Before and after the neck: to understand the vertebrate body elaboration from the perspective of muscle, skeleton, and nerve development

• Project/Area Number: 15H02416
• Research Category: Grant-in-Aid for Scientific Research (A)
• Allocation Type: Single-year Grants
• Section: 一般
• Research Field: Evolutionary biology
• Research Institution: Institute of Physical and Chemical Research
• Principal Investigator: Shigeru Kuratani 国立研究開発法人理化学研究所, 倉谷形態進化研究室, 主任研究員 (00178089)
• Project Period (FY): 2015-04-01 – 2018-03-31
• Project Status: Completed (Fiscal Year 2017)
• Budget Amount: ¥39,520,000 (Direct Cost: ¥30,400,000、Indirect Cost: ¥9,120,000); Fiscal Year 2017: ¥11,700,000 (Direct Cost: ¥9,000,000、Indirect Cost: ¥2,700,000); Fiscal Year 2016: ¥11,700,000 (Direct Cost: ¥9,000,000、Indirect Cost: ¥2,700,000); Fiscal Year 2015: ¥16,120,000 (Direct Cost: ¥12,400,000、Indirect Cost: ¥3,720,000)
• Keywords: 脊椎動物 / 頭頸部 / 発生 / 進化 / 中胚葉 / 体節 / 体壁 / ナメクジウオ / 形態進化 / 比較発生学 / 実験発生学

Outline of Final Research Achievements

ertebrate can be characterized by the posession of head mesoderm. Through the study using amphioxus, an animal with no head mesoderm, we found that the head mesoderm was obtained by topographical shift of early mesoderm, and by suppression of this shift, we showed that amphioxus-like mesoderm could be partially phenocopied in vertebrate embryos.
Cyclostomes were employed for the study of hypobranchial muscle (HBM) development. Through comparative embryological observation and cell lineage analyses, we found that the deep position of the gnathostome HBM arose as the result of the shift of relative positions between the pharynx and pericardium. In the ongoing study using elasmobranch (shark) embryos, we have so far found that two paralogues of Lbx genes are diferentially expressed in the anterior and posterior portions of the HBM primordia. Of those, cyclostome HBM resemble more the posterior. We suspect gene duplication behind the novel patterns of muscle.

Research Products

• [Int'l Joint Research] Uppsala University(スウェーデン)
• [Int'l Joint Research] Academia Sinica(台湾)
• [Int'l Joint Research] Chinese Academy of Sciences(中国)
• [Int'l Joint Research] University of Essex(英国)
• [Int'l Joint Research] Universidad de Navarra(スペイン)
• [Journal Article] Hagfish and lamprey Hox genes reveal conservation of temporal colinearity in vertebrates (2018)
  • Author(s): Pascual-Anaya J, Sato I, Sugahara F, Higuchi S, Paps J, Ren Y, Takagi W, Ruiz-Villalba A, Ota KG, Wang W, Kuratani S.
  • Journal Title: Nat Ecol Evol. Volume: 2 Issue: 5 Pages: 859-866
  • DOI: 10.1038/s41559-018-0526-2
  • Peer Reviewed / Int'l Joint Research
• [Journal Article] Expansions, diversification, and interindividual copy number variations of AID/APOBEC family cytidine deaminase genes in lampreys. (2018)
  • Author(s): Holland SJ, Berghuis LM, King JJ, Iyer LM, Sikora K, Fifield H, Peter S, Quinlan EM, Sugahara F, Shingate P, Trancoso I, Iwanami N, Temereva E, Strohmeier C, Kuratani S, Venkatesh B, Evanno G, Aravind L, Schorpp M, Larijani M, Boehm T.
  • Journal Title: Proc Natl Acad Sci U S A. Volume: 115 Issue: 14
  • DOI: 10.1073/pnas.1720871115
  • Peer Reviewed / Open Access / Int'l Joint Research
• [Journal Article] Development of hypobranchial muscles with special reference to the evolution of the vertebrate neck. (2018)
  • Author(s): Adachi N, Pascual-Anaya J, Hirai T, Higuchi S, Kuratani S.
