HLA遺伝子多型からみた日本人集団の混合的起源
HLA領域の遺伝的多型は医学分野のみならず，人類の移住，混合，自然選択や遺伝的適応といった進化過程を推測する集団遺伝学においても重要なツールとして用いられている。日本人の起源については諸説あるが，約30,000年前の後期旧石器時代に日本列島へと移住してきた狩猟採集民である縄文人と，紀元前1,000年から西暦300年頃に朝鮮半島から日本列島へと移住してきた弥生人の祖先が混血して，現在の日本人集団の起源になったとする“混合モデル”が有力であると考えられている。本稿では，日本人集団の混合的起源を支持する遺伝的知見について概説するとともに，HLA遺伝子多型から日本人の祖先集団を推測する遺伝的痕跡の探索...
Saved in:
| Published in | MHC（日本組織適合性学会誌） Vol. 21; no. 1; pp. 37 - 44 |
|---|---|
| Main Authors | , , , , |
| Format | Journal Article |
| Language | Japanese |
| Published |
日本組織適合性学会
20.04.2014
|
| Subjects | |
| Online Access | Get full text |
| ISSN | 2186-9995 2187-4239 |
| DOI | 10.12667/mhc.21.37 |
Cover
| Abstract | HLA領域の遺伝的多型は医学分野のみならず，人類の移住，混合，自然選択や遺伝的適応といった進化過程を推測する集団遺伝学においても重要なツールとして用いられている。日本人の起源については諸説あるが，約30,000年前の後期旧石器時代に日本列島へと移住してきた狩猟採集民である縄文人と，紀元前1,000年から西暦300年頃に朝鮮半島から日本列島へと移住してきた弥生人の祖先が混血して，現在の日本人集団の起源になったとする“混合モデル”が有力であると考えられている。本稿では，日本人集団の混合的起源を支持する遺伝的知見について概説するとともに，HLA遺伝子多型から日本人の祖先集団を推測する遺伝的痕跡の探索について，最新の研究を紹介したい。 |
|---|---|
| AbstractList | HLA領域の遺伝的多型は医学分野のみならず，人類の移住，混合，自然選択や遺伝的適応といった進化過程を推測する集団遺伝学においても重要なツールとして用いられている。日本人の起源については諸説あるが，約30,000年前の後期旧石器時代に日本列島へと移住してきた狩猟採集民である縄文人と，紀元前1,000年から西暦300年頃に朝鮮半島から日本列島へと移住してきた弥生人の祖先が混血して，現在の日本人集団の起源になったとする“混合モデル”が有力であると考えられている。本稿では，日本人集団の混合的起源を支持する遺伝的知見について概説するとともに，HLA遺伝子多型から日本人の祖先集団を推測する遺伝的痕跡の探索について，最新の研究を紹介したい。 HLA領域の遺伝的多型は医学分野のみならず, 人類の移住, 混合, 自然選択や遺伝的適応といった進化過程を推測する集団遺伝学においても重要なツールとして用いられている. 日本人の起源については諸説あるが, 約30,000年前の後期旧石器時代に日本列島へと移住してきた狩猟採集民である縄文人と, 紀元前1,000年から西暦300年頃に朝鮮半島から日本列島へと移住してきた弥生人の祖先が混血して, 現在の日本人集団の起源になったとする“混合モデル”が有力であると考えられている. 本稿では, 日本人集団の混合的起源を支持する遺伝的知見について概説するとともに, HLA遺伝子多型から日本人の祖先集団を推測する遺伝的痕跡の探索について, 最新の研究を紹介したい. 「はじめに」 HLA領域はヒト第6染色体短腕の3.6Mbにわたる主要組織適合遺伝子複合体に相当する領域である. HLA領域は免疫機能に関わる多数の遺伝子を有し, ヒトゲノムで最も多型性に富む領域である. |
| Author | 中岡, 博史 猪子, 英俊 井ノ上, 逸朗 光永, 滋樹 細道, 一善 |
| Author_xml | – sequence: 1 fullname: 細道, 一善 organization: 国立遺伝学研究所 総合遺伝研究系 人類遺伝研究部門 – sequence: 1 fullname: 光永, 滋樹 organization: 東海大学医学部 基礎医学系 分子生命科学 – sequence: 1 fullname: 中岡, 博史 organization: 国立遺伝学研究所 総合遺伝研究系 人類遺伝研究部門 – sequence: 1 fullname: 井ノ上, 逸朗 organization: 国立遺伝学研究所 総合遺伝研究系 人類遺伝研究部門 – sequence: 1 fullname: 猪子, 英俊 organization: 東海大学医学部 基礎医学系 分子生命科学 |
| BookMark | eNo1kM1Kw0AUhQdRsNZufI_UuTOTSQKClKKtUHCj62EyndiENpWkLlxGRJRuuhNafxbqTt1JoQg-TJpq38JodXPO4cL9LueuoeWwG2qENgCXgXBubXZaqkygTK0lVCBgWwYj1Fn-zdxwHMdcRaU49l3MMOMWBqeAtuuNyjyZTN9vs5dB9jjM7vpp0k_PrtLkI03uZ9dPs5vn6WQyH11ko4c0eZ2Nx9ng8nN4_vU2nk0G62jFk-1Yl_68iA53dw6qdaOxX9urVhpGQDhjhgaMPerSJm-6HmVKetw1TYs62qa2wlgq6hHJPWAe17ZrccfhJlEUbJPLpgJaRLUFt6ObvpLtbtj2Qy2C7kkU5neFUpC373YEwcAExgQwiEWkVi4sfwVwDiwnbS1IQdyTR1ocR35HRqdCRj1ftbXIMYKAgB_JV__HqiUjEUj6DRQygJ0 |
| ContentType | Journal Article |
| Copyright | 2014 日本組織適合性学会 |
| Copyright_xml | – notice: 2014 日本組織適合性学会 |
| CorporateAuthor | 国立遺伝学研究所総合遺伝研究系人類遺伝研究部門 東海大学医学部基礎医学系分子生命科学 |
| CorporateAuthor_xml | – name: 国立遺伝学研究所総合遺伝研究系人類遺伝研究部門 – name: 東海大学医学部基礎医学系分子生命科学 |
| DOI | 10.12667/mhc.21.37 |
| DatabaseTitleList | |
| DeliveryMethod | fulltext_linktorsrc |
| EISSN | 2187-4239 |
| EndPage | 44 |
| ExternalDocumentID | cc1mhcom_2014_002101_014_0037_00442316614 article_mhc_21_1_21_37_article_char_ja |
