytoadhesion mechanisms. It is well known that the pathogenesis mechanisms involved in severe malaria induced by <italic>P. falciparum</italic> are associated with tissue- and organ-specific sequestration of IE (<xref rid="B85" ref-type="bibr">Schofield and Grau, 2005</xref>). More specifically, <italic>P. falciparum</italic> MiP may lead to severe PM as a result of IE sequestration in the placenta, which is mediated by the PfEMP1 variant VAR2CSA (which binds to placental CSA). The wide range of PfEMP1 variants known to be encoded by 50–60 <italic>var</italic> genes (<xref rid="B38" ref-type="bibr">Kraemer and Smith, 2006</xref>) have no known homologs in murine parasites (<xref rid="B30" ref-type="bibr">Hall et al., 2005</xref>). In fact, only some genetic signatures are similar between the murine and human <italic>Plasmodium</italic> species that encode alternate variable surface antigens (VSA), which were grouped in the multigenic <italic>pir</italic> superfamily (<italic>Plasmodium</italic> interspersed repeats) (<xref rid="B35" ref-type="bibr">Janssen et al., 2004</xref>; <xref rid="B30" ref-type="bibr">Hall et al., 2005</xref>). Nevertheless, murine parasite strains such as <italic>P. berghei</italic> have been shown to accumulate in specific tissues such as brain, fat, lung and spleen in a CD36-dependent and independent manner, which is also a well conserved pathogenesis mechanisms of severe malaria induced by <italic>P. falciparum</italic> (<xref rid="B25" ref-type="bibr">Franke-Fayard et al., 2010</xref>). In the same extent, the expression of alternate VSA and the binding capacity to placental CSA observed in murine parasites support the conclusions regarding the similarities between the pathogenesis mechanisms involved in murine and human disease (<xref rid="B30" ref-type="bibr">Hall et al., 2005</xref>; <xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>; <xref rid="B47" ref-type="bibr">Marinho et al., 2009</xref>; <xref rid="B33" ref-type="bibr">Hviid et al., 2010</xref>). Distinct mechanisms of pathogenesis might also be directly linked with the biology of the parasites. As such, it is advisable to consider that the usage of murine parasites needs to be adjusted with caution since specific strains might better mimic the distinct diseases associated with the unrelated biology of human parasites (e. g. <italic>P. falciparum</italic> or <italic>P. vivax</italic>). Accordingly, disease severity in mice tends to be higher in <italic>P. berghei</italic> ANKA causing lethal anemia and cerebral malaria in C57BL/6 for instance (resembling <italic>P. falciparum</italic> in humans); however, it has a tropism for reticulocytes (similar to <italic>P. vivax</italic>). Differently, <italic>P. chabaudi</italic> invades mature erythrocytes and is responsible for a less severe pathology shown by the resistance to infection observed in BALB/c and C57BL/6 mice. In the same extent, different parasites might also have different tendencies to accumulation in specific tissues, being better used to study specific diseases and host-pathogen interactions (reviewed in <xref rid="B40" ref-type="bibr">Lamb et al., 2006</xref>).</p><p>Despite the aforementioned host and parasite characteristics, one must carefully translate findings obtained in murine models to humans, especially regarding preclinical observations, due to the observed differences between both mammals. Nevertheless, the following sections will depict some of the current knowledge acquired using murine models of MiP (research details for each model are presented in <xref rid="T1" ref-type="table">Table 1</xref>) during the past decades while describing the parallel characteristics existing between humans and rodents in regard to poor pregnancy outcomes and MiP placental pathology (summarized list in <xref rid="T2" ref-type="table">Table 2</xref>).</p><table-wrap id="T1" position="float"><label>TABLE 1</label><caption><p>Summary of the principal findings and observations done using murine models of malaria in pregnancy.</p></caption><table frame="hsides" rules="groups" cellspacing="5" cellpadding="5"><thead><tr><td valign="top" align="left" rowspan="1" colspan="1"><bold>Author(s)</bold></td><td valign="top" align="left" rowspan="1" colspan="1"><bold><italic>Plasmodium</italic> strain</bold></td><td valign="top" align="left" rowspan="1" colspan="1"><bold>Murine strain</bold></td><td valign="top" align="left" rowspan="1" colspan="1"><bold>Recrudescence</bold></td><td valign="top" align="left" rowspan="1" colspan="1"><bold>Anemia</bold></td><td valign="top" align="left" rowspan="1" colspan="1"><bold>Maternal mortality</bold></td><td valign="top" align="left" rowspan="1" colspan="1"><bold>Abortion Resorption Stillbirth Preterm delivery</bold></td><td valign="top" align="left" rowspan="1" colspan="1"><bold>Litter size Newborn/fetal weight/health</bold></td><td valign="top" align="left" rowspan="1" colspan="1"><bold>Placental abnormalities/alterations</bold></td></tr></thead><tbody><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B97" ref-type="bibr">van Zon and Eling,1980</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic> K173</td><td valign="top" align="left" rowspan="1" colspan="1">Swiss Albino</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Primigravidae</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ In recrudescence</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">C3H/StZ</td><td valign="top" align="left" rowspan="1" colspan="1">↓multigravidae</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↑ primigravidae</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">BALB/c</td><td valign="top" align="left" rowspan="1" colspan="1">↑ due to</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">B10LP</td><td valign="top" align="left" rowspan="1" colspan="1">↓immunity</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B99" ref-type="bibr">van Zon et al.,1982</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic> K173</td><td valign="top" align="left" rowspan="1" colspan="1">Swiss Albino</td><td valign="top" align="left" rowspan="1" colspan="1">→ To pregnancy</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ In recrudescence</td><td valign="top" align="left" rowspan="1" colspan="1">Preterm delivery ↑ in recrudescence</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B61" ref-type="bibr">Oduola et al.,1982</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">A/J</td><td valign="top" align="left" rowspan="1" colspan="1">→ To pregnancy</td><td valign="top" align="left" rowspan="1" colspan="1">↑ In pregnancy</td><td valign="top" align="left" rowspan="1" colspan="1">↑ In early infection</td><td valign="top" align="left" rowspan="1" colspan="1">Stillbirth → to early infection</td><td valign="top" align="left" rowspan="1" colspan="1">Normal litter size</td><td valign="top" align="left" rowspan="1" colspan="1">Impaired labyrinth structure</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">NK65</td><td valign="top" align="left" rowspan="1" colspan="1">ICR</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ in late infection</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ Weight → to late infection</td><td valign="top" align="left" rowspan="1" colspan="1">Trophoblast barrier thickening</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↑ parturition in late infection</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↑ spleen weight</td><td valign="top" align="left" rowspan="1" colspan="1">IEs/Hz</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">No congenital malaria</td><td valign="top" align="left" rowspan="1" colspan="1">MΦ</td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B98" ref-type="bibr">van Zon et al.,1985</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">Swiss Albino</td><td valign="top" align="left" rowspan="1" colspan="1">→ To pregnancy</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">K173</td><td valign="top" align="left" rowspan="1" colspan="1">B10LP</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B101" ref-type="bibr">Vinayak et al.,1986</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic> NICD</td><td valign="top" align="left" rowspan="1" colspan="1">Swiss Albino</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ In early infection</td><td valign="top" align="left" rowspan="1" colspan="1">Stillbirth → to mid gestation infection</td><td valign="top" align="left" rowspan="1" colspan="1">↓ Litter size due to resorption</td><td valign="top" align="left" rowspan="1" colspan="1">Hyperplasia</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">Death before parturition</td><td valign="top" align="left" rowspan="1" colspan="1">↑ resorption</td><td valign="top" align="left" rowspan="1" colspan="1">↓ weight</td><td valign="top" align="left" rowspan="1" colspan="1">↓ placental sinusoids</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↑ Spleen weight</td><td valign="top" align="left" rowspan="1" colspan="1">IEs/Hz</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">No congenital malaria</td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B62" ref-type="bibr">Oduola et al.,1986</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">Sprague-Dawley</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">Hyperplasia</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">NK65</td><td valign="top" align="left" rowspan="1" colspan="1">ICR</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">Necrosis/fibrin</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">Impaired labyrinth structure</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">Trophoblast barrier thickening</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">IEs/Hz</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">MΦ/Leu</td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B22" ref-type="bibr">Desowitz et al.,1989</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">Wistar</td><td valign="top" align="left" rowspan="1" colspan="1">No recrudescence</td><td valign="top" align="left" rowspan="1" colspan="1">↑ In pregnancy</td><td valign="top" align="left" rowspan="1" colspan="1">↑ In early infection</td><td valign="top" align="left" rowspan="1" colspan="1">No preterm delivery</td><td valign="top" align="left" rowspan="1" colspan="1">Normal litter size</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Placental parasitemia</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">NYU-2</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ in mid gestation infection with ↑ parasitemia at term</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B32" ref-type="bibr">Hioki et al.,1990</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">BALB/c</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ In pregnancy</td><td valign="top" align="left" rowspan="1" colspan="1">↑ In early infection (death before term)</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">NK65</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↑ in mid gestation infection</td><td valign="top" align="left" rowspan="1" colspan="1">↓ in late infection (live until term)</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B68" ref-type="bibr">Pathak et al.,1990</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">Swiss Albino</td><td valign="top" align="left" rowspan="1" colspan="1">No recrudescence</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Maternal mortality</td><td valign="top" align="left" rowspan="1" colspan="1">Stillbirth → to mid gestation infection</td><td valign="top" align="left" rowspan="1" colspan="1">↓ Weight</td><td valign="top" align="left" rowspan="1" colspan="1">Hyperplasia</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">NICD</td><td valign="top" align="left" rowspan="1" colspan="1">VRC</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ in treated pregnant mice</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ Litter size</td><td valign="top" align="left" rowspan="1" colspan="1">IEs</td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B69" ref-type="bibr">Pavia and