  • Journal Title: Zoological Lett Volume: 4 Issue: 1 Pages: 5-5
  • DOI: 10.1186/s40851-018-0087-x
  • Peer Reviewed / Open Access
• [Journal Article] The neural crest and evolution of the head/trunk interface in vertebrates (2018)
  • Author(s): Kuratani S, Kusakabe R, Hirasawa T
  • Journal Title: Developmental Biology Volume: in press Pages: 30922-30923
  • DOI: 10.1016/j.ydbio.2018.01.017
  • Peer Reviewed / Open Access
• [Journal Article] Evolution of the vertebrate neurocranium: problems of the premandibular domain and the origin of the trabecula. (2018)
  • Author(s): Kuratani S, Ahlberg PE.
  • Journal Title: Zoological Lett. Volume: 4 Issue: 1 Pages: 1-1
  • DOI: 10.1186/s40851-017-0083-6
  • Peer Reviewed / Open Access / Int'l Joint Research
• [Journal Article] Metamerism in cephalochordates and the problem of the vertebrate head. (2017)
  • Author(s): Onai T, Adachi N, Kuratani S.
  • Journal Title: Int J Dev Biol Volume: 61 Issue: 10-11-12 Pages: 621-632
  • DOI: 10.1387/ijdb.170121to
  • Peer Reviewed
• [Journal Article] 1.Anatomical integration of the sacral-hindlimb unit coordinated by GDF11 underlies variation in hindlimb positioning in tetrapods. (2017)
  • Author(s): Matsubara Y., Hirasawa T., Egawa S., Hattori A., Suganuma T., Kohara Y., Nagai T., Tamura K., Kuratani S., Kuroiwa A., Suzuki T
  • Journal Title: Nature Ecology and Evolution Volume: １ Issue: 9 Pages: 1392-1399
  • DOI: 10.1038/s41559-017-0247-y
  • Peer Reviewed
• [Journal Article] Migratory appendicular muscles precursor cells in the common ancestor to all vertebrates. (2017)
  • Author(s): Eri Okamoto, Rie Kusakabe, Shigehiro Kuraku, Susumu Hyodo, Alexandre Robert-Moreno, Koh Onimaru, James Sharpe, Shigeru Kuratani and Mikiko Tanaka
  • Journal Title: Nat. Ecol. Evol. Volume: 1 Issue: 11 Pages: 1731-1736
  • DOI: 10.1038/s41559-017-0330-4
  • Peer Reviewed / Int'l Joint Research
• [Journal Article] Constrained vertebrate evolution by pleiotropic genes. (2017)
  • Author(s): Hu H, Uesaka M, Guo S, Shimai K, Lu TM, Li F, Fujimoto S, Ishikawa M, Liu S, Sasagawa Y, Zhang G, Kuratani S, Yu JK, Kusakabe TG, Khaitovich P, Irie N; EXPANDE Consortium.
  • Journal Title: Nat Ecol Evol Volume: 1 Issue: 11 Pages: 1722-1730
  • DOI: 10.1038/s41559-017-0318-0
  • Peer Reviewed / Int'l Joint Research
• [Journal Article] Nitric Oxide regulates mouth development in amphioxus. (2017)
  • Author(s): Annona G, Caccavale F, Pascual-Anaya J, Kuratani S, De Luca P, Palumbo A, D'Aniello S.
  • Journal Title: Sci Rep Volume: 7 Issue: 1 Pages: 8432-8432
  • DOI: 10.1038/s41598-017-08157-w
  • Peer Reviewed / Int'l Joint Research
• [Journal Article] Differing contributions of the first and second pharyngeal arches to tympanic membrane formation in the mouse and chick. (2017)
  • Author(s): Furutera T, Takechi M, Kitazawa T, Takei J, Yamada T, Vu Hoang T, Rijli FM, Kurihara H, Kuratani S, Iseki S.
  • Journal Title: Development. Volume: 144 Issue: 18 Pages: 3315-3324
  • DOI: 10.1242/dev.149765
  • Peer Reviewed / Int'l Joint Research
• [Journal Article] Expression patterns of Sema3A in developing amniote limbs: With reference to the diversification of peripheral nerve innervation. (2017)
  • Author(s): Noguchi K, Ishikawa R, Kawaguchi M, Miyoshi K, Kawasaki T, Hirata T, Fukui M, Kuratani S, Tanaka M, Murakami Y.
