We are enjoying the research to explore the biological significance of sperm cells and eukaryotic cilia and flagella. The main experimental materials used in this lab are tunicates (Ciona intestinalis), salmon, trout, flounder, sea urchin, snails, comb jelly, Japanese spiny lobster, etc, most of which can be collected near our lab.

MEMBERS (2019)

Kazuo Inaba: Professor 【USJI Report】

Kogiku Shiba: Assistant Professor

Nana Terauchi: JSPS Postdoctoral Fellow

Ryuji Yanase: Postdoctoral Fellow

Misa Shibata: Postdoctoral Fellow

Kei Jokura: Graduate student, Graduate School of Life and Environmental Sciences

Sayaka Yamaguchi: Graduate student, Graduate School of Life and Environmental Sciences

Masae Ohata: Technical Assistant


Biology of Sperm: Molecular characterization of sperm flagella, Molecular mechanism of flagellar motility and its regulation, Sperm activation and chemotaxis by egg-derived substances, Ubiquitin/proteasome system in sperm function, EST/genome/poroteomic analyses of testis-expressed genes and proteins*, Molecular mechanism of spermatogenesis, Comparative biochemistry of sperm protein, Molecular diversity of sperm protein.

Biology of cilia and flagella: Structure and function of axonemal dyneins, Regulation of flagellar motility by protein kinase and phosphatase, Molecular architecture of axoneme, cDNA and proteomic analysis of axonemal protein, in vitro assembly of axoneme, Functions of cilia and flagella in development, CNS and immune system, Phylogenetic analysis of ciliary and flagellar proteins. Evolution and diversity of cilia and flagella.

Post-genomic analysis of marine invertebrates: Proteomic analysis of embryonic development and adult tissues of the ascidian Ciona intestinalis, Ciona intestinalis protein database (CIPRO**), application of the system for proteomic analysis in Ciona to other marine invertebrates, such as sea urchins

*We have established a system for peptide mass finger printing using MSCITS and PerMS.

** http://cipro.ibio.jp/current/



  1. Miyata M, Robinson RC, Uyeda TQP, Fukumori Y, Fukushima SI, Haruta S, Homma M, Inaba K, Ito M, Kaito C, Kato K, Kenri T, Kinosita Y, Kojima S, Minamino T, Mori H, Nakamura S, Nakane D, Nakayama K, Nishiyama M, Shibata S, Shimabukuro K, Tamakoshi M, Taoka A, Tashiro Y, Tulum I, Wada H, Wakabayashi KI. Tree of motility - A proposed history of motility systems in the tree of life. Genes Cells. 2020 Jan;25(1):6-21. doi: 10.1111/gtc.12737. Review.
  2. Morohoshi A, Miyata H, Shimada K, Nozawa K, Matsumura T, Yanase R, Shiba K, Inaba K, Ikawa M. Nexin-Dynein regulatory complex component DRC7 but not FBXL13 is required for sperm flagellum formation and male fertility in mice. PLoS Genet. 2020 Jan 21;16(1):e1008585. doi: 10.1371/journal.pgen.1008585. eCollection 2020 Jan.


  1. Jokura K, Shibata D, Yamaguchi K, Shiba K, Makino Y, Shigenobu S, Inaba K.
    CTENO64 Is Required for Coordinated Paddling of Ciliary Comb Plate in Ctenophores. Curr Biol. 2019 Oct 21;29(20):3510-3516.e4. doi: 10.1016/j.cub.2019.08.059. Epub 2019 Oct 10.
  2. Sasaki K, Shiba K, Nakamura A, Kawano N, Satouh Y, Yamaguchi H, Morikawa M, Shibata D, Yanase R, Jokura K, Nomura M, Miyado M, Takada S, Ueno H, Nonaka S, Baba T, Ikawa M, Kikkawa M, Miyado K, Inaba K. Calaxin is required for cilia-driven determination of vertebrate laterality. Commun Biol. 2019 Jun 20;2:226. doi: 10.1038/s42003-019-0462-y. eCollection 2019.
  3. Nakayama T, Nomura M, Takano Y, Tanifuji G, Shiba K, Inaba K, Inagaki Y, Kawata M. Single-cell genomics unveiled a cryptic cyanobacterial lineage with a worldwide distribution hidden by a dinoflagellate host. Proc Natl Acad Sci U S A. 2019 Jun 24. pii: 201902538. doi: 10.1073/pnas.1902538116. [Epub ahead of print]
  4. Inaba K. Regulatory Mechanism of Axonemal Dynein. In: Handbook of Dynein (Second Edition) (K Hirose, Ed), CRC Press, 2019 May; pp.375-410, ISBN: 9789814800013.
  5. Itou T, Kanno M, Suyama Y, Inaba K, Aoki MN. Opening the black box: microspatial patterns of zoospore dispersal, parentage, and selfing in the kelp Ecklonia cava as revealed by microsatellite markers. Journal of Applied Phycology, First Online: 06 April 2019. doi: 10.1007/s10811-019-01790-0
  6. Kinoshita-Terauchi N, Shiba K, Umezawa T, Matsuda F, Motomura T, Inaba K. A brown algal sex pheromone reverses the sign of phototaxis by cAMP/Ca2+-dependent signaling in the male gametes of Mutimo cylindricus (Cutleriaceae). J Photochem Photobiol B. 2019 Mar;192:113-123. doi: 10.1016/j.jphotobiol.2019.01.010. Epub 2019 Jan 29.
  7. Nomura M, Atsuji K, Hirose K, Shiba K, Yanase R, Nakayama T, Ishida KI, Inaba K. Microtubule stabilizer reveals requirement of Ca2+-dependent conformational changes of microtubules for rapid coiling of haptonema in haptophyte algae. Biol Open. 2019 Feb 11;8(2). pii: bio036590. doi: 10.1242/bio.036590.