Niederbuhl,1991</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. yoelii</italic> YM</td><td valign="top" align="left" rowspan="1" colspan="1">ICR</td><td valign="top" align="left" rowspan="1" colspan="1">No recrudescence</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Maternal mortality in non-immunized pregnancies</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">Delivery of dead pups in non-immunized pregnancies</td><td valign="top" align="center" rowspan="1" colspan="1">X</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. yoelii</italic> 17X</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ with ↓ parasite virulence</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">No congenital malaria</td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B94" ref-type="bibr">Tegoshi et al.,1992</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">Wistar</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">Hyperplasia</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">NYU-2</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">Necrosis/fibrin</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">Impaired labyrinth structure</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">Trophoblast barrier thickening</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↑ IEs after late infection</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">MΦ/Leu</td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B2" ref-type="bibr">Adachi et al.,2000</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">BALB/c</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">Congenital malaria</td><td valign="top" align="center" rowspan="1" colspan="1">X</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">ANKA</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B70" ref-type="bibr">Poovassery and Moore,2006</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. chabaudi</italic></td><td valign="top" align="left" rowspan="1" colspan="1">C57BL/6</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ In pregnancy → to</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Mortality → pregnancy</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Abortion/resorption at mid gestation → to early infection</td><td valign="top" align="left" rowspan="1" colspan="1">↓ Fetal viability</td><td valign="top" align="left" rowspan="1" colspan="1">↑ IEs at mid gestation</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">AS</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↑ parasitemia</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ monocyte accumulation</td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B59" ref-type="bibr">Neres et al.,2008</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">BALB/c</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Mortality → pregnancy</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Abortion/resorption during infection</td><td valign="top" align="left" rowspan="1" colspan="1">↓ Fetal/birth weight</td><td valign="top" align="left" rowspan="1" colspan="1">Hyperplasia</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">ANKA</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↑ preterm delivery during infection</td><td valign="top" align="left" rowspan="1" colspan="1">↓ fetal blood flow/content</td><td valign="top" align="left" rowspan="1" colspan="1">Necrosis/fibrin</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ viability</td><td valign="top" align="left" rowspan="1" colspan="1">Impaired labyrinth structure</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">Trophoblast barrier thickening</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">IEs/Hz</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">MΦ</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ Placental vasculature</td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B71" ref-type="bibr">Poovassery and Moore,2009</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. chabaudi</italic></td><td valign="top" align="left" rowspan="1" colspan="1">C57BL/6</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Abortion at mid gestation → to early infection</td><td valign="top" align="left" rowspan="1" colspan="1">↓ Fetal viability</td><td valign="top" align="left" rowspan="1" colspan="1">IEs/Hz</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">AS</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B72" ref-type="bibr">Poovassery et al.,2009</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. chabaudi</italic></td><td valign="top" align="left" rowspan="1" colspan="1">C57BL/6</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ In pregnancy → to</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Mortality → pregnancy</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Abortion/resorption at mid gestation → to early infection</td><td valign="top" align="left" rowspan="1" colspan="1">↓ Fetal viability → to IFN-γ/TNF-α</td><td valign="top" align="left" rowspan="1" colspan="1">Fibrin thrombi</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">AS</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↑ parasitemia</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">Placental hemorrhage</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">Impaired labyrinth structure</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">Monocytes/macrophages</td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B50" ref-type="bibr">Megnekou et al.,2009</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">BALB/c</td><td valign="top" align="left" rowspan="1" colspan="1">→ To pregnancy</td><td valign="top" align="left" rowspan="1" colspan="1">→ To recrudescence</td><td valign="top" align="left" rowspan="1" colspan="1">→ To recrudescence</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">→ To recrudescence</td><td valign="top" align="left" rowspan="1" colspan="1">iRBCs/Hz</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">K173</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↑ primigravidae</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↑ due to ↓ immunity</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ Litter size</td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ Weight</td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B47" ref-type="bibr">Marinho et al.,2009</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">BALB/c</td><td valign="top" align="left" rowspan="1" colspan="1">→ To pregnancy</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">→ To recrudescence</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">→ To recrudescence</td><td valign="top" align="left" rowspan="1" colspan="1">Trophoblast barrier thickening</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">ANKA</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↑ primigravidae</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ Litter size</td><td valign="top" align="left" rowspan="1" colspan="1">IEs</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ multigravidae</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ Weight</td><td valign="top" align="left" rowspan="1" colspan="1">MΦ/Tc/NKc</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ Primigravidae</td><td valign="top" align="left" rowspan="1" colspan="1">↓ Placental vascular spaces</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↑ Multigravidae</td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B90" ref-type="bibr">Silver et al.,2010</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">BALB/c</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Abortion/resorption at late-gestation → to mid gestation infection</td><td valign="top" align="left" rowspan="1" colspan="1">↓ Fetal weight at late gestation</td><td valign="top" align="center" rowspan="1" colspan="1">X</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">ANKA</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ fetal viability</td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B84" ref-type="bibr">Sarr et al.,2012</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. chabaudi</italic></td><td valign="top" align="left" rowspan="1" colspan="1">A/J</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ In pregnant A/J mice when compared to C57BL/6</td><td valign="top" align="left" rowspan="1" colspan="1">↑ In pregnant A/J mice when compared to C57BL/6</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Abortion/resorption in A/J and C57BL/6</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Placental IEs in A/J when comparing to C57BL/6</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">AS</td><td valign="top" align="left" rowspan="1" colspan="1">C57BL/6</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B75" ref-type="bibr">Rodrigues-Duarte et al.,2012</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">C57BL/6</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ In pregnant mice infected with <italic>P. berghei</italic> NK65</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Stillbirth → mid gestation infection</td><td valign="top" align="left" rowspan="1" colspan="1">↓ Fetal weight at late gestation</td><td valign="top" align="left" rowspan="1" colspan="1">Necrosis/fibrin</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">K173</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ newborn viability</td><td valign="top" align="left" rowspan="1" colspan="1">Impaired labyrinth structure</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">NK65</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">Trophoblast barrier thickening</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">ANKAΔpm4</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">IEs</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ Placental vascular spaces</td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B4" ref-type="bibr">Avery et al.,2012</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. chabaudi</italic></td><td valign="top" align="left" rowspan="1" colspan="1">C57BL/6</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ In pregnancy → to</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Abortion at mid gestation → to early infection → to coagulation and impaired fibrinolysis</td><td valign="top" align="left" rowspan="1" colspan="1">↓ Fetal viability</td><td valign="top" align="left" rowspan="1" colspan="1">Necrosis/fibrin</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">AS</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↑ parasitemia</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">Impaired labyrinth structure</td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B17" ref-type="bibr">Conroy et al.,2013</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">BALB/c</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Fetal weight and ↑ viability in infected C5aR<sup>–/–</sup> mice</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Placental vascular remodeling in infected C5aR<sup>–/–</sup> mice</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">ANKA</td><td valign="top" align="left" rowspan="1" colspan="1">Wildtype</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">C5aR<sup>–/–</sup></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B19" ref-type="bibr">de Moraes et al.,2013</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">BALB/c ♀</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">Impaired labyrinth structure</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">ANKA</td><td valign="top" align="left" rowspan="1" colspan="1">C57BL/6 ♂</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">Trophoblast barrier thickening</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">Placental vasculature remodeling</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">IEs in ↓ blood flow areas</td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B7" ref-type="bibr">Barboza et al.,2014</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">C57BL/6</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Fetal weight in infected MyD88<sup>–/–</sup> mice</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Placental