{"id":3326,"date":"2018-08-16T06:00:49","date_gmt":"2018-08-15T21:00:49","guid":{"rendered":"https:\/\/www.muscle-hypertrophy.com\/?p=3326"},"modified":"2018-08-18T15:04:44","modified_gmt":"2018-08-18T06:04:44","slug":"%e3%82%bf%e3%82%a4%e3%83%81%e3%83%b3%e3%81%afwinding-filament%e3%81%aa%e3%81%ae%e3%81%8b%ef%bc%9f%e7%ad%8b%e5%8f%8e%e7%b8%ae%e3%81%ab%e3%81%8a%e3%81%91%e3%82%8b%e6%96%b0%e3%81%9f%e3%81%aa%e3%81%ad","status":"publish","type":"post","link":"https:\/\/www.muscle-hypertrophy.com\/?p=3326","title":{"rendered":"\u30bf\u30a4\u30c1\u30f3\u306f’winding filament’\u306a\u306e\u304b\uff1f\u7b4b\u53ce\u7e2e\u306b\u304a\u3051\u308b\u65b0\u305f\u306a\u306d\u3058\u308c"},"content":{"rendered":"

\u3000\u53ce\u7e2e\u306b\u95a2\u4e0e\u3059\u308b\u86cb\u767d\u8cea\u306f\u3053\u308c\u307e\u3067\u30a2\u30af\u30c1\u30f3\u3068\u30df\u30aa\u30b7\u30f3\u3068\u3055\u308c\u3066\u304d\u305f\uff0e\u3057\u304b\u3057\u6700\u8fd1\uff0c\u4f38\u5f35\u6027\u53ce\u7e2e\uff08\u30a8\u30ad\u30bb\u30f3\u30c8\u30ea\u30c3\u30af\uff0c\u30cd\u30ac\u30c6\u30a3\u30d6\uff09\u69d8\u5f0f\u306b\u304a\u3044\u3066\u30bf\u30a4\u30c1\u30f3\u306e\u679c\u305f\u3059\u5f79\u5272\u304c\u660e\u3089\u304b\u306b\u306a\u308a\u3064\u3064\u3042\u308b\uff0e\u4eca\u56de\u306f\u30bf\u30a4\u30c1\u30f3\u306b\u3064\u3044\u3066\u8ff0\u3079\u305f\u8ad6\u6587\u3092\u7d39\u4ecb\u3059\u308b\uff0e<\/p>\n

\u3000\u8a33\u306b\u63fa\u308c\u304c\u898b\u3089\u308c\u308b\u304c\u3054\u5bb9\u8d66\u3092\uff0e\u65b0\u898f\u306e\u5206\u91ce\u306e\u7ffb\u8a33\u306b\u306f\u6c17\u3092\u4f7f\u3046\uff0e’winding filament’ \u306f\u7121\u7406\u306b\u8a33\u3059\u3068\u300c\u5dfb\u304d\u3064\u304d\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u300d\u3068\u3067\u3082\u306a\u308b\u3060\u308d\u3046\u304c\uff0c\u304a\u305d\u3089\u304f\u82f1\u8a9e\u306e\u307e\u307e\u306b\u3057\u3066\u304a\u3044\u305f\u65b9\u304c\u5b9a\u7740\u3057\u3084\u3059\u3044\u3060\u308d\u3046\uff0e\u3053\u306e\u7528\u8a9e\u306b\u3064\u3044\u3066\u306f\u5c71\u672c\u7fa9\u5fb3\u6c0f\u304c\u8457\u66f8\u306b\u304a\u3044\u3066\u8a00\u53ca\u3057\u3066\u3044\u308b\uff0e<\/p>\n

<\/a>\"\"<\/p>\n

\u3000\u7b4b\u53ce\u7e2e\u306e\u969b\u306b\u30a2\u30af\u30c1\u30f3\u3068\u30df\u30aa\u30b7\u30f3\u304c ATP \u3092\u6d88\u8cbb\u3057\u3066\u52d5\u529b\u3092\u767a\u751f\u3055\u305b\u308b\u3053\u3068\u306f\u3088\u304f\u77e5\u3089\u308c\u3066\u304a\u308a\uff0c\u3053\u308c\u3092\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u6ed1\u8d70\u8aac<\/strong>\u3068\u3044\u3046\uff0e\u5b9f\u306f\u6570\u5341\u5e74\u524d\u304b\u3089\u3053\u308c\u4ee5\u5916\u306b\u3082\u5f3e\u6027\u7dda\u7dad\u306e\u5b58\u5728\u3092\u793a\u5506\u3059\u308b\u5b9f\u9a13\u7d50\u679c\u304c\u5f97\u3089\u308c\u3066\u304a\u308a, ‘elastic filament’ \u3068\u547c\u79f0\u3055\u308c\u3066\u3044\u305f\u304c\uff0c\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u6ed1\u8d70\u8aac\u3067\u591a\u304f\u306e\u4eba\u304c\u7d0d\u5f97\u3057\u3066\u3057\u307e\u3044\uff0c\u7814\u7a76\u306f\u7d30\u3005\u3068\u3057\u304b\u7d9a\u3044\u3066\u3044\u306a\u304b\u3063\u305f\uff0e<\/p>\n

\u3000\u4e00\u65b9\uff0c\u30c8\u30ec\u30fc\u30cb\u30fc\u306e\u9593\u3067\u306f\u30dd\u30b8\u30c6\u30a3\u30d6\u53ce\u7e2e\u3088\u308a\u3082\u30cd\u30ac\u30c6\u30a3\u30d6\u53ce\u7e2e\u306e\u65b9\u304c\u91cd\u3044\u91cd\u91cf\u3092\u6271\u3048\u308b\u3053\u3068\u304c\u7d4c\u9a13\u7684\u306b\u5206\u304b\u3063\u3066\u3044\u305f\u304c\uff0c\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u6ed1\u8d70\u8aac\u3067\u306f\u3053\u306e\u77db\u76fe\u3092\u8aac\u660e\u3067\u304d\u306a\u304b\u3063\u305f\uff0e<\/p>\n

\u3000\u81ea\u52d5\u8eca\u306b\u4f8b\u3048\u3066\u307f\u3088\u3046\uff0e\u30a8\u30f3\u30b8\u30f3\u306e\u56de\u8ee2\u306f\u30b7\u30e3\u30d5\u30c8\u3092\u901a\u3058\u3066\u30ae\u30e4\u30dc\u30c3\u30af\u30b9\u306b\u4f1d\u9054\u3055\u308c\u308b\uff0e\u30a2\u30af\u30c1\u30f3\u3068\u30df\u30aa\u30b7\u30f3\u306f\u30a8\u30f3\u30b8\u30f3\u3068\u30ae\u30e4\u30dc\u30c3\u30af\u30b9\u306e\u30bb\u30c3\u30c8\u3067\u3042\u308b\uff0e\u305d\u3057\u3066\uff0c\u30a8\u30f3\u30b8\u30f3\u3082\u30df\u30aa\u30b7\u30f3\u3082\uff0c\u4e00\u65b9\u5411\u306b\u3057\u304b\u52d5\u529b\u3092\u767a\u751f\u3067\u304d\u306a\u3044\uff0e\u3053\u308c\u306f\u30ae\u30e4\u30dc\u30c3\u30af\u30b9\u306b\u30ed\u30fc\u30ae\u30e4\u3057\u304b\u642d\u8f09\u3057\u3066\u304a\u3089\u305a\uff0c\u30ea\u30d0\u30fc\u30b9\u30ae\u30e4\u304c\u5b58\u5728\u3057\u306a\u3044\u306e\u3068\u540c\u3058\u3067\u3042\u308b\uff0e\u305d\u3093\u306a\u8eca\u306f\u30d0\u30c3\u30af\u304c\u3067\u304d\u306a\u3044\uff01\u30cd\u30ac\u30c6\u30a3\u30d6\u53ce\u7e2e\u3068\u306f\uff0c\u9ad8\u901f\u9053\u8def\u3092\u8d70\u3063\u3066\u3044\u308b\u6700\u4e2d\u306b\u30ae\u30e4\u3092\u30ea\u30d0\u30fc\u30b9\u306b\u5165\u308c\u308b\u3088\u3046\u306a\u3082\u306e\u3067\u3042\u308b\uff0e\u30aa\u30fc\u30c8\u30de\u8eca\u306f\u305d\u3082\u305d\u3082\u30ae\u30e4\u304c\u30ea\u30d0\u30fc\u30b9\u306b\u5165\u3089\u306a\u3044\u3088\u3046\u5b89\u5168\u88c5\u7f6e\u304c\u7d44\u307f\u8fbc\u307e\u308c\u3066\u3044\u308b\u304c\uff0c\u30de\u30cb\u30e5\u30a2\u30eb\u8eca\u306a\u3089\u3042\u3063\u3068\u3044\u3046\u9593\u306b\u30af\u30e9\u30c3\u30c1\u304c\u713c\u304d\u4ed8\u3044\u3066\u30a8\u30f3\u30b9\u30c8\u3057\u3066\u3057\u307e\u3046\uff0e<\/p>\n

\u3000\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u6ed1\u8d70\u8aac\u3067\u306f\u30df\u30aa\u30b7\u30f3\u982d\u90e8\u306e\u30d1\u30ef\u30fc\u30b9\u30c8\u30ed\u30fc\u30af\u306f\u30dd\u30b8\u30c6\u30a3\u30d6\u53ce\u7e2e\u306e\u969b\u306b\u3057\u304b\u52d5\u529b\u3092\u767a\u751f\u3067\u304d\u306a\u3044\u306f\u305a\u3067\u3042\u308b\u304c\uff0c\u5b9f\u969b\u306b\u306f\u30cd\u30ac\u30c6\u30a3\u30d6\u53ce\u7e2e\u306e\u6642\u306b\u3082\u52d5\u529b\u306f\u767a\u751f\u3057\u3066\u304a\u308a\uff0c\u3057\u304b\u3082\u30cd\u30ac\u30c6\u30a3\u30d6\u53ce\u7e2e\u306e\u65b9\u304c\u51fa\u529b\u304c\u5927\u304d\u3044\uff0e\u3067\u306f\uff0c\u305d\u306e\u52d5\u529b\u6e90\u306f\u4f55\u306a\u306e\u304b\uff1f\u305d\u3057\u3066\u3053\u306e\u5dee\u5206\u306e\u52d5\u529b\u306f\u3069\u3053\u304b\u3089\u6765\u3066\u3044\u308b\u306e\u304b\uff1f<\/p>\n

