ライフサイエンスコーパス: P granule

1 ermline and is a component of germ granules (P granules).

2 r side of nuclear pores, and associated with P granules.

3 trate that nascent mRNA traffics directly to P granules.

4 including C. elegans, where they are called P granules.

5 RNA helicase and a constitutive component of P granules.

6 ocalizes to germline nuage structures called P granules.

7 the stability, localization, and function of P granules.

8 sm and to ribonucleoprotein particles called P granules.

9 viously identified as critical components of P granules.

10 nd localizes to nuage-like structures called P granules.

11 ctures normally limited to germline-specific P granules.

12 at least some Sm proteins are components of P granules.

13 nt to send homogenously distributed mRNAs to P granules.

14 but does not affect the initial assembly of P granules.

15 RNA, are present at least transiently within P granules.

16 mplex accompanied by the mis-localization of P granules.

17 elegans contain distinctive granules called P granules.

18 of which contribute to localized assembly of P granules.

19 germline blastomeres, and is a component of P granules.

20 rotein PGL-3 with RNA drives the assembly of P granules.

21 mponent of germline-specific granules called P granules.

22 he surface of PGL condensates at the core of P granules.

23 ranscripts by Argonaute proteins enriched in P granules.

24 s and in the posterior cytoplasm surrounding P granules.

25 protein that forms low dynamic assemblies on P granules.

26 d to promote the perinuclear localization of P granules.

27 s alongside the piRNA and siRNA machinery at P granules.

28 d in phase separation and a key component of P granules.

29 -seq on dissected germlines with and without P granules.

30 highly enriched at the NPCs associated with P granules.

31 he helicase DRH-3, act to antagonize RNA and P-granule accumulation in the germ line.

32 asymmetry of cellular components, including P granules, also control GLP-1 asymmetry in the early em

33 ind and anchor the Vasa homolog GLH-1 within P granules and are jointly required for coalescence of M

34 of vertebrate Nup98 (CeNup98) is enriched in P granules and associates with the translationally repre

35 n-containing DEAD box helicase, localizes in P granules and cytoplasmic foci that are enriched in RSD

36 is required to localize FBF-2 to perinuclear P granules and for efficient binding of FBF-2 to its mRN

37 RDE-12 colocalizes with WAGO-1 in germline P granules and in cytoplasmic and perinuclear foci in so

38 efective)-3 and MEG-4, which are enriched in P granules and in the posterior cytoplasm surrounding P

39 n at the two-cell stage, and localization of P granules and MEX-5 during the first and subsequent cel

40 zes to distinct subcellular foci adjacent to P granules and Mutator foci, two phase-separated condens

41 illance, may serve to tune mRNA flow through P granules and other nuage condensates.

42 rmline and early embryo, and is localized to P granules and other possible mRNA-protein particles.

43 P granules and other RNA/protein bodies are membrane-les

44 asymmetrically localized proteins, including P granules and PAR-3, are normal in early let-99 embryos

45 -sized blastomeres, improper localization of P granules and SKN-1 protein, and abnormal second divisi

46 Our observations reveal similarities between P granules and stress granules and identify intrinsicall

47 f diverse membraneless organelles, including P granules and stress granules in the cytoplasm and nucl

48 tress causes additional mRNAs to localize to P granules and translational activation correlates with

49 -2 mutant worms, and knockdown of individual P-granule and other germ-line genes in daf-2 young adult

50 rulation from a maternal RNA associated with P granules, and is required for the efficient incorporat

51 By injecting a fluorescently labelled anti-P-granule antibody into the C. elegans germline syncitiu

52 P granules appear to sequester large amounts of mRNA in

53 on of nuclear pore complexes and perinuclear P granules are altered in the absence of EGO-1, effects

54 most of the C. elegans life cycle, however, P granules are associated with clusters of nuclear pore

55 However, during most of development P granules are associated with germ cell nuclei.

56 In the cytoplasm, germ-line or P granules are circulated by an actomyosin-driven founta

57 C. elegans P granules are conserved cytoplasmic ribonucleoprotein c

58 P granules are cytoplasmic bodies in oocytes and early e

59 P granules are cytoplasmic structures of unknown functio

60 Caenorhabditis elegans, germ granules called P granules are directly inherited from mother to daughte

61 P granules are germ-cell-specific cytoplasmic structures

62 In Caenorhabditis elegans, P granules are germline-specific, RNA-containing granule

63 ot increase in the young adult germline when P granules are impaired.