vascular spaces in infected MyD88<sup>–/–</sup> mice</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">NK65</td><td valign="top" align="left" rowspan="1" colspan="1">Wildtype</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↑ survival in newborn from infected MyD88<sup>–/–</sup> mice</td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">MyD88<sup>–/–</sup></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B87" ref-type="bibr">Sharma and Shukla,2014</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">BALB/c</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↓ In pregnant-infected mice treated with CQ/SP</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Weight and ↑ viability in newborn from mice treated with CQ/SP</td><td valign="top" align="left" rowspan="1" colspan="1">↓ IEs, ↓ Hz, and normal placental histology in mice treated with CQ/SP</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">NK65</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B43" ref-type="bibr">Lima et al.,2014</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">C57BL/6</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">Accumulation of iRBCs mature forms in the placenta</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. chabaudi</italic></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">IEs uptake by trophoblast</td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B83" ref-type="bibr">Sarr et al.,2015</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. chabaudi</italic></td><td valign="top" align="left" rowspan="1" colspan="1">A/J</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Abortion/resorption in A/J and C57BL/6</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Tc, Bc, NKc in conceptus from infected A/J and C57BL/6 mice</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">AS</td><td valign="top" align="left" rowspan="1" colspan="1">C57BL/6</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↑ MΦ in the placental junctional zone of A/J mice</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">Apoptosis present in immune cells and spongiotrophoblast</td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B86" ref-type="bibr">Sharma et al.,2016</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. chabaudi</italic></td><td valign="top" align="left" rowspan="1" colspan="1">C57BL/6</td><td valign="top" align="left" rowspan="1" colspan="1">→ To pregnancy</td><td valign="top" align="left" rowspan="1" colspan="1">→ To recrudescence</td><td valign="top" align="left" rowspan="1" colspan="1">↑ In reinfections but not in recrudescence</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Stillbirth in reinfections and → to high parasitemia</td><td valign="top" align="left" rowspan="1" colspan="1">Normal litter size</td><td valign="top" align="left" rowspan="1" colspan="1">IEs in recrudescence</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">CB</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">→ to CB sporozoites</td><td valign="top" align="left" rowspan="1" colspan="1">→ to reinfection</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↓ weight and ↓ malaria susceptibility in reinfection</td><td valign="top" align="left" rowspan="1" colspan="1">No histologic alterations</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. chabaudi</italic></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">AS</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B6" ref-type="bibr">Barboza et al.,2017</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">C57BL/6</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↑ Fetal weight in infected TLR4<sup>–/–</sup> mice</td><td valign="top" align="left" rowspan="1" colspan="1">IEs</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">NK65</td><td valign="top" align="left" rowspan="1" colspan="1">Wildtype</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">MΦ/NΦ/Dc</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">TLR2<sup>–/–</sup></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">TLR4<sup>–/–</sup></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">TLR9<sup>–/–</sup></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">↑ placental vascular spaces in infected TLR4<sup>–/–</sup> mice</td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B76" ref-type="bibr">Rodrigues-Duarte et al.,2018</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">C57BL/6</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↓ Stillbirth in TLR4<sup>–/–</sup> and IFNAR1<sup>–/–</sup> infected pregnant mice carrying TLR4<sup>+/–</sup> and IFNAR1<sup>+/–</sup> progeny</td><td valign="top" align="left" rowspan="1" colspan="1">Wildtype and TLR4<sup>–/–</sup> infected mice with similar litter weight ↑ weight of IFNAR1<sup>+/–</sup> fetuses from infected IFNAR1<sup>–/–</sup> mice</td><td valign="top" align="center" rowspan="1" colspan="1">X</td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">NK65</td><td valign="top" align="left" rowspan="1" colspan="1">Wildtype</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">TLR4<sup>–/–</sup></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">IFNAR1<sup>–/–</sup></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="9" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><xref rid="B5" ref-type="bibr">Barboza et al.,2019</xref></td><td valign="top" align="left" rowspan="1" colspan="1"><italic>P. berghei</italic></td><td valign="top" align="left" rowspan="1" colspan="1">C57BL/6</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="center" rowspan="1" colspan="1">X</td><td valign="top" align="left" rowspan="1" colspan="1">↓ Fetal weight in MyD88<sup>+/–</sup> progeny born from MyD88<sup>–/–</sup></td><td valign="top" align="left" rowspan="1" colspan="1">↓ Placental vascular spaces in MyD88<sup>+/–</sup> placentas from MyD88<sup>–/–</sup></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">NK65</td><td valign="top" align="left" rowspan="1" colspan="1">Wildtype</td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1">MyD88<sup>–/–</sup></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="left" rowspan="1" colspan="1"></td></tr></tbody></table><table-wrap-foot><attrib><italic>The current table content aims to summarize research conducted during the past four decades that was used to inform this review. In summary, <italic>Plasmodium</italic> and murine strains are described for each reference, as well as observations made regarding pregnancy outcomes and placental histology. Information regarding symbology is depicted as follows: Increased (↑), decreased (↓), associated (→), males (♂), females (♀), infected erythrocytes (IEs), hemozoin (Hz), leukocytes (Leu), monocytes/macrophages (MΦ), T cells (Tc), B cells (Bc), NK cells (NKc), dendritic cells (Dc), neutrophils (NΦ), chloroquine (CQ), sulphadoxine-pyrimethamine (SP).</italic></attrib></table-wrap-foot></table-wrap><table-wrap id="T2" position="float"><label>TABLE 2</label><caption><p>Compilation of references mentioning the main pathologic manifestations of malaria in pregnancy observed in humans and murine models.</p></caption><table frame="hsides" rules="groups" cellspacing="5" cellpadding="5"><thead><tr><td valign="top" align="left" rowspan="1" colspan="1"></td><td valign="top" align="justify" rowspan="1" colspan="1"><bold>Human MiP</bold></td><td valign="top" align="justify" rowspan="1" colspan="1"><bold>Murine MiP</bold></td></tr></thead><tbody><tr><td valign="top" align="left" rowspan="1" colspan="1"><bold>Pregnancy outcome</bold></td><td valign="top" align="justify" rowspan="1" colspan="1"></td><td valign="top" align="justify" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="3" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1">Recrudescence</td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B28" ref-type="bibr">Giobbia et al., 2005</xref>; <xref rid="B48" ref-type="bibr">Mayor et al., 2009</xref>; <xref rid="B41" ref-type="bibr">Laochan et al., 2015</xref>; <xref rid="B3" ref-type="bibr">Al Hammadi et al., 2017</xref>; <xref rid="B46" ref-type="bibr">Malvy et al.,2018</xref></td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B97" ref-type="bibr">van Zon and Eling, 1980</xref>; <xref rid="B61" ref-type="bibr">Oduola et al., 1982</xref>; <xref rid="B99" ref-type="bibr">van Zon et al., 1982</xref>, <xref rid="B98" ref-type="bibr">1985</xref>; <xref rid="B47" ref-type="bibr">Marinho et al., 2009</xref>; <xref rid="B50" ref-type="bibr">Megnekou et al., 2009</xref>; <xref rid="B86" ref-type="bibr">Sharma et al.,2016</xref></td></tr><tr><td valign="top" align="left" colspan="3" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1">Anemia</td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B12" ref-type="bibr">Brabin et al., 2001</xref>; <xref rid="B79" ref-type="bibr">Rogerson et al.,2003</xref></td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B61" ref-type="bibr">Oduola et al., 1982</xref>; <xref rid="B22" ref-type="bibr">Desowitz et al., 1989</xref>; <xref rid="B32" ref-type="bibr">Hioki et al., 1990</xref>; <xref rid="B50" ref-type="bibr">Megnekou et al., 2009</xref>; <xref rid="B72" ref-type="bibr">Poovassery et al., 2009</xref>; <xref rid="B4" ref-type="bibr">Avery et al., 2012</xref>; <xref rid="B84" ref-type="bibr">Sarr et al., 2012</xref>; <xref rid="B86" ref-type="bibr">Sharma et al.,2016</xref></td></tr><tr><td valign="top" align="left" colspan="3" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1">Maternal mortality</td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B45" ref-type="bibr">Luxemburger et al., 1997</xref>; <xref rid="B12" ref-type="bibr">Brabin et al., 2001</xref>; <xref rid="B89" ref-type="bibr">Shulman et al., 2002</xref>; <xref rid="B60" ref-type="bibr">Nosten et al., 2004</xref>; <xref rid="B53" ref-type="bibr">Menéndez et al., 2008</xref>; <xref rid="B77" ref-type="bibr">Rogerson et al.,2018</xref></td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B97" ref-type="bibr">van Zon and Eling, 1980</xref>; <xref rid="B61" ref-type="bibr">Oduola et al., 1982</xref>; <xref rid="B99" ref-type="bibr">van Zon et al., 1982</xref>; <xref rid="B101" ref-type="bibr">Vinayak et al., 1986</xref>; <xref rid="B22" ref-type="bibr">Desowitz et al., 1989</xref>; <xref rid="B32" ref-type="bibr">Hioki et al., 1990</xref>; <xref rid="B68" ref-type="bibr">Pathak et al., 1990</xref>; <xref rid="B69" ref-type="bibr">Pavia and Niederbuhl, 1991</xref>; <xref rid="B70" ref-type="bibr">Poovassery and Moore, 2006</xref>; <xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>; <xref rid="B47" ref-type="bibr">Marinho et al., 2009</xref>; <xref rid="B50" ref-type="bibr">Megnekou et al., 2009</xref>; <xref rid="B72" ref-type="bibr">Poovassery et al., 2009</xref>; <xref rid="B75" ref-type="bibr">Rodrigues-Duarte et al., 2012</xref>; <xref rid="B84" ref-type="bibr">Sarr et al., 2012</xref>; <xref rid="B87" ref-type="bibr">Sharma and Shukla, 2014</xref>; <xref rid="B86" ref-type="bibr">Sharma et al.,2016</xref></td></tr><tr><td valign="top" align="left" colspan="3" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1">Stillbirth</td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B51" ref-type="bibr">Menendez, 1995</xref>; <xref rid="B21" ref-type="bibr">Desai et al.,2007</xref></td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B61" ref-type="bibr">Oduola et al., 1982</xref>; <xref rid="B101" ref-type="bibr">Vinayak et al., 1986</xref>; <xref rid="B68" ref-type="bibr">Pathak et al., 1990</xref>; <xref rid="B75" ref-type="bibr">Rodrigues-Duarte et al., 2012</xref>, <xref rid="B76" ref-type="bibr">2018</xref>; <xref rid="B86" ref-type="bibr">Sharma et al.,2016</xref></td></tr><tr><td valign="top" align="left" colspan="3" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1">Abortion</td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B21" ref-type="bibr">Desai et al.,2007</xref></td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B70" ref-type="bibr">Poovassery and Moore, 2006</xref>, <xref rid="B71" ref-type="bibr">2009</xref>; <xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>; <xref