\u3000\u96fb\u5b50\u9855\u5fae\u93e1\u306b\u3088\u308b\u89b3\u5bdf\u3067\u306f\uff0c\u30df\u30aa\u30b7\u30f3\u306e\u672b\u7aef\u306f\u30d5\u30ea\u30fc\u3067\u306f\u306a\u304f, Z \u76e4\u306b\u7e4b\u304e\u6b62\u3081\u308b\u86cb\u767d\u8cea\u306e\u5b58\u5728\u304c\u6307\u6458\u3055\u308c\u3066\u304a\u308a\u5f3e\u6027\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3068\u3057\u3066\u6a5f\u80fd\u3057\u3066\u3044\u308b\u306b\u9055\u3044\u306a\u3044\u3068\u8a00\u308f\u308c\u3066\u304d\u305f\uff0e\u3053\u306e\u86cb\u767d\u8cea\u306f Y \u5b57\u578b\u3092\u3057\u3066\u3044\u308b\u305f\u3081\u514d\u75ab\u30b0\u30ed\u30d6\u30ea\u30f3 (Ig) \u3068\u8868\u73fe\u3055\u308c\u308b\u304c\uff0c\u3082\u3061\u308d\u3093\u5f62\u8cea\u7d30\u80de\u304b\u3089\u5206\u6ccc\u3055\u308c\u308b\u672c\u7269\u306e\u514d\u75ab\u30b0\u30ed\u30d6\u30ea\u30f3\u3067\u306f\u306a\u3044\uff0e\u305d\u3082\u305d\u3082\u5927\u304d\u3055\u304c\u5168\u304f\u9055\u3046\uff0e\u672c\u7269\u306e\u514d\u75ab\u30b0\u30ed\u30d6\u30ea\u30f3\u306e\u5206\u5b50\u91cf\u306f\u305f\u3063\u305f 25,000 \u3060\u304c\uff0c\u4eca\u8a71\u984c\u306b\u3057\u3066\u3044\u308b\u30bf\u30a4\u30c1\u30f3\u306f 370 \u4e07\u3082\u3042\u308b\uff0e<\/p>\n

\u3000\u96fb\u5b50\u9855\u5fae\u93e1\u306e\u89e3\u50cf\u5ea6\u306f\u7406\u8ad6\u4e0a\u306f 0.1 nm \u3068\u3055\u308c\u3066\u3044\u308b\u304c\uff0c\u771f\u7a7a\u4e2d\u3067\u64ae\u5f71\u3057\u306a\u3051\u308c\u3070\u306a\u3089\u306a\u3044\u3068\u3044\u3046\u5236\u7d04\u306e\u305f\u3081\uff0c\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u306b\u5916\u529b\u3092\u52a0\u3048\u308b\u3068\u3069\u306e\u3088\u3046\u306b\u3075\u308b\u307e\u3046\u306e\u304b\u306b\u3064\u3044\u3066\u306f\u3088\u304f\u5206\u304b\u3063\u3066\u3044\u306a\u304b\u3063\u305f\uff0e\u305d\u306e\u305f\u3081\uff0c\u69d8\u3005\u306a\u4eee\u8aac\u304c\u7acb\u3066\u3089\u308c\u305f\uff0e<\/p>\n

\u3000\u3053\u306e\u7591\u554f\u3092\u8aac\u660e\u3057\u3046\u308b\u6a5f\u5e8f\u3068\u3057\u3066\uff0c\u3053\u306e ‘winding filament’ \u4eee\u8aac\u304c\u6709\u529b\u8996\u3055\u308c\u3066\u304d\u305f\uff0e<\/p>\n

\u3000\u3056\u3063\u304f\u308a\u8aac\u660e\u3059\u308b\u3068\uff0c\u30dd\u30b8\u30c6\u30a3\u30d6\u53ce\u7e2e\u306e\u969b\u306b\u30df\u30aa\u30b7\u30f3\u304c\u767a\u751f\u3059\u308b\u52d5\u529b\u306e\u5927\u90e8\u5206\u306f\u53ce\u7e2e\u305d\u306e\u3082\u306e\u306b\u6d88\u8cbb\u3055\u308c\u308b\u304c\uff0c\u305d\u306e\u4e00\u90e8\u304c\u5f3e\u6027\u30dd\u30c6\u30f3\u30b7\u30e3\u30eb\u30a8\u30ca\u30b8\u30fc\u3068\u3057\u3066\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u5185\u306b\u84c4\u7a4d\u3055\u308c\uff0c\u30cd\u30ac\u30c6\u30a3\u30d6\u53ce\u7e2e\u306e\u969b\u306b\u653e\u51fa\u3055\u308c\u308b\u306e\u3067\u3042\u308b\uff0e\u305d\u306e\u84c4\u7a4d\u306e\u6a5f\u5e8f\u3092\u6700\u521d\u306b\u8aac\u660e\u3057\u305f\u306e\u304c ‘winding filament’ \u4eee\u8aac\u3067\u3042\u308b\uff0e\u6700\u521d\u306b\u300c\u5dfb\u304d\u3064\u304d\u300d\u3068\u66f8\u3044\u305f\u304c\uff0c\u30bf\u30a4\u30c1\u30f3\u304c\u30a2\u30af\u30c1\u30f3\u306b\u5dfb\u304d\u3064\u3044\u3066\u5f3e\u6027\u30dd\u30c6\u30f3\u30b7\u30e3\u30eb\u30a8\u30ca\u30b8\u30fc\u3092\u84c4\u7a4d\u3057\u3066\u3044\u308b\u306e\u3067\u306f\u306a\u3044\u304b\uff0c\u305d\u3057\u3066\u30cd\u30ac\u30c6\u30a3\u30d6\u53ce\u7e2e\u306e\u969b\u306b\u306f\u5dfb\u304d\u3064\u3044\u305f\u30bf\u30a4\u30c1\u30f3\u304c\u307b\u3069\u3051\u3066\u84c4\u7a4d\u3057\u305f\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u958b\u653e\u3059\u308b\u305f\u3081\u306b\u30dd\u30b8\u30c6\u30a3\u30d6\u53ce\u7e2e\u3088\u308a\u3082\u5f37\u3044\u51fa\u529b\u304c\u5f97\u3089\u308c\u308b\u306e\u3067\u306f\u306a\u3044\u304b\uff0c\u3068\u3044\u3046\u306e\u304c\u672c\u8ad6\u6587\u306e\u8da3\u65e8\u3067\u3042\u308b\uff0e<\/p>\n

\u3000\u3057\u304b\u3057\u5b9f\u306f\u3053\u306e ‘winding filament’ \u4eee\u8aac\u306f\uff0c\u6700\u65b0\u306e\u7814\u7a76\u3067\u5426\u5b9a\u3055\u308c\u3064\u3064\u3042\u308b\u3088\u3046\u306b\u898b\u3048\u308b\uff0e\u539f\u5b50\u9593\u529b\u9855\u5fae\u93e1\u304a\u3088\u3073\u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u306b\u3088\u308b\u7814\u7a76\u3067\u306f\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u306e\u6298\u308a\u305f\u305f\u307f\u304c\u95a2\u308f\u308b\uff0c\u3044\u308f\u3086\u308b titin folding \u4eee\u8aac<\/a> \u304c\u512a\u52e2\u3067\u3042\u308b\uff0e\u3053\u306e\u5206\u91ce\u306f\u30db\u30c3\u30c8\u306a\u9818\u57df\u3067\u3042\u308a\uff0c\u4eca\u5f8c\u306e\u7814\u7a76\u304c\u5f85\u305f\u308c\u308b\uff0e<\/p>\n

<\/p>\n

Is titin a \u2018winding filament\u2019? A new twist on muscle contraction<\/a> Proc Biol Sci. 2012 Mar 7; 279(1730): 981\u2013990.<\/h3>\n

\u8981\u7d04<\/h3>\n

Recent studies have demonstrated a role for the elastic protein titin in active muscle, but the mechanisms by which titin plays this role remain to be elucidated. In active muscle, Ca2+<\/sup>-binding has been shown to increase titin stiffness, but the observed increase is too small to explain the increased stiffness of parallel elastic elements upon muscle activation. We propose a \u2018winding filament\u2019 mechanism for titin’s role in active muscle. First, we hypothesize that Ca2+<\/sup>-dependent binding of titin’s N2A region to thin filaments increases titin stiffness by preventing low-force straightening of proximal immunoglobulin domains that occurs during passive stretch. This mechanism explains the difference in length dependence of force between skeletal myofibrils and cardiac myocytes. Second, we hypothesize that cross-bridges serve not only as motors that pull thin filaments towards the M-line, but also as rotors that wind titin on the thin filaments, storing elastic potential energy in PEVK during force development and active stretch. Energy stored during force development can be recovered during active shortening. The winding filament hypothesis accounts for force enhancement during stretch and force depression during shortening, and provides testable predictions that will encourage new directions for research on mechanisms of muscle contraction.<\/p>\n

\u30ad\u30fc\u30ef\u30fc\u30c9<\/h3>\n

connectin, force enhancement, force depression, history dependence of force production, thin filament rotation, titin\u2013actin interactions<\/p>\n