64 zation has focused on the early embryo, when P granules are located in the cytoplasm.

65 de that, despite their liquid-like behavior, P granules are non-homogeneous structures whose assembly

66 P granules are non-membrane-bound organelles found in th

67 P granules are non-membrane-bound RNA-protein compartmen

68 nrich maternal transcripts, but surprisingly P granules are not essential for germ cell fate specific

69 Nematode P granules are paradigmatic liquid droplet granules and

70 During most of germline development, P granules are perinuclear and associate with clusters o

71 P granules are perinuclear condensates in C. elegans ger

72 Cytoplasmic structures called P granules are present in the fertilized egg and are seg

73 anule segregation challenges the belief that P granules are responsible for determining the germline

74 P granules are RNA granules that form in the posterior o

75 P granules are RNA-rich protein condensates that share t

76 During early embryogenesis, P granules are segregated asymmetrically into those blas

77 e are no signs of cytoplasmic movements, and P granules are segregated differently.

78 P granules are still present in glh-1/4(RNAi) sterile wo

79 germ granule" counterparts in other animals, P granules are thought to act as determinants of the ide

80 We show that P granules are tightly associated with nuclear pores and

81 oncentration gradient and the segregation of P granules are two parallel manifestations of MEG-3's re

82 bution and the development of the germ line, P granules are widely thought to function in some aspect

83 of the Caenorhabditis elegans zygote, germ (P) granules are disassembled in the anterior cytoplasm a

84 ns are nearly equal in size, and cytoplasmic P-granules are not properly localized to germline precur

85 orhabditis elegans, these structures, termed P granules, are partitioned to the germline P cells duri

86 In C. elegans, germ granules, called P granules, are segregated to the germline precursor cel

87 Using nematode "P-granules" as a paradigm, we focus on the PGL granule s

88 Caenorhabditis elegans germ granule, called P granule, as a model system, we employed a proximity-ba

89 elles, such as the endoplasmic reticulum and P granules, as vectors for the segregation of informatio

90 legans zygote, germline RNA granules, called P granules, assemble preferentially in the posterior cyt

91 Our findings provide strong evidence that P granule assembly and disassembly are governed by phase

92 estigate the functional relationship between P granule assembly and function.

93 PGL-1 dimerization lies at the heart of both P granule assembly and function.

94 strate that LAF-1 is important for promoting P granule assembly and provide insight into the mechanis

95 utations of those interface residues prevent P granule assembly in vivo, de-repress PGL-1 tethered mR

96 ilar spatial and temporal characteristics to P granule assembly in vivo.

97 ompetition mechanism can drive a gradient of P granule assembly with similar spatial and temporal cha

98 en identified factors that are essential for P granule assembly, notably EGGD-1 and EGGD-2, two putat

99 ndicating the GLHs function independently in P granule assembly.

100 oteins as drivers of RNA condensation during P granule assembly.

101 In Caenorhabditis elegans embryos, germ (P) granule assembly requires MEG-3, an intrinsically dis

102 transport, and mRNA homeostasis converge on P-granule assembly and function.

103 ins with P granules, suggesting a pathway of P-granule assembly in which the GLHs are upstream of the

104 uggest that DEPS-1 is a key component of the P-granule assembly pathway and that its roles include pr

105 dimer forms a fundamental building block for P-granule assembly.

106 ars to operate in later P cells (P2 and P3): P granules associate with the nucleus and move with it t

107 ously within cells, localizing to membranes, P granules (associated with progenitor germ cells in the

108 Finally, we identify the P granule-associated Argonaute WAGO-1 as crucial for rep

109 We have identified a new P-granule-associated protein, DEPS-1, the loss of which

110 We present evidence that P granule asymmetry depends on RNA-induced phase separat

111 ntribute significantly to the maintenance of P granule asymmetry.

112 suppressing MEG-3 phase separation to drive P granule asymmetry.

113 Both GLH proteins localize in the P granules at all stage of germ-line development.

114 omplex localizes to Mutator foci adjacent to P granules at the nuclear periphery in germ cells.