rid="B72" ref-type="bibr">Poovassery et al., 2009</xref>; <xref rid="B90" ref-type="bibr">Silver et al., 2010</xref>; <xref rid="B4" ref-type="bibr">Avery et al., 2012</xref>; <xref rid="B84" ref-type="bibr">Sarr et al.,2012</xref>, <xref rid="B83" ref-type="bibr">2015</xref></td></tr><tr><td valign="top" align="left" colspan="3" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1">Preterm delivery</td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B52" ref-type="bibr">Menendez et al., 2000</xref>; <xref rid="B21" ref-type="bibr">Desai et al., 2007</xref>; <xref rid="B54" ref-type="bibr">Moore et al.,2017</xref></td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B99" ref-type="bibr">van Zon et al., 1982</xref>; <xref rid="B59" ref-type="bibr">Neres et al.,2008</xref></td></tr><tr><td valign="top" align="left" colspan="3" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1">Reduced newborn/fetal weight</td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B52" ref-type="bibr">Menendez et al., 2000</xref>; <xref rid="B79" ref-type="bibr">Rogerson et al., 2003</xref>; <xref rid="B60" ref-type="bibr">Nosten et al., 2004</xref>; <xref rid="B21" ref-type="bibr">Desai et al., 2007</xref>; <xref rid="B95" ref-type="bibr">Umbers et al.,2011</xref></td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B61" ref-type="bibr">Oduola et al., 1982</xref>; <xref rid="B101" ref-type="bibr">Vinayak et al., 1986</xref>; <xref rid="B68" ref-type="bibr">Pathak et al., 1990</xref>; <xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>; <xref rid="B47" ref-type="bibr">Marinho et al., 2009</xref>; <xref rid="B50" ref-type="bibr">Megnekou et al., 2009</xref>; <xref rid="B90" ref-type="bibr">Silver et al., 2010</xref>; <xref rid="B75" ref-type="bibr">Rodrigues-Duarte et al., 2012</xref>, <xref rid="B76" ref-type="bibr">2018</xref>; <xref rid="B17" ref-type="bibr">Conroy et al., 2013</xref>; <xref rid="B7" ref-type="bibr">Barboza et al., 2014</xref>, <xref rid="B6" ref-type="bibr">2017</xref>, <xref rid="B5" ref-type="bibr">2019</xref>; <xref rid="B87" ref-type="bibr">Sharma and Shukla, 2014</xref>; <xref rid="B86" ref-type="bibr">Sharma et al.,2016</xref></td></tr><tr><td valign="top" align="left" colspan="3" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1">Congenital malaria</td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B73" ref-type="bibr">Rai et al., 2015</xref>; <xref rid="B11" ref-type="bibr">Bhatia et al.,2016</xref></td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B2" ref-type="bibr">Adachi et al.,2000</xref></td></tr><tr><td valign="top" align="left" colspan="3" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1"><bold>Placental histology</bold></td><td valign="top" align="justify" rowspan="1" colspan="1"></td><td valign="top" align="justify" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="3" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1">Parasite accumulation</td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B102" ref-type="bibr">Walter et al., 1982</xref>; <xref rid="B8" ref-type="bibr">Beeson et al., 2002</xref>; <xref rid="B9" ref-type="bibr">Beeson and Brown, 2004</xref>; <xref rid="B57" ref-type="bibr">Muthusamy et al.,2004</xref>, <xref rid="B58" ref-type="bibr">2007</xref></td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B61" ref-type="bibr">Oduola et al., 1982</xref>, <xref rid="B62" ref-type="bibr">1986</xref>; <xref rid="B101" ref-type="bibr">Vinayak et al., 1986</xref>; <xref rid="B22" ref-type="bibr">Desowitz et al., 1989</xref>; <xref rid="B68" ref-type="bibr">Pathak et al., 1990</xref>; <xref rid="B94" ref-type="bibr">Tegoshi et al., 1992</xref>; <xref rid="B70" ref-type="bibr">Poovassery and Moore, 2006</xref>, <xref rid="B71" ref-type="bibr">2009</xref>; <xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>; <xref rid="B47" ref-type="bibr">Marinho et al., 2009</xref>; <xref rid="B50" ref-type="bibr">Megnekou et al., 2009</xref>; <xref rid="B75" ref-type="bibr">Rodrigues-Duarte et al., 2012</xref>; <xref rid="B84" ref-type="bibr">Sarr et al., 2012</xref>; <xref rid="B19" ref-type="bibr">de Moraes et al., 2013</xref>; <xref rid="B43" ref-type="bibr">Lima et al., 2014</xref>; <xref rid="B87" ref-type="bibr">Sharma and Shukla, 2014</xref>; <xref rid="B86" ref-type="bibr">Sharma et al., 2016</xref>; <xref rid="B6" ref-type="bibr">Barboza et al.,2017</xref></td></tr><tr><td valign="top" align="left" colspan="3" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1">Hemozoin deposition</td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B13" ref-type="bibr">Bulmer et al., 1993</xref>; <xref rid="B34" ref-type="bibr">Ismail et al.,2000</xref></td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B61" ref-type="bibr">Oduola et al., 1982</xref>, <xref rid="B62" ref-type="bibr">1986</xref>; <xref rid="B101" ref-type="bibr">Vinayak et al., 1986</xref>; <xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>; <xref rid="B50" ref-type="bibr">Megnekou et al., 2009</xref>; <xref rid="B71" ref-type="bibr">Poovassery and Moore,2009</xref></td></tr><tr><td valign="top" align="left" colspan="3" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1">Immune cells infiltrate</td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B102" ref-type="bibr">Walter et al., 1982</xref>; <xref rid="B64" ref-type="bibr">Ordi et al., 1998</xref>; <xref rid="B34" ref-type="bibr">Ismail et al., 2000</xref>; <xref rid="B78" ref-type="bibr">Rogerson et al., 2007</xref>; <xref rid="B66" ref-type="bibr">Othoro et al., 2008</xref>; <xref rid="B91" ref-type="bibr">Souza et al.,2013</xref></td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B61" ref-type="bibr">Oduola et al., 1982</xref>, <xref rid="B62" ref-type="bibr">1986</xref>; <xref rid="B94" ref-type="bibr">Tegoshi et al., 1992</xref>; <xref rid="B70" ref-type="bibr">Poovassery and Moore, 2006</xref>; <xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>; <xref rid="B47" ref-type="bibr">Marinho et al., 2009</xref>; <xref rid="B72" ref-type="bibr">Poovassery et al., 2009</xref>; <xref rid="B6" ref-type="bibr">Barboza et al.,2017</xref></td></tr><tr><td valign="top" align="left" colspan="3" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1">Fibrin Necrosis</td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B102" ref-type="bibr">Walter et al., 1982</xref>; <xref rid="B34" ref-type="bibr">Ismail et al., 2000</xref>; <xref rid="B91" ref-type="bibr">Souza et al.,2013</xref></td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B62" ref-type="bibr">Oduola et al., 1986</xref>; <xref rid="B94" ref-type="bibr">Tegoshi et al., 1992</xref>; <xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>; <xref rid="B72" ref-type="bibr">Poovassery et al., 2009</xref>; <xref rid="B4" ref-type="bibr">Avery et al., 2012</xref>; <xref rid="B75" ref-type="bibr">Rodrigues-Duarte et al.,2012</xref></td></tr><tr><td valign="top" align="left" colspan="3" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1">Syncytial knots Trophoblast rupture</td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B34" ref-type="bibr">Ismail et al., 2000</xref>; <xref rid="B18" ref-type="bibr">Crocker et al., 2004</xref>; <xref rid="B91" ref-type="bibr">Souza et al.,2013</xref></td><td valign="top" align="justify" rowspan="1" colspan="1"></td></tr><tr><td valign="top" align="left" colspan="3" rowspan="1"><hr></hr></td></tr><tr><td valign="top" align="left" rowspan="1" colspan="1">Trophoblast membrane thickening</td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B34" ref-type="bibr">Ismail et al., 2000</xref>; <xref rid="B91" ref-type="bibr">Souza et al.,2013</xref></td><td valign="top" align="justify" rowspan="1" colspan="1"><xref rid="B61" ref-type="bibr">Oduola et al., 1982</xref>, <xref rid="B62" ref-type="bibr">1986</xref>; <xref rid="B94" ref-type="bibr">Tegoshi et al., 1992</xref>; <xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>; <xref rid="B47" ref-type="bibr">Marinho et al., 2009</xref>; <xref rid="B75" ref-type="bibr">Rodrigues-Duarte et al., 2012</xref>; <xref rid="B19" ref-type="bibr">de Moraes et al., 2013</xref>; <xref rid="B7" ref-type="bibr">Barboza et al.,2014</xref>, <xref rid="B6" ref-type="bibr">2017</xref>, <xref rid="B5" ref-type="bibr">2019</xref></td></tr></tbody></table><table-wrap-foot><attrib><italic>Distinct pregnancy outcomes and placental histopathologic alterations are depicted. References used to inform this review are cited in the table according to their contribution in unraveling specific clinical and histologic manifestations of both human and murine malaria in pregnancy.</italic></attrib></table-wrap-foot></table-wrap></sec></sec><sec id="S3"><title>Contribution of Murine Models to the Study of MiP</title><sec id="S3.SS1"><title>Understanding Recrudescence Using Murine Models of MiP</title><p>Currently, we have been facing the emerging problem of drug resistance acquisition observed in <italic>P. falciparum</italic> infections, the outcome of which might be the occurrence of recrudescence (without a new infection episode) as a consequence of subcurative therapy (<xref rid="B16" ref-type="bibr">Cattamanchi et al., 2003</xref>; <xref rid="B10" ref-type="bibr">Berrevoets et al., 2013</xref>). In addition to treatment incapacity to clear circulating parasites, the immune depression observed in specific situations, such as during pregnancy, might elicit the emergence of recrudescent parasites that are either dormant or at submicroscopic levels (<xref rid="B28" ref-type="bibr">Giobbia et al., 2005</xref>; <xref rid="B48" ref-type="bibr">Mayor et al., 2009</xref>; <xref rid="B41" ref-type="bibr">Laochan et al., 2015</xref>; <xref rid="B3" ref-type="bibr">Al Hammadi et al., 2017</xref>; <xref rid="B46" ref-type="bibr">Malvy et al., 2018</xref>). In this context, some of the first available studies using murine models of MiP appeared during the early 1980s, when van Zon and Eling first described recrudescence in pregnant mice infected with <italic>P. berghei</italic> (<xref rid="B97" ref-type="bibr">van Zon and Eling, 1980</xref>; <xref rid="B99" ref-type="bibr">van Zon et al., 1982</xref>). Females of different mouse strains (Swiss, C3H/StZ, BALB/c, and B10LP) challenged with <italic>P. berghei</italic> K173 were treated with chloroquine and sulfadiazine to decrease parasitemia and promote the acquisition of immunity before mating. Afterward, pregnancy was shown to induce recrudescence, which was clearly shown to be strain- and gravidity-dependent (<xref rid="B97" ref-type="bibr">van Zon and Eling, 1980</xref>) and linked to preterm delivery and maternal mortality (<xref rid="B99" ref-type="bibr">van Zon et al., 1982</xref>). Consequently, the pioneering results enabled the authors to draw brief conclusions on similarities observed between human and murine recrudescence by determining that the phenomenon could be associated with (1) the drugs’ incapacity to completely clear the parasite in a previous malaria episode and (2) the particular immune status associated with pregnancy, recapitulating the features of recrudescence in human MiP (<xref rid="B28" ref-type="bibr">Giobbia et al., 2005</xref>; <xref rid="B3" ref-type="bibr">Al Hammadi et al., 2017</xref>; <xref rid="B46" ref-type="bibr">Malvy et al., 2018</xref>). Pregnancy-associated immune modulation was then shown to be linked to increased production of corticoids during murine gestation, thereby facilitating the onset of recrudescence (<xref rid="B99" ref-type="bibr">van Zon et al., 1982</xref>, <xref rid="B98" ref-type="bibr">1985</xref>). Notwithstanding the relevance of van Zon and coworkers’ findings, parasite reappearance in maternal circulation was often ensured by experimental reinfection, raising questions about the translational reliability of murine recrudescence results obtained in these models (<xref rid="B97" ref-type="bibr">van Zon and Eling, 1980</xref>; <xref rid="B99" ref-type="bibr">van Zon et al., 1982</xref>).