  1. Yoshida K, Shiba K, Sakamoto A, Ikenaga J, Matsunaga S, Inaba K, Yoshida M. Ca2+ efflux via plasma membrane Ca2+-ATPase mediates chemotaxis in ascidian sperm. Sci Rep. 2018 Nov 9;8(1):16622. doi: 10.1038/s41598-018-35013-2.
  2. Harvey BP, Agostini S, Wada S, Inaba K, Hall-Spencer JM. Dissolution: The Achilles’ Heel of the Triton Shell in an Acidifying Ocean. Front. Mar. Sci. 2018 Oct 12; 5:371. doi: 10.3389/fmars.2018.00371
  3. Mizotani Y, Suzuki M, Hotta K, Watanabe H, Shiba K, Inaba K, Tashiro E, Oka K, Imoto M. 14-3-3εa directs the pulsatile transport of basal factors toward the apical domain for lumen growth in tubulogenesis. Proc Natl Acad Sci U S A. 2018 Sep 18;115(38):E8873-E8881. doi: 10.1073/pnas.1808756115. Epub 2018 Aug 29.
  4. Momose T, De Cian A, Shiba K, Inaba K, Giovannangeli C, Concordet JP. High doses of CRISPR/Cas9 ribonucleoprotein efficiently induce gene knockout with low mosaicism in the hydrozoan Clytia hemisphaerica through microhomology-mediated deletion. Sci Rep. 2018 Aug 6;8(1):11734. doi: 10.1038/s41598-018-30188-0.
  5. Kamiya R, Shiba K, Inaba K, Kato-Minoura T. Release of sticky glycoproteins from Chlamydomonas flagella during microsphere translocation on the surface membrane. Zoolog Sci. 2018 Apr; doi: 10.2108/zs180025 [Early view]
  6. Hill RW, Armstrong EJ, Inaba K, Morita M, Tresguerres M, Stillman JH, Roa JN, Kwan GT. Acid secretion by the boring organ of the burrowing giant clam, Tridacna crocea. Biol Lett. 2018 Jun;14(6). pii: 20180047. doi: 10.1098/rsbl.2018.0047.
  7. Inaba K, Shiba K. Microscopic analysis of sperm movement: links to mechanisms and protein components. Microscopy (Oxf). 2018 Jun 1;67(3):144-155. doi: 10.1093/jmicro/dfy021.
  8. Ugwu SI, Shiba K, Inaba K, Morita M. A Unique Seminal Plasma Protein, Zona Pellucida 3-Like Protein, has Ca2+ -Dependent Sperm Agglutination Activity. Zoolog Sci. 2018 Apr;35(2):161-171. doi: 10.2108/zs170150.
  9. Morita N, Inaba K, Saito Y. Post-Embryonic Development and Genital-Complex Formation in Three Species of Polyclad Flatworms. Zoolog Sci. 2018 Feb;35(1):28-38. doi: 10.2108/zs170114.
  10. Shojima T, Hou F, Takahashi Y, Matsumura Y, Okai M, Nakamura A, Mizuno K, Inaba K, Kojima M, Miyakawa T, Tanokura M. Crystal structure of a Ca2+-dependent regulator of flagellar motility reveals the open-closed structural transition. Sci Rep. 2018 Jan 31;8(1):2014. doi: 10.1038/s41598-018-19898-7.


  1. Inaba K. Biochemical purification of axonemal and cytoplasmic dyneins. In: Dyneins - Dynein Mechanics, Dysfunction, and Disease (2nd Edition) (SM King, Ed), pp. 89-112, Nov 2017. eBook ISBN: 9780128097014.
  2. Mizuno K, Shiba K, Yaguchi J, Shibata D, Yaguchi S, Prulière G, Chenevert J, Inaba K. Calaxin establishes basal body orientation and coordinates movement of monocilia in sea urchin embryos. Sci Rep. 2017 Sep 7;7(1):10751. doi: 10.1038/s41598-017-10822-z.
  3. Yamamoto R, Obbineni JM, Alford LM, Ide T, Owa M, Hwang J, Kon T, Inaba K, James N, King SM, Ishikawa T, Sale WS, Dutcher SK. Chlamydomonas DYX1C1/PF23 is essential for axonemal assembly and proper morphology of inner dynein arms. PLoS Genet. 2017 Sep 11;13(9):e1006996. doi: 10.1371/journal.pgen.1006996. eCollection 2017 Sep.
  4. L. Valdés (eds), M Cargo, H. Oksfeldt, HE Garcia, L Horn, K Inaba, L Inniss, K Isensee, B Kelly, YH Lee, J Mees, G Reed, S Pae, L Pikula, P Pissierssens, L Raymond, M Scraper, A Simcock, AH Trios. UNESCO. Global Ocean Science Report—The current status of ocean science around the world. UNESCO Publishing, Paris, ISBN 978-92-3-100226-7 (July 2017).
  5. Shiba K, Inaba K. Inverse relationship of Ca2+-dependent flagellar response between animal sperm and prasinophyte algae. J Plant Res. 2017 May;130(3):465-473. doi: 10.1007/s10265-017-0931-7.


  1. Konno A, Ikegami K, Konishi Y, Yang HJ, Abe M, Yamazaki M, Sakimura K, Yao I, Shiba K, Inaba K, Setou M. Doublet 7 shortening, doublet 5-preferential poly-Glu reduction, and beating stall of sperm flagella in Ttll9-/- mice. J Cell Sci. 2016 Jul 15;129(14):2757-66. doi: 10.1242/jcs.185983. Epub 2016 Jun 2.
  2. Yaguchi J, Takeda N, Inaba K, Yaguchi S. Cooperative Wnt-Nodal Signals Regulate the Patterning of Anterior Neuroectoderm. PLoS Genet. 2016 Apr 21;12(4):e1006001. doi: 10.1371/journal.pgen.1006001. eCollection 2016 Apr.
  3. Kinoshita N, Shiba K, Inaba K, Fu G, Nagasato C and Motomura T. Flagellar waveforms of gametes in the brown alga Ectocarpus siliculosus. Eur J Phycol, 2016 April (2015 Dec, online): 51(2): 139-148.
    doi: 10.1080/09670262.2015.1109144.


