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, _:b66507468 . _:b66507411 rdf:type ns4:Section . @prefix dc: . _:b66507411 dc:title "introduction" ; ns4:contains _:b66507432 , _:b66507433 , _:b66507434 , _:b66507435 , _:b66507436 , _:b66507437 , _:b66507438 , _:b66507439 , _:b66507424 , _:b66507425 , _:b66507426 , _:b66507427 , _:b66507428 , _:b66507429 , _:b66507430 , _:b66507431 , _:b66507448 , _:b66507449 , _:b66507450 , _:b66507451 , _:b66507452 , _:b66507453 , _:b66507454 , _:b66507455 , _:b66507440 , _:b66507441 , _:b66507442 , _:b66507443 , _:b66507444 , _:b66507445 , _:b66507446 , _:b66507447 , _:b66507416 , _:b66507417 , _:b66507418 , _:b66507419 , _:b66507420 , _:b66507421 , _:b66507422 , _:b66507423 , _:b66507412 , _:b66507413 , _:b66507414 , _:b66507415 , _:b66507464 , _:b66507465 , _:b66507466 , _:b66507467 , _:b66507456 , _:b66507457 , _:b66507458 , _:b66507459 , _:b66507460 , _:b66507461 , _:b66507462 , _:b66507463 . _:b66507412 rdf:type ns1:Context ; rdf:value "is essential for the localized activation of signaling pathways that coordinate many aspects of embryonic development, including asymmetric cell division, epithelial organization,\u00A0and embryo axis establishment (Goldstein and Macara, >>2007<<, St Johnston and Ahringer, 2010)." ; ns1:mentions . _:b66507413 rdf:type ns1:Context ; rdf:value "activation of signaling pathways that coordinate many aspects of embryonic development, including asymmetric cell division, epithelial organization,\u00A0and embryo axis establishment (Goldstein and Macara, 2007, St Johnston and Ahringer, >>2010<<). Although the precise details vary between\u00A0systems, in most cases the conserved PDZ domain proteins PAR-3 and PAR-6, the small guanosine triphosphatase (GTPase) CDC-42 and atypical protein kinase C (aPKC) act together to establish" ; ns1:mentions . _:b66507414 rdf:type ns1:Context ; rdf:value "and PAR-6, the small guanosine triphosphatase (GTPase) CDC-42 and atypical protein kinase C (aPKC) act together to establish polarity on one side of the cell and drive asymmetry of a range of downstream pathways (reviewed in Goehring, >>2014<<, McCaffrey and Macara, 2012, Suzuki et\u00A0al., 2004, Ziomek et\u00A0al., 1982)." ; ns1:mentions . _:b66507415 rdf:type ns1:Context ; rdf:value "CDC-42 and atypical protein kinase C (aPKC) act together to establish polarity on one side of the cell and drive asymmetry of a range of downstream pathways (reviewed in Goehring, 2014, McCaffrey and Macara, 2012, Suzuki et\u00A0al., >>2004<<, Ziomek et\u00A0al., 1982)." ; ns1:mentions . _:b66507416 rdf:type ns1:Context ; rdf:value "protein kinase C (aPKC) act together to establish polarity on one side of the cell and drive asymmetry of a range of downstream pathways (reviewed in Goehring, 2014, McCaffrey and Macara, 2012, Suzuki et\u00A0al., 2004, Ziomek et\u00A0al., >>1982<<)." ; ns1:mentions . _:b66507417 rdf:type ns1:Context ; rdf:value "Together, aPARs and pPARs define the anterior-posterior axis of the zygote and orchestrate an asymmetric division that restricts germline determinants to the posterior daughter cell (Beatty et\u00A0al., >>2010<<, Boyd et\u00A0al., 1996, Etemad-Moghadam et\u00A0al., 1995, Gotta et\u00A0al., 2001, Guo and Kemphues, 1995, Hoege et\u00A0al., 2010, Kay and Hunter, 2001, Kumfer et\u00A0al., 2010, Tabuse et\u00A0al., 1998, Watts et\u00A0al., 1996)." ; ns1:mentions . _:b66507418 rdf:type ns1:Context ; rdf:value "Together, aPARs and pPARs define the anterior-posterior axis of the zygote and orchestrate an asymmetric division that restricts germline determinants to the posterior daughter cell (Beatty et\u00A0al., 2010, Boyd et\u00A0al., >>1996<<, Etemad-Moghadam et\u00A0al., 1995, Gotta et\u00A0al., 2001, Guo and Kemphues, 1995, Hoege et\u00A0al., 2010, Kay and Hunter, 2001, Kumfer et\u00A0al., 2010, Tabuse et\u00A0al., 1998, Watts et\u00A0al., 1996)." ; ns1:mentions . _:b66507419 rdf:type ns1:Context ; rdf:value "and pPARs define the anterior-posterior axis of the zygote and orchestrate an asymmetric division that restricts germline determinants to the posterior daughter cell (Beatty et\u00A0al., 2010, Boyd et\u00A0al., 1996, Etemad-Moghadam et\u00A0al., >>1995<<, Gotta et\u00A0al., 2001, Guo and Kemphues, 1995, Hoege et\u00A0al., 2010, Kay and Hunter, 2001, Kumfer et\u00A0al., 2010, Tabuse et\u00A0al., 1998, Watts et\u00A0al., 1996)." ; ns1:mentions . _:b66507420 rdf:type ns1:Context ; rdf:value "the anterior-posterior axis of the zygote and orchestrate an asymmetric division that restricts germline determinants to the posterior daughter cell (Beatty et\u00A0al., 2010, Boyd et\u00A0al., 1996, Etemad-Moghadam et\u00A0al., 1995, Gotta et\u00A0al., >>2001<<, Guo and Kemphues, 1995, Hoege et\u00A0al., 2010, Kay and Hunter, 2001, Kumfer et\u00A0al., 2010, Tabuse et\u00A0al., 1998, Watts et\u00A0al., 1996)." ; ns1:mentions . _:b66507421 rdf:type ns1:Context ; rdf:value "axis of the zygote and orchestrate an asymmetric division that restricts germline determinants to the posterior daughter cell (Beatty et\u00A0al., 2010, Boyd et\u00A0al., 1996, Etemad-Moghadam et\u00A0al., 1995, Gotta et\u00A0al., 2001, Guo and Kemphues, >>1995<<, Hoege et\u00A0al., 2010, Kay and Hunter, 2001, Kumfer et\u00A0al., 2010, Tabuse et\u00A0al., 1998, Watts et\u00A0al., 1996)." ; ns1:mentions . _:b66507422 rdf:type ns1:Context ; rdf:value "orchestrate an asymmetric division that restricts germline determinants to the posterior daughter cell (Beatty et\u00A0al., 2010, Boyd et\u00A0al., 1996, Etemad-Moghadam et\u00A0al., 1995, Gotta et\u00A0al., 2001, Guo and Kemphues, 1995, Hoege et\u00A0al., >>2010<<, Kay and Hunter, 2001, Kumfer et\u00A0al., 2010, Tabuse et\u00A0al., 1998, Watts et\u00A0al., 1996)." ; ns1:mentions . _:b66507423 rdf:type ns1:Context ; rdf:value "division that restricts germline determinants to the posterior daughter cell (Beatty et\u00A0al., 2010, Boyd et\u00A0al., 1996, Etemad-Moghadam et\u00A0al., 1995, Gotta et\u00A0al., 2001, Guo and Kemphues, 1995, Hoege et\u00A0al., 2010, Kay and Hunter, >>2001<<, Kumfer et\u00A0al., 2010, Tabuse et\u00A0al., 1998, Watts et\u00A0al., 1996)." ; ns1:mentions . _:b66507424 rdf:type ns1:Context ; rdf:value "restricts germline determinants to the posterior daughter cell (Beatty et\u00A0al., 2010, Boyd et\u00A0al., 1996, Etemad-Moghadam et\u00A0al., 1995, Gotta et\u00A0al., 2001, Guo and Kemphues, 1995, Hoege et\u00A0al., 2010, Kay and Hunter, 2001, Kumfer et\u00A0al., >>2010<<, Tabuse et\u00A0al., 1998, Watts et\u00A0al., 1996)." ; ns1:mentions . _:b66507425 rdf:type ns1:Context ; rdf:value "to the posterior daughter cell (Beatty et\u00A0al., 2010, Boyd et\u00A0al., 1996, Etemad-Moghadam et\u00A0al., 1995, Gotta et\u00A0al., 2001, Guo and Kemphues, 1995, Hoege et\u00A0al., 2010, Kay and Hunter, 2001, Kumfer et\u00A0al., 2010, Tabuse et\u00A0al., >>1998<<, Watts et\u00A0al., 1996)." ; ns1:mentions . _:b66507426 rdf:type ns1:Context ; rdf:value "daughter cell (Beatty et\u00A0al., 2010, Boyd et\u00A0al., 1996, Etemad-Moghadam et\u00A0al., 1995, Gotta et\u00A0al., 2001, Guo and Kemphues, 1995, Hoege et\u00A0al., 2010, Kay and Hunter, 2001, Kumfer et\u00A0al., 2010, Tabuse et\u00A0al., 1998, Watts et\u00A0al., >>1996<<).Figure" ; ns1:mentions . _:b66507427 rdf:type ns1:Context ; rdf:value "See recent reviews (Goehring, >>2014<<, Hoege and Hyman, 2013, Motegi and Seydoux, 2013) for more information." ; ns1:mentions . _:b66507428 rdf:type ns1:Context ; rdf:value "See recent reviews (Goehring, 2014, Hoege and Hyman, >>2013<<, Motegi and Seydoux, 2013) for more information." ; ns1:mentions . _:b66507429 rdf:type ns1:Context ; rdf:value "See recent reviews (Goehring, 2014, Hoege and Hyman, 2013, Motegi and Seydoux, >>2013<<) for more information." ; ns1:mentions . _:b66507430 rdf:type ns1:Context ; rdf:value "First, the centrosome induces actomyosin cortical flow away from the newly defined posterior pole, which transports membrane-associated aPAR proteins into the anterior (Cheeks et\u00A0al., >>2004<<, Goehring et\u00A0al., 2011b, Mayer et\u00A0al., 2010, Munro et\u00A0al., 2004)." ; ns1:mentions . _:b66507431 rdf:type ns1:Context ; rdf:value "First, the centrosome induces actomyosin cortical flow away from the newly defined posterior pole, which transports membrane-associated aPAR proteins into the anterior (Cheeks et\u00A0al., 2004, Goehring et\u00A0al., 2011b, Mayer et\u00A0al., 2010, Munro et\u00A0al., 2004)." ; ns1:mentions . _:b66507432 rdf:type ns1:Context ; rdf:value "First, the centrosome induces actomyosin cortical flow away from the newly defined posterior pole, which transports membrane-associated aPAR proteins into the anterior (Cheeks et\u00A0al., 2004, Goehring et\u00A0al., 2011b, Mayer et\u00A0al., >>2010<<, Munro et\u00A0al., 2004)." ; ns1:mentions . _:b66507433 rdf:type ns1:Context ; rdf:value "induces actomyosin cortical flow away from the newly defined posterior pole, which transports membrane-associated