| GroupedDBID | ABJNI ALMA_UNASSIGNED_HOLDINGS JSF KQ8 MOJWN RJT |
| ID | FETCH-LOGICAL-j2644-e100f3b3d6dbf34caf6b55739e838c00ac3f2a6f14f6e8b7699652c31856adc13 |
| ISSN | 2186-9995 |
| IngestDate | Thu Jul 10 16:21:12 EDT 2025 Wed Sep 03 06:30:22 EDT 2025 |
| IsDoiOpenAccess | true |
| IsOpenAccess | true |
| IsPeerReviewed | false |
| IsScholarly | false |
| Issue | 1 |
| Language | Japanese |
| LinkModel | OpenURL |
| MergedId | FETCHMERGED-LOGICAL-j2644-e100f3b3d6dbf34caf6b55739e838c00ac3f2a6f14f6e8b7699652c31856adc13 |
| OpenAccessLink | https://www.jstage.jst.go.jp/article/mhc/21/1/21_37/_article/-char/ja |
| PageCount | 8 |
| ParticipantIDs | medicalonline_journals_cc1mhcom_2014_002101_014_0037_00442316614 jstage_primary_article_mhc_21_1_21_37_article_char_ja |
| PublicationCentury | 2000 |
| PublicationDate | 20140420 |
| PublicationDateYYYYMMDD | 2014-04-20 |
| PublicationDate_xml | – month: 04 year: 2014 text: 20140420 day: 20 |
| PublicationDecade | 2010 |
| PublicationTitle | MHC（日本組織適合性学会誌） |
| PublicationTitleAlternate | 日本組織適合性学会誌 |
| PublicationYear | 2014 |
| Publisher | 日本組織適合性学会 |
| Publisher_xml | – name: 日本組織適合性学会 |
| References | 13) Tajima A, Hayami M, Tokunaga K, et al.: Genetic origins of the Ainu inferred from combined DNA analyses of maternal and paternal lineages. Journal of Human Genetics (49): 187–193, 2004. 8) Yamaguchi-Kabata Y, Nakazono K, Takahashi A, et al.: Japanese population structure, based on SNP genotypes from 7003 individuals compared to other ethnic groups: Effects on population-based association studies. American Journal of Human Genetics (83): 445–456, 2008. 21) Karafet T, Xu L, Du R, et al.: Paternal population history of East Asia: Sources, patterns, and microevolutionary processes. American Journal of Human Genetics (69): 615–628, 2001. 28) Shiina T, Suzuki S, Ozaki Y, et al.: Super high resolution for single molecule-sequence-based typing of classical HLA loci at the 8-digit level using next generation sequencers. Tissue Antigens. (80): 305–316, 2012. 11) Hatta Y, Ohashi J, Imanishi T, et al.: HLA genes and haplotypes in Ryukyuans suggest recent gene flow to the Okinawa islands. Human Biology (71): 353–365, 1999. 16) Lee KW, Oh DH, Lee C, et al.: Allelic and haplotypic diversity of HLA-A, -B, -C, -DRB1, and -DQB1 genes in the Korean population. Tissue Antigens (65): 437–447, 2005. 4) Hanihara K: Dual structure model for the population history of the Japanese. Japan Review (2):1–33, 1991. 10) Tokunaga K, Imanishi T, Takahashi K, et al.: On the origin and dispersal of East Asian populations as viewed from HLA haplotypes. In Akazawa T, Szathmary EJE (Eds.), Prehistoric Mongoloid Dispersals, Oxford University Press, Oxford. 1996. 2) Robinson J, Waller MJ, Parham P, et al.: IMGT/HLA and IMGT/MHC: Sequence databases for the study of the major histocompatibility complex. Nucleic Acids Research (31): 311–314, 2003. 14) Tokunaga K, Ishikawa Y, Ogawa A, et al.: Sequence-based association analysis of HLA class I and II alleles in Japanese supports conservation of common haplotypes. Immunogenetics (46): 199–205, 1997. 