{"id":3488,"date":"2018-09-13T06:00:30","date_gmt":"2018-09-12T21:00:30","guid":{"rendered":"https:\/\/www.muscle-hypertrophy.com\/?p=3488"},"modified":"2019-04-25T08:25:25","modified_gmt":"2019-04-24T23:25:25","slug":"%e7%ad%8b%e5%8f%8e%e7%b8%ae%e3%81%ab%e3%81%8a%e3%81%91%e3%82%8b%e4%b8%bb%e8%a6%81%e3%81%aa%e9%a7%86%e5%8b%95%e5%8a%9b%e3%81%a8%e3%81%97%e3%81%a6%e3%81%ae%e3%82%bf%e3%82%a4%e3%83%81%e3%83%b3%e8%9b%8b","status":"publish","type":"post","link":"https:\/\/www.muscle-hypertrophy.com\/?p=3488","title":{"rendered":"\u7b4b\u53ce\u7e2e\u306b\u304a\u3051\u308b\u4e3b\u8981\u306a\u99c6\u52d5\u529b\u3068\u3057\u3066\u306e\u30bf\u30a4\u30c1\u30f3\u86cb\u767d\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u50cd\u304d"},"content":{"rendered":"<p>\u3000\u30bf\u30a4\u30c1\u30f3\u306e PEVK \u30c9\u30e1\u30a4\u30f3\u304c\u30a2\u30af\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306b\u5dfb\u304d\u3064\u3044\u3066\u5f3e\u6027\u529b\u3092\u767a\u63ee\u3059\u308b\u3068\u3044\u3046 &#8216;winding filament&#8217; \u4eee\u8aac\uff08<a href=\"https:\/\/www.muscle-hypertrophy.com\/?p=3326\" rel=\"noopener noreferrer\" target=\"_blank\">\u30bf\u30a4\u30c1\u30f3\u306f&#8217;winding filament&#8217;\u306a\u306e\u304b\uff1f\u7b4b\u53ce\u7e2e\u306b\u304a\u3051\u308b\u65b0\u305f\u306a\u306d\u3058\u308c<\/a>\uff09\u306b\u306f\u4e00\u5b9a\u306e\u8aac\u5f97\u529b\u304c\u3042\u3063\u305f\u3082\u306e\u306e\uff0c\u6700\u65b0\u306e\u89b3\u6e2c\u6280\u8853\u306f\u305d\u306e\u4eee\u8aac\u3092\u5426\u5b9a\u3057\u3064\u3064\u3042\u308b\uff0e<\/p>\n<p>\u3000\u6b66\u5668\u3068\u306a\u3063\u305f\u306e\u306f<a href=\"https:\/\/annex.jsap.or.jp\/photonics\/kogaku\/public\/42-02-sougouhoukoku.pdf\" rel=\"noopener noreferrer\" target=\"_blank\">\u539f\u5b50\u9593\u529b\u9855\u5fae\u93e1<\/a>\u3068\u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u3068\u3044\u3046\u30ca\u30ce\u30e1\u30fc\u30c8\u30eb\u30b5\u30a4\u30ba\u306e\u89b3\u6e2c\u6280\u8853\u304a\u3088\u3073\u64cd\u4f5c\u6280\u8853\u304c\u767a\u5c55\u3057\u305f\u3053\u3068\u3067\u3042\u308b\uff0e\u79d1\u5b66\u6280\u8853\u306e\u767a\u5c55\u304c\u6b21\u306e\u79d1\u5b66\u6280\u8853\u306e\u767a\u5c55\u3092\u4fc3\u3059\u3068\u3044\u3046\u6307\u6570\u95a2\u6570\u7684\u306a\u6280\u8853\u767a\u5c55\u306e\u6642\u4ee3\u306b\u6211\u3005\u306f\u3044\u308b. &#8220;Technology will always win.&#8221; \u3068\u3044\u3046\u540d\u8a00\u3092\u565b\u307f\u7de0\u3081\u3066\u3044\u308b\uff0e<\/p>\n<figure id=\"attachment_3555\" aria-describedby=\"caption-attachment-3555\" style=\"width: 374px\" class=\"wp-caption aligncenter\"><a href=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/AtomicForceMicroscopy.jpg\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/AtomicForceMicroscopy.jpg\" alt=\"\" width=\"374\" height=\"252\" class=\"size-full wp-image-3555\" srcset=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/AtomicForceMicroscopy.jpg 374w, https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/AtomicForceMicroscopy-300x202.jpg 300w\" sizes=\"auto, (max-width: 374px) 100vw, 374px\" \/><\/a><figcaption id=\"caption-attachment-3555\" class=\"wp-caption-text\">https:\/\/warwick.ac.uk\/fac\/sci\/physics\/current\/postgraduate\/regs\/mpagswarwick\/ex5\/techniques\/structural\/afm\/<br \/>\u539f\u5b50\u9593\u529b\u9855\u5fae\u93e1\u306e\u63a2\u91dd<\/figcaption><\/figure>\n<p>\u3000\u73fe\u5728\u3067\u306f\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u306e\u6298\u308a\u305f\u305f\u307f\uff08\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\uff09\u3068\u4f38\u5c55\uff08\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\uff09\u304c\u5f3e\u6027\u30a8\u30cd\u30eb\u30ae\u30fc\u306e\u6839\u6e90\u3067\u306f\u306a\u3044\u304b\u3068\u8003\u3048\u3089\u308c\u3066\u304a\u308a, <strong>titin folding \u4eee\u8aac<\/strong>\u3068\u3044\u3046\uff0e\u4eca\u56de\u306f\u305d\u306e\u6839\u62e0\u3068\u306a\u308b\u8ad6\u6587\u3092\u7d39\u4ecb\u3059\u308b\uff0e<\/p>\n<p>\u3000\u6b63\u76f4\u306a\u3068\u3053\u308d\uff0c\u4ffa\u306f\u5316\u5b66\u3084\u7269\u7406\u5b66\u306e\u5c02\u9580\u5bb6\u3067\u306f\u306a\u3044\uff0e\u82f1\u8a9e\u8ad6\u6587\u3092\u548c\u8a33\u3059\u308b\u306b\u3042\u305f\u308a\uff0c\u5c02\u9580\u7528\u8a9e\u306e\u30aa\u30f3\u30d1\u30ec\u30fc\u30c9\u306b\u840e\u7e2e\u3057\u304c\u3061\u3067\u3042\u308b\uff0e\u3053\u306e\u5206\u91ce\u306e\u82f1\u548c\u8f9e\u5178\u304c\u6b32\u3057\u3044\u3068\u5207\u5b9f\u306b\u9858\u3063\u3066\u3044\u308b\uff0e<\/p>\n<p><!--more--><\/p>\n<h3><a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC5957538\/\" rel=\"noopener noreferrer\" target=\"_blank\">The work of titin protein folding as a major driver in muscle contraction<\/a> Annu Rev Physiol. 2018 February 10; 80: 327\u2013351. doi:10.1146\/annurev-physiol-021317-121254.<\/h3>\n<h3>Abstract<\/h3>\n<p>Single molecule atomic force microscopy and magnetic tweezers experiments have demonstrated that titin Ig domains are capable of folding against a pulling force, generating mechanical work which exceeds that produced by a myosin motor. We hypothesize that upon muscle activation, formation of actomyosin crossbridges reduces the force on titin causing entropic recoil of the titin polymer and triggering the folding of the titin Ig domains. In the physiological force range of 4\u201315 pN under which titin operates in muscle, the folding contraction of a single Ig domain can generate 200% of the work of entropic recoil, and occurs at forces which exceed the maximum stalling force of single myosin motors. Thus titin operates like a mechanical battery storing elastic energy efficiently by unfolding Ig domains, and delivering the charge back by folding when the motors are activated during a contraction. We advance the hypothesis that titin folding and myosin activation act as inextricable partners during muscle contraction. <\/p>\n<h3>Keywords<\/h3>\n<p>Muscle contraction; titin; protein folding; polymer physics; single molecule; force spectroscopy<\/p>\n<h3>\u8981\u7d04<\/h3>\n<p>\u3000\u5358\u5206\u5b50\u539f\u5b50\u9593\u529b\u9855\u5fae\u93e1\u304a\u3088\u3073\u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u5b9f\u9a13\u304c\u660e\u3089\u304b\u306b\u3057\u305f\u306e\u306f\uff0c\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u304c\u727d\u5f15\u529b\u306b\u5bfe\u6297\u3059\u308b\u3053\u3068\u304c\u3067\u304d\u308b\u3068\u3044\u3046\u3053\u3068\u3067\u3042\u308a\uff0c\u305d\u308c\u306f\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306b\u3088\u308a\u7523\u751f\u3055\u308c\u308b\u4ed5\u4e8b\u3092\u8d85\u3048\u308b\u6a5f\u68b0\u7684\u4ed5\u4e8b\u3092\u767a\u751f\u3055\u305b\u308b\uff0e\u6211\u3005\u306f\u7b4b\u6d3b\u52d5\u3092\u4eee\u5b9a\u3057\u305f\uff0c\u30a2\u30af\u30c1\u30f3-\u30df\u30aa\u30b7\u30f3\u67b6\u6a4b\u5f62\u6210\u304c\u30bf\u30a4\u30c1\u30f3\u30dd\u30ea\u30de\u30fc\u306e\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc\u53cd\u52d5\u3092\u5f15\u304d\u8d77\u3053\u3057\uff0c\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u306e\u6298\u308a\u305f\u305f\u307f\u3092\u60f9\u8d77\u3059\u308b\u3053\u3068\u3067\u30bf\u30a4\u30c1\u30f3\u306b\u304b\u304b\u308b\u529b\u3092\u6e1b\u5c11\u3055\u305b\u308b\u3068\uff0e\u7b4b\u8089\u5185\u3067\u30bf\u30a4\u30c1\u30f3\u304c\u52d5\u4f5c\u3059\u308b\u72b6\u6cc1\u4e0b\u3067\u306e 4-15 pN \u3068\u3044\u3046\u751f\u7406\u5b66\u7684\u306a\u529b\u306e\u7bc4\u56f2\u3067\u306f\uff0c\u5358\u4e00\u306e Ig \u30c9\u30e1\u30a4\u30f3\u306e\u53ce\u7e2e\u306e\u6298\u308a\u305f\u305f\u307f\u306f\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc\u53cd\u52d5\u306e 200 % \u306e\u4ed5\u4e8b\u3092\u767a\u751f\u3055\u305b\u308b\u3053\u3068\u304c\u3067\u304d\uff0c\u5358\u4e00\u306e\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u3092\u5931\u901f\u3055\u305b\u308b\u6700\u5927\u306e\u529b\u3092\u8d85\u3048\u308b\u529b\u3092\u767a\u751f\u3055\u305b\u308b\u3053\u3068\u304c\u3067\u304d\u308b\uff0e\u3086\u3048\u306b\u30bf\u30a4\u30c1\u30f3\u306f\u6a5f\u68b0\u5f0f\u30d0\u30c3\u30c6\u30ea\u30fc\u306e\u3088\u3046\u306b\u52d5\u4f5c\u3057, Ig \u30c9\u30e1\u30a4\u30f3\u3092\u4f38\u5c55\u3059\u308b\u3053\u3068\u3067\u5f3e\u6027\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u52b9\u7387\u7684\u306b\u84c4\u7a4d\u3057\uff0c\u53ce\u7e2e\u671f\u306b\u30e2\u30fc\u30bf\u30fc\u304c\u6d3b\u6027\u5316\u3057\u305f\u969b\u306b\u6298\u308a\u305f\u305f\u307f\u306b\u3088\u308a\u96fb\u8377\u3092\u518d\u5145\u96fb\u3059\u308b\uff0e\u6211\u3005\u306f\u3053\u306e\u4eee\u8aac\u3092\u767a\u5c55\u3055\u305b\u305f\uff0e\u3064\u307e\u308a, titin folding \u304a\u3088\u3073\u30df\u30aa\u30b7\u30f3\u6d3b\u6027\u5316\u306f\u7b4b\u53ce\u7e2e\u306b\u304a\u3044\u3066\u4e0d\u53ef\u5206\u306e\u30d1\u30fc\u30c8\u30ca\u30fc\u3068\u3057\u3066\u3075\u308b\u307e\u3046\u306e\u3067\u3042\u308b\u3068\uff0e<\/p>\n<h3>Introduction<\/h3>\n<p>\u3000\u524d\u4e16\u7d00\u306e\u307b\u3068\u3093\u3069\u306e\u671f\u9593\uff0c\u4eba\u4f53\u306b\u304a\u3051\u308b\u6700\u5927\u306e\u86cb\u767d\u8cea\u3067\u3042\u308b\u30bf\u30a4\u30c1\u30f3\u306e\u5b58\u5728\u306f\u77e5\u3089\u308c\u3066\u3044\u306a\u304b\u3063\u305f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/914784\" rel=\"noopener noreferrer\" target=\"_blank\">1<\/a>). \u9a5a\u304f\u3053\u3068\u3067\u306f\u306a\u3044\u304c\uff0c\u7b4b\u53ce\u7e2e\u306e\u7406\u8ad6\u306f\u5f53\u6642\u77e5\u3089\u308c\u3066\u3044\u305f\u5206\u5b50\u3067\u3042\u308b\u30a2\u30af\u30c1\u30f3\u3068\u30df\u30aa\u30b7\u30f3\u306b\u57fa\u3065\u3044\u3066\u3044\u305f\uff0e\u3057\u304b\u3057\uff0c\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u6ed1\u8d70\u8aac\u306e\u767a\u898b\u306e\u524d\u591c\u306b\u306f\uff0c\u3044\u304f\u3064\u304b\u7af6\u5408\u3059\u308b\u4eee\u8aac\u304c\u5b58\u5728\u3057\u3066\u304a\u308a\uff0c\u4e0d\u601d\u8b70\u306a\u3053\u3068\u306b\uff0c\u7b4b\u53ce\u7e2e\u306b\u304a\u3051\u308b\u30bf\u30a4\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e\u3088\u3046\u306a\u5f79\u5272\u3092\u4e88\u8a00\u3057\u305f\u3082\u306e\u3082\u3042\u3063\u305f\uff0e1952 \u5e74\uff0c\u7269\u7406\u5316\u5b66\u8005\u306e Henry Erying \u3068\u89aa\u3057\u304b\u3063\u305f\u7814\u7a76\u8005\u306e Milton Polissar \u306f\u904b\u52d5\u901f\u5ea6\u7406\u8ad6\u3092\u7528\u3044\u3066 100 \u500b\u306e\u72ec\u7acb\u3057\u305f\u53ce\u7e2e\u5358\u4f4d\u304b\u3089\u306a\u308b\u5206\u7bc0\u72b6\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e\u5b58\u5728\u3092\u793a\u5506\u3057\u3066\u304a\u308a\uff0c\u305d\u306e\u53ce\u7e2e\u5358\u4f4d\u306f\u529b\u306e\u5b58\u5728\u4e0b\u306b\u6a5f\u68b0\u7684\u4ed5\u4e8b\u3092\u306a\u3057\u3066\u77ed\u7e2e\u3059\u308b\u3053\u3068\u304c\u3067\u304d\u308b\u3068\u3044\u3046\u3082\u306e\u3067\u3042\u3063\u305f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/14914890\" rel=\"noopener noreferrer\" target=\"_blank\">2<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/14914891\" rel=\"noopener noreferrer\" target=\"_blank\">3<\/a>,<a href=\"https:\/\/europepmc.org\/abstract\/med\/14914892\" rel=\"noopener noreferrer\" target=\"_blank\">4<\/a>,<a href=\"https:\/\/www.physiology.org\/doi\/abs\/10.1152\/ajplegacy.1952.168.3.805?journalCode=ajplegacy\" rel=\"noopener noreferrer\" target=\"_blank\">5<\/a>). \u5f7c\u306e\u30a2\u30a4\u30c7\u30a2\u306f\uff0c\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u6ed1\u8d70\u8aac\u304c\u5341\u5206\u306a\u5b9f\u9a13\u7d50\u679c (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC1357552\/\" rel=\"noopener noreferrer\" target=\"_blank\">6<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/5921536\" rel=\"noopener noreferrer\" target=\"_blank\">7<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/13165697\" rel=\"noopener noreferrer\" target=\"_blank\">8<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/4939977\" rel=\"noopener noreferrer\" target=\"_blank\">9<\/a>) \u3092\u4f34\u3063\u3066\u767a\u5c55\u3057\u3066\u304d\u305f\u6642\u306b\u306f\u9053\u306e\u7aef\u306b\u843d\u3061\u3066\u3057\u307e\u3063\u305f\uff0e\u9b45\u60d1\u7684\u306a\u904b\u547d\u306e\u306d\u3058\u308c\u306b\u3088\u308a\uff0c\u6211\u3005\u306f\u3053\u3093\u306b\u3061\uff0c\u4e21\u65b9\u306e\u4eee\u8aac\u304c\u6b63\u3057\u304f\uff0c\u4e21\u65b9\u306e\u6a5f\u5e8f\u304c\u7b4b\u53ce\u7e2e\u306b\u5bc4\u4e0e\u3059\u308b\u3053\u3068\u3092\u77e5\u3063\u3066\u3044\u308b\uff0e\u4eca\u3084\uff0c\u6b21\u306e\u70b9\u3092\u78ba\u7acb\u3059\u308b\u6c7a\u5b9a\u7684\u306a\u6839\u62e0\u304c\u5b58\u5728\u3059\u308b\uff0e\u3064\u307e\u308a\uff0c\u7b4b\u30b5\u30eb\u30b3\u30e1\u30a2\u306f\u4e09\u3064\u306e\u4e3b\u8981\u306a\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\uff0c\u3064\u307e\u308a\u30bf\u30a4\u30c1\u30f3\uff0c\u30a2\u30af\u30c1\u30f3\u304a\u3088\u3073\u30df\u30aa\u30b7\u30f3\u304b\u3089\u306a\u308a\uff0c\u5171\u306b\u5f3e\u6027\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u84c4\u7a4d\u3057\uff0c\u6a5f\u68b0\u7684\u4ed5\u4e8b\u3092\u5b9f\u884c\u3059\u308b\u305f\u3081\u306b\u50cd\u304f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/26854230\" rel=\"noopener noreferrer\" target=\"_blank\">10<\/a>). \u7b4b\u8089\u306b\u304a\u3051\u308b\u4e09\u756a\u76ee\u306e\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3068\u3057\u3066\u306e\u30bf\u30a4\u30c1\u30f3\u306e\u6839\u62e0\u306f\uff0c\u6700\u521d\u306f\u7591\u3044\u306e\u76ee\u3067\u63a1\u7528\u3055\u308c (<a href=\"http:\/\/rspb.royalsocietypublishing.org\/content\/160\/981\/442\" rel=\"noopener noreferrer\" target=\"_blank\">11<\/a>), \u30b5\u30eb\u30b3\u30e1\u30a2\u5185\u90e8\u306e\u30bf\u30a4\u30c1\u30f3\u306e\u5b58\u5728\u304c\u5e83\u304f\u53d7\u3051\u5165\u308c\u3089\u308c\u308b\u307e\u3067\u306b\u306f\u6570\u5341\u5e74\u3092\u8981\u3057\u305f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/934326\" rel=\"noopener noreferrer\" target=\"_blank\">12<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/7287636\" rel=\"noopener noreferrer\" target=\"_blank\">13<\/a>). \u30df\u30aa\u30b7\u30f3\u306e\u767a\u898b\u3068\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u6ed1\u8d70\u8aac (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/8663555\" rel=\"noopener noreferrer\" target=\"_blank\">14<\/a>) \u306e\u767a\u5c55\u306e\u9593\u3067\u4e00\u4e16\u7d00\u8fd1\u304f\u304c\u7d4c\u904e\u3057\uff0c\u6211\u3005\u306f\u9593\u9055\u3044\u306a\u304f\u30bf\u30a4\u30c1\u30f3\u306e\u751f\u7406\u5b66\u7684\u6a5f\u80fd\u306e\u7406\u89e3\u306e\u521d\u671f\u6bb5\u968e\u306b\u9054\u3057\u305f\u3070\u304b\u308a\u3067\u3042\u308b\uff0e<\/p>\n<p>\u3000\u30bf\u30a4\u30c1\u30f3\u306f\u4eba\u4f53\u306b\u304a\u3044\u3066\u6700\u5927\u7d1a\u306e\u86cb\u767d\u8cea\u3067\u3042\u308a, 370 \u4e07\u30c0\u30eb\u30c8\u30f3 (<a href=\"https:\/\/www.ahajournals.org\/doi\/abs\/10.1161\/hh2301.100981\" rel=\"noopener noreferrer\" target=\"_blank\">15<\/a>) \u3068\u65e2\u77e5\u306e\u30a2\u30a4\u30bd\u30d5\u30a9\u30fc\u30e0\u3067\u6700\u5927\u306e\u5206\u5b50\u91cf\u3092\u6301\u3064\u5358\u4e00\u306e\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u9396\u304b\u3089\u306a\u308b\uff0e\u5404\u30bf\u30a4\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306f\u534a\u30b5\u30eb\u30b3\u30e1\u30a2\u5168\u4f53\u306b\u308f\u305f\u308a\uff0c\u305d\u308c\u3089\u306f 4 \u3064\u306e\u7570\u306a\u308b\u9818\u57df\u306b\u5206\u5272\u3055\u308c\uff0c\u9855\u5fae\u93e1\u3067\u5bb9\u6613\u306b\u8a8d\u8b58\u3067\u304d\u308b (<strong>Figure 1A<\/strong>). \u7b4b\u8089\u306e I-band \u306f\u9577\u3055\u304c\u53ef\u5909\u3067\u4e3b\u306b\u30a2\u30af\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u304b\u3089\u306a\u308a\uff0c\u30bf\u30a4\u30c1\u30f3\u3068\u5e73\u884c\u306b\u8d70\u884c\u3057\u3066\u3044\u308b\uff0e\u30b5\u30eb\u30b3\u30e1\u30a2\u306e\u7dca\u5f35\u306b\u4f9d\u5b58\u3057\u3066\uff0c\u30a2\u30af\u30c1\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e\u3042\u308b\u90e8\u4f4d\u304c A-band \u5185\u90e8\u306b\u5411\u304b\u3063\u3066\u8d70\u884c\u3059\u308b\uff0e\u305d\u306e A-band \u306f\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u672b\u7aef\u306b\u3088\u308a\u533a\u753b\u3055\u308c\u3066\u304a\u308a\uff0c\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3068\u306f 294 \u500b\u306e\u30df\u30aa\u30b7\u30f3\u5206\u5b50\u306e\u675f\u306e\u3053\u3068\u3067\u3042\u308b\u304c\uff0c\u591a\u304f\u306e\u810a\u690e\u52d5\u7269\u306b\u304a\u3044\u3066 1.65 &micro;m \u306e\u9577\u3055\u3067\u898b\u51fa\u3055\u308c\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/11296141\" rel=\"noopener noreferrer\" target=\"_blank\">16<\/a>). \u7b4b\u8089\u306e\u6a2a\u65ad\u9762\u3092\u96fb\u5b50\u9855\u5fae\u93e1\u3067\u89b3\u5bdf\u3059\u308b\u3068\uff0c\u7dda\u7dad\u304c\u516d\u89d2\u683c\u5b50\u72b6\u306b\u914d\u5217\u3057\u3066\u304a\u308a, 6 \u672c\u306e\u30a2\u30af\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u304c\u5404\u30df\u30aa\u30b7\u30f3\u306e\u675f\u306e\u5468\u56f2\u3092\u53d6\u308a\u56f2\u3093\u3067\u3044\u308b\u306e\u304c\u89b3\u5bdf\u3067\u304d\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/9562033\" rel=\"noopener noreferrer\" target=\"_blank\">17<\/a>). \u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306f M-line \u5185\u306e\u86cb\u767d\u8cea\u3067\u4e92\u3044\u306b\u56fa\u5b9a\u3055\u308c\u3066\u304a\u308a\uff0c\u4e00\u65b9\u30a2\u30af\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306f Z \u76e4\u5185\u306e\u30a2\u30eb\u30d5\u30a1\u30a2\u30af\u30c1\u30cb\u30f3\u306b\u3088\u308a\u4fc2\u7559\u3055\u308c\u3066\u3044\u308b\uff0e<\/p>\n<figure id=\"attachment_3510\" aria-describedby=\"caption-attachment-3510\" style=\"width: 525px\" class=\"wp-caption aligncenter\"><a href=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFolding-Figure1.png\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFolding-Figure1-1024x877.png\" alt=\"\" width=\"525\" height=\"450\" class=\"size-large wp-image-3510\" srcset=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFolding-Figure1-1024x877.png 1024w, https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFolding-Figure1-300x257.png 300w, https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFolding-Figure1-768x658.png 768w, https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFolding-Figure1.png 1422w\" sizes=\"auto, (max-width: 525px) 100vw, 525px\" \/><\/a><figcaption id=\"caption-attachment-3510\" class=\"wp-caption-text\"><strong>Figure 