\u3000\u6700\u65b0\u306e\u7814\u7a76\u306f\uff0c\u6d3b\u52d5\u3057\u3066\u3044\u308b\u7b4b\u8089\u306b\u304a\u3051\u308b\u5f3e\u6027\u86cb\u767d\u30bf\u30a4\u30c1\u30f3\u306e\u5f79\u5272\u3092\u660e\u3089\u304b\u306b\u3057\u3066\u304d\u305f\uff0e\u3057\u304b\u3057\uff0c\u30bf\u30a4\u30c1\u30f3\u304c\u3053\u306e\u5f79\u5272\u3092\u3069\u306e\u3088\u3046\u306b\u6f14\u3058\u308b\u306e\u304b\u306b\u3064\u3044\u3066\u306f\u89e3\u660e\u3055\u308c\u3066\u3044\u306a\u3044\uff0e\u6d3b\u52d5\u7684\u306a\u7b4b\u8089\u306b\u304a\u3044\u3066\u306f Ca2+<\/sup> \u7d50\u5408\u304c\u30bf\u30a4\u30c1\u30f3\u306e\u786c\u5ea6\u3092\u5897\u52a0\u3055\u305b\u308b\u3053\u3068\u304c\u793a\u3055\u308c\u3066\u3044\u308b\u304c\uff0c\u89b3\u5bdf\u3055\u308c\u305f\u5897\u52a0\u306f\u5c0f\u3055\u3059\u304e\u3066\u7b4b\u8089\u6d3b\u52d5\u4e0a\u306e\u4e26\u884c\u5f3e\u6027\u8981\u7d20\u306e\u786c\u5ea6\u306e\u5897\u52a0\u3092\u8aac\u660e\u3059\u308b\u306b\u306f\u8db3\u308a\u306a\u3044\uff0e\u6211\u3005\u306f\u6d3b\u52d5\u3059\u308b\u7b4b\u8089\u306b\u304a\u3051\u308b\u30bf\u30a4\u30c1\u30f3\u306e\u5f79\u5272\u306b ‘winding filament’ \u6a5f\u5e8f\u3092\u63d0\u6848\u3059\u308b\uff0e\u307e\u305a\u6211\u3005\u306f\u4eee\u5b9a\u3059\u308b, Ca2+<\/sup> \u4f9d\u5b58\u6027\u306b\u30bf\u30a4\u30c1\u30f3\u306e N2A \u90e8\u4f4d\u304c\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306b\u7d50\u5408\u3059\u308b\u3053\u3068\u304c\uff0c\u53d7\u52d5\u7684\u30b9\u30c8\u30ec\u30c3\u30c1\u306e\u9593\u306b\u8d77\u304d\u308b\u514d\u75ab\u30b0\u30ed\u30d6\u30ea\u30f3\u8fd1\u4f4d\u30c9\u30e1\u30a4\u30f3\u306e\u5f31\u3044\u529b\u306e\u30b9\u30c8\u30ec\u30c3\u30c1\u3092\u963b\u5bb3\u3059\u308b\u3053\u3068\u306b\u3088\u308a\u30bf\u30a4\u30c1\u30f3\u306e\u786c\u5ea6\u3092\u5897\u52a0\u3055\u305b\u308b\uff0e\u3053\u306e\u6a5f\u5e8f\u306f\uff0c\u9aa8\u683c\u7b4b\u539f\u7dda\u7dad\u3068\u5fc3\u7b4b\u7d30\u80de\u306e\u9593\u306e\u529b\u306e\u9577\u3055\u4f9d\u5b58\u6027\u306e\u9055\u3044\u3092\u8aac\u660e\u3059\u308b\uff0e\u6b21\u306b\u6211\u3005\u306f\u4eee\u5b9a\u3059\u308b\uff0c\u67b6\u6a4b\u306f\u30e2\u30fc\u30bf\u30fc\u3068\u3057\u3066\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3092 M-line \u306b\u5f15\u304d\u5bc4\u305b\u308b\u3060\u3051\u3067\u306a\u304f\uff0c\u30bf\u30a4\u30c1\u30f3\u3092\u306d\u3058\u3063\u3066\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306b\u5dfb\u304d\u3064\u3051\u308b\u30ed\u30fc\u30bf\u30fc\u3068\u3057\u3066\u3082\u50cd\u304d\uff0c\u7b4b\u51fa\u529b\u5897\u5927\u3068\u30a2\u30af\u30c6\u30a3\u30d6\u77ed\u7e2e\u306e\u9593\u306b PEVK \u306b\u304a\u3044\u3066\u5f3e\u6027\u30dd\u30c6\u30f3\u30b7\u30e3\u30eb\u30a8\u30ca\u30b8\u30fc\u3092\u84c4\u7a4d\u3059\u308b\u306e\u3067\u3042\u308b\u3068\uff0e\u7b4b\u51fa\u529b\u5897\u5927\u306e\u9593\u306b\u84c4\u7a4d\u3055\u308c\u305f\u30a8\u30cd\u30eb\u30ae\u30fc\u306f\u30a2\u30af\u30c6\u30a3\u30d6\u77ed\u7e2e\u306e\u9593\u306b\u56de\u5fa9\u3057\u3046\u308b\uff0eWinding filament \u4eee\u8aac\u306f\u30b9\u30c8\u30ec\u30c3\u30c1\u4e2d\u306e\u529b\u306e\u5897\u5927\u3068\u77ed\u7e2e\u4e2d\u306e\u529b\u306e\u6291\u5236\u3092\u8aac\u660e\u3057\uff0c\u7b4b\u53ce\u7e2e\u306e\u6a5f\u5e8f\u306e\u7814\u7a76\u306e\u65b0\u3057\u3044\u65b9\u5411\u6027\u3092\u5968\u52b1\u3059\u308b\u691c\u8a3c\u53ef\u80fd\u306a\u6307\u6a19\u3092\u63d0\u4f9b\u3059\u308b\uff0e<\/p>\n

\u5c0e\u5165<\/h3>\n

\u3000\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u6ed1\u8d70\u8aac [1<\/a>,2<\/a>] \u306e\u516c\u958b\u4ee5\u964d\uff0c\u7b4b\u53ce\u7e2e\u306e\u5206\u5b50\u30e1\u30ab\u30cb\u30ba\u30e0 [3<\/a>,4<\/a>] \u306e\u89e3\u660e\u306b\u306f\u8457\u3057\u3044\u9032\u5c55\u304c\u3042\u3063\u305f\uff0e\u3053\u306e\u9032\u5c55\u306b\u3082\u304b\u304b\u308f\u3089\u305a\uff0c\u7b4b\u8089\u306e\u6a5f\u80fd\u306e\u3044\u304f\u3064\u304b\u306e\u554f\u984c\u306b\u95a2\u3059\u308b\u8aac\u660e\u304c\u307e\u3060\u89e3\u660e\u3055\u308c\u3066\u304a\u3089\u305a [5<\/a>], \u305d\u308c\u306b\u306f\u4f38\u5c55\u6642\u306b\u529b\u304c\u5897\u5f37\u3059\u308b\u3053\u3068 [6<\/a>], \u77ed\u7e2e\u6642\u306b\u7b4b\u529b\u304c\u4f4e\u4e0b\u3059\u308b\u3053\u3068 [6<\/a>],\u30a2\u30af\u30c6\u30a3\u30d6\u306a\u4f38\u5c55\u6642\u306e\u4f4e\u30b3\u30b9\u30c8\u306a\u7b4b\u529b\u7523\u751f [7<\/a>], \u9ad8\u3044\u71b1\u529b\u5b66\u7684\u52b9\u7387\u3067\u306e\u7b4b\u306e\u6d3b\u52d5\u7684\u77ed\u7e2e [8<\/a>] \u304c\u542b\u307e\u308c\u308b\uff0e\u3053\u308c\u3089\u304a\u3088\u3073\u305d\u308c\u3089\u306e\u89b3\u5bdf\u306e\u7d50\u679c\uff0c\u5143\u306e\u4eee\u8aac\u306e\u4fee\u6b63 [9<\/a>] \u304a\u3088\u3073\u5225\u306e\u4eee\u8aac\u306e\u767a\u5c55 [10<\/a>] \u306b\u3064\u306a\u304c\u3063\u305f\uff0e<\/p>\n

\u3000\u6211\u3005\u306f\u63fa\u52d5\u3059\u308b\u67b6\u6a4b\u306e\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u6ed1\u8d70\u8aac\u306b\u65b0\u3057\u3044\u4eee\u8aac\u3092\u4ed8\u3051\u52a0\u3048\u3066\u63d0\u6848\u3059\u308b\uff0e\u7b4b\u30b5\u30eb\u30b3\u30e1\u30a2\u306e\u4e8c\u6bb5\u968e\u306e ‘winding filament’ \u30e2\u30c7\u30eb\u306f\uff0c\u305d\u306e\u4e2d\u3067\u30bf\u30a4\u30c1\u30f3\u304c Ca2+<\/sup> \u6d41\u5165\u306b\u3088\u308a\u6d3b\u6027\u5316\u3059\u308b\u306e\u3060\u304c\uff0c\u305d\u306e\u7d50\u679c\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u304c\u67b6\u6a4b\u306b\u3088\u308a\u640d\u50b7\u3057\uff0c\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e\u7ffb\u8ee2\u3060\u3051\u3067\u306a\u304f\u56de\u8ee2\u3082\u8d77\u3053\u3059\uff0e\u3053\u306e\u4eee\u8aac\u304c\uff0c\u7b4b\u529b\u306e\u5897\u5f37\u304a\u3088\u3073\u6e1b\u8870\u3068\u540c\u69d8\u306b\uff0c\u7b4b\u751f\u7406\u5b66\u306b\u304a\u3051\u308b\u4ed6\u306e\u672a\u89e3\u660e\u306e\u89b3\u5bdf\u3092\u3082\u7406\u89e3\u3059\u308b\u4e0a\u3067\u7d0d\u5f97\u3067\u304d\u308b\u67a0\u7d44\u307f\u3092\u63d0\u4f9b\u3059\u308b\u3053\u3068\u3082\u793a\u305d\u3046\uff0e<\/p>\n

\u3000\u65e9\u671f\u306e\u69cb\u9020\u7814\u7a76 [2<\/a>] \u304b\u3089\uff0c\u30bf\u30a4\u30c1\u30f3\u69d8\u7dda\u7dad\u306e\u5b58\u5728\u304c\u63a8\u6e2c\u3055\u308c\u3066\u3044\u305f\u306b\u3082\u304b\u304b\u308f\u3089\u305a\uff0c\u30b3\u30cd\u30af\u30c1\u30f3\u3068\u3057\u3066\u3082\u77e5\u3089\u308c\u308b\u30bf\u30a4\u30c1\u30f3\u304c\u767a\u898b\u3055\u308c\u305f\u306e\u306f\uff0c\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u6ed1\u8d70\u8aac\u304c\u767a\u5c55\u3057\u3066\u304b\u3089 20 \u5e74\u4ee5\u4e0a\u3082\u5f8c\u3060\u3063\u305f [11<\/a>,12<\/a>]. \u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u6ed1\u8d70-\u67b6\u6a4b\u63fa\u52d5\u8aac\u306f\u30bf\u30a4\u30c1\u30f3\u3092\u8003\u616e\u305b\u305a\u306b\u9032\u5c55\u3057\u305f\uff0e\u30bf\u30a4\u30c1\u30f3\u306e\u65e9\u671f\u306e\u7814\u7a76\u306f\uff0c\u30b5\u30eb\u30b3\u30e1\u30a2\u96c6\u7a4d\u306e\u7dad\u6301 [13<\/a>] \u304a\u3088\u3073\u6a2a\u7d0b\u7b4b\u306e\u53d7\u52d5\u7684\u5f35\u529b\u3078\u306e\u5bc4\u4e0e [14<\/a>] \u306b\u304a\u3051\u308b\u305d\u306e\u5f79\u5272\u3092\u78ba\u7acb\u3057\u305f\uff0e\u73fe\u5728\u306e\u7814\u7a76\u306f\u7b4b\u539f\u7dda\u7dad\u306e\u96c6\u5408\u306e\u5236\u5fa1 [15<\/a>] \u304a\u3088\u3073\u7d30\u80de\u5185\u30b7\u30b0\u30ca\u30eb [16<\/a>] \u306b\u304a\u3051\u308b\u30bf\u30a4\u30c1\u30f3\u306e\u5f79\u5272\u306b\u7126\u70b9\u3092\u5f53\u3066\u3066\u3044\u308b\uff0e\u305d\u306e\u793a\u5506\u3059\u308b\u3068\u3053\u308d\u306b\u3088\u308b\u3068\uff0c\u30bf\u30a4\u30c1\u30f3\u306f\u4f11\u6b62\u671f\u3060\u3051\u3067\u306a\u304f\u6d3b\u52d5\u6642\u306e\u7b4b\u8089\u306b\u304a\u3044\u3066\u3082\u3070\u306d\u3068\u3057\u3066\u6a5f\u80fd\u3059\u308b\u53ef\u80fd\u6027\u304c\u3042\u308b [17<\/a>,18<\/a>,19<\/a>,20<\/a>]. \u3057\u304b\u3057\u306a\u304c\u3089\uff0c\u672a\u3060\u306b\uff0c\u30bf\u30a4\u30c1\u30f3\u304c\u3044\u304b\u306b\u3057\u3066\u305d\u306e\u3088\u3046\u306a\u5f79\u5272\u3092\u679c\u305f\u3059\u306e\u304b\u306b\u3064\u3044\u3066\u306e\u8aac\u5f97\u529b\u306e\u3042\u308b\u6a5f\u5e8f\u306f\u63d0\u4f9b\u3055\u308c\u3066\u3044\u306a\u3044\uff0e<\/p>\n