23) Goebel T, Waters MR, O’Rourke DH.: The late pleistocene dispersal of modern humans in the Americas. Science (319): 1497–1502, 2008. 3) Vina MA, Hollenbach JA, Lyke KE, et al.: Tracking human migrations by the analysis of the distribution of HLA alleles, lineages and haplotypes in closed and open populations. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences (367): 820–829, 2012. 7) Tanaka M, Cabrera VM, Gonzalez AM, et al.: Mitochondrial genome variation in eastern Asia and the peopling of Japan. Genome Research (14): 1832–1850, 2004. 19) HUGO Pan-Asian SNP Consortium, Abdulla MA, Ahmed I, et al.: Mapping human genetic diversity in Asia. Science (326): 1541–1545, 2009. 26) Hosomichi K, Jinam TA, Mitsunaga S, et al.: Phase-defined complete sequencing of the HLA genes by next-generation sequencing. BMC Genomics (14): 355, 2013. 27) Wang C, Krishnakumar S, Wilhelmy J, et al.: High-throughput, high-fidelity HLA genotyping with deep sequencing. Proceedings of the National Academy of Sciences of the United States of America (109): 8676–8681, 2012. 6) Hammer MF, Karafet TM, Park H, et al.: Dual origins of the Japanese: Common ground for hunter-gatherer and farmer Y chromosomes. Journal of Human Genetics (51): 47–58, 2006. 12) Bannai M, Ohashi J, Harihara S, et al.: Analysis of HLA genes and haplotypes in Ainu (from hokkaido, northern Japan) supports the premise that they descent from upper paleolithic populations of East Asia. Tissue Antigens (55): 128–139, 2000. 1) Shiina T, Hosomichi K, Inoko H, et al.: The HLA genomic loci map: Expression, interaction, diversity and disease. Journal of Human Genetics (54): 15–39, 2009. 5) Omoto K, Saitou N: Genetic origins of the Japanese: A partial support for the dual structure hypothesis. American Journal of Physical Anthropology (102): 437–446, 1997. 17) Nakaoka H, Mitsunaga S, Hosomichi K, et al.: Detection of ancestry informative HLA alleles confirms the admixed origins of Japanese population. PLoS One (8): e60793, 2013. 25)Reich D, Patterson N, Campbell D, et al.: Reconstructing native American population history. Nature (488): 370–374, 2012. 20) Cavalli-Sforza LL, Menozzi P, Piazza A: The history and geography of human genes. Princeton University Press, Princeton. 1994. 24) Rasmussen M, Li Y, Lindgreen S, et al.: Ancient human genome sequence of an extinct Palaeo-Eskimo. Nature (463): 757–762, 2010. 9) Yamaguchi-Kabata Y, Tsunoda T, Kumasaka N, et al.: Genetic differences in the two main groups of the Japanese population based on autosomal SNPs and haplotypes. Journal of Human Genetics (57): 326–334, 2012. 18) Kawashima M, Ohashi J, Nishida N, et al.: Evolutionary analysis of classical HLA class I and II genes suggests that recent positive selection acted on DPB1*04:01 in Japanese population. PLoS One (7): e46806, 2012. 22)Di D, Sanchez-Mazas A: Challenging views on the peopling history of East Asia: The story according to HLA markers. American Journal of Physical Anthropology (145): 81–96, 2011. 15) Saito S, Ota S, Yamada E, et al.: Allele frequencies and haplotypic associations defined by allelic DNA typing at HLA class I and class II loci in the Japanese population. Tissue Antigens (56): 522–529, 2000. |