  1. Matsuzaki M, Mizushima S, Hiyama G, Hirohashi N, Shiba K, Inaba K, Suzuki T, Dohra H, Ohnishi T, Sato Y, Kohsaka T, Ichikawa Y, Atsumi Y, Yoshimura T, Sasanami T. Lactic acid is a sperm motility inactivation factor in the sperm storage tubules. Sci Rep. 2015 Dec 1;5:17643. doi: 10.1038/srep17643.
  2. Inaba K. Japanese marine biological stations: preface to the special issue. Reg Stud Mar Sci, 2015 Nov; 2:154–157. doi:10.1016/j.rsma.2015.09.005
  3. Agostini S, Wada S, Kon K, Omori A, Kohtsuka H, Fujimura H, Tsuchiya Y, Sato T, Shinagawa H, Yamada Y, Inaba K. Geochemistry of two shallow CO2 seeps in Shikine Island (Japan) and their potential for ocean acidification research. Reg Stud Mar Sci, 2015 Nov; 2:45-53. doi:10.1016/j.rsma.2015.07.004
  4. Cosson J, Prokopchuk G, Inaba K. The flagellar mechanics of spermatozoa and its regulation. In: Flagellar Mechanics and Sperm Guidance (Cosson J, Ed), Bentham Science Publishers, 2015 Nov; pp.3-134. doi: 10.2174/97816810812811150101
  5. Yoshida M, Inaba K. Sperm chemotaxis in urochordates. In: Flagellar Mechanics and Sperm Guidance (Cosson J, Ed), Bentham Science Publishers, 2015 Nov; pp.183-207. doi: 10.2174/97816810812811150101
  6. Inaba K, Kutomi O, Shiba K, Cosson J. Sperm guidance: comparison with motility regulation in bikont species. In: Flagellar Mechanics and Sperm Guidance (Cosson J, Ed), Bentham Science Publishers, 2015 Nov; pp.349-389. doi: 10.2174/97816810812811150101
  7. Miyata H, Satouh Y, Mashiko D, Muto M, Nozawa K, Shiba K, Fujihara Y, Isotani A, Inaba K, Ikawa M. Sperm calcineurin inhibition prevents mouse fertility with implications for male contraceptive. Science. 2015 Oct 23;350(6259):442-5. doi: 10.1126/science.aad0836.
  8. Inaba K, Mizuno K. Sperm Dysfunction and Ciliopathy. Reprod Med Biol, 2015 Oct 14; online. doi: 10.1007/s12522-015-0225-5
  9. Konno A, Shiba K, Cai C, Inaba K. Branchial cilia and sperm flagella recruit distinct axonemal components. PLoS One. 2015 May 11;10(5):e0126005. doi: 10.1371/journal.pone.0126005.
  10. Inaba K. Calcium sensors of ciliary outer arm dynein: functions and phylogenetic considerations for eukaryotic evolution. Cilia. 2015 Apr 30;4:6. doi: 10.1186/s13630-015-0015-z.
  11. Miyashiro D, Shiba K, Miyashita T, Baba SA, Yoshida M, Kamimura S. Chemotactic response with a constant delay-time mechanism in Ciona spermatozoa revealed by a high time resolution analysis of flagellar motility. Biol Open. 2015 Jan 8;4(2):109-18. doi: 10.1242/bio.20137351.
  12. Satou Y, Hirayama K, Mita K, Fujie M, Chiba S, Yoshida R, Endo T, Sasakura Y, Inaba K, Satoh N. Sustained heterozygosity across a self-incompatibility locus in an inbred ascidian. Mol Biol Evol. 2015 Jan;32(1):81-90. doi: 10.1093/molbev/msu268.


  1. Mizushima S, Hiyama G, Shiba K, Inaba K, Dohra H, Ono T, Shimada K, Sasanami T. The birth of quail chicks after intracytoplasmic sperm injection. Development. 2014 Oct;141(19):3799-806. doi: 10.1242/dev.111765.
  2. Yaguchi S, Yaguchi J, Inaba K. bicaudal-C is required for the formation of anterior neurogenic ectoderm in the sea urchin embryo. Sci Rep. 2014 Oct 31;4:6852. doi: 10.1038/srep06852.
  3. Alavi SM, Postlerová-Maňásková P, Hatef A, Pšenička M, Pěknicová J, Inaba K, Ciereszko A, Linhart O. Protease in sturgeon sperm and the effects of protease inhibitors on sperm motility and velocity. Fish Physiol Biochem. 2014 Oct;40(5):1393-8. doi: 10.1007/s10695-014-9933-8.
  4. Yokoe M, Sano M, Shibata H, Shibata D, Takayama-Watanabe E, Inaba K, Watanabe A. Sperm Proteases that May Be Involved in the Initiation of Sperm Motility in the Newt, Cynops pyrrhogaster. Int J Mol Sci. 2014 Aug 28;15(9):15210-24. doi: 10.3390/ijms150915210.
  5. Sung CG, Kim TW, Park YG, Kang SG, Inaba K, Shiba K, Choi TS, Moon SD, Litvin S, Lee KT, Lee JS. Species and gamete-specific fertilization success of two sea urchins under near future levels of pCO2. Journal of Marine Systems. 2014 Sep;137: 67-73. doi:10.1016/j.jmarsys.2014.04.013
  6. Shiba K, Inaba K. Distinct roles of soluble and transmembrane adenylyl cyclases in the regulation of flagellar motility in ciona sperm. Int J Mol Sci. 2014 Jul 28;15(8):13192-208. doi: 10.3390/ijms150813192.
  7. Shiba K, Shibata D, Inaba K. Autonomous changes in the swimming direction of sperm in the gastropod Strombus luhuanus. J Exp Biol. 2014 Mar 15;217(Pt 6):986-96. doi: 10.1242/jeb.095398.
  8. Alavi SM, Matsumura N, Shiba K, Itoh N, Takahashi KG, Inaba K, Osada M. Roles of extracellular ions and pH in 5-HT-induced sperm motility in marine bivalve. Reproduction. 2014 Feb 3;147(3):331-45. doi: 10.1530/REP-13-0418. Print 2014 Mar.
  9. Inaba K, Mizuno K, Shiba K. Structure, Function, and Phylogenetic Consideration of Calaxin. Sexual Reproduction in Animals and Plants (ed. H. Sawada, N. Inoue, M. Iwano). Springer. 2014 Jan, pp 49-57.