\u3000\u6211\u3005\u306f\u307e\u305a\uff0c\u7b4b\u30b5\u30eb\u30b3\u30e1\u30a2\u5185\u90e8\u306e\u30bf\u30a4\u30c1\u30f3\u306e\u914d\u7f6e\u306b\u3064\u3044\u3066\uff0c\u7b4b\u8089\u306e\u53d7\u52d5\u7684\u5f35\u529b\u3078\u306e\u30bf\u30a4\u30c1\u30f3\u306e\u8ca2\u732e\u306b\u3064\u3044\u3066\uff0c\u3055\u3089\u306b\u6d3b\u52d5\u6642\u306e\u7b4b\u8089\u306b\u304a\u3051\u308b\u529b\u306e\u7523\u751f\u3078\u306e\u30bf\u30a4\u30c1\u30f3\u306e\u8ca2\u732e\u306b\u3064\u3044\u3066\u793a\u3057\u305f\u6700\u65b0\u306e\u7814\u7a76\u3092\u30ec\u30d3\u30e5\u30fc\u3059\u308b\uff0e\u7d9a\u3044\u3066\uff0c\u6211\u3005\u306f\u4e8c\u6bb5\u968e\u306e winding filament \u4eee\u8aac\u3092\u767a\u5c55\u3055\u305b\u308b\uff0e\u305d\u3053\u3067\u306f Ca2+<\/sup> \u6d41\u5165\u304c\u30bf\u30a4\u30c1\u30f3\u3092\u300c\u6d3b\u6027\u5316\u300d\u3057\u3066 Ca2+<\/sup> \u4f9d\u5b58\u6027\u306b N2A \u304c\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306b\u7d50\u5408\u3059\u308b\u306e\u3060\u304c\uff0c\u30bf\u30a4\u30c1\u30f3\u306e\u3070\u306d\u3092\u77ed\u7e2e\u304a\u3088\u3073\u786c\u5316\u3055\u305b\uff0c\u67b6\u6a4b\u304c\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3092\u7ffb\u8ee2\u3055\u305b\u308b\u3060\u3051\u3067\u306a\u304f\u56de\u8ee2\u3055\u305b\u305f\u5f8c\uff0c\u30bf\u30a4\u30c1\u30f3\u306f\u305d\u308c\u3089\u306b\u5dfb\u304d\u3064\u304d\uff0c\u5f3e\u6027\u30dd\u30c6\u30f3\u30b7\u30e3\u30eb\u30a8\u30ca\u30b8\u30fc\u3092 PEVK \u5185\u306b\u84c4\u7a4d\u3059\u308b\uff0e\u6700\u5f8c\u306b\uff0c\u6211\u3005\u306f\u7b4b\u8089\u306e\u3075\u308b\u307e\u3044\u3092\u7406\u89e3\u3059\u308b\u305f\u3081\u306b winding filament \u4eee\u8aac\u306e\u610f\u7fa9\u306b\u3064\u3044\u3066\u8b70\u8ad6\u3057\u3088\u3046\uff0e<\/p>\n

\u7b4b\u30b5\u30eb\u30b3\u30e1\u30a2\u5185\u306e\u30bf\u30a4\u30c1\u30f3\u306e\u914d\u7f6e<\/h3>\n

\u3000420 \u4e07\u30c0\u30eb\u30c8\u30f3\u306b\u3082\u53ca\u3076\uff0c\u30bf\u30a4\u30c1\u30f3\u306f\u65e2\u77e5\u306e\u86cb\u767d\u8cea\u306e\u4e2d\u3067\u6700\u5927\u7d1a\u3067\u3042\u308a [21<\/a>], \u6a2a\u7d0b\u7b4b\u306e\u4e2d\u3067 3 \u756a\u76ee\u306b\u8c4a\u5bcc\u306a\u86cb\u767d\u8cea\u3067\u3042\u308b [22<\/a>]. \u30bf\u30a4\u30c1\u30f3\u306f Z \u76e4\u304b\u3089 M-line \u307e\u3067\uff0c\u30b5\u30eb\u30b3\u30e1\u30a2\u306e\u534a\u5206\u306e\u9577\u3055\u306b\u307e\u305f\u304c\u308b (1 µm) [15<\/a>]. \u5404\u30bf\u30a4\u30c1\u30f3\u5206\u5b50 (Figure 1a<\/strong>) \u306f 1 \u672c\u306e\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3068 A-band \u306b\u304a\u3044\u3066\u7d50\u5408\u3057, 1 \u672c\u306e\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3068 I-band \u306b\u304a\u3044\u3066\u7d50\u5408\u3059\u308b [23<\/a>]. \u5404\u30b5\u30eb\u30b3\u30e1\u30a2\u306e\u534a\u5206\u306b\u304a\u3044\u3066\uff0c\u5404\u3005\u306e\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306f 6 \u672c\u306e\u30bf\u30a4\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e\u30b3\u30a2\u3092\u53d6\u308a\u56f2\u307f\uff0c\u5404\u3005\u306e\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306f 3 \u672c\u306e\u30bf\u30a4\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3068\u76f8\u4e92\u4f5c\u7528\u3057\uff0c\u30bf\u30a4\u30c1\u30f3\u306f\u96a3\u63a5\u3059\u308b\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u304b\u3089\u653e\u5c04\u72b6\u306b\u4f38\u3073\u308b [24<\/a>]. \u975e\u4f38\u5f35\u6027\u306e\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u672b\u7aef\u306f\uff0c6 \u672c\u306e\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u306e\u514d\u75ab\u30b0\u30ed\u30d6\u30ea\u30f3 (Ig) \u9060\u4f4d\u30c9\u30e1\u30a4\u30f3\u304b\u3089\u69cb\u6210\u3055\u308c\u308b\u304c, A\/I \u5206\u5c90\u90e8\u306b\u304a\u3051\u308b\u5404\u3005\u306e\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e\u5148\u7aef\u304b\u3089\u51fa\u73fe\u3059\u308b [24<\/a>]. Ig \u30c9\u30e1\u30a4\u30f3\u9593\u306e\u76f8\u4e92\u4f5c\u7528\u306f\uff0c\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e\u516d\u89d2\u683c\u5b50\u306e\u4e2d\u5fc3\u306b\u304a\u3051\u308b\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u672b\u7aef\u306e\u8ef8\u65b9\u5411\u306e\u4f4d\u7f6e\u3092\u7dad\u6301\u3057\uff0c\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306f\u5404\u3005\u306e\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3092\u53d6\u308a\u56f2\u3080 [24<\/a>]. <\/p>\n

\"\"<\/a>
https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC2119535\/pdf\/jc1203711.pdf
\u30b5\u30eb\u30b3\u30e1\u30a2\u9577 2.5 µm \u306b\u304a\u3051\u308b\u5fc3\u7b4b\u7d30\u80de Z line \u8fd1\u508d\u306e\u96fb\u5b50\u9855\u5fae\u93e1\u306b\u3088\u308b\u7acb\u4f53\u5199\u771f\uff0e\u4e0a\u4e0b\u306b\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u304c\u3042\u308a\uff0c\u4e2d\u9593\u306b Z line \u304c\u3042\u308b\uff0e\u305d\u306e\u9593\u306b\u5f3e\u6027\u7dda\u7dad\u304c\u898b\u3048\u308b\uff0e<\/figcaption><\/figure>\n

\u3000\u30bf\u30a4\u30c1\u30f3\u306e\u30ab\u30eb\u30dc\u30ad\u30b7\u672b\u7aef\u306f M-line \u306b\u4f4d\u7f6e\u3057\uff0c\u305d\u3053\u3067\u306f\u30bf\u30a4\u30c1\u30f3\u306f\u9006\u306e\u6975\u6027\u306e\u91cd\u8907\u3067\u3082\u3063\u3066\u534a\u30b5\u30eb\u30b3\u30e1\u30a2\u3068\u96a3\u63a5\u3057\u3066\u3044\u308b [15<\/a>]. A-band \u306b\u304a\u3044\u3066\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u306f\u30d5\u30a3\u30d6\u30ed\u30cd\u30af\u30c1\u30f3 (FNIII) \u304a\u3088\u3073 Ig \u30c9\u30e1\u30a4\u30f3\u306e\u7e70\u308a\u8fd4\u3057\u304b\u3089\u69cb\u6210\u3055\u308c\uff0c\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3068\u5f37\u56fa\u306b\u7d50\u5408\u3057\u3066\u3044\u308b\u305f\u3081\u306b\u4f38\u9577\u4e0d\u80fd\u3067\u3042\u308b [25<\/a>]. I-band \u306b\u304a\u3044\u3066\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u306f Ig \u30bb\u30b0\u30e1\u30f3\u30c8\u306e\u8fd1\u4f4d\u8155\u304a\u3088\u3073\u9060\u4f4d\u8155\u3088\u308a\u69cb\u6210\u3055\u308c, PEVK \u90e8\u4f4d\u3068\u96a3\u63a5\u3057\u3066\u3044\u308b\uff0eIg \u8fd1\u4f4d\u8155\u304a\u3088\u3073 PEVK \u30bb\u30b0\u30e1\u30f3\u30c8\u306f\u76f4\u5217\u306e\u3070\u306d\u3068\u3057\u3066\u6a5f\u80fd\u3059\u308b [14<\/a>,26<\/a>]. Z \u76e4\u306b\u304a\u3044\u3066\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u306f\u30b5\u30eb\u30b3\u30e1\u30a2\u306b\u96a3\u63a5\u3059\u308b\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8, α \u30a2\u30af\u30c1\u30cb\u30f3\u304a\u3088\u3073\u53cd\u5e73\u884c\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u306b\u4fc2\u7559\u3055\u308c\u3066\u3044\u308b [27<\/a>]. Z \u76e4\u304a\u3088\u3073 M-line \u4e21\u8005\u306b\u304a\u3051\u308b\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u306e\u91cd\u8907\u306f\u30bf\u30a4\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u30b7\u30b9\u30c6\u30e0\u3092\u4f5c\u308a\u51fa\u3057\uff0c\u305d\u308c\u306f\u5168\u3066\u306e\u7b4b\u539f\u7dda\u7dad\u306e\u5168\u9577\u306b\u6e21\u308b\u3082\u306e\u3067\u3042\u308b [15<\/a>]. <\/p>\n