1. \u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u539f\u5b50\u9593\u529b\u5206\u5149\u6cd5\u306f\u727d\u5f15\u529b\u306b\u6297\u3059\u308b\u30bf\u30a4\u30c1\u30f3\u518d\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3092\u793a\u3059. (A)<\/strong> \u7d04 2.6 \u03bcm \u306e\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u3067\u306e\u53b3\u683c\u306a\u9aa8\u683c\u7b4b\u7dda\u7dad\u3092\u63cf\u5199\u3059\u308b\u6a21\u5f0f\u56f3\uff0e\u3053\u306e\u4f38\u5c55\u306b\u304a\u3044\u3066\uff0c\u30bf\u30a4\u30c1\u30f3\u306f 4 pN \u306e\u4eee\u8aac\u7684\u306a\u529b\u306e\u4e0b\u306b\u3042\u308a\uff0c\u3059\u3079\u3066\u306e Ig \u30c9\u30e1\u30a4\u30f3\u306f\u6298\u308a\u7573\u307e\u308c\u3066\u3044\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/16230467\" rel=\"noopener noreferrer\" target=\"_blank\">87<\/a>). \u3042\u308b\u30b5\u30eb\u30b3\u30e1\u30a2\u306e\u534a\u5206\u306b\u30ba\u30fc\u30e0\u30a4\u30f3\u3057\u3066\u307f\u308b: \u30bf\u30a4\u30c1\u30f3\u306f\u5358\u4e00\u306e\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u3067\u3042\u308a, Z \u76e4\uff08N \u672b\u7aef\uff09\u304b\u3089 M \u30d0\u30f3\u30c9\uff08C \u672b\u7aef\uff09\u306b\u5e83\u304c\u308a\uff0c\u6298\u308a\u305f\u305f\u307e\u308c\u305f\u30c9\u30e1\u30a4\u30f3\uff08\u5857\u308a\u3064\u3076\u3057\u305f\u5186\uff09\u304a\u3088\u3073\u975e\u69cb\u9020\u9818\u57df\uff08\u7070\u8272\u306e\u7dda\uff09\u306e\u914d\u5217\u3068\u3057\u3066\u914d\u7f6e\u3055\u308c\u308b. \u30bf\u30a4\u30c1\u30f3\u306f 4 \u3064\u306e\u9818\u57df\u304b\u3089\u306a\u308b: N \u672b\u7aef\u30bb\u30b0\u30e1\u30f3\u30c8\u306f\uff08\u7dd1\u8272\u306e\u5186\uff09\uff0c\u514d\u75ab\u30b0\u30ed\u30d6\u30ea\u30f3 (Ig) \u69d8\u30c9\u30e1\u30a4\u30f3\u304b\u3089\u306a\u308a\uff0c\u30bf\u30a4\u30c1\u30f3\u3092 Z \u76e4\u306b\u4fc2\u7559\u3057\u3066\u3044\u308b. I-band \u30bb\u30b0\u30e1\u30f3\u30c8\u306f\u5f3e\u6027\u8981\u7d20\u3067\u3042\u308a\u4e8c\u7a2e\u985e\u306e\u30c9\u30e1\u30a4\u30f3\u3092\u6709\u3059\u308b\uff0e\u4e00\u5bfe\u306e Ig \u30c9\u30e1\u30a4\u30f3\uff08\u8d64\u3044\u5186\uff09\u3068\u672c\u8cea\u7684\u306b\u7121\u79e9\u5e8f\u306a\u69cb\u9020 (N2-B \u304a\u3088\u3073 PEVK) \u3067\u3042\u308b\uff0e A-band \u9818\u57df\u306f\uff08\u767d\u3044\u5186\uff09\u306f Ig \u304a\u3088\u3073 III \u578b\u30d5\u30a3\u30d6\u30ed\u30cd\u30af\u30c1\u30f3\u30c9\u30e1\u30a4\u30f3\u304b\u3089\u69cb\u6210\u3055\u308c\u3066\u304a\u308a\uff0c\u30bf\u30a4\u30c1\u30f3\u3092\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306b\u5805\u56fa\u306b\u7d50\u5408\u3059\u308b\u8cac\u4efb\u3092\u6709\u3059\u308b\uff08\u5099\u8003\uff0e\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306f I-band \u3092\u62e1\u5927\u3059\u308b\u305f\u3081\u5207\u308a\u8a70\u3081\u3089\u308c\u3066\u3044\u308b\uff09\uff0e\u6700\u5f8c\u306b, C \u672b\u7aef\u30bb\u30b0\u30e1\u30f3\u30c8\uff08\u9752\u3044\u5186\uff09\u306f 10 \u500b\u306e Ig \u30c9\u30e1\u30a4\u30f3\u3092\u6709\u3057\u3066\u304a\u308a\uff0c\u30bf\u30a4\u30c1\u30f3\u3092 M-band \u306e\u86cb\u767d\u8cea\u306b\u4fc2\u7559\u3057\u3066\u3044\u308b\uff0e\u4f1d\u7d71\u7684\u306b\uff0c\u30bf\u30a4\u30c1\u30f3\u306f\u53d7\u52d5\u7684\u5f35\u529b\u3092\u8abf\u7bc0\u3059\u308b\u306e\u307f\u3067\uff0c\u3059\u3079\u3066\u306e\u6a5f\u68b0\u7684\u4ed5\u4e8b\u306f\u30a2\u30af\u30c1\u30f3\u30df\u30aa\u30b7\u30f3\u306e\u67b6\u6a4b\u5f62\u6210\u306b\u3088\u308a\u306a\u3055\u308c\u308b\u3068\u8003\u3048\u3089\u308c\u3066\u304d\u305f\uff0e\u7570\u306a\u308b\u30bf\u30a4\u30c1\u30f3\u5207\u7247\u306e\u76f8\u5bfe\u7684\u306a\u9577\u3055\u306f\u4e00\u5b9a\u306e\u7e2e\u5c3a\u3067\u63cf\u304b\u308c\u3066\u3044\u306a\u3044\u3053\u3068\u306b\u7559\u610f\u3055\u308c\u305f\u3044. <strong>(B)<\/strong> \u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u5b9f\u9a13\u306e\u5178\u578b\u7684\u306a\u8a18\u9332\uff0e\u6700\u521d\u306b\u9ad8\u3044\u529b\u306e\u6ce2 (105 pN) \u306f 8 \u500b\u306e Ig \u30c9\u30e1\u30a4\u30f3\u3092\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3055\u305b\uff0c\u4e0a\u5411\u304d\u306e\u30b9\u30c6\u30c3\u30d7\uff08\u9ed2\u306e\u77e2\u5370\uff09\u3068\u3057\u3066\u8a18\u9332\u3055\u308c\uff0c\u305d\u306e\u9577\u3055\u306f 25 nm \u3067\u3042\u308b (<strong>Figure 2A<\/strong> \u53c2\u7167). \u529b\u3092\u6e1b\u5c11\u3055\u305b\u308b\u3068 (5 pN), \u305d\u306e\u5206\u5b50\u306f\u307e\u305a\u81ea\u767a\u7684\u306b\u5f3e\u6027\u53ce\u7e2e\u3057 (1), \u5f15\u304d\u7d9a\u3044\u3066\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53ce\u7e2e\u304c\u8d77\u304d\u308b (2). \u305d\u3053\u3067\u306f\u500b\u3005\u306e\u30c9\u30e1\u30a4\u30f3\u306f\u96e2\u6563\u7684\u306a\u4e0b\u65b9\u30b9\u30c6\u30c3\u30d7\u3068\u3057\u3066\u518d\u6298\u308a\u305f\u305f\u307f\u3055\u308c\uff08\u8d64\u77e2\u5370\uff09\uff0c\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u306f\u77ed\u7e2e\u3059\u308b\uff0e\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u30d1\u30eb\u30b9\u306f\u975e\u751f\u7406\u7684\u306a\u9ad8\u3044\u529b\u3067\u5b9f\u65bd\u3055\u308c, 8 \u500b\u306e\u30c9\u30e1\u30a4\u30f3\u306f\u6570\u79d2\u9593\u3067\u4f38\u5c55\u3055\u308c\u305f\uff0e\u7070\u8272\u306e\u7834\u7dda\u306e\u8ecc\u8de1\u306f 10 pN \u306e\u751f\u7406\u7684\u306a\u529b\u3067\u306e\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u8ecc\u9053\u3092\u793a\u3057\uff0c\u5b8c\u5168\u306a\u4f38\u5c55\u306b 6 \u6642\u9593\u3092\u8981\u3057\u305f\uff0e\u305d\u306e\u529b\u304c 5 pN \u306b\u6e1b\u5c11\u3059\u308b\u3068\uff0c\u5f3e\u6027\u53ce\u7e2e\u306e\u5f37\u3055\u306f\u66f4\u306b\u5c0f\u3055\u304f\u306a\u3063\u305f\u304c\uff0c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53ce\u7e2e\u306f\u540c\u3058\u307e\u307e\u3060\u3063\u305f\uff0e (<strong>Inset<\/strong>) \u5358\u5206\u5b50\u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u5b9f\u9a13\u3092\u793a\u3059\u6a21\u5f0f\u56f3\uff0e\u30bf\u30a4\u30c1\u30f3\u30dd\u30ea\u86cb\u767d\u8cea\u306f HaloTag \u5171\u6709\u7d50\u5408\u306e\u5316\u5b66\u7684\u6027\u8cea\u306b\u3088\u308a\u30ac\u30e9\u30b9\u8868\u9762\u306b\u4fc2\u7559\u3055\u308c\uff0c\u30d3\u30aa\u30c1\u30f3-\u30b9\u30c8\u30ec\u30d7\u30c8\u30a2\u30d3\u30b8\u30f3\u7d50\u5408\u3092\u7528\u3044\u305f\u8d85\u5e38\u78c1\u6027\u30d3\u30fc\u30ba\u306b\u56fa\u7740\u3055\u308c\u305f\uff0e\u305d\u306e\u529b\u306f\uff0c\u78c1\u6c17\u30d3\u30fc\u30ba\u3092\u542b\u3080\u6d41\u4f53\u30c1\u30e3\u30f3\u30d0\u30fc\u306e\u4e0a\u306b\u4e00\u5bfe\u306e\u6c38\u4e45\u78c1\u77f3\u3092\u914d\u7f6e\u3059\u308b\u3053\u3068\u306b\u3088\u3063\u3066\u5236\u5fa1\u3055\u308c\u308b\uff0e<\/figcaption><\/figure>\n<p>\u3000\u30bf\u30a4\u30c1\u30f3\u306f\u7b4b\u8089\u306e\u3053\u308c\u3089 4 \u3064\u306e\u9818\u57df\u3067\u5bb9\u6613\u306b\u898b\u3064\u304b\u308b\uff0e\u30bf\u30a4\u30c1\u30f3\u306e N \u672b\u7aef\u306f 9 \u3064\u306e\u514d\u75ab\u30b0\u30ed\u30d6\u30ea\u30f3 (Ig) \u30c9\u30e1\u30a4\u30f3\u304b\u3089\u306a\u308a (<strong>Figure 1A<\/strong>, \u7dd1\u8272), Z \u76e4\u306b\u86cb\u767d\u8cea\u3067\u7d50\u5408\u3057\u3066\u3044\u308b\uff0eI-band \u9818\u57df\u306e\u30bf\u30a4\u30c1\u30f3 (<strong>Figure 1A<\/strong>, \u8d64) \u306f\u9078\u629e\u7684\u306b\u30b9\u30d7\u30e9\u30a4\u30b9\u3055\u308c\u305f\u30a8\u30af\u30bd\u30f3\u304b\u3089\u306a\u308a, 100 \u500b\u306b\u3082\u53ca\u3076 Ig \u30c9\u30e1\u30a4\u30f3\u3092\u30b3\u30fc\u30c9\u3057\u3066\u304a\u308a\uff0c\u672c\u8cea\u7684\u306b\u7121\u79e9\u5e8f\u306a\u9818\u57df\u3068\u540c\u69d8\u3067\uff0c\u3044\u308f\u3086\u308b N2B \u914d\u5217\u304a\u3088\u3073 PEVK \u914d\u5217\u3068\u547c\u3070\u308c\uff0c\u30d7\u30ed\u30ea\u30f3\uff0c\u30b0\u30eb\u30bf\u30df\u30f3\u9178\uff0c\u30d0\u30ea\u30f3\u304a\u3088\u3073\u30ea\u30b8\u30f3\u6b8b\u57fa\u306b\u5bcc\u3080\uff0e\u7279\u8a18\u3059\u3079\u304d\u91cd\u8981\u306a\u3053\u3068\u3067\u3042\u308b\u304c\uff0c\u9078\u629e\u7684\u30b9\u30d7\u30e9\u30a4\u30b7\u30f3\u30b0\u306b\u3088\u308a I-band \u3067\u898b\u3064\u304b\u308b Ig \u30c9\u30e1\u30a4\u30f3\u304a\u3088\u3073\u975e\u69cb\u9020\u7684\u30a2\u30df\u30ce\u9178\u306e\u6570\u306f\uff0c\u5404\u7b4b\u8089\u7fa4\u306b\u3088\u308a\u7570\u306a\u308b\u3068\u8003\u3048\u3089\u308c\u3066\u3044\u308b\uff0e\u6700\u9577\u306e\u9aa8\u683c\u7b4b\u30a2\u30a4\u30bd\u30d5\u30a9\u30fc\u30e0\u306f 100 \u500b\u306e Ig \u30c9\u30e1\u30a4\u30f3\u3092\u6709\u3059\u308b\u306e\u306b\uff0c\u5fc3\u7b4b\u7d44\u7e54\u3067\u652f\u914d\u7684\u306a N2B \u30bf\u30a4\u30c1\u30f3\u306e\u30a2\u30a4\u30bd\u30d5\u30a9\u30fc\u30e0\u306f\u305f\u3063\u305f 47 \u500b\u306e Ig \u30c9\u30e1\u30a4\u30f3\u3068 735 \u500b\u306e\u975e\u69cb\u9020\u30a2\u30df\u30ce\u9178\u6b8b\u57fa\u3092\u6709\u3059\u308b\u306b\u904e\u304e\u306a\u3044\uff0e\u30bf\u30a4\u30c1\u30f3\u306e\u30a2\u30a4\u30bd\u30d5\u30a9\u30fc\u30e0\u306e\u5927\u304d\u3055\u306f\uff0c\u3044\u304b\u306a\u308b\u6240\u4e0e\u306e\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u3067\u30bf\u30a4\u30c1\u30f3\u304c\u529b\u3092\u7d4c\u9a13\u3059\u308b\u304b\u3092\u7406\u89e3\u3059\u308b\u305f\u3081\u306b\u91cd\u8981\u3067\u3042\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/10625306\" rel=\"noopener noreferrer\" target=\"_blank\">18<\/a>). \u30bf\u30a4\u30c1\u30f3\u306e A-band \u9818\u57df\u306f (<strong>Figure 1A<\/strong>, \u7070\u8272) Ig \u30c9\u30e1\u30a4\u30f3\u304a\u3088\u3073\u30d5\u30a3\u30d6\u30ed\u30cd\u30af\u30c1\u30f3\u30c9\u30e1\u30a4\u30f3\u304b\u3089\u306a\u308a\uff0c\u305d\u308c\u3089\u306f\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306b\u56fa\u304f\u7d50\u5408\u3057\u3066\u3044\u308b\uff0e\u6700\u5f8c\u306b\uff0c\u30bf\u30a4\u30c1\u30f3\u306e M-band \u90e8\u4f4d (<strong>Figure 1A<\/strong>, \u9752) \u306f\u30bf\u30a4\u30c1\u30f3\u507d\u30ad\u30ca\u30fc\u30bc (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/24850911\" rel=\"noopener noreferrer\" target=\"_blank\">19<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/15802564\" rel=\"noopener noreferrer\" target=\"_blank\">20<\/a>) \u304a\u3088\u3073 10 \u500b\u306e Ig \u30c9\u30e1\u30a4\u30f3\u304b\u3089\u306a\u308a\uff0c\u96a3\u63a5\u3059\u308b\u30bf\u30a4\u30c1\u30f3\u304a\u3088\u3073\u4ed6\u306e\u69cb\u9020\u86cb\u767d\u8cea\u3092\u7d50\u5408\u3057\u3066\u3044\u308b\uff0e\u3053\u306e\u30ec\u30d3\u30e5\u30fc\u306f\u4e3b\u306b\u30bf\u30a4\u30c1\u30f3\u306e\u5f3e\u6027 I-band \u5207\u7247\u306e\u529b\u306e\u767a\u751f\u80fd\u529b\u306b\u95a2\u4fc2\u3057\u3066\u3044\u308b\u304c\uff0c\u30bf\u30a4\u30c1\u30f3\u3068\u7b4b\u8089\u306e\u5fae\u7d30\u69cb\u9020\u3092\u8b70\u8ad6\u3057\u3066\u3044\u308b\u512a\u79c0\u306a\u8cc7\u6e90\u306f\u591a\u6570\u5b58\u5728\u3059\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/24625729\" rel=\"noopener noreferrer\" target=\"_blank\">21<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/28232805\" rel=\"noopener noreferrer\" target=\"_blank\">22<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/14506471\" rel=\"noopener noreferrer\" target=\"_blank\">23<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/20551232\" rel=\"noopener noreferrer\" target=\"_blank\">24<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/24655499\" rel=\"noopener noreferrer\" target=\"_blank\">25<\/a>). <\/p>\n<p>\u3000\u7b4b\u53ce\u7e2e\u306e\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u6ed1\u8d70\u8aac\u306b\u304a\u3044\u3066\u306f, ATP \u52a0\u6c34\u5206\u89e3\u3068\u7121\u6a5f\u30ea\u30f3\u9178\u306e\u653e\u51fa\u304c\u30df\u30aa\u30b7\u30f3\u9838\u90e8\u306e\u69cb\u9020\u5909\u5316\u3092\u3082\u305f\u3089\u3057\uff0c\u30df\u30aa\u30b7\u30f3\u982d\u90e8\u304c\u30a2\u30af\u30c1\u30f3\u306b\u7d50\u5408\u3059\u308b\uff0e\u3053\u306e\u52d5\u4f5c\u304c\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3092\u52d5\u304b\u3057\uff0c\u30a2\u30af\u30c1\u30f3\u306b\u5bfe\u3059\u308b\u529b\u3092\u767a\u751f\u3055\u305b\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/4939977\" rel=\"noopener noreferrer\" target=\"_blank\">9<\/a>). \u7b4b\u53ce\u7e2e\u306e\u969b\u306b\u30df\u30aa\u30b7\u30f3\u304c\u5358\u72ec\u3067\u30a2\u30af\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3092\u6ed1\u8d70\u3055\u305b, ATP \u306e\u5316\u5b66\u30a8\u30cd\u30eb\u30ae\u30fc\u304c\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u5468\u671f\u306b\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u4f9b\u7d66\u3059\u308b\u3068\u3044\u3046\u70b9\u306f\u9593\u9055\u3044\u306a\u3044 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/8139653\" rel=\"noopener noreferrer\" target=\"_blank\">26<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/5639798\" rel=\"noopener noreferrer\" target=\"_blank\">27<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/20689017\" rel=\"noopener noreferrer\" target=\"_blank\">28<\/a>,<a href=\"https:\/\/www.nature.com\/articles\/ncomms16036\" rel=\"noopener noreferrer\" target=\"_blank\">29<\/a>) \u304c\uff0c\u77ed\u7e2e\u6642\u306e\u30bf\u30a4\u30c1\u30f3\u306e\u5f79\u5272\u306f\u307b\u3068\u3093\u3069\u7406\u89e3\u3055\u308c\u3066\u304a\u3089\u305a\uff0c\u660e\u78ba\u306b\u3059\u308b\u305f\u3081\u306b\u6301\u7d9a\u7684\u306a\u7126\u70b9\u3092\u5f53\u3066\u308b\u3053\u3068\u304c\u7b4b\u8089\u30b3\u30df\u30e5\u30cb\u30c6\u30a3\u306b\u6c42\u3081\u3089\u308c\u3066\u3044\u305f\uff0e\u30bf\u30a4\u30c1\u30f3\u304c\u3044\u304b\u306b\u3057\u3066\u7b4b\u53ce\u7e2e\u306e\u969b\u306b\u529b\u3092\u767a\u751f\u3059\u308b\u306e\u304b\u306b\u3064\u3044\u3066\u306f\uff0c\u3044\u304f\u3064\u304b\u7af6\u5408\u3059\u308b\u4eee\u8aac\u304c\u5b58\u5728\u3059\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/28232805\" rel=\"noopener noreferrer\" target=\"_blank\">22<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/27252165\" rel=\"noopener noreferrer\" target=\"_blank\">30<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/11948204\" rel=\"noopener noreferrer\" target=\"_blank\">31<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC3259925\/\" rel=\"noopener noreferrer\" target=\"_blank\">32<\/a>) \u304c\uff0c\u7b4b\u7dda\u7dad\u30ec\u30d9\u30eb\u3067\u5206\u5b50\u6a5f\u69cb\u3092\u89e3\u660e\u3059\u308b\u8a66\u307f\u306f\u5931\u6557\u3057\u3066\u304d\u305f\uff0e\u904e\u53bb 20 \u5e74\u9593\u306b\u308f\u305f\u308a\uff0c\u30bf\u30f3\u30d1\u30af\u8cea\u5de5\u5b66\u3068\u7d44\u307f\u5408\u308f\u305b\u305f\u5206\u5b50\u9593\u529b\u5206\u5149\u6cd5\u306b\u304a\u3051\u308b\u9032\u6b69\u306b\u3088\u308a\uff0c\u529b\u306e\u4e0b\u3067\u306e\u30bf\u30a4\u30c1\u30f3\u306e\u529b\u5b66\u306e\u7814\u7a76\u304c\u9032\u3093\u3067\u304d\u305f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/26854230\" rel=\"noopener noreferrer\" target=\"_blank\">10<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/10097099\" rel=\"noopener noreferrer\" target=\"_blank\">33<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/15017000\" rel=\"noopener noreferrer\" target=\"_blank\">34<\/a>,<a href=\"http:\/\/science.sciencemag.org\/content\/276\/5315\/1112\" rel=\"noopener noreferrer\" target=\"_blank\">35<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/12198551\" rel=\"noopener noreferrer\" target=\"_blank\">36<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/9148804\" rel=\"noopener noreferrer\" target=\"_blank\">37<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/15123816\" rel=\"noopener noreferrer\" target=\"_blank\">38<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/9153398\" rel=\"noopener noreferrer\" target=\"_blank\">39<\/a>). \u5206\u5b50\u9593\u529b\u5206\u5149\u6cd5\u306e\u30c7\u30fc\u30bf\u306e\u793a\u3059\u3068\u3053\u308d\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u306f Ig \u30c9\u30e1\u30a4\u30f3\u5206\u5b50\u306e\u591a\u304f\u306e\u8155\u3092\u4f38\u5c55\u3057\u305f\u308a\u518d\u5ea6\u6298\u308a\u305f\u305f\u3093\u3060\u308a\u3059\u308b\u3053\u3068\u3067\u5f3e\u6027\u30a8\u30cd\u30eb\u30ae\u30fc\u306e\u307b\u3068\u3093\u3069\u3092\u84c4\u7a4d\u3057\uff0c\u63d0\u4f9b\u3059\u308b\u3068\u3044\u3046\u3082\u306e\u3067\u3042\u308b\uff0e\u4eca\u3084\u6211\u3005\u306e\u8cac\u52d9\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u6298\u308a\u305f\u305f\u307f\u306e\u529b\u5b66\u3092\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u6ed1\u8d70\u8aac\u306e\u67a0\u7d44\u307f\u306b\u7d44\u307f\u8fbc\u3080\u3053\u3068\u3067\uff0c\u7b4b\u53ce\u7e2e\u306e\u77e5\u898b\u3092\u767a\u5c55\u3055\u305b\u308b\u3053\u3068\u3067\u3042\u308b\uff0e\u3055\u3042\uff0c\u3084\u3063\u3066\u307f\u3088\u3046\uff0e<\/p>\n<h3>Studies of titin under force<\/h3>\n<p>\u3000\u5316\u5b66\u7684\u30fb\u71b1\u7684\u5909\u6027\u306a\u3069\u306e\u4f1d\u7d71\u7684\u306a\u904e\u534a\u6570\u306e\u751f\u5316\u5b66\u6280\u8853\u306f\u7b4b\u8089\u5185\u306e\u30bf\u30a4\u30c1\u30f3\u306e\u6a5f\u80fd\u3092\u7406\u89e3\u3059\u308b\u306b\u306f\u5b9f\u7528\u7684\u3067\u306f\u306a\u3044. \u751f\u4f53\u5185\u3067\u30bf\u30a4\u30c1\u30f3\u306f 15 pN \u307e\u3067\u306e\u6a5f\u68b0\u7684\u8ca0\u8377\u306e\u3082\u3068\u3067\u52d5\u4f5c\u3059\u308b\u304c\uff0c\u305d\u308c\u306f Z \u76e4\u3078\u306e\u4ed8\u7740\u3068\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u5148\u7aef\u3068\u306e\u9593\u306e\u30d9\u30af\u30c8\u30eb\u306b\u6cbf\u3063\u3066\u9069\u7528\u3055\u308c\u308b\u8ca0\u8377\u3067\u3042\u308b\uff0e\u539f\u5b50\u9593\u529b\u9855\u5fae\u93e1 (AFM) \u3084\u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u304a\u3088\u3073\u5149\u30d4\u30f3\u30bb\u30c3\u30c8\u306a\u3069\u306e\u5358\u5206\u5b50\u9593\u529b\u5206\u5149\u6cd5\u6280\u8853\u306f\uff0c\u751f\u4f53\u5185\u3067\u7d4c\u9a13\u3059\u308b\u306e\u3068\u540c\u3058\u5354\u8abf\u53cd\u5fdc\u3092\u901a\u3058\u3066\u30bf\u30a4\u30c1\u30f3\u306e\u30ca\u30ce\u30b9\u30b1\u30fc\u30eb\u306e\u6a5f\u5e8f\u3092\u6398\u308a\u4e0b\u3052\uff0c\u904e\u534a\u6570\u306e\u6280\u8853\u3088\u308a\u3082\u3063\u3068\u751f\u7406\u5b66\u7684\u306a\u6d1e\u5bdf\u3092\u3082\u305f\u3089\u3057\u3066\u304f\u308c\u308b\uff0e\u86cb\u767d\u8cea\u30ec\u30d9\u30eb\u306e\u30b3\u30f3\u30d5\u30a9\u30e1\u30fc\u30b7\u30e7\u30f3\u5909\u5316\u3092\u5f15\u304d\u8d77\u3053\u3059\u7d30\u80de\u30d7\u30ed\u30bb\u30b9\u306e\u591a\u69d8\u6027\u3092\u4fee\u98fe\u3059\u308b\u529b\u306e\u65b0\u3057\u3044\u89b3\u70b9\u306b\u3088\u308a\uff0c\u3053\u308c\u3089\u306e\u624b\u6cd5\u306f\u6025\u901f\u306b\u7d30\u80de\u306e\u6a5f\u5e8f\u3092\u7406\u89e3\u3059\u308b\u305f\u3081\u306e\u4e0d\u53ef\u6b20\u306e\u6280\u8853\u3068\u306a\u308a\u3064\u3064\u3042\u308b\uff0e<\/p>\n<p>\u3000\u86cb\u767d\u8cea\u5206\u5b50\u9593\u529b\u5206\u5149\u6cd5\u306e\u5b9f\u9a13\u306e\u5b9f\u65bd\u306b\u306f\uff0c\u56fa\u5b9a\u8868\u9762\u3068\u9855\u5fae\u93e1\u30d7\u30ed\u30fc\u30d6\u306e\u9593\u306e\u5358\u4e00\u306e\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u4e00\u672c\u9396\u3092\u5fc5\u8981\u3068\u3057\uff0c\u305d\u306e\u30d7\u30ed\u30fc\u30d6\u306f\u5f35\u529b\u3092\u304b\u3051\u3089\u308c\u308b\u3088\u3046\u914d\u7f6e\u3067\u304d\u308b\u3082\u306e\u3067\u3042\u308b\uff0eAFM \u7814\u7a76\u306e\u305f\u3081\u306b\uff0c\u5f35\u529b\u30d7\u30ed\u30fc\u30d6\u306f\u7a92\u5316\u30b1\u30a4\u7d20\u88fd\u306e\u30de\u30a4\u30af\u30ed\u30ab\u30f3\u30c1\u30ec\u30d0\u30fc\u3067\u3042\u308a\uff0c\u86cb\u767d\u8cea\u672b\u7aef\u306b\u5171\u6709\u7d50\u5408\u3067\u3082\u975e\u5171\u6709\u7d50\u5408\u3067\u3082\u53d6\u308a\u4ed8\u3051\u308b\u3053\u3068\u304c\u3067\u304d\u308b (<a href=\"https:\/\/www.nature.com\/articles\/nprot.2013.056\" rel=\"noopener noreferrer\" target=\"_blank\">40<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/23909704\" rel=\"noopener noreferrer\" target=\"_blank\">41<\/a>). \u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u306b\u3064\u3044\u3066\u306f\uff0c\u78c1\u754c\u3092\u53d7\u3051\u305f\u78c1\u6027\u4f53\u7c92\u5b50\u3092\u4ed8\u7740\u3055\u305b\u305f\u86cb\u767d\u8cea\u3092\u4e00\u672c\u9396\u306b\u5f15\u304d\u4f38\u3070\u3059 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/26854230\" rel=\"noopener noreferrer\" target=\"_blank\">10<\/a>,<a href=\"https:\/\/www.nature.com\/articles\/nprot.2013.056\" rel=\"noopener noreferrer\" target=\"_blank\">40<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/21244848\" rel=\"noopener noreferrer\" target=\"_blank\">42<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/25726700\" rel=\"noopener noreferrer\" target=\"_blank\">43<\/a>). \u5149\u30d4\u30f3\u30bb\u30c3\u30c8\u306f\u30ec\u30fc\u30b6\u30fc\u30c8\u30e9\u30c3\u30d7\u3092\u7528\u3044\u308b\u3082\u306e\u3067\uff0c\u8a98\u96fb\u4f53\u7c92\u5b50\u3092\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u306b\u4ed8\u7740\u3055\u305b\u529b\u3092\u9069\u7528\u3059\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/8413650\" rel=\"noopener noreferrer\" target=\"_blank\">44<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/24357719\" rel=\"noopener noreferrer\" target=\"_blank\">45<\/a>). \u3059\u3079\u3066\u306e\u4f8b\u3067\uff0c\u5f35\u529b\u30d7\u30ed\u30fc\u30d6\u306e\u4f4d\u7f6e\u306f\u9ad8\u901f\u30ec\u30fc\u30b6\u30fc\u691c\u51fa\u5668\u307e\u305f\u306f\u30d3\u30c7\u30aa\u9855\u5fae\u93e1\u3067\u30ca\u30ce\u30e1\u30fc\u30bf\u30fc\uff0c\u307e\u305f\u306f\u6700\u4f4e\u3067\u3082\u30aa\u30f3\u30b0\u30b9\u30c8\u30ed\u30fc\u30e0\u7cbe\u5ea6\u3067\u8a08\u6e2c\u3055\u308c\u308b\uff0e\u4f38\u5c55\u306b\u3088\u308b\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u9577\u306e\u6f38\u9032\u7684\u306a\u5909\u5316\uff0c\u307e\u305f\u306f\u30c9\u30e1\u30a4\u30f3\u306e\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304a\u3088\u3073\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306b\u3088\u308b\u7a81\u7136\u306e\u5909\u5316\u306f\u9ad8\u3044\u6642\u7a7a\u9593\u89e3\u50cf\u5ea6\u3067\u8a08\u6e2c\u53ef\u80fd\u3067\u3042\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/18511917\" rel=\"noopener noreferrer\" target=\"_blank\">46<\/a>,<a href=\"https:\/\/www.nature.com\/articles\/ncomms6885\" rel=\"noopener noreferrer\" target=\"_blank\">47<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/26588570\" rel=\"noopener noreferrer\" target=\"_blank\">48<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/27409974\" rel=\"noopener noreferrer\" target=\"_blank\">49<\/a>). <\/p>\n<p>\u3000\u521d\u671f\u306e\u5358\u5206\u5b50\u9593\u529b\u5206\u5149\u6cd5\u306e\u5b9f\u9a13\u3067\u306f, DNA (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/8079175\" rel=\"noopener noreferrer\" target=\"_blank\">50<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/10201403\" rel=\"noopener noreferrer\" target=\"_blank\">51<\/a>) \u3068\u591a\u7cd6\u985e (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/9036852\" rel=\"noopener noreferrer\" target=\"_blank\">52<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/9872313\" rel=\"noopener noreferrer\" target=\"_blank\">53<\/a>) \u306e\u6a5f\u5e8f\uff0c\u53d7\u5bb9\u4f53\u3068\u30ea\u30ac\u30f3\u30c9\u306e\u89e3\u96e2 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/8153628\" rel=\"noopener noreferrer\" target=\"_blank\">54<\/a>), \u30e2\u30fc\u30bf\u30fc\u86cb\u767d\u529b\u5b66 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/8139653\" rel=\"noopener noreferrer\" target=\"_blank\">26<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/8413650\" rel=\"noopener noreferrer\" target=\"_blank\">44<\/a>), \u305d\u3057\u3066\u9175\u7d20\u306e\u30b3\u30f3\u30d5\u30a9\u30e1\u30fc\u30b7\u30e7\u30f3\u5909\u5316 (<a href=\"http:\/\/science.sciencemag.org\/content\/265\/5178\/1577\" rel=\"noopener noreferrer\" target=\"_blank\">55<\/a>) \u3092\u8abf\u67fb\u3057\u305f\uff0e\u3057\u304b\u3057\uff0c\u3054\u304f\u521d\u671f\u306e\u30ca\u30ce\u30e1\u30fc\u30c8\u30eb\u89e3\u50cf\u5ea6\u3067\u306e\u53ef\u9006\u7684\u306a\u86cb\u767d\u8cea\u306e\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u8a18\u9332\u306f\uff0c\u30de\u30a4\u30af\u30ed\u30e1\u30fc\u30c8\u30eb\u9577\u306e\u30bf\u30a4\u30c1\u30f3\u3092\u57fa\u8cea\u3068\u3057\u3066\u5b9f\u884c\u3055\u308c\u305f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/9148804\" rel=\"noopener noreferrer\" target=\"_blank\">37<\/a>). AFM \u3092\u7528\u3044\u3066\uff0c\u30a6\u30b7\u306e\u7d44\u7e54\u304b\u3089\u7cbe\u88fd\u3055\u308c\u305f\u5929\u7136\u306e\u30bf\u30a4\u30c1\u30f3\u306f\u9ad8\u3044\u529b\u3067\u4f38\u5c55\u3055\u308c\uff0c\u30ab\u30f3\u30c1\u30ec\u30d0\u30fc\u5148\u7aef\u3092\u5f15\u304d\u4e0a\u3052\u9762\u306b\u623b\u3059\u3053\u3068\u3067\u518d\u30d5\u30a9\u30fc\u30eb\u30c9\u3067\u304d\u308b\u3088\u3046\u306b\u3057\u305f\uff0e\u30bf\u30a4\u30c1\u30f3\u306f\u3053\u308c\u3089\u306e\u5b9f\u9a13\u306e\u305f\u3081\u306e\u81ea\u7136\u306e\u9078\u629e\u3067\u3042\u3063\u305f\uff0e\u3068\u3044\u3046\u306e\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u306f\u591a\u304f\u306e\u30c9\u30e1\u30a4\u30f3\u304c\u307b\u307c\u540c\u3058\u5927\u304d\u3055\u306e\u514d\u75ab\u30b0\u30ed\u30d6\u30ea\u30f3\u304b\u3089\u306a\u308a\uff0c\u4e00\u5b9a\u901f\u5ea6\u3067\u5358\u5206\u5b50\u3092\u727d\u5f15\u3059\u308b\u5b9f\u9a13\u306b\u304a\u3044\u3066\u7279\u5fb4\u7684\u306a\u300c\u92f8\u6b6f\u72b6\u300d\u30d1\u30bf\u30fc\u30f3\u3092\u5448\u3057\u3066\u3044\u305f\u304b\u3089\u3067\u3042\u308b\uff0e\u540c\u69d8\u306e\u7d50\u679c\u304c\u5149\u30d4\u30f3\u30bb\u30c3\u30c8\u3092\u7528\u3044\u3066\u5929\u7136\u306e\u30bf\u30a4\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3092\u9ad8\u3044\u529b\u3067\u4f38\u5c55\u3059\u308b\u5b9f\u9a13\u3067\u793a\u3055\u308c\u3066\u304a\u308a\uff0c\u86cb\u767d\u8cea\u306e\u6bb5\u968e\u7684\u69d8\u5f0f\u307e\u305f\u306f\u6f38\u5897\u69d8\u5f0f\u304c\u5f97\u3089\u308c\uff0c\u3053\u308c\u306f\u5358\u4e00\u30c9\u30e1\u30a4\u30f3\u306e\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304c\u8d77\u304d\u305f\u3053\u3068\u3092\u793a\u5506\u3057\u3066\u3044\u308b (<a href=\"http:\/\/science.sciencemag.org\/content\/276\/5315\/1112\" rel=\"noopener noreferrer\" target=\"_blank\">35<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/9153398\" rel=\"noopener noreferrer\" target=\"_blank\">39<\/a>). \u907a\u4f1d\u5b50\u7d44\u63db\u3048\u86cb\u767d\u8cea\u5de5\u5b66\u304a\u3088\u3073\u5206\u5b50\u30af\u30ed\u30fc\u30cb\u30f3\u30b0\u6280\u8853\u306f\uff0c\u975e\u5e38\u306b\u8a73\u7d30\u306b\u7814\u7a76\u3055\u308c\u305f\u5f3e\u6027\u30bf\u30a4\u30c1\u30f3 I-band \u306e\u4f4d\u7f6e\u3092\u3088\u308a\u5c0f\u3055\u304f\uff0c\u3088\u308a\u5236\u5fa1\u53ef\u80fd\u306b\u3057\u3066\u3044\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/10097099\" rel=\"noopener noreferrer\" target=\"_blank\">33<\/a>,<a href=\"http:\/\/www.pnas.org\/content\/96\/20\/11288\" rel=\"noopener noreferrer\" target=\"_blank\">56<\/a>). \u3053\u308c\u306b\u3088\u308a Ig \u30c9\u30e1\u30a4\u30f3\u5185\u90e8\u306e\u6a5f\u68b0\u7684\u968e\u5c64\u304c\u660e\u3089\u304b\u306b\u306a\u308a\uff0c\u8fd1\u4f4d Ig \u30c9\u30e1\u30a4\u30f3\uff08N \u672b\u7aef\u306b\u8fd1\u3044\u90e8\u4f4d\uff09\u306e\u6a5f\u68b0\u7684\u5b89\u5b9a\u6027\u304c I-band \u306e\u9060\u4f4d Ig \u30c9\u30e1\u30a4\u30f3\u3088\u308a\u3082\u4f4e\u3044\u3053\u3068\u304c\u5206\u304b\u3063\u305f\uff0e\u3055\u3089\u306b, HaloTag \u30a2\u30f3\u30ab\u30fc\u306e\u86cb\u767d\u8cea\u306e N \u307e\u305f\u306f C \u672b\u7aef\u3078\u306e\u5171\u6709\u7d50\u5408\u306e\u5de5\u5b66\u306f\uff0c\u5f37\u3044\u529b\u3067\u9577\u6642\u9593\u306e\u7814\u7a76\u306e\u305f\u3081\u306e\u907a\u4f1d\u5b50\u7d44\u307f\u63db\u3048\u30bf\u30a4\u30c1\u30f3\u306e\u7279\u7570\u7684\u56fa\u5b9a\u5316\u3078\u306e\u9053\u3092\u958b\u3044\u305f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/23909704\" rel=\"noopener noreferrer\" target=\"_blank\">41<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/27409974\" rel=\"noopener noreferrer\" target=\"_blank\">49<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/28470663\" rel=\"noopener noreferrer\" target=\"_blank\">57<\/a>). <\/p>\n<p>\u3000\u904e\u53bb 20 \u5e74\u9593\u306b\u308f\u305f\u308a\uff0c\u5206\u5b50\u9593\u529b\u5206\u5149\u6cd5\u306f\u8a08\u88c5\u3068\u6280\u8853\u306b\u304a\u3044\u3066\u5927\u304d\u306a\u6539\u5584\u306e\u6069\u6075\u3092\u53d7\u3051\u305f\uff0e\u96fb\u4f4d\u56fa\u5b9a\u6cd5\u3092\u9451\u307f\u308b\u306b\uff0c\u5358\u4e00\u306e\u30a4\u30aa\u30f3\u30c1\u30e3\u30cd\u30eb\u30b2\u30fc\u30c8\u306e\u5b9f\u9a13\u306e\u521d\u3081\u3066\u306e\u6210\u529f\u3092\u652f\u3048\u305f\u3082\u306e\u3067\u3042\u308b\u304c, AFM \u306e\u305f\u3081\u306e\u539f\u5b50\u9593\u529b\u56fa\u5b9a\u6cd5\u306e\u767a\u5c55\u306f\u86cb\u767d\u8cea\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u30a8\u30cd\u30eb\u30ae\u30fc\u8ad6\u306b\u3064\u3044\u3066\u6211\u3005\u306e\u7406\u89e3\u3092\u5927\u304d\u304f\u9032\u6b69\u3055\u305b\u305f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/15017000\" rel=\"noopener noreferrer\" target=\"_blank\">34<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/15123816\" rel=\"noopener noreferrer\" target=\"_blank\">38<\/a>,<a href=\"http:\/\/www.pnas.org\/content\/98\/2\/468\" rel=\"noopener noreferrer\" target=\"_blank\">58<\/a>). \u539f\u5b50\u9593\u529b\u56fa\u5b9a\u5206\u5149\u6cd5\u306f\u660e\u767d\u306b\uff0c\u5b9a\u7fa9\u3055\u308c\u305f\u727d\u5f15\u529b\u306e\u4e0b\u3067\u306e\uff0c\u5358\u4e00\u306e\u86cb\u767d\u8cea\u30da\u30d7\u30c1\u30c9\u306e\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304a\u3088\u3073\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u8ecc\u8de1\u306b\u4ed8\u304d\u6dfb\u3044\uff0c\u30aa\u30f3\u30b0\u30b9\u30c8\u30ed\u30fc\u30e0\u5358\u4f4d\u306e\u89e3\u50cf\u5ea6\u3067\u86cb\u767d\u8cea\u306e\u672b\u7aef\u304b\u3089\u672b\u7aef\u307e\u3067\u306e\u9577\u3055\u3092\u8ffd\u8de1\u3059\u308b\uff0e\u3053\u308c\u306b\u3088\u308a\uff0c\u529b\u5b66\u3068\u5916\u529b\u4f9d\u5b58\u6027\u306e\u86cb\u767d\u8cea\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3084\u4ed6\u306e\u751f\u5316\u5b66\u53cd\u5fdc\u3068\u306e\u7814\u7a76\u304c\u9032\u3093\u3060 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/16645035\" rel=\"noopener noreferrer\" target=\"_blank\">59<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/17972886\" rel=\"noopener noreferrer\" target=\"_blank\">60<\/a>). \u3082\u3063\u3068\u6700\u8fd1\u3067\u306f\uff0c\u53d7\u52d5\u7684\u306b\u7121\u9650\u306e\u5e2f\u57df\u5e45\u3067\u52d5\u4f5c\u3059\u308b\u86cb\u767d\u8cea\u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u306e\u767a\u5c55\u306b\u3088\u308a, 15 pN \u4ee5\u4e0b\u306e\u529b\u3067\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u306b\u304a\u3051\u308b\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u30fb\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u9077\u79fb\u5e73\u8861\u3092\u76f4\u63a5\u89b3\u6e2c\u3067\u304d\u308b\u3088\u3046\u306b\u306a\u3063\u305f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/26854230\" rel=\"noopener noreferrer\" target=\"_blank\">10<\/a>,<a href=\"http:\/\/Dynamics of Equilibrium Folding and Unfolding Transitions of Titin Immunoglobulin Domain under Constant Forces.\" rel=\"noopener noreferrer\" target=\"_blank\">43<\/a>). \u3053\u306e\u7a2e\u306e\u5b9f\u9a13\u306f\uff0c\u751f\u4f53\u5185\u3067\u52d5\u4f5c\u3059\u308b\u306e\u3068\u6975\u3081\u3066\u3088\u304f\u4f3c\u305f\u72b6\u614b\u3067\u306e\u30bf\u30a4\u30c1\u30f3\u3092\u63a2\u7d22\u3059\u308b\u305f\u3081\u306e\u81ea\u7136\u306a\u30b7\u30ca\u30ea\u30aa\u3092\u63d0\u4f9b\u3057\u3066\u304f\u308c\u308b. <strong>Figure 1B<\/strong> \u306b\u5b9f\u9a13\u4f8b\u3092\u56f3\u793a\u3057\u305f\u304c\uff0c\u4eca\u65e5\u6211\u3005\u304c\u30bf\u30a4\u30c1\u30f3\u306e\u6a5f\u5e8f\u306b\u3064\u3044\u3066\u77e5\u3063\u3066\u3044\u308b\u3053\u3068\u306e\u591a\u304f\u3092\u8981\u7d04\u3057\u3066\u3042\u308b\uff0e\u3053\u306e\u5b9f\u9a13\u306b\u304a\u3044\u3066\u306f 8 \u672c\u306e\u8155\u3092\u6301\u3064\u5358\u4e00\u306e\u86cb\u767d\u8cea Ig 91 (I91) \u30c9\u30e1\u30a4\u30f3\u304c\u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u88c5\u7f6e\u306e\u4e2d\u3067\u4e00\u5b9a\u306e\u4f38\u5c55\u529b\u306e\u4e0b\u306b\u56fa\u5b9a\u3055\u308c\u3066\u3044\u308b\uff0e\u91cd\u8981\u306a\u3053\u3068\u306a\u306e\u3067\u66f8\u3044\u3066\u304a\u304f\u304c\uff0c\u3053\u306e I91 \u30c9\u30e1\u30a4\u30f3\u306f\u53e4\u3044\u6587\u732e (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/12198551\" rel=\"noopener noreferrer\" target=\"_blank\">36<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC1697845\/\" rel=\"noopener noreferrer\" target=\"_blank\">61<\/a>) \u3067\u8a00\u53ca\u3055\u308c\u3066\u3044\u308b I27 \u30c9\u30e1\u30a4\u30f3\u3068\u540c\u4e00\u3067\u3042\u308a\uff0c\u547d\u540d\u6cd5\u304c\u5909\u5316\u3057\u305f\u306e\u307f\u3067\u3042\u308b\uff0e\u305d\u306e\u86cb\u767d\u8cea\u306f\u5f53\u521d, 105 pN \u3082\u306e\u975e\u751f\u7406\u7684\u306a\u5f37\u3044\u529b\u3067\u3042\u3063\u3068\u3044\u3046\u9593\u306b\u3059\u3079\u3066\u306e Ig \u30c9\u30e1\u30a4\u30f3\u304c\u4f38\u5c55\u3057\u3066\u3057\u307e\u3063\u305f\uff0e\u3053\u308c\u306f\u6700\u521d\u306b\u5f3e\u6027\u306e\u4f38\u5c55\u304c\u8d77\u3053\u308a\uff0c\u5f15\u304d\u7d9a\u3044\u3066\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3068 8 \u3064\u306e Ig \u30c9\u30e1\u30a4\u30f3\u306e\u4f38\u9577\u306b\u306f\u305f\u3063\u305f 20 \u79d2\u9593\u3057\u304b\u304b\u304b\u3089\u306a\u304b\u3063\u305f\uff0e\u5bfe\u7167\u7684\u306b, 10 pN \u306e\u751f\u7406\u7684\u306a\u529b\u3067\u306f I91 \u306e\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306f\u6570\u6642\u9593\u3092\u8981\u3057\u305f (<strong>Figure 1B<\/strong> \u7070\u8272\u70b9\u7dda). \u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u306b\u304b\u304b\u308b\u529b\u304c 5 pN \u306b\u307e\u3067\u6e1b\u5c11\u3059\u308b\u3068\uff0c\u30bf\u30a4\u30c1\u30f3\u306f\u5373\u6642\u5f3e\u529b\u30b3\u30a4\u30eb\u306b\u5909\u5316\u3057\u3066\u5f15\u304d\u7d9a\u304d\u6bb5\u968e\u7684\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53cd\u5fdc\u3092\u793a\u3057\u305f\uff0e<\/p>\n<p>\u3000\u539f\u5b50\u9593\u529b\u5206\u5149\u6cd5\u306b\u3088\u308a\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c9\u3057\u305f\u6700\u521d\u306e\u86cb\u767d\u8cea\u304c\u30bf\u30a4\u30c1\u30f3\u3067\u3042\u3063\u305f\u305f\u3081\uff0c\u5f15\u304d\u7d9a\u3044\u3066\u86cb\u767d\u8cea\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u7269\u7406\u5b66\u3092\u7406\u89e3\u3059\u308b\u30e2\u30c7\u30eb\u3068\u306a\u3063\u305f\uff0e\u30bf\u30a4\u30c1\u30f3\u306e\u3088\u3046\u306b\u7403\u72b6\u306b\u6298\u308a\u305f\u305f\u307e\u308c\u308b\u3053\u3068\u306e\u6a5f\u68b0\u7684\u5b89\u5b9a\u6027\u306f\u9759\u96fb\u76f8\u4e92\u4f5c\u7528\uff0c\u6c34\u7d20\u7d50\u5408\u304a\u3088\u3073\u758e\u6c34\u76f8\u4e92\u4f5c\u7528\u306b\u4f9d\u5b58\u3057\uff0c\u86cb\u767d\u8cea\u306e\u914d\u5217\u304a\u3088\u3073\u4f4d\u76f8\u306b\u3088\u308a\u652f\u914d\u3055\u308c\u308b. <strong>Figure 2A<\/strong> \u306b, 6 pN \u306e\u6a5f\u68b0\u7684\u529b\u3067\u53ef\u9006\u7684\u306b\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3059\u308b\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u306e\u8a73\u7d30\u56f3\u3092\u793a\u3059\uff0e\u529b\u306e\u5927\u90e8\u5206\u306f\u6700\u521d\u306e (A-A&#8217;) \u3068\u6700\u5f8c\u306e (G) \u30d9\u30fc\u30bf\u9396\u3068\u306e\u9593\u306e\u6c34\u7d20\u7d50\u5408\u306b\u3088\u308a\u7dad\u6301\u3055\u308c (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/17525328\" rel=\"noopener noreferrer\" target=\"_blank\">62<\/a>), \u5076\u767a\u7684\u306b\u30c9\u30e1\u30a4\u30f3\u306e\u5b8c\u5168\u306a\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304c\u8d77\u304d\u308b\uff0e\u3053\u308c\u304c\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u306b\u3068\u3063\u3066\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304c\u8d77\u304d\u308b\u6a5f\u5e8f\u3068\u8003\u3048\u3089\u308c\u3066\u304a\u308a, I91 \u3068\u306e\u9ad8\u3044\u69cb\u9020\u7684\u985e\u4f3c\u6027\u304c\u8a8d\u3081\u3089\u308c\u308b\uff0e\u4e00\u65e6\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304c\u8d77\u304d\u308b\u3068\uff0c\u305d\u306e\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u306f\u30e9\u30f3\u30c0\u30e0\u30b3\u30a4\u30eb\u3068\u306a\u308b\uff0e\u305d\u306e\u610f\u5473\u3059\u308b\u3068\u3053\u308d\u306f\uff0c\u672b\u7aef\u9593\u306e\u8ddd\u96e2\u306f\uff0c\u3088\u304f\u77e5\u3089\u308c\u3066\u3044\u308b\u30dd\u30ea\u30de\u30fc\u7269\u7406\u5b66\u6cd5\u5247\u306b\u3088\u308b\u5f35\u529b\u3068\u898f\u6a21\u306b\u3088\u308a\u5b9a\u7fa9\u3055\u308c\u308b\u3068\u3044\u3046\u3053\u3068\u3067\u3042\u308b (<a href=\"https:\/\/amzn.to\/2Morq2V\" rel=\"noopener noreferrer\" target=\"_blank\">63<\/a>). \u4f8b\u3048\u3070, 6 pN \u306b\u304a\u3044\u3066\uff0c\u6298\u308a\u305f\u305f\u307e\u308c\u305f I91 \u30c9\u30e1\u30a4\u30f3\u306f\u672b\u7aef\u304b\u3089\u672b\u7aef\u3078\u306e\u9577\u3055\u304c\u7d04 4 nm \u3067\u3042\u308a\uff08\u539f\u5b50\u5ea7\u6a19\u304b\u3089\u63a8\u6e2c\uff0e\u8a73\u7d30\u306f PDB 1WAA \u53c2\u7167\u306e\u3053\u3068(<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC2643529\/\" rel=\"noopener noreferrer\" target=\"_blank\">64<\/a>)\uff09\uff0c\u4e00\u65b9\u3067\u4f38\u5c55\u3055\u308c\u305f\u30dd\u30ea\u30de\u30fc\u306f 17 nm \u306e\u9577\u3055\u3092\u6301\u3064\uff0e\u3086\u3048\u306b\uff0c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3068\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u9077\u79fb\u671f\u306b\u89b3\u5bdf\u3055\u308c\u305f\u30b9\u30c6\u30c3\u30d7\u306f 6 pN \u306e\u529b\u306e\u4e0b\u3067\u306f 13 nm \u306e\u30b5\u30a4\u30ba\u3092\u6709\u3059\u308b. <strong>Figure 1B<\/strong> \u3082\u307e\u305f 5.0 pN, 10.0 pN \u304a\u3088\u3073 105 pN \u306e\u529b\u3067\u306e\u3053\u306e\u5927\u304d\u3055\u306e\u5909\u5316\u3092\u793a\u3057\u3066\u3044\u308b\uff0e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304a\u3088\u3073\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u9077\u79fb\u3092\u30b9\u30ad\u30e3\u30f3\u3059\u308b\u3068\u5e83\u7bc4\u56f2\u306e\u529b\u304c\u81ea\u7531\u7d50\u5408\u9396\u30e2\u30c7\u30eb\u306b\u9069\u5408\u3059\u308b\u30c7\u30fc\u30bf\u3092\u8a31\u5bb9\u3059\u308b\u3053\u3068\u304c\u5206\u304b\u308b (<a href=\"https:\/\/archive.org\/details\/FloryF.1953PrinciplesOfPolymerChemistry\" rel=\"noopener noreferrer\" target=\"_blank\">63<\/a>). \u305d\u306e\u8f2a\u90ed\u9577\u306f 25 nm \u3067\u3042\u308a, Kuhn \u9577\u306f 0.9 nm \u3067\u3042\u308b (<strong>Figure 2B<\/strong>, \u5b9f\u7dda). \u30b4\u30e0\u30d0\u30f3\u30c9\u306e\u7dda\u5f62\u5f3e\u6027\u3068\u306f\u5bfe\u7167\u7684\u306b\uff0c\u30dd\u30ea\u30de\u30fc\u306e\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc\u53cd\u52d5\u306f\u9ad8\u5ea6\u306b\u975e\u7dda\u5f62\u6027\u3067\u3042\u308a\uff0c\u751f\u7406\u5b66\u7684\u306a\u529b\u306e\u7bc4\u56f2\u3092\u8d85\u3048\u3066\u305d\u306e\u6700\u5927\u4f38\u5c55\u306e 50 % \u306b\u3082\u9054\u3059\u308b\uff0e\u3053\u306e\u9ad8\u3044\u529b\u306b\u304a\u3044\u3066\u306f\uff0c\u86cb\u767d\u8cea\u306e\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u30b9\u30c6\u30c3\u30d7\u30b5\u30a4\u30ba\u306f\u6f38\u8fd1\u7684\u306b\u8f2a\u90ed\u9577\u306b\u8fd1\u3065\u304f\uff0e<\/p>\n<figure id=\"attachment_3552\" aria-describedby=\"caption-attachment-3552\" style=\"width: 525px\" class=\"wp-caption aligncenter\"><a href=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure2.png\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure2-1024x719.png\" alt=\"\" width=\"525\" height=\"369\" class=\"size-large wp-image-3552\" srcset=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure2-1024x719.png 1024w, https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure2-300x211.png 300w, https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure2-768x539.png 768w, https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure2.png 1325w\" sizes=\"auto, (max-width: 525px) 100vw, 525px\" \/><\/a><figcaption id=\"caption-attachment-3552\" class=\"wp-caption-text\"><strong>Figure 2<\/strong><br \/><strong>Ig \u30c9\u30e1\u30a4\u30f3\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306b\u3088\u3063\u3066\u767a\u751f\u3057\u305f\u6a5f\u68b0\u7684\u4ed5\u4e8b\u306f\uff0c\u30d5\u30a9\u30fc\u30b9\u30af\u30e9\u30f3\u30d7\u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u3092\u7528\u3044\u3066\u5b9a\u91cf\u53ef\u80fd\u3067\u3042\u308b<\/strong><br \/><strong>(A)<\/strong> \u6a5f\u68b0\u7684\u5f35\u529b\u306e\u4e0b\u3067\u306e Ig \u30c9\u30e1\u30a4\u30f3\u306e\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\uff0c\u518d\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0. 6 pN \u307e\u3067\u306e\u727d\u5f15\u529b\u306e\u4e0b\u3067\u306f, Ig \u30c9\u30e1\u30a4\u30f3\u306f\uff0c\u30dd\u30ea\u30de\u30fc\u7269\u7406\u5b66\u306e\u30b9\u30b1\u30fc\u30ea\u30f3\u30b0\u5247\u306b\u5f93\u3044\uff0c\u5e73\u8861\u72b6\u614b\u3067\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304a\u3088\u3073\u518d\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3057\uff0c\u7d04 13 nm \u4f38\u9577\u307e\u305f\u306f\u77ed\u7e2e\u3059\u308b. <strong>(B)<\/strong> \u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u30fb\u518d\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u9593\u306b I10 \u304a\u3088\u3073 I91 \u306e\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u306e\u529b\u306e\u95a2\u6570\u3068\u3057\u3066\u6e2c\u5b9a\u3055\u308c\u305f\u30b9\u30c6\u30c3\u30d7\u30b5\u30a4\u30ba\u306e\u5927\u304d\u3055\uff0e\u30b9\u30c6\u30c3\u30d7\u30b5\u30a4\u30ba\u306e\u4f9d\u5b58\u6027\u306f, Kuhn \u9577 lk = 0.9 nm, \u8f2a\u90ed\u9577 Lc = 25 nm \u306e\u81ea\u7531\u7d50\u5408\u9396\u30e2\u30c7\u30eb\uff08\u5b9f\u7dda\uff09\u3067\u3088\u304f\u8a18\u8ff0\u3067\u304d\u308b\uff0e <strong>(C)<\/strong> \u6240\u4e0e\u306e\u727d\u5f15\u529b\u3067\uff0c\u6298\u308a\u7573\u307e\u308c\u305f\u30c9\u30e1\u30a4\u30f3\u306e\u5e73\u8861\u5206\u753b\uff08\u9752\u3044\u5186\uff09\u304b\u3089\u8a08\u7b97\u3055\u308c\u305f, Ig \u30c9\u30e1\u30a4\u30f3\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u78ba\u7387\uff0e\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u306f, 4 pN \u306e\u4f4e\u3044\u529b\u3067\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3057, 10 pN \u306e\u9ad8\u3044\u529b\u3067\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3059\u308b\u3053\u3068\u304c\u3067\u304d\u308b\uff0e\u305d\u306e\u30c7\u30fc\u30bf\u306f, 6 pN \u307e\u3067\u306e\u4e2d\u9593\u70b9\u3092\u6709\u3059\u308b\u30b7\u30b0\u30e2\u30a4\u30c9\u95a2\u6570\uff08\u5b9f\u7dda\uff09\u306b\u3088\u3063\u3066\u3088\u304f\u8868\u3059\u3053\u3068\u304c\u3067\u304d\u308b\uff0e <strong>(D)<\/strong> \u30bf\u30a4\u30c1\u30f3 Ig\u30c9\u30e1\u30a4\u30f3\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306b\u3088\u308a\u751f\u6210\u3055\u308c\u308b\u4e88\u60f3\u3055\u308c\u308b\u4ed5\u4e8b\u306f\uff0c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u78ba\u7387\uff08\u9752\u7dda\uff09\u3068\u3001\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53ce\u7e2e\u306b\u3088\u3063\u3066\u884c\u308f\u308c\u305f\u4ed5\u4e8b\uff08\u8d64\u7dda\uff09\u3068\u306e\u7a4d\u3068\u3057\u3066\u5f97\u3089\u308c\uff0c\u30b9\u30c6\u30c3\u30d7\u30b5\u30a4\u30ba\u306b\u9069\u5408\u3059\u308b\u3088\u3046\u7528\u3044\u305f\u81ea\u7531\u7d50\u5408\u9396\u30e2\u30c7\u30eb\u306e\u6570\u5024\u7a4d\u5206\u304b\u3089\u5f97\u3089\u308c\u308b\uff0e\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u306f 5.7 pN \u3067 46 zJ \u306e\u30d4\u30fc\u30af\u4ed5\u4e8b\u3092\u751f\u6210\u3057\uff0c\u305d\u308c\u306f\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306b\u3088\u308a\u4f9b\u7d66\u3055\u308c\u308b\u30a8\u30cd\u30eb\u30ae\u30fc\u306b\u5339\u6575\u3059\u308b\uff0e<\/figcaption><\/figure>\n<p>\u3000\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u306e\u305d\u308c\u305e\u308c\u304c\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3059\u308b\u969b\u306e\u7387\u306f\u5927\u5e45\u306b\u7570\u306a\u308b\uff0e\u305d\u306e\u305f\u3081\u751f\u7406\u7684\u306a\u4f38\u5c55\u4e2d\u306b\u306f\uff0c\u5168\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u306e\u4e00\u90e8\u306e\u307f\u304c\u4f38\u5c55\u3055\u308c\u308b\u53ef\u80fd\u6027\u304c\u3042\u308b. I-band \u304b\u3089\u63a1\u53d6\u3057\u305f 8 \u672c\u8155\u306e Ig \u30c9\u30e1\u30a4\u30f3\u306e\u91ce\u751f\u578b\u30bb\u30b0\u30e1\u30f3\u30c8\u3092\u30af\u30ed\u30fc\u30cb\u30f3\u30b0\u3057\u3066\u307f\u308b\u3068\uff0c\u30bf\u30a4\u30c1\u30f3\u5185\u90e8\u306e\u6a5f\u68b0\u7684\u968e\u5c64\u304c\u660e\u3089\u304b\u306b\u306a\u3063\u305f\uff0e\u8fd1\u4f4d I-band \u5185\u90e8\u306e Ig \u30c9\u30e1\u30a4\u30f3\u306f Z \u76e4\u306b\u8fd1\u3044\u307b\u3069\uff0c\u9060\u4f4d I-band Ig \u30c9\u30e1\u30a4\u30f3\u3088\u308a\u3082\u306f\u308b\u304b\u306b\u5c0f\u3055\u3044\u529b\u3067\u4f38\u5c55\u3057\uff0c\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306b\u8fd1\u3044 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/12198551\" rel=\"noopener noreferrer\" target=\"_blank\">36<\/a>). \u3086\u3048\u306b, I91 \u304c 10 pN \u3067\u4f38\u5c55\u3059\u308b\u306e\u306b\u6570\u6642\u9593\u3092\u8981\u3059\u308b\u306b\u3082\u304b\u304b\u308f\u3089\u305a (43,65), \u8fd1\u4f4d Ig \u30c9\u30e1\u30a4\u30f3 10 (I10) \u306f\u306f\u308b\u304b\u306b\u65e9\u304f\u4f38\u5c55\u3059\u308b\uff0eI10 \u3067\u306e\u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u5b9f\u9a13\u3067\u660e\u3089\u304b\u306b\u306a\u3063\u305f\u306e\u306f, 6.0 pN \u306e\u529b\u3067\u306f, 20 \u5206\u9593\u3067\u7d04 66 \u56de\u306e\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u30a4\u30d9\u30f3\u30c8\u304c\u767a\u751f\u3057\uff0c\u305d\u308c\u305e\u308c\u306e\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u30a4\u30d9\u30f3\u30c8\u9593\u306b\u306f 15 \u79d2\u9593\u304c\u3042\u308b\u3068\u3044\u3046\u3053\u3068\u3067\u3042\u308b\uff0e\u305d\u308c\u3086\u3048 10 pN \u306b\u304a\u3044\u3066\u306f, I10 \u306f\u6570\u79d2\u9593\u3067\u4f38\u5c55\u3059\u308b\u3068\u63a8\u6e2c\u3055\u308c\u308b\u304c\uff0c\u3053\u308c\u306f\u5b9f\u9a13\u3067\u691c\u8a3c\u3059\u308b\u5fc5\u8981\u304c\u3042\u308b\uff0e\u91ce\u751f\u578b\u306e I4-I11 \u30bb\u30b0\u30e1\u30f3\u30c8\u3092\u7528\u3044\u305f\u5b9f\u9a13\u3067 5.0-6.6 pN \u9593\u306e\u529b\u3067\u306f\uff0c\u7d20\u65e9\u3044\u4f38\u5c55\u30a4\u30d9\u30f3\u30c8\u304c\u89b3\u6e2c\u3055\u308c\uff0c\u305d\u308c\u3086\u3048\u6a5f\u68b0\u7684\u5b89\u5b9a\u6027\u306f\u4f4e\u3044\u3068\u3055\u308c\u305f\uff0e\u3053\u308c\u306f\u8fd1\u4f4d Ig \u30c9\u30e1\u30a4\u30f3\u304c\u751f\u7406\u7684\u306a\u529b\u306e\u7bc4\u56f2\u3067\u5bb9\u6613\u306b\u4f38\u5c55\u3059\u308b\u3068\u3044\u3046\u8996\u70b9\u3092\u652f\u6301\u3057\u3066\u3044\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/26854230\" rel=\"noopener noreferrer\" target=\"_blank\">10<\/a>). <\/p>\n<p>\u3000\u30bf\u30a4\u30c1\u30f3\u30c9\u30e1\u30a4\u30f3\u306e\u4f38\u5c55\u7d4c\u8def\u306b\u95a2\u3057\u3066\u306f\u8c4a\u5bcc\u306a\u60c5\u5831\u304c\u3042\u308b\u306b\u3082\u304b\u304b\u308f\u3089\u305a\uff0c\u6298\u308a\u305f\u305f\u307f\u7d4c\u8def\u306e\u89b3\u5bdf\u306f\u3055\u3089\u306b\u898b\u3064\u304b\u308a\u306b\u304f\u304f, force clamp \u6cd5\u306e\u3088\u3046\u306a\u6280\u8853\u306e\u767a\u5c55\u3092\u5f85\u305f\u306d\u3070\u306a\u3089\u306a\u304b\u3063\u305f (<a href=\"http:\/\/www.pnas.org\/content\/98\/2\/468\" rel=\"noopener noreferrer\" target=\"_blank\">58<\/a>). \u305d\u306e force clamp \u6cd5\u306b\u3088\u308a\uff0c\u30e9\u30f3\u30c0\u30e0\u30b3\u30a4\u30eb\u304b\u3089\u81ea\u7136\u306a\u6298\u308a\u305f\u305f\u307f\u3078\u306e\u86cb\u767d\u8cea\u306e\u5f62\u614b\u306e\u5909\u5f62\u306e\u9593\u306b\uff0c\u4e09\u3064\u306e\u7570\u306a\u308b\u9077\u79fb\u304c\u3042\u308b\u3053\u3068\u304c\u660e\u3089\u304b\u306b\u306a\u3063\u305f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/15017000\" rel=\"noopener noreferrer\" target=\"_blank\">34<\/a>). \u305d\u306e\u7d50\u679c\uff0c\u5f93\u6765\u306e\u4e00\u62ec\u6cd5\u3092\u7528\u3044\u3066\u898b\u3089\u308c\u305f\u306e\u3068\u306f\u5168\u304f\u7570\u306a\u308b\u56f3\u304c\u63cf\u304b\u308c\u308b\u3053\u3068\u3068\u306a\u3063\u305f\uff0e\u6700\u521d\u306b\uff0c\u3042\u308b\u86cb\u767d\u8cea\u306f\u529b\u306e\u5909\u5316\u306b\u5bfe\u5fdc\u3057\u3066\u529b\u306e\u4e0b\u3067\u5f3e\u6027\u53ce\u7e2e\u529b\u3092\u7d4c\u9a13\u3057\u3066\u4f38\u5c55\u3059\u308b. <strong>Figure 1B<\/strong> \u3092\u53c2\u7167\u3055\u308c\u305f\u3044\uff0e\u529b\u304c\u6e1b\u5f31\u3059\u308b\u3068\u30dd\u30ea\u30de\u30fc\u5185\u306e\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc\u304c\u5927\u5e45\u306b\u5897\u5927\u3057\uff0c\u86cb\u767d\u8cea\u304c\u6298\u308a\u305f\u305f\u307e\u308c\u308b\u3088\u3046\u306b\u3059\u308b\u306b\u9055\u3044\u306a\u3044\uff0e\u3053\u306e\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc\u3092\u9664\u53bb\u3059\u308b\u306b\u306f\u30a8\u30cd\u30eb\u30ae\u30fc\u969c\u58c1\u3092\u6a2a\u65ad\u3059\u308b\u5fc5\u8981\u304c\u3042\u308a\uff0c\u305d\u306e\u969c\u58c1\u306f\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc\u969c\u58c1\u3068\u3057\u3066\u77e5\u3089\u308c\u308b\u727d\u5f15\u529b\u306b\u3088\u308a\u4f5c\u3089\u308c\u308b (<a href=\"https:\/\/www.sciencedirect.com\/science\/article\/pii\/S0006291X10020310\" rel=\"noopener noreferrer\" target=\"_blank\">66<\/a>). \u3053\u306e\u9077\u79fb\u306f\uff0c\u30e2\u30eb\u30c6\u30f3\u30fb\u30b0\u30ed\u30d3\u30e5\u30fc\u30eb\u9077\u79fb\u3078\u306e\u6298\u308a\u305f\u305f\u307f\u3068\u3057\u3066\u77e5\u3089\u308c\u3066\u304a\u308a\uff0c\u758e\u6c34\u6027\u6b8b\u57fa\u306e\u86cb\u767d\u8cea\u30b3\u30a2\u3078\u306e\u9694\u96e2\u306b\u3088\u308a\u99c6\u52d5\u3055\u308c\uff0c\u305d\u306e\u7d50\u679c <strong>Figure 1B<\/strong> \u306b\u793a\u3059\u3088\u3046\u306b\u4e0b\u5411\u304d\u306e\u6bb5\u968e\u3068\u306a\u308b\u3068\u8003\u3048\u3089\u308c\u3066\u3044\u308b\uff0e\u914d\u7f6e\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc\u306e\u3055\u3089\u306a\u308b\u6e1b\u5c11\u306f\uff0c\u30e2\u30eb\u30c6\u30f3\u30fb\u30b0\u30ed\u30d3\u30e5\u30fc\u30eb\u304b\u3089\u81ea\u7136\u72b6\u614b\u3078\u306e\u767a\u5c55\u306e\u904e\u7a0b\u3092\u901a\u3058\u3066\u9054\u6210\u3055\u308c\u308b\uff0e\u305d\u308c\u306f\u6a5f\u68b0\u7684\u30af\u30e9\u30f3\u30d7\u306b\u304a\u3051\u308b\u6c34\u7d20\u7d50\u5408\u306e\u5f62\u6210\u3092\u901a\u3058\u3066\u306a\u3055\u308c\u308b\u3088\u3046\u3067\u3042\u308b\uff0e\u3053\u306e\u30e2\u30eb\u30c6\u30f3\u30fb\u30b0\u30ed\u30d3\u30e5\u30fc\u30eb\u306e\u81ea\u7136\u9077\u79fb\u306f\u86cb\u767d\u8cea\u672b\u7aef\u9593\u306e\u9577\u3055\u306e\u5fae\u5c0f\u5909\u5316\u3092\u5f15\u304d\u8d77\u3053\u3057\uff0c\u30e2\u30eb\u30c6\u30f3\u30fb\u30b0\u30ed\u30d3\u30e5\u30fc\u30eb\u3068\u81ea\u7136\u72b6\u614b\u3068\u306e\u9593\u3067\u5287\u7684\u306b\u6a5f\u68b0\u7684\u5b89\u5b9a\u6027\u304c\u7570\u306a\u308b\u3053\u3068\u3067\uff0c\u7570\u306a\u308b\u72b6\u614b\u304c\u691c\u51fa\u3055\u308c\u308b\u3053\u3068\u306b\u306a\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/19541635\" rel=\"noopener noreferrer\" target=\"_blank\">67<\/a>). \u5316\u5b66\u7684\u5909\u6027\u3084 phi-value analysis \u306e\u4e00\u62ec\u6cd5\u306e\u7814\u7a76\u3067\u60f3\u5b9a\u3055\u308c\u308b\u3053\u306e\u53e4\u5178\u7684\u306a\u300c\u4e8c\u72b6\u614b\u300d\u30e2\u30c7\u30eb\u306f\uff0c\u4f38\u5c55\u3055\u308c\u6298\u308a\u305f\u305f\u307e\u308c\u305f\u30e2\u30eb\u30c6\u30f3\u30fb\u30b0\u30ed\u30d3\u30e5\u30fc\u30eb\u3084\uff0c\u5358\u5206\u5b50\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u5b9f\u9a13\u3067\u89b3\u6e2c\u3055\u308c\u308b\u81ea\u7136\u72b6\u614b\u3068\u306f\u76f8\u5bb9\u308c\u306a\u3044\uff0e\u5358\u5206\u5b50\u539f\u5b50\u9593\u529b\u5206\u5149\u6cd5\u306f\uff0c\u3053\u308c\u3089\u5f93\u6765\u306e\u624b\u6cd5\u3088\u308a\u3082\uff0c\u3053\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u7d4c\u8def\u306b\u3064\u3044\u3066\u3088\u308a\u591a\u304f\u306e\u8a73\u7d30\u3092\u3082\u305f\u3089\u3057\u3066\u304f\u308c\u308b\uff0e<\/p>\n<h3>Protein folding performs mechanical work<\/h3>\n<p>\u3000Force-clamp \u539f\u5b50\u9593\u529b\u9855\u5fae\u93e1\u306e\u767a\u660e\u4ee5\u524d\u306b\u306f\uff0c\u529b\u306e\u4e0b\u3067\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u8a08\u6e2c\u306f\u975e\u5e38\u306b\u5358\u7d14\u306a\u3082\u306e\u3057\u304b\u306a\u304b\u3063\u305f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/10097099\" rel=\"noopener noreferrer\" target=\"_blank\">33<\/a>). \u3053\u308c\u3089\u306e\u5b9f\u9a13\u3067\u306f\uff0c\u516b\u3064\u306e\u8155\u306e\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u3092\u542b\u3080\u86cb\u767d\u8cea\u306e\u69cb\u6210\u304c\uff0c\u4e00\u5b9a\u901f\u5ea6\u3067\u306e AFM \u30ab\u30f3\u30c1\u30ec\u30d0\u30fc\u306e\u727d\u5f15\u306b\u3088\u308b\u6700\u521d\u306e\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3068\u306a\u3063\u305f\uff0e\u5f15\u304d\u7d9a\u3044\u3066\u30ab\u30f3\u30c1\u30ec\u30d0\u30fc\u5148\u7aef\u304c\u8a66\u6599\u8868\u9762\u304b\u3089\u306e\u529b\u3067\u3054\u304f\u77ed\u3044\u8ddd\u96e2\uff08\u6570\u5341\u30ca\u30ce\u30e1\u30fc\u30c8\u30eb\uff09\u4e0b\u304c\u308a\uff0c\u63a5\u89e6\u306a\u3057\u306b\u3054\u304f\u77ed\u6642\u9593\u305d\u3053\u306b\u3068\u3069\u307e\u308b\uff0e\u3053\u306e\u904e\u7a0b\u304c\u86cb\u767d\u8cea\u306e residual force \u306e\u6b8b\u7559\u52b9\u679c\u3067\u3042\u308b\uff0e\u30ab\u30f3\u30c1\u30ec\u30d0\u30fc\u304c\u518d\u5ea6\u727d\u5f15\u3055\u308c\u308b\u3053\u3068\u3067\uff0c\u305d\u306e\u6298\u308a\u305f\u305f\u307e\u308c\u305f\u30c9\u30e1\u30a4\u30f3\u6570\u304c\u8a08\u6e2c\u3067\u304d\u308b\uff0e\u305d\u308c\u3089\u306e\u5b9f\u9a13\u304b\u3089\u660e\u3089\u304b\u306b\u306a\u3063\u305f\u306e\u306f\uff0c\u518d\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u7387\u304c\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c9\u3057\u3066\u3044\u308b\u86cb\u767d\u8cea\u306e\u5f1b\u7de9\u306e\u7a0b\u5ea6\u306b\u5f37\u304f\u4f9d\u5b58\u3059\u308b\u3068\u3044\u3046\u3053\u3068\u3067\u3042\u308a (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/10097099\" rel=\"noopener noreferrer\" target=\"_blank\">33<\/a>), \u518d\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u7387\u306f residual force \u306b\u6025\u6fc0\u306b\u4f9d\u5b58\u3059\u308b\u3068\u306e\u30a8\u30d3\u30c7\u30f3\u30b9\u304c\u5f97\u3089\u308c\u305f\uff0e\u3057\u304b\u3057\uff0c\u305d\u308c\u3089\u521d\u671f\u306e\u5b9f\u9a13\u306b\u304a\u3044\u3066\u306f\uff0c\u86cb\u767d\u8cea\u304c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3059\u308b\u529b\u306e\u4e0b\u3067\u306f\uff0c\u76f4\u63a5\u8a08\u6e2c\u304c\u3067\u304d\u306a\u304b\u3063\u305f\uff0eForce-clamp AFM \u306e\u767a\u5c55\u306f\uff0c\u4e16\u754c\u3067\u521d\u3081\u3066\u6a5f\u68b0\u7684\u306a\u529b\u306e\u4e0b\u3067\u306e\u86cb\u767d\u8cea\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u5b8c\u5168\u3067\u92ed\u3044\u76f4\u63a5\u89b3\u5bdf\u3092\u3082\u305f\u3089\u3057\u3066\u304f\u308c\u305f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/15017000\" rel=\"noopener