115 In the C. elegans germline, cytoplasmic P granules at the nuclear pores and perinuclear Mutator

116 s not required for posterior localization of P granules at the one-cell stage, it is required for pro

117 encoding a key autophagy regulator (ectopic P-granules autophagy protein 5) implicated in the format

118 e have identified a mutant (pptr-1) in which P granules become unstable during mitosis and P granule

119 escuing GFP::alg-3 transgene is localized to P granules beginning at the late pachytene stage of male

120 The Mutator foci are adjacent to P granules but are not dependent on core P-granule compo

121 Thus, RNAi likely does not require intact P granules but instead relies on particular P-granule fa

122 heavily targeted by piRNAs and accumulate in P granules but maintain expression.

123 st cause and effect, we severely compromised P granules by simultaneously knocking down factors that

124 cal and ultrastructural data suggesting that P granules can associate, or remain associated, with por

125 We found that compromising P granules causes germ cells to express neuronal and mus

126 Instead, we found that impairing P granules causes sperm-specific mRNAs to become highly

127 FBF-2 activity depends on the P granule component PGL-1.

128 is present in germline blastomeres and is a P granule component, although MEX-1 is a cytoplasmic pro

129 lting defects, is identified here as a novel P-granule component and a binding partner of GLH-1 (Germ

130 rst larval stage onward and is unusual for a P-granule component in lacking recognizable RNA binding

131 Loss of the P-granule component pgl-1 in meg-1 mutants increases ger

132 his report demonstrates that PAN-1 is also a P-granule component that is essential for fertility.

133 is dependent on downstream RNAi factors and P granule components but is independent of the viral sen

134 t is thought that asymmetric partitioning of P granule components during mitosis is essential to dist

135 Despite symmetric partitioning of P granule components, pptr-1 mutants segregate a germlin

136 While loss of some P-granule components causes worms to be defective in RNA

137 to P granules but are not dependent on core P-granule components or other RNAi pathway factors for t

138 ression of target mRNA and requires germline P-granule components.

139 e proteins GLH-1, GLH-2, and GLH-4 and other P-granule components.

140 his question, we investigate the response of P granule condensates in living cells to temperature cha

141 These proteins promote P granule condensation, form granules independently of M

142 together, our work expands the repertoire of P granule constituents and provides new insights into th

143 All the known protein components of P granules contain putative RNA-binding motifs, suggesti

144 e positions of certain structures within the P granules correspond to the positions of pores on the n

145 port that co-clustering of nuclear pores and P granules depends on FG repeat-containing nucleoporins

146 Localization of MEG-1 to P granules depends upon the membrane-bound protein MES-1

147 We show that perinuclear P granules differ from cytoplasmic P granules in many re

148 We observe that P granules dissolve upon increasing the temperature and

149 Because of the correlation between P granule distribution and the development of the germ l

150 flow is also lost in these embryos, although P granules do become localized to the posterior pole aft

151 asymmetric cortical contractions are absent, P granules do not localize, and cortical PAR-3 does not

152 e family, which encode protein components of P granules, do not appear essential for RNA to concentra

153 C. elegans germ granules, known as P granules, do not appear to be required for primordial

154 :Halo in the posterior cytoplasm surrounding P granules does not appear to contribute significantly t

155 encode proteins that localize exclusively to P granules during embryonic germline segregation.

156 segregate a germline that uniquely expresses P granules during postembryonic development.

157 Asymmetric segregation of P granules during the first four divisions of the Caenor

158 accelerate spatially regulated growth of the P granule emulsion.

159 In Caenorhabditis elegans embryos, P granules enrich maternal transcripts, but surprisingly

160 sociates with the translationally repressed, P granule-enriched mRNA nos-2 (nanos homolog).

161 and translational activation correlates with P granule exit for two mRNAs coding for germ cell fate r

162 Our results suggest that P granules extend the NPC environment in the germ line a

163 Our findings also support the view that P granules facilitate mRNA silencing by providing an env

164 P granules but instead relies on particular P-granule factors.

165 This is suggested by the loss of P granules from germ cells that transform into somatic c

166 In the germline, PAN-1 uniquely localizes to P granules from the first larval stage onward and is unu

167 f eggd-1 and eggd-2 results in separation of P granules from the nuclear envelope, germline atrophy,

168 We conclude that P granule function requires both assembly and localized

169 CAR-1 localizes to P-granules (germ-line specific ribonucleoprotein particl

170 m line, and balance each other in regulating P granule growth and localization.

171 hway engages with the membraneless organelle P granule in Caenorhabditis elegans.

