메뉴 건너뛰기




Volumn 108, Issue 1, 2012, Pages 68-74

Chromosomal dynamics of nucleolar organizer regions (NORs) in the house mouse: Micro-evolutionary insights

Author keywords

house mouse; mus musculus; NOR; phylogeny; radiation; rDNA

Indexed keywords

ADAPTIVE RADIATION; CHROMOSOME; COLONIZATION; EVOLUTIONARY BIOLOGY; GEOGRAPHICAL DISTRIBUTION; HERITABILITY; HYBRIDIZATION; MOLECULAR ANALYSIS; PHYLOGENETICS; PHYLOGENY; POLYMORPHISM; RODENT; SUBSPECIES; TAXONOMY;

EID: 83655201191     PISSN: 0018067X     EISSN: 13652540     Source Type: Journal    
DOI: 10.1038/hdy.2011.105     Document Type: Article
Times cited : (48)

References (43)
  • 1
    • 0037337784 scopus 로고    scopus 로고
    • Phylogenetic position and description of a new species of subgenus Mus (Rodentia, Mammalia) from Thailand
    • DOI 10.1046/j.1463-6409.2003.00108.x
    • Auffray JC, Orth A, Catalan J, Gonzalez JP, Desmarais E, Bonhomme F (2003). Phyloge- netic position and description of a new species of subgenus Mus (Rodentia, Mammalia) from Thailand. Zool Scripta32: 119-127. (Pubitemid 36317388)
    • (2003) Zoologica Scripta , vol.32 , Issue.2 , pp. 119-127
    • Auffray, J.-C.1    Orth, A.2    Catalan, J.3    Gonzalez, J.-P.4    Desmarais, E.5    Bonhomme, F.6
  • 2
    • 0025675195 scopus 로고
    • The house mouse progression in Eurasia: A palaeontological and archaeozoological approach
    • Auffray JC, Vanlerberghe F, Britton-Davidian J (1990). The house mouse progression in Eurasia: a palaeontological and archaeozoological approach. Biol JLinn Soc41: 13-25.
    • (1990) Biol JLinn Soc , vol.41 , pp. 13-25
    • Auffray, J.C.1    Vanlerberghe, F.2    Britton-Davidian, J.3
  • 3
    • 0003576389 scopus 로고    scopus 로고
    • Logiciel Sous Windows TM Pour la Génétique Des Populations Université de Montpel- lier II: Montpellier, France
    • Belkhir K, Borsa P, Chikhi L, Raufaste N, Bonhomme F (1996-2004). GENETIX 4.05, Logiciel Sous Windows TM Pour la Génétique Des Populations. Université de Montpel- lier II: Montpellier, France.
    • (1996) GENETIX 4.05
    • Belkhir, K.1    Borsa, P.2    Chikhi, L.3    Raufaste, N.4    Bonhomme, F.5
  • 4
    • 33749631235 scopus 로고    scopus 로고
    • The boundary of macaque rDNA is constituted by low-copy sequences conserved during evolution
    • DOI 10.1016/j.ygeno.2006.05.001, PII S0888754306001455
    • Bodega B, Cardone MF, Rocchi M, Neusser M, Orzan F, Rossi E et al. (2006). Theboundary of macaque r DNA is constituted by low-copy sequences conserved during evolution. Genomics88: 564-571. (Pubitemid 44540242)
    • (2006) Genomics , vol.88 , Issue.5 , pp. 564-571
    • Bodega, B.1    Cardone, M.F.2    Rocchi, M.3    Meneveri, R.4    Marozzi, A.5    Ginelli, E.6
  • 5
    • 34548309601 scopus 로고    scopus 로고
    • Species-wide distribution of highly polymorphic minisatellite markers suggests past and present genetic exchanges among house mouse subspecies
    • Bonhomme F, Rivals E, Orth A, Grant GR, Jeffreys AJ, Bois PRJ (2007). Species-wide distribution of highly polymorphic minisatellite markers suggests past and present genetic exchanges among house mouse subspecies. Genome Biol8: R80.