noreferrer\" target=\"_blank\">34<\/a>). \u30e6\u30d3\u30ad\u30c1\u30f3\u30c9\u30e1\u30a4\u30f3\u8155\u304b\u3089\u306a\u308b\u86cb\u767d\u8cea\u69cb\u6210\u3092\u7528\u3044\u3066, \u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c9\u3057\u305f\u86cb\u767d\u8cea\u3092\u727d\u5f15\u3059\u308b\u529b\u3092\u300c\u6025\u505c\u6b62\u300d\u3059\u308b\u3068\uff0c\u305d\u306e\u86cb\u767d\u8cea\u306f\u307e\u305a\u5f3e\u6027\u53ce\u7e2e\u3057\uff0c\u6b21\u3044\u3067\u9045\u5ef6\u306e\u5f8c\uff0c\u727d\u5f15\u529b\u306b\u62b5\u6297\u3057\u3066\u6a5f\u68b0\u7684\u4ed5\u4e8b\u3092\u884c\u3044\uff0c\u5b8c\u5168\u306b\u6298\u308a\u305f\u305f\u307e\u308c\u305f\u9577\u3055\u306b\u77ed\u7e2e\u3059\u308b\u3053\u3068\u3092 Fernandez \u3068 Li \u304c\u8a3c\u660e\u3057\u305f\uff0e\u529b\u306e\u6025\u505c\u6b62\u5f8c\uff0c\u7d9a\u3044\u3066\u9ad8\u3044\u529b\u306e\u30d1\u30eb\u30b9\u304c\u6298\u308a\u305f\u305f\u307e\u308c\u305f\u30c9\u30e1\u30a4\u30f3\u306e\u6307\u7d0b\u3092\u660e\u3089\u304b\u306b\u3057\uff0c\u3053\u308c\u3089\u306e\u6298\u308a\u305f\u305f\u307f\u306e\u8ecc\u8de1\u306f\uff0c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u904e\u7a0b\u304c\u771f\u5b9f\u306e\u89b3\u5bdf\u3067\u3042\u308b\u3053\u3068\u3092\u6559\u3048\u3066\u304f\u308c\u308b\uff0e\u3088\u308a\u77ed\u6642\u9593\u306e\u6025\u505c\u6b62\u3067\u306e\u4f3c\u305f\u5b9f\u9a13\u3067\u306f\u6a5f\u68b0\u7684\u306b\u8106\u5f31\u306a\u72b6\u614b\u306e\u5b58\u5728\u304c\u660e\u3089\u304b\u306b\u306a\u3063\u305f\uff0e\u305d\u3053\u3067\u306f\uff0c\u5b8c\u5168\u306b\u6298\u308a\u305f\u305f\u307e\u308c\u305f\u86cb\u767d\u8cea\u306e\u524d\u99c6\u4f53\u3068\u3057\u3066\u4fdd\u5b58\u3055\u308c\uff0c\u30e2\u30eb\u30c6\u30f3\u30fb\u30b0\u30ed\u30d3\u30e5\u30fc\u30eb\u72b6\u614b\u3092\u30b3\u30d4\u30fc\u3057\u305f\u72b6\u6cc1\u306b\u4f3c\u3066\u3044\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/19541635\" rel=\"noopener noreferrer\" target=\"_blank\">67<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/7656002\" rel=\"noopener noreferrer\" target=\"_blank\">68<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/28097712\" rel=\"noopener noreferrer\" target=\"_blank\">69<\/a>). \u3053\u308c\u306f\u529b\u306e\u4e0b\u3067\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u7269\u7406\u5b66\u3092\u660e\u3089\u304b\u306b\u3057\u305f\u3082\u306e\u3067\uff0c\u305d\u3053\u3067\u306f\u529b\u306e\u6025\u505c\u6b62\u306b\u53cd\u5fdc\u3057\u3066\u86cb\u767d\u8cea\u306f\u307e\u305a\u4f38\u9577\u3057\u305f\u72b6\u614b\u304b\u3089\u6298\u308a\u305f\u305f\u307e\u308c\u305f\u72b6\u614b\u306b\u5f3e\u6027\u53ce\u7e2e\u3057\uff0c\u305d\u306e\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc\u3092\u5927\u5e45\u306b\u5897\u52a0\u3055\u305b\u308b\uff0e\u305d\u306e\u6298\u308a\u305f\u305f\u307f\u72b6\u614b\u304b\u3089\uff0c\u86cb\u767d\u8cea\u306f\u3055\u3089\u306b\u53ce\u7e2e\u3057\u3066\u6bb5\u968e\u69d8\u5f0f\u3067\u77ed\u7e2e\u3057\uff0c\u4f4e\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc\u306e\u30e2\u30eb\u30c6\u30f3\u30fb\u30b0\u30ed\u30d3\u30e5\u30fc\u30eb\u72b6\u614b\u306b\u306a\u308b\uff0e\u3053\u306e\u30e2\u30eb\u30c6\u30f3\u30fb\u30b0\u30ed\u30d3\u30e5\u30fc\u30eb\u72b6\u614b\u306b\u304a\u3044\u3066\u86cb\u767d\u8cea\u306f\u300c\u6210\u719f\u300d\u3057\u3066\u6a5f\u68b0\u7684\u306b\u81ea\u7136\u306a\u5f62\u614b\u306b\u79fb\u884c\u3057\uff0c\u6c34\u7d20\u7d50\u5408\u306e\u30cd\u30c3\u30c8\u30ef\u30fc\u30af\u3092\u518d\u69cb\u7bc9\u3057\u3066\u6a5f\u68b0\u7684\u306b\u5b89\u5b9a\u3057\u305f\u72b6\u614b\u306b\u306a\u308b\uff0e\u3053\u306e\u30b7\u30ca\u30ea\u30aa\u306b\u304a\u3044\u3066\u306f\uff0c\u5927\u90e8\u5206\u306e\u6298\u308a\u305f\u305f\u307f\u53ce\u7e2e\u306f\u6298\u308a\u305f\u305f\u307f\u72b6\u614b\u304b\u3089\u30e2\u30eb\u30c6\u30f3\u30fb\u30b0\u30ed\u30d3\u30e5\u30fc\u30eb\u72b6\u614b\u3078\u306e\u86cb\u767d\u8cea\u306e\u52d5\u304d\u3068\u3057\u3066\u767a\u751f\u3059\u308b\u304c\uff0c\u6c34\u7d20\u7d50\u5408\u30cd\u30c3\u30c8\u30ef\u30fc\u30af\u306e\u518d\u69cb\u7bc9\u306f\u86cb\u767d\u8cea\u306e\u672b\u7aef\u9593\u306e\u8ddd\u96e2\u3092\u305d\u308c\u307b\u3069\u5909\u5316\u3055\u305b\u306a\u3044 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/19541635\" rel=\"noopener noreferrer\" target=\"_blank\">67<\/a>). \u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u6cd5\u3092\u7528\u3044\u3066\uff0cI10 \u3068 I91 \u306e\u6298\u308a\u305f\u305f\u307f\u8ecc\u9053\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304a\u3088\u3073\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u30b9\u30c6\u30c3\u30d7\u30b5\u30a4\u30ba\u3092 30 pN \u672a\u6e80\u306e\u529b\u3067\u8a08\u6e2c\u3057\u3066\u307f\u308b\u3068 (<strong>Figure 2B<\/strong>), \u30e2\u30eb\u30c6\u30f3\u30fb\u30b0\u30ed\u30d3\u30e5\u30fc\u30eb\u72b6\u614b\u306e\u3082\u3063\u3068\u5b8c\u5168\u306a\u6839\u62e0\u304c\u5f97\u3089\u308c\u308b (<a href=\"https:\/\/www.cell.com\/biophysj\/abstract\/S0006-3495(15)03306-8\" rel=\"noopener noreferrer\" target=\"_blank\">70<\/a>). \u8a73\u7d30\u306b\u8abf\u3079\u3089\u308c\u305f\u306e\u306f\u3053\u308c\u3089\u305f\u3063\u305f\u4e8c\u3064\u306e\u30c9\u30e1\u30a4\u30f3\u306b\u904e\u304e\u306a\u3044\u304c, 4-10 pN \u306e\u540c\u3058\u7bc4\u56f2\u306e\u529b\u306e\u4e2d\u3067\uff0c\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u4e2d\u306e\u5287\u7684\u306a\u6a5f\u68b0\u7684\u5b89\u5b9a\u6027\u306e\u9055\u3044\u306b\u3082\u304b\u304b\u308f\u3089\u305a\uff0c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u9077\u79fb\u3092\u7d4c\u9a13\u3057\u3066\u3044\u308b\u3088\u3046\u306b\u898b\u3048\u308b\uff0e\u304a\u305d\u3089\u304f\u3053\u308c\u306f\u5404 Ig \u30c9\u30e1\u30a4\u30f3\u304c\u540c\u6570\u306e\u6b8b\u57fa\u3092\u6709\u3057\u3066\u304a\u308a\uff0c\u30e2\u30eb\u30c6\u30f3\u30fb\u30b0\u30ed\u30d3\u30e5\u30fc\u30eb\u9077\u79fb\u3078\u306e\u6298\u308a\u305f\u305f\u307f\u306e\u9593\uff0c\u5927\u307e\u304b\u306b\u540c\u91cf\u306e\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc\u3092\u653e\u96fb\u3059\u308b\u304b\u3089\u306b\u9055\u3044\u306a\u3044\uff0e\u3057\u304b\u3057\uff0c\u69cb\u6210\u30eb\u30fc\u30d7\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/18472109\" rel=\"noopener noreferrer\" target=\"_blank\">71<\/a>) \u3084\u70b9\u7a81\u7136\u5909\u7570\u304c\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u7d4c\u8def\u306b\u5f71\u97ff\u3059\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/11101892\" rel=\"noopener noreferrer\" target=\"_blank\">72<\/a>) \u3053\u3068\u304c\u660e\u3089\u304b\u306b\u306a\u3063\u305f\u305f\u3081, Ig \u30c9\u30e1\u30a4\u30f3\u914d\u5217\u306e\u591a\u69d8\u6027\u304c\u751f\u7406\u7684\u7bc4\u56f2\u5185\u3067\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u7387\u306e\u9593\u3067\u306e\u968e\u5c64\u3092\u5f62\u6210\u3059\u308b\u7d50\u679c\u3068\u306a\u3063\u305f\u306e\u3067\u3042\u308d\u3046\uff0e\u3053\u308c\u306f\u5b9f\u9a13\u7684\u306b\u691c\u8a3c\u3055\u308c\u306a\u3051\u308c\u3070\u306a\u3089\u306a\u3044\u3053\u3068\u3060\u304c\uff0c\u3053\u306e\u70b9\u306b\u304a\u3044\u3066\u6839\u62e0\u306e\u793a\u5506\u3059\u308b\u3068\u3053\u308d\u306f\uff0c\u7b4b\u53ce\u7e2e\u6642\u306b\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u306f\u72ed\u3044\u529b\u306e\u7bc4\u56f2\u3067\u6298\u308a\u305f\u305f\u307e\u308c\u308b\u3068\u3044\u3046\u3053\u3068\u3067\u3042\u308b\uff0e\u3053\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53ce\u7e2e\u304c AFM \u3067\u89b3\u5bdf\u3055\u308c\u308b\u3068\uff0c\u6301\u7d9a\u7684\u304b\u3064\u307b\u3068\u3093\u3069\u7a00\u306b\u6bb5\u968e\u3092\u793a\u3059\u3053\u3068\u304c\u5206\u304b\u3063\u305f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/15017000\" rel=\"noopener noreferrer\" target=\"_blank\">34<\/a>). \u5bfe\u7167\u7684\u306b\uff0c\u540c\u3058\u5b9f\u9a13\u3067\u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u6cd5\u3092\u7528\u3044\u308b\u3068\uff0c\u6700\u521d\u306e\u5f3e\u6027\u53ce\u7e2e\u5f8c\uff0c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53ce\u7e2e\u6642\u306b\u660e\u78ba\u306a\u6bb5\u968e\uff08\u7684\u53ce\u7e2e\uff09\u3092\u898b\u305b\u305f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/26854230\" rel=\"noopener noreferrer\" target=\"_blank\">10<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/25726700\" rel=\"noopener noreferrer\" target=\"_blank\">43<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/27409974\" rel=\"noopener noreferrer\" target=\"_blank\">49<\/a>). AFM \u3068\u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u6cd5\u3067\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u8ecc\u9053\u306e\u5f62\u304c\u7570\u306a\u308b\u7406\u7531\u306f\uff0c\u3059\u3079\u3066\u306e\u5358\u5206\u5b50\u539f\u5b50\u9593\u529b\u5206\u5149\u6cd5\u306e\u5b9f\u9a13\u306b\u304a\u3044\u3066\uff0c\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u306e\u529b\u5b66\u304c\u30c6\u30b6\u30fc\u4f9d\u5b58\u6027\u3067\u3042\u308b\u3068\u3044\u3046\u3053\u3068\u3067\u3042\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC3437906\/\" rel=\"noopener noreferrer\" target=\"_blank\">73<\/a>). \u305d\u306e\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u306f\u63a2\u91dd\u304c\u63a5\u89e6\u3059\u308b\u307b\u3069\u901f\u304f\u52d5\u304f\u3053\u3068\u304c\u3067\u304d\u308b\u306e\u307f\u3067\uff0c\u727d\u5f15\u3059\u308b\u65b9\u5411\u306e\u9396\u306e\u529b\u5b66\u306f\uff0c\u63a2\u91dd\u3068\u540c\u3058\u6709\u52b9\u306a\u62e1\u6563\u4fc2\u6570\u3092\u6709\u3059\u308b\uff0e\u4e09\u89d2\u5f62\u306e\u30ab\u30f3\u30c1\u30ec\u30d0\u30fc\u306f\u901a\u5e38\u5358\u5206\u5b50\u306e\u5b9f\u9a13\u3067\u7528\u3044\u3089\u308c\uff0c\u9577\u3055\u3068\u5e45\u304c\u305d\u308c\u305e\u308c 310 &micro;m \u304a\u3088\u3073 20 &micro;m \u3067\u3042\u308a\uff0c\u62e1\u6563\u4fc2\u6570\u306f 1,200 nm<sup>2<\/sup> \u3068\u6e2c\u5b9a\u3055\u308c\u3066\u3044\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC3437906\/\" rel=\"noopener noreferrer\" target=\"_blank\">73<\/a>). \u30ab\u30f3\u30c1\u30ec\u30d0\u30fc\u306e\u62e1\u6563\u4fc2\u6570\u3092\u7528\u3044\u305f\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u30d6\u30e9\u30a6\u30f3\u904b\u52d5\u529b\u5b66\u30b7\u30df\u30e5\u30ec\u30fc\u30b7\u30e7\u30f3\u306f\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u8ecc\u9053\u306e\u518d\u73fe\u6027\u3092\u53ef\u80fd\u3068\u3057, force-clamp AFM \u3092\u7528\u3044\u305f\u3082\u306e\u3068\u8fd1\u4f3c\u3057\u3066\u3044\u305f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/21050839\" rel=\"noopener noreferrer\" target=\"_blank\">66<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC2877348\/\" rel=\"noopener noreferrer\" target=\"_blank\">74<\/a>). \u5bfe\u7167\u7684\u306b\uff0c\u76f4\u5f84 2.8 &micro;m \u306e\u81ea\u7531\u6d6e\u52d5\u7403\u4f53\u7c92\u5b50\u306f 175,000 nm<sup>2<\/sup> \u307e\u3067\u306e\u62e1\u6563\u4fc2\u6570\u3092\u6301\u3064\uff0e\u78c1\u6c17\u30d3\u30fc\u30ba\u304c\u8a66\u6599\u8868\u9762\u3068\u8fd1\u63a5\u3057\u3066\u3044\u3066\u305d\u306e\u5024\u3092\u82e5\u5e72\u6e1b\u5f31\u3059\u308b\u306b\u3082\u304b\u304b\u308f\u3089\u305a\u3067\u3042\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/27369870\" rel=\"noopener noreferrer\" target=\"_blank\">75<\/a>,<a href=\"https:\/\/www.sciencedirect.com\/science\/article\/pii\/0009250961800353\" rel=\"noopener noreferrer\" target=\"_blank\">76<\/a>). \u3053\u308c\u306f\u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u3067\u898b\u3089\u308c\u308b\u6298\u308a\u305f\u305f\u307f\u8ecc\u9053\u306e\u9055\u3044\u306b\u5bc4\u4e0e\u3057\u3066\u3044\u308b\uff0e\u3053\u3053\u3067\u6ce8\u610f\u304c\u5fc5\u8981\u3067\u3042\u308b\uff0e\u7f8e\u3057\u3044\u6bb5\u968e\u7684\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u8ecc\u9053\u304c\u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u6cd5\u3067\u89b3\u6e2c\u3055\u308c\uff0c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u30a8\u30cd\u30eb\u30ae\u30fc\u306e\u98a8\u666f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/25796331\" rel=\"noopener noreferrer\" target=\"_blank\">77<\/a>) \u304a\u3088\u3073\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u4ed5\u4e8b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/26854230\" rel=\"noopener noreferrer\" target=\"_blank\">10<\/a>) \u306e\u753b\u50cf\u304c\u3088\u308a\u826f\u3044\u89e3\u50cf\u5ea6\u3067\u5f97\u3089\u308c\u305f\u304b\u3089\u3068\u8a00\u3063\u3066\uff0c\u305d\u308c\u304c\u751f\u4f53\u5185\u3067\u306e\u30bf\u30a4\u30c1\u30f3\u86cb\u767d\u8cea\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u771f\u5b9f\u306e\u62e1\u6563\u306e\u3075\u308b\u307e\u3044\u3092\u8868\u3057\u3066\u3044\u308b\u308f\u3051\u3067\u306f\u306a\u3044\uff0e\u30bf\u30a4\u30c1\u30f3\u304c Z \u76e4\u3068 M-line \u3068\u3044\u3046\u5de8\u5927\u306a\u5206\u5b50\u69cb\u9020\u306b\u4fc2\u7559\u3055\u308c\u3066\u3044\u308b\u3053\u3068\u304b\u3089\uff0c\u307e\u305f\uff0c\u30b7\u30e3\u30da\u30ed\u30f3\u3092\u542b\u3080\u6df7\u96d1\u3057\u305f\u74b0\u5883\u3067\u52d5\u4f5c\u3059\u308b\u3053\u3068\u304b\u3089\uff0c\u751f\u4f53\u5185\u3067\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53ce\u7e2e\u306f AFM \u3067\u89b3\u6e2c\u3055\u308c\u308b\u3088\u3046\u306a\u6ed1\u3089\u304b\u306a\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53ce\u7e2e\u306e\u3088\u3046\u306b\u3082\u3063\u3068\u62e1\u6563\u306e\u5236\u7d04\u3055\u308c\u305f\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u3075\u308b\u307e\u3044\u306b\u4f3c\u3066\u3044\u308b\u53ef\u80fd\u6027\u304c\u3042\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/15017000\" rel=\"noopener noreferrer\" target=\"_blank\">34<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC2877348\/\" rel=\"noopener noreferrer\" target=\"_blank\">74<\/a>). <\/p>\n<p>\u3000\u6211\u3005\u306f\u63d0\u6848\u3059\u308b\uff0e\u6b63\u5e38\u306a\u7b4b\u6d3b\u52d5\u6642\u306e\u30bf\u30a4\u30c1\u30f3\u3078\u306e\u8ca0\u8377\u306e\u751f\u7406\u7684\u306a\u529b\u306e\u7bc4\u56f2\u3068\u306f\uff0c\u6298\u308a\u305f\u305f\u307f\u3068\u4f38\u5c55\u306e\u5e73\u8861\u72b6\u614b\u3067 Ig \u30c9\u30e1\u30a4\u30f3\u304c\u5b58\u5728\u3059\u308b\u7bc4\u56f2\u3092\u8d85\u3048\u308b\u529b\u306e\u7bc4\u56f2\u3067\u3042\u308b\u3068\uff0e\u3053\u308c\u306f\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u78ba\u7387\u3092\u901a\u3058\u3066\u5b9a\u91cf\u3055\u308c\u308b\u53ef\u80fd\u6027\u304c\u3042\u308a\uff0c\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u304c\u4efb\u610f\u306e\u529b\u3067\u6298\u308a\u305f\u305f\u307e\u308c\u308b\u6642\u9593\u306e\u5272\u5408\u3092\u8a08\u6e2c\u3059\u308b\u3082\u306e\u3067\u3042\u308b\uff0e\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3 I10 \u304a\u3088\u3073 I91 \u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u78ba\u7387\u3092\u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u306e\u539f\u5b50\u9593\u529b\u5206\u5149\u6cd5\u3067\u8a08\u6e2c\u3057, <strong>Figure 2C<\/strong> \uff08\u9752\u4e38\uff09\u306b\u8981\u7d04\u3057\u3066\u3042\u308a\uff0c\u30b7\u30b0\u30e2\u30a4\u30c9\u95a2\u6570\u306b\u9069\u5408\u3057\u3066\u304a\u308a\uff0c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53cd\u5fdc\u306e midpoint \u3068 steepness \u3092\u8a18\u8ff0\u3057\u3066\u3042\u308b (\u9752\u7dda, <strong>Figure 2C<\/strong>, <strong>Figure 2D<\/strong>). \u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u304c 4 pN \u4ee5\u4e0b\u306e\u529b\u3067\u5b8c\u5168\u306b\u6298\u308a\u305f\u305f\u307e\u308c, 10 pN \u3088\u308a\u5927\u304d\u3044\u529b\u3067\u306f\u6298\u308a\u305f\u305f\u307e\u308c\u306a\u3044\u3053\u3068\u306f\u660e\u3089\u304b\u3067\u3042\u308b\uff0e\u73fe\u5728\u3067\u306f\u3053\u306e\u529b\u306e\u7bc4\u56f2\u306f 15 pN \u307e\u3067\u62e1\u5927\u3057\u3066\u3044\u308b\u3068\u306e\u6839\u62e0\u304c\u3042\u308b\uff0e\u30bf\u30a4\u30c1\u30f3\u306e\u3053\u306e I-band \u9818\u57df\u306f\u30b7\u30b9\u30c6\u30a4\u30f3\u304c\u8c4a\u5bcc\u3067 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/24630725\" rel=\"noopener noreferrer\" target=\"_blank\">78<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/14596801\" rel=\"noopener noreferrer\" target=\"_blank\">79<\/a>), \u3053\u308c\u3089\u306e\u591a\u304f\u306e\u6b8b\u57fa\u306f\u30b8\u30b9\u30eb\u30d5\u30a3\u30c9\u7d50\u5408\u3092\u5f62\u6210\u3057\uff0c\u30dd\u30ea\u30de\u30fc\u529b\u5b66\u3092\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u72b6\u614b\u306b\u5909\u3048\u308b\uff0e\u305d\u306e\u7d50\u679c\uff0c\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u304c 15 pN \u307e\u3067\u306e\u3088\u308a\u9ad8\u3044\u529b\u3067\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c9\u3057\u3084\u3059\u304f\u306a\u308a\uff0c\u6298\u308a\u305f\u305f\u307f\u72b6\u614b\u306b\u304a\u3051\u308b\u6a5f\u68b0\u7684\u5b89\u5b9a\u6027\u3092\u3088\u308a\u5931\u308f\u305b\u3084\u3059\u304f\u3059\u308b\u52b9\u679c\u304c\u3042\u308b (<a href=\"https:\/\/www.cell.com\/biophysj\/abstract\/S0006-3495(16)33473-7?code=cell-site\" rel=\"noopener noreferrer\" target=\"_blank\">80<\/a>). <\/p>\n<p>\u3000Force-clamp \u5206\u5149\u6cd5\u304c\u4ed8\u52a0\u7684\u306b\u53ef\u80fd\u306b\u3057\u305f\u5358\u7d14\u306a\u5b9a\u91cf\u5316\u3068\u3057\u3066\uff0c\u30bf\u30a4\u30c1\u30f3\u30c9\u30e1\u30a4\u30f3\u304c\u9069\u7528\u3055\u308c\u305f\u529b\u306b\u6297\u3057\u3066\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3057\u3066\u5b9f\u884c\u3059\u308b\u53ce\u7e2e\u306e\u4ed5\u4e8b\u91cf\u304c\u3042\u308b\uff0e\u3044\u304b\u306a\u308b\u529b\u3092\u4e0e\u3048\u3066\u3082\uff0c\u30b7\u30b9\u30c6\u30e0\u4e0a\u307e\u305f\u306f\u30b7\u30b9\u30c6\u30e0\u306b\u3088\u308a\u306a\u3055\u308c\u308b\u53ce\u7e2e\u306e\u4ed5\u4e8b\u91cf\u306f\uff0c\u529b\u3068\u30b9\u30c6\u30c3\u30d7\u30b5\u30a4\u30ba\u306e\u7a4d\u3067\u3042\u308b. I10 \u304a\u3088\u3073 I91 \u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306b\u3088\u308a\u306a\u3055\u308c\u308b\u4ed5\u4e8b\u306f <strong>Figure 2D<\/strong> \u306e\u8d64\u3044\u66f2\u7dda\u3067\u793a\u3057\u305f\u304c\uff0c\u529b\u306b\u5bfe\u5fdc\u3059\u308b\u81ea\u7531\u7d50\u5408\u9396\u66f2\u7dda\u306e\u7a4d\u5206\u3067\u3042\u308b\uff0e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u78ba\u7387\u3067\u306a\u3055\u308c\u305f\u53ce\u7e2e\u4ed5\u4e8b\u3092\u4ee3\u8868\u3059\u308b\u4e57\u7b97\u66f2\u7dda\u306f\uff0c\u4e0e\u3048\u3089\u308c\u305f\u529b\u3067\u306e\u86cb\u767d\u8cea\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306b\u3088\u308b\u30a8\u30cd\u30eb\u30ae\u30fc\u56de\u5fa9\u306e\u671f\u5f85\u5024\u3092\u4e0e\u3048\u3066\u304f\u308c\u308b (\u9ed2\u306e\u66f2\u7dda, <strong>Figure 