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_:b420373414 , _:b420373415 , _:b420373464 , _:b420373465 , _:b420373466 , _:b420373467 , _:b420373468 , _:b420373469 , _:b420373470 , _:b420373471 , _:b420373456 , _:b420373457 , _:b420373458 , _:b420373459 , _:b420373460 , _:b420373461 , _:b420373462 , _:b420373463 , _:b420373448 , _:b420373449 , _:b420373450 , _:b420373451 , _:b420373452 , _:b420373453 , _:b420373454 , _:b420373455 , _:b420373440 , _:b420373441 , _:b420373442 , _:b420373443 , _:b420373444 , _:b420373445 , _:b420373446 , _:b420373447 , _:b420373480 , _:b420373481 , _:b420373482 , _:b420373483 , _:b420373484 , _:b420373485 , _:b420373472 , _:b420373473 , _:b420373474 , _:b420373475 , _:b420373476 , _:b420373477 , _:b420373478 , _:b420373479 ; ns1:pmcid "PMC0" ; bibo:doi "10.1084%2Fjem.20091268" . @prefix ns4: . ns4:contains _:b6576623 , _:b6576609 . _:b6576609 rdf:type ns4:Section . @prefix dc: . _:b6576609 dc:title "materials and methods" ; ns4:contains _:b6576610 , _:b6576611 , _:b6576612 , _:b6576613 , _:b6576614 , _:b6576615 , _:b6576616 , _:b6576617 , _:b6576618 , _:b6576619 , _:b6576620 , _:b6576621 , _:b6576622 . _:b6576610 rdf:type ns1:Context ; rdf:value "RELM-\u03B2\u2212 mice (Hogan et al., >>2006<<) were bred onto a C57BL/6 background for nine generations before use in our studies." ; ns1:mentions . _:b6576611 rdf:type ns1:Context ; rdf:value "Hybridomas that secrete neutralizing rat IgG1 anti\u2013mouse IFN-\u03B3 mAb (XMG-6; Mosmann and Fong, >>1989<<) and an isotype-matched control mAb (GL113) were obtained originally from DNAX." ; ns1:mentions . _:b6576612 rdf:type ns1:Context ; rdf:value "The hybridoma that secretes GK1.5 (rat IgG2b anti-CD4; Miller and Jenkins, >>1985<<) was obtained from the American Type Culture Collection." ; ns1:mentions . _:b6576613 rdf:type ns1:Context ; rdf:value "L-4/anti\u2013IL-4 mAb complexes (IL-4C) were prepared by mixing rmIL-4 with the rat IgG1 anti-IL-4 mAb 11B11 (Finkelman et al., >>1993<<; Shea-Donohue et al., 2001) at a 2:1 molar (1:5 weight) ratio (Urban et al., 1995)." ; ns1:mentions . _:b6576614 rdf:type ns1:Context ; rdf:value "L-4/anti\u2013IL-4 mAb complexes (IL-4C) were prepared by mixing rmIL-4 with the rat IgG1 anti-IL-4 mAb 11B11 (Finkelman et al., 1993; Shea-Donohue et al., >>2001<<) at a 2:1 molar (1:5 weight) ratio (Urban et al., 1995). Mice were injected i.p with IL-4C containing 5 \u00B5g of IL-4 and 25 \u00B5g of 11B11." ; ns1:mentions . _:b6576615 rdf:type ns1:Context ; rdf:value "L-4/anti\u2013IL-4 mAb complexes (IL-4C) were prepared by mixing rmIL-4 with the rat IgG1 anti-IL-4 mAb 11B11 (Finkelman et al., 1993; Shea-Donohue et al., 2001) at a 2:1 molar (1:5 weight) ratio (Urban et al., >>1995<<). Mice were injected i.p with IL-4C containing 5 \u00B5g of IL-4 and 25 \u00B5g of 11B11. IL-4C slowly dissociates, releasing biologically active IL-4 for 3\u20135 d." ; ns1:mentions . _:b6576616 rdf:type ns1:Context ; rdf:value "Relative rates of in vivo IL-4 and IFN-\u03B3 secretion were determined by the in vivo cytokine capture assay (IVCCA; Finkelman and Morris, >>1999<<). Injected biotin-labeled anti-cytokine mAbs in this assay form complexes with the secreted cytokines they specifically bind that have a much longer in vivo half-life than free cytokines." ; ns1:mentions . _:b6576617 rdf:type ns1:Context ; rdf:value "and the bound complex was detected with biotin-labeled affinity-purified goat anti\u2013IL-13R\u03B12 antibody followed by HRP-SA