172 erine-rich proteins regulate the dynamics of P granules in C. elegans embryos.

173 5 regulates the formation and dissolution of P granules in Caenorhabditis elegans embryos, yet the th

174 is the presence of germ granules, including P granules in Caenorhabditis elegans, which are typicall

175 LHs, are components of the germline-specific P granules in Caenorhabditis elegans.

176 (A) polymerase (PAP) that is associated with P granules in early embryos.

177 ng previous studies showing that cytoplasmic P granules in embryos contain RNA.

178 the Piwi-like protein PRG-1 is localized to P granules in germ cells entering spermatogenesis and is

179 rinuclear P granules differ from cytoplasmic P granules in many respects, including structure, stabil

180 required for proper cortical association of P granules in P1.

181 mutant (meg-3 meg-4) that does not assemble P granules in primordial germ cells loses competence for

182 servations support a "safe harbor" model for P granules in protecting germline transcripts from piRNA

183 kdown of LAF-1 results in the dissolution of P granules in the early embryo, with an apparent submicr

184 tructural analysis of the nuclear-associated P granules in the germ cells of the adult hermaphrodite

185 rotein- and RNA-rich nuclear pore-associated P granules in the germline, where they are thought to su

186 Finally, we show that nuclear-associated P granules in the gonad contain RNA, complementing previ

187 o specific mRNAs have been identified within P granules in the gonad.

188 titutive components of the germline-specific P granules in the nematode Caenorhabditis elegans and ar

189 A well-studied example of condensates is P granules in the roundworm Caenorhabditis elegans that

190 nules (for example, polar granules in flies, P granules in worms) are ribonucleoprotein particles imp

191 Germ granules (P granules) in C. elegans are required for fertility and

192 roteins that associate with germ-line nuage (P granules), including the Piwi-clade argonaute PRG-1, h

193 ntified a specific nucleoporin essential for P granule integrity and function.

194 y, we demonstrate that localized assembly of P granules is controlled by MEG-3, an intrinsically diso

195 ld elevated and the organization of GLH-1 in P granules is grossly disrupted.

196 Hence, asymmetric partitioning of maternal P granules is not essential to specify germ cell fate.

197 Localization to P granules is not required for translational repression

198 Sterility in the absence of P granules is often accompanied by the misexpression of

199 it has not been established whether loss of P granules is the cause or effect of cell fate transform

200 One potential role for P granules is to maintain germline fate and totipotency.

201 known location of the germline determinants, P granules, leading us to speculate that they may be ass

202 We demonstrate that P granules, like NPCs, are held together by weak hydroph

203 We reconstitute P granule-like droplets in vitro using a single protein

204 rhabditis elegans protein LAF-1, which forms P granule-like droplets in vitro.

205 se found in P granules, phase separates into P granule-like droplets in vitro.

206 found that translational repression preceded P granule localization and could occur independently of

207 loss of SmE function also caused defects in P granule localization and premature division in early g

208 Germline P granule localization at the time of cytoplasmic flow i

209 disruption of Sm activity caused defects in P granule localization to the germ cell precursors durin

210 lations between low translational status and P granule localization within the progenitor germ lineag

211 ut neurite-like projections, suggesting that P granules maintain totipotency and germline identity by

212 1 mutants exhibit multiple germline defects: P-granule mis-segregation in embryos, underproliferation

213 We discover a toroidal P granule morphology, which encircles the other germ gra

214 in embryos, underproliferation and aberrant P-granule morphology in larval germ cells, and ultimatel

215 on of P(2) and P(3) to which the spindle and P granules must move to ensure normal division asymmetry

216 are comprised of sub-domains referred to as P granules, Mutator foci, Z granules, and SIMR foci.

217 We show here that P granules normally contain a low level of RNA, and desc

218 However, we did not find evidence that P granules normally sequester aberrant mRNAs, or mRNAs t

219 Studying the P granules of Caenorhabditis elegans, we find that conde

220 y, cytoplasmic and colocalizes with PGL-1 in P granules of germline precursor cells.

221 Two components of the germ-line-specific P granules of the nematode Caenorhabditis elgans have be

222 ally controlled segregation of the germ line P granules or the pattern of cell division through the f

223 ulin mRNA and rRNA are either not present in P granules, or are present at relatively low levels.