aPAR proteins into the anterior (Cheeks et\u00A0al., 2004, Goehring et\u00A0al., 2011b, Mayer et\u00A0al., 2010, Munro et\u00A0al., >>2004<<). Second, centrosomal microtubules promote local loading of PAR-2 in the posterior." ; ns1:mentions . _:b66507434 rdf:type ns1:Context ; rdf:value "PAR-2 then recruits PAR-1, which drives posterior exclusion of aPARs through phosphorylation of\u00A0PAR-3 (Boyd et\u00A0al., >>1996<<, Hao et\u00A0al., 2006, Motegi et\u00A0al., 2011)." ; ns1:mentions . _:b66507435 rdf:type ns1:Context ; rdf:value "PAR-2 then recruits PAR-1, which drives posterior exclusion of aPARs through phosphorylation of\u00A0PAR-3 (Boyd et\u00A0al., 1996, Hao et\u00A0al., >>2006<<, Motegi et\u00A0al., 2011)." ; ns1:mentions . _:b66507436 rdf:type ns1:Context ; rdf:value "PAR-2 then recruits PAR-1, which drives posterior exclusion of aPARs through phosphorylation of\u00A0PAR-3 (Boyd et\u00A0al., 1996, Hao et\u00A0al., 2006, Motegi et\u00A0al., >>2011<<). Following this \u201Cestablishment phase,\u201D the zygote enters a \u201Cmaintenance phase\u201D during which mutual antagonism between anterior and posterior PARs ensures their continued asymmetric localizations (Boyd et\u00A0al., 1996, Cuenca et\u00A0al., 2003," ; ns1:mentions . _:b66507437 rdf:type ns1:Context ; rdf:value "Following this \u201Cestablishment phase,\u201D the zygote enters a \u201Cmaintenance phase\u201D during which mutual antagonism between anterior and posterior PARs ensures their continued asymmetric localizations (Boyd et\u00A0al., >>1996<<, Cuenca et\u00A0al., 2003, Etemad-Moghadam et\u00A0al., 1995, Guo and Kemphues, 1995, Tabuse et\u00A0al., 1998, Watts et\u00A0al., 1996)." ; ns1:mentions . _:b66507438 rdf:type ns1:Context ; rdf:value "Following this \u201Cestablishment phase,\u201D the zygote enters a \u201Cmaintenance phase\u201D during which mutual antagonism between anterior and posterior PARs ensures their continued asymmetric localizations (Boyd et\u00A0al., 1996, Cuenca et\u00A0al., >>2003<<, Etemad-Moghadam et\u00A0al., 1995, Guo and Kemphues, 1995, Tabuse et\u00A0al., 1998, Watts et\u00A0al., 1996)." ; ns1:mentions . _:b66507439 rdf:type ns1:Context ; rdf:value "phase,\u201D the zygote enters a \u201Cmaintenance phase\u201D during which mutual antagonism between anterior and posterior PARs ensures their continued asymmetric localizations (Boyd et\u00A0al., 1996, Cuenca et\u00A0al., 2003, Etemad-Moghadam et\u00A0al., >>1995<<, Guo and Kemphues, 1995, Tabuse et\u00A0al., 1998, Watts et\u00A0al., 1996)." ; ns1:mentions . _:b66507440 rdf:type ns1:Context ; rdf:value "enters a \u201Cmaintenance phase\u201D during which mutual antagonism between anterior and posterior PARs ensures their continued asymmetric localizations (Boyd et\u00A0al., 1996, Cuenca et\u00A0al., 2003, Etemad-Moghadam et\u00A0al., 1995, Guo and Kemphues, >>1995<<, Tabuse et\u00A0al., 1998, Watts et\u00A0al., 1996)." ; ns1:mentions . _:b66507441 rdf:type ns1:Context ; rdf:value "phase\u201D during which mutual antagonism between anterior and posterior PARs ensures their continued asymmetric localizations (Boyd et\u00A0al., 1996, Cuenca et\u00A0al., 2003, Etemad-Moghadam et\u00A0al., 1995, Guo and Kemphues, 1995, Tabuse et\u00A0al., >>1998<<, Watts et\u00A0al., 1996)." ; ns1:mentions . _:b66507442 rdf:type ns1:Context ; rdf:value "mutual antagonism between anterior and posterior PARs ensures their continued asymmetric localizations (Boyd et\u00A0al., 1996, Cuenca et\u00A0al., 2003, Etemad-Moghadam et\u00A0al., 1995, Guo and Kemphues, 1995, Tabuse et\u00A0al., 1998, Watts et\u00A0al., >>1996<<)." ; ns1:mentions . _:b66507443 rdf:type ns1:Context ; rdf:value "Anterior PAR protein function is mediated through the kinase activity of PKC-3, which can phosphorylate PAR-1, PAR-2, and LGL-1 and drive their dissociation from the membrane (Beatty et\u00A0al., >>2010<<, Hao et\u00A0al., 2006, Hoege et\u00A0al., 2010, Hurov et\u00A0al., 2004, Motegi et\u00A0al., 2011)." ; ns1:mentions . _:b66507444 rdf:type ns1:Context ; rdf:value "Anterior PAR protein function is mediated through the kinase activity of PKC-3, which can phosphorylate PAR-1, PAR-2, and LGL-1 and drive their dissociation from the membrane (Beatty et\u00A0al., 2010, Hao et\u00A0al., >>2006<<, Hoege et\u00A0al., 2010, Hurov et\u00A0al., 2004, Motegi et\u00A0al., 2011). PAR-3, PAR-6, and CDC-42 are all required for proper PKC-3 membrane localization (Gotta et\u00A0al., 2001, Kay and Hunter, 2001, Schonegg and Hyman, 2006, Tabuse et\u00A0al., 1998)." ; ns1:mentions . _:b66507445 rdf:type ns1:Context ; rdf:value "Anterior PAR protein function is mediated through the kinase activity of PKC-3, which can phosphorylate PAR-1, PAR-2, and LGL-1 and drive their dissociation from the membrane (Beatty et\u00A0al., 2010, Hao et\u00A0al., 2006, Hoege et\u00A0al., >>2010<<, Hurov et\u00A0al., 2004, Motegi et\u00A0al., 2011). PAR-3, PAR-6, and CDC-42 are all required for proper PKC-3 membrane localization (Gotta et\u00A0al., 2001, Kay and Hunter, 2001, Schonegg and Hyman, 2006, Tabuse et\u00A0al., 1998)." ; ns1:mentions . _:b66507446 rdf:type ns1:Context ; rdf:value "function is mediated through the kinase activity of PKC-3, which can phosphorylate PAR-1, PAR-2, and LGL-1 and drive their dissociation from the membrane (Beatty et\u00A0al., 2010, Hao et\u00A0al., 2006, Hoege et\u00A0al., 2010, Hurov et\u00A0al., >>2004<<, Motegi et\u00A0al., 2011). PAR-3, PAR-6, and CDC-42 are all required for proper PKC-3 membrane localization (Gotta et\u00A0al., 2001, Kay and Hunter, 2001, Schonegg and Hyman, 2006, Tabuse et\u00A0al., 1998)." ; ns1:mentions . _:b66507447 rdf:type ns1:Context ; rdf:value "through the kinase activity of PKC-3, which can phosphorylate PAR-1, PAR-2, and LGL-1 and drive their dissociation from the membrane (Beatty et\u00A0al., 2010, Hao et\u00A0al., 2006, Hoege et\u00A0al., 2010, Hurov et\u00A0al., 2004, Motegi et\u00A0al., >>2011<<). PAR-3, PAR-6, and CDC-42 are all required for proper PKC-3 membrane localization (Gotta et\u00A0al., 2001, Kay and Hunter, 2001, Schonegg and Hyman, 2006, Tabuse et\u00A0al., 1998)." ; ns1:mentions . _:b66507448 rdf:type ns1:Context ; rdf:value "PAR-3, PAR-6, and CDC-42 are all required for proper PKC-3 membrane localization (Gotta et\u00A0al., >>2001<<, Kay and Hunter, 2001, Schonegg and Hyman, 2006, Tabuse et\u00A0al., 1998)." ; ns1:mentions . _:b66507449 rdf:type ns1:Context ; rdf:value "PAR-3, PAR-6, and CDC-42 are all required for proper PKC-3 membrane localization (Gotta et\u00A0al., 2001, Kay and Hunter, >>2001<<, Schonegg and Hyman, 2006, Tabuse et\u00A0al., 1998)." ; ns1:mentions . _:b66507450 rdf:type ns1:Context ; rdf:value "PAR-3, PAR-6, and CDC-42 are all required for proper PKC-3 membrane localization (Gotta et\u00A0al., 2001, Kay and Hunter, 2001, Schonegg and Hyman, >>2006<<, Tabuse et\u00A0al., 1998)." ; ns1:mentions . _:b66507451 rdf:type ns1:Context ; rdf:value "PAR-3, PAR-6, and CDC-42 are all required for proper PKC-3 membrane localization (Gotta et\u00A0al., 2001, Kay and Hunter, 2001, Schonegg and Hyman, 2006, Tabuse et\u00A0al., >>1998<<). Consequently, loss of any of these four proteins results in identical zygote polarity phenotypes: posterior PAR\u00A0proteins are found uniformly on the embryo membrane and the first cell division is symmetric, leading to cell fate defects" ; ns1:mentions . _:b66507452 rdf:type ns1:Context ; rdf:value "results in identical zygote polarity phenotypes: posterior PAR\u00A0proteins are found uniformly on the embryo membrane and the first cell division is symmetric, leading to cell fate defects and embryo lethality (Etemad-Moghadam et\u00A0al., >>1995<<, Kay and Hunter, 2001, Tabuse et\u00A0al., 1998, Watts et\u00A0al., 1996)." ; ns1:mentions . _:b66507453 rdf:type ns1:Context ; rdf:value "zygote polarity phenotypes: posterior PAR\u00A0proteins are found uniformly on the embryo membrane and the first cell division is symmetric, leading to cell fate defects and embryo lethality (Etemad-Moghadam et\u00A0al., 1995, Kay and Hunter, >>2001<<, Tabuse et\u00A0al., 1998, Watts et\u00A0al., 1996)." ; ns1:mentions . _:b66507454 rdf:type ns1:Context ; rdf:value "posterior PAR\u00A0proteins are found uniformly on the embryo membrane and the first cell division is symmetric, leading to cell fate defects and embryo lethality (Etemad-Moghadam et\u00A0al., 1995, Kay and Hunter, 2001, Tabuse et\u00A0al., >>1998<<, Watts et\u00A0al., 1996)." ; ns1:mentions . _:b66507455 rdf:type ns1:Context ; rdf:value "proteins are found uniformly on the embryo membrane and the first cell division is symmetric, leading to cell fate defects and embryo lethality (Etemad-Moghadam et\u00A0al., 1995, Kay and Hunter, 2001, Tabuse et\u00A0al., 1998, Watts et\u00A0al., >>1996<<)." ; ns1:mentions . _:b66507456 rdf:type ns1:Context ; rdf:value "This similarity of aPAR mutant phenotypes, their co-segregation within the anterior domain, and their ability to interact with