| References_xml | – reference: 28) Shiina T, Suzuki S, Ozaki Y, et al.: Super high resolution for single molecule-sequence-based typing of classical HLA loci at the 8-digit level using next generation sequencers. Tissue Antigens. (80): 305–316, 2012. – reference: 3) Vina MA, Hollenbach JA, Lyke KE, et al.: Tracking human migrations by the analysis of the distribution of HLA alleles, lineages and haplotypes in closed and open populations. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences (367): 820–829, 2012. – reference: 16) Lee KW, Oh DH, Lee C, et al.: Allelic and haplotypic diversity of HLA-A, -B, -C, -DRB1, and -DQB1 genes in the Korean population. Tissue Antigens (65): 437–447, 2005. – reference: 4) Hanihara K: Dual structure model for the population history of the Japanese. Japan Review (2):1–33, 1991. – reference: 18) Kawashima M, Ohashi J, Nishida N, et al.: Evolutionary analysis of classical HLA class I and II genes suggests that recent positive selection acted on DPB1*04:01 in Japanese population. PLoS One (7): e46806, 2012. – reference: 19) HUGO Pan-Asian SNP Consortium, Abdulla MA, Ahmed I, et al.: Mapping human genetic diversity in Asia. Science (326): 1541–1545, 2009. – reference: 1) Shiina T, Hosomichi K, Inoko H, et al.: The HLA genomic loci map: Expression, interaction, diversity and disease. Journal of Human Genetics (54): 15–39, 2009. – reference: 12) Bannai M, Ohashi J, Harihara S, et al.: Analysis of HLA genes and haplotypes in Ainu (from hokkaido, northern Japan) supports the premise that they descent from upper paleolithic populations of East Asia. Tissue Antigens (55): 128–139, 2000. – reference: 20) Cavalli-Sforza LL, Menozzi P, Piazza A: The history and geography of human genes. Princeton University Press, Princeton. 1994. – reference: 14) Tokunaga K, Ishikawa Y, Ogawa A, et al.: Sequence-based association analysis of HLA class I and II alleles in Japanese supports conservation of common haplotypes. Immunogenetics (46): 199–205, 1997. – reference: 17) Nakaoka H, Mitsunaga S, Hosomichi K, et al.: Detection of ancestry informative HLA alleles confirms the admixed origins of Japanese population. PLoS One (8): e60793, 2013. – reference: 15) Saito S, Ota S, Yamada E, et al.: Allele frequencies and haplotypic associations defined by allelic DNA typing at HLA class I and class II loci in the Japanese population. Tissue Antigens (56): 522–529, 2000. – reference: 8) Yamaguchi-Kabata Y, Nakazono K, Takahashi A, et al.: Japanese population structure, based on SNP genotypes from 7003 individuals compared to other ethnic groups: Effects on population-based association studies. American Journal of Human Genetics (83): 445–456, 2008. – reference: 13) Tajima A, Hayami M, Tokunaga K, et al.: Genetic origins of the Ainu inferred from combined DNA analyses of maternal and paternal lineages. Journal of Human Genetics (49): 187–193, 2004. – reference: 27) Wang C, Krishnakumar S, Wilhelmy