a Bacteriorhodopsin Halorhodopsin Channelrhodopsin b D96 2 D85 D212 7 R82 Na',K',Ca2,H All-trans retinal chromophore C d 40m H134R N297Q E129S BR-EYFP BR-TS-EYFP T98S 10m 8E1629T2850 V156K《 2E140S V281K BR-TS-EYFP-ER、 (eBR)