</p><p>Despite the promising advances made during that period, recrudescence in murine models was poorly addressed until the last decade. Only in 2009, two studies have addressed questions about the topic (<xref rid="B47" ref-type="bibr">Marinho et al., 2009</xref>; <xref rid="B50" ref-type="bibr">Megnekou et al., 2009</xref>). By reevaluating the pioneering experimental model implemented by van Zon and coworkers, the team of Megnekou addressed the production of specific anti-VSA antibodies in pregnant BALB/c mice immunized against <italic>P. berghei</italic> K173. Although previously immunized, protection was lost during pregnancy due to the recrudescence of parasites expressing specific VSA. Consequently, protection was shown to be gravidity-dependent since the less susceptible multiparous mice have raised antibodies against parasite-specific VSA during their previous pregnancies (<xref rid="B50" ref-type="bibr">Megnekou et al., 2009</xref>). However, some parasite reappearances were once again ensured by experimental infections, raising the same translational limitations discussed above. During that same year, a study from <xref rid="B47" ref-type="bibr">Marinho et al. (2009)</xref> established a model to study recrudescence in BALB/c mice, however, without recurring to mid-gestation reinfection. Accordingly, non-pregnant mice were infected with <italic>P. berghei</italic> ANKA and administered a subcurative treatment to clear apparent parasitemia. Mice were then mated approximately 1 month later and analyzed for recrudescence, which occurred in up to 58% of pregnant mice from gestational day (G)12 onward (<xref rid="B47" ref-type="bibr">Marinho et al., 2009</xref>). Maternal mortality and parasitemia were shown to be increased in primigravidae, which diminished with increased gravidity, supporting the observations of <xref rid="B50" ref-type="bibr">Megnekou et al. (2009)</xref> regarding acquired immunity in subsequent pregnancies.</p><p>Nevertheless, both recrudescent and newly inoculated parasites constitute a risk for highly susceptible pregnant women (<xref rid="B41" ref-type="bibr">Laochan et al., 2015</xref>). Accordingly, a recent study has shown that <italic>P. chabaudi</italic> CB sporozoites are capable of inducing recrudescence in pregnant C57BL/6 mice, being more closely related to the human parasite life cycle. However, poor pregnancy outcomes, such as maternal mortality and reduced birth weight, were likely to occur upon heterologous reinfection with <italic>P. chabaudi</italic> AS rather than in the cases of recrudescence. Preacquired immunity against the recrudescent parasite might justify why only reinfection with heterologous parasite lead to poor pregnancy outcomes. Nonetheless, exposure to heterologous <italic>P. chabaudi</italic> AS ensures newborn protection against postnatal infection, reliably resembling the outcomes of human MiP (<xref rid="B86" ref-type="bibr">Sharma et al., 2016</xref>).</p><p>Although observed in murine MiP, some studies have reported no recrudescence in protocols of treatment and immunization used in mice to control infection before pregnancy (<xref rid="B68" ref-type="bibr">Pathak et al., 1990</xref>; <xref rid="B69" ref-type="bibr">Pavia and Niederbuhl, 1991</xref>). Explanations might rely on the (1) drug efficiency to clear circulating parasite, (2) usage of less virulent strains (e.g., <italic>P. berghei</italic> NICD and <italic>P. yoelii</italic> 17X), or (3) less susceptible mice strains (e.g., Swiss Albino VRC). Nonetheless, despite some controversial experimental settings, findings associated with the aforementioned murine models have revealed some hidden biological features associated with malaria recrudescence during pregnancy that otherwise would be impossible to address.</p></sec><sec id="S3.SS2"><title>Understanding Poor Pregnancy Outcomes Using Murine Models of MiP</title><p>A multitude of outcomes can occur as a result of developing MiP. When considering the areas of <italic>P. falciparum</italic> frequent and stable transmission, pregnancy-associated consequences might depend on several aspects such as infection trimester, gravidity and maternal health status that ultimately dictate the onset of maternal anemia, fetal and maternal mortality, abortion, and reduced birth weight as consequences of IUGR and/or preterm delivery (<xref rid="B21" ref-type="bibr">Desai et al., 2007</xref>; <xref rid="B95" ref-type="bibr">Umbers et al., 2011</xref>; <xref rid="B77" ref-type="bibr">Rogerson et al., 2018</xref>). Outcomes in human pregnancy are accessible for study; however, in addition to ethical constraints, illogical and incorrect correlations might be drawn from imprecise information given by patients enrolling in these prospective studies, such as imprecise time of infection or clinical status. Fittingly, murine models provided the opportunity to investigate MiP outcomes in a controlled experimental setting.</p><sec id="S3.SS2.SSS1"><title>Maternal Anemia</title><p>Maternal anemia that develops during MiP is considered a significant risk factor for poor pregnancy outcomes and is closely associated with increased parasite burden and reduced birth weight (<xref rid="B79" ref-type="bibr">Rogerson et al., 2003</xref>). Accordingly, several distinct experimental murine models accurately recapitulate anemia observed during human MiP. Similarly, <italic>P. chabaudi</italic> AS infection potentiates the onset of anemia (reduced hematocrit percentage) in pregnant C57BL/6 mice. Anemia was therefore linked to a rapid increase in parasitemia, which peaks abruptly in pregnant mice (<xref rid="B70" ref-type="bibr">Poovassery and Moore, 2006</xref>; <xref rid="B72" ref-type="bibr">Poovassery et al., 2009</xref>; <xref rid="B4" ref-type="bibr">Avery et al., 2012</xref>; <xref rid="B84" ref-type="bibr">Sarr et al., 2012</xref>). This phenomenon was also shown to occur in different murine strains, such as BALB/c mice (<xref rid="B32" ref-type="bibr">Hioki et al., 1990</xref>) and Wistar rats (<xref rid="B22" ref-type="bibr">Desowitz et al., 1989</xref>), infected with <italic>P. berghei</italic> NK65 and NYU-2, respectively, in which hemoglobin and hematocrit decreased as a result of increased parasitemia. Reproducibility of anemia is expected among different experimental models since hemolysis constitutes an intrinsic step in the <italic>Plasmodium</italic> spp. life cycle in both rodents and humans. Nevertheless, this critical parameter has been poorly investigated in murine malaria, in which its relationship with infection timing, gravidity, and pregnancy outcomes were never addressed as in humans (<xref rid="B12" ref-type="bibr">Brabin et al., 2001</xref>; <xref rid="B79" ref-type="bibr">Rogerson et al., 2003</xref>).</p></sec><sec id="S3.SS2.SSS2"><title>Maternal Mortality</title><p>Maternal mortality seems to be a less frequent outcome in pregnancies complicated by malaria (<xref rid="B53" ref-type="bibr">Menéndez et al., 2008</xref>; <xref rid="B77" ref-type="bibr">Rogerson et al., 2018</xref>). Maternal death associated with <italic>Plasmodium</italic> spp. infection tends to occur more frequently in areas of low malaria transmission due to the reduced level of premunition (<xref rid="B60" ref-type="bibr">Nosten et al., 2004</xref>). Nevertheless, pregnant women with malaria who live in endemic areas are also prone to die due to severe anemia (<xref rid="B12" ref-type="bibr">Brabin et al., 2001</xref>; <xref rid="B89" ref-type="bibr">Shulman et al., 2002</xref>). Likewise, both pregnant mice and rats infected with murine <italic>Plasmodium</italic> spp. strains were shown to die during gestation, which was dependent on the parasite strain, infection timing, and host intrinsic characteristics. Regarding parasite strains, it was demonstrated that <italic>P. berghei</italic> (<xref rid="B97" ref-type="bibr">van Zon and Eling, 1980</xref>; <xref rid="B61" ref-type="bibr">Oduola et al., 1982</xref>; <xref rid="B99" ref-type="bibr">van Zon et al., 1982</xref>; <xref rid="B101" ref-type="bibr">Vinayak et al., 1986</xref>; <xref rid="B22" ref-type="bibr">Desowitz et al., 1989</xref>; <xref rid="B32" ref-type="bibr">Hioki et al., 1990</xref>; <xref rid="B68" ref-type="bibr">Pathak et al., 1990</xref>; <xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>; <xref rid="B47" ref-type="bibr">Marinho et al., 2009</xref>; <xref rid="B50" ref-type="bibr">Megnekou et al., 2009</xref>; <xref rid="B75" ref-type="bibr">Rodrigues-Duarte et al., 2012</xref>; <xref rid="B87" ref-type="bibr">Sharma and Shukla, 2014</xref>), <italic>P. yoelii</italic> (<xref rid="B69" ref-type="bibr">Pavia and Niederbuhl, 1991</xref>), and <italic>P. chabaudi</italic> (<xref rid="B70" ref-type="bibr">Poovassery and Moore, 2006</xref>, <xref rid="B71" ref-type="bibr">2009</xref>; <xref rid="B72" ref-type="bibr">Poovassery et al., 2009</xref>; <xref rid="B84" ref-type="bibr">Sarr et al., 2012</xref>; <xref rid="B86" ref-type="bibr">Sharma et al., 2016</xref>) induce maternal death in a considerable variety of experimental settings. However, only one study has directly compared this outcome induced by different parasites, clearly showing increased mortality rates in pregnant C57BL/6 mice infected with <italic>P. berghei</italic> NK65 compared to K173 and ANKAΔpm4 strains (<xref rid="B75" ref-type="bibr">Rodrigues-Duarte et al., 2012</xref>). One must consider that mortality rates also depend on host susceptibility to infection since, for instance, pregnant A/J mice experience a higher risk of mortality than C57BL/6 mice when infected with <italic>P. chabaudi</italic> AS (<xref rid="B84" ref-type="bibr">Sarr et al., 2012</xref>). These findings support the notion that maternal survival is dependent on both parasite and host intrinsic features. Similarly, mortality was shown to increase in pregnant C57BL/6 and BALB/c mice infected with <italic>P. chabaudi</italic> (<xref rid="B70" ref-type="bibr">Poovassery and Moore, 2006</xref>; <xref rid="B72" ref-type="bibr">Poovassery et al., 2009</xref>) or <italic>P. berghei</italic> (<xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>; <xref rid="B75" ref-type="bibr">Rodrigues-Duarte et al., 2012</xref>), respectively, when compared to their non-pregnant counterparts. This is a well-known phenomenon that also occurs in humans who are primarily in areas of unstable malaria transmission (<xref rid="B45" ref-type="bibr">Luxemburger et al., 1997</xref>; <xref rid="B60" ref-type="bibr">Nosten et al., 2004</xref>). Infection timing was also shown to be linked to maternal mortality, which primarily increases in cases of early gestation infection. This was demonstrated to occur in Wistar rats (<xref rid="B22" ref-type="bibr">Desowitz et al., 1989</xref>), as well as in Swiss Albino, A/J, ICR, and BALB/c mice (<xref rid="B61" ref-type="bibr">Oduola et al., 1982</xref>; <xref rid="B101" ref-type="bibr">Vinayak et al., 1986</xref>; <xref rid="B32" ref-type="bibr">Hioki et al., 1990</xref>) infected with <italic>P. berghei</italic>. However, mid-gestation infection was shown to lead to increased survival with a higher number of animals achieving parturition. Interestingly, it is unclear whether this occurs in humans and if first trimester infections lead to increased maternal mortality rates (<xref rid="B21" ref-type="bibr">Desai et al., 2007</xref>; <xref rid="B77" ref-type="bibr">Rogerson et al., 2018</xref>). Nevertheless, regardless of the infection trimester, mortality is highly dependent on gravidity and is more prevalent in primigravidae than in multiparous women (<xref rid="B53" ref-type="bibr">Menéndez et al., 2008</xref>). Equally, mice tend to experience the same effect, which is likely to be dependent on immunity acquired during previous gestations (<xref rid="B97" ref-type="bibr">van Zon and Eling, 1980</xref>; <xref rid="B47" ref-type="bibr">Marinho et al., 2009</xref>). Accordingly, immunity seems to be pivotal in reduced murine mortality, as ICR mice immunized with attenuated <italic>P. yoelii</italic> before pregnancy had improved survival rates when challenged with the lethal <italic>P. yoelii</italic> 17X strain (<xref rid="B69" ref-type="bibr">Pavia and Niederbuhl, 1991</xref>). Although a direct causality has never been proved, increased mortality observed in murine models of MiP seem to be associated with the development of systemic infection measured by increased peripheral parasitemia that correlates with decreased body weight and patent patterns of anemia (<xref rid="B72" ref-type="bibr">Poovassery et al., 2009</xref>; <xref rid="B84" ref-type="bibr">Sarr et al., 2012</xref>).