  • Journal Title: Dev Growth Differ. Volume: 59(4) Issue: 4 Pages: 270-285
  • DOI: 10.1111/dgd.12364
  • Peer Reviewed / Int'l Joint Research
• [Journal Article] Outflow tract septation and the aortic arch system in reptiles: lessons for understanding the mammalian heart (2017)
  • Author(s): Poelmann RE, Gittenberger-de Groot AC, Biermans MWM, Dolfing AI, Jagessar A, van Hattum S, Hoogenboom A, Wisse LJ, Vicente-Steijn R, de Bakker MAG, Vonk FJ, Hirasawa T, Kuratani S, Richardson MK
  • Journal Title: EvoDevo Volume: 8 Issue: 1 Pages: 9-9
  • DOI: 10.1186/s13227-017-0072-z
  • Peer Reviewed / Open Access / Int'l Joint Research
• [Journal Article] Reconstructing the ancestral vertebrate brain. (2017)
  • Author(s): Sugahara F, Murakami Y, Pascual-Anaya J, Kuratani S.
  • Journal Title: Dev Growth Differ. Volume: 59(4) Issue: 4 Pages: 163-174
  • DOI: 10.1111/dgd.12347
  • Peer Reviewed / Open Access / Int'l Joint Research
• [Journal Article] Expression patterns of Sema3A in developing amniote limbs: with reference to the diversification of peripheral nerve innervation (2017)
  • Author(s): Noguchi, K., Ishikawa, R., Miyoshi, K., Kawaguchi, M., Kawasaki, T., Hirata, T., Fukui, M., Kuratani, S., Tanaka, M., and Murakami, Y.
  • Journal Title: Dev. Growth Diff. Volume: 印刷中
  • Peer Reviewed
• [Journal Article] Reconstruction of the ancestral vertebrate brain. (2017)
  • Author(s): Sugahara, F., Murakami, Y., Pascual-Anaya, J., and Kuratani, S.
  • Journal Title: Dev. Growth Diff. Volume: 印刷中
  • Peer Reviewed
• [Journal Article] Paleontological studies integrated into a new evolutionary zoology. (2017)
  • Author(s): Kuratani, S., and Fukatsu, T.
  • Journal Title: Zool. Sci. Volume: 34 Issue: 1 Pages: 1-4
  • DOI: 10.2108/zs160203
  • NAID: 40021108431
  • Peer Reviewed
• [Journal Article] Palaeospondylus as a primitive hagfish. (2016)
  • Author(s): Hirasawa, T., Oisi, Y., and Kuratani, S.
  • Journal Title: Zool. Lett. Volume: 2 Issue: 1 Pages: 20-20
  • DOI: 10.1186/s40851-016-0057-0
  • Peer Reviewed / Open Access
• [Journal Article] On the vagal cardiac nerves, with special reference to the early evolution of the head-trunk-interface. (2016)
  • Author(s): Higashiyama, H., Hirasawa, T., Oisi, Y., Sugahara, F., Hyodo, S., Kanai, Y., and Kuratani, S.
  • Journal Title: J. Morphol. Volume: 277 Issue: 9 Pages: 1146-1158
  • DOI: 10.1002/jmor.20563
  • Peer Reviewed
• [Journal Article] Evolution of the vertebrate cranium: viewed from the hagfish developmental studies. (2016)
  • Author(s): Kuratani, S., Oisi, Y., and Ota, K. G.
  • Journal Title: Zool. Sci. Volume: 33 Issue: 3 Pages: 229-238
  • DOI: 10.2108/zs150187
  • NAID: 210000168430
  • Peer Reviewed / Int'l Joint Research
• [Journal Article] What are head cavities? - History of studies on the vertebrate head segmentation. (2016)
  • Author(s): Kuratani, S., and Adachi, N.
  • Journal Title: Zool. Sci. Volume: 33 Issue: 3 Pages: 213-228
  • DOI: 10.2108/zs150181
  • NAID: 40020859763
  • Peer Reviewed
• [Journal Article] News & Views: Getting the measure of a monster. (2016)
  • Author(s): Kuratani, S., and Hirasawa, T
  • Journal Title: Nature Volume: 532 Issue: 7600 Pages: 447-448
  • DOI: 10.1038/nature17885
  • Peer Reviewed
• [Journal Article] Morphology and development of extra-ocular muscles in the lamprey reveals the ancestral head structure and its developmental mechanism of vertebrates. (2016)
  • Author(s): Suzuki, G. D., Fukumoto, Y., Kusakabe, R., Yamazaki, Y., Kosaka, J., Kuratani, S., and Wada, H.