    • (2007) Genome Biol , vol.8
    • Bonhomme, F.1    Rivals, E.2    Orth, A.3    Grant, G.R.4    Jeffreys, A.J.5    Bois, P.R.J.6
  • 6
    • 45649085741 scopus 로고    scopus 로고
    • Location and expression of ribosomal RNA genes in grasshoppers: Abundance of silent and cryptic loci
    • Cabrero J, Camacho JPM (2008). Location and expression of ribosomal RNA genes in grasshoppers: abundance of silent and cryptic loci. Chrom Res 16: 595-607.
    • (2008) Chrom Res , vol.16 , pp. 595-607
    • Cabrero, J.1    Camacho, J.P.M.2
  • 7
    • 75749083320 scopus 로고    scopus 로고
    • NOR dispersion, telomeric sequence detection in centromeric regions and meiotic multivalent configurations in species of the Aplastodiscus albofrenatus group (Anura, Hylidae
    • Carvalho KA, Garcia PCA, Recco-Pimentel SM (2009). NOR dispersion, telomeric sequence detection in centromeric regions and meiotic multivalent configurations in species of the Aplastodiscus albofrenatus group (Anura, Hylidae). Cytogenet Genome Res 126: 359-367.
    • (2009) Cytogenet Genome Res , vol.126 , pp. 359-367
    • Carvalho, K.A.1    Garcia, P.C.A.2    Recco-Pimentel, S.M.3
  • 8
    • 79955796348 scopus 로고    scopus 로고
    • Are ribosomal DNA clusters rearrangement hotspots? A case study in the genus Mus (Rodentia, Muridae
    • Cazaux B, Catalan J, Veyrunes F, Douzery EJP, Britton-Davidian J (2011). Are ribosomal DNA clusters rearrangement hotspots? A case study in the genus Mus (Rodentia, Muridae). BMCEvol Biol11: 124.
    • (2011) BMCEvol Biol , vol.11 , pp. 124
    • Cazaux, B.1    Catalan, J.2    Veyrunes, F.3    Douzery, E.J.P.4    Britton-Davidian, J.5
  • 9
    • 77957357252 scopus 로고    scopus 로고
    • Distribution of 45S and 5S r DNA sites in 23 species of Eleocharis (Cyperaceae)
    • da Silva CRM, Quintas CC, Vanzela ALL (2010). Distribution of 45S and 5S r DNA sites in 23 species of Eleocharis (Cyperaceae). Genetica 138: 951-957.
    • (2010) Genetica , vol.138 , pp. 951-957
    • Da Silva, C.R.M.1    Quintas, C.C.2    Vanzela, A.L.L.3
  • 10
    • 33846438184 scopus 로고    scopus 로고
    • Ribosomal DNA locus evolution in Nemesia: Transposition rather than structural rearrangement as the key mechanism?
    • DOI 10.1007/s10577-006-1092-z
    • Datson PM, Murray BG (2006). Ribosomal DNA locus evolution in Nemesia: transposition rather than structural rearrangement as the key mechanism? Chrom Res 14: 845-857. (Pubitemid 46144229)
    • (2006) Chromosome Research , vol.14 , Issue.8 , pp. 845-857
    • Datson, P.M.1    Murray, B.G.2
  • 12
    • 0035994597 scopus 로고    scopus 로고
    • Evidence for mitochondrial lineage originating from the Arabian peninsula in the Madagascar house mouse (Mus musculus
    • Duplantier J-M, Orth A, Catalan J, Bonhomme F (2002). Evidence for mitochondrial lineage originating from the Arabian peninsula in the Madagascar house mouse (Mus musculus). Heredity89: 154-158.
    • (2002) Heredity , vol.89 , pp. 154-158
    • Duplantier, J.-M.1    Orth, A.2    Catalan, J.3    Bonhomme, F.4
  • 13
    • 83655213182 scopus 로고    scopus 로고
    • Isolation and geneflow: Inferring the speciation history of European house mice
    • in press
    • Duvaux L, Boulesteix M, Belkhir K, Boursot P (in press) Isolation and geneflow: inferring the speciation history of European house mice. Mol Ecol.