\"\"<\/a>
Figure 1. (a) \u9aa8\u683c\u7b4b\u30b5\u30eb\u30b3\u30e1\u30a2\u306e\u534a\u5206\u3067\u30bf\u30a4\u30c1\u30f3\u306e\u914d\u7f6e\u3092\u56f3\u793a\uff08\u9ec4\u8272\uff0e\u8d64\u3067 N2A \u30bb\u30b0\u30e1\u30f3\u30c8\u3092\u793a\u3059\uff09\uff0e\u5404\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u306f I-band \u306b\u304a\u3044\u3066\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\uff08\u9752\uff09\u306b\u7d50\u5408\u3057\u3066\u304a\u308a\uff0cA-band \u306b\u304a\u3044\u3066\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\uff08\u7dd1\uff09\u306b\u7d50\u5408\u3057\u3066\u3044\u308b\uff0e\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306f\u5358\u7d14\u5316\u3057\u3066\u4e8c\u91cd\u3089\u305b\u3093\u3068\u3057\u3066\u63cf\u3044\u3066\u3042\u308b\u304c\uff0c\u810a\u690e\u52d5\u7269\u306e\u9aa8\u683c\u7b4b\u306b\u304a\u3044\u3066\u306f\uff0c\u5b9f\u969b\u306b\u306f\u4e09\u91cd\u3089\u305b\u3093\u3067\u3042\u308b\uff0eN2A \u30bb\u30b0\u30e1\u30f3\u30c8\u306f\u8fd1\u4f4d\u8155\u306e Ig \u30bb\u30b0\u30e1\u30f3\u30c8\u3068 PEVK \u30bb\u30b0\u30e1\u30f3\u30c8\u306e\u9593\u306b\u5c40\u5728\u3059\u308b\uff0e(b,c) Ca2+<\/sup> \u4f9d\u5b58\u6027\u306e N2A \u306e\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3078\u306e\u7d50\u5408\uff0e(b) \u4f4e\u30ab\u30eb\u30b7\u30a6\u30e0\u6fc3\u5ea6\u3067\u306e\u305f\u308b\u307f\u306e\u9577\u3055\u3067\u306e\u5b89\u9759\u6642\u30b5\u30eb\u30b3\u30e1\u30a2 (pCa = 9). (c) Ca2+<\/sup> \u6d41\u5165\u3067\u306e (pCa = 4.5), \u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\uff08\u9752\uff09\u3078\u306e N2A \u30bf\u30a4\u30c1\u30f3\u7d50\u5408\uff0c\u6d3b\u52d5\u6027\u306e\u30b5\u30eb\u30b3\u30e1\u30a2\u306b\u304a\u3044\u3066\u77ed\u7e2e\u3057\u30bf\u30a4\u30c1\u30f3\u3070\u306d\u3092\u786c\u5316\u3059\u308b\uff0e<\/figcaption><\/figure>\n

\u30bf\u30a4\u30c1\u30f3\u306f\u7b4b\u8089\u306e\u53d7\u52d5\u7684\u5f35\u529b\u306b\u5bc4\u4e0e\u3059\u308b<\/h3>\n

\u3000\u591a\u304f\u306e\u7814\u7a76\u304c\u7121\u50b7\u306e\u7b4b\u539f\u7dda\u7dad\u306b\u304a\u3051\u308b\u53d7\u52d5\u7684\u7b4b\u529b\u3068\u4f38\u9577\u306e\u95a2\u9023\u540c\u69d8\uff0c\u500b\u3005\u306e\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u304a\u3088\u3073\u305d\u306e\u90e8\u5206\u306b\u3064\u3044\u3066\u8a18\u8ff0\u3057\u3066\u304d\u305f [14<\/a>,20<\/a>,28<\/a>]. \u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u304c\u77ed\u3044\u307b\u3069\uff0c\u6298\u308a\u7573\u307e\u308c\u305f I-band \u30bf\u30a4\u30c1\u30f3\u306e Ig \u30c9\u30e1\u30a4\u30f3\u3092\u771f\u3063\u76f4\u3050\u306b\u3059\u308b\u53d7\u52d5\u7684\u5f35\u529b\u306e\u5897\u52a0\u306f\u5c11\u306a\u3044 [28<\/a>,29<\/a>]. \u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u304c\u9577\u3044\u307b\u3069\uff0cPEVK \u30c9\u30e1\u30a4\u30f3\u304c\u5f15\u304d\u4f38\u3070\u3055\u308c\uff0c\u53d7\u52d5\u7684\u5f35\u529b\u306f\u6025\u6fc0\u306b\u5897\u52a0\u3059\u308b [26<\/a>,30<\/a>]. PEVK \u306f\u5c0f\u3055\u306a\u529b\u306b\u304a\u3044\u3066\u306f entropic spring \u3068\u3057\u3066\u3075\u308b\u307e\u3044\uff0c\u9006\u306b\u5f37\u3044\u529b\u306b\u304a\u3044\u3066\u306f enthalpic spring \u307e\u305f\u306f Hookean spring \u3068\u3057\u3066\u3075\u308b\u307e\u3046 [14<\/a>,31<\/a>]. \u30b5\u30eb\u30b3\u30e1\u30a2\u304c\u4f11\u6b62\u9577 2.1-3.5 µm \u304b\u3089\u4f38\u5c55\u3055\u308c\u305f\u969b\u306b\u306f, PEVK \u306e\u9577\u3055\u306f 10 \u500d\u4ee5\u4e0a\u306b\u5897\u52a0\u3059\u308b [14<\/a>]. \u751f\u7406\u7684\u306a\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u306b\u304a\u3044\u3066\u306f, PEVK \u306e\u4f38\u9577\u304c\u9aa8\u683c\u7b4b\u539f\u7dda\u7dad\u304a\u3088\u3073\u5fc3\u7b4b\u7d30\u80de\u306e\u53d7\u52d5\u7684\u5f3e\u6027\u3092\u5927\u304d\u304f\u898f\u5b9a\u3059\u308b [28<\/a>,32<\/a>]. \u53d7\u52d5\u7684\u5f35\u529b\u304c\u5897\u52a0\u3059\u308b\u5c40\u9762\u3067\u306f\uff0c\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u306f\u7570\u306a\u308b\u7b4b\u8089\u3067\u767a\u73fe\u3059\u308b\u30bf\u30a4\u30c1\u30f3\u306e\u30a2\u30a4\u30bd\u30d5\u30a9\u30fc\u30e0\u3092\u5909\u5316\u3055\u305b\u308b [28<\/a>]. <\/p>\n

\u6d3b\u52d5\u4e2d\u306e\u7b4b\u8089\u306b\u304a\u3044\u3066\u30bf\u30a4\u30c1\u30f3\u306e\u5f79\u5272\u306f\u3042\u308b\u306e\u304b\uff1f<\/h3>\n

\u3000\u6d3b\u52d5\u4e2d\u306e\u7b4b\u7dda\u7dad\u306b\u304a\u3044\u3066, Ca2+<\/sup> \u6d41\u5165\u306f\u975e\u67b6\u6a4b\u69cb\u9020\uff0c\u304a\u305d\u3089\u304f\u30bf\u30a4\u30c1\u30f3\u306e\u5f35\u529b\u3068\u525b\u6027\u3092\u5897\u52a0\u3055\u305b\u308b [19<\/a>]. PEVK \u65ad\u7247\u3068\u4e00\u500b\u306e\u7b4b\u7dda\u7dad\u306e\u4e21\u8005\u306b\u304a\u3044\u3066, Labeit \u3089\u306f Ca2+<\/sup> \u6d41\u5165\u304c\u30bf\u30a4\u30c1\u30f3\u306e\u525b\u6027\u3092\u5897\u52a0\u3055\u305b\u308b\u3060\u3051\u3067\u306a\u304f\uff0c\u7b4b\u539f\u7dda\u7dad\u5185\u306e Ca2+<\/sup> \u5897\u52a0\u304c force-extension curve \u306e\u5de6\u65b9\u504f\u4f4d\u3092\u8d77\u3053\u3059\u3053\u3068\u3092\u793a\u3057\u305f [20<\/a>]. \u3057\u304b\u3057\uff0c\u3053\u308c\u3089\u306e\u8a66\u9a13\u3067\u89b3\u5bdf\u3055\u308c\u305f\u30bf\u30a4\u30c1\u30f3\u525b\u6027\u3067\u306e Ca2+<\/sup> \u306e\u52b9\u679c\u306f 10 \u500d\u7a0b\u5ea6\u3067\u3042\u308a\uff0c\u30ab\u30eb\u30b7\u30a6\u30e0\u6d3b\u6027\u5316\u306b\u304a\u3044\u3066\u7b4b\u7dda\u7dad\u306e\u4e26\u884c\u5f3e\u6027\u525b\u6027\u3067\u89b3\u5bdf\u3055\u308c\u308b\u5897\u52a0\u3092\u8003\u616e\u3059\u308b\u3068\u5c0f\u3055\u3059\u304e\u308b [19<\/a>]. <\/p>\n