  • Journal Title: Zoological Letters Volume: in press
  • Peer Reviewed / Open Access
• [Journal Article] Evolution of the vertebrate cranium: viewed from the hagfish developmental studies. (2016)
  • Author(s): Kuratani, S., Oisi, Y., and Ota, K. G.
  • Journal Title: Zoological Science Volume: ih press
  • NAID: 210000168430
  • Peer Reviewed / Int'l Joint Research
• [Journal Article] What are head cavities? - History of studies on the vertebrate head segmentation (2016)
  • Author(s): Kuratani, S., and Adachi, N.
  • Journal Title: Zoological Science Volume: in press
  • NAID: 40020859763
  • Peer Reviewed
• [Journal Article] Evidence from cyclostomes for complex regionalization of the ancestral vertebrate brain (2016)
  • Author(s): Fumiaki Sugahara, Juan Pascual-Anaya, Yasuhiro Oisi, Shigehiro Kuraku, Shin-IchiAota, Noritaka Adachi, Wataru Takagi, Tamami Hirai, Noboru Sato, Yasunori Murakami, Shigeru Kuratani
  • Journal Title: Nature Volume: 531 Issue: 7592 Pages: 97-100
  • DOI: 10.1038/nature16518
  • Peer Reviewed / Open Access / Int'l Joint Research
• [Journal Article] Launch of Zoological Letters. (2016)
  • Author(s): Fukatsu T, Kuratani S.
  • Journal Title: Zoological Science Volume: 33 Issue: 1 Pages: 1-5
  • DOI: 10.2108/zs150160
  • NAID: 40020738702
  • Peer Reviewed / Open Access
• [Journal Article] Developmental mechanisms of the tympanic membrane in mammals and non-mammalian amniotes (2016)
  • Author(s): Takechi, M., Kitazawa, T., Hirasawa, T., Hirai, T., Iseki, S., Kurihara, H., Kuratani, S.
  • Journal Title: Congenital Anomalies Volume: 56 Issue: 1 Pages: 12-17
  • DOI: 10.1111/cga.12132
  • Peer Reviewed / Open Access / Acknowledgement Compliant
• [Journal Article] Expansion of the neck reconstituted the shoulder-diaphragm in amniote evolution (2016)
  • Author(s): Hirasawa T, Fujimoto S, Kuratani S
  • Journal Title: Development, Growth & Differentiation Volume: 58 Issue: 1 Pages: 143-153
  • DOI: 10.1111/dgd.12243
  • Peer Reviewed / Open Access
• [Journal Article] On the origin of vertebrate somites (2016)
  • Author(s): Onai T, Aramaki T, Inomata H, Hirai T, Kuratani S.
  • Journal Title: Zoological Letters Volume: 1 Issue: 1 Pages: 33-33
  • DOI: 10.1186/s40851-015-0033-0
  • Peer Reviewed / Open Access
• [Journal Article] Ancenstral mesodermal reorganization and evolution of the vertebrate head (2016)
  • Author(s): Onai, T., Aramaki, T., Inomata, H., Hirai, T., and Kuratani, S.
  • Journal Title: Zoological Letters Volume: 1 Issue: 1 Pages: 29-29
  • DOI: 10.1186/s40851-015-0030-3
  • Peer Reviewed / Open Access
• [Journal Article] Evolution of retinoic acid receptors in chordates: insights from three lamprey species, Lampetra fluviatilis, Petromyzon marinus and Lethenteron japonicum. (2015)
  • Author(s): Campo-Paysaa, F., Jandzik, D., Takio-Ogawa, Y., Cattell, M. V, Neef, H. C., Langeland, J. A., Kuratani, S., Medeiros, D. M., Mazan, S., Kuraku, S., Laudet, V., and Schubert, M.