and luminogenic substrate (Khodoun et al., >>2007<<)." ; ns1:mentions . _:b6576618 rdf:type ns1:Context ; rdf:value "Relative expression was calculated with the 1/\u0394\u0394ct method using vimentin as the housekeeping gene (Livak and Schmittgen, >>2001<<). Primer sequences used were the following: vimentin, forward 5\u2032-TGACCGGCTTGTATGCTATC-3\u2032 and reverse 5\u2032-CAGTGTGAGCCAGGATATAG-3\u2032; RELM-\u03B1, forward 5\u2032-AGATGGGCCTCCTGCCCTGCTGGG-3\u2032 and reverse 5\u2032-ACCTGGTGACGGGCGACGACGGTT-3\u2032; RELM-\u03B2, forward" ; ns1:mentions . _:b6576619 rdf:type ns1:Context ; rdf:value "Maintenance and recovery of mouse-adapted N. brasiliensis has been described previously (Urban et al., >>2001<<). T. spiralis L1 were recovered by 1% pepsin/1% HCl digests of muscle from mice infected with this parasite for 90\u2013120 d. Naive mice were inoculated by oral gavage with 200 L3 in 0.2 ml PBS/0.2% agarose (BD). H. polygyrus\u2013infective" ; ns1:mentions . _:b6576620 rdf:type ns1:Context ; rdf:value "H. polygyrus\u2013infective ensheathed infective\u2013stage larvae (L3) were propagated and stored at 4\u00B0C until use (Urban et al., >>1995<<). For primary inoculation, mice received 200 L3 by oral gavage. For secondary inoculation, mice were first treated with the anthelminthic pyrantel pamoate 14 d after primary inoculation, allowed to rest for 28 d, and then rechallenged by" ; ns1:mentions . _:b6576621 rdf:type ns1:Context ; rdf:value "with rhodamine B (1 mg/ml in saline) 1 h before sacrifice (Bansemir and Sukhdeo, >>2001<<). Worms were photographed with a fluorescence microscope (Nikon) using an SPT Diagnostics imaging system and Simple PCI C-Imaging systems software." ; ns1:mentions . _:b6576622 rdf:type ns1:Context ; rdf:value "The ATPLite kit (Perkin Elmer) was used to measure the ATP content of individual H. polygyrus adult worms (Ishiwata and Watanabe, >>2007<<)." ; ns1:mentions . _:b6576623 rdf:type ns4:Section ; dc:title "results and discussion" ; ns4:contains _:b6576632 , _:b6576633 , _:b6576634 , _:b6576635 , _:b6576636 , _:b6576637 , _:b6576638 , _:b6576639 , _:b6576624 , _:b6576625 , _:b6576626 , _:b6576627 , _:b6576628 , _:b6576629 , _:b6576630 , _:b6576631 , _:b6576656 , _:b6576657 , _:b6576658 , _:b6576659 , _:b6576648 , _:b6576649 , _:b6576650 , _:b6576651 , _:b6576652 , _:b6576653 , _:b6576654 , _:b6576655 , _:b6576640 , _:b6576641 , _:b6576642 , _:b6576643 , _:b6576644 , _:b6576645 , _:b6576646 , _:b6576647 . _:b6576624 rdf:type ns1:Context ; rdf:value "Results that IECs constitutively express IL-4R\u03B1 (Reinecker and Podolsky, >>1995<<) and can promote Th2 immunity (Zaph et al., 2007) prompted us to investigate whether IEC activation by IL-4/IL-13 promotes GI nematode expulsion." ; ns1:mentions . _:b6576625 rdf:type ns1:Context ; rdf:value "Results that IECs constitutively express IL-4R\u03B1 (Reinecker and Podolsky, 1995) and can promote Th2 immunity (Zaph et al., >>2007<<) prompted us to investigate whether IEC activation by IL-4/IL-13 promotes GI nematode expulsion." ; ns1:mentions . _:b6576626 rdf:type ns1:Context ; rdf:value "Thus, IL-4/IL-13 effects on IEC account for most, but not all, of the defect in worm expulsion previously observed in mice deficient in both IL-4 and IL-13 (Fallon et al., >>2002<<). Villin-Cre expression by IL-4R\u03B1flox/\u2212 mice did not affect IL-4 or IFN-\u03B3 production (Fig. 1 C) but was associated with increased IL-13 (Fig. 1 D), probably as a result of increased