one another in a wide range of systems (Izumi et\u00A0al., >>1998<<, Joberty et\u00A0al., 2000, Lin et\u00A0al., 2000, Qiu et\u00A0al., 2000) has led to consideration of an effective aPAR unit comprising PAR-3, PAR-6, PKC-3, and CDC-42." ; ns1:mentions . _:b66507457 rdf:type ns1:Context ; rdf:value "This similarity of aPAR mutant phenotypes, their co-segregation within the anterior domain, and their ability to interact with one another in a wide range of systems (Izumi et\u00A0al., 1998, Joberty et\u00A0al., >>2000<<, Lin et\u00A0al., 2000, Qiu et\u00A0al., 2000) has led to consideration of an effective aPAR unit comprising PAR-3, PAR-6, PKC-3, and CDC-42." ; ns1:mentions . _:b66507458 rdf:type ns1:Context ; rdf:value "This similarity of aPAR mutant phenotypes, their co-segregation within the anterior domain, and their ability to interact with one another in a wide range of systems (Izumi et\u00A0al., 1998, Joberty et\u00A0al., 2000, Lin et\u00A0al., >>2000<<, Qiu et\u00A0al., 2000) has led to consideration of an effective aPAR unit comprising PAR-3, PAR-6, PKC-3, and CDC-42." ; ns1:mentions . _:b66507459 rdf:type ns1:Context ; rdf:value "similarity of aPAR mutant phenotypes, their co-segregation within the anterior domain, and their ability to interact with one another in a wide range of systems (Izumi et\u00A0al., 1998, Joberty et\u00A0al., 2000, Lin et\u00A0al., 2000, Qiu et\u00A0al., >>2000<<) has led to consideration of an effective aPAR unit comprising PAR-3, PAR-6, PKC-3, and CDC-42." ; ns1:mentions . _:b66507460 rdf:type ns1:Context ; rdf:value "and aPKC localize to distinct regions of the apical domain: PAR-3 is primarily junctional, while PAR-6 and aPKC are more apical and, together with CDC-42 and Crumbs, exclude PAR-1 and LGL from the apical domain (Betschinger et\u00A0al., >>2003<<, Harris and Peifer, 2005, Morais-De-Sa et\u00A0al., 2010, Yamanaka et\u00A0al., 2006, Yamanaka et\u00A0al., 2003)." ; ns1:mentions . _:b66507461 rdf:type ns1:Context ; rdf:value "regions of the apical domain: PAR-3 is primarily junctional, while PAR-6 and aPKC are more apical and, together with CDC-42 and Crumbs, exclude PAR-1 and LGL from the apical domain (Betschinger et\u00A0al., 2003, Harris and Peifer, >>2005<<, Morais-De-Sa et\u00A0al., 2010, Yamanaka et\u00A0al., 2006, Yamanaka et\u00A0al., 2003)." ; ns1:mentions . _:b66507462 rdf:type ns1:Context ; rdf:value "domain: PAR-3 is primarily junctional, while PAR-6 and aPKC are more apical and, together with CDC-42 and Crumbs, exclude PAR-1 and LGL from the apical domain (Betschinger et\u00A0al., 2003, Harris and Peifer, 2005, Morais-De-Sa et\u00A0al., >>2010<<, Yamanaka et\u00A0al., 2006, Yamanaka et\u00A0al., 2003)." ; ns1:mentions . _:b66507463 rdf:type ns1:Context ; rdf:value "junctional, while PAR-6 and aPKC are more apical and, together with CDC-42 and Crumbs, exclude PAR-1 and LGL from the apical domain (Betschinger et\u00A0al., 2003, Harris and Peifer, 2005, Morais-De-Sa et\u00A0al., 2010, Yamanaka et\u00A0al., >>2006<<, Yamanaka et\u00A0al., 2003)." ; ns1:mentions . _:b66507464 rdf:type ns1:Context ; rdf:value "PAR-6 and aPKC are more apical and, together with CDC-42 and Crumbs, exclude PAR-1 and LGL from the apical domain (Betschinger et\u00A0al., 2003, Harris and Peifer, 2005, Morais-De-Sa et\u00A0al., 2010, Yamanaka et\u00A0al., 2006, Yamanaka et\u00A0al., >>2003<<)." ; ns1:mentions . _:b66507465 rdf:type ns1:Context ; rdf:value "Supporting this hypothesis, PAR-6 and PKC-3 only partially co-localize with PAR-3 in wild-type embryos, but co-localize strongly when CDC-42 is depleted (Beers and Kemphues, >>2006<<, Hung and Kemphues, 1999, Tabuse et\u00A0al., 1998)." ; ns1:mentions . _:b66507466 rdf:type ns1:Context ; rdf:value "Supporting this hypothesis, PAR-6 and PKC-3 only partially co-localize with PAR-3 in wild-type embryos, but co-localize strongly when CDC-42 is depleted (Beers and Kemphues, 2006, Hung and Kemphues, >>1999<<, Tabuse et\u00A0al., 1998)." ; ns1:mentions . _:b66507467 rdf:type ns1:Context ; rdf:value "Supporting this hypothesis, PAR-6 and PKC-3 only partially co-localize with PAR-3 in wild-type embryos, but co-localize strongly when CDC-42 is depleted (Beers and Kemphues, 2006, Hung and Kemphues, 1999, Tabuse et\u00A0al., >>1998<<). However, it remains unclear whether these observations reflect the existence of discrete functional modules and, if so, what their respective functions are." ; ns1:mentions . _:b66507468 rdf:type ns4:Section ; dc:title "results" ; ns4:contains _:b66507536 , _:b66507537 , _:b66507538 , _:b66507524 , _:b66507525 , _:b66507526 , _:b66507527 , _:b66507520 , _:b66507521 , _:b66507522 , _:b66507523 , _:b66507532 , _:b66507533 , _:b66507534 , _:b66507535 , _:b66507528 , _:b66507529 , _:b66507530 , _:b66507531 , _:b66507476 , _:b66507477 , _:b66507478 , _:b66507479 , _:b66507472 , _:b66507473 , _:b66507474 , _:b66507475 , _:b66507484 , _:b66507485 , _:b66507486 , _:b66507487 , _:b66507480 , _:b66507481 , _:b66507482 , _:b66507483 , _:b66507469 , _:b66507470 , _:b66507471 , _:b66507508 , _:b66507509 , _:b66507510 , _:b66507511 , _:b66507504 , _:b66507505 , _:b66507506 , _:b66507507 , _:b66507516 , _:b66507517 , _:b66507518 , _:b66507519 , _:b66507512 , _:b66507513 , _:b66507514 , _:b66507515 , _:b66507492 , _:b66507493 , _:b66507494 , _:b66507495 , _:b66507488 , _:b66507489 , _:b66507490 , _:b66507491 , _:b66507500 , _:b66507501 , _:b66507502 , _:b66507503 , _:b66507496 , _:b66507497 , _:b66507498 , _:b66507499 . _:b66507469 rdf:type ns1:Context ; rdf:value "First, we examined a previously identified temperature-sensitive allele of pkc-3, ne4246 (Fievet et\u00A0al., >>2012<<), which alters a conserved Asp residue (D386V) close to the active site." ; ns1:mentions . _:b66507470 rdf:type ns1:Context ; rdf:value "In parallel, we examined PKC-3 inhibitors in permeable, perm-1(RNAi) embryos (Carvalho et\u00A0al., >>2011<<) to identify compounds that yielded a PKC-3 deficient polarity phenotype." ; ns1:mentions . _:b66507471 rdf:type ns1:Context ; rdf:value "One compound, CRT0103390 (CRT90), a derivative of CRT0066854 (Figures S2A\u2013S2C) (Kj\u00E6r et\u00A0al., >>2013<<, Dorsey et\u00A0al., 2013) resulted in embryos that progressed normally through the cell cycle but showed loss of PAR-2 asymmetry and divided symmetrically (Figure\u00A01E and Movie S1)." ; ns1:mentions . _:b66507472 rdf:type ns1:Context ; rdf:value "Normally, PAR-3, PAR-6, and PKC-3 are efficiently segregated into the anterior during the polarity establishment phase and remain asymmetric until cytokinesis (Figures 1A and 2A\u20132F) (Cuenca et\u00A0al., >>2003<<). When PKC-3 is depleted by RNAi of pkc-3, PAR-6 fails to localize to the membrane (Figure\u00A02A) (Hung and Kemphues, 1999). By contrast, PAR-3 remains membrane associated and segregates into the anterior, but this population is generally" ; ns1:mentions . _:b66507473 rdf:type ns1:Context ; rdf:value "When PKC-3 is depleted by RNAi of pkc-3, PAR-6 fails to localize to the membrane (Figure\u00A02A) (Hung and Kemphues, >>1999<<). By contrast, PAR-3 remains membrane associated and segregates into the anterior, but this population is generally reduced compared to wild-type and is lost as the cell proceeds through mitosis (Figure\u00A02E) (Tabuse et\u00A0al., 1998).Figure" ; ns1:mentions . _:b66507474 rdf:type ns1:Context ; rdf:value "By contrast, PAR-3 remains membrane associated and segregates into the anterior, but this population is generally reduced compared to wild-type and is lost as the cell proceeds through mitosis (Figure\u00A02E) (Tabuse et\u00A0al., >>1998<<).Figure" ; ns1:mentions . _:b66507475 rdf:type ns1:Context ; rdf:value "However, we found that PAR-1 kinase activity, as measured by MEX-5 mobility (Griffin et\u00A0al., >>2011<<), appears normal in PKC-3-inhibited zygotes (Figures S2D\u2013S2F)." ; ns1:mentions . _:b66507476 rdf:type ns1:Context ; rdf:value "Contrary to what has been observed in Drosophila epithelia, where aPKC activity promotes decoupling of PAR-3 from PAR-6/aPKC and their targeting to distinct sites (Morais-De-Sa et\u00A0al., >>2010<<), here we observe the opposite:" ; ns1:mentions . _:b66507477 rdf:type ns1:Context ; rdf:value "PKC-3 and PAR-6 normally require both PAR-3 and CDC-42 to localize stably to the membrane (Beers and Kemphues, >>2006<<, Sailer et\u00A0al., 2015)." ; ns1:mentions . _:b66507478 rdf:type ns1:Context ; rdf:value "PKC-3 and PAR-6 normally require both PAR-3 and CDC-42 to localize stably to the membrane (Beers and Kemphues, 2006, Sailer et\u00A0al., >>2015<<). The dependency on PAR-3 is stronger: PKC-3 and PAR-6 fail to localize to the membrane in PAR-3-depleted zygotes (par-3(RNAi) in Figures 3A\u20133F) (Tabuse et\u00A0al., 1998). By contrast, in CDC-42-depleted zygotes, PKC-3 and PAR-6 initially" ; ns1:mentions . _:b66507479 rdf:type ns1:Context ; rdf:value "The dependency on PAR-3 is stronger: PKC-3 and PAR-6 fail to localize to the