  1. Hirohashi N, Alvarez L, Shiba K, Fujiwara E, Iwata Y, Mohri T, Inaba K, Chiba K, Ochi H, Supuran CT, Kotzur N, Kakiuchi Y, Kaupp UB, Baba SA. Sperm from Sneaker Male Squids Exhibit Chemotactic Swarming to CO2. Curr Biol. 2013 May 6;23(9):775-81. doi: 10.1016/j.cub.2013.03.040.
  2. Jin Y, Yaguchi S, Shiba K, Yamada L, Yaguchi J, Shibata D, Sawada H, Inaba K. Glutathione transferase theta in apical ciliary tuft regulates mechanical reception and swimming behavior of Sea Urchin Embryos. Cytoskeleton 2013 Aug 1;70:453–470. doi: 10.1002/cm.21127.
  3. Takahashi T, Kutsuzawa M, Shiba K, Takayama-Watanabe E, Inaba K, Watanabe A. Distinct Ca2+ channels maintain a high motility state of the sperm that may be needed for penetration of egg jelly of the newt, Cynops pyrrhogaster. Dev Growth Differ. 2013 Sep;55(7):657-67. doi: 10.1111/dgd.12073. Epub 2013 Aug 25.
  4. Hiyama G, Matsuzaki M, Mizushima S, Dohra H, Ikegami K, Yoshimura T, Shiba K, Inaba K, Sasanami T. Sperm activation by heat shock protein 70 supports the migration of sperm released from sperm storage tubules in Japanese quail (Coturnix japonica). Reproduction. 2013 Dec 19;147(2):167-78. doi: 10.1530/REP-13-0439. Print 2014.


  1. Kikuchi A, Shiba K, Ozawa T, Nakano K, Inaba K, Numata O. Black Tea High-Molecular-Weight Polyphenol Increases the Motility of Sea Urchin Sperm by Activating Mitochondrial Respiration.Biosci Biotechnol Biochem. 2012 Dec 23;76(12):2321-4.
  2. Mizuno K, Shiba K, Okai M, Takahashi Y, Shitaka Y, Oiwa K, Tanokura M, Inaba K. Calaxin drives sperm chemotaxis by Ca2+-mediated direct modulation of a dynein motor. Proc Natl Acad Sci U S A. 2012 Dec 11;109(50):20497-502.
  3. Mohri H, Inaba K, Ishijima S, Baba SA. Tubulin-dynein system in flagellar and ciliary movement.Proc Jpn Acad Ser B Phys Biol Sci. 2012;88(8):397-415.
  4. Yuan Y, Tanabe T, Maekawa F, Inaba K, Maeda Y, Itoh N, Takahashi KG, Osada M. Isolation and functional characterization for oocyte maturation and sperm motility of the oocyte maturation arresting factor from the Japanese scallop, Patinopecten yessoensis.Gen Comp Endocrinol. 2012 Dec 1;179(3):350-7.
  5. Itou T, Kanno M, Suyama ., Kamiyama A, Sakamoto SX, Kijima A, Inaba K, Aoki MN. Development of 12 polymorphic microsatellite DNA markers for the kelp Ecklonia cava (Phaeophyceae, Laminariales). Conservation Genetics Resources 2012 Jun; 4(2):459-461.
  6. Saito T, Shiba K, Inaba K, Yamada L, Sawada H. Self-incompatibility response induced by calcium increase in sperm of the ascidian Ciona intestinalis. Proc Natl Acad Sci U S A. 2012 Mar 13;109(11):4158-62.
  7. Yaguchi J, Angerer LM, Inaba K, Yaguchi S. Zinc finger homeobox is required for the differentiation of serotonergic neurons in the sea urchin embryo.Dev Biol. 2012 Mar 1;363(1):74-83.
  8. Inaba K. Regulatory subunits of axonemal dynein. Handbook of Dynein (eds. Hirose, Amos). PanStanford Publishing, (2012), pp. 303-324.
  9. Shiba K, Mizuno K and Inaba K. Molecular comparison of the axonemal components between sperm flagella and Chlamydomonas flagella. Spermatology: new horizons in the 21 century (ed. M. Moriwsawa). Adthree Publishing Co. Ltd (2012), pp.30-40.


  1. Inaba K. Sperm flagella: comparative and phylogenetic perspectives of protein components. Mol Hum Reprod. 2011 Aug;17(8):524-38.
  2. Nakachi M, Nakajima A, Nomura M, Yonezawa K, Ueno K, Endo T, Inaba K. Proteomic profiling reveals compartment-specific, novel functions of ascidian sperm proteins. Mol Reprod Dev. 2011 Jul;78(7):529-49.
  3. Yaguchi S, Yaguchi J, Wei Z, Jin Y, Angerer LM, Inaba K. Fez function is required to maintain the size of the animal plate in the sea urchin embryo. Development. 2011 Oct;138(19):4233-43.
  4. Zhu L, Inaba K. Lipid rafts function in Ca(2+) signaling responsible for activation of sperm motility and chemotaxis in the ascidian Ciona intestinalis. Mol Reprod Dev. 2011 Aug 16.