worm burden. Despite increased IL-13 levels in" ; ns1:mentions . _:b6576627 rdf:type ns1:Context ; rdf:value "1 F), a goblet cell\u2013produced molecule that is normally up-regulated by worm infection (Artis et al., >>2004<<). RELM-\u03B2 expression increased greatly in the small intestine of N. brasiliensis\u2013inoculated mice after larvae migrated to that organ. In contrast, pulmonary RELM-\u03B2 expression increased only marginally in these mice (Fig. 2 A). Conversely," ; ns1:mentions . _:b6576628 rdf:type ns1:Context ; rdf:value "Because RELM-\u03B2 binds to chemosensory organs of the intestinal worms Trichuris muris and Strongyloides stercoralis and inhibits S. stercoralis chemotaxis in vitro (Artis et al., >>2004<<), and because intact chemotaxis might be required for gut lumen-dwelling worms to locate and feed on host epithelium, we tested whether expulsion of worms that reside in the intestinal lumen is defective in RELM-\u03B2\u2212/\u2212 mice." ; ns1:mentions . _:b6576629 rdf:type ns1:Context ; rdf:value "To evaluate whether RELM-\u03B2 also promotes expulsion of worms that normally establish a chronic infection of the gut lumen, WT and RELM-\u03B2\u2212/\u2212 were inoculated with H. polygyrus (Urban et al., >>1991<<). In contrast to N. brasilensis, H. polygyrus has two intestinal niches." ; ns1:mentions . _:b6576630 rdf:type ns1:Context ; rdf:value "Larvae encyst in the intestinal muscularis (days 0\u20138) and emerge afterward as adults that reside only in the intestinal lumen, where they survive >90 d during primary infection in most mouse strains (Su and Dobson, >>1997<<). During a second infection, however, IL-4R\u03B1\u2013mediated signaling promotes destruction of some encysted H. polygyrus larvae by inducing macrophage production of arginase (Anthony et al., 2006) and then induces expulsion from the gut lumen" ; ns1:mentions . _:b6576631 rdf:type ns1:Context ; rdf:value "During a second infection, however, IL-4R\u03B1\u2013mediated signaling promotes destruction of some encysted H. polygyrus larvae by inducing macrophage production of arginase (Anthony et al., >>2006<<) and then induces expulsion from the gut lumen of the remaining worms that survive beyond day 7 (Anthony et al., 2006)." ; ns1:mentions . _:b6576632 rdf:type ns1:Context ; rdf:value "destruction of some encysted H. polygyrus larvae by inducing macrophage production of arginase (Anthony et al., 2006) and then induces expulsion from the gut lumen of the remaining worms that survive beyond day 7 (Anthony et al., >>2006<<). As expected, RELM-\u03B2 deficiency neither increases H. polygyrus survival during the initial 7 d of a second infection, when encysted larvae are surrounded by arginase-expressing cells (Fig." ; ns1:mentions . _:b6576633 rdf:type ns1:Context ; rdf:value "Although immunity against N. brasiliensis and H. polygyrus is CD4+ T cell dependent (Voehringer et al., >>2006<<), the CD4+ T cell requirement is restricted to IL-4/IL-13 production, inasmuch as treatment with a long-acting form of IL-4 (IL-4C) induces worm expulsion by CD4+ T cell\u2013depleted mice (Urban et al., 1995)." ; ns1:mentions . _:b6576634 rdf:type ns1:Context ; rdf:value "dependent (Voehringer et al., 2006), the CD4+ T cell requirement is restricted to IL-4/IL-13 production, inasmuch as treatment with a long-acting form of IL-4 (IL-4C) induces worm expulsion by CD4+ T cell\u2013depleted mice (Urban et al., >>1995<<). IL-4C\u2013induced T cell\u2013independent worm