membrane in PAR-3-depleted zygotes (par-3(RNAi) in Figures 3A\u20133F) (Tabuse et\u00A0al., >>1998<<). By contrast, in CDC-42-depleted zygotes, PKC-3 and PAR-6 initially localize to the membrane and segregate to the anterior, but their membrane localization is gradually lost during the maintenance phase, becoming weaker and more uniform" ; ns1:mentions . _:b66507480 rdf:type ns1:Context ; rdf:value "segregate to the anterior, but their membrane localization is gradually lost during the maintenance phase, becoming weaker and more uniform as zygotes approach cytokinesis (cdc-42(RNAi) Figures 3A\u20133F and Movie S4)(Beers and Kemphues, >>2006<<, Gotta et\u00A0al., 2001, Motegi and Sugimoto, 2006, Sailer et\u00A0al., 2015, Schonegg and Hyman, 2006)." ; ns1:mentions . _:b66507481 rdf:type ns1:Context ; rdf:value "but their membrane localization is gradually lost during the maintenance phase, becoming weaker and more uniform as zygotes approach cytokinesis (cdc-42(RNAi) Figures 3A\u20133F and Movie S4)(Beers and Kemphues, 2006, Gotta et\u00A0al., >>2001<<, Motegi and Sugimoto, 2006, Sailer et\u00A0al., 2015, Schonegg and Hyman, 2006)." ; ns1:mentions . _:b66507482 rdf:type ns1:Context ; rdf:value "localization is gradually lost during the maintenance phase, becoming weaker and more uniform as zygotes approach cytokinesis (cdc-42(RNAi) Figures 3A\u20133F and Movie S4)(Beers and Kemphues, 2006, Gotta et\u00A0al., 2001, Motegi and Sugimoto, >>2006<<, Sailer et\u00A0al., 2015, Schonegg and Hyman, 2006)." ; ns1:mentions . _:b66507483 rdf:type ns1:Context ; rdf:value "lost during the maintenance phase, becoming weaker and more uniform as zygotes approach cytokinesis (cdc-42(RNAi) Figures 3A\u20133F and Movie S4)(Beers and Kemphues, 2006, Gotta et\u00A0al., 2001, Motegi and Sugimoto, 2006, Sailer et\u00A0al., >>2015<<, Schonegg and Hyman, 2006)." ; ns1:mentions . _:b66507484 rdf:type ns1:Context ; rdf:value "phase, becoming weaker and more uniform as zygotes approach cytokinesis (cdc-42(RNAi) Figures 3A\u20133F and Movie S4)(Beers and Kemphues, 2006, Gotta et\u00A0al., 2001, Motegi and Sugimoto, 2006, Sailer et\u00A0al., 2015, Schonegg and Hyman, >>2006<<). Importantly, depletion of PAR-1 or PAR-2, which invade the anterior in the absence of PAR-3 or CDC-42 (Etemad-Moghadam et\u00A0al., 1995, Gotta et\u00A0al., 2001, Kay and Hunter, 2001, Schonegg and Hyman, 2006), fails to rescue PAR-6 membrane" ; ns1:mentions . _:b66507485 rdf:type ns1:Context ; rdf:value "Importantly, depletion of PAR-1 or PAR-2, which invade the anterior in the absence of PAR-3 or CDC-42 (Etemad-Moghadam et\u00A0al., >>1995<<, Gotta et\u00A0al., 2001, Kay and Hunter, 2001, Schonegg and Hyman, 2006), fails to rescue PAR-6 membrane localization in these conditions (Figure\u00A0S4)." ; ns1:mentions . _:b66507486 rdf:type ns1:Context ; rdf:value "Importantly, depletion of PAR-1 or PAR-2, which invade the anterior in the absence of PAR-3 or CDC-42 (Etemad-Moghadam et\u00A0al., 1995, Gotta et\u00A0al., >>2001<<, Kay and Hunter, 2001, Schonegg and Hyman, 2006), fails to rescue PAR-6 membrane localization in these conditions (Figure\u00A0S4)." ; ns1:mentions . _:b66507487 rdf:type ns1:Context ; rdf:value "Importantly, depletion of PAR-1 or PAR-2, which invade the anterior in the absence of PAR-3 or CDC-42 (Etemad-Moghadam et\u00A0al., 1995, Gotta et\u00A0al., 2001, Kay and Hunter, >>2001<<, Schonegg and Hyman, 2006), fails to rescue PAR-6 membrane localization in these conditions (Figure\u00A0S4)." ; ns1:mentions . _:b66507488 rdf:type ns1:Context ; rdf:value "Importantly, depletion of PAR-1 or PAR-2, which invade the anterior in the absence of PAR-3 or CDC-42 (Etemad-Moghadam et\u00A0al., 1995, Gotta et\u00A0al., 2001, Kay and Hunter, 2001, Schonegg and Hyman, >>2006<<), fails to rescue PAR-6 membrane localization in these conditions (Figure\u00A0S4)." ; ns1:mentions . _:b66507489 rdf:type ns1:Context ; rdf:value "Consistent with previous work showing that PAR-6/aPKC are typically associated with an active, guanosine triphosphate (GTP)-bound form of CDC-42 (Atwood et\u00A0al., >>2007<<, Gotta et\u00A0al., 2001, Joberty et\u00A0al., 2000, Lin et\u00A0al., 2000, Qiu et\u00A0al., 2000), we found that decreasing CDC-42/GTP by depletion of the CDC-42 GEF, CGEF-1, reduces membrane association of PAR-6/PKC-3 in pkc-3(ts) embryos compared with" ; ns1:mentions . _:b66507490 rdf:type ns1:Context ; rdf:value "Consistent with previous work showing that PAR-6/aPKC are typically associated with an active, guanosine triphosphate (GTP)-bound form of CDC-42 (Atwood et\u00A0al., 2007, Gotta et\u00A0al., >>2001<<, Joberty et\u00A0al., 2000, Lin et\u00A0al., 2000, Qiu et\u00A0al., 2000), we found that decreasing CDC-42/GTP by depletion of the CDC-42 GEF, CGEF-1, reduces membrane association of PAR-6/PKC-3 in pkc-3(ts) embryos compared with wild-type, while" ; ns1:mentions . _:b66507491 rdf:type ns1:Context ; rdf:value "Consistent with previous work showing that PAR-6/aPKC are typically associated with an active, guanosine triphosphate (GTP)-bound form of CDC-42 (Atwood et\u00A0al., 2007, Gotta et\u00A0al., 2001, Joberty et\u00A0al., >>2000<<, Lin et\u00A0al., 2000, Qiu et\u00A0al., 2000), we found that decreasing CDC-42/GTP by depletion of the CDC-42 GEF, CGEF-1, reduces membrane association of PAR-6/PKC-3 in pkc-3(ts) embryos compared with wild-type, while leaving PAR-3 levels" ; ns1:mentions . _:b66507492 rdf:type ns1:Context ; rdf:value "Consistent with previous work showing that PAR-6/aPKC are typically associated with an active, guanosine triphosphate (GTP)-bound form of CDC-42 (Atwood et\u00A0al., 2007, Gotta et\u00A0al., 2001, Joberty et\u00A0al., 2000, Lin et\u00A0al., >>2000<<, Qiu et\u00A0al., 2000), we found that decreasing CDC-42/GTP by depletion of the CDC-42 GEF, CGEF-1, reduces membrane association of PAR-6/PKC-3 in pkc-3(ts) embryos compared with wild-type, while leaving PAR-3 levels unchanged (Figures 3G and" ; ns1:mentions . _:b66507493 rdf:type ns1:Context ; rdf:value "with previous work showing that PAR-6/aPKC are typically associated with an active, guanosine triphosphate (GTP)-bound form of CDC-42 (Atwood et\u00A0al., 2007, Gotta et\u00A0al., 2001, Joberty et\u00A0al., 2000, Lin et\u00A0al., 2000, Qiu et\u00A0al., >>2000<<), we found that decreasing CDC-42/GTP by depletion of the CDC-42 GEF, CGEF-1, reduces membrane association of PAR-6/PKC-3 in pkc-3(ts) embryos compared with wild-type, while leaving PAR-3 levels unchanged (Figures 3G and S5A\u2013S5E)." ; ns1:mentions . _:b66507494 rdf:type ns1:Context ; rdf:value "PKC-3 remains dependent on PAR-6 in PKC-3-inhibited embryos, consistent with PAR-6 being required to mediate the interactions of the PAR-6/PKC-3 heterodimer with PAR-3 and/or CDC-42 (Figures 3A and 3D\u20133F; Movie S4) (Joberty et\u00A0al., >>2000<<, Qiu et\u00A0al., 2000), as well as our general findings that PKC-3 and PAR-6 respond similarly in all assays described." ; ns1:mentions . _:b66507495 rdf:type ns1:Context ; rdf:value "on PAR-6 in PKC-3-inhibited embryos, consistent with PAR-6 being required to mediate the interactions of the PAR-6/PKC-3 heterodimer with PAR-3 and/or CDC-42 (Figures 3A and 3D\u20133F; Movie S4) (Joberty et\u00A0al., 2000, Qiu et\u00A0al., >>2000<<), as well as our general findings that PKC-3 and PAR-6 respond similarly in all assays described." ; ns1:mentions . _:b66507496 rdf:type ns1:Context ; rdf:value "Previous work has shown that the efficient segregation of anterior PAR proteins is due to advective transport by anteriorly-directed actomyosin cortical flow (Cheeks et\u00A0al., >>2004<<, Goehring et\u00A0al., 2011b, Munro et\u00A0al., 2004)." ; ns1:mentions . _:b66507497 rdf:type ns1:Context ; rdf:value "Previous work has shown that the efficient segregation of anterior PAR proteins is due to advective transport by anteriorly-directed actomyosin cortical flow (Cheeks et\u00A0al., 2004, Goehring et\u00A0al., 2011b, Munro et\u00A0al., 2004)." ; ns1:mentions . _:b66507498 rdf:type ns1:Context ; rdf:value "Previous work has shown that the efficient segregation of anterior PAR proteins is due to advective transport by anteriorly-directed actomyosin cortical flow (Cheeks et\u00A0al., 2004, Goehring et\u00A0al., 2011b, Munro et\u00A0al., >>2004<<). Because PAR-3 continues to be segregated efficiently in PKC-3-inhibited zygotes, we reasoned that PKC-3 inhibition may selectively alter the molecular organization of PAR-6 and PKC-3 at the membrane relative to PAR-3, which would be" ; ns1:mentions . _:b66507499 rdf:type ns1:Context ; rdf:value "To investigate these possibilities, we imaged PAR-3, PAR-6, and PKC-3 at the membrane using variable-angle epifluorescence microscopy (VAEM or pseudo-TIRF) (Konopka and Bednarek, >>2008<<). All three proteins exhibit a distinct clustered appearance during the polarity establishment phase, consistent with reports of non-homogenous distributions of PAR proteins at the membrane (Figures 