    • Mol Ecol
    • Duvaux, L.1    Boulesteix, M.2    Belkhir, K.3    Boursot, P.4
  • 14
    • 34247899003 scopus 로고    scopus 로고
    • Finely orchestrated movements: Evolution of the ribosomal RNA genes
    • DOI 10.1534/genetics.107.071399
    • Eickbush TH, Eickbush DG (2007). Finely orchestrated movements: Evolution of the ribosomal RNA genes. Genetics 175: 477-485. (Pubitemid 46798252)
    • (2007) Genetics , vol.175 , Issue.2 , pp. 477-485
    • Eickbush, T.H.1    Eickbush, D.G.2
  • 15
    • 0031948760 scopus 로고    scopus 로고
    • Centromeric incompatibilities in the hybrid zone between house mouse subspecies from Denmark: Evidence from patterns of NOR activity
    • Fel-Clair F, Catalan J, Lenormand T, Britton-Davidian J (1998). Centromeric incompat- ibilities in the hybrid zone between house mouse subspecies from Denmark: evidence from patterns of NOR activity. Evolution52: 592-603. (Pubitemid 28222729)
    • (1998) Evolution , vol.52 , Issue.2 , pp. 592-603
    • Fel-Clair, F.1    Catalan, J.2    Lenormand, T.3    Britton-Davidian, J.4
  • 16
    • 0022203256 scopus 로고
    • Phylogenies and the comparative method
    • Felsenstein J (1985). Phylogenies and the comparative method. Am Nat 125: 1-15.
    • (1985) Am Nat , vol.125 , pp. 1-15
    • Felsenstein, J.1
  • 17
    • 0032734531 scopus 로고    scopus 로고
    • A molecularcytogenetic analysis of thetribe Bovini (Artiodactyla: Bovidae: Bovinae) with an emphasisonsexchromosomemorphologyand NOR distribution
    • Gallagher DSJ, Davis SK, De Donato M, Burzlaff JD, Womack JE, Taylor JF et al. (1999). A molecularcytogenetic analysis of thetribe Bovini (Artiodactyla: Bovidae: Bovinae) with an emphasisonsexchromosomemorphologyand NOR distribution. Chrom Res7: 481-492.
    • (1999) Chrom Res , vol.7 , pp. 481-492
    • Gallagher, D.S.J.1    Davis, S.K.2    De Donato, M.3    Burzlaff, J.D.4    Womack, J.E.5    Taylor, J.F.6
  • 18
    • 57349140938 scopus 로고    scopus 로고
    • Inferringthe history of speciation in house mice from autosomal, X-linked, Y-linked and mitochondrial genes
    • Geraldes A, Basset P, Gibson B, Smith KL, Harr B, Yu HTet al. (2008). Inferringthe history of speciation in house mice from autosomal, X-linked, Y-linked and mitochondrial genes. Mol Ecol 17: 5349-5363.
    • (2008) Mol Ecol , vol.17 , pp. 5349-5363
    • Geraldes, A.1    Basset, P.2    Gibson, B.3    Smith, K.L.4    Harr, B.5    Htet Al., Y.6
  • 19
    • 0035338424 scopus 로고    scopus 로고
    • Human rDNA: Evolutionary patterns within the genes and tandem arrays derived from multiple chromosomes
    • DOI 10.1006/geno.2001.6540
    • Gonzalez IL, Sylvester JE (2001). Human r DNA: evolutionarypatternswithinthegenesand tandem arrays derived from multiple chromosomes. Genomics 73: 255-263. (Pubitemid 32463721)
    • (2001) Genomics , vol.73 , Issue.3 , pp. 255-263
    • Gonzalez, I.L.1    Sylvester, J.E.2
  • 20
    • 0242594489 scopus 로고    scopus 로고
    • Complete sequence of the 45-kb mouse ribosomal DNA repeat: Analysis of the intergenic spacer
    • DOI 10.1016/S0888-7543(03)00199-X
    • Grozdanov P, Geogiev O, Karagyozov L (2003). Complete sequence of the 45-kb mouse ribosomal DNA repeat: analysis of the intergenic spacer. Genomics82: 637-643. (Pubitemid 37373720)
    • (2003) Genomics , vol.82 , Issue.6 , pp. 637-643
    • Grozdanov, P.1    Georgiev, O.2    Karagyozov, L.3
  • 21
    • 38149113813 scopus 로고    scopus 로고
    • Identification of r RNAgene loci in the wild mouse (Mus musculus molossinus) captured at Hachioji, Tokyo
    • Ito T, Osawa S, Shibata H, Kanda N (2007). Identification of r RNAgene loci in the wild mouse (Mus musculus molossinus) captured at Hachioji, Tokyo. J Vet Med Sci 69: 1277-1279.