J, et al.: High-throughput, high-fidelity HLA genotyping with deep sequencing. Proceedings of the National Academy of Sciences of the United States of America (109): 8676–8681, 2012. – reference: 9) Yamaguchi-Kabata Y, Tsunoda T, Kumasaka N, et al.: Genetic differences in the two main groups of the Japanese population based on autosomal SNPs and haplotypes. Journal of Human Genetics (57): 326–334, 2012. – reference: 24) Rasmussen M, Li Y, Lindgreen S, et al.: Ancient human genome sequence of an extinct Palaeo-Eskimo. Nature (463): 757–762, 2010. – reference: 23) Goebel T, Waters MR, O’Rourke DH.: The late pleistocene dispersal of modern humans in the Americas. Science (319): 1497–1502, 2008. – reference: 7) Tanaka M, Cabrera VM, Gonzalez AM, et al.: Mitochondrial genome variation in eastern Asia and the peopling of Japan. Genome Research (14): 1832–1850, 2004. – reference: 5) Omoto K, Saitou N: Genetic origins of the Japanese: A partial support for the dual structure hypothesis. American Journal of Physical Anthropology (102): 437–446, 1997. – reference: 2) Robinson J, Waller MJ, Parham P, et al.: IMGT/HLA and IMGT/MHC: Sequence databases for the study of the major histocompatibility complex. Nucleic Acids Research (31): 311–314, 2003. – reference: 22)Di D, Sanchez-Mazas A: Challenging views on the peopling history of East Asia: The story according to HLA markers. American Journal of Physical Anthropology (145): 81–96, 2011. – reference: 21) Karafet T, Xu L, Du R, et al.: Paternal population history of East Asia: Sources, patterns, and microevolutionary processes. American Journal of Human Genetics (69): 615–628, 2001. – reference: 26) Hosomichi K, Jinam TA, Mitsunaga S, et al.: Phase-defined complete sequencing of the HLA genes by next-generation sequencing. BMC Genomics (14): 355, 2013. – reference: 11) Hatta Y, Ohashi J, Imanishi T, et al.: HLA genes and haplotypes in Ryukyuans suggest recent gene flow to the Okinawa islands. Human Biology (71): 353–365, 1999. – reference: 6) Hammer MF, Karafet TM, Park H, et al.: Dual origins of the Japanese: Common ground for hunter-gatherer and farmer Y chromosomes. Journal of Human Genetics (51): 47–58, 2006. – reference: 10) Tokunaga K, Imanishi T, Takahashi K, et al.: On the origin and dispersal of East Asian populations as viewed from HLA haplotypes. In Akazawa T, Szathmary EJE (Eds.), Prehistoric Mongoloid Dispersals, Oxford University Press, Oxford. 1996. – reference: 25)Reich D, Patterson N, Campbell D, et al.: Reconstructing native American population history. Nature (488): 370–374, 2012. |
| SSID | ssib040467019 ssib008886877 ssib012948844 ssj0000866597 |
| Score | 1.6205113 |
| Snippet | HLA領域の遺伝的多型は医学分野のみならず，人類の移住，混合，自然選択や遺伝的適応といった進化過程を推測する集団遺伝学においても重要なツールとして用いられている。... HLA領域の遺伝的多型は医学分野のみならず, 人類の移住, 混合, 自然選択や遺伝的適応といった進化過程を推測する集団遺伝学においても重要なツールとして用いられている. ... |
| SourceID | medicalonline jstage |
| SourceType | Publisher |
| StartPage | 37 |
| SubjectTerms | ヒト白血球抗原 日本人集団 混合モデル 移住 進化 |
| Title | HLA遺伝子多型からみた日本人集団の混合的起源 |