expulsion is RELM-\u03B2 dependent; IL-4C induced N. brasiliensis expulsion in anti-CD4 mAb-treated WT but not RELM-\u03B2\u2212/\u2212 (Fig." ; ns1:mentions . _:b6576635 rdf:type ns1:Context ; rdf:value "RELM-\u03B2, which acts on mouse macrophages to promote IFN-\u03B3\u2013driven intestinal inflammation (Nair et al., >>2008<<) in addition to its direct effect on worm chemotaxis, might expel worms through effects on the parasite or the host." ; ns1:mentions . _:b6576636 rdf:type ns1:Context ; rdf:value "between these possibilities, H. polygyrus adult worms were isolated from chronically infected mice, exposed to PBS, rRELM-\u03B2, or rRELM-\u03B1, a closely related molecule which limits Th2 responses in worm-infected mice (Nair et al., >>2009<<; Pesce et al., 2009), washed, reinoculated into naive WT and IL-4R\u03B1\u2013deficient hosts, and evaluated for survival and fecundity." ; ns1:mentions . _:b6576637 rdf:type ns1:Context ; rdf:value "possibilities, H. polygyrus adult worms were isolated from chronically infected mice, exposed to PBS, rRELM-\u03B2, or rRELM-\u03B1, a closely related molecule which limits Th2 responses in worm-infected mice (Nair et al., 2009; Pesce et al., >>2009<<), washed, reinoculated into naive WT and IL-4R\u03B1\u2013deficient hosts, and evaluated for survival and fecundity." ; ns1:mentions . _:b6576638 rdf:type ns1:Context ; rdf:value "In contrast, in vitro treatment of worms with IL-4, RELM-\u03B1 (which differs from RELM-\u03B2 by absence of a cysteine residue; Patel et al., >>2004<<), or heat-denatured RELM-\u03B2 had no detectable effect (Fig." ; ns1:mentions . _:b6576639 rdf:type ns1:Context ; rdf:value "Because rRELM-\u03B2 interferes with chemotaxis in vitro (Artis et al., >>2004<<), it might promote expulsion of adult worms by blocking their ability to sense their source of nutrition." ; ns1:mentions . _:b6576640 rdf:type ns1:Context ; rdf:value "Furthermore, RELM-\u03B2 pretreatment of H. polygyrus adults inhibited their in vivo ingestion of rhodamine B\u2013labeled host IEC (Fig. 5 D; Bansemir and Sukhdeo, >>2001<<)." ; ns1:mentions . _:b6576641 rdf:type ns1:Context ; rdf:value "worms by inhibiting their ability to locate their food source, it should not affect worm parasites that live entirely or partially within host intestinal epithelium, such as Trichinella spiralis or Trichuris muris (Gagliardo et al., >>2002<<). These worms would not need to locate IEC and would likely be at least partially protected from RELM-\u03B2 secreted into the gut lumen." ; ns1:mentions . _:b6576642 rdf:type ns1:Context ; rdf:value "Indeed, RELM-\u03B2\u2212/\u2212 has recently been shown to have normal protective immunity against T. muris (Nair et al., >>2008<<) and we find no defect in expulsion of T. spiralis in RELM-\u03B2\u2212/\u2212 (Fig." ; ns1:mentions . _:b6576643 rdf:type ns1:Context ; rdf:value "S4), although protective immunity against both worms is IL-4R\u03B1 dependent (Else et al., >>1994<<; Urban et al., 2001)." ; ns1:mentions . _:b6576644 rdf:type ns1:Context ; rdf:value "S4), although protective immunity against both worms is IL-4R\u03B1 dependent (Else et al., 1994; Urban et al., >>2001<<). Combined, these observations provide evidence that IL-4/IL-13 induction of IEC RELM-\u03B2 production is selectively important for expulsion of worms that live in the intestinal lumen." ; ns1:mentions . _:b6576645 rdf:type ns1:Context ; rdf:value "The role of IEC in host protection is complex and