The University of Osaka - Department of Ophthalmology, The University of Osaka - Department of Ophthalmology, Kobe University - Division of Pathogenetic Signaling, The University of Osaka - Laboratory for Molecular and Developmental Biology, Kobe University - Division of Pathogenetic Signaling, Kobe University - Division of Pathogenetic Signaling, RIKEN Center for Integrative Medical Sciences - RIKEN-IFOM Joint Laboratory for Cancer Genomics, RIKEN Center for Integrative Medical Sciences - RIKEN-IFOM Joint Laboratory for Cancer Genomics, Kindai University - Department of Ophthalmology, The University of Osaka - Laboratory for Molecular and Developmental Biology, The University of Osaka - Department of Ophthalmology and Kobe University - Division of Pathogenetic Signaling