</p><p>Considering the discussion above, it is clear that mortality during pregnancy depends on a wide range of factors that ultimately will affect host survival. Most of them are commonly recapitulated in rodent experimental models in which experiments have contributed with important findings in this regard. However, one must carefully consider the often-fatal outcomes of murine infections induced by highly lethal parasites that are dissimilar to the infrequent lethality observed in human MiP.</p></sec><sec id="S3.SS2.SSS3"><title><italic>In uterus</italic> Death and Abortion</title><p><italic>In uterus</italic> death and abortion are rather infrequent outcomes of pregnancies complicated by malaria. However, there is a clear association between MiP development and stillbirth incidence, which is noted as being higher in women with perceivable placental parasitemia (<xref rid="B21" ref-type="bibr">Desai et al., 2007</xref>). In contrast, the association between abortion and MiP seems to be rarely discussed, with only scarce reports of this event occurring in low transmission areas, probably resulting from early trimester infections (<xref rid="B51" ref-type="bibr">Menendez, 1995</xref>; <xref rid="B21" ref-type="bibr">Desai et al., 2007</xref>). It is important to highlight that logistical constraints imposed by cultural and socioeconomic boundaries are the reason by which it is difficult to assess this particular pregnancy outcome. Interestingly, the incidence of abortion seems to be much more frequent in murine models than in humans. Accordingly, <italic>P. chabaudi</italic> AS infection of pregnant C57BL/6 mice revealed a rather frequent pattern of abortion and reduced fetal viability at mid-gestation (G10-12) when the parasite was inoculated immediately after conception. This event overlaps with the peripheral parasitemia peak and massive accumulation of parasites inside the placenta (<xref rid="B70" ref-type="bibr">Poovassery and Moore, 2006</xref>). As expected, the parasitemia peak and abortion were frequently observed in mice having increased production of cytokines in both peripheral blood (e.g., IL-1β and IL-10) and placenta (e.g., IFN-γ) (<xref rid="B71" ref-type="bibr">Poovassery and Moore, 2009</xref>). In a follow-up study, it was shown that fetal loss and reduced viability in pregnant C57BL/6 mice infected with <italic>P. chabaudi</italic> AS were markedly influenced by IFN-γ and TNF-α production (<xref rid="B72" ref-type="bibr">Poovassery et al., 2009</xref>) and increased coagulopathy and impaired fibrinolysis (<xref rid="B4" ref-type="bibr">Avery et al., 2012</xref>). In the same extent, abortion was shown to occur concomitantly with apoptosis of inflammatory cells and spongiotrophoblast in the junctional zone (between the uterus and the labyrinth), which has been discussed to occur in a way that is dependent on TNF-α signaling (<xref rid="B83" ref-type="bibr">Sarr et al., 2015</xref>). Altogether, these findings established a clear link between the immune response triggered against malaria and poor pregnancy outcomes. Moreover, models of abortion are also characterized by early infection with <italic>P. chabaudi</italic>, but stillbirth has also been observed in Swiss Albino (<xref rid="B101" ref-type="bibr">Vinayak et al., 1986</xref>; <xref rid="B68" ref-type="bibr">Pathak et al., 1990</xref>), BALB/c (<xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>), and C57BL/6 (<xref rid="B75" ref-type="bibr">Rodrigues-Duarte et al., 2012</xref>) mice infected with <italic>P. berghei</italic> at mid-gestation. In these experimental models, reduced fetal viability without clear abortion observed at late gestation somewhat reflects stillbirth occurring in humans during the third gestational trimester (<xref rid="B21" ref-type="bibr">Desai et al., 2007</xref>).</p><p>Notably, it is important to highlight that translating these findings to human research may raise some serious controversies. Human stillbirth is often described as a dead conceptus that was expelled or removed from the womb 22 weeks after conception, being no longer considered an abortion (less than 22 weeks) (<xref rid="B42" ref-type="bibr">Lawn et al., 2016</xref>). In contrast, in the aforementioned murine models, there is no parallel measure at which fetal viability was assessed that considers gestational age. Vaginal secretions containing blood are normally taken as an indicative of early abortion, followed by the observation of necrotic structures (resorptions) that lack fetal or placental morphology (<xref rid="B70" ref-type="bibr">Poovassery and Moore, 2006</xref>; <xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>). On the other hand, fetal viability is normally evaluated closer to term, after touching <italic>in uterus</italic> or delivered conceptus with pliers. Fetuses that fail to react are considered stillbirths (<xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>). Therefore, definitions in these situations are often misleading, and conclusions should be made with extreme caution since abortion and stillbirth etiology may vary between rodent and human MiP.</p></sec><sec id="S3.SS2.SSS4"><title>Preterm Birth</title><p>Preterm birth, which encompasses every live birth that occurs before the 37th week of gestation (<xref rid="B29" ref-type="bibr">Goldenberg et al., 2009</xref>), is a more frequent consequence and is one of the leading causes of reduced birth weight associated with MiP (<xref rid="B52" ref-type="bibr">Menendez et al., 2000</xref>; <xref rid="B21" ref-type="bibr">Desai et al., 2007</xref>; <xref rid="B54" ref-type="bibr">Moore et al., 2017</xref>). Interestingly, the occurrence of spontaneous delivery in mice or rats in the context of MiP is either poorly addressed or considerably infrequent. One study reported the occurrence of preterm delivery of dead pups from pregnant BALB/c mice infected with <italic>P. berghei</italic> ANKA (<xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>). However, this definition does not seem to fit the circumstance since preterm birth assumes that the progeny is born alive before term, which is unlikely to occur before G19 (reviewed elsewhere <xref rid="B49" ref-type="bibr">McCarthy et al., 2018</xref>). To the same extent, murine models have been questioned regarding their capacity to reproduce human preterm birth due to inaccurate measurements and non-standardized markers.</p><p>In this regard, evaluation of preterm birth in murine models of MiP is probably far from replicating human pathology. However, a better evaluation of this outcome may lead to improved disease models with more reliable translation capacities (<xref rid="B49" ref-type="bibr">McCarthy et al., 2018</xref>).</p></sec><sec id="S3.SS2.SSS5"><title>Progeny Outcomes and Development</title><p>Progeny outcomes and development will ultimately be severely affected as a result of the aforementioned clinical and obstetric manifestations during MiP. Often resulting from IUGR and/or preterm delivery, reduced birth weight contributes significantly to postnatal mortality and impaired child development across the malaria endemicity spectrum and is clearly more frequent in women during their first pregnancies (<xref rid="B60" ref-type="bibr">Nosten et al., 2004</xref>; <xref rid="B21" ref-type="bibr">Desai et al., 2007</xref>). It is unclear how MiP truly leads to growth restriction, yet evidences point to a multitude of factors such as angiogenic imbalance, endocrine dysregulation, deficiencies in transplacental nutrient transportation, severe inflammation and placental insufficiency (<xref rid="B95" ref-type="bibr">Umbers et al., 2011</xref>). During the 1980s, some studies reported the first observations of altered birth weight in mice suffering from MiP, which was found to be significantly diminished in pups from A/J, ICR (<xref rid="B61" ref-type="bibr">Oduola et al., 1982</xref>) and Swiss albino (<xref rid="B101" ref-type="bibr">Vinayak et al., 1986</xref>) litters born from mice infected with <italic>P. berghei</italic> NK65 and NICD, respectively. It seems that impaired progeny development is a rather well-conserved outcome of murine MiP since it was reported in BALB/c (<xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>; <xref rid="B47" ref-type="bibr">Marinho et al., 2009</xref>; <xref rid="B50" ref-type="bibr">Megnekou et al., 2009</xref>; <xref rid="B90" ref-type="bibr">Silver et al., 2010</xref>) and C57BL/6 (<xref rid="B75" ref-type="bibr">Rodrigues-Duarte et al., 2012</xref>; <xref rid="B86" ref-type="bibr">Sharma et al., 2016</xref>) mice infected with a wide variety of parasite strains, suggesting the existence of conserved pathogenesis mechanisms (<xref rid="B75" ref-type="bibr">Rodrigues-Duarte et al., 2012</xref>). This pathologic outcome seems to be avoided with on-time administration of antimalarial drugs such as chloroquine and sulphadoxine-pyrimethamine, which were shown to abrogate systemic parasitemia, therefore improving offspring weight at delivery (<xref rid="B87" ref-type="bibr">Sharma and Shukla, 2014</xref>). Additionally, as it occurs in humans, progeny birth weight improved in mouse multigravidae. This trait was clearly shown to be gravidity-dependent, as pups belonging to the first litter were much smaller on average than those born from subsequent pregnancies when the corresponding dams were induced with MiP by recrudescent <italic>P. berghei</italic> ANKA (<xref rid="B47" ref-type="bibr">Marinho et al., 2009</xref>).</p><p>Further research conducted in the last decade revealed some hidden aspects of reduced birth weight etiology that would not be revealed without the wide range of genetically manipulated mice strains. Strikingly, it seems that proper fetal development is impaired upon the activation of specific components linked to innate immunity, such as complement system receptor (C5aR), Toll-like receptor 4 (TLR4), type I interferon receptor 1 (IFNAR1), and adaptor protein myeloid differentiation factor 88 (MyD88). BALB/c C5aR<sup>–/–</sup> (<xref rid="B17" ref-type="bibr">Conroy et al., 2013</xref>), C57BL/6 TLR4<sup>–/–</sup> (<xref rid="B6" ref-type="bibr">Barboza et al., 2017</xref>; <xref rid="B76" ref-type="bibr">Rodrigues-Duarte et al., 2018</xref>), C57BL/6 IFNAR<sup>–/–</sup> (<xref rid="B76" ref-type="bibr">Rodrigues-Duarte et al., 2018</xref>), and C57BL/6 MyD88<sup>–/–</sup> (<xref rid="B7" ref-type="bibr">Barboza et al., 2014</xref>, <xref rid="B5" ref-type="bibr">2019</xref>) mice were shown to have litters with normal fetal weight, development and survival, even though being infected with <italic>P. berghei</italic> ANKA (in BALB/c) or <italic>P. berghei</italic> NK65 (in C57BL/6). These findings substantially advanced the field by strikingly implying the innate immune system on MiP-associated reduced birth weight pathogenesis, enabling future preclinical trials of anti-inflammatory drugs (e.g., the TLR4 antagonist IAXO) (<xref rid="B6" ref-type="bibr">Barboza et al., 2017</xref>) to be used concomitantly with antimalarial therapies to prevent MiP-associated poor pregnancy outcomes. However, progress in this field should also be done with extreme caution due to possible undesired side effects resulting from adjunctive therapies to treat severe malaria such as those observed in trials conducted with anti-TNF-α therapy to treat cerebral malaria and others (<xref rid="B96" ref-type="bibr">van Hensbroek et al., 1996</xref>; <xref rid="B100" ref-type="bibr">Varo et al., 2018</xref>).