includes a critical inducer function (production of the Th2-promoting cytokine TSLP in mice infected with Trichuris muris; Zaph et al., >>2007<<) as well as the effector function shown here." ; ns1:mentions . _:b6576646 rdf:type ns1:Context ; rdf:value "Such a mechanism most likely works in concert with other effects of IL-4R signaling, including increased smooth muscle responsiveness (Zhao et al., >>2003<<, 2008), establishment of a mucus barrier (Kuperman et al., 2002; Ramalingam et al., 2008), CD8+ T cell proliferation (Morris et al., 2009), chemokine secretion (Fulkerson et al., 2004), and adhesion molecule expression (Hickey et al.," ; ns1:mentions . _:b6576647 rdf:type ns1:Context ; rdf:value "Such a mechanism most likely works in concert with other effects of IL-4R signaling, including increased smooth muscle responsiveness (Zhao et al., 2003, >>2008<<), establishment of a mucus barrier (Kuperman et al., 2002; Ramalingam et al., 2008), CD8+ T cell proliferation (Morris et al., 2009), chemokine secretion (Fulkerson et al., 2004), and adhesion molecule expression (Hickey et al., 1999)," ; ns1:mentions . _:b6576648 rdf:type ns1:Context ; rdf:value "Such a mechanism most likely works in concert with other effects of IL-4R signaling, including increased smooth muscle responsiveness (Zhao et al., 2003, 2008), establishment of a mucus barrier (Kuperman et al., >>2002<<; Ramalingam et al., 2008), CD8+ T cell proliferation (Morris et al., 2009), chemokine secretion (Fulkerson et al., 2004), and adhesion molecule expression (Hickey et al., 1999), that together protect hosts against different worm parasites." ; ns1:mentions . _:b6576649 rdf:type ns1:Context ; rdf:value "a mechanism most likely works in concert with other effects of IL-4R signaling, including increased smooth muscle responsiveness (Zhao et al., 2003, 2008), establishment of a mucus barrier (Kuperman et al., 2002; Ramalingam et al., >>2008<<), CD8+ T cell proliferation (Morris et al., 2009), chemokine secretion (Fulkerson et al., 2004), and adhesion molecule expression (Hickey et al., 1999), that together protect hosts against different worm parasites." ; ns1:mentions . _:b6576650 rdf:type ns1:Context ; rdf:value "other effects of IL-4R signaling, including increased smooth muscle responsiveness (Zhao et al., 2003, 2008), establishment of a mucus barrier (Kuperman et al., 2002; Ramalingam et al., 2008), CD8+ T cell proliferation (Morris et al., >>2009<<), chemokine secretion (Fulkerson et al., 2004), and adhesion molecule expression (Hickey et al., 1999), that together protect hosts against different worm parasites." ; ns1:mentions . _:b6576651 rdf:type ns1:Context ; rdf:value "smooth muscle responsiveness (Zhao et al., 2003, 2008), establishment of a mucus barrier (Kuperman et al., 2002; Ramalingam et al., 2008), CD8+ T cell proliferation (Morris et al., 2009), chemokine secretion (Fulkerson et al., >>2004<<), and adhesion molecule expression (Hickey et al., 1999), that together protect hosts against different worm parasites." ; ns1:mentions . _:b6576652 rdf:type ns1:Context ; rdf:value "2008), establishment of a mucus barrier (Kuperman et al., 2002; Ramalingam et al., 2008), CD8+ T cell proliferation (Morris et al., 2009), chemokine secretion (Fulkerson et al., 2004), and adhesion molecule expression (Hickey et al., >>1999<<), that together protect hosts against different worm parasites." ; ns1:mentions . _:b6576653 rdf:type ns1:Context ; rdf:value "Taken in the context