    • (2007) J Vet Med Sci , vol.69 , pp. 1277-1279
    • Ito, T.1    Osawa Shibata, S.H.2    Kanda, N.3
  • 22
    • 54349098760 scopus 로고    scopus 로고
    • Mapping of r RNAgene loci in the mice, Mus musculus molossinus (Japan) captured and Mus musculus musculus (Russia) by double color FISH
    • Ito T, Tsuchiya K, Osawa S, Shibata H, Kanda N (2008). Mapping of r RNAgene loci in the mice, Mus musculus molossinus (Japan) captured and Mus musculus musculus (Russia) by double color FISH. J Vet Med Sci 70: 997-1000.
    • (2008) J Vet Med Sci , vol.70 , pp. 997-1000
    • Ito, T.1    Tsuchiya, K.2    Osawa, S.3    Shibata, H.4    Kanda, N.5
  • 23
    • 3142776157 scopus 로고    scopus 로고
    • Karyotypic evolution of Apodemus (Muridae, Rodentia) inferred from comparative FISH analyses
    • DOI 10.1023/B:CHRO.0000034103.05528.83
    • Matsubara K, Nishida-Umehara C, Tsuchiya K, Nukaya D, Matsuda Y (2004). Karyotypic evolution of Apodemus (Muridae, Rodentia) inferred from comparative FISH analyses. Chrom Res 12: 383-395. (Pubitemid 38932378)
    • (2004) Chromosome Research , vol.12 , Issue.4 , pp. 383-395
    • Matsubara, K.1    Nishida-Umehara, C.2    Tsuchiya, K.3    Nukaya, D.4    Matsuda, Y.5
  • 24
    • 77949269660 scopus 로고    scopus 로고
    • Evolutionary dynamics of r DNA clusters on chromosomes of moths and butterflies (Lepidoptera)
    • Nguyen P, Sahara K, Yoshido A, Marec F (2010). Evolutionary dynamics of r DNA clusters on chromosomes of moths and butterflies (Lepidoptera). Genetica 138: 343-354.
    • (2010) Genetica , vol.138 , pp. 343-354
    • Nguyen, P.1    Sahara, K.2    Yoshido, A.3    Marec, F.4
  • 25
    • 54249129804 scopus 로고    scopus 로고
    • Phylogenetic position of the saola (Pseudoryx nghetinhensis) inferred from cytogenetic analysis of eleven species of Bovidae
    • Nguyen TT, Aniskin VM, Gerbault-Seureau M, Planton H, Renard JP, Nguyen BX et al. (2008). Phylogenetic position of the saola (Pseudoryx nghetinhensis) inferred from cytogenetic analysis of eleven species of Bovidae. Cytogenet Genome Res 122: 41-54.
    • (2008) Cytogenet Genome Res , vol.122 , pp. 41-54
    • Nguyen, T.T.1    Aniskin, V.M.2    Gerbault-Seureau, M.3    Planton, H.4    Renard, J.P.5    Nguyen, B.X.6
  • 26
    • 0036023028 scopus 로고    scopus 로고
    • Natural hybridisation between two sympatric species of mice Mus musculus domesticus L. and Mus spretus Lataste
    • DOI 10.1016/S1631-0691(02)01413-0
    • Orth A, Belkhir K, Britton-Davidian J, Boursot P, Benazzou T, Bonhomme F(2002). Natural hybridization between two sympatric species of mice Mus musculus domesticus L and Mus spretus Lataste. C R Acad Sci 325: 89-97. (Pubitemid 34855206)
    • (2002) Comptes Rendus - Biologies , vol.325 , Issue.2 , pp. 89-97
    • Orth, A.1    Belkhir, K.2    Britton-Davidian, J.3    Boursot, P.4    Benazzou, T.5    Bonhomme, F.6
  • 27
    • 0030203863 scopus 로고    scopus 로고
    • TREEVIEW: An application to display phylogenetic trees on personal computers
    • Page RDM (1996). TREEVIEW: an application to display phylogenetic trees on personal computers. Comp Appl Biosci 12: 357-358.