\u3000\u7b4b\u8089\u306b\u304a\u3051\u308b\u529b\u306e\u5897\u5f37\u306f\u30a2\u30af\u30c6\u30a3\u30d6\u30b9\u30c8\u30ec\u30c3\u30c1\u5f8c\u306e\u5b9a\u5e38\u72b6\u614b\u306e\u529b\u306e\u5897\u52a0\u3067\u3042\u308a\uff0c\u5bfe\u5fdc\u3059\u308b\u9577\u3055\u306e\u7b49\u5c3a\u6027\u306e\u529b\u3092\u8d85\u3048 [6<\/a>], \u305d\u308c\u306f\u53d7\u52d5\u7684\u8981\u7d20\u306e\u52d5\u54e1\u306b\u3088\u308b [33<\/a>]. \u3055\u3089\u306b\uff0c\u30a2\u30af\u30c6\u30a3\u30d6\u30b9\u30c8\u30ec\u30c3\u30c1\u3057\u305f\u7b4b\u7dda\u7dad\u306e\u975e\u30a2\u30af\u30c6\u30a3\u30d6\u5316\u5f8c\u306e\u53d7\u52d5\u7684\u7b4b\u529b\u306f\uff0c\u53d7\u52d5\u7684\u30b9\u30c8\u30ec\u30c3\u30c1\u5f8c\u3084\uff0c\u5bfe\u5fdc\u3059\u308b\u9577\u3055\u306e\u7b49\u5c3a\u6027\u53ce\u7e2e\u304b\u3089\u306e\u975e\u30a2\u30af\u30c6\u30a3\u30d6\u5316\u5f8c\u306b\u7523\u751f\u3055\u308c\u308b\u7b4b\u529b\u3088\u308a\u5f37\u304f [18<\/a>], \u3055\u3089\u306b\u306f ‘passive force enhancement’ \u304c\u53d7\u52d5\u7684\u8981\u7d20\uff0c\u3044\u308f\u3086\u308b\u30bf\u30a4\u30c1\u30f3\u306e\u52d5\u54e1\u306b\u3088\u308b\u3053\u3068\u304c\u793a\u5506\u3055\u308c\u305f [18<\/a>]. Joumaa \u3089\u306f\u7b4b\u539f\u7dda\u7dad\u306b\u304a\u3051\u308b\u53d7\u52d5\u7684\u7b4b\u529b\u5897\u5f37\u3092\u8a08\u6e2c\u3057\uff0c\u6d3b\u52d5\u7684\u7b4b\u529b\u7523\u751f\u306f\u30c8\u30ed\u30dd\u30cb\u30f3 C \u306e\u9664\u53bb\u306b\u3088\u308a\u963b\u5bb3\u3055\u308c\u305f [34<\/a>]. Labeit \u3089\u306e\u3088\u3046\u306b [20], \u5f7c\u3089\u3082\u307e\u305f Ca2+<\/sup> \u8a98\u767a\u6027\u306e\u30bf\u30a4\u30c1\u30f3\u306b\u3088\u308b\u525b\u6027\u306e\u5897\u52a0\u3092\u89b3\u5bdf\u3057\u305f\u304c\uff0c\u3057\u304b\u3057\u305d\u306e\u5897\u52a0\u306f\u53d7\u52d5\u7684\u7b4b\u529b\u5897\u5f37\u3092\u8003\u616e\u3059\u308b\u3068\u5c0f\u3055\u3059\u304e\u305f\uff0e\u5f7c\u3089\u306e\u63d0\u6848\u306b\u3088\u308b\u3068\uff0c\u53d7\u52d5\u7684\u7b4b\u529b\u5897\u52a0\u306f Ca2+<\/sup> \u6d41\u5165\u3060\u3051\u3067\u306a\u304f\u67b6\u6a4b\u5f62\u6210\u3084\u6d3b\u52d5\u7684\u7b4b\u529b\u7523\u751f\u3092\u8981\u3059\u308b\u3068\u3044\u3046\u3082\u306e\u3067\u3042\u308b [34<\/a>]. \u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3068\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3068\u306e\u91cd\u306a\u308a\u3092\u8d85\u3048\u3066\uff08\u30b5\u30eb\u30b3\u30e1\u30a2\u9577 > 3.8 µm\uff09\u4f38\u5c55\u3055\u308c\u305f\u7b4b\u539f\u7dda\u7dad\u306b\u304a\u3044\u3066\uff0cLeonard \u3068 Herzog \u3089\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u525b\u6027\u306e\u6d3b\u52d5\u4f9d\u5b58\u6027\u306e\u5897\u52a0\u304a\u3088\u3073\u5f35\u529b\u4f9d\u5b58\u6027\u306e\u5897\u52a0\u306e\u4e21\u8005\u306e\u30a8\u30d3\u30c7\u30f3\u30b9\u3092\u898b\u51fa\u3057\u305f [35<\/a>]. \u3053\u308c\u3089\u306e\u30c7\u30fc\u30bf\u306e\u793a\u3059\u3068\u3053\u308d\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u525b\u6027\u306f Ca2+<\/sup> \u6d41\u5165\u304a\u3088\u3073\u6d3b\u52d5\u3059\u308b\u7b4b\u8089\u306b\u304a\u3051\u308b\u529b\u306e\u5897\u52a0\u306b\u3088\u308b<\/strong>\uff0c\u3068\u3044\u3046\u3053\u3068\u3067\u3042\u308b\uff0eWinding filament \u4eee\u8aac\u306f\uff0c\u4ee5\u4e0b\u3067\u8ff0\u3079\u308b\u304c\uff0c\u6d3b\u52d5\u3059\u308b\u30b5\u30eb\u30b3\u30e1\u30a2\u3067\u306e\u30bf\u30a4\u30c1\u30f3\u306b\u3088\u308b\u5f35\u529b\u3078\u306e Ca2+<\/sup> \u6d41\u5165\u304a\u3088\u3073\u67b6\u6a4b\u30b5\u30a4\u30af\u30ea\u30f3\u30b0\u306e\u5f71\u97ff\u306e\u5206\u5b50\u30e1\u30ab\u30cb\u30ba\u30e0\u3092\u63d0\u4f9b\u3059\u308b\u3082\u306e\u3067\u3042\u308b\uff0e<\/p>\n

Ca2+<\/sup>\u4f9d\u5b58\u6027\u306e\u30bf\u30a4\u30c1\u30f3\u306e\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3078\u306e\u7d50\u5408<\/h3>\n

\u3000N2A \u90e8\u4f4d (Figure 1b, c) \u306f Ca2+<\/sup> \u4f9d\u5b58\u6027\u306b\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306b\u7d50\u5408\u3059\u308b\u9593\uff0c\u30bf\u30a4\u30c1\u30f3\u525b\u6027\u3092\u4fee\u98fe\u3059\u308b\u306b\u3042\u305f\u3063\u3066\u7406\u60f3\u7684\u306a\u4f4d\u7f6e\u306b\u3042\u308b\uff0e\u3053\u306e\u90e8\u4f4d\u3067\u30bf\u30a4\u30c1\u30f3\u304c\u30a2\u30af\u30c1\u30f3\u306b\u7d50\u5408\u3059\u308b\u3053\u3068\u306f I-band \u306b\u304a\u3051\u308b Ig \u30c9\u30e1\u30a4\u30f3\u8fd1\u4f4d\u8155\u3092\u307e\u3063\u3059\u3050\u306b\u3059\u308b\u5c0f\u3055\u306a\u529b\u3092\u5236\u9650\u3059\u308b\u53ef\u80fd\u6027\u304c\u3042\u308a, I-band \u306f\u5f1b\u7de9\u3057\u305f\u9577\u3055\u306e\u7b4b\u539f\u7dda\u7dad\u306e\u53d7\u52d5\u7684\u30b9\u30c8\u30ec\u30c3\u30c1\u306b\u3088\u308a\u81ea\u7136\u306b\u767a\u751f\u3059\u308b [14<\/a>]. \u3055\u3089\u306b, Ca2+<\/sup> \u306b\u3088\u308a\u6d3b\u6027\u5316\u3057\u305f\u30b5\u30eb\u30b3\u30e1\u30a2\u304c\u4f38\u5c55\u3055\u308c\u308b\u6642, PEVK \u30bb\u30b0\u30e1\u30f3\u30c8\u306f\u5f37\u3044\u529b\u3067\u5f15\u304d\u4f38\u3070\u3059\u3053\u3068\u306b\u306a\u308b\uff0e\u4eee\u306b Ca2+<\/sup> \u4f9d\u5b58\u6027\u306e N2A \u30bf\u30a4\u30c1\u30f3\u3068\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u9593\u306e\u7d50\u5408\u304c\u963b\u5bb3\u3055\u308c\u308c\u3070, active force production \u306f\u77ed\u3044\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u3067\u6e1b\u8870\u3059\u308b\u306f\u305a\u3067\u3042\u308b\uff0e\u30bf\u30a4\u30c1\u30f3\u5185\u3067\u767a\u751f\u3057\u305f\u3044\u304b\u306a\u308b\u7dca\u5f35\u3082\uff0c\u5f37\u3044\u529b\u3067 PEVK \u30bb\u30b0\u30e1\u30f3\u30c8\u3092\u4f38\u9577\u3059\u308b\u3088\u308a\u3082\u3080\u3057\u308d\u5c0f\u3055\u306a\u529b\u3067 Ig \u30bb\u30b0\u30e1\u30f3\u30c8\u6900\u3092\u771f\u3063\u76f4\u3050\u306b\u3059\u308b\u306f\u305a\u3067\u3042\u308b\uff0e\u3086\u3048\u306b\uff0c\u30bf\u30a4\u30c1\u30f3\u306e\u5168\u3066\u306e active force \u306b\u5bfe\u3059\u308b\u3044\u304b\u306a\u308b\u8ca2\u732e\u3082\u6e1b\u5c11\u3059\u308b\u3060\u308d\u3046\uff0e<\/p>\n

\u3000\u53ef\u80fd\u6027\u306e\u3042\u308b\u30a8\u30d3\u30c7\u30f3\u30b9\u306e\u793a\u5506\u3059\u308b\u3068\u3053\u308d\u3067\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u306f Ca2+<\/sup> \u5b58\u5728\u4e0b\u306b\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306b\u7d50\u5408\u3059\u308b\uff0e\u3057\u304b\u3057\uff0c\u5de8\u5927\u306a\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u5185\u90e8\u306e\u7d50\u5408\u90e8\u4f4d\u306e\u5c40\u5728\u306f\u5206\u304b\u3063\u3066\u3044\u306a\u3044\uff0e\u9aa8\u683c\u7b4b\u306b\u304a\u3044\u3066\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u5206\u89e3\u7523\u7269 (T2) \u306f\u30a2\u30af\u30c1\u30f3\u306b\u7d50\u5408\u3057, Ca2+<\/sup> \u4f9d\u5b58\u6027\u306e\u65b9\u6cd5\u3067\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3092\u9ad8\u3044\u89aa\u548c\u6027\u306b\u518d\u69cb\u7bc9\u3059\u308b [36<\/a>]. \u3053\u306e T2 \u65ad\u7247\u306f\u7d04 200 \u4e07\u30c0\u30eb\u30c8\u30f3\uff0c\u7d04 920 nm \u3067\u3042\u308a, C \u672b\u7aef, A-band FNIII, Ig \u30c9\u30e1\u30a4\u30f3, PEVK \u304a\u3088\u3073 N2A \u90e8\u4f4d\u306e\u4e00\u90e8\u3092\u542b\u3080 [37<\/a>]. \u3053\u306e N2A \u90e8\u4f4d\u306b\u306f\u30a8\u30d4\u30c8\u30fc\u30d7\u304c\u3042\u308a\uff0c\u305d\u308c\u306f p94\/calpain3 \u3068\u7d50\u5408\u3059\u308b [38<\/a>]. \u3053\u306e T2 \u65ad\u7247\u306f p94\/calpain3 \u304c\u30bf\u30a4\u30c1\u30f3\u3092 N2A \u90e8\u4f4d\u8fd1\u508d\u306e\u7d50\u5408\u90e8\u4f4d\u306b\u6d88\u5316\u3059\u308b\u969b\u306b\u5f62\u6210\u3055\u308c\u308b\u3068\u8003\u3048\u3089\u308c\u3066\u3044\u308b\uff0eCa2+<\/sup> \u6b20\u4e4f\u4e0b\u3067\u306f T2 \u3068\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u9593\u306b\u306f\u5f31\u3044\u76f8\u4e92\u4f5c\u7528\u3057\u304b\u8d77\u304d\u306a\u3044\u304c, Ca2+<\/sup> \u304c\u5897\u52a0\u3059\u308b (pCa = 4.5) \u3068, T2 \u306e\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3078\u306e\u89aa\u548c\u6027\u304c\u5897\u52a0\u3059\u308b [36<\/a>]. \u5de8\u5927\u306a T2 \u65ad\u7247\u5185\u90e8\u306e Ca2+<\/sup> \u4f9d\u5b58\u6027\u30a2\u30af\u30c1\u30f3\u7d50\u5408\u90e8\u4f4d\u306e\u5c40\u5728\u306f\u307e\u3060\u540c\u5b9a\u3055\u308c\u3066\u3044\u306a\u3044\uff0e<\/p>\n