of previous observations, our studies demonstrate the remarkable versatility of functions mediated by IL-4R signaling in host protection against nematode parasites (Finkelman et al., >>1997<<). Signaling through this receptor appears to protect against T. spiralis by stimulating isotype switching to IgE, promoting mast cell development, and increasing sensitivity to mast cell\u2013produced mediators (Strait et al., 2003), to" ; ns1:mentions . _:b6576654 rdf:type ns1:Context ; rdf:value "Signaling through this receptor appears to protect against T. spiralis by stimulating isotype switching to IgE, promoting mast cell development, and increasing sensitivity to mast cell\u2013produced mediators (Strait et al., >>2003<<), to protect against lethal inflammation caused by Schistosoma mansoni by causing macrophages to produce arginase (Herbert et al., 2004), to protect against N. brasiliensis by inducing IEC to differentiate into goblet cells that produce" ; ns1:mentions . _:b6576655 rdf:type ns1:Context ; rdf:value "mast cell development, and increasing sensitivity to mast cell\u2013produced mediators (Strait et al., 2003), to protect against lethal inflammation caused by Schistosoma mansoni by causing macrophages to produce arginase (Herbert et al., >>2004<<), to protect against N. brasiliensis by inducing IEC to differentiate into goblet cells that produce RELM-\u03B2, to protect against H. polygyrus first through an arginase-dependent mechanism directed against larvae encysted in the gut wall" ; ns1:mentions . _:b6576656 rdf:type ns1:Context ; rdf:value "N. brasiliensis by inducing IEC to differentiate into goblet cells that produce RELM-\u03B2, to protect against H. polygyrus first through an arginase-dependent mechanism directed against larvae encysted in the gut wall (Anthony et al., >>2006<<) and then through a RELM-\u03B2\u2013dependent mechanism directed against luminal adults, and to protect against Trichuris muris by promoting the proliferation and shedding of the IEC in which these worms embed (Cliffe et al., 2005)." ; ns1:mentions . _:b6576657 rdf:type ns1:Context ; rdf:value "et al., 2006) and then through a RELM-\u03B2\u2013dependent mechanism directed against luminal adults, and to protect against Trichuris muris by promoting the proliferation and shedding of the IEC in which these worms embed (Cliffe et al., >>2005<<). Further still, these mechanisms are distinct from IL-4R\u03B1\u2013mediated RELM-\u03B1 production that down-modulates worm-driven lung inflammation (Nair et al., 2009;Pesce et al., 2009)." ; ns1:mentions . _:b6576658 rdf:type ns1:Context ; rdf:value "Further still, these mechanisms are distinct from IL-4R\u03B1\u2013mediated RELM-\u03B1 production that down-modulates worm-driven lung inflammation (Nair et al., >>2009<<;Pesce et al., 2009)." ; ns1:mentions . _:b6576659 rdf:type ns1:Context ; rdf:value "Further still, these mechanisms are distinct from IL-4R\u03B1\u2013mediated RELM-\u03B1 production that down-modulates worm-driven lung inflammation (Nair et al., 2009;Pesce et al., >>2009<<)." ; ns1:mentions . _:b420372994 rdf:type ns1:RelevantBibliographicResource . @prefix xsd: . _:b420372994 ns1:RelevantScore "21"^^xsd:nonNegativeInteger ; ns1:hasRelevantPaperId . _:b420372995 rdf:type ns1:RelevantBibliographicResource ; ns1:RelevantScore "20"^^xsd:nonNegativeInteger ; ns1:hasRelevantPaperId . _:b420372996 rdf:type ns1:RelevantBibliographicResource ; 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