4A and 5A, Establishment) (Munro et\u00A0al." ; ns1:mentions . _:b66507500 rdf:type ns1:Context ; rdf:value "three proteins exhibit a distinct clustered appearance during the polarity establishment phase, consistent with reports of non-homogenous distributions of PAR proteins at the membrane (Figures 4A and 5A, Establishment) (Munro et\u00A0al., >>2004<<, Robin et\u00A0al., 2014, Sailer et\u00A0al., 2015)." ; ns1:mentions . _:b66507501 rdf:type ns1:Context ; rdf:value "a distinct clustered appearance during the polarity establishment phase, consistent with reports of non-homogenous distributions of PAR proteins at the membrane (Figures 4A and 5A, Establishment) (Munro et\u00A0al., 2004, Robin et\u00A0al., >>2014<<, Sailer et\u00A0al., 2015)." ; ns1:mentions . _:b66507502 rdf:type ns1:Context ; rdf:value "appearance during the polarity establishment phase, consistent with reports of non-homogenous distributions of PAR proteins at the membrane (Figures 4A and 5A, Establishment) (Munro et\u00A0al., 2004, Robin et\u00A0al., 2014, Sailer et\u00A0al., >>2015<<). Similar to previous analysis of PAR-6 (Munro et\u00A0al., 2004), we find that clusters of PAR-6, PAR-3, and PKC-3 move in a highly directional manner in the direction of cortical flow, coinciding with increasing overall asymmetry (Figure\u00A04B" ; ns1:mentions . _:b66507503 rdf:type ns1:Context ; rdf:value "Similar to previous analysis of PAR-6 (Munro et\u00A0al., >>2004<<), we find that clusters of PAR-6, PAR-3, and PKC-3 move in a highly directional manner in the direction of cortical flow, coinciding with increasing overall asymmetry (Figure\u00A04B and Movie S5)." ; ns1:mentions . _:b66507504 rdf:type ns1:Context ; rdf:value "While aPAR clusters have been noted (Hung and Kemphues, >>1999<<, Munro et\u00A0al., 2004, Sailer et\u00A0al., 2015), the relationship between clustered and non-clustered PAR proteins and their ability to segregate in response to flow has not been explored." ; ns1:mentions . _:b66507505 rdf:type ns1:Context ; rdf:value "While aPAR clusters have been noted (Hung and Kemphues, 1999, Munro et\u00A0al., >>2004<<, Sailer et\u00A0al., 2015), the relationship between clustered and non-clustered PAR proteins and their ability to segregate in response to flow has not been explored." ; ns1:mentions . _:b66507506 rdf:type ns1:Context ; rdf:value "While aPAR clusters have been noted (Hung and Kemphues, 1999, Munro et\u00A0al., 2004, Sailer et\u00A0al., >>2015<<), the relationship between clustered and non-clustered PAR proteins and their ability to segregate in response to flow has not been explored." ; ns1:mentions . _:b66507507 rdf:type ns1:Context ; rdf:value "PAR-3 contains a conserved CR1 oligomerization domain, which is required for membrane binding and is targeted by PAR-1 kinase to induce displacement form the membrane (Benton and St Johnston, >>2003<<, Feng et\u00A0al., 2007, Li et\u00A0al., 2010, Mizuno et\u00A0al., 2003)." ; ns1:mentions . _:b66507508 rdf:type ns1:Context ; rdf:value "PAR-3 contains a conserved CR1 oligomerization domain, which is required for membrane binding and is targeted by PAR-1 kinase to induce displacement form the membrane (Benton and St Johnston, 2003, Feng et\u00A0al., >>2007<<, Li et\u00A0al., 2010, Mizuno et\u00A0al., 2003)." ; ns1:mentions . _:b66507509 rdf:type ns1:Context ; rdf:value "PAR-3 contains a conserved CR1 oligomerization domain, which is required for membrane binding and is targeted by PAR-1 kinase to induce displacement form the membrane (Benton and St Johnston, 2003, Feng et\u00A0al., 2007, Li et\u00A0al., >>2010<<, Mizuno et\u00A0al., 2003)." ; ns1:mentions . _:b66507510 rdf:type ns1:Context ; rdf:value "a conserved CR1 oligomerization domain, which is required for membrane binding and is targeted by PAR-1 kinase to induce displacement form the membrane (Benton and St Johnston, 2003, Feng et\u00A0al., 2007, Li et\u00A0al., 2010, Mizuno et\u00A0al., >>2003<<). We reasoned that this domain would be required for clustering; however, because mutations in the CR1 domain disrupt membrane binding (Figure\u00A04D, WT versus \u0394CR1), assessing clustering and segregation of a PAR-3\u0394CR1 mutant requires an" ; ns1:mentions . _:b66507511 rdf:type ns1:Context ; rdf:value "coincides temporally with a decrease in the prominence of PAR-3 clusters and an overall reduction in PAR-3 membrane localization (Figures 4A and 4C) as well as an increase in anterior CDC-42 activity (Figures S5C\u2013S5E) (Kumfer et\u00A0al., >>2010<<). We therefore speculated that the mix of\u00A0diffuse and clustered PAR-6/PKC-3 observed during establishment phase may reflect the distinct CDC-42- and PAR-3-dependent assemblies that we describe above." ; ns1:mentions . _:b66507512 rdf:type ns1:Context ; rdf:value "Depletion of CDC-42 is known to increase co-localization of PAR-6 with PAR-3 during polarity establishment (Beers and Kemphues, >>2006<<). We find that this also increases overall clustering of PAR-6 (Figures 5A and 5B)." ; ns1:mentions . _:b66507513 rdf:type ns1:Context ; rdf:value "To confirm that this reduction in clustering and segregation is due to favoring CDC-42-dependent assemblies, we examined the effect of expressing CDC-42 (Q61L), which stabilizes the active GTP-bound form of CDC-42 (Aceto et\u00A0al., >>2006<<, Ziman et\u00A0al., 1991)." ; ns1:mentions . _:b66507514 rdf:type ns1:Context ; rdf:value "this reduction in clustering and segregation is due to favoring CDC-42-dependent assemblies, we examined the effect of expressing CDC-42 (Q61L), which stabilizes the active GTP-bound form of CDC-42 (Aceto et\u00A0al., 2006, Ziman et\u00A0al., >>1991<<). Unlike PKC-3 inhibition, CDC-42 (Q61L) does not efficiently bypass the normal dependence of PKC-3 on PAR-3 (Figures S5F\u2013S5J)." ; ns1:mentions . _:b66507515 rdf:type ns1:Context ; rdf:value "Because PAR proteins are known to regulate actomyosin dynamics (Cheeks et\u00A0al., >>2004<<, Munro et\u00A0al., 2004), and changes in flow velocities could, in principle, affect advective transport (Goehring et\u00A0al., 2011b), we wanted to confirm that clustering rather than potential changes in flow velocity were the cause of reduced" ; ns1:mentions . _:b66507516 rdf:type ns1:Context ; rdf:value "Because PAR proteins are known to regulate actomyosin dynamics (Cheeks et\u00A0al., 2004, Munro et\u00A0al., >>2004<<), and changes in flow velocities could, in principle, affect advective transport (Goehring et\u00A0al., 2011b), we wanted to confirm that clustering rather than potential changes in flow velocity were the cause of reduced segregation" ; ns1:mentions . _:b66507517 rdf:type ns1:Context ; rdf:value "Because PAR proteins are known to regulate actomyosin dynamics (Cheeks et\u00A0al., 2004, Munro et\u00A0al., 2004), and changes in flow velocities could, in principle, affect advective transport (Goehring et\u00A0al., 2011b), we wanted to confirm that clustering rather than potential changes in flow velocity were the cause of reduced segregation efficiency." ; ns1:mentions . _:b66507518 rdf:type ns1:Context ; rdf:value "Finally, to test whether restoring flows could rescue efficient segregation of PKC-3, we\u00A0used RNAi to target the RhoGAPs, RGA-3/4, which results in excess actomyosin contractility and increased cortical flow rates (Fievet et\u00A0al., >>2012<<, Schonegg et\u00A0al., 2007)." ; ns1:mentions . _:b66507519 rdf:type ns1:Context ; rdf:value "restoring flows could rescue efficient segregation of PKC-3, we\u00A0used RNAi to target the RhoGAPs, RGA-3/4, which results in excess actomyosin contractility and increased cortical flow rates (Fievet et\u00A0al., 2012, Schonegg et\u00A0al., >>2007<<). Despite fully rescuing the moderate reduction in asymmetry of PAR-3 observed in pkc-3(ts) embryos to levels indistinguishable from wild-type, RGA-3/4 depletion failed to restore asymmetry of PKC-3 (Figures S6B and S6C)." ; ns1:mentions . _:b66507520 rdf:type ns1:Context ; rdf:value "While CDC-42 is generally thought to play an activating role (Atwood et\u00A0al., >>2007<<, Gotta et\u00A0al., 2001, Joberty et\u00A0al., 2000, Lin et\u00A0al., 2000, Qiu et\u00A0al., 2000), the roles for PAR-3 and PAR-6 are less clear and may vary in different contexts (Achilleos et\u00A0al., 2010, Atwood et\u00A0al., 2007, David et\u00A0al., 2013, Graybill et" ; ns1:mentions . _:b66507521 rdf:type ns1:Context ; rdf:value "While CDC-42 is generally thought to play an activating role (Atwood et\u00A0al., 2007, Gotta et\u00A0al., >>2001<<, Joberty et\u00A0al., 2000, Lin et\u00A0al., 2000, Qiu et\u00A0al., 2000), the roles for PAR-3 and PAR-6 are less clear and may vary in different contexts (Achilleos et\u00A0al., 2010, Atwood et\u00A0al., 2007, David et\u00A0al., 2013, Graybill et\u00A0al., 2012, Lin et\u00A0al." ; ns1:mentions . _:b66507522 rdf:type ns1:Context ; rdf:value "While CDC-42 is generally thought to play an activating role (Atwood et\u00A0al., 2007, Gotta et\u00A0al., 