</p><p>Moreover, the postnatal scenario of human MiP might rarely include the appearance of congenital malaria, whose onset is tightly controlled by transplacental transmission of maternal antibodies to the fetus, resulting in isolated and scarce events often diagnosed as neonatal sepsis (<xref rid="B73" ref-type="bibr">Rai et al., 2015</xref>; <xref rid="B11" ref-type="bibr">Bhatia et al., 2016</xref>). Conceptus infection might occur due to blood exchanges at delivery or during pregnancy due to loss of villi integrity and syncytium rupture (<xref rid="B18" ref-type="bibr">Crocker et al., 2004</xref>; <xref rid="B74" ref-type="bibr">Robbins and Bakardjiev, 2012</xref>; <xref rid="B91" ref-type="bibr">Souza et al., 2013</xref>). By contrast, the phenomenon is thought to be even rarer in rodents. In related studies, the authors have clearly stated that no congenital malaria was observed, despite an increased fetal spleen weight, which might an indicative of fetal infection (<xref rid="B61" ref-type="bibr">Oduola et al., 1982</xref>; <xref rid="B101" ref-type="bibr">Vinayak et al., 1986</xref>). However, work by <xref rid="B2" ref-type="bibr">Adachi et al. (2000)</xref> addressed this question under the assumption that both mice and rats rarely transmit the parasite to their offspring. Accordingly, the authors detected parasites by nested PCR in a few pups born from pregnant BALB/c mice infected with <italic>P. berghei</italic>. These findings support the rareness of the event due to the lack of a uniform presence of parasites, even in pups within the same litter. The hemotrichorial layer in the murine placenta presents a much thicker barrier against transplacental passage of parasites than the human hemomonochorial placenta (<xref rid="B18" ref-type="bibr">Crocker et al., 2004</xref>), which might explain the reduced incidence of murine congenital malaria. Nevertheless, discrepancies in existing reports contraindicate the usage of murine models in this regard, identifying them as unsuitable for studying this rare disease outcome.</p><p>In conclusion, one may face challenges when translating pregnancy outcome findings obtained in murine models to human pathology. Nevertheless, significant achievements have been made in recent years, despite the considerable diversity of experimental settings and non-standardized methodologies. Improving them will certainly lead to enriched results and more accurate and meaningful conclusions taken from murine models of MiP.</p></sec></sec><sec id="S3.SS3"><title>Understanding Placental Pathology Using Murine Models of MiP</title><p>Most of the aforementioned outcomes occurring in pregnancies complicated by malaria are strongly associated with the dysregulation of placental homeostasis due to a significant accumulation of parasite-infected erythrocytes inside this organ, which overlaps with the onset of PM (<xref rid="B88" ref-type="bibr">Sharma and Shukla, 2017</xref>). Accordingly, several cytological and histological abnormalities are easily observed in infected placentas in addition to parasite accumulation, such as the malarial pigment hemozoin, immune cells, syncytial knots, fibrin deposition, necrosis and placental barrier thickening (<xref rid="B102" ref-type="bibr">Walter et al., 1982</xref>; <xref rid="B34" ref-type="bibr">Ismail et al., 2000</xref>; <xref rid="B91" ref-type="bibr">Souza et al., 2013</xref>). Although despite being PM hallmarks, not all of these histopathologic features are accurately shown by murine models of MiP (schematically represented in <xref ref-type="fig" rid="F3">Figure 3</xref>).</p><sec id="S3.SS3.SSS1"><title>Parasitized Erythrocytes</title><p>Parasitized erythrocytes tend to selectively accumulate in the placenta (<xref ref-type="fig" rid="F1">Figures 1C,D</xref>), achieving greater parasitemia levels than those observed in the peripheral blood of pregnant infected women (<xref rid="B8" ref-type="bibr">Beeson et al., 2002</xref>; <xref rid="B57" ref-type="bibr">Muthusamy et al., 2004</xref>). It is unquestionable that murine parasites also accumulate inside the placenta (<xref ref-type="fig" rid="F2">Figure 2D</xref>) since a large number of studies of placental histopathology noted this finding for a wide range of strains (<xref rid="T1" ref-type="table">Table 1</xref>). Notably, <xref rid="B70" ref-type="bibr">Poovassery and Moore (2006)</xref> have demonstrated that <italic>P. chabaudi</italic> AS parasites also accumulate preferentially inside the placenta of C57BL/6 mice, leading to greater placental parasitemia than that observed in peripheral blood. Interestingly, these murine parasite populations were shown to be mainly constituted by mature forms with an almost null percentage of early ring-stage parasites (<xref rid="B50" ref-type="bibr">Megnekou et al., 2009</xref>), which is in line with observations made during <italic>P. falciparum</italic> infections (<xref rid="B8" ref-type="bibr">Beeson et al., 2002</xref>). Similarly, intravital microscopy studies performed in placentas from pregnant mice infected either with <italic>P. berghei</italic> or <italic>P. chabaudi</italic> also suggested that those that accumulate are indeed the mature forms of the parasite (<xref rid="B43" ref-type="bibr">Lima et al., 2014</xref>), which preferably adhere to the trophoblast in areas of low blood flow inside the maternal blood sinusoids (<xref rid="B19" ref-type="bibr">de Moraes et al., 2013</xref>). As observed for <italic>P. falciparum</italic> regarding its preferential adhesion to CSA and controversial binding to hyaluronic acid (HA) (<xref rid="B9" ref-type="bibr">Beeson and Brown, 2004</xref>; <xref rid="B58" ref-type="bibr">Muthusamy et al., 2007</xref>), the adhesion of <italic>P. berghei</italic> was also shown to be dependent on placental CSA and HA as demonstrated by adhesion assays of mice IEs to placental sections treated with chondroitinase or hyaluronidase (<xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>; <xref rid="B47" ref-type="bibr">Marinho et al., 2009</xref>). These findings once again support similar pathogenesis mechanisms between human and murine PM despite the lack of known PfEMP1 homologs (<xref rid="B33" ref-type="bibr">Hviid et al., 2010</xref>).</p></sec><sec id="S3.SS3.SSS2"><title>The Malarial Pigment Hemozoin</title><p>The malarial pigment hemozoin, a byproduct of hemoglobin catabolism performed by <italic>Plasmodium</italic> spp. to detoxify free heme (<xref rid="B63" ref-type="bibr">Olivier et al., 2014</xref>), is frequently observed in placentas from infected pregnant women (<xref ref-type="fig" rid="F1">Figures 1E,F</xref>) and is often used as an indicator of PM, even in the absence of detectable parasites (<xref rid="B13" ref-type="bibr">Bulmer et al., 1993</xref>; <xref rid="B34" ref-type="bibr">Ismail et al., 2000</xref>). Similarly, placental infection by murine parasites also leads to hemozoin deposition (<xref ref-type="fig" rid="F2">Figure 2F</xref>). Oduola and coworkers first observed this phenomenon in histologic sections of placentas from different murine backgrounds infected with <italic>P. berghei</italic> NK65. Shortly thereafter, the pigment was exclusively observed in maternal blood sinusoids, whose concentrations increased with augmented parasite loads. Accordingly, hemozoin-containing monocytes were also frequently observed (<xref rid="B61" ref-type="bibr">Oduola et al., 1982</xref>, <xref rid="B62" ref-type="bibr">1986</xref>). In addition, hemozoin was also detected in trophoblast giant cells, suggesting an active phagocytic process (<xref rid="B71" ref-type="bibr">Poovassery and Moore, 2009</xref>). The pigment is frequently detected under polarized light microscopy (<xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>; <xref rid="B50" ref-type="bibr">Megnekou et al., 2009</xref>) and is used to indicate past-chronic PM in humans according to the diagnostic criteria of <xref rid="B13" ref-type="bibr">Bulmer et al. (1993)</xref>. To our knowledge, this event was rarely observed in murine models (<xref rid="B62" ref-type="bibr">Oduola et al., 1986</xref>). The fact that hemozoin was rarely observed alone in infected placentas suggests that the experimental settings from current models fail to reproduce past-chronic infections and are only able to reproduce acute PM.</p></sec><sec id="S3.SS3.SSS3"><title>Immune Cell Infiltrate</title><p>Immune cell infiltrate, which occurs as a response to parasite/hemozoin accumulation inside the placenta (<xref ref-type="fig" rid="F1">Figure 1D</xref>), is one of the key hallmarks of the disease and is often associated with poor pregnancy outcomes (<xref rid="B52" ref-type="bibr">Menendez et al., 2000</xref>; <xref rid="B79" ref-type="bibr">Rogerson et al., 2003</xref>; <xref rid="B95" ref-type="bibr">Umbers et al., 2011</xref>). Monocytes/macrophages are the most abundant constituents of these inflammatory infiltrates, and recruitment is widely correlated with placental production of MIP-1α, MCP-1, I-309, and IL-8 chemokines (<xref rid="B1" ref-type="bibr">Abrams et al., 2003</xref>). Not surprisingly, the same event was observed in placentas from mice (<xref ref-type="fig" rid="F2">Figure 2E</xref>) and rats infected with <italic>P. berghei</italic>, especially in those containing detectable placental parasites (<xref rid="B61" ref-type="bibr">Oduola et al., 1982</xref>, <xref rid="B62" ref-type="bibr">1986</xref>; <xref rid="B94" ref-type="bibr">Tegoshi et al., 1992</xref>). Notably, the infiltrate, which was mainly composed of monocytes (CD11b<sup>+ve</sup> cells detected by cytochemistry) and macrophages (CD11b<sup>+ve</sup> cells detected by cytochemistry and <italic>Cd68</italic>- and <italic>Mgl2</italic>-expressing cells identified by qPCR), was linked to the production of some attractant chemokines, such as MIP-1α and MCP-1 (<xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>; <xref rid="B47" ref-type="bibr">Marinho et al., 2009</xref>; <xref rid="B83" ref-type="bibr">Sarr et al., 2015</xref>; <xref rid="B6" ref-type="bibr">Barboza et al., 2017</xref>). However, the event is far from reflecting chronic intervillositis observed in humans (<xref rid="B64" ref-type="bibr">Ordi et al., 1998</xref>), probably due to the shorter gestational period in rodents. Additionally, some authors have commented on the scarcity of this event, especially in placentas from C57BL/6 mice infected with <italic>P. chabaudi</italic>, which tend to have fewer accumulated monocytes/macrophages due to reduced placental parasite burden (<xref rid="B70" ref-type="bibr">Poovassery and Moore, 2006</xref>; <xref rid="B72" ref-type="bibr">Poovassery et al., 2009</xref>). Early reports on different leukocyte populations in infected murine placentas noted the existence of mononuclear (<xref rid="B94" ref-type="bibr">Tegoshi et al., 1992</xref>) and polymorphonuclear (<xref rid="B62" ref-type="bibr">Oduola et al., 1986</xref>) cells. Molecular biology methods were later used to dissect these cell types, dividing them into dendritic cells (<italic>Mgl2</italic>), neutrophils (<italic>Ncf2</italic>), NK cells (<italic>Klrd1</italic>), T (<italic>Cd3e</italic>), and B (<italic>Cd22</italic>) lymphocytes (gene expression quantification by qPCR) (<xref rid="B47" ref-type="bibr">Marinho et al., 2009</xref>; <xref rid="B83" ref-type="bibr">Sarr et al., 2015</xref>; <xref rid="B6" ref-type="bibr">Barboza et al., 2017</xref>). Remarkably, this is in line with observations performed in human PM regarding the placental accumulation of NK cells, T lymphocytes and other non-specified polymorphonuclear cells (<xref rid="B65" ref-type="bibr">Ordi et al., 2001</xref>; <xref rid="B78" ref-type="bibr">Rogerson et al., 2007</xref>; <xref rid="B66" ref-type="bibr">Othoro et al., 2008</xref>). Nevertheless, despite the clear accumulation of immune cells in murine placentas, its association with poor pregnancy outcomes remains to be elucidated in experimental models that oppose current knowledge of human pathology (<xref rid="B79" ref-type="bibr">Rogerson et al., 2003</xref>; <xref rid="B95" ref-type="bibr">Umbers et al., 2011</xref>).