  • Journal Title: EvoDevo Volume: 6 Issue: 1 Pages: 18-18
  • DOI: 10.1186/s13227-015-0016-4
  • Peer Reviewed / Open Access / Int'l Joint Research
• [Journal Article] Developmental genetic bases behind the independent origin of the tympanic membrane in mammals and diapsids. (2015)
  • Author(s): Kitazawa T, Takechi M, Hirasawa T, Adachi N, Narboux-Neme N, Kume H, Maeda K, Hirai T, Miyagawa-Tomita S, Kurihara Y, Hitomi J, Levi G, Kuratani S, Kurihara H
  • Journal Title: Nature Communications Volume: 6 Issue: 1 Pages: 6853-6853
  • DOI: 10.1038/ncomms7853
  • Peer Reviewed / Open Access / Acknowledgement Compliant
• [Journal Article] On the homology of the shoulder girdle in turtles. (2015)
  • Author(s): Nagashima, H., Sugahara, F., Takechi, M., Sato, N., and Kuratani, S.
  • Journal Title: J. Exp. Zool. (Mol. Dev. Evol.). Volume: 324 Issue: 3 Pages: 244-254
  • DOI: 10.1002/jez.b.22584
  • Peer Reviewed
• [Journal Article] The evolutionary origin of the turtle shell grounded on the axial arrest of the embryonic rib cage. (2015)
  • Author(s): Hirasawa, T., Pascual-Anaya, J., Kamezaki, N., Taniguchi, N., Mine, K., and Kuratani, S.
  • Journal Title: J. Exp. Zool. (Mol. Dev. Evol. Volume: 324 Issue: 3 Pages: 194-207
  • DOI: 10.1002/jez.b.22579
  • Peer Reviewed
• [Presentation] “Does the ontogeny recapitulate phylogeny?“（個体発生は系統発生を繰り返すのか？) (2017)
  • Author(s): Shigeru Kuratani
  • Organizer: 50th Annual Meeting for the Japanese Society of Developmental Biologists
  • Int'l Joint Research / Invited
• [Presentation] Acquistion of The Turtle Shell: Changes in Developmental Program Behind The Evolutionary Novelty (2017)
  • Author(s): Shigeru Kuratani
  • Organizer: 18th International Congress of Developmental Biology
  • Int'l Joint Research / Invited
• [Presentation] かたちの進化と発生 (2017)
  • Author(s): 倉谷 滋
  • Organizer: 名古屋大学EGER グリーン自然科学レクチャー 2017
  • Invited
• [Presentation] Development of the hagfish and origin of the gnathostomehead (2017)
  • Author(s): Shigeru Kuratani
  • Organizer: Zoological Letters Symposium:Early Evolution and Diversity of Vertebrates
  • Place of Presentation: 理化学研究所CDB (神戸市）
  • Int'l Joint Research
• [Presentation] Timing of skeletal muscle development during primary body wall transformation in amniote embryos. (2016)
  • Author(s): Tatsuya Hirasawa, Satoko Fujimoto, Shigeru Kuratani
  • Organizer: CDB Symposium 2015 Time in Development.
  • Place of Presentation: 理化学研究所CDB(神戸市)
  • Year and Date: 2016-03-23
  • Int'l Joint Research
• [Presentation] 脊椎動物の初期進化を考える-発生からみたボディプランのオリジン (2016)
  • Author(s): 倉谷 滋
  • Organizer: The 10th Nich Neuro-Angiology Conference 2016
  • Place of Presentation: 大阪科学技術センター(大阪市）
  • Invited
• [Presentation] カメを作った発生プログラムの進化 (2016)
  • Author(s): 倉谷 滋
  • Organizer: 化石研究会第34回総会・学術大会 シンポジウム「カメ類の起源と進化を考える」
  • Place of Presentation: 早稲田大学・早稲田キャンパス(東京都新宿区）
  • Invited
• [Presentation] Evolution and development of the head in agnathans and fishes (2016)
  • Author(s): Kuratani S, Sugahara F, Ohta K, Oisi Y.
  • Organizer: The 11th International Congress of Vertebrate Morphology
  • Place of Presentation: ソルトレイクシティ、アメリカ
  • Int'l Joint Research / Invited
• [Presentation] 脊椎動物の比較形態学と進化発生学 (2016)
  • Author(s): 倉谷 滋
  • Organizer: 日本進化学会第18回大会
  • Place of Presentation: 東京工業大学大岡山キャンパス（東京都目黒区）
• [Presentation] Development of hagfish and early evolution of vertebrates (2016)
  • Author(s): Shigeru Kuratani
  • Organizer: International Conference on Lamprey Immune System.