  1. Konno A, Kaizu M, Hotta K, Horie T, Sasakura Y, Ikeo K, Inaba K. Distribution and structural diversity of cilia in tadpole larvae of the ascidian Ciona intestinalis. Dev Biol. 2010 Jan 1;337(1):42-62.
  2. Yamazaki Y, Akashi R, Banno Y, Endo T, Ezura H, Fukami-Kobayashi K, Inaba K, Isa T, Kamei K, Kasai F, Kobayashi M, Kurata N, Kusaba M, Matuzawa T, Mitani S, Nakamura T, Nakamura Y, Nakatsuji N, Naruse K, Niki H, Nitasaka E, Obata Y, Okamoto H, Okuma M, Sato K, Serikawa T, Shiroishi T, Sugawara H, Urushibara H, Yamamoto M, Yaoita Y, Yoshiki A, Kohara Y. NBRP databases: databases of biological resources in Japan. Nucleic Acids Res. 2010 Jan;38(Database issue):D26-32.
  3. Sasakura Y, Suzuki MM, Hozumi A, Inaba K, Satoh N. Maternal factor-mediated epigenetic gene silencing in the ascidian Ciona intestinalis. Mol Genet Genomics. 2010 Jan;283(1):99-110.
  4. Konno A, Padma P, Ushimaru Y, Inaba K. Multidimensional analysis of uncharacterized sperm proteins in Ciona intestinalis: EST-based analysis and functional immunoscreening of testis-expressed genes. Zoolog Sci. 2010 Feb;27(2):204-15.
  5. Yaguchi S, Yaguchi J, Wei Z, Shiba K, Angerer LM, Inaba K. ankAT-1 is a novel gene mediating the apical tuft formation in the sea urchin embryo. . Dev Biol. 2010 Dec 1;348(1):67-75.
  6. Endo T, Ueno K, Yonezawa K, Mineta K, Hotta K, Satou Y, Yamada L, Ogasawara M, Takahashi H, Nakajima A, Nakachi M, Nomura M, Yaguchi J, Sasakura Y, Yamasaki C, Sera M, Yoshizawa AC, Imanishi T, Taniguchi H, Inaba K. CIPRO 2.5: Ciona intestinalis protein database, a unique integrated repository of large-scale omics data, bioinformatic analyses and curated annotation, with user rating and reviewing functionality. Nucleic Acids Res. 2011 Jan;39(Database issue):D807-14.


  1. Sasakura Y, Inaba K, Satoh N, Kondo M, Akasaka K. Ciona intestinalis and Oxycomanthus japonicus, representatives of marine invertebrates. Exp Anim. 2009 Oct;58(5):459-69.
  2. Inaba K and Mizuno K. Purification of dyneins from sperm flagella. Axonemal dynein-mediated motility and IFT (King and Pazour, eds), Methods Cell Biol. 2009;92:49-63
  3. Satouh Y, Inaba K. Proteomic characterization of sperm radial spokes identifies a novel spoke protein with an ubiquitin domain. FEBS Lett. 2009 Jul 7;583(13):2201-7.
  4. Mizuno K, Padma P, Konno A, Satouh Y, Ogawa K, Inaba K.
    A novel neuronal calcium sensor family protein, calaxin, is a potential Ca2+-dependent regulator for outer arm dynein of metazoan cilia and flagella.
    Biol Cell. 2009 Feb;101(2):91-103.
  5. Nomura M, Nakajima A, Inaba K.
    Proteomic profiles of embryonic development in the ascidian Ciona intestinalis.
    Dev Biol. 2009 Jan 15;325(2):468-81.


  1. Satou Y, Mineta K, Ogasawara M, Sasakura Y, Shoguchi E, Ueno K, Yamada L, Matsumoto J, Wasserscheid J, Dewar K, Wiley GB, Macmil SL, Roe BA, Zeller RW, Hastings KE, Lemaire P, Lindquist E, Endo T, Hotta K, Inaba K.
    Improved genome assembly and evidence-based global gene model set for the chordate Ciona intestinalis: new insight into intron and operon populations.
    Genome Biol. 2008 Oct 14;9(10):R152. [Epub ahead of print]
  2. Kondoh E, Konno A, Inaba K, Oishi T, Murata M, Yoshida M.
    Valosin-containing protein/p97 interacts with sperm-activating and sperm-attracting factor (SAAF) in the ascidian egg and modulates sperm-attracting activity.
    Dev Growth Differ. 2008 Oct;50(8):665-73.
  3. Hozumi A, Padma P, Toda T, Ide H, Inaba K.
    Molecular characterization of axonemal proteins and signaling molecules responsible for chemoattractant-induced sperm activation in Ciona intestinalis.
    Cell Motil Cytoskeleton. 2008 Mar;65(3):249-67.
  4. Sasakura Y, Konno A, Mizuno K, Satoh N, Inaba K.
    Enhancer detection in the ascidian Ciona intestinalis with transposase-expressing lines of Minos.
    Dev Dyn. 2008 Jan;237(1):39-50.


  1. Hotta K, Mitsuhara K, Takahashi H, Inaba K, Oka K, Gojobori T, Ikeo K.
    A web-based interactive developmental table for the ascidian Ciona intestinalis, including 3D real-image embryo reconstructions: I. From fertilized egg to hatching larva.
    Dev Dyn. 2007 Jul;236(7):1790-805.
  2. Awazu S, Matsuoka T, Inaba K, Satoh N, Sasakura Y.
    High-throughput enhancer trap by remobilization of transposon Minos in Ciona intestinalis.
    Genesis. 2007 May;45(5):307-17.
  3. Inaba K, Nomura M, Nakajima A, Hozumi A.
    Functional proteomics in Ciona intestinalis: a breakthrough in the exploration of the molecular and cellular mechanism of ascidian development.
    Dev Dyn. 2007 Jul;236(7):1782-9.
  4. Inaba K.
    Molecular basis of sperm flagellar axonemes: structural and evolutionary aspects.
    Ann N Y Acad Sci. 2007 Apr;1101:506-26.