</p></sec><sec id="S3.SS3.SSS4"><title>Placental Fibrinoid Necrosis</title><p>Placental fibrinoid necrosis normally occurs as a consequence of extensive placental tissue damage caused during <italic>Plasmodium</italic> spp. infection (<xref ref-type="fig" rid="F1">Figure 1B</xref>). In fact, fibrin deposition is initiated to promote placental tissue repair but soon becomes cytotoxic, leading to necrosis (<xref rid="B102" ref-type="bibr">Walter et al., 1982</xref>; <xref rid="B34" ref-type="bibr">Ismail et al., 2000</xref>) and poor pregnancy outcomes such as premature delivery and reduced birth weight (<xref rid="B52" ref-type="bibr">Menendez et al., 2000</xref>; <xref rid="B4" ref-type="bibr">Avery et al., 2012</xref>). Likewise, pioneering studies from the 1980s revealed the presence of fibrinoid necrosis in placental sections from mice and rats infected with <italic>P. berghei</italic>, although without clear conclusions on its consequences (<xref rid="B62" ref-type="bibr">Oduola et al., 1986</xref>; <xref rid="B94" ref-type="bibr">Tegoshi et al., 1992</xref>). Later, fibrinoid necrosis was reported in placentas from <italic>P. berghei</italic>-infected BALB/c mice (<xref ref-type="fig" rid="F2">Figure 2B</xref>), which delivered litters with a patently reduced birth weight phenotype (<xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>), and fibrin thrombi in placentas from C57BL/6 mice infected with <italic>P. chabaudi</italic> that experienced spontaneous abortion (<xref rid="B72" ref-type="bibr">Poovassery et al., 2009</xref>). Discussion was taken to the point in which fibrin deposition in maternal blood sinusoids would significantly impair placental capability to perform physiological tasks such as respiration and nutrient exchanges, probably due to clotting, blood arrest, necrosis and trophoblast death. In a subsequent study, <xref rid="B4" ref-type="bibr">Avery et al. (2012)</xref> showed by western blot that fibrin deposits were increased in placentas from C57BL/6 mice infected with <italic>P. chabaudi</italic>, which occurred in parallel with an upregulation of coagulation-associated genes. These findings established an important association between impaired fibrinolysis and coagulation and the poor pregnancy outcomes of MiP.</p></sec><sec id="S3.SS3.SSS5"><title>Labyrinth Disarrangement</title><p>Labyrinth disarrangement will ultimately reflect murine placental dysfunction, which is characterized by particular histological alterations that have considerable differences from human PM, mostly due to the existence of a widely different villi structure (<xref ref-type="fig" rid="F1">Figures 1A,B</xref> vs. <xref ref-type="fig" rid="F2">Figures 2A–D</xref>). Of note, two features of human PM that have no counterparts in infected murine placentas are syncytiotrophoblast rupture (previously discussed in the context of congenital malaria) and syncytial knots (<xref rid="B34" ref-type="bibr">Ismail et al., 2000</xref>; <xref rid="B91" ref-type="bibr">Souza et al., 2013</xref>). These protrusions of syncytial nuclear aggregates, which have been associated with hypoxia and oxidative stress in human placentas (<xref rid="B31" ref-type="bibr">Heazell et al., 2007</xref>), were discussed as having no similar structure in murine placentas that could be detected under light or electron microscopy (<xref rid="B94" ref-type="bibr">Tegoshi et al., 1992</xref>). Syncytial knots, which are considered an accumulation of degrading nuclei, was once erroneously described as a phenomenon of trophoblast hyperplasia [discussed elsewhere (<xref rid="B31" ref-type="bibr">Heazell et al., 2007</xref>)]. To our knowledge, the latter was never clearly discussed in the context of human PM. Accordingly, there are unclear reports of trophoblast hyperplasia occurring in the placentas of mice (<xref rid="B62" ref-type="bibr">Oduola et al., 1986</xref>; <xref rid="B101" ref-type="bibr">Vinayak et al., 1986</xref>; <xref rid="B68" ref-type="bibr">Pathak et al., 1990</xref>; <xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>) and rats (<xref rid="B62" ref-type="bibr">Oduola et al., 1986</xref>; <xref rid="B94" ref-type="bibr">Tegoshi et al., 1992</xref>) infected with <italic>P. berghei</italic>. Nevertheless, evidence of this event includes unclear microscopy images that fail to address the apparent enlargement/swelling of tissue that occurs due to cell proliferation. However, there is a striking thickening of the trophoblast basal membrane that partially overlaps with the concept of tissue swelling. In human PM, this frequent phenomenon can be qualitatively analyzed (<xref rid="B34" ref-type="bibr">Ismail et al., 2000</xref>) or more accurately quantified as the distance that separates fetal capillaries from villi outer membrane (<xref rid="B91" ref-type="bibr">Souza et al., 2013</xref>). Regardless of the methodology used, this parameter was found to be significantly thicker during human PM and, being discussed as strongly influencing transplacental transport of vital compounds. Similarly, this was qualitatively analyzed and reported in mouse and rat placentas infected with <italic>P. berghei</italic>, which was hypothesized to be a consequence of the fibrotic process resulting from massive tissue repair (<xref rid="B62" ref-type="bibr">Oduola et al., 1986</xref>; <xref rid="B94" ref-type="bibr">Tegoshi et al., 1992</xref>). Later, some other works have developed methods to indirectly quantify trophoblast thickening, taking advantage of the sinusoidal nature of mouse placentas. Accordingly, maternal blood areas were quantified, and the reduction of vascular spaces in placentas from BALB/c mice infected with <italic>P. berghei</italic> ANKA was considered a proxy for basal membrane thickening (<xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>). This supports the conclusion that reduced maternal blood spaces due to trophoblast membrane thickening and maternal sinusoidal remodeling (<xref rid="B19" ref-type="bibr">de Moraes et al., 2013</xref>) would ultimately lead to placental insufficiency and impaired transplacental transport of nutrients [discussed elsewhere (<xref rid="B59" ref-type="bibr">Neres et al., 2008</xref>; <xref rid="B19" ref-type="bibr">de Moraes et al., 2013</xref>)]. The same methodology was further used to determine that distinct parasites inflict different magnitudes of circulatory impairment and membrane thickening in placentas from C57BL/6 mice (<xref rid="B75" ref-type="bibr">Rodrigues-Duarte et al., 2012</xref>). The etiology of the event was further addressed in <italic>P. berghei</italic> NK65-infected C57BL/6 TLR4<sup>–/–</sup> and MyD88<sup>–/–</sup> KO mice, which had blood sinusoidal areas similar to those observed in non-infected pregnant mice (<xref rid="B7" ref-type="bibr">Barboza et al., 2014</xref>, <xref rid="B6" ref-type="bibr">2017</xref>). These findings established a logical link between host innate immunity and placental pathology, once again supporting the notion that outcomes of MiP mostly result from damage and homeostatic dysregulation inflicted mostly by factors of an immunologic nature.</p></sec></sec></sec><sec id="S4"><title>Conclusion</title><p>Clearly, the development of murine models that recapitulate traits from human MiP has definitively contributed to the current understanding of this disease. However, there are still few reports that truly reveal some of the pathogenesis mechanisms of MiP. Only in the last decade have some studies clearly unraveled some hidden molecular mechanisms of MiP, such as innate immunity activation and its contributions to poor pregnancy outcomes (<xref rid="B72" ref-type="bibr">Poovassery et al., 2009</xref>; <xref rid="B17" ref-type="bibr">Conroy et al., 2013</xref>; <xref rid="B7" ref-type="bibr">Barboza et al., 2014</xref>, <xref rid="B6" ref-type="bibr">2017</xref>, <xref rid="B5" ref-type="bibr">2019</xref>; <xref rid="B76" ref-type="bibr">Rodrigues-Duarte et al., 2018</xref>). Indeed, the advent of genetic engineering and the capacity to generate a wide range of knockout mice were definite turning points from which we are still benefiting. Until this point, most studies have tried to establish proper experimental settings that would ultimately validate murine models as suitable for studying MiP. First, striking similarities are observed regarding evolutionary and developmental traits between murine and human placentas, from which one might conclude that physiologically, both would behave in a particularly similar way (<xref rid="B27" ref-type="bibr">Georgiades et al., 2002</xref>; <xref rid="B103" ref-type="bibr">Wildman et al., 2006</xref>). Additionally, murine parasites exhibit a group of characteristics that support the conclusion that murine pathology would somewhat resemble human MiP, despite the well-known differences between rodent <italic>Plasmodium</italic> species and <italic>P. falciparum</italic> biology (<xref rid="B40" ref-type="bibr">Lamb et al., 2006</xref>; <xref rid="B33" ref-type="bibr">Hviid et al., 2010</xref>). Nevertheless, one might consider the patent limitations of conclusions regarding disease outcomes, which can later limit the translational capacities of observed results. As such, refinement of experimental design and standardization of methodology is necessary for the improvement of such models (<xref ref-type="boxed-text" rid="BX1">BOX 1</xref>). Resolving these gaps will certainly enrich research in the field, possibly reaffirming the usage of murine models to address more specific and complex questions implicit in drug preclinical trials and vaccine development (<xref rid="B23" ref-type="bibr">Doritchamou et al., 2017</xref>). Research in the field has indeed overcome several milestones due to the usage of rodent models with much progress that is still to come.</p><boxed-text id="BX1" position="float"><caption><title>Box 1. Suggestions to improve future murine models of MiP.</title></caption><p>Evolution has grouped murine animals and humans in similar phylogenetic clades according to some striking similarities. However, it has also separated them especially from an anatomically perspective. Some differences cannot be surpassed; yet, our understanding of some diseases and complex biological processes can be improved with the refinement of experimental design and standardization of analysis methods when using animal models such as experimental murine models of MiP. As such, our understanding of MiP could be improved by addressing some of the following points:</p><list list-type="simple" id="S3.I1"><list-item><label>•</label><p>Understanding the adhesion mechanism of murine parasite strains.</p></list-item><list-item><label>•</label><p>Identification of VAR2CSA functional homologs in murine parasites.</p></list-item><list-item><label>•</label><p>Standardization of protocols using specific species of murine parasites to differently address specific aspects of uncomplicated or severe MiP (e.g., usage of less or more virulent parasite strains, tropism for reticulocytes or mature erythrocytes, preferential tissue for accumulation).</p></list-item><list-item><label>•</label><p>Standardized definitions and accurate measures for abortion, preterm delivery, stillbirth and placental malaria in the context of murine MiP.</p></list-item><list-item><label>•</label><p>Investigating the possibility of congenital malaria in murine models of MiP.</p></list-item></list></boxed-text></sec><sec id="S6"><title>Author Contributions</title><p>AB drafted the manuscript, compiled information from the literature, and designed the figures and tables. MP drafted the manuscript and gathered information from the literature. SE supervised and reviewed the manuscript. CM supervised and reviewed the manuscript and designed the figures and tables.</p></sec><sec id="conf1"><title>Conflict of Interest Statement</title><p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p></sec></body><back><fn-group><fn fn-type="financial-disclosure"><p><bold>Funding.</bold> This work was primarily funded by the São Paulo Research Foundation – FAPESP (Grant Nos. 2018/20468-0 to CM and 2017/05782-8 to SE). AB and MP were supported by fellowships from the FAPESP (2017/03939-7 and 2012/10081-5, respectively).</p></fn></fn-group><ref-list><title>References</title><ref id="B1"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Abrams</surname><given-names>E. 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