2D<\/strong>). \u3053\u306e\u5206\u5e03\u304c\u793a\u3059\u3068\u3053\u308d\u306f\uff0c\u30bf\u30a4\u30c1\u30f3 Ig \u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304c, 5.7 pN \u306e\u8ca0\u8377\u306b\u5bfe\u3057\u3066 47 zJ \u3068\u671f\u5f85\u3055\u308c\u308b\u6700\u5927\u306e\u4ed5\u4e8b\u3092\u767a\u751f\u3055\u305b\u308b\u3068\u3044\u3046\u3053\u3068\u3067\u3042\u308b\uff0e10 pN \u306b\u8fd1\u3044\u9ad8\u3044\u8ca0\u8377\u3067\u306f\uff0c\u30a8\u30cd\u30eb\u30ae\u30fc\u56de\u5fa9\u306f\u6700\u5927\u5316\u3059\u308b\u304c\uff0c\u30c9\u30e1\u30a4\u30f3\u306e\u6298\u308a\u305f\u305f\u307f\u983b\u5ea6\u306f\u6e1b\u5c11\u3059\u308b\uff0e\u4ee3\u308f\u308a\u306b\uff0c\u8efd\u3044\u8ca0\u8377\u306b\u304a\u3044\u3066\u306f\u30a8\u30cd\u30eb\u30ae\u30fc\u56de\u5fa9\u306f\u6700\u5c0f\u5316\u3059\u308b\uff0e\u3068\u3044\u3046\u306e\u306f\uff0c\u86cb\u767d\u8cea\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u30b9\u30c6\u30c3\u30d7\u30b5\u30a4\u30ba\u304c\u5c0f\u3055\u304f\uff0c\u307b\u3068\u3093\u3069\u306e\u30a8\u30cd\u30eb\u30ae\u30fc\u304c\u5f3e\u6027\u53ce\u7e2e\u3067\u6d88\u8cbb\u3055\u308c\u3066\u3057\u307e\u3046\u304b\u3089\u3067\u3042\u308b\uff0e\u3086\u3048\u306b\u6211\u3005\u306f\uff0c\u751f\u4f53\u5185\u3067\u5358\u4e00\u306e\u30bf\u30a4\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306b\u304b\u304b\u308b\u529b\u3092\u5b9a\u7fa9\u3067\u304d\u308c\u3070\uff0c\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3092\u901a\u3058\u3066\u84c4\u7a4d\u3055\u308c\u89e3\u653e\u3055\u308c\u308b\u30a8\u30cd\u30eb\u30ae\u30fc\u91cf\u3092\u4e88\u6e2c\u3067\u304d\u308b\uff0e<\/p>\n<h3>How the three filaments work together<\/h3>\n<p>\u3000\u5358\u5206\u5b50\u306e\u5b9f\u9a13\u306f\uff0c\u86cb\u767d\u8cea\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u30a8\u30cd\u30eb\u30ae\u30fc\u7406\u8ad6\u306b\u3064\u3044\u3066\u7406\u89e3\u3059\u308b\u306e\u307f\u306a\u3089\u305a\uff0c\u5358\u4e00\u306e\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306b\u3088\u308b\u529b\u306e\u767a\u751f\u306e\u57fa\u790e\u306b\u3064\u3044\u3066\u7406\u89e3\u3059\u308b\u305f\u3081\u306e\u5f37\u56fa\u306a\u57fa\u76e4\u3092\u63d0\u4f9b\u3057\u3066\u304d\u305f\uff0e\u3053\u308c\u306f\u30bf\u30a4\u30c1\u30f3 Ig \u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u53ce\u7e2e\u7279\u6027\u3068\u30df\u30aa\u30b7\u30f3\u30d1\u30ef\u30fc\u30b9\u30c8\u30ed\u30fc\u30af\u3068\u306e\u9593\u306e\u516c\u5e73\u306a\u6bd4\u8f03\u3092\u610f\u5473\u3057\u3066\u3044\u308b\uff0e\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306e\u72ec\u594f\u307e\u305f\u306f\u5408\u594f\u306b\u304a\u3051\u308b\u5358\u5206\u5b50\u539f\u5b50\u9593\u529b\u5206\u5149\u6cd5\u306f, ATP \u3067\u99c6\u52d5\u3055\u308c\u308b\u30e2\u30fc\u30bf\u30fc\u304c\u5b9f\u884c\u3067\u304d\u308b\u5229\u7528\u53ef\u80fd\u306a\u4ed5\u4e8b\u91cf\u306b\u56fa\u6709\u306e\u5236\u9650\u304c\u3042\u308b\u3053\u3068\u3092\u660e\u3089\u304b\u306b\u3057\u305f\uff0e\u4eee\u306b\u5341\u5206\u306b\u5927\u304d\u306a\u8ca0\u8377\u304c\u30e2\u30fc\u30bf\u30fc\u306b\u304b\u304b\u308b\u3068\uff0c\u305d\u306e\u30e2\u30fc\u30bf\u30fc\u51fa\u529b\u306f\u30bc\u30ed\u306b\u4f4e\u4e0b\u3057\u3066\u3057\u307e\u3046\uff0e\u81ea\u8ee2\u8eca\u3067\u5742\u9053\u3092\u767b\u308b\u6642\uff0c\u7269\u7406\u5b66\u306e\u3053\u306e\u4e00\u304b\u3051\u3089\u3092\u75db\u611f\u3055\u305b\u3089\u308c\u308b\uff0e\u521d\u671f\u306e\u7b4b\u8089\u306e\u30d1\u30a4\u30aa\u30cb\u30a2\u3067\u3042\u308b A. V. Hill \u306f\u3053\u306e\u52b9\u679c\u3092\u6349\u3048\u3066\u5f7c\u306e\u6709\u540d\u306a\u300cHill \u306e\u516c\u5f0f\u300d\u3068\u3057\u305f\uff0e\u3064\u307e\u308a\uff0c\u53ce\u7e2e\u901f\u5ea6\u3068\u8ca0\u8377\u3068\u306e\u9593\u306e\u53cc\u66f2\u7dda\u95a2\u4fc2\u3092\u8a18\u8ff0\u3057\u305f\u3082\u306e\u3067\u3042\u308b (<a href=\"http:\/\/rspb.royalsocietypublishing.org\/content\/126\/843\/136\" rel=\"noopener noreferrer\" target=\"_blank\">81<\/a>). \u8a66\u9a13\u7ba1\u5185\u306e\u5b9f\u9a13\u304c\u660e\u3089\u304b\u306b\u3057\u305f\u3068\u3053\u308d\u3067\u306f\uff0c\u514d\u8377\u72b6\u614b\u306e\u7b4b\u8089\u306e\u901f\u5ea6\u306f\u6975\u3081\u3066\u901f\u3044\u304c\uff0c\u5c11\u3057\u3067\u3082\u8ca0\u8377\u3092\u304b\u3051\u308b\u3068, II \u578b\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306e\u6ed1\u8d70\u901f\u5ea6\u304c\u6025\u6fc0\u306b\u4f4e\u4e0b\u3057\u3066\u3057\u307e\u3046\uff0e\u9aa8\u683c\u7b4b\u306e\u30df\u30aa\u30b7\u30f3\u306e\u6a5f\u5e8f\u3092\u63a2\u7d22\u3059\u308b\u7cbe\u5bc6\u306a\u5b9f\u9a13\u3067\u660e\u3089\u304b\u306b\u306a\u3063\u305f\u306e\u306f\uff0c\u7b49\u5c3a\u6027\u72b6\u614b\u4e0b\u3067\u30e2\u30fc\u30bf\u30fc 1 \u500b\u304c\u7523\u751f\u3059\u308b\u529b\u306e\u5e73\u5747\u5024\u306f 3.4 pN \u3067\u3042\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/8139653\" rel=\"noopener noreferrer\" target=\"_blank\">26<\/a>). \u307e\u308c\u306b\uff0c\u3053\u306e\u30df\u30aa\u30b7\u30f3\u304c\u6301\u7d9a\u7684\u306b\u767a\u751f\u3055\u305b\u308b\u529b\u304c 6-7 pN \u306e\u7bc4\u56f2\u306b\u9077\u79fb\u3059\u308b\u3053\u3068\u304c\u3042\u308b\uff0e\u3055\u3089\u306b\u6700\u8fd1\u3067\u306f\uff0c\u5358\u5206\u5b50\u306e\u529b-\u901f\u5ea6\u95a2\u4fc2\u3092\u5b9a\u7fa9\u3059\u308b\u3053\u3068\u3092\u76ee\u7684\u3068\u3057\u3066\uff0c\u540c\u3058\u5b9f\u9a13\u304c 8 \u500b\u307e\u3067\u306e\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u3092\u675f\u306b\u3057\u3066\u5b9f\u65bd\u3055\u308c\u305f (<strong>Figure 3A<\/strong>, \u7dd1\u8272\u66f2\u7dda) (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/16169988\" rel=\"noopener noreferrer\" target=\"_blank\">82<\/a>). \u3053\u306e\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306f\u3084\u306f\u308a 6.0 pN \u306e\u8ca0\u8377\u3092\u9069\u7528\u3059\u308b\u3068\u30a2\u30af\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e\u5909\u4f4d\u304c\u6b63\u5473\u30bc\u30ed\u306b\u306a\u3063\u3066\u3057\u307e\u3044\uff0c\u3053\u306e\u5206\u5b50\u30de\u30b7\u30f3\u306e\u5b9f\u884c\u306b\u306f\u4e0a\u9650\u5024\u304c\u5b58\u5728\u3059\u308b\u3053\u3068\u3092\u518d\u78ba\u8a8d\u3057\uff0c\u305d\u308c\u306f\u9ad8\u89e3\u50cf\u5ea6\u5149\u30d4\u30f3\u30bb\u30c3\u30c8\u6cd5\u3067\u3082\u518d\u78ba\u8a8d\u3055\u308c\u305f (<a href=\"https:\/\/www.nature.com\/articles\/ncomms16036\" rel=\"noopener noreferrer\" target=\"_blank\">29<\/a>). \u4ed6\u306b\u30ad\u30cd\u30b7\u30f3, V \u578b\u30df\u30aa\u30b7\u30f3\uff0c\u30c0\u30a4\u30cb\u30f3\u306e\u5931\u901f\u3059\u308b\u529b\u306f\u3059\u3079\u3066 6-7 pN \u306e\u7bc4\u56f2\u306b\u53ce\u307e\u3063\u3066\u3044\u308b\u3053\u3068\u304c\u793a\u3055\u308c\u3066\u3044\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/10408448\" rel=\"noopener noreferrer\" target=\"_blank\">83<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC1305667\/\" rel=\"noopener noreferrer\" target=\"_blank\">84<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/18045537\" rel=\"noopener noreferrer\" target=\"_blank\">85<\/a>). \u3053\u308c\u3089\u306e\u5b9f\u9a13\u306f force clamp \u6cd5\u3092\u7528\u3044\u3066\u5909\u63db\u5668\u306e\u5fdc\u7b54\u3092\u6e1b\u5f31\u3055\u305b\uff0c\u5206\u5b50\u30e2\u30fc\u30bf\u30fc\u306e\u52d5\u529b\u767a\u751f\u3092\u6700\u5927\u5316\u3057\u3066\u3044\u308b\uff0e\u3053\u308c\u3089\u306e\u30c7\u30fc\u30bf\u304b\u3089\u8ad6\u7406\u7684\u306b\u5c0e\u304b\u308c\u308b\u7d50\u679c\u3068\u306f, 6 pN \u306e\u529b\u3067\u306f II \u578b\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306f\u30a2\u30af\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306b\u53ce\u7e2e\u306e\u4ed5\u4e8b\u3092\u767a\u751f\u3055\u305b\u308b\u3053\u3068\u304c\u3067\u304d\u306a\u3044\u3068\u3044\u3046\u3053\u3068\u3067\u3042\u308b\uff0e\u5bfe\u7167\u7684\u306b\uff0c\u30bf\u30a4\u30c1\u30f3\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304c 6 pN \u306e\u8ca0\u8377\u306b\u304a\u3044\u3066\u30d4\u30fc\u30af\u30d1\u30ef\u30fc\u3092\u63d0\u4f9b\u3059\u308b\u3053\u3068\u3092\u6211\u3005\u306f\u793a\u3057\u305f\u304c\uff0c\u305d\u3053\u3067\u306f II \u578b\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306f\u5931\u901f\u3057\u3066\u3044\u305f (<strong>Figure 2D<\/strong>, <strong>Figure 3A<\/strong>) (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/26854230\" rel=\"noopener noreferrer\" target=\"_blank\">10<\/a>). <\/p>\n<figure id=\"attachment_3598\" aria-describedby=\"caption-attachment-3598\" style=\"width: 525px\" class=\"wp-caption aligncenter\"><a href=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure3.jpg\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure3-756x1024.jpg\" alt=\"\" width=\"525\" height=\"711\" class=\"size-large wp-image-3598\" srcset=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure3-756x1024.jpg 756w, https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure3-222x300.jpg 222w, https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure3-768x1040.jpg 768w, https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure3.jpg 1084w\" sizes=\"auto, (max-width: 525px) 100vw, 525px\" \/><\/a><figcaption id=\"caption-attachment-3598\" class=\"wp-caption-text\"><strong>Figure 3<\/strong><br \/><strong>\u30bf\u30a4\u30c1\u30f3\u306f\uff0c\u7b4b\u8089\u304c\u30df\u30aa\u30b7\u30f3\u306e\u5931\u901f\u3059\u308b\u529b\u3092\u8d85\u3048\u308b\u8ca0\u8377\u3067\u50cd\u304f\u3053\u3068\u3092\u53ef\u80fd\u306b\u3059\u308b<\/strong><br \/><strong>(A)<\/strong> \u5358\u5206\u5b50\u7814\u7a76\u304b\u3089\u306e\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u3068\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u306e\u30a8\u30cd\u30eb\u30ae\u30fc\u8ad6\u306e\u6bd4\u8f03: 8 \u500b\u307e\u3067\u306e\u30df\u30aa\u30b7\u30f3\u982d\u90e8\u304c\u5149\u30c8\u30e9\u30c3\u30d7\u4e2d\u306e\u30a2\u30af\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3092\u63a8\u9032\u3057\u305f\u5b9f\u9a13\u304b\u3089\u306e\u30c7\u30fc\u30bf\u306f\uff0c\u30df\u30aa\u30b7\u30f3\u304c 6 pN \u306e\u8ca0\u8377\u306b\u5bfe\u3057\u3066\u5931\u901f\u3057\u3066\u3044\u308b\u3053\u3068\u3092\u793a\u3057\u3066\u3044\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/16169988\" rel=\"noopener noreferrer\" target=\"_blank\">82<\/a>). \u305d\u306e\u30c7\u30fc\u30bf\u306f A.V.Hill \u306e\u516c\u5f0f\uff08\u7dd1\u8272\u306e\u66f2\u7dda\uff09\u306b\u5408\u81f4\u3059\u308b\uff0e\u3042\u308b\u3044\u306f\uff0c\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u306f 5.7 pN \u307e\u3067\uff08\u9752\u8272\u66f2\u7dda\u306e\u30d4\u30fc\u30af\uff09\u306e\u529b\u3067 46 zJ \u306e\u4ed5\u4e8b\u306e\u6700\u9069\u306a\u4f9b\u7d66\u3092\u9054\u6210\u3059\u308b\uff0e\u3057\u305f\u304c\u3063\u3066, 6-12 pN \uff08\u7070\u8272\u306e\u9670\u5f71\u90e8\u5206\uff09\u306e\u8ca0\u8377\u3067\u306f\uff0c\u30df\u30aa\u30b7\u30f3\u306e\u51fa\u529b\u304c\u30de\u30a4\u30ca\u30b9\u3067\u3042\u308b\u306e\u306b\u5bfe\u3057\u3066\uff0c\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306f\u6709\u7528\u306a\u53ce\u7e2e\u306e\u4ed5\u4e8b\u3092\u767a\u751f\u3055\u305b\u308b\u3053\u3068\u304c\u3067\u304d\u308b. <strong>(B)<\/strong> \u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306b\u3088\u3063\u3066\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u304c\u524d\u9032\u3059\u308b: (1) \u7b4b\u8089\u304c\u6d3b\u6027\u5316\u3055\u308c\u308b\u524d\u306b\u30bf\u30a4\u30c1\u30f3\u306f 12 pN \u306e\u53d7\u52d5\u7684\u8ca0\u8377\u5168\u4f53\u3092\u8ca0\u62c5\u3057\uff0c\u305d\u306e\u7d50\u679c, Ig \u30c9\u30e1\u30a4\u30f3\u306e\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304c\u8d77\u3053\u308b\uff0e (2) \u30ab\u30eb\u30b7\u30a6\u30e0\u304c\u653e\u51fa\u3055\u308c\u308b\u3068\uff0c\u30a2\u30af\u30c8\u30df\u30aa\u30b7\u30f3\u67b6\u6a4b\u304c\u5f62\u6210\u3055\u308c\u308b\u304c\uff0c\u305d\u306e\u30e2\u30fc\u30bf\u30fc\u306f\u5931\u901f\u3057\u3066\u3044\u308b\u305f\u3081, 6 pN \u306e\u529b\u3057\u304b\u767a\u751f\u3055\u305b\u308b\u3053\u3068\u304c\u3067\u304d\u306a\u3044\uff0e\u3057\u304b\u3057\uff0c\u30e2\u30fc\u30bf\u30fc\u306b\u3088\u3063\u3066\u767a\u751f\u3057\u305f\u529b\u306f\u30bf\u30a4\u30c1\u30f3\u30c9\u30e1\u30a4\u30f3\u304c\u53d7\u3051\u308b\u8ca0\u8377\u3092 6 pN \u306b\u6e1b\u5c11\u3055\u305b\uff0c\u7d50\u679c\u3068\u3057\u3066 10 nm \u307e\u3067\u306e\u518d\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u30b9\u30c6\u30c3\u30d7\u3092\u3082\u305f\u3089\u3059. (3) \u86cb\u767d\u8cea\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u30d7\u30ed\u30bb\u30b9\u306f\u30bf\u30a4\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3092\u77ed\u7e2e\u3057\uff0c\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306e\u8ca0\u62c5\u3092\u8efd\u6e1b\u3057\u3066\u30d1\u30ef\u30fc\u30b9\u30c8\u30ed\u30fc\u30af\u306e\u5b8c\u4e86\u3092\u53ef\u80fd\u306b\u3059\u308b\uff0e\u3053\u306e\u30e1\u30ab\u30cb\u30ba\u30e0\u304b\u3089\uff0c\u30bf\u30a4\u30c1\u30f3\u306f\u67b6\u6a4b\u5f62\u6210\u306a\u3057\u3067\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3059\u308b\u3053\u3068\u304c\u3067\u304d\u306a\u3044\u3060\u3051\u3067\u306a\u304f\uff0c\u30d1\u30ef\u30fc\u30b9\u30c8\u30ed\u30fc\u30af\u306e\u5b8c\u4e86\u3082\u307e\u305f\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306b\u4f9d\u5b58\u3059\u308b\u3053\u3068\u3082\u660e\u3089\u304b\u3067\u3042\u308b\uff0e\u5f93\u3063\u3066\uff0c\u30bf\u30a4\u30df\u30f3\u3068\u30df\u30aa\u30b7\u30f3\u306f\u3001\u9ad8\u3044\u8ca0\u8377\u306b\u5bfe\u3057\u3066\u30b5\u30eb\u30b3\u30e1\u30a2\u3092\u77ed\u7e2e\u3059\u308b\u305f\u3081\u306b\u624b\u3092\u643a\u3048\u3066\u50cd\u304f\uff0e<\/figcaption><\/figure>\n<p>\u3000\u7b4b\u8089\u53ce\u7e2e\u6642\u306b\u3053\u308c\u304c\u767a\u751f\u3059\u308b\u969b\u306e\u3082\u3063\u3068\u3082\u3089\u3057\u3044\u30b7\u30ca\u30ea\u30aa\u306f\uff0c\u5404\u30bf\u30a4\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u304c 12 pN \u306e\u529b\u3092\u652f\u3048\u308b\u305f\u3081\u306b\uff0c\u3044\u3064\u30b5\u30eb\u30b3\u30e1\u30a2\u304c\u53d7\u52d5\u7684\u306b\u8ca0\u8377\u3055\u308c\u308b\u304b\u3067\u3042\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/9580564\" rel=\"noopener noreferrer\" target=\"_blank\">86<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/16230467\" rel=\"noopener noreferrer\" target=\"_blank\">87<\/a>). \u7b4b\u8089\u306e\u6d3b\u52d5\u524d\u5f8c\u306b\u304a\u3044\u3066\u30bf\u30a4\u30c1\u30f3\u3068\u30df\u30aa\u30b7\u30f3\u9593\u3067\uff0c\u3069\u3046\u3084\u3063\u3066\u3053\u306e\u8ca0\u8377\u3092\u5206\u914d\u3057\u3066\u3044\u308b\u306e\u3060\u308d\u3046\uff1f\u4eee\u306b 147 \u672c\u306e\u53cc\u982d\u306e\u30df\u30aa\u30b7\u30f3\u5206\u5b50\u304c\u534a\u5206\u306e\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u5185\u306b\u3042\u308a\uff0c\u307e\u305f\u30b5\u30eb\u30b3\u30e1\u30a2\u534a\u5206\u3042\u305f\u308a 6 \u500b\u306e\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u304c\u3042\u308b\u3068\u8003\u3048\u308b\u306a\u3089\uff0c\u30bf\u30a4\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3042\u305f\u308a\u306e\u30df\u30aa\u30b7\u30f3\u5206\u5b50\u306e\u7d76\u5bfe\u7684\u306a\u6700\u5927\u6bd4\u306f 25 \u3067\u3042\u308b\uff0e\u3057\u304b\u3057\uff0c\u7b4b\u77ed\u7e2e\u4e2d\u306b\u3059\u3079\u3066\u306e\u30df\u30aa\u30b7\u30f3\u982d\u90e8\u304c\u30a2\u30af\u30c1\u30f3\u7d50\u5408\u90e8\u4f4d\u306b\u7d50\u5408\u3057\u3066\u3044\u308b\u308f\u3051\u3067\u306f\u306a\u3044\uff0e\u7b4b\u30df\u30aa\u30b7\u30f3\u306f\u975e\u9023\u7d9a\u30e2\u30fc\u30bf\u30fc\u3067\u3042\u308a\uff0c\u53ce\u7e2e\u4e2d\u306b\u30df\u30aa\u30b7\u30f3\u982d\u90e8\u304c\u30a2\u30af\u30c1\u30f3\u306b\u7d50\u5408\u3059\u308b\u30d1\u30fc\u30bb\u30f3\u30c6\u30fc\u30b8\u306f\u7d50\u5408\u90e8\u4f4d\u306b\u7d50\u5408\u53ef\u80fd\u306a\u6700\u5927\u6570\u306e 10-20 % \u7a0b\u5ea6\u3068\u63a8\u5b9a\u3055\u308c\u3066\u304d\u305f (<a href=\"http:\/\/jgp.rupress.org\/content\/143\/3\/387\" rel=\"noopener noreferrer\" target=\"_blank\">88<\/a>). \u53ce\u7e2e\u4e2d\u306e\u7b4b\u7dda\u7dad\u306e\u8ffd\u5f93\u3092\u8a08\u6e2c\u3059\u308b\u3053\u3068\u3067\uff0c1 \u672c\u306e\u592a\u3044\u7dda\u7dad\u3042\u305f\u308a 84 \u500b (~ 24 %) \u307e\u3067\u306e\u30df\u30aa\u30b7\u30f3\u982d\u90e8\u304c\u5f93\u4e8b\u3057\u3066\u3044\u308b\u53ef\u80fd\u6027\u304c\u3042\u308b\u3068\u8a08\u7b97\u3055\u308c\uff0c\u7d50\u5408\u3057\u305f\u982d\u90e8\u306e\u6570\u304c\u6025\u901f\u306b\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e\u30aa\u30fc\u30d0\u30fc\u30e9\u30c3\u30d7\u304a\u3088\u3073\u77ed\u7e2e\u901f\u5ea6\u306e\u4e21\u8005\u3092\u5909\u5316\u3055\u305b\u3066\u3044\u305f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/18022371\" rel=\"noopener noreferrer\" target=\"_blank\">89<\/a>). \u3057\u304b\u3057\uff0c\u30bf\u30a4\u30c1\u30f3\u306e\u5f3e\u529b\u6027\u306f\u3053\u306e\u5b9f\u9a13\u3067\u306f\u8003\u616e\u3055\u308c\u3066\u304a\u3089\u305a\uff0c\u305d\u306e\u30c9\u30e1\u30a4\u30f3\u304c\u6298\u308a\u305f\u305f\u307e\u308c\u3066\u30bf\u30a4\u30c1\u30f3\u306f\u786c\u5316\u3057\u3066\u3044\u308b\u3068\u3057\u3066\uff0c\u53ce\u7e2e\u3057\u3066\u3044\u308b\u7b4b\u8089\u3067\u5f93\u4e8b\u3057\u3066\u3044\u308b\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306e\u6570\u3092\u904e\u5270\u63a8\u5b9a\u3059\u308b\u7d50\u679c\u3068\u306a\u3063\u305f\uff0e\u30a6\u30b5\u30ae\u306e\u5927\u8170\u7b4b\u306e 3.1 &micro;m \u3084\u30d2\u30c8\u4f38\u5c55\u7b4b\u306e 3.4 &micro;m \u306a\u3069\u751f\u7406\u7684\u306a\u30b5\u30eb\u30b3\u30e1\u30a2\u306e\u4f38\u5c55\u3067\u306f\uff0c\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3068\u7d30\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e\u30aa\u30fc\u30d0\u30fc\u30e9\u30c3\u30d7\u306e\u7a0b\u5ea6\u304c\u6e1b\u5c11\u3059\u308b\u3068\u30df\u30aa\u30b7\u30f3\u982d\u90e8\u306e\u7d50\u5408\u6570\u304c\u5c11\u306a\u304f\u306a\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/26854230\" rel=\"noopener noreferrer\" target=\"_blank\">10<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/18600262\" rel=\"noopener noreferrer\" target=\"_blank\">90<\/a>). \u3086\u3048\u306b\uff0c\u305d\u306e\u3088\u3046\u306a\u4f38\u5c55\u3055\u308c\u305f\u9577\u3055\u3067\u306f\uff0c\u53ce\u7e2e\u3092\u652f\u3048\u308b\u30bf\u30a4\u30c1\u30f3\u5206\u5b50 1 \u500b\u3042\u305f\u308a\u306e\u30df\u30aa\u30b7\u30f3\u982d\u90e8\u306e\u6570\u306f\u305f\u3063\u305f 1-2 \u500b\u306b\u904e\u304e\u306a\u3044\u3068\u8a08\u7b97\u3055\u308c\u305f\uff0e<\/p>\n<p>\u3000\u4eee\u8aac\u306e\u30b7\u30ca\u30ea\u30aa\u306b\u623b\u308b\u304c\uff0c\u30bf\u30a4\u30c1\u30f3\u306f\u7b4b\u53ce\u7e2e\u306b\u5148\u7acb\u3063\u3066 12 pN \u306e\u8ca0\u8377\u5168\u4f53\u3092\u652f\u3048\uff0c\u8fd1\u4f4d Ig \u30c9\u30e1\u30a4\u30f3\u3092\u52d5\u54e1\u3057\u3066\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c9\u72b6\u614b\u306b\u3059\u308b (<strong>Figure 3B<\/strong>, \u4e0a\u90e8\u30d1\u30cd\u30eb). \u4f8b\u3048\u3070\u91ce\u7403\u306e\u6295\u624b\u306e\u30ef\u30a4\u30f3\u30c9\u30a2\u30c3\u30d7\u4e2d\u306e\u3088\u3046\u306b\uff0c\u3053\u308c\u306f\u7b4b\u8089\u306e\u7206\u767a\u7684\u53ce\u7e2e\u306b\u5148\u7acb\u3063\u3066\u8d77\u304d\u308b\u53ef\u80fd\u6027\u304c\u3042\u308b\uff0e\u7b4b\u8089\u306e\u6d3b\u52d5\u5f8c\u306e\u6570\u30df\u30ea\u79d2\u9593\u3067\uff0c\u6700\u5927\u6570\u306e\u67b6\u6a4b\u304c\u5f62\u6210\u3055\u308c\u3066\u30b5\u30eb\u30b3\u30e1\u30a2\u5185\u306e\u4e09\u3064\u306e\u7b4b\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u304b\u3089\u306a\u308b\u6a5f\u68b0\u7684\u30cd\u30c3\u30c8\u30ef\u30fc\u30af\u306b\u8ca0\u8377\u304c\u5206\u914d\u3055\u308c\u308b\uff0e\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306f\u6700\u5927 6 pN \u306e\u52d5\u529b\u3092\u767a\u751f\u3055\u305b\u308b\u304c\uff0c\u30b5\u30eb\u30b3\u30e1\u30a2\u3092\u77ed\u7e2e\u3055\u305b\u308b\u307b\u3069\u5341\u5206\u3067\u306f\u306a\u304f\uff0c\u5168\u3066\u306e\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u306b\u304b\u304b\u308b\u8ca0\u8377\u3092 6 pN \u306b\u6e1b\u5f31\u3055\u305b\u308b (<strong>Figure 3B<\/strong>, \u4e2d\u90e8\u30d1\u30cd\u30eb). \u3053\u306e\u7d50\u679c\u307e\u305a\u975e\u5e38\u306b\u6025\u901f\u306b\u5168\u3066\u306e\u4f38\u5c55\u3055\u308c\u305f\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u304c\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc\u53cd\u52d5\u3092\u8d77\u3053\u3057\uff0c\u7d9a\u3044\u3066\u30bf\u30a4\u30c1\u30f3\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304c\u8d77\u304d\u3066\u6700\u5927\u53ef\u80fd\u306a\u4ed5\u4e8b\u3092\u306a\u3057\u3066\u529b\u3092\u56de\u5fa9\u3055\u305b\u308b (<strong>Figure 3A<\/strong>, \u9752\u66f2\u7dda\u30d4\u30fc\u30af). \u5f93\u6765\u306e\u4eee\u8aac\u3067\u306f\u30bf\u30a4\u30c1\u30f3\u306f\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc\u53cd\u52d5\u7d4c\u7531\u3067\u306e\u307f\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u56de\u5fa9\u3059\u308b\u3068\u307f\u306a\u3055\u308c\u3066\u3044\u305f\u304c\uff0c\u305d\u3046\u3067\u306f\u306a\u304f\uff0c\u3053\u3053\u3067\u56de\u5fa9\u3059\u308b\u30a8\u30cd\u30eb\u30ae\u30fc\u306e\u5927\u534a\u306f\u5b9f\u969b\u306b\u306f\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53ce\u7e2e\u306b\u7531\u6765\u3057\u3066\u3044\u308b\u306e\u3067\u3042\u308b. 12 pN \u306e\u529b\u306b\u304a\u3044\u3066\uff0c\u6298\u308a\u305f\u305f\u307e\u308c\u305f\u72b6\u614b\u304b\u3089\u30e9\u30f3\u30c0\u30e0\u30b3\u30a4\u30eb\u306b\u884c\u304f\u3053\u3068\u3067\u9054\u6210\u3055\u308c\u308b\u8f2a\u90ed\u9577\u306e\u5909\u5316\u306f 15.8 nm \u3067\u3042\u308b. 6 pN \u307e\u3067\u306e\u529b\u3092\u6e1b\u5f31\u3055\u305b\u308b\u306b\u304a\u3044\u3066, 5.9 nm \u306e\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc\u53cd\u52d5\u306b\u5f15\u304d\u7d9a\u3044\u3066 9.9 nm \u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53ce\u7e2e\u304c\u8d77\u304d\u308b (<strong>Figure 3B<\/strong>, \u4e2d\u90e8\u30d1\u30cd\u30eb). \u86cb\u767d\u8cea\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306f\uff0c\u4f38\u5c55\u3055\u308c\u305f\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u304b\u3089\u306e\u56de\u5fa9\u306b\u3088\u308b\u30a8\u30cd\u30eb\u30ae\u30fc\u306e 63 % \u3092\u542b\u3080\u4e00\u65b9\uff0c\u5f3e\u6027\u53ce\u7e2e\u306f 37 % \u3092\u63d0\u4f9b\u3059\u308b\u306b\u904e\u304e\u306a\u3044\uff0e<strong>\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53ce\u7e2e\u3053\u305d\uff0c\u751f\u7406\u5b66\u7684\u306a\u529b\u306e\u7bc4\u56f2\u3067\u306e\u30bf\u30a4\u30c1\u30f3\u306e\u5f3e\u6027\u304a\u3088\u3073\u30a8\u30cd\u30eb\u30ae\u30fc\u84c4\u7a4d\u306e\u4e3b\u8981\u306a\u8981\u7d20\u306b\u4ed6\u306a\u3089\u306a\u3044<\/strong>\uff0e\u3086\u3048\u306b\uff0c\u3053\u306e\u30b7\u30ca\u30ea\u30aa\u3067\u306f, 12 pN \u4ee5\u4e0b\u306e\u8ca0\u8377\u3067\u306f\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3057\u305f Ig \u30c9\u30e1\u30a4\u30f3\u304c\u5931\u901f\u3057\u305f\u30e2\u30fc\u30bf\u30fc\u3092\u89e3\u653e\u3057\uff0c\u30b5\u30eb\u30b3\u30e1\u30a2\u306b\u306f\u4f38\u9577\u3059\u308b\u3088\u308a\u3080\u3057\u308d\u53ce\u7e2e\u3059\u308b\u3053\u3068\u3092\u8a31\u3059 (<strong>Figure 3B<\/strong>, \u4e0b\u90e8\u30d1\u30cd\u30eb). 12 pN \u3067\u30bf\u30a4\u30c1\u30f3\u3092\u4f38\u5c55\u3055\u305b\u308b\u3053\u3068\u3067\u84c4\u7a4d\u3055\u308c\u308b\u30a8\u30cd\u30eb\u30ae\u30fc\u306f\uff0c\u30e2\u30fc\u30bf\u30fc\u304c\u5f93\u4e8b\u3059\u308b\u6642\u306b\u306e\u307f\uff0c\u623b\u3063\u3066\u304f\u308b\uff0e\u3053\u306e\u7d20\u6674\u3089\u3057\u3044\u30e1\u30ab\u30cb\u30ba\u30e0\u306f, II \u578b\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u304c\u5931\u901f\u3057\u305f\u529b\u3092\u8d85\u3048\u308b\u8ca0\u8377\u3067\u3082\uff0c\u30b5\u30eb\u30b3\u30e1\u30a2\u304c\u52d5\u529b\u3092\u63d0\u4f9b\u3059\u308b\u53ef\u80fd\u6027\u3092\u5bb9\u6613\u306b\u62e1\u5f35\u3059\u308b\u3082\u306e\u3067\u3042\u308b\uff0e<\/p>\n<h3>Titin folding dictates the optimal sarcomere length of a contracting muscle<\/h3>\n<p>\u3000\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53ce\u7e2e\u306e\u5b58\u5728\u306f\uff0c\u7b4b\u751f\u7406\u5b66\u306e\u6559\u7fa9\u306e\u518d\u8a2a\u3092\u4fdd\u8a3c\u3059\u308b\u53ef\u80fd\u6027\u304c\u3042\u308b\uff0e\u4f8b\u3048\u3070\uff0c\u6b21\u306e\u3053\u3068\u306f\u4e00\u822c\u306b\u53d7\u3051\u5165\u308c\u3089\u308c\u3066\u3044\u308b\uff0e\u3064\u307e\u308a\uff0c\u4e0b\u80a2\u306e active force \u3068\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u3068\u306e\u95a2\u4fc2\u306f\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3068\u7d30\u3044\u30d5\u30a9\u30e9\u30e1\u30f3\u30c8\u306e\u30aa\u30fc\u30d0\u30fc\u30e9\u30c3\u30d7\u306e\u6e1b\u5c11\u306b\u3088\u308b\uff0c\u3068\u3044\u3046\u3053\u3068\u3067\u3042\u308b\uff0e\u3057\u304b\u3057\uff0c\u30d2\u30c8\u9aa8\u683c\u7b4b\u306b\u304a\u3044\u3066\u30b5\u30eb\u30b3\u30e1\u30a2\u9577 1.7-2.6 &micro;m \u3067\u4e0a\u80a2\u306b\u304a\u3051\u308b\u529b\u306f\u65e2\u306b\u6700\u5927\u306b\u8fd1\u3044 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/10233103\" rel=\"noopener noreferrer\" target=\"_blank\">91<\/a>). \u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306f\u305b\u3044\u305c\u3044 1.6 &micro;m \u307e\u3067\u306e\u9577\u3055\u3067\u3042\u308a\uff0c\u305d\u306e\u305f\u3081\u4e0a\u80a2\u306e\u3053\u306e\u90e8\u4f4d\u306b\u304a\u3044\u3066\u6e1b\u5c11\u3057\u305f\u529b\u306f\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e Z \u76e4\u306b\u5bfe\u3059\u308b\u5727\u7e2e\u3067\u306f\u306a\u3055\u305d\u3046\u3067\u3042\u308b\uff0e\u5225\u306e\u8aac\u660e\u3068\u3057\u3066\u53ef\u80fd\u6027\u304c\u3042\u308b\u306e\u306f\uff0c\u3053\u306e\u90e8\u4f4d\u3067\u306e\u7b49\u5c3a\u6027\u306e\u529b\u304c\u6e1b\u5c11\u3059\u308b\u306e\u306f\u5168\u3066\u306e\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u304c\u6298\u308a\u305f\u305f\u307e\u308c\u305f\u307e\u307e\u3067\u3042\u308b\u304b\u3089\uff0c\u3068\u3044\u3046\u3082\u306e\u304c\u3042\u308b\uff0e\u3086\u3048\u306b\uff0c\u30bf\u30a4\u30c1\u30f3\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u306e\u758e\u6c34\u6027\u5d29\u58ca\u306b\u3088\u308a\u767a\u751f\u3059\u308b\u4ed8\u52a0\u7684\u306a\u53ce\u7e2e\u529b\u3068\u3044\u3046\u3082\u306e\u306f\u5b58\u5728\u3057\u306a\u3044\uff0e\u4eee\u8aac\u7684\u306a\u9aa8\u683c\u7b4b\u3092\u8003\u3048\u3066\u307f\u3088\u3046\uff0e\u30bf\u30a4\u30c1\u30f3\u30a2\u30a4\u30bd\u30d5\u30a9\u30fc\u30e0\u304c\u4e2d\u9593\u7684\u306a\u9577\u3055\u3068\u786c\u5ea6\u3067\u5b58\u5728\u3059\u308b\u3068\u3059\u308b (<strong>Figure 4A<\/strong>, \u9752\u66f2\u7dda). \u3053\u306e\u30bf\u30a4\u30c1\u30f3\u30a2\u30a4\u30bd\u30d5\u30a9\u30fc\u30e0\u306f\u7b4b\u8089\u304b\u3089\u306e\u6700\u9ad8\u306e\u30d1\u30d5\u30a9\u30fc\u30de\u30f3\u30b9\u3092\u53ef\u80fd\u3068\u3059\u308b\uff0e\u3068\u3044\u3046\u306e\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u78ba\u7387\u306b\u304a\u3051\u308b\u6700\u5927\u306e\u5909\u5316\u306f\uff0c\u6700\u5927\u306e active force \u767a\u751f\u3059\u308b\u90e8\u4f4d\u5185\u3067\u307e\u3063\u3059\u3050\u306b\u843d\u3061\u8fbc\u3080\u304b\u3089\u3067\u3042\u308b (Figure 4A, \u30d4\u30f3\u30af\u9818\u57df). \u30b0\u30ec\u30fc\u9818\u57df\u3067\u306e\u6700\u5927\u7b49\u5c3a\u529b\u306e\u8a08\u6e2c\u3067\u306f\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u95a2\u4e0e\u3092\u6b20\u3044\u3066\u3044\u308b\uff0e\u3068\u3044\u3046\u306e\u306f\uff0c\u8a72\u5f53\u3059\u308b\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u3067\u306e\u30bf\u30a4\u30c1\u30f3\u306f\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306b\u3082\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306b\u3082\u3044\u305a\u308c\u306e\u72b6\u614b\u3082\u3042\u308a\u3046\u308b\u304b\u3089\u3067\u3042\u308b (<strong>Figure 4A<\/strong>, \u30b0\u30ec\u30fc\u9818\u57df). \u3053\u308c\u3089\u306e\u4e00\u898b\u5358\u7d14\u306a\u5b9f\u9a13\u306f\u7b4b\u751f\u7406\u5b66\u306e\u57fa\u790e\u3092\u652f\u6301\u3057\u3066\u304a\u308a\uff0c\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u6587\u8108\u3067\u8003\u616e\u3059\u308b\u3068\uff0c\u305d\u3046\u7c21\u5358\u306a\u3082\u306e\u3067\u306f\u306a\u3044\uff0e<\/p>\n<figure id=\"attachment_3620\" aria-describedby=\"caption-attachment-3620\" style=\"width: 525px\" class=\"wp-caption aligncenter\"><a href=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure4.png\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure4-1024x388.png\" alt=\"\" width=\"525\" height=\"199\" class=\"size-large wp-image-3620\" srcset=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure4-1024x388.png 1024w, https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure4-300x114.png 300w, https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure4-768x291.png 768w, https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure4.png 1419w\" sizes=\"auto, (max-width: 525px) 100vw, 525px\" \/><\/a><figcaption id=\"caption-attachment-3620\" class=\"wp-caption-text\"><strong>Figure 4<\/strong><br \/><strong>\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306f\u751f\u7406\u5b66\u7684\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u3067\u751f\u3058\u308b<\/strong><br \/><strong>(A)<\/strong> \u30d2\u30c8\u306e\u9aa8\u683c\u7b4b\uff08\u9ed2\u3044\u66f2\u7dda\uff09\u306e\u30a2\u30af\u30c6\u30a3\u30d6\u30d5\u30a9\u30fc\u30b9\u5bfe\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u306e\u95a2\u4fc2 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/10233103\" rel=\"noopener noreferrer\" target=\"_blank\">91<\/a>) \u306f\uff0c\u4e2d\u9593\u9577\u306e\u30bf\u30a4\u30c1\u30f3\u30a2\u30a4\u30bd\u30d5\u30a9\u30fc\u30e0\u3092\u6709\u3059\u308b\u4eee\u60f3\u306e\u7b4b\u8089\u306b\u3064\u3044\u3066\uff0c\u30bf\u30a4\u30c1\u30f3\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u78ba\u7387\u3068\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u306e\u95a2\u4fc2\uff08\u7070\u8272\u306e\u66f2\u7dda\uff09\u3068\u91cd\u306a\u308b\uff0e\u3053\u3053\u3067\uff0c\u30a2\u30af\u30c6\u30a3\u30d6\u30d5\u30a9\u30fc\u30b9\u304c\u6700\u5927\u3068\u306a\u308b\u7bc4\u56f2\u3067\u306f\uff0c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u78ba\u7387\u304c\u6025\u6fc0\u306b 0 \u304b\u3089 1 \u306b\u5909\u5316\u3057\uff0c\u7b4b\u53ce\u7e2e\u529b\u304c\u5927\u304d\u304f\u5897\u5927\u3059\u308b\uff0e\u30a2\u30af\u30c6\u30a3\u30d6\u30d5\u30a9\u30fc\u30b9\u66f2\u7dda\u306e\u4e0a\u6607\u5c40\u9762\u3067\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u306f\u6298\u308a\u305f\u305f\u307e\u308c\u305f\u307e\u307e\u3067\u3042\u308a\uff0c\u7d04 2.5 \u03bcm \u672a\u6e80\u306e\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u3067\u529b\u306e\u767a\u751f\u306b\u5bc4\u4e0e\u3059\u308b\u3053\u3068\u304c\u3067\u304d\u306a\u3044\uff0e\u4e0b\u884c\u5c40\u9762\u3067\u306f\uff0c3.1 \u03bcm \u3088\u308a\u3082\u5927\u304d\u3044\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u3067\u306f\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304c\u307b\u3068\u3093\u3069\u306a\u3044\u305f\u3081\uff0c\u30bf\u30a4\u30c1\u30f3\u306f\u4e3b\u306b\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc\u53cd\u52d5\u5fa9\u5143\u529b\u3092\u63d0\u4f9b\u3059\u308b\uff0e\u3059\u3079\u3066\u306e\u7b4b\u8089\u304c\u30a2\u30af\u30c6\u30a3\u30d6\u30d5\u30a9\u30fc\u30b9\u5bfe\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u306e\u95a2\u4fc2\u3067\u52d5\u4f5c\u3059\u308b\u308f\u3051\u3067\u306f\u306a\u3044\u306e\u3067\uff0c\u5404\u7b4b\u8089\u7fa4\u306f\uff0c\u305d\u306e\u7279\u5b9a\u306e\u7b4b\u8089\u7fa4\u306e\u4f5c\u52d5\u7bc4\u56f2\u306b\u5165\u308b\u3088\u3046\u306b\uff0c\u9078\u629e\u7684\u306b\u30b9\u30d7\u30e9\u30a4\u30b9\u3055\u308c\u305f\u30bf\u30a4\u30c1\u30f3\u30a2\u30a4\u30bd\u30d5\u30a9\u30fc\u30e0\u3092\u6709\u3059\u308b\u53ef\u80fd\u6027\u304c\u9ad8\u3044 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/16230467\" rel=\"noopener noreferrer\" target=\"_blank\">87<\/a>). <strong>(B)<\/strong> \u30bf\u30a4\u30c1\u30f3 I-band \u304c\u591a\u304f\u306e\u96e2\u6563 Ig \u30c9\u30e1\u30a4\u30f3\u306b\u5206\u7bc0\u5316\u3057\u3066\u3044\u308b\u3053\u3068\u306f\uff0c\u7279\u5225\u306a\u6750\u6599\u7279\u6027\u3092\u4ed8\u4e0e\u3059\u308b\uff0e\u6b63\u5e38\u306a\u751f\u7406\u7684\u6761\u4ef6\u4e0b\uff08\u56fa\u4f53\u306e\u9752\u8272\u306e\u66f2\u7dda\uff09\u306e\u30bf\u30a4\u30c1\u30f3\u306f, 4-15 pN \uff08\u7070\u8272\u306e\u9670\u5f71\u9818\u57df\uff09\u306e\u751f\u7406\u5b66\u7684\u306a\u529b\u306e\u7bc4\u56f2\u5185\u3067\u305d\u306e Ig \u30c9\u30e1\u30a4\u30f3\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304a\u3088\u3073\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3092\u7d4c\u9a13\u3059\u308b\uff0e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53cd\u5fdc\u3092\u5b8c\u5168\u306b\u963b\u6b62\u3059\u308b\u3053\u3068\u306b\u3088\u308a\uff0c\u30bf\u30a4\u30c1\u30f3\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u78ba\u7387\uff0c\u3072\u3044\u3066\u306f\u5f3e\u529b\u7279\u6027\u3092\u5909\u3048\u308b\u3053\u3068\u304c\u3067\u304d\u308b\uff08\u4f8b\uff0eS-\u30b0\u30eb\u30bf\u30c1\u30aa\u30cb\u30eb\u5316\uff0c\u9752\u8272\u306e\u7834\u7dda\u66f2\u7dda\uff09\uff0e\u9178\u5316\u7684\u30b9\u30c8\u30ec\u30b9\u307e\u305f\u306f\u9175\u7d20\u3092\u4ecb\u3057\u305f\u5206\u5b50\u5185\u30b8\u30b9\u30eb\u30d5\u30a3\u30c9\u7d50\u5408\u306e\u5c0e\u5165\u306f\uff0c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53cd\u5fdc\u304c\u8d77\u3053\u308b\u529b\u3092\u5897\u52a0\u3055\u305b\u308b\uff08\u91d1\u8272\u306e\u66f2\u7dda\uff09\uff0e\u5c0f\u5206\u5b50\u307e\u305f\u306f\u30b7\u30e3\u30da\u30ed\u30f3\u306e\u7d50\u5408\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u306e\u6a5f\u68b0\u7684\u5b89\u5b9a\u6027\u3092\u5897\u52a0\u3055\u305b, Ig \u30c9\u30e1\u30a4\u30f3\u3092\u3088\u308a\u4f38\u5c55\u4e0d\u80fd\u306b\u3057\uff0c\u30bf\u30a4\u30c1\u30f3\u306e\u5168\u4f53\u7684\u306a\u525b\u6027\u3092\u5897\u52a0\u3055\u305b\u308b\uff08\u8d64\u8272\u66f2\u7dda\u7834\u7dda\uff09\uff0e\u5b9f\u969b\u306b\u3001\u30bf\u30a4\u30c1\u30f3\u306e\u5f3e\u529b\u6027\u306f\u3053\u308c\u3089\u306e\u66f2\u7dda\u306e\u3059\u3079\u3066\u306e\u6df7\u5408\u7269\u3067\u3042\u308a\uff0c\u7b4b\u8089\u306e\u4ee3\u8b1d\u72b6\u614b\u306b\u4f9d\u5b58\u3059\u308b\uff0e<\/figcaption><\/figure>\n<h3>Why protein folding?