2001, Joberty et\u00A0al., >>2000<<, Lin et\u00A0al., 2000, Qiu et\u00A0al., 2000), the roles for PAR-3 and PAR-6 are less clear and may vary in different contexts (Achilleos et\u00A0al., 2010, Atwood et\u00A0al., 2007, David et\u00A0al., 2013, Graybill et\u00A0al., 2012, Lin et\u00A0al., 2000, McCaffrey and" ; ns1:mentions . _:b66507523 rdf:type ns1:Context ; rdf:value "While CDC-42 is generally thought to play an activating role (Atwood et\u00A0al., 2007, Gotta et\u00A0al., 2001, Joberty et\u00A0al., 2000, Lin et\u00A0al., >>2000<<, Qiu et\u00A0al., 2000), the roles for PAR-3 and PAR-6 are less clear and may vary in different contexts (Achilleos et\u00A0al., 2010, Atwood et\u00A0al., 2007, David et\u00A0al., 2013, Graybill et\u00A0al., 2012, Lin et\u00A0al., 2000, McCaffrey and Macara, 2009," ; ns1:mentions . _:b66507524 rdf:type ns1:Context ; rdf:value "While CDC-42 is generally thought to play an activating role (Atwood et\u00A0al., 2007, Gotta et\u00A0al., 2001, Joberty et\u00A0al., 2000, Lin et\u00A0al., 2000, Qiu et\u00A0al., >>2000<<), the roles for PAR-3 and PAR-6 are less clear and may vary in different contexts (Achilleos et\u00A0al., 2010, Atwood et\u00A0al., 2007, David et\u00A0al., 2013, Graybill et\u00A0al., 2012, Lin et\u00A0al., 2000, McCaffrey and Macara, 2009, Wirtz-Peitz et\u00A0al.," ; ns1:mentions . _:b66507525 rdf:type ns1:Context ; rdf:value "thought to play an activating role (Atwood et\u00A0al., 2007, Gotta et\u00A0al., 2001, Joberty et\u00A0al., 2000, Lin et\u00A0al., 2000, Qiu et\u00A0al., 2000), the roles for PAR-3 and PAR-6 are less clear and may vary in different contexts (Achilleos et\u00A0al., >>2010<<, Atwood et\u00A0al., 2007, David et\u00A0al., 2013, Graybill et\u00A0al., 2012, Lin et\u00A0al., 2000, McCaffrey and Macara, 2009, Wirtz-Peitz et\u00A0al., 2008)." ; ns1:mentions . _:b66507526 rdf:type ns1:Context ; rdf:value "role (Atwood et\u00A0al., 2007, Gotta et\u00A0al., 2001, Joberty et\u00A0al., 2000, Lin et\u00A0al., 2000, Qiu et\u00A0al., 2000), the roles for PAR-3 and PAR-6 are less clear and may vary in different contexts (Achilleos et\u00A0al., 2010, Atwood et\u00A0al., >>2007<<, David et\u00A0al., 2013, Graybill et\u00A0al., 2012, Lin et\u00A0al., 2000, McCaffrey and Macara, 2009, Wirtz-Peitz et\u00A0al., 2008)." ; ns1:mentions . _:b66507527 rdf:type ns1:Context ; rdf:value "et\u00A0al., 2007, Gotta et\u00A0al., 2001, Joberty et\u00A0al., 2000, Lin et\u00A0al., 2000, Qiu et\u00A0al., 2000), the roles for PAR-3 and PAR-6 are less clear and may vary in different contexts (Achilleos et\u00A0al., 2010, Atwood et\u00A0al., 2007, David et\u00A0al., >>2013<<, Graybill et\u00A0al., 2012, Lin et\u00A0al., 2000, McCaffrey and Macara, 2009, Wirtz-Peitz et\u00A0al., 2008)." ; ns1:mentions . _:b66507528 rdf:type ns1:Context ; rdf:value "al., 2001, Joberty et\u00A0al., 2000, Lin et\u00A0al., 2000, Qiu et\u00A0al., 2000), the roles for PAR-3 and PAR-6 are less clear and may vary in different contexts (Achilleos et\u00A0al., 2010, Atwood et\u00A0al., 2007, David et\u00A0al., 2013, Graybill et\u00A0al., >>2012<<, Lin et\u00A0al., 2000, McCaffrey and Macara, 2009, Wirtz-Peitz et\u00A0al., 2008)." ; ns1:mentions . _:b66507529 rdf:type ns1:Context ; rdf:value "et\u00A0al., 2000, Lin et\u00A0al., 2000, Qiu et\u00A0al., 2000), the roles for PAR-3 and PAR-6 are less clear and may vary in different contexts (Achilleos et\u00A0al., 2010, Atwood et\u00A0al., 2007, David et\u00A0al., 2013, Graybill et\u00A0al., 2012, Lin et\u00A0al., >>2000<<, McCaffrey and Macara, 2009, Wirtz-Peitz et\u00A0al., 2008)." ; ns1:mentions . _:b66507530 rdf:type ns1:Context ; rdf:value "2000, Qiu et\u00A0al., 2000), the roles for PAR-3 and PAR-6 are less clear and may vary in different contexts (Achilleos et\u00A0al., 2010, Atwood et\u00A0al., 2007, David et\u00A0al., 2013, Graybill et\u00A0al., 2012, Lin et\u00A0al., 2000, McCaffrey and Macara, >>2009<<, Wirtz-Peitz et\u00A0al., 2008)." ; ns1:mentions . _:b66507531 rdf:type ns1:Context ; rdf:value "the roles for PAR-3 and PAR-6 are less clear and may vary in different contexts (Achilleos et\u00A0al., 2010, Atwood et\u00A0al., 2007, David et\u00A0al., 2013, Graybill et\u00A0al., 2012, Lin et\u00A0al., 2000, McCaffrey and Macara, 2009, Wirtz-Peitz et\u00A0al., >>2008<<)." ; ns1:mentions . _:b66507532 rdf:type ns1:Context ; rdf:value "and CDC-42-associated assemblies in\u00A0vivo, we targeted PKC-3 to the membrane by fusing it to the C1B domain of human PKC\u03B1, which can be induced to bind the membrane by the addition of phorbol ester (Figure\u00A06A) (Lekomtsev et\u00A0al., >>2012<<). This bypasses the membrane-binding requirement of PKC-3 on PAR-3, PAR-6, and CDC-42, allowing us to test their contribution to PKC-3 activity by monitoring membrane removal of the PKC-3 target, PAR-2." ; ns1:mentions . _:b66507533 rdf:type ns1:Context ; rdf:value "The failure to completely remove PAR-2 in polarized zygotes is not simply due to PAR-2 being concentrated in a domain, because ectopic PAR-2 domains that form in meiotic arrest mutants, e.g., mei-1 and emb-27 (Wallenfang and Seydoux, >>2000<<), are rapidly cleared (Figure\u00A0S7)." ; ns1:mentions . _:b66507534 rdf:type ns1:Context ; rdf:value "Thus PAR-3 has two roles in vivo: it promotes PKC-3 membrane targeting while at the same time limiting its activation, reconciling in\u00A0vivo reports whereby PAR-3 positively regulates PAR polarity (Achilleos et\u00A0al., >>2010<<, McCaffrey and Macara, 2009) with data indicating that PAR-3 can inhibit PKC-3 activity in\u00A0vitro (Graybill et\u00A0al., 2012, Lin et\u00A0al., 2000, Soriano et\u00A0al., 2016)." ; ns1:mentions . _:b66507535 rdf:type ns1:Context ; rdf:value "has two roles in vivo: it promotes PKC-3 membrane targeting while at the same time limiting its activation, reconciling in\u00A0vivo reports whereby PAR-3 positively regulates PAR polarity (Achilleos et\u00A0al., 2010, McCaffrey and Macara, >>2009<<) with data indicating that PAR-3 can inhibit PKC-3 activity in\u00A0vitro (Graybill et\u00A0al., 2012, Lin et\u00A0al., 2000, Soriano et\u00A0al., 2016)." ; ns1:mentions . _:b66507536 rdf:type ns1:Context ; rdf:value "its activation, reconciling in\u00A0vivo reports whereby PAR-3 positively regulates PAR polarity (Achilleos et\u00A0al., 2010, McCaffrey and Macara, 2009) with data indicating that PAR-3 can inhibit PKC-3 activity in\u00A0vitro (Graybill et\u00A0al., >>2012<<, Lin et\u00A0al., 2000, Soriano et\u00A0al., 2016)." ; ns1:mentions . _:b66507537 rdf:type ns1:Context ; rdf:value "reconciling in\u00A0vivo reports whereby PAR-3 positively regulates PAR polarity (Achilleos et\u00A0al., 2010, McCaffrey and Macara, 2009) with data indicating that PAR-3 can inhibit PKC-3 activity in\u00A0vitro (Graybill et\u00A0al., 2012, Lin et\u00A0al., >>2000<<, Soriano et\u00A0al., 2016)." ; ns1:mentions . _:b66507538 rdf:type ns1:Context ; rdf:value "reports whereby PAR-3 positively regulates PAR polarity (Achilleos et\u00A0al., 2010, McCaffrey and Macara, 2009) with data indicating that PAR-3 can inhibit PKC-3 activity in\u00A0vitro (Graybill et\u00A0al., 2012, Lin et\u00A0al., 2000, Soriano et\u00A0al., >>2016<<)." ; ns1:mentions . _:b66507539 rdf:type ns4:Section ; dc:title "discussion" ; ns4:contains _:b66507552 , _:b66507553 , _:b66507554 , _:b66507555 , _:b66507556 , _:b66507557 , _:b66507544 , _:b66507545 , _:b66507546 , _:b66507547 , _:b66507548 , _:b66507549 , _:b66507550 , _:b66507551 , _:b66507540 , _:b66507541 , _:b66507542 , _:b66507543 . _:b66507540 rdf:type ns1:Context ; rdf:value "In previous work, we showed that the diffusion and membrane dissociation rates of aPARs were in principle sufficient to explain segregation in response to flows (Goehring et\u00A0al., 2011b). Here we show that segregation of aPARs is directly linked to PAR-3-dependent clustering." ; ns1:mentions . _:b66507541 rdf:type ns1:Context ; rdf:value "This fact, coupled with PAR-3 exclusion from the posterior by\u00A0PAR-1-dependent phosphorylation (Motegi et\u00A0al., >>2011<<), supports a model in which PAR-3 is responsible for sensing asymmetry-generating cues." ; ns1:mentions . _:b66507542 rdf:type ns1:Context ; rdf:value "Importantly, once it is asymmetric, PAR-3 provides a landmark for polarized loading of PAR-6/PKC-3, explaining recent observations that PAR-6 loads preferentially in the anterior of polarized embryos (Sailer et\u00A0al., >>2015<<)." ; ns1:mentions . _:b66507543 rdf:type ns1:Context ; rdf:value "Because the inhibitory role of PAR-3 appears to be broadly conserved (David et\u00A0al., >>2013<<, Graybill et\u00A0al., 2012, Lin et\u00A0al., 2000, McCaffrey et\u00A0al., 2012, Soriano et\u00A0al., 2016), this apparent paradoxical role of PAR-3 in promoting formation of membrane-associated aPKC complexes, yet\u00A0also suppressing PKC-3 activity, may be a" ; ns1:mentions . _:b66507544 rdf:type ns1:Context ; rdf:value "Because the inhibitory role of PAR-3 appears to be