224 In the Caenorhabditis elegans germline, P granules overlay large clusters of nuclear pores and t

225 We show that, in early P cells (P0 and P1), P-granule partitioning is achieved primarily by their mi

226 protein LAF-1, a DDX3 RNA helicase found in P granules, phase separates into P granule-like droplets

227 genes fall into specific classes with shared P-granule phenotypes, allowing us to better understand h

228 ressed gene mex-1 disrupt the segregation of P granules, prevent the formation of germ cells, and cau

229 e reversed orientation of polarity proteins, P granules, pronuclei migration and asymmetric cell divi

230 These findings suggest that P granules protect germline integrity through two differ

231 reveals an interaction with the constitutive P granule protein DEPS-1.

232 the intrinsically disordered RGG domain from P granule protein LAF-1.

233 xperiment was also found for RtoK mutants of P-granule protein LAF-1, underscoring that, to a degree,

234 iation of IFE-1 with P granules requires the P-granule protein PGL-1.

235 icase-1), a constitutive, germline-specific, P-granule protein.

236 granules become unstable during mitosis and P granule proteins and RNAs are distributed equally to s

237 repeat domains are found in the VASA-related P-granule proteins GLH-1, GLH-2, and GLH-4 and other P-g

238 (i) We could not detect P-granule proteins in the somatic cells of daf-2 mutants

239 nts ectopically misexpress germline-specific P-granule proteins in their somatic cells, suggesting a

240 (iii) Simultaneous removal of multiple P-granule proteins or the entire germ-line program from

241 eat domain is found in several nucleolar and P-granule proteins, but how it influences their phase se

242 Our results suggest that nuclear-associated P granules provide a perinuclear compartment where newly

243 We find that P granules recruit mRNAs by condensation with the disord

244 The association of IFE-1 with P granules requires the P-granule protein PGL-1.

245 P granules, ribonucleoprotein (RNP) complexes specific t

246 aracterize the transcriptome and assembly of P granules, RNA granules in the C. elegans germ plasm.

247 Here we analyze a key P granule scaffolding protein, PGL-1, to investigate the

248 the regulatory subunit of phosphatase 2A, in P granule segregation challenges the belief that P granu

249 P granule segregation depends on MEG (maternal-effect ge

250 Our data suggests AIR-1 plays a role in P-granule segregation and the association of the germlin

251 e C. elegans germline syncitium, we followed P-granule segregation in live embryos using laser-scanni

252 mes-1 are required for the normal pattern of P-granule segregation in P2.

253 n addition we find that Dcp2 is localized to P-granules, showing that Dcp2 is stored and/or active in

254 eats in C. elegans and uncover nucleolar and P-granule-specific RG/RGG motifs.

255 Most of the analysis of P granule structure and localization has focused on the

256 protein, DEPS-1, the loss of which disrupts P-granule structure and function.

257 sociation of the PGL family of proteins with P granules, suggesting a pathway of P-granule assembly i

258 on of PIE-1 remarkably parallels that of the P granules, suggesting that the localizations of these t

259 enters for ribonucleoprotein networks within P granules that help recruit and balance essential RNA p

260 is also disassembly or degradation of those P granules that remain in the cytoplasm destined for the

261 ired for the proper localization of PGL-1 to P granules, the accumulation of glh-1 mRNA and protein,

262 A core component of P granules, the endo-siRNA-binding Argonaute protein CSR

263 t appear essential for RNA to concentrate in P granules; these proteins may instead function in event

264 ong-standing question: Are mRNAs directed to P granules to be translationally repressed, or do they a

265 Loss of CeNup98 causes P granules to disperse in the cytoplasm and to release n

266 that segregates critical factors such as the P granules to one side of the uncleaved embryo [3,4].

267 Instead, we found that CSR-1 functions with P granules to prevent MSP and sperm-specific mRNAs from

268 ants by immunostaining or by expression of a P-granule transgene.

269 The germ-line-specific P granules were also mislocalized at the two-cell stage.

270 ding components of C. elegans germ granules (P granules) were up-regulated in daf-2 mutant worms, and

271 tative ATP-dependent enzymes localize to the P granules, which are nonmembranous complexes of protein

272 eveloping mitotic spindle and its associated P granules within P(2) and P(3), or provides an orientat

273 Therefore, understanding how P granules work is critical to understanding how cellula

274 s of perinuclear germ granule compartments - P granules, Z granules, SIMR foci, and Mutator foci - mu