  1. Hozumi A, Satouh Y, Makino Y, Toda Y, Ide H, Ogawa K, King SM and Inaba K.
    Molecular Characterization of Ciona Sperm Outer Arm Dynein Reveals Multiple Components Related to Outer Arm Docking ComplexProtein 2.
    Cell Motil Cytoskeleton,2006,Oct;63(10):591-603.
  2. Ushimaru Y, Konno A, Kaizu M, Ogawa K, Satoh N and Inaba K.
    Association of a 66 kDa homolog of Chlamydomonas DC2,a subunit of outer arm docking complex,with outer arm dynein of sperm flagella in the ascidian Ciona intestinalis.
    Zool.Sci,2006 Aug;23(8):679-87.
  3. Tanabe T, Osada M, Kyozuka K, Inaba K. Kijima A.
    A nobel oocyte maturation arresting factor in the central nervous system of scallops inhibits serotonin-induced oocyte maturation and spawning of bivalve mollusks.
    Gen Comp Endocrinol.2006,147:352-361.
  4. Ogawa K and Inaba K.
    Ap58: a novel in situ outer dynein arm-binding protein.
    Biochem Biophys Res Commun.2005.343:385-390.


  1. Satouh Y, Padma P, Toda T, Satoh N, Ide H, Inaba K.
    Molecular Characterization of Radial Spoke Subcomplex Containing Radial Spoke Protein 3 and Heat Shock Protein 40 in Sperm Flagella of the Ascidian Ciona intestinalis.
    Mol Biol Cell. 2005, Feb; 16:626-36.
  2. Murata Y, Iwasaki H, Sasaki M, Inaba K, Okamura Y.
    Phosphoinositide phosphatase activity coupled to an intrinsic voltage sensor.
    Nature. 2005 Jun 30;435(7046):1239-1243.


  1. Hozumi A, Satouh Y, Ishibe D, Kaizu M, Konno A, Ushimaru Y, Toda T, Inaba K.
    Local database and the search program for proteomic analysis of sperm proteins in the ascidian Ciona intestinalis.
    Biochem Biophys Res Commun. 2004 Jul 9;319(4):1241-6.
  2. Kho KH, Inaba K.
    Expression of 17beta-hydroxysteroid dehydrogenease in testis of the ascidian Ciona intestinalis [corrected]
    Mol Cells. 2004 Oct 31;18(2):171-176. Erratum in: Mol Cells. 2005 Apr 30;19(2):301.


  1. P. Padma, Y. Satouh, K. Wakabayashi, A.Hozumi, Y. Ushimaru, R. Kamiya, K. Inaba.
    Identification of a novel leucine-rich repeat protein as a component of flagellar radial spoke in the ascidian Ciona intestinalis.
    Mol. Biol. Cell (2003), 14, 774-485.
  2. Inaba K, Dreanno C, Cosson J.
    Control of flatfish sperm motility by CO2 and carbonic anhydrase.
    Cell Motil Cytoskeleton. (2003) 55:174-87.
  3. Itoh A, Inaba K, Ohtake H, Fujinoki M, Morisawa M.
    Characterization of a cAMP-dependent protein kinase catalytic subunit from rainbow trout spermatozoa.
    Biochem Biophys Res Commun. (2003) 305:855-61.
  4. Krasznai, Z., Morisawa, M., Krasznai, ZT., Morisawa,S., Inaba, K., Bazsane, ZK., Rubovszky, B., Bodnar, B., Borsos, A. and Marian, T.
    Gadolinium, a Mechano-Sensitive Channel Blocker, Inhibits Osmosis-Initiated Motility of Sea- and Freshwater Fish Sperm, But Does Not Affect Human or Ascidian Sperm Motility.
    Cell Motility and the Cytoskeleton (2003) 55:232-243
  5. Inaba, K.
    Molecular architecture of sperm flagella: molecules for motility and signaling.
    Zool. Sci. (2003), 20, 1043-1056.
  6. Casey, DM., Inaba, K., Pazour, G.J., Takada, S., Wakabayashi, K., Wilkerson, CG., Kamiya, R., and Witman, G.B.
    DC3, the 21-kDa subunit of the outer dynein arm-docking complex (ODA-DC), is a novel EF-hand protein important for assembly of both the outer arm and the ODA-DC.
    Mol Biol Cell. 2003 Sep;14(9):3650-63. Epub 2003 Jun 27.
  7. Itoh, A., Fujinoki, M., Kawamura, T., Inaba, K., Ohtake, H., Shimizu, N. and Morisawa, M
    Purification and Characterization of the 15-kDa Protein from the Sperm Flagella of Salmonid Fishes,
    Biomed. Res. (2003), 24, 153-164
  8. Ogawa, K. and Inaba, K
    Smoac: Sperm motility activating complex formed by t-complex distorters.
    Biochem Biophys Res Commun. (2003),310, 1155-1159.