\u3000\u6211\u3005\u306f\u3053\u306e Ca2+<\/sup> \u4f9d\u5b58\u6027\u30a2\u30af\u30c1\u30f3\u7d50\u5408\u90e8\u4f4d\u304c\u30bf\u30a4\u30c1\u30f3\u306e N2A \u5185\u306e T2 \u65ad\u7247\u306e N \u672b\u7aef\u306b\u5c40\u5728\u3059\u308b\u3068\u63d0\u6848\u3059\u308b (Figure 1b,c<\/strong>). \u7a81\u7136\u5909\u7570\u30de\u30a6\u30b9 (mdm) \u306e\u30d2\u30e9\u30e1\u7b4b\u306b\u304a\u3051\u308b\u6211\u3005\u306e\u6700\u8fd1\u306e\u7814\u7a76\u304c\u793a\u5506\u3059\u308b\u3068\u3053\u308d\u3067\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u306e\u30a8\u30d4\u30c8\u30fc\u30d7\u306f, N2A \u9818\u57df\u306e\u5185\u90e8\u307e\u305f\u306f\u8fd1\u508d\u306b\u3042\u308a, Ca2+<\/sup> \u5b58\u5728\u4e0b\u306b\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306b\u7d50\u5408\u3059\u308b\u3068\u3044\u3046\u3082\u306e\u3067\u3042\u308b [39]. \u30de\u30a6\u30b9\u306b\u304a\u3044\u3066\u306f mdm \u7a81\u7136\u5909\u7570\u306f\u30bf\u30a4\u30c1\u30f3\u907a\u4f1d\u5b50 N2A \u9818\u57df\u306e 779 \u5869\u57fa\u5bfe\u306e\u6b20\u5931\u3092\u7279\u5fb4\u3068\u3059\u308b [40<\/a>]. \u91ce\u751f\u578b\u30de\u30a6\u30b9\u306e\u7b4b\u8089\u306b\u304a\u3044\u3066\u6025\u901f\u306b\u514d\u8377\u3059\u308b\u9593, force-displacement \u66f2\u7dda\u306f\u4f11\u6b62\u72b6\u614b\u306e\u7b4b\u8089\u3068\u6bd4\u8f03\u3057\u3066\u77ed\u3044\u65b9\u5411\u306b\u6d3b\u52d5\u3092\u30b7\u30d5\u30c8\u3055\u305b\u308b\uff0e\u4f11\u6b62\u72b6\u614b\u306b\u6bd4\u3079\u3066\u6d3b\u52d5\u671f\u3067\u306f\u30d2\u30e9\u30e1\u7b4b\u3067\u306e\u30bf\u30a4\u30c1\u30f3\u3070\u306d\u306f\u7d04 10 % \u77ed\u7e2e\u3057, 2.5 \u500d\u786c\u304f\u306a\u308b\uff0e\u5bfe\u7167\u7684\u306b, mdm \u30de\u30a6\u30b9\u304b\u3089\u306e\u6d3b\u52d5\u3059\u308b\u7b4b\u8089\u3067\u306e\u30bf\u30a4\u30c1\u30f3\u3070\u306d\u306e\u9577\u3055\u3068\u525b\u6027\u306e\u5909\u5316\u306f\u89b3\u6e2c\u3055\u308c\u305a, N2A \u9818\u57df\u306b\u304a\u3051\u308b\u591a\u304f\u306e\u30a2\u30df\u30ce\u9178\u306e\u6b20\u640d\u304c\u8d77\u304d\u3066\u3044\u308b [39]. \u3053\u308c\u3089\u306e\u89b3\u5bdf\u304b\u3089, Ca2+<\/sup> \u6d41\u5165\u3067\u306e\u30bf\u30a4\u30c1\u30f3 N2A \u306e\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3078\u306e\u7d50\u5408\u304c\u30bf\u30a4\u30c1\u30f3\u3070\u306d\u3092\u77ed\u7e2e\u3057\u525b\u6027\u3092\u5897\u3059\u3068\u3044\u3046\u4eee\u8aac\u304c\u6210\u308a\u7acb\u3064\uff0e<\/p>\n

Ca2+<\/sup> \u4f9d\u5b58\u6027\u306e N2A\u2013ACTIN \u76f8\u4e92\u4f5c\u7528\u306f\u9577\u3055-\u5f35\u529b\u95a2\u4fc2\u306b\u5bc4\u4e0e\u3059\u308b<\/h3>\n

\u3000\u4eee\u306b N2A \u30bf\u30a4\u30c1\u30f3\u304c\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u5168\u9577\u306b\u308f\u305f\u3063\u3066\u8907\u6570\u306e\u7d50\u5408\u90e8\u4f4d\u3092\u6301\u3064\u306a\u3089\uff0c\u7b4b\u8089\u306e\u529b\u3068\u901f\u5ea6 [41<\/a>] \u306f\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u304c\u5897\u52a0\u3057\u305f\u6642\u3068\u76f8\u5bfe\u7684\u306b\u540c\u7b49\u3060\u3063\u305f\u3060\u308d\u3046 (Figure 2a,b<\/strong>). \u3053\u306e\u65b9\u6cd5\u3067\u306f, Ca2+<\/sup> \u4f9d\u5b58\u6027 N2A \u30bf\u30a4\u30c1\u30f3\u306e\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3078\u306e\u7d50\u5408\u306f\u30a2\u30af\u30c6\u30a3\u30d6\u306a\u9577\u3055-\u5f35\u529b\u95a2\u4fc2\u306b\u304a\u3051\u308b\u30d7\u30e9\u30c8\u30fc\u3092\u5f37\u5316\u3057\uff0c\u305d\u308c\u306f\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e\u4e2d\u9593\u306b\u3042\u308b\u81ea\u7531\u9818\u57df\u306e\u67b6\u6a4b\u306b\u5bfe\u5fdc\u3059\u308b [42<\/a>]. <\/p>\n

\u3000\u7b4b\u751f\u7406\u5b66\u306b\u304a\u3051\u308b\u672a\u89e3\u6c7a\u306e\u7591\u554f\u3068\u3057\u3066\uff0c\u306a\u305c\u5fc3\u7b4b\u306b\u306f\uff0c\u9aa8\u683c\u7b4b\u3068\u5bfe\u7167\u7684\u306b\uff0c\u30a2\u30af\u30c6\u30a3\u30d6\u306a\u9577\u3055-\u5f35\u529b\u95a2\u4fc2\u306b\u304a\u3051\u308b\u30d7\u30e9\u30c8\u30fc\u304c\u5b58\u5728\u3057\u306a\u3044\u306e\u304b\uff0c\u3068\u3044\u3046\u3082\u306e\u304c\u3042\u308b\uff0e\u305d\u306e\u30d7\u30e9\u30c8\u30fc\u306f\uff0c\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u9577\u306e\u69d8\u3005\u306a\u57fa\u672c\u304b\u3089 [43<\/a>] \u9aa8\u683c\u7b4b\u306b\u304a\u3051\u308b\u3088\u308a\u3082\u5fc3\u7b4b\u306b\u304a\u3044\u3066\u6700\u4f4e\u3067\u3082\u5e83\u304f\u307e\u305f\u306f\u3088\u308a\u5e45\u5e83\u304f\u5b58\u5728\u3059\u308b\u3082\u306e\u3068\u4e88\u60f3\u3055\u308c\u3066\u3044\u308b [42<\/a>]. \u9aa8\u683c\u7b4b\u3068\u306f\u5bfe\u7167\u7684\u306b\uff0c\u30bf\u30a4\u30c1\u30f3\u306e N2A \u30a2\u30a4\u30bd\u30d5\u30a9\u30fc\u30e0\u304c\u767a\u73fe\u3059\u308b\u3068\u3053\u308d\u3067\u306f\uff0c\u30de\u30a6\u30b9\u3068\u30e9\u30c3\u30c8\u306e\u5fc3\u5ba4\u3067\u306e\u5fc3\u7b4b\u7d30\u80de\u306f N2B \u30a2\u30a4\u30bd\u30d5\u30a9\u30fc\u30e0\u306e\u307f\u3092\u767a\u73fe\u3057\u3066\u304a\u308a [44<\/a>], Ca2+<\/sup> \u4f9d\u5b58\u6027\u306e\u30a2\u30af\u30c1\u30f3\u3078\u306e\u7d50\u5408\u304c\u5b58\u5728\u3057\u306a\u3044 [45<\/a>]. \u305d\u308c\u3086\u3048, mdm \u30de\u30a6\u30b9\u304b\u3089\u306e\u7b4b\u8089\u306e\u3088\u3046\u306b\uff0c\u5fc3\u7b4b\u7d30\u80de\u306f N2A \u30bf\u30a4\u30c1\u30f3\u306e\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3078\u306e\u7d50\u5408\u3092\u6b20\u3044\u3066\u3044\u308b\u3068\u4e88\u60f3\u3055\u308c\u3066\u3044\u308b\uff0e\u30e9\u30c3\u30c8\u3068\u30de\u30a6\u30b9\u306e\u5fc3\u7b4b\u7d30\u80de\u306f active force \u306b\u304a\u3051\u308b\u30d7\u30e9\u30c8\u30fc\u3092\u5448\u3055\u306a\u3044 [46<\/a>,47<\/a>]. \u5bfe\u7167\u7684\u306b\uff0c\u9c52\u304b\u3089\u306e\u5fc3\u7b4b\u7d30\u80de\u306f\u3088\u308a\u5de8\u5927\u306a N2AB \u30a2\u30a4\u30bd\u30d5\u30a9\u30fc\u30e0\u3092\u767a\u73fe\u3057\u3066\u304a\u308a, N2A \u3068 N2B \u306e\u4e21\u8005\u3092\u4f75\u305b\u6301\u3063\u3066\u3044\u308b\u306e\u3060\u304c\uff0c\u3086\u3048\u306b Ca2+<\/sup> \u4f9d\u5b58\u6027\u306e\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3078\u306e\u7d50\u5408\u3092\u5448\u3059\u308b\u3068\u4e88\u60f3\u3055\u308c\u308b\uff0eN2AB \u30a2\u30a4\u30bd\u30d5\u30a9\u30fc\u30e0\u3067\u63d0\u4f9b\u3055\u308c\u305f\u9c52\u5fc3\u7b4b\u3078\u306e\u8ffd\u5f93\u306e\u5897\u52a0\u306f\u62e1\u5f35\u671f\u306b\u304a\u3051\u308b\u3088\u308a\u5927\u304d\u306a\u4f38\u5c55\u3092\u8a31\u3059\u3053\u3068\u306b\u306a\u308b\uff0e\u3053\u306e\u4eee\u8aac\u306e\u4e88\u60f3\u3059\u308b\u3068\u3053\u308d\u306f\uff0c\u9c52\u5fc3\u7b4b\u7d30\u80de\u306f\uff0c\u9aa8\u683c\u7b4b\u306e\u3088\u3046\u306b\uff0c\u9577\u3055-\u5f35\u529b\u95a2\u4fc2\u306b\u304a\u3044\u3066\u30d7\u30e9\u30c8\u30fc\u3092\u5448\u3059\u308b\u3068\u3044\u3046\u3082\u306e\u3067\u3042\u308b [48<\/a>]. <\/p>\n