broadly conserved (David et\u00A0al., 2013, Graybill et\u00A0al., >>2012<<, Lin et\u00A0al., 2000, McCaffrey et\u00A0al., 2012, Soriano et\u00A0al., 2016), this apparent paradoxical role of PAR-3 in promoting formation of membrane-associated aPKC complexes, yet\u00A0also suppressing PKC-3 activity, may be a general feature of aPKC" ; ns1:mentions . _:b66507545 rdf:type ns1:Context ; rdf:value "Because the inhibitory role of PAR-3 appears to be broadly conserved (David et\u00A0al., 2013, Graybill et\u00A0al., 2012, Lin et\u00A0al., >>2000<<, McCaffrey et\u00A0al., 2012, Soriano et\u00A0al., 2016), this apparent paradoxical role of PAR-3 in promoting formation of membrane-associated aPKC complexes, yet\u00A0also suppressing PKC-3 activity, may be a general feature of aPKC regulation." ; ns1:mentions . _:b66507546 rdf:type ns1:Context ; rdf:value "Because the inhibitory role of PAR-3 appears to be broadly conserved (David et\u00A0al., 2013, Graybill et\u00A0al., 2012, Lin et\u00A0al., 2000, McCaffrey et\u00A0al., >>2012<<, Soriano et\u00A0al., 2016), this apparent paradoxical role of PAR-3 in promoting formation of membrane-associated aPKC complexes, yet\u00A0also suppressing PKC-3 activity, may be a general feature of aPKC regulation." ; ns1:mentions . _:b66507547 rdf:type ns1:Context ; rdf:value "Because the inhibitory role of PAR-3 appears to be broadly conserved (David et\u00A0al., 2013, Graybill et\u00A0al., 2012, Lin et\u00A0al., 2000, McCaffrey et\u00A0al., 2012, Soriano et\u00A0al., >>2016<<), this apparent paradoxical role of PAR-3 in promoting formation of membrane-associated aPKC complexes, yet\u00A0also suppressing PKC-3 activity, may be a general feature of aPKC regulation." ; ns1:mentions . _:b66507548 rdf:type ns1:Context ; rdf:value "Measurements elsewhere suggest that the distance these activated assemblies travel is on the order of 5\u201310\u00A0\u03BCm (Goehring et\u00A0al., 2011a, Robin et\u00A0al., 2014), consistent with the PKC-3 gradient extending further into the posterior than PAR-3 during the establishment phase even in wild-type embryos (see Figure\u00A02I)." ; ns1:mentions . _:b66507549 rdf:type ns1:Context ; rdf:value "Measurements elsewhere suggest that the distance these activated assemblies travel is on the order of 5\u201310\u00A0\u03BCm (Goehring et\u00A0al., 2011a, Robin et\u00A0al., >>2014<<), consistent with the PKC-3 gradient extending further into the posterior than PAR-3 during the establishment phase even in wild-type embryos (see Figure\u00A02I)." ; ns1:mentions . _:b66507550 rdf:type ns1:Context ; rdf:value "This is compatible with a model in which aPKC undergoes asymmetric membrane loading but symmetric dissociation (Robin et\u00A0al., >>2014<<)." ; ns1:mentions . _:b66507551 rdf:type ns1:Context ; rdf:value "During polarity establishment, when cortical flow is the major cue for anterior PAR segregation, PAR-3 clustering is prominent (Cheeks et\u00A0al., >>2004<<, Goehring et\u00A0al., 2011b, Munro et\u00A0al., 2004)." ; ns1:mentions . _:b66507552 rdf:type ns1:Context ; rdf:value "During polarity establishment, when cortical flow is the major cue for anterior PAR segregation, PAR-3 clustering is prominent (Cheeks et\u00A0al., 2004, Goehring et\u00A0al., 2011b, Munro et\u00A0al., 2004)." ; ns1:mentions . _:b66507553 rdf:type ns1:Context ; rdf:value "During polarity establishment, when cortical flow is the major cue for anterior PAR segregation, PAR-3 clustering is prominent (Cheeks et\u00A0al., 2004, Goehring et\u00A0al., 2011b, Munro et\u00A0al., >>2004<<). As the system enters the maintenance phase, flow ceases and continued aPAR segregation becomes dependent on the activity of PAR-1 and CHIN-1 (Beatty et\u00A0al., 2013, Guo and Kemphues, 1995, Kumfer et\u00A0al., 2010, Sailer et\u00A0al., 2015)." ; ns1:mentions . _:b66507554 rdf:type ns1:Context ; rdf:value "As the system enters the maintenance phase, flow ceases and continued aPAR segregation becomes dependent on the activity of PAR-1 and CHIN-1 (Beatty et\u00A0al., >>2013<<, Guo and Kemphues, 1995, Kumfer et\u00A0al., 2010, Sailer et\u00A0al., 2015)." ; ns1:mentions . _:b66507555 rdf:type ns1:Context ; rdf:value "As the system enters the maintenance phase, flow ceases and continued aPAR segregation becomes dependent on the activity of PAR-1 and CHIN-1 (Beatty et\u00A0al., 2013, Guo and Kemphues, >>1995<<, Kumfer et\u00A0al., 2010, Sailer et\u00A0al., 2015)." ; ns1:mentions . _:b66507556 rdf:type ns1:Context ; rdf:value "As the system enters the maintenance phase, flow ceases and continued aPAR segregation becomes dependent on the activity of PAR-1 and CHIN-1 (Beatty et\u00A0al., 2013, Guo and Kemphues, 1995, Kumfer et\u00A0al., >>2010<<, Sailer et\u00A0al., 2015)." ; ns1:mentions . _:b66507557 rdf:type ns1:Context ; rdf:value "As the system enters the maintenance phase, flow ceases and continued aPAR segregation becomes dependent on the activity of PAR-1 and CHIN-1 (Beatty et\u00A0al., 2013, Guo and Kemphues, 1995, Kumfer et\u00A0al., 2010, Sailer et\u00A0al., >>2015<<). Notably, clustering appears to be reduced during this phase, which Dickinson et\u00A0al. (2017) show is dependent on PLK-1. This change in PAR molecular organization potentially reflects the shift in the spatial signals to which the PAR" ; ns1:mentions . _:b66507558 rdf:type ns4:Section ; dc:title "star\u00E2\u0098\u0085methods" ; ns4:contains _:b66507592 , _:b66507593 , _:b66507588 , _:b66507589 , _:b66507590 , _:b66507591 , _:b66507584 , _:b66507585 , _:b66507586 , _:b66507587 , _:b66507580 , _:b66507581 , _:b66507582 , _:b66507583 , _:b66507576 , _:b66507577 , _:b66507578 , _:b66507579 , _:b66507572 , _:b66507573 , _:b66507574 , _:b66507575 , _:b66507568 , _:b66507569 , _:b66507570 , _:b66507571 , _:b66507564 , _:b66507565 , _:b66507566 , _:b66507567 , _:b66507560 , _:b66507561 , _:b66507562 , _:b66507563 , _:b66507559 . _:b66507559 rdf:type ns1:Context ; rdf:value "REAGENT or RESOURCESOURCEIDENTIFIERAntibodiesRabbit Anti-PAR-2(Dong et\u00A0al., >>2007<<)N/ARabbit Anti-PAR-6(Gotta et\u00A0al., 2001)N/ARat Anti-PKC-3(Tabuse et\u00A0al., 1998)N/AMouse Anti-PAR-3Developmental Studies Hybridoma BankP4A1; RRID:" ; ns1:mentions . _:b66507560 rdf:type ns1:Context ; rdf:value "REAGENT or RESOURCESOURCEIDENTIFIERAntibodiesRabbit Anti-PAR-2(Dong et\u00A0al., 2007)N/ARabbit Anti-PAR-6(Gotta et\u00A0al., >>2001<<)N/ARat Anti-PKC-3(Tabuse et\u00A0al., 1998)N/AMouse Anti-PAR-3Developmental Studies Hybridoma BankP4A1; RRID:" ; ns1:mentions . _:b66507561 rdf:type ns1:Context ; rdf:value "REAGENT or RESOURCESOURCEIDENTIFIERAntibodiesRabbit Anti-PAR-2(Dong et\u00A0al., 2007)N/ARabbit Anti-PAR-6(Gotta et\u00A0al., 2001)N/ARat Anti-PKC-3(Tabuse et\u00A0al., >>1998<<)N/AMouse Anti-PAR-3Developmental Studies Hybridoma BankP4A1; RRID: AB_528424Mouse Anti-\u03B1TubulinSigmaDM1A (T9026); RRID:" ; ns1:mentions . _:b66507562 rdf:type ns1:Context ; rdf:value "elegans: KK1063[lgl-1::gfp]: it256 [lgl-1::gfp\u00A0+ unc-119(+)]; unc-119(ed4) III; lgl-1(tm2616) X(Beatty et\u00A0al., >>2010<<)WB Strain:" ; ns1:mentions . _:b66507563 rdf:type ns1:Context ; rdf:value "elegans: OD70[mCherry::PH-PLC\u03941]: unc-119(ed3) III; ltIs44[pie-1p-mCherry::PH(PLC1\u03941) +unc-119(+)] V(Kachur et\u00A0al., >>2008<<)WB Strain:" ; ns1:mentions . _:b66507564 rdf:type ns1:Context ; rdf:value "elegans: SA131[gfp::cdc-42]: unc-119(ed3)III; tjIs 6[Ppie-1::gfp::cdc-42+unc-119(+)](Motegi and Sugimoto, >>2006<<)WB Strain:" ; ns1:mentions . _:b66507565 rdf:type ns1:Context ; rdf:value "elegans: TH129[gfp::par-2]: unc-119(ed3)III;ddIs25[GFP::F58B6.3;unc-119(+)];(Schonegg et\u00A0al., >>2007<<)TH129C.\u00A0elegans: TH159[mCherry-cdc-42]: unc-119(ed3)III; ddls46[WRM0625bA11 GLCherry::cdc-42; Cbr-unc-119(+)]Tony HymanTH159C.\u00A0elegans: TH209[mCherry::par-2]: unc-119(ed3)III; ddIs31[pie-1p::mCherry::par-2; unc-119(+)](Schonegg et\u00A0al.," ; ns1:mentions . _:b66507566 rdf:type ns1:Context ; rdf:value "elegans: TH209[mCherry::par-2]: unc-119(ed3)III; ddIs31[pie-1p::mCherry::par-2; unc-119(+)](Schonegg et\u00A0al., >>2007<<)TH209C.\u00A0elegans: TY3558[gfp::his-11; b-tubulin::gfp]: ruls[pie-1::GFPhis-11] III; ojIs1 [\u03B2-tubulin::GFP]CGCWB Strain: TY3558C.\u00A0elegans: UE37[pie-1::gfp]: axEx73 [pie-1p::pie-1::GFP\u00A0+ rol-6(su1006) + N2 genomic DNA]; tubulin mCherryCarrie" ; ns1:mentions . _:b66507567 rdf:type ns1:Context ; rdf:value "elegans: WH423[mCherry::cdc-42(Q61L)]: Ppie-1::mcherry::cdc-42(Q61L)(Kumfer et\u00A0al., >>2010<<)WH423C.\u00A0elegans: WH497[gfp::chin-1]: ojls69[pie-1::mGFP::chin-1\u00A0+ unc-119(+)]CGCWB Strain: WH497C.\u00A0elegans: WH517[gfp::wsp-1]: ojIs40 [Ppie-1::gfp::GBDwsp-1\u00A0+ unc-119(+)]CGCWB Strain: WH517C.\u00A0elegans: WM150[pkc-3(ts)]: pkc-3(ne4246)" ; ns1:mentions . _:b66507568 rdf:type ns1:Context ; rdf:value "elegans: WM150[pkc-3(ts)]: pkc-3(ne4246) II(Fievet et\u00A0al., >>2012<<)WM150C.