  • Place of Presentation: 大連、中国
  • Int'l Joint Research / Invited
• [Presentation] Craniofacial development and evolution in vertebrates (2016)
  • Author(s): Shigeru Kuratani
  • Organizer: The Joint meeting of The 22nd International Congress of Zoology and the 87th meeting of Zoological Society of Japan,
  • Place of Presentation: 沖縄科学技術大学院大学（沖縄県恩納村）
  • Int'l Joint Research
• [Presentation] Embryology of the hagfish and the early evolution of vertebrates (2016)
  • Author(s): Shigeru Kuratani
  • Organizer: The Joint meeting of The 22nd International Congress of Zoology and the 87th meeting of Zoological Society of Japan
  • Place of Presentation: 沖縄科学技術大学院大学（沖縄県恩納村）
  • Int'l Joint Research / Invited
• [Presentation] カメの甲はどうやって出来たか (2015)
  • Author(s): 倉谷 滋
  • Organizer: 日本進化学会第17回大会 シンポジウム「複合適応形質進化の遺伝子基盤解明
  • Place of Presentation: 中央大学（東京都）
  • Year and Date: 2015-08-20
• [Presentation] Hagfish development and early evolution of vertebrates (2015)
  • Author(s): Shigeru Kuratani
  • Organizer: Palaeontological and Molecular Approaches to Early Vertebrate Evolution
  • Place of Presentation: Ｕｐｐｓａｌａ (Sweden)
  • Year and Date: 2015-05-12
  • Int'l Joint Research / Invited
• [Book] 新版・動物進化形態学 - Evolutionary Morphology: Bauplan and Embryonic Development of Vertebrates (2017)
  • Author(s): 倉谷 滋
  • Total Pages: 768
  • Publisher: 東京大学出版会.
• [Book] 分節幻想：脊椎動物の起源をめぐる科学思想史 (2016)
  • Author(s): 倉谷 滋
  • Total Pages: 864
  • Publisher: 工作舎
• [Book] 生物学はどこまで進むのかin:「これからどうする」 (2016)
  • Author(s): 倉谷 滋（分担）
  • Total Pages: 3
  • Publisher: 岩波書店.
• [Book] 変容する形態 - 比較形態学のすすめ」in「科学者の目、科学の芽」 (2016)
  • Author(s): 倉谷 滋（分担）
  • Total Pages: 2
  • Publisher: 岩波科学ライブラリー
• [Book] In Situ Hybridization Methods, Neuromethods, vol. 99 Chapter 13 In Situ Hybridization Methods, Neuromethods, vol. 99 Chapter 13 Gene Expression Analysis of Lamprey Embryos. (2015)
  • Author(s): Giselbert Hauptmann 編集 Fumiaki Sugahara, Yasunori Muratakami, and Shigeru Kutratani
  • Total Pages: 16
  • Publisher: Springer Protocols. Humana Press
• [Book] In situ Hybridization Methods, Neuromethods, Vol.99. Chapter12 Analysis of Embryonic Gene Expression Patterns in the Hagfish. (2015)
  • Author(s): Giselbert Hauptmann 編集 Yasuhiro Oisi, Osamu Kakitani, Shigeru Kuratani, and Kinya G. Ota
  • Total Pages: 16
  • Publisher: Springer Protocols. Humana Press
• [Book] 形態学-形づくりにみる動物進化のシナリオ (2015)
  • Author(s): 倉谷 滋
  • Total Pages: 211
  • Publisher: 丸善出版
• [Book] 「カメの甲の初期進化：約2億5000万年前に何が起きたのか？」in 「細胞工学 別冊 進化の謎をゲノムで解く」 (2015)
  • Author(s): 監修 長谷部光泰（分担：平沢 達矢、Juan Pascual-Anaya, 倉谷 滋）
  • Total Pages: 10
  • Publisher: 秀潤社
• [Remarks] 研究所HP
  • URL: http://www.riken.jp/research/labs/chief/evol_morphol/
• [Remarks] 研究室HP
  • URL: http://www.cdb.riken.jp/emo/japanese/indexj.html
• [Remarks] 理化学研究所 倉谷形態進化研究室
  • URL: http://www.riken.jp/research/labs/chief/evol_morphol/
• [Remarks] 研究室オリジナルHP
  • URL: http://www.cdb.riken.jp/emo/japanese/indexj.html