  1. Inaba K., Padma, P., Satouh, Y., Shin-i, T., Kohara, Y., Satoh, N. and Satou, Y.
    EST analysis of gene expression in testis of the ascidian, Ciona intestinalis.
    Mol. Reprod. Develop., (2002), 62, 431-445
  2. Satou, Y., Yamada, L., Mochizuki, Y., Takatori, N., Kawashima, T., Sasaki, A., Hamaguchi, M., Awazu, S., Yagi, K., Sasakura, Y., Nakayama, A., Inaba, K., Satoh, N.
    A cDNA resource from the basal chordate Ciona intestinalis.
    genesis (2002) 33, 153-154
  3. Inaba, K.
    Dephosphorylation of Tctex2-related dynein light chain by type 2A protein phosphatase.
    Biochem Biophys Res Commun (2002) 297, 800-805
  4. Yoshida M, Murata M, Inaba K, Morisawa M.
    A chemoattractant for ascidian spermatozoa is a sulfated steroid.
    Proc Natl Acad Sci USA (2002) 99, 14831-14836
  5. Dehal P, Satou Y, Campbell RK, Chapman J, Degnan B, De Tomaso A, Davidson B, Di Gregorio A, Gelpke M, Goodstein DM, Harafuji N, Hastings KE, Ho I, Hotta K, Huang W, Kawashima T, Lemaire P, Martinez D, Meinertzhagen IA, Necula S, Nonaka M, Putnam N, Rash S, Saiga H, Satake M, Terry A, Yamada L, Wang HG, Awazu S, Azumi K, Boore J, Branno M, Chin-Bow S, DeSantis R, Doyle S, Francino P, Keys DN, Haga S, Hayashi H, Hino K, Imai KS, Inaba K, Kano S, Kobayashi K, Kobayashi M, Lee BI, Makabe KW, Manohar C, Matassi G, Medina M, Mochizuki Y, Mount S, Morishita T, Miura S, Nakayama A, Nishizaka S, Nomoto H, Ohta F, Oishi K, Rigoutsos I, Sano M, Sasaki A, Sasakura Y, Shoguchi E, Shin-i T, Spagnuolo A, Stainier D, Suzuki MM, Tassy O, Takatori N, Tokuoka M, Yagi K, Yoshizaki F, Wada S, Zhang C, Hyatt PD, Larimer F, Detter C, Doggett N, Glavina T, Hawkins T, Richardson P, Lucas S, Kohara Y, Levine M, Satoh N, Rokhsar DS.
    The draft genome of Ciona intestinalis: insights into chordate and vertebrate origins.
    Science (2002) 298(5601):2157-67


  1. Kho, K.H., Tanimoto, S., Inaba K., Oka, Y. and Morisawa, M.
    Transmembrane cell signaling for the initiation of trout sperm motility: role of ion channel and membrane hyperpolarization for cyclic AMP synthesis.
    Zool. Sci. (2001), 18, 919-928
  2. Padma, P, Hozumi, A, Ogawa, K. and Inaba K..
    Molecular cloning and characterization of a thioredoxin/NDPK related dynein intermediate chain from the ascidian, Ciona intestinalis.
    Gene, (2001), 275, 177-183.
  3. Itoh, A., Inaba K., Fujinoki, M. and Morisawa, M.
    Motility-associated and Cyclic AMP-dependent Protein Phosphorylation in the Sperm of the Chum Salmon, Oncorhynchus keta.
    Biomed. Res. (2001), 22, 241-248.
  4. Inaba, K.
    High level of protein ubiquitination in ascidian sperm.
    In The biology of ascidians. (Sawada, Yokosawa, Lambert, eds,) (2001) pp.74-79.,Springer-Verlag.


  1. Nomura, M., Inaba K. and Morisawa, M.
    Cyclic AMP- and calmodulin-dependent phosphorylation of axonemal 21 kDa and 26 kDa proteins is a prerequisite for SAAF-induced motile activation in ascidian spermatozoa.
    Develop. Growth. Differ. (2000), 42, 129-138.
  2. Inaba K.
    Conformational Changes of Dynein: Mapping and Sequence Analysis of ATP/Vanadate-Dependent Trypsin-Sensitive Sites on the Outer Arm Dynein b Heavy Chain from Sea Urchin Sperm Flagella.
    J. Biochem. (2000), 127, 1115-1120
  3. Inaba, K.
    Molecular mechanism of the regulation of sperm motility - The role of protein phosphorylation in dynein-driven microtubule sliding.
    Proceedings of the JSPS-DOST symposium on Current Trends in Molecular Biology. (2000) pp.65-67
  4. Nomura M, Inaba K., and Morisawa M.
    Phosphorylation of axonemal 21 kDa and 26 kDa proteins modulates activation of sperm motility in the ascidian, Ciona intestinalis.
    Zygote. (2000) 8 Suppl 1, S59-60.


  1. Yoshida, K., Inaba K., Ohtake, H. and Morisawa, M.
    Purification and characterization of prolyl endopeptidase from the Pacific herring, Clupea pallasi and its role in the activation of sperm motility.
    Develop. Growth & Differ (1999) 41, 217-225.
  2. Inaba K., Kagami, O. and Ogawa, K.
    Tctex2-related outer arm dynein light chain is phosphorylated at activation of sperm motility.
    Biochem. Biophys. Res. Commun. (1999) 256, 177-183.
  3. Izumi, H., Marian, T., Inaba K., Oka, Y. and Morisawa, M.
    Membrane Hyperpolarization by sperm-activating and -attracting factor (SAAF) increases cAMP level and activates sperm motility in the ascidian Ciona intestinalis.
    Dev. Biol. (1999) 213, 246-256
  4. Mohri, H., Inaba K., Kubo-Irie, M., Takai, H. and Toyoshima, Y.Y.
    Characterization of outer arm dynein in sea anemone, Anthopleura midori.
    Cell Motil. Cytoskel., (1999), 44, 202-208.


  1. Inaba K., Morisawa, S. and Morisawa, M.
    Proteasomes regulate the motility of salmonid fish sperm through modulation of cAMP-dependnet phosphorylation of an outer arm dynein light chain.
    J. Cell Sci. (1998) 111, 1105-1115
  2. Ohkawa, K., Inaba, K., and Morisawa, M.
    Characterization of the 26S proteasome from sperm of chum salmon and sea urchin.
    J. Reprod. Develop. (1998) 44, 22.
  3. Nomura, M., Yoshida, M., Inaba, K. and Morisawa, M.
    Calmodulin (CaM)-mediated sperm activation in the ascidian, Ciona intestinalis.
    J. Reprod. Develop. (1998) 44, 52.