\u00A0elegans:" ; ns1:mentions . _:b66507569 rdf:type ns1:Context ; rdf:value "elegans: WS5018[gfp::cdc-42]: cdc-42(gk388); opIs295 [cdc-42p::gfp::cdc-42(genomic)::cdc-42 3'UTR\u00A0+ unc-119(+)] II.(Neukomm et\u00A0al., >>2014<<)WB Strain: WS5018Oligonucleotidespkc-3(genomic) fwd:" ; ns1:mentions . _:b66507570 rdf:type ns1:Context ; rdf:value "Feeding RNAi: perm-1Source BioScienceWB Clone: sjj_T01H3.4Ahringer Feeding RNAi: rga-3Source BioScienceWB Clone: sjj_K09H11.3Ahringer Feeding RNAi: cgef-1Source BioScienceWB Clone: sjj_C14A11.3Feeding RNAi: mlc-4(Redemann et\u00A0al., >>2010<<)N/AFeeding RNAi: control(ctl)This paperN/APH-GBP gBlock (sequence on request)IDT DNAN/Actl (RNAi):" ; ns1:mentions . _:b66507571 rdf:type ns1:Context ; rdf:value "elegans strains were maintained on nematode growth media (NGM) under standard conditions (Brenner, >>1974<<) at 16\u00B0C or 20\u00B0C unless otherwise indicated." ; ns1:mentions . _:b66507572 rdf:type ns1:Context ; rdf:value "Following the scheme of (Lekomtsev et\u00A0al., >>2012<<), a codon-optimized (Redemann et\u00A0al., 2011) sequence encoding the C1B domain from human PKC\u03B1 (GenScript) was inserted into pTH699 via BamHI and SmaI to generate a sfgfp:" ; ns1:mentions . _:b66507573 rdf:type ns1:Context ; rdf:value "Following the scheme of (Lekomtsev et\u00A0al., 2012), a codon-optimized (Redemann et\u00A0al., >>2011<<) sequence encoding the C1B domain from human PKC\u03B1 (GenScript) was inserted into pTH699 via BamHI and SmaI to generate a sfgfp:" ; ns1:mentions . _:b66507574 rdf:type ns1:Context ; rdf:value "Both plasmids were introduced by biolistic bombardment into HT1593 worms (Praitis et\u00A0al., >>2001<<), yielding NWG0012 and NWG0016." ; ns1:mentions . _:b66507575 rdf:type ns1:Context ; rdf:value "The membrane-tethered GFP-binding protein (PH-GBP) was generated by combining amino acids 1-175 corresponding to the PH domain of rat PH-PLC\u03941 (Audhya et\u00A0al., >>2005<<) and VHH4GFP (Caussinus et\u00A0al., 2011) coupled by a SGQGGSGGSGGS linker. The resulting sequence was codon-optimized (CAI\u00A0= 0.49) and a single GFP intron inserted as described (Redemann et\u00A0al., 2011)." ; ns1:mentions . _:b66507576 rdf:type ns1:Context ; rdf:value "The membrane-tethered GFP-binding protein (PH-GBP) was generated by combining amino acids 1-175 corresponding to the PH domain of rat PH-PLC\u03941 (Audhya et\u00A0al., 2005) and VHH4GFP (Caussinus et\u00A0al., >>2011<<) coupled by a SGQGGSGGSGGS linker. The resulting sequence was codon-optimized (CAI\u00A0= 0.49) and a single GFP intron inserted as described (Redemann et\u00A0al., 2011)." ; ns1:mentions . _:b66507577 rdf:type ns1:Context ; rdf:value "The resulting sequence was codon-optimized (CAI\u00A0= 0.49) and a single GFP intron inserted as described (Redemann et\u00A0al., >>2011<<). A synthetic gBlock (IDT DNA) encoding the PH-GBP was PCR amplified and cloned in frame with a C-terminal codon-optimized mKate2 under the control of the mex-5 promoter and nmy-2 3\u2019 UTR in a MosSCI vector containing wild-type unc-119" ; ns1:mentions . _:b66507578 rdf:type ns1:Context ; rdf:value "The resulting plasmid (pNG0018) was inserted at the ttTi5605 mos1 site locus of DP38 worms via CRISPR after mutating the sgRNA/PAM site following the method described (pNG0019) (Dickinson et\u00A0al., >>2013<<). Modified worms were crossed into DR466 to generate a stable male line expressing PH-GBP (NWG0047). To rescue membrane localization of PAR-3 variants, we crossed NWG0047 with KK1216 (par-3::gfp) or KK973 (par-3\u0394cr1::gfp) lines. We were" ; ns1:mentions . _:b66507579 rdf:type ns1:Context ; rdf:value "HEK-293 are female and were obtained from Cell Production, Cancer Research UK (CRUK) and cultivated in DMEM (Dulbecco\u2019s modified Eagle\u2019s medium), 10% FBS (fetal bovine serum) and penicillin\u2013streptomycin (Invitrogen) (Kj\u00E6r et\u00A0al., >>2013<<)." ; ns1:mentions . _:b66507580 rdf:type ns1:Context ; rdf:value "RNAi by feeding was performed similar to described methods (Kamath et\u00A0al., >>2003<<). Briefly, HT115(DE3) bacterial feeding clones were inoculated from LB agar plates to LB liquid cultures and grown overnight at 37\u00B0C in the presence of 10\u00A0\u03BCg/ml carbenicillin." ; ns1:mentions . _:b66507581 rdf:type ns1:Context ; rdf:value "par-3, par-6, cdc-42, pkc-3, perm-1, rga-3/4, cgef-1 and emb-27 clones are from the Ahringer library (Kamath et\u00A0al., >>2003<<). mlc-4 is from Redemann et\u00A0al., 2010. A control RNAi clone was generated by synthesizing a random 500bp sequence using the Matlab random number generator with no homology to the worm genome, cloned into Bgl-II / HindIII sites of L4440" ; ns1:mentions . _:b66507582 rdf:type ns1:Context ; rdf:value "par-3, par-6, cdc-42, pkc-3, perm-1, rga-3/4, cgef-1 and emb-27 clones are from the Ahringer library (Kamath et\u00A0al., 2003). mlc-4 is from Redemann et\u00A0al., >>2010<<. A control RNAi clone was generated by synthesizing a random 500bp sequence using the Matlab random number generator with no homology to the worm genome, cloned into Bgl-II / HindIII sites of L4440 (Addgene, plasmid#1654), and transformed" ; ns1:mentions . _:b66507583 rdf:type ns1:Context ; rdf:value "All drug treatment experiments were performed in 10 to 50% perm-1(RNAi) (Carvalho et\u00A0al., >>2011<<). Drugs were dissolved in DMSO and used at the following concentrations: phorbol 12-myristate 13-acetate (PMA, Sigma-Aldrich, P1585-1MG), 100\u00A0\u03BCM; CRT90 (CRT0103390, Cancer Research Technology LTD), 10\u00A0\u03BCM." ; ns1:mentions . _:b66507584 rdf:type ns1:Context ; rdf:value "Immunofluorescence was performed as previously described (Andrews and Ahringer, >>2007<<). Briefly, gravid hermaphrodite worms were washed and then transferred to a 7\u00A0\u03BCl drop of M9 on a 0.1% poly-lysine coated well." ; ns1:mentions . _:b66507585 rdf:type ns1:Context ; rdf:value "Embryos were dissected in 2-4\u00A0\u03BCl of M9 buffer (22 mM KH2PO4, 42 mM NaHPO4, 86 mM NaCl and 1 mM MgSO4) on a coverslip and mounted under 2% agarose pads (Zipperlen et\u00A0al., >>2001<<) or dissected in Shelton\u2019s Growth Medium (Edgar and Goldstein, 2012) and mounted with 16-21\u00A0\u03BCm polystyrene beads between a slide and coverslip and sealed with VALAP (Goehring et\u00A0al., 2011a)." ; ns1:mentions . _:b66507586 rdf:type ns1:Context ; rdf:value "were dissected in 2-4\u00A0\u03BCl of M9 buffer (22 mM KH2PO4, 42 mM NaHPO4, 86 mM NaCl and 1 mM MgSO4) on a coverslip and mounted under 2% agarose pads (Zipperlen et\u00A0al., 2001) or dissected in Shelton\u2019s Growth Medium (Edgar and Goldstein, >>2012<<) and mounted with 16-21\u00A0\u03BCm polystyrene beads between a slide and coverslip and sealed with VALAP (Goehring et\u00A0al., 2011a)." ; ns1:mentions . _:b66507587 rdf:type ns1:Context ; rdf:value "under 2% agarose pads (Zipperlen et\u00A0al., 2001) or dissected in Shelton\u2019s Growth Medium (Edgar and Goldstein, 2012) and mounted with 16-21\u00A0\u03BCm polystyrene beads between a slide and coverslip and sealed with VALAP (Goehring et\u00A0al., 2011a). 16-18\u00A0\u03BCm beads were used for cortex imaging to maximize imaging surface. In all other cases, 21\u00A0\u03BCm beads were used to minimize compression effects on development." ; ns1:mentions . _:b66507588 rdf:type ns1:Context ; rdf:value "For C1B targeting experiments, two sides of the coverslip were left unsealed to create a flow chamber (Goehring et\u00A0al., 2011a) and PMA washed in at the indicated times." ; ns1:mentions . _:b66507589 rdf:type ns1:Context ; rdf:value "All image analysis was performed in Fiji (ImageJ)(Schindelin et\u00A0al., >>2012<<) and Matlab (Mathworks)." ; ns1:mentions . _:b66507590 rdf:type ns1:Context ; rdf:value "m script, part of the feature detection and particle tracking package from the Kilfoil Lab (Pelletier et\u00A0al., >>2009<<). Embryos were automatically detected and partitioned into 3 domains (Anterior, Middle, Posterior) and normalized anterior vs posterior particle densities used for ASI calculation." ; ns1:mentions . _:b66507591 rdf:type ns1:Context ; rdf:value "Costes\u2019 Mask and intensity correlation quotient (Li et\u00A0al., >>2004<<) were obtained using JaCOP plug-in in Fiji." ; ns1:mentions . _:b66507592 rdf:type ns1:Context ; rdf:value "Using the Kymograph Plugin in Fiji (Seitz and Surrey, >>2006<<), we generated kymographs for individual embryos by tracing a segmented line along the cortex starting at the origin of flow." ; ns1:mentions . _:b66507593 rdf:type ns1:Context ; rdf:value "Measurement of cortical flow in wild-type embryos expressing GFP fusions to PAR-3, PKC-3 or CDC-42 yielded a velocity of 7.1+/-1.4\u00A0\u03BCm/min (n=22), consistent with previously published values (7.66+/-1.0\u00A0\u03BCm/min, n=6)(Munro et\u00A0al., >>2004<<)." ; ns1:mentions . _:b633036173 rdf:type ns1:RelevantBibliographicResource . @prefix xsd: . _:b633036173 ns1:RelevantScore "22"^^xsd:nonNegativeInteger ; 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