    • (1996) Comp Appl Biosci , vol.12 , pp. 357-358
    • Page, R.D.M.1
  • 28
    • 3242795199 scopus 로고    scopus 로고
    • Within- and between-individual sequence variation among ITS1 copies in the meadow grasshopper Chorthippus parallelus indicates frequent intrachromosomal gene conversion
    • DOI 10.1093/molbev/msh163
    • Parkin EJ, Butlin RK (2004). Within- and between-individual sequence variation among ITS1 copies in the meadow grasshopper Chorthippus parallelus indicates frequent intrachromosomal gen conversion. Mol Biol Evol 21: 1595-1601. (Pubitemid 38971770)
    • (2004) Molecular Biology and Evolution , vol.21 , Issue.8 , pp. 1595-1601
    • Parkin, E.J.1    Butlin, R.K.2
  • 30
    • 0031662638 scopus 로고    scopus 로고
    • Genetic variation and phylogeography of central asian and other house mice, including a major new mitochondrial lineage in Yemen
    • Prager EM, Orrego C, Sage RD (1998). Genetic variation and phylogeography of central Asian and other house mice, including a major new mitochondrial lineage in Yemen. Genetics 150: 835-861. (Pubitemid 28463059)
    • (1998) Genetics , vol.150 , Issue.2 , pp. 835-861
    • Prager, E.M.1    Orrego, C.2    Sage, R.D.3
  • 31
    • 38149012877 scopus 로고    scopus 로고
    • Phylogeographyand postglacial expansion of Mus musculus domesticus inferred from mitochondrial DNA coalescent from Iran to Europe
    • Rajabi-Maham H, Orth A, Bonhomme F (2008). Phylogeographyand postglacial expansion of Mus musculus domesticus inferred from mitochondrial DNA coalescent from Iran to Europe. Mol Ecol 17: 627-641.
    • (2008) Mol Ecol , vol.17 , pp. 627-641
    • Rajabi-Maham, H.1    Orth, A.2    Bonhomme, F.3
  • 32
    • 15944428236 scopus 로고    scopus 로고
    • AgNOR variability among Robertsonian races of the house mouse from the island of Madeira: Implications for patterns of Rb fusion formation and genetic differentiation
    • DOI 10.1111/j.1095-8312.2005.00456.x
    • Ramalhinho MG, Braz C, Catalan J, Mathias ML, Britton-Davidian J (2005). Ag NOR variability among Robertsonian races of the house mouse from the island of Madeira: implications for patterns of Rb fusion formation and genetic differentiation. Biol JLinn Soc 84: 585-591. (Pubitemid 40441060)
    • (2005) Biological Journal of the Linnean Society , vol.84 , Issue.3 , pp. 585-591
    • Ramalhinho, G.1    Braz, C.2    Catalan, J.3    Mathias, M.L.4    Britton-Davidian, J.5
  • 33
    • 44349083668 scopus 로고    scopus 로고
    • Repetitive DNA and chromosomal rearrangements: Speciation-related events in plant genomes
    • DOI 10.1159/000121084
    • Raskina O, Barber JC, Nevo E, Belyayev A (2008). Repetitive DNA and chromosomal rearrangements: speciation-related events in plant genomes. Cytogenet Genome Res 120: 351-357. (Pubitemid 351738041)
    • (2008) Cytogenetic and Genome Research , vol.120 , Issue.3-4 , pp. 351-357
    • Raskina, O.1    Barber, J.C.2    Nevo, E.3    Belyayev, A.4
  • 34
    • 55749111236 scopus 로고    scopus 로고
    • Hemiplasy and homoplasy in the karyotypic phylogenies of mammals
    • Robinson TJ, Ruiz-Herrera A, Avise J (2008). Hemiplasy and homoplasy in the karyotypic phylogenies of mammals. Proc Natl Acad Sci 105: 14477-14481.
    • (2008) Proc Natl Acad Sci , vol.105 , pp. 14477-14481
    • Robinson, T.J.1    Ruiz-Herrera, A.2    Avise, J.3
  • 35
    • 0030330244 scopus 로고    scopus 로고
    • Genetic mapping of 18S ribosomal RNA-related loci to mouse Chromosomes 5, 6, 9, 12, 17, 18, 19, and X
    • Rowe LB, Janaswami PM, Barter ME, Birkenmeier EH (1996). Genetic mapping of 18S ribosomal RNA-related loci to mouse chromosomes 5, 6, 9, 12, 17, 18, 19, and X. Mamm Genome 7: 886-889. (Pubitemid 126439170)
    • (1996) Mammalian Genome , vol.7 , Issue.12 , pp. 886-889
    • Rowe, L.B.1    Janaswami, P.M.2    Barter, M.E.3    Birkenmeier, E.H.4
  • 37
    • 0025364752 scopus 로고
    • Variation in the distribution of silver-staining Nucleolar Organizer Regions on the chromosomes of the wild mouse, Mus musculus
    • Suzuki H, Kurihara Y, Moriwaki K (1990). Variation in the distribution of silver-staining Nucleolar Organizer Regions on the chromosomes of the wild mouse, Mus musculus. Mol Biol Evol7: 271-282.