| URI | https://www.jstage.jst.go.jp/article/mhc/21/1/21_37/_article/-char/ja http://mol.medicalonline.jp/en/journal/download?GoodsID=cc1mhcom/2014/002101/014&name=0037-0044j |
| Volume | 21 |
| hasFullText | 1 |
| inHoldings | 1 |
| isFullTextHit | |
| isPrint | |
| ispartofPNX | 日本組織適合性学会誌, 2014/04/20, Vol.21(1), pp.37-44 |
| journalDatabaseRights | – providerCode: PRVAFT databaseName: Open Access Digital Library customDbUrl: eissn: 2187-4239 dateEnd: 99991231 omitProxy: true ssIdentifier: ssj0000866597 issn: 2186-9995 databaseCode: KQ8 dateStart: 20130101 isFulltext: true titleUrlDefault: http://grweb.coalliance.org/oadl/oadl.html providerName: Colorado Alliance of Research Libraries |
| link | http://utb.summon.serialssolutions.com/2.0.0/link/0/eLvHCXMwrR1Na9VAMNR6EUQUFesXPbgnSU2yyWb3pJvXlOIXCC30FrJ5CfLgtaLtxdsTEaWX3oTWj4N6U29SeAj-EW-v72n_hTO7m_eBPdQihDDZnZ3Znd3NzCa7M45zrcyDZl5w3xWR13RDFTKXR0q5VBTCUxVnVMdYunefLS6Ht1eilaljP8d2LW2sq7ni6YHnSo7Sq5AG_YqnZP-hZ4dEIQFg6F-4Qw_D_VB9vHhXklSAMUgSAEKSNIiYJ2lE5DwRngZCIiQCIiU8ISlFZAsEhAubkixYQADAiIiJjDTQILKhKUvNQhCREM40wYRIakvJFJGTGC_M8gjnJI2RNQ9JykkS6SyGREzcztogPohdjPhYEADgGNo2SjFGnOEWDRnbxkpWN18iOwl8h19_NRnI9CwZ9GvVuK7xORJBmkAtHOED80jLhmEpwNL48JBo0TEioUVihK8pSS34JCDSN_hAYt4Kny9AxgQ-iCHSwoMrresiTUGBtUq4FYjZV1x_mPH1fp7AG06l_y6_kWbAOGIuWPZmO0BZp8UuenMcV23m8PnEFDZ6yjjasRaPccD5ly4F0w1_57cfFnOBP1eXmPBNbkd-BjhZ4Gc-3mic1cl4cDBrwerleAA6FwOr3HkwettzzhkfnYIGwxOUycjYDD3Q4XWsAG1XoWtGHS1p2H7rbxhremNYT7AlW7CyQpcZJ9vmb6fxOjNmOC6ddk7ZFd-sNLU940y18rPOTZi6-53u3ve3_S9b_Y_b_Xebvc5m79mrXudHr_N-8PrT4M3nvW53f-dFf-dDr_N1sLvb33r5a_v572-7g-7WOWd5IV1qLLo2lonbwiWHW_qeV1FFMX5bRcMir5iKopiKklNeeF5e0CrIWeWHFSu5ipkQLAoK9G3A8mbh0_PO9OraannBmVW0KmMl4OI09KpQBYVivOnnhSoUrI9mnMi0P3tkHNZkh-umGefWhLgy-6Z7khWFD0XX2hkO80x_JvIzA1LcvhLCuPPR0L94RNaXnBOjGXTZmV5_vFFeAWN_XV3VY-YPuufMPw |
| linkProvider | Colorado Alliance of Research Libraries |
| openUrl | ctx_ver=Z39.88-2004&ctx_enc=info%3Aofi%2Fenc%3AUTF-8&rfr_id=info%3Asid%2Fsummon.serialssolutions.com&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=HLA%E9%81%BA%E4%BC%9D%E5%AD%90%E5%A4%9A%E5%9E%8B%E3%81%8B%E3%82%89%E3%81%BF%E3%81%9F%E6%97%A5%E6%9C%AC%E4%BA%BA%E9%9B%86%E5%9B%A3%E3%81%AE%E6%B7%B7%E5%90%88%E7%9A%84%E8%B5%B7%E6%BA%90&rft.jtitle=%E6%97%A5%E6%9C%AC%E7%B5%84%E7%B9%94%E9%81%A9%E5%90%88%E6%80%A7%E5%AD%A6%E4%BC%9A%E8%AA%8C&rft.au=%E7%B4%B0%E9%81%93%2C+%E4%B8%80%E5%96%84&rft.au=%E5%85%89%E6%B0%B8%2C+%E6%BB%8B%E6%A8%B9&rft.au=%E4%B8%AD%E5%B2%A1%2C+%E5%8D%9A%E5%8F%B2&rft.au=%E4%BA%95%E3%83%8E%E4%B8%8A%2C+%E9%80%B8%E6%9C%97&rft.date=2014-04-20&rft.pub=%E6%97%A5%E6%9C%AC%E7%B5%84%E7%B9%94%E9%81%A9%E5%90%88%E6%80%A7%E5%AD%A6%E4%BC%9A&rft.issn=2186-9995&rft.eissn=2187-4239&rft.volume=21&rft.issue=1&rft.spage=37&rft.epage=44&rft_id=info:doi/10.12667%2Fmhc.21.37&rft.externalDocID=article_mhc_21_1_21_37_article_char_ja |
| thumbnail_l | http://covers-cdn.summon.serialssolutions.com/index.aspx?isbn=/lc.gif&issn=2186-9995&client=summon |
| thumbnail_m | http://covers-cdn.summon.serialssolutions.com/index.aspx?isbn=/mc.gif&issn=2186-9995&client=summon |
| thumbnail_s | http://covers-cdn.summon.serialssolutions.com/index.aspx?isbn=/sc.gif&issn=2186-9995&client=summon |