\u3000\u5fc3\u7b4b\u3068\u9aa8\u683c\u7b4b\u306e\u91cd\u8981\u306a\u9055\u3044\u306f\uff0c\u5fc3\u7b4b\u306f\u30a2\u30af\u30c6\u30a3\u30d6\u306a\u6642\u306b\u3082\u6c7a\u3057\u3066\u4f38\u5c55\u3055\u308c\u306a\u3044\u3053\u3068\u3067\u3042\u308b\uff0e\u305f\u3068\u3048\u62e1\u5f35\u671f\u306b\u53d7\u52d5\u7684\u306a\u9577\u3055\u3092\u7d4c\u3066\u3044\u305f\u3068\u3057\u3066\u3082\u3067\u3042\u308b [49<\/a>]. \u3086\u3048\u306b, Ca2+<\/sup> \u4f9d\u5b58\u6027\u306e\u30bf\u30a4\u30c1\u30f3\u306e\u525b\u6027\u306e\u5897\u52a0\u306f\u5fc3\u7b4b\u3067\u306f\u5fc5\u8981\u306a\u3044\uff0e\u5bfe\u7167\u7684\u306b\uff0c\u9aa8\u683c\u7b4b\u306f\u4f38\u5c55\u306e\u9593\u306f\u6052\u5e38\u7684\u306b\u30a2\u30af\u30c6\u30a3\u30d6\u3067\u3042\u308a [49<\/a>,50<\/a>,51<\/a>], N2A \u3068\u3044\u3046\u30bf\u30a4\u30c1\u30f3\u306e\u4e00\u3064\u306e\u30a2\u30a4\u30bd\u30d5\u30a9\u30fc\u30e0\u3092\u6301\u3061, Ca2+<\/sup> \u6d3b\u6027\u5316\u3067\u306e\u525b\u6027\u304c\u5897\u52a0\u3059\u308b\u3068\u4eee\u5b9a\u3055\u308c\u308b\uff0e<\/p>\n

\"\"<\/a>
Figure 2. Winding filament \u4eee\u8aac\u306e\u6982\u7565\u56f3\uff0e(a,b) Ca2+<\/sup> \u4f9d\u5b58\u6027\u306e N2A \u306e\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3078\u306e\u7d50\u5408\u306f\u9577\u3055\u3068\u5f35\u529b\u306e\u95a2\u4fc2\u306b\u5bc4\u4e0e\u3059\u308b\uff0e\u4eee\u306b N2A (red) \u304c\u975e\u9078\u629e\u7684\u306b\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\uff08\u9752\uff09\u306b\u5bfe\u3057\u3066 Ca2+<\/sup> \u5b58\u5728\u4e0b\u306b\u7d50\u5408\u3057\u305f\u3089\uff0c\u305d\u3057\u3066\u4eee\u306b\u305d\u306e\u7d50\u5408\u90e8\u4f4d\u304c Ca2+<\/sup> \u6d41\u5165\u6642\u306b\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u306b\u4f9d\u5b58\u6027\u3067\u3042\u308c\u3070\uff0c\u30a6\u30b5\u30ae\u306e\u5927\u8170\u7b4b\u306b\u304a\u3044\u3066 (a) 2.4 µm \u3068 (b) 2.6 µm \u306e\u9593\u3067\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u3067\u306e\u6d3b\u52d5\u6027\u306e\u529b\u306b\u306f\u30d7\u30e9\u30c8\u30fc\u304c\u793a\u5506\u3055\u308c\u308b\uff0e(c,d) \u67b6\u6a4b\u306e\u30b5\u30a4\u30af\u30eb\u306f\u30bf\u30a4\u30c1\u30f3\u306e\u306d\u3058\u308c\u308b\u7d50\u679c\u3068\u306a\u308b\uff0e(c) \u67b6\u6a4b\u306e\u30b5\u30a4\u30af\u30eb\u306f PEVK \u3092\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306b\u5dfb\u304d\u3064\u3051\u308b\uff08\u77e2\u5370\u306f\u56de\u8ee2\u65b9\u5411\u3092\u793a\u3059\uff09\uff0e\u3053\u306e\u30e2\u30c7\u30eb\u3067\u306f\uff0c\u5dfb\u304d\u3064\u3051\u89d2\u5ea6\u306f\u30b5\u30eb\u30b3\u30e1\u30a2\u306e\u5e7e\u4f55\u5b66\u7684\u5f62\u72b6\u306b\u306e\u307f\u4f9d\u5b58\u3059\u308b\uff0e(d) \u6d3b\u52d5\u4e2d\u306e\u30b5\u30eb\u30b3\u30e1\u30a2\u306e\u4f38\u9577\u306f PEVK \u30bb\u30b0\u30e1\u30f3\u30c8\u3092\u4f38\u5c55\u3057\uff0c\u30a2\u30af\u30c6\u30a3\u30d6\u306a\u529b\u3092\u5897\u5f37\u3059\u308b\uff0e<\/figcaption><\/figure>\n

\u67b6\u6a4b\u30b5\u30a4\u30af\u30eb\u306f\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3092\u56de\u8ee2\u3055\u305b\u308b\u304b\uff1f<\/h3>\n

\u3000\u6d3b\u52d5\u6642\u306e\u7b4b\u30b5\u30eb\u30b3\u30e1\u30a2\u306b\u304a\u3044\u3066\uff0c\u67b6\u6a4b\u306f\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3092\u7ffb\u8ee2\u3055\u305b\u308b\u306e\u3068\u540c\u69d8\u306b\u56de\u8ee2\u3055\u305b\u308b\u3060\u308d\u3046\uff0e\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3068\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e\u69cb\u9020\u304c\u4e0e\u3048\u3089\u308c\u308b\u3068, Morgan \u306f\uff0c\u30a2\u30af\u30c1\u30f3\u30e2\u30ce\u30de\u30fc\u3068\u96a3\u63a5\u3059\u308b 3 \u672c\u306e\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u9593\u306e\u7acb\u4f53\u7279\u7570\u7684\u306a\u7d50\u5408\u3092\u7dad\u6301\u3059\u308b\u306b\u306f\uff0c\u30df\u30aa\u30b7\u30f3\u982d\u90e8\u304c\u30a2\u30af\u30c1\u30f3\u30e2\u30ce\u30de\u30fc\u4e00\u3064\u5206\u306e\u9577\u3055 (5.5 nm) \u3092\u7ffb\u8ee2\u3059\u308b\u969b\u306b\uff0c\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u304c\u7d04 28° \u56de\u8ee2\u3059\u308b\u5fc5\u8981\u304c\u3042\u308b\u3053\u3068\u3092\u793a\u3057\u305f [10<\/a>]. \u3053\u308c\u306f 71.5 nm \u306e\u7ffb\u8ee2\u3054\u3068\u306b\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u304c 1 \u56de\u8ee2\u3059\u308b\u3053\u3068\u3092\u793a\u3057\u3066\u3044\u308b\uff0e<\/p>\n

\u3000\u30a2\u30af\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e\u30e1\u30ed\u30df\u30aa\u30b7\u30f3\u91cd\u9396\u306b\u3088\u308b\u56de\u8ee2\u306f in vivo \u3067\u89b3\u5bdf\u3055\u308c\u3066\u304d\u305f [52<\/a>,53<\/a>,54<\/a>]. Nishizaka \u3089\u306f\u30df\u30aa\u30b7\u30f3\u982d\u90e8\u304c\u30a2\u30af\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e\u9577\u8ef8\u306b\u5bfe\u3057\u3066\u53f3\u624b\u7cfb\u306e\u30c8\u30eb\u30af\u3092\u751f\u307f\u51fa\u3057\uff0c\u30a2\u30af\u30c1\u30f3\u4e8c\u91cd\u3089\u305b\u3093\u306b\u5bfe\u3057\u3066\u53f3\u56de\u8ee2\u306e\u3072\u306d\u308a\u3092\u52a0\u3048\u3066\u5dfb\u304d\u4e0a\u3052\u3066\u3044\u308b\u3053\u3068\u3092\u89b3\u5bdf\u3057\u305f [53<\/a>]. In vivo \u306b\u304a\u3044\u3066\u306f\uff0c\u30a2\u30af\u30c1\u30f3\u3068\u30df\u30aa\u30b7\u30f3\u9593\u306e\u76f8\u4e92\u4f5c\u7528\u306f\u7b4b\u30b5\u30eb\u30b3\u30e1\u30a2\u306b\u304a\u3051\u308b\u3088\u308a\u3082\u3082\u3063\u3068\u62e1\u6563\u3057\u3066\u304a\u308a\uff0c\u30a2\u30af\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306f 1 µm \u306e\u7ffb\u8ee2\u3054\u3068\u306b\u5b8c\u5168\u306b\u4e00\u56de\u8ee2\u3059\u308b [54<\/a>]. <\/p>\n