    • (1990) Mol Biol Evo , vol.17 , pp. 271-282
    • Suzuki, H.1    Kurihara, Y.2    Moriwaki, K.3
  • 38
    • 0022655697 scopus 로고
    • Evolutionary implication of heterogeneity of the nontranscribed spacer region of ribosomal DNA repeating units in various subspecies of Mus musculus
    • Suzuki H, Miyashita N, Moriwaki K, Kominami R, Muramatsu M, Kanehisa Tet al. (1986). Evolutionary implications of heterogeneity of the nontranscribed spacer region of ribosomal DNA repeating units in various subspecies of Mus musculus. Mol Biol Evol 3: 126-137. (Pubitemid 16118385)
    • (1986) Molecular Biology and Evolution , vol.3 , Issue.2 , pp. 126-137
    • Suzuki, H.1    Miyashita, N.2    Moriwaki, K.3
  • 39
    • 68149153157 scopus 로고    scopus 로고
    • Geographical localised bursts of ribosomal DNA mobility in the grasshopper Podismapedestris
    • Veltsos P, Keller I, Nichols RA (2009). Geographical localised bursts of ribosomal DNA mobility in the grasshopper Podismapedestris. Heredity 103: 54-61.
    • (2009) Heredity , vol.103 , pp. 54-61
    • Veltsos, P.1    Keller, I.2    Nichols, R.A.3
  • 40
    • 33646833572 scopus 로고    scopus 로고
    • High-resolution single-copy gene fluorescence in situ hybridization and its use in the construction of a cytogenetic map of maize chromosome 9
    • DOI 10.1105/tpc.105.037838
    • Wang C-JR, Harper L, Cande Z (2006). High-resolution single-copy gene fluorescence in situ hybridization and its use in the construction of a cytogenetic map of maize chromosome 9. Plant Cell 18: 529-544. (Pubitemid 43956159)
    • (2006) Plant Cell , vol.18 , Issue.3 , pp. 529-544
    • Wang, C.-J.R.1    Harper, L.2    Cande, W.Z.3
  • 42
    • 84925588054 scopus 로고    scopus 로고
    • Origin and genetic status of Mus musculus molossinus: A typical example for reticulate evolution in the genus Mus
    • In: Macholan M, Baird S, Munclinger P, Piálek J (eds) Cambridge University Press: Cambridge, UK
    • Yonekawa H, Sato JJ, Suzuki H, Moriwaki K (2012). Origin and genetic status of Mus musculus molossinus: a typical example for reticulate evolution in the genus Mus. In: Macholan M, Baird S, Munclinger P, Piálek J (eds). Evolution of the House Mouse. Cambridge University Press: Cambridge, UK.
    • (2012) Evolution of the House Mouse
    • Yonekawa, H.1    Sato, J.J.2    Suzuki, H.3    Moriwaki, K.4
  • 43
    • 1942441730 scopus 로고    scopus 로고
    • Transcriptional repression mechanisms of nucleolus organizer regions (NORs) in humans and chimpanzees
    • DOI 10.1023/B:CHRO.0000021911.43225.eb
    • Zavala Guillén AK, Hirai Y, Tanoue T, Hirai H (2004). Transcriptional repression mechan- isms of nucleolus organizer regions (NORs) in humans and chimpanzees. Chrom Res 12: 225-237. (Pubitemid 38507849)
    • (2004) Chromosome Research , vol.12 , Issue.3 , pp. 225-237
    • Zavala Guillén, A.K.1    Hirai, Y.2    Tanoue, T.3    Hirai, H.4


* 이 정보는 Elsevier사의 SCOPUS DB에서 KISTI가 분석하여 추출한 것입니다.