  1. Koutoulis, A., Pazour, G.J., Wilkerson, C.G., Inaba K., Sheng, H., Takada, S. and Witman, G.B.
    The Chlamydomonas reinhardtii ODA3 gene encodes a protein of the outer dynein arm docking complex (ODA-DC).
    J. Cell Biol. (1997) 137, 1069-1080.
  2. Inaba K., Buerano, C.C., Natividad, F.F. and Morisawa, M.
    Degradation of vitellogenins by 170 kDa trypsin-like protease in the plasma of the tilapia, Oreochromis niloticus.
    Comp. Biochem. Physiol. (1997) 118B, 85-90.
  3. Ohkawa, K., Inaba K. and Morisawa, M.
    Purification and characterization of 26 S proteasome from sperm flagella of chum salmon and its roles in the regulation of sperm motility.
    Biomed. Res. (1997) 18, 353-363
  4. Inaba, K., Ohkawa, K., Morisawa, S. and Morisawa, M.
    Proteasome-dependent regulatory pathway for the phosphorylation of a dynein light chain in chum salmon sperm. Proceedings of Twelfth International Symposium in Conjunction with Award of the International Prize for Biology, Biology of Reproduction,
    J. Reprod. Develop. (1997) 43s, 135-136


  1. Ogawa, K., Takai, H., Ogiwara, A., Yokota, E., Shimizu, T., Inaba K. and Mohri, H. : Is outer arm dynein intermediate chain 1 multifunctional? Mol. Biol. Cell (1996) 7, 1895-1907
  2. Inaba K., Ohkawa, K., and Morisawa, M. : Identification of tubulins associated with the 950 kDa protease from salmon sperm. Biomed. Res. (1996) 17, 87-93
  3. Mukai, T., Sato, T., Naruse, K., Inaba K., Shima, A. and Morisawa, M. : Genetic relationships of the genus Tridentiger (Pisces, Gobiidae) based on allozyme polymorphism . Zool. Sci. (1996) 13, 175-183.


  1. Buerano, C.C., Inaba K., Natividad, F.F. and Morisawa, M. : Vitellogenins of Oreochromis niloticus: Identification, isolation and biochemical and immunochemical characterizations. J. Exp. Zool. (1995) 273, 59-69.
  2. Inaba, K. and Morisawa, M. : Identification of a 10 S trypsin-like protease complex that cross-reacts with anti-proteasome antibody in sea urchin egg jelly. Arch. Biochem. Biophys. (1995) 319, 177-184.
  3. Inaba, K. : ATP-dependent conformational changes of dynein: Evidence for changes in the interaction of dynein heavy chain with the intermediate chain. J. Biochem. (1995) 117, 903-907.


  1. Yoshida, M., Inaba, K., Ishida, K. and Morisawa, M. : Calcium and cyclic AMP mediate sperm activation, but Ca2+ alone contributes sperm chemotaxis in the ascidian, Ciona savignyi. Develop. Growth & Differ.(1994) 36, 589-595
  2. Inaba, K. : Cross-linking of ß heavy chain subunit of dynein from sea urchin sperm flagella by dimethyl suberimidate. Biomed. Res. (1994) 15, 123-126.
  3. Jin, Z.X., Inaba, K., Manaka, K., Morisawa, M. and Hayashi, H. : Monoclonal antibodies against the protein complex that contains the flagellar movement initiating-phosphoprotein of Oncorhynchus keta. J. Biochem. (1994), 115, 885-890.


  1. Yoshida, M., Inaba, K. and Morisawa, M. : Sperm chemotaxis during the process of fertilization in the ascidian, Ciona savygni and Ciona intestinalis. Dev. Biol. (1993) 157, 497-506
  2. Inaba, K., Akazome, Y. and Morisawa, M. : Purification of proteasomes from salmonid fish sperm and their localization along sperm flagella. J. Cell Sci. (1993) 104, 907-915


  1. Inaba, K., Akazome, Y. and Morisawa, M.: Two high molecular mass proteases from sea urchin sperm. Biochem. Biophys. Res. Commun. (1992) 182, 667-674
  2. Inaba, K. and Morisawa, M. : Chymotrypsin-like protease activity associated with demembranated chum salmon sperm. Biol. Cell (1992) 76, 329-333


  1. Inaba, K., Ogawa, K. and Mohri, H. : Mapping of ATP-dependent trypsin-sensitive sites on the βchain of outer-arm dynein from sea urchin sperm flagella. J. Biochem. (1991) 110, 795-801
  2. Inaba, K. and Morisawa, M. : A chymotrypsin-like protease involved in motility of sperm in salmonid fish. Biomed. Res. (1991) 12, 435-437


  1. Inaba, K., Ono, M. and Mohri, H. : Conformational changes of the βchain of the outer-arm dynein from sea urchin sperm flagella coupled with ATP hydrolysis. J. Biochem. (1990) 108, 663-668


  1. Inaba, K. and Mohri, H. : Dynamic conformational changes of 21S dynein ATPase coupled with ATP hydrolysis revealed by proteolytic digestion. J. Biol. Chem. (1989) 264, 8384-8388
  2. Inaba, K. and Mohri, H. : Two states of the conformation of 21S outer arm dynein coupled with ATP hydrolysis. J. Biochem. (1989) 106, 249-354
  3. Inaba, K., Okuno, M., and Mohri, H. : Anthraniloyl ATP, a fluorescent analog of ATP, as a substrate for dynein ATPase and flagellar motility. Arch. Biochem. Biophys. (1989) 274, 209-215


  1. Inaba, K., Mohri, T. and Mohri, H. : B-band protein in sea urchin sperm flagella. Cell Motil. Cytoskel. (1988) 10, 506-517



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