<\/h3>\n<p>\u3000\u7b4b\u8089\u306e\u53ef\u9006\u7684\u5909\u5f62\u306f\uff0c\u304a\u305d\u3089\u304f\u4ed6\u306e\u7d44\u7e54\u3068\u306f\u6bd4\u985e\u306e\u306a\u3044\u6750\u6599\u7279\u6027\u3067\u3042\u308b\uff0e\u7b4b\u8089\u306f\u6750\u6599\u75b2\u52b4\u3084\u30d2\u30b9\u30c6\u30ea\u30b7\u30b9\u306a\u3057\u306b\u4f38\u5c55\u3068\u5f1b\u7de9\u306e\u6301\u7d9a\u7684\u306a\u30b5\u30a4\u30af\u30eb\u3092\u7d4c\u9a13\u3057\u306a\u304f\u3066\u306f\u306a\u3089\u306a\u3044\uff0e\u3044\u304f\u3064\u304b\u306e\u7b4b\u8089\u7fa4\u3067\u306f\uff0c\u5b8c\u5168\u53ce\u7e2e\u304b\u3089\u5b8c\u5168\u4f38\u5c55\u72b6\u614b\u306b\u81f3\u308b\u3068\u30b5\u30eb\u30b3\u30e1\u30a2\u306e\u9577\u3055\u304c 300 nm \u306b\u307e\u3067\u4f38\u5c55\u3059\u308b\u53ef\u80fd\u6027\u304c\u3042\u308a\uff0c\u3053\u308c\u306f\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u306e 10 % \u306e\u5897\u52a0\u306b\u3042\u305f\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/18600262\" rel=\"noopener noreferrer\" target=\"_blank\">90<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC5858182\/\" rel=\"noopener noreferrer\" target=\"_blank\">92<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC5920519\/\" rel=\"noopener noreferrer\" target=\"_blank\">93<\/a>). \u53ef\u9006\u7684\u306a\u7b4b\u5f3e\u6027\u306b\u5bc4\u4e0e\u3059\u308b\u305d\u306e\u86cb\u767d\u8cea\u306f\uff0c\u76f8\u53cd\u3059\u308b\u4e8c\u3064\u306e\u5f79\u5272\u3092\u6e80\u305f\u3055\u306a\u304f\u3066\u306f\u306a\u3089\u306a\u3044\uff0e\u3064\u307e\u308a\uff0c\u7b4b\u8089\u306f\u5bb9\u6613\u306b\u5909\u5f62\u3067\u304d\uff0c\u62ee\u6297\u7b4b\u306e\u53ce\u7e2e\u3092\u59a8\u5bb3\u3057\u306a\u3044\u7a0b\u5ea6\u306b\u306f\u67d4\u8edf\u3067\u306a\u304f\u3066\u306f\u306a\u3089\u306a\u3044\u304c\uff0c\u7b4b\u8089\u904e\u4f38\u5c55\u3084\u7b4b\u53ce\u7e2e\u6642\u306b\u89e3\u653e\u3055\u308c\u308b\u81a8\u5927\u306a\u5f3e\u6027\u30a8\u30cd\u30eb\u30ae\u30fc\u84c4\u7a4d\u306b\u3088\u308b\u640d\u50b7\u3092\u907f\u3051\u308b\u7a0b\u5ea6\u306b\u306f\u56fa\u304f\u306a\u304f\u3066\u306f\u306a\u3089\u306a\u3044\uff0e\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306f\uff0c\u6b63\u3057\u304f\u69cb\u9020\u5316\u3055\u308c\u308c\u3070\uff0c\u5f3e\u6027\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u84c4\u7a4d\u3057\u89e3\u653e\u3059\u308b\u305f\u3081\u306e\u512a\u308c\u305f\u6750\u6599\u3067\u3042\u308b\u3053\u3068\u304c\u5206\u304b\u3063\u305f\uff0e<\/p>\n<p>\u3000\u591a\u304f\u306e\u7d30\u80de\u9aa8\u683c\u69cb\u9020\u306f\u7b4b\u8089\u304c\u52d5\u4f5c\u3059\u308b\u7bc4\u56f2\u306e\u529b\u3067\u306f\u786c\u3044\uff0e\u5fae\u5c0f\u7ba1\u306f\u30df\u30ea\u30e1\u30fc\u30c8\u30eb\u4ee5\u4e0a\u306e\u6c38\u7d9a\u6027\u306e\u9577\u3055\u3092\u6709\u3057\u3066\u3044\u308b\u4e00\u65b9\u3067, f-\u30a2\u30af\u30c1\u30f3\u306f 10 \u30df\u30af\u30ed\u30f3\u306e\u6c38\u7d9a\u6027\u306e\u9577\u3055\u3092\u6709\u3057\u3066\u3044\u308b\u306e\u307f\u3067\u3042\u308b (<a href=\"http:\/\/jcb.rupress.org\/content\/120\/4\/923\" rel=\"noopener noreferrer\" target=\"_blank\">94<\/a>). \u30b1\u30e9\u30c1\u30f3\u3084\u30d3\u30e1\u30f3\u30c1\u30f3\u306e\u3088\u3046\u306a\u4e2d\u9593\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306f\u6570\u767e\u30ca\u30ce\u30e1\u30fc\u30c8\u30eb\u304b\u3089\u30de\u30a4\u30af\u30ed\u30e1\u30fc\u30c8\u30eb\u307e\u3067\u306e\u6c38\u7d9a\u6027\u306e\u9577\u3055\u3092\u6709\u3059\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC1303373\/\" rel=\"noopener noreferrer\" target=\"_blank\">95<\/a>). I \u578b\u30b3\u30e9\u30fc\u30b2\u30f3\u306f\u4e09\u3064\u306e\u72ec\u7acb\u3057\u305f\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u9396\u306e\u4e09\u91cd\u3089\u305b\u3093\u3067\u69cb\u6210\u3055\u308c\u308b\u304c\uff0c\u6570\u5343\u30ca\u30ce\u30e1\u30fc\u30c8\u30eb\u3082\u306e\u6c38\u7d9a\u6027\u306e\u9577\u3055\u3092\u6709\u3059\u308b\u3068\u8003\u3048\u3089\u308c\u3066\u3044\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/23907817\" rel=\"noopener noreferrer\" target=\"_blank\">96<\/a>). \u3053\u308c\u3089\u306e\u7d20\u6750\u306e\u3046\u3061\u3067\u6709\u7528\u306a\u5f3e\u529b\u6027\u3092\u84c4\u7a4d\u3059\u308b\u30e1\u30ab\u30cb\u30ba\u30e0\u3092\u6709\u3059\u308b\u3082\u306e\u306f\u5b58\u5728\u3057\u306a\u3044\uff0e\u7406\u7531\u306f\u5358\u7d14\u3067\uff0c\u305d\u308c\u3089\u306f\u786c\u3059\u304e\u308b\u306e\u3067\u3042\u308b\uff0e\u6b63\u5e38\u306a\u7b4b\u8089\u306e\u52d5\u4f5c\u6642\u306b\u5358\u4e00\u306e\u86cb\u767d\u8cea\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u304c\u7d4c\u9a13\u3059\u308b\u529b\u306e\u7bc4\u56f2 (1-15 pN) \u3092\u8d85\u3048\u308b\u3068\uff0c\u3053\u308c\u3089\u306e\u86cb\u767d\u8cea\u306f\u65e2\u306b\u5b8c\u5168\u306b\u4f38\u5c55\u3055\u308c\u3066\u3044\u308b\u304c\uff0c\u786c\u3044\u7aff\u306e\u3088\u3046\u306b\uff0c\u7b4b\u8089\u306e\u4f38\u3073\u306b\u62b5\u6297\u3059\u308b\u3060\u308d\u3046\uff0e\u4ee3\u308f\u308a\u306b\uff0c\u975e\u69cb\u9020\u7684\u307e\u305f\u306f\u4f38\u5c55\u3057\u305f\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u9396\u306f\u591a\u69d8\u306a\u6280\u8853\u3067\u7279\u5fb4\u3065\u3051\u3089\u308c\u3066\u304d\u305f\u3057, Kuhn \u9577\u306e\u7bc4\u56f2\u306f 0.8 nm \u304b\u3089 1.2 mn \u3067\u3042\u308b\u3068\u793a\u3055\u308c\u305f\uff08\u6c38\u7d9a\u6027\u306e\u9577\u3055\u306f 0.4 nm \u304b\u3089 0.6 nm \u306b\u7b49\u3057\u3044\uff09. 5 pN \u4ee5\u4e0b\u306e\u529b\u3067\u306f\uff0c\u3042\u308b\u30e9\u30f3\u30c0\u30e0\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u30b3\u30a4\u30eb\u306f\u307b\u307c Hookean \u3070\u306d\u3068\u3057\u3066\u3075\u308b\u307e\u3046\u3060\u308d\u3046\u304c\uff0c\u3088\u308a\u5927\u304d\u306a\u529b\u3067\u306f\uff0c\u305d\u306e\u30e9\u30f3\u30c0\u30e0\u30b3\u30a4\u30eb\u306f\u975e\u7dda\u5f62\u306e\u3075\u308b\u307e\u3044\u3092\u793a\u3057\uff0c\u4f38\u5c55\u304c\u5897\u52a0\u3059\u308b\u306b\u3064\u308c\u3066\u3088\u308a\u6025\u901f\u306b\u529b\u3092\u5897\u3059\uff0e\u6700\u8fd1\u306e\u767a\u898b\u3067\u306f\uff0c\u751f\u7406\u7684\u306a\u7b4b\u8089\u306e\u6d3b\u52d5\u6642\u306e\u30bf\u30a4\u30c1\u30f3\u306f 4-15 pN \u306e\u529b\u306e\u4e0b\u3067\u306f\u6b21\u306e\u7591\u554f\u3092\u63d0\u793a\u3059\u308b\uff0e\u3053\u306e\u529b\u306e\u7bc4\u56f2\u3067\uff0c\u5358\u7d14\u306a\u30e9\u30f3\u30c0\u30e0\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u30b3\u30a4\u30eb\u3088\u308a\u3082\u591a\u304f\u306e\u5f3e\u6027\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u84c4\u7a4d\u3059\u308b\u3053\u3068\u304c\u3067\u304d\u308b\u3088\u3046\u306a\u5f62\u614b\u3092\uff0c\u3042\u308b\u7d20\u6750\u306b\u53d6\u3089\u305b\u308b\u3053\u3068\u306f\u53ef\u80fd\u306a\u306e\u3060\u308d\u3046\u304b\uff1f<\/p>\n<p>\u3000\u30bf\u30a4\u30c1\u30f3\u306e\u69cb\u9020\u306f\u3053\u306e\u554f\u984c\u306b\u5bfe\u3057\u3066\uff0c\u975e\u69cb\u9020\u30bb\u30b0\u30e1\u30f3\u30c8 (PEVK, N2B) \u304a\u3088\u3073\u591a\u304f\u306e\u30bf\u30f3\u30c7\u30e0\u304b\u3089\u306a\u308b Ig \u30c9\u30e1\u30a4\u30f3\u306e\u4e21\u8005\u3067\u3082\u3063\u3066\u7279\u5225\u306a\u56de\u7b54\u3092\u63d0\u4f9b\u3057\u3066\u304f\u308c\u308b. Ig \u30c9\u30e1\u30a4\u30f3\u3078\u5206\u7bc0\u5316\u3057\u305f\u30bf\u30a4\u30c1\u30f3 I-band \u306f\u7b4b\u8089\u306e\u5f3e\u529b\u6027\u3092\u751f\u7406\u7684\u306a\u529b\u306e\u7bc4\u56f2\u3067\u5927\u304d\u304f\u5897\u52a0\u3055\u305b\u308b\uff0e\u305d\u306e\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u306f\u81a8\u5927\u306a\u30a8\u30cd\u30eb\u30ae\u30fc (W = F&Delta;L) \u3092\u84c4\u7a4d\u3059\u308b\uff0e\u3053\u3053\u3067 F \u306f FJC \u306b\u3088\u308a\u5b9a\u7fa9\u3055\u308c\u308b\u529b\u3067\u3042\u308a L \u306f\u89b3\u6e2c\u3055\u308c\u308b\u9577\u3055\u3067\u3042\u308b\uff0e\u4eee\u306b\u30bf\u30a4\u30c1\u30f3\u304c\u975e\u69cb\u9020\u30da\u30d7\u30c1\u30c9\u306e\u307f\u542b\u6709\u3059\u308b\u3068\u4eee\u5b9a\u3059\u308b\u3068, 12 pN \u304b\u3089 6 pN \u3078\u306e\u529b\u306e\u6e1b\u5f31\u306f\u305d\u306e\u4f38\u5c55\u9577\u306e 63 % \u306e\u5d29\u58ca\u306b\u7531\u6765\u3059\u308b\u3053\u3068\u306b\u306a\u308b. 90 \u500b\u307e\u3067\u306e\u30a2\u30df\u30ce\u9178\u3092\u542b\u6709\u3059\u308b\u975e\u69cb\u9020\u30bb\u30b0\u30e1\u30f3\u30c8\u306b\u3068\u3063\u3066\uff0c\u305d\u306e\u9577\u3055\u304c 15.7 nm \u304b\u3089 9.9 nm \u306b\u7e2e\u9000\u3059\u308b\u3053\u3068\u306b\u306a\u308a\uff0c\u305d\u306e\u7d50\u679c\u89e3\u653e\u3055\u308c\u308b\u5f3e\u6027\u30a8\u30cd\u30eb\u30ae\u30fc\u306f 34.8 zJ \u3068\u306a\u308b\uff0e\u4eca\u5ea6\u306f\u540c\u3058 90 \u500b\u306e\u30a2\u30df\u30ce\u9178\u3067\u5206\u7bc0\u5316\u3057\u3066\u6298\u308a\u305f\u305f\u3080\u3053\u3068\u306e\u3067\u304d\u308b\u3082\u306e\u3092\u8003\u3048\u3066\u307f\u3088\u3046\uff0e12 \u304b\u3089 6 pN \u306b\u529b\u304c\u6e1b\u5f31\u3059\u308b\u3068\uff0c\u307e\u305a 34.8 zJ\u3000\u304c\u653e\u51fa\u3055\u308c\uff0c\u6b21\u306b\u4ed8\u52a0\u7684\u306a 59.2 zJ \u304c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306b\u4f34\u3063\u3066\u653e\u51fa\u3055\u308c\u308b\uff0e\u30a8\u30cd\u30eb\u30ae\u30fc\u56de\u5fa9\u306e\u5408\u8a08\u306f\u4eca\u3084 94 zJ \u306b\u5897\u52a0\u3057, 3 \u5206\u306e 2 \u306f\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53ce\u7e2e\u306b\u7531\u6765\u3059\u308b\u3053\u3068\u306b\u306a\u308b. Ig \u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3068\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306f\u30bf\u30a4\u30c1\u30f3\u306b\u30c0\u30a4\u30ca\u30df\u30c3\u30af\u306a\u9069\u5fdc\u3092\u8a31\u3057\u3066\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u306e\u5e45\u5e83\u3044\u7bc4\u56f2\u306b\u308f\u305f\u308b\u5f35\u529b\u3092\u63d0\u4f9b\u3059\u308b\u3060\u3051\u3067\u306f\u306a\u304f\uff0c\u7b4b\u8089\u5185\u306e\u30a8\u30cd\u30eb\u30ae\u30fc\u84c4\u7a4d\u3068\u89e3\u653e\u3092\u5897\u3084\u3059\u3053\u3068\u306b\u3082\u306a\u308b\uff0e\u30a8\u30cd\u30eb\u30ae\u30fc\u84c4\u7a4d\u3068\u89e3\u653e\u306f\uff0c\u975e\u5e38\u306b\u7cbe\u5bc6\u306a\u529b\u306e\u7bc4\u56f2\u5185\u3067\u306e\u307f\u53ef\u80fd\u3067\u3042\u308a\uff0c\u305d\u3053\u3067\u306f\u86cb\u767d\u8cea\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3068\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304c\u8d77\u304d\u3066\u3044\u308b\uff0e\u6211\u3005\u306f\uff0c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u78ba\u7387\u3092\u307e\u305f\u3044\u3067\uff0c\u30bf\u30a4\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e\u5f3e\u6027\u30dd\u30c6\u30f3\u30b7\u30e3\u30eb\u306e\u84c4\u7a4d\u3092\u6d3b\u7528\u3059\u308b\u3088\u3046\u306a\u9577\u3055\u3068\u529b\u3067\u7b4b\u8089\u304c\u52d5\u4f5c\u3059\u308b\u3088\u3046\u8abf\u6574\u3055\u308c\u3066\u3044\u308b\u3068\u63d0\u6848\u3059\u308b\uff0e\u3055\u3089\u306b\uff0c\u7d30\u80de\u30b7\u30b0\u30ca\u30ea\u30f3\u30b0\u3092\u901a\u3058\u3066\u30bf\u30a4\u30c1\u30f3\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3092\u64cd\u4f5c\u3059\u308b\u3053\u3068\u3067\uff0c\u7b4b\u8089\u306f\u5f3e\u6027\u30a8\u30cd\u30eb\u30ae\u30fc\u84c4\u7a4d\u3092\u5909\u66f4\u3059\u308b\u3053\u3068\u304c\u3067\u304d\u308b\uff0e<\/p>\n<h3>The titin folding reaction can be biochemically tuned<\/h3>\n<p>\u3000\u30bf\u30a4\u30c1\u30f3\u306f\u7b4b\u8089\u306e\u7d14\u7c8b\u306a\u6a5f\u68b0\u7684\u86cb\u767d\u8cea\u3067\u3082\u306a\u3051\u308c\u3070\u69cb\u9020\u86cb\u767d\u8cea\u3067\u3082\u306a\u3044\uff0e\u305d\u308c\u306f\u7b4b\u8089\u306b\u304a\u3051\u308b\u5316\u5b66\u7684\u304b\u3064\u7269\u7406\u7684\u30b9\u30c8\u30ec\u30b9\u306e\u7a4d\u5206\u5668\u3067\u3082\u3042\u308a\uff0c\u30b7\u30b0\u30ca\u30eb\u7d4c\u8def\u306e\u4e00\u90e8\u3067\u3042\u308a\uff0c\u6700\u7d42\u7684\u306b\u306f\u907a\u4f1d\u5b50\u767a\u73fe\u306b\u5f71\u97ff\u3059\u308b\uff0e\u30bf\u30a4\u30c1\u30f3\u306e\u5f3e\u6027\u30bb\u30b0\u30e1\u30f3\u30c8\u306f\u7ffb\u8a33\u5f8c\u4fee\u98fe\u306e\u4e3b\u8981\u306a\u6a19\u7684\u3067\u3042\u308b\u3068\u793a\u3055\u308c\u305f\uff0e\u30bf\u30a4\u30c1\u30f3\u306e\u975e\u69cb\u9020\u90e8\u4f4d\u5185\u306e\u6b8b\u57fa\u306e\u30ea\u30f3\u9178\u5316\u306f\u30bf\u30a4\u30c1\u30f3\u306e\u5f3e\u6027\u306b\u304a\u3044\u3066\u306f\u7a4f\u3084\u304b\u306a\u5909\u5316\u3067\u3042\u308b\u3068\u793a\u3055\u308c\u305f\u4e00\u65b9\u3067 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC2764991\/\" rel=\"noopener noreferrer\" target=\"_blank\">97<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/23764881\" rel=\"noopener noreferrer\" target=\"_blank\">98<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC4701010\/\" rel=\"noopener noreferrer\" target=\"_blank\">99<\/a>), \u3055\u3089\u306b\u5287\u7684\u306a\u5909\u5316\u306f I-band \u306e Ig \u30c9\u30e1\u30a4\u30f3\u306e\u6a19\u7684\u5316\u306b\u3088\u308a\u9054\u6210\u3055\u308c\u308b\uff0e\u30de\u30a6\u30b9\u5fc3\u7b4b\u6897\u585e\u30e2\u30c7\u30eb\u306b\u304a\u3044\u3066\u306f\u865a\u8840\u518d\u704c\u6d41\u969c\u5bb3\u306b\u3088\u308b\u9178\u5316\u30b9\u30c8\u30ec\u30b9\u306f\u30b5\u30eb\u30b3\u30e1\u30a2\u86cb\u767d\u8cea\u306e S-\u30b0\u30eb\u30bf\u30c1\u30aa\u30cb\u30eb\u5316\u3092\u62db\u304d\uff0c\u305d\u308c\u306b\u306f\u975e\u5e38\u306b\u5de8\u5927\u306a\u5206\u5b50\u91cf\u306e\u86cb\u767d\u8cea\u304c\u542b\u307e\u308c\uff0c\u30bf\u30a4\u30c1\u30f3\u3067\u3042\u308b\u3068\u4e3b\u5f35\u3055\u308c\u3066\u3044\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC3659404\/\" rel=\"noopener noreferrer\" target=\"_blank\">100<\/a>). \u30bf\u30a4\u30c1\u30f3 I-band \u306e Ig \u30c9\u30e1\u30a4\u30f3\u306f\u4e0d\u91e3\u308a\u5408\u3044\u306b\u9ad8\u3044\u30b7\u30b9\u30c6\u30a4\u30f3\u6b8b\u57fa\u3092\u542b\u6709\u3057\u3066\u3044\u308b\uff0e\u3057\u304b\u3057\uff0c\u3053\u308c\u3089\u5927\u534a\u306e\u30b7\u30b9\u30c6\u30a4\u30f3\u5074\u9396\u306f\u8b0e\u3081\u3044\u3066\u3044\u308b\uff0e\u305d\u308c\u3089\u306f Ig \u6298\u308a\u7573\u307f\u306e\u758e\u6c34\u6027\u30b3\u30a2\u306b\u57cb\u3081\u8fbc\u307e\u308c\u3066\u304a\u308a\uff0c\u6eb6\u5a92\u306b\u30a2\u30af\u30bb\u30b9\u3067\u304d\u306a\u3044\u3053\u3068\u3092\u610f\u5473\u3057\u3066\u3044\u308b\uff0e\u305d\u308c\u3089\u306f Ig \u30c9\u30e1\u30a4\u30f3\u304c\u4f38\u5c55\u3057\u59cb\u3081\u306a\u3044\u9650\u308a, S-\u30b0\u30eb\u30bf\u30c1\u30aa\u30cb\u30eb\u5316\u3067\u304d\u306a\u3044. Alegre-Cebollada \u3068\u305d\u306e\u4ef2\u9593\u304c\u793a\u3057\u305f\u3068\u3053\u308d\u3067\u306f\uff0c\u5358\u5206\u5b50 force-clamp AFM \u3092\u7528\u3044\u3066\uff0c\u5358\u5206\u5b50\u30bf\u30a4\u30c1\u30f3 I91 \u30c9\u30e1\u30a4\u30f3\u306e S-\u30b0\u30eb\u30bf\u30c1\u30aa\u30cb\u30eb\u5316\u304c\u529b\u306b\u3088\u308a\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3057\u305f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/24630725\" rel=\"noopener noreferrer\" target=\"_blank\">78<\/a>). \u3042\u308b\u91cd\u5927\u306a\u767a\u898b\u304c\u3042\u3063\u305f. S-\u30b0\u30eb\u30bf\u30c1\u30aa\u30cb\u30eb\u5316\u304c I91 \u30c9\u30e1\u30a4\u30f3\u306e\u758e\u6c34\u6027\u30b3\u30a2\u5185\u90e8\u306b\u597d\u307e\u3057\u304f\u306a\u3044\u7acb\u4f53\u69cb\u9020\u3092\u4f5c\u308a\u51fa\u3057\uff0c\u305d\u308c\u304c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3092\u963b\u5bb3\u3057\u305f\u306e\u3067\u3042\u308b\uff0e\u3086\u3048\u306b\u305d\u308c\u306f\u9178\u5316\u578b\u30b0\u30eb\u30bf\u30c1\u30aa\u30f3\u306f\u5358\u4e00\u5fc3\u7b4b\u5185\u306e\u53d7\u52d5\u7684\u5f35\u529b\u3092\u5927\u5e45\u306b\u6e1b\u5c11\u3055\u305b\u308b\u3053\u3068\u304c\u3067\u304d\u308b\u3068\u8a3c\u660e\u3059\u308b\u3053\u3068\u306b\u306a\u3063\u305f\u304c\uff0c\u5fc3\u7b4b\u7d30\u80de\u304c\u4f38\u5c55\u3057\u3066 Ig \u30c9\u30e1\u30a4\u30f3\u3092\u52d5\u54e1\u3057\uff0c\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u72b6\u614b\u306b\u306a\u3063\u305f\u5f8c\u306b\u9650\u3089\u308c\u308b\uff0e\u30ea\u30f3\u9178\u5316\u306e\u3088\u3046\u306b, S-\u30b0\u30eb\u30bf\u30c1\u30aa\u30cb\u30eb\u5316\u306f\u53ef\u9006\u7684\u3067\u3042\u308a\uff0c\u7b4b\u7d44\u7e54\u5185\u90e8\u3067\u9175\u7d20\u7684\u306a\u5207\u65ad\u3084\u5909\u6027\u3092\u901a\u3058\u3066\u30b0\u30eb\u30bf\u30c1\u30aa\u30f3\u3092\u6e1b\u5c11\u3055\u305b\u308b\u3053\u3068\u3055\u3048\u3059\u308b\uff0e\u3057\u304b\u3057\uff0c\u30ea\u30f3\u9178\u5316\u306f\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u306e\u786c\u5ea6\u307e\u305f\u306f Kuhn \u9577\u306b\u304a\u3044\u3066\u5c0f\u3055\u306a\u5909\u5316\u3057\u304b\u3082\u305f\u3089\u3055\u306a\u3044\u4e00\u65b9\u3067, Ig \u6298\u308a\u7573\u307f\u306e S-\u30b0\u30eb\u30bf\u30c1\u30aa\u30cb\u30eb\u5316\u3084\u4ed6\u306e\u975e\u7279\u7570\u7684\u9178\u5316\u306f\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53cd\u5fdc\u306b\u5927\u898f\u6a21\u306a\u9589\u585e\u3092\u3082\u305f\u3089\u3059\u3053\u3068\u304c\u3067\u304d\u308b\uff0e\u4eee\u306b\u5168\u30bf\u30a4\u30c1\u30f3\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304c\u963b\u5bb3\u3055\u308c\u305f\u3089\uff0c\u305d\u306e\u5f3e\u529b\u7279\u6027\u306f <strong>Figure 4B<\/strong> \u306b\u793a\u3057\u305f\u9752\u306e\u70b9\u7dda\u306e\u3088\u3046\u306b\u3075\u308b\u307e\u3046\u3060\u308d\u3046\uff0e\u53d7\u52d5\u7684\u5f35\u529b\u306e\u55aa\u5931\u306f\u4ed6\u306e\u30b5\u30eb\u30b3\u30e1\u30a2\u86cb\u767d\u8cea\u306b\u3088\u308a\u611f\u77e5\u3055\u308c\u308b\u3068\u3053\u308d\u3068\u306a\u308a\uff0c\u7d30\u80de\u30b7\u30b0\u30ca\u30ea\u30f3\u30b0\u306b\u5909\u63db\u3055\u308c\u308b\u7d50\u679c\u3068\u306a\u308b\u3068\u4e88\u60f3\u3055\u308c\u308b\uff0e\u4ed6\u306e\u4e3b\u8981\u306a\u30bf\u30a4\u30c1\u30f3\u306e\u9178\u5316\u7684\u4fee\u98fe\u306f\uff0c\u5206\u5b50\u5185\u306e\u30b8\u30b9\u30eb\u30d5\u30a3\u30c9\u7d50\u5408\u3067\u3042\u308b\uff0e\u305d\u306e\u30b8\u30b9\u30eb\u30d5\u30a3\u30c9\u7d50\u5408\u306f\uff0c\u6a5f\u68b0\u7684\u5b89\u5b9a\u6027\u3084\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u7387\u3078\u306e\u5927\u5e45\u306a\u5f71\u97ff\u306a\u3057\u306b, Ig \u30c9\u30e1\u30a4\u30f3\u304c\u6298\u308a\u305f\u305f\u307e\u308c\u308b\u7b87\u6240\u3067\u306e\uff0c\u5927\u5e45\u306a\u529b\u306e\u5897\u52a0\u3092\u53ef\u80fd\u3068\u3059\u308b (<a href=\"https:\/\/www.cell.com\/biophysj\/abstract\/S0006-3495(16)33473-7?code=cell-site\" rel=\"noopener noreferrer\" target=\"_blank\">80<\/a>). \u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u78ba\u7387\u30b7\u30d5\u30c8\u306e\u7d50\u679c\uff0c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53cd\u5fdc\u304c\u53ce\u7e2e\u6642\u306b\u3088\u308a\u9ad8\u3044\u529b\u3092\u767a\u751f\u3055\u305b\u308b\u7d50\u679c\u3068\u306a\u308b (<strong>Figure 4B<\/strong>, \u91d1\u8272\u66f2\u7dda). \u9178\u5316\u30b9\u30c8\u30ec\u30b9\u306b\u52a0\u3048\u3066\uff0c\u30a2\u30b7\u30c9\u30fc\u30b7\u30b9\u304c\u7b4b\u8089\u306e\u6052\u5e38\u6027\u306b\u5bfe\u3059\u308b\u4e00\u822c\u7684\u306a\u6442\u52d5\u3067\u3042\u308a\uff0c\u86cb\u767d\u8cea\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306b\u76f4\u63a5\u7684\u306a\u7d50\u679c\u3092\u3082\u305f\u3089\u3059 (<a href=\"https:\/\/www.cell.com\/biophysj\/abstract\/S0006-3495(16)31293-0?code=cell-site\" rel=\"noopener noreferrer\" target=\"_blank\">101<\/a>). pH \u306e\u5909\u5316\u306f\u591a\u304f\u306e\u5074\u9396\u306e\u30d7\u30ed\u30c8\u30f3\u5316\u3092\u5909\u5316\u3055\u305b\uff0c\u305d\u308c\u306f\u9806\u756a\u306b\u9759\u96fb\u6c17\u529b\u304a\u3088\u3073\u758e\u6c34\u529b\u3092\u5909\u5316\u3055\u305b\uff0c\u86cb\u767d\u8cea\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3092\u99c6\u52d5\u3057\uff0c\u6a5f\u68b0\u7684\u5b89\u5b9a\u6027\u3092\u63d0\u4f9b\u3059\u308b\uff0e<strong>\u30a2\u30b7\u30c9\u30fc\u30b7\u30b9\u306f\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3092\u963b\u5bb3\u3059\u308b\u304b\u30df\u30b9\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3092\u8a98\u5c0e\u3057\uff0c\u305d\u308c\u306f\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53ce\u7e2e\u306e\u62e1\u5927\u3092\u6e1b\u5f31\u3055\u305b\u3066\u7b4b\u5f3e\u6027\u306e\u5909\u5316\u3092\u3082\u305f\u3089\u3059<\/strong> (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC3897184\/\" rel=\"noopener noreferrer\" target=\"_blank\">102<\/a>). \u65b0\u305f\u306a\u8996\u70b9\u306f\u6b21\u306e\u3088\u3046\u306a\u3082\u306e\u3067\u3082\u3042\u308b\uff0e\u3064\u307e\u308a\uff0c\u7b4b\u7279\u7570\u7684\u30b7\u30e3\u30da\u30ed\u30f3\u306f\u7269\u7406\u7684\u4f38\u5c55\u306b\u5bfe\u3057\u3066\u30bf\u30a4\u30c1\u30f3\u306e I-band \u30bb\u30b0\u30e1\u30f3\u30c8\u306b\u52d5\u54e1\u3055\u308c\u308b\u53ef\u80fd\u6027\u304c\u3042\u308a (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/14676215\" rel=\"noopener noreferrer\" target=\"_blank\">103<\/a>), \u3042\u308b\u7a2e\u306e\u7b4b\u75be\u60a3\u306b\u304a\u3044\u3066\u30ec\u30d9\u30eb\u304c\u3072\u3069\u304f\u5909\u66f4\u3055\u308c\u308b\u5206\u5b50\u30b7\u30e3\u30da\u30ed\u30f3\u304c\u3042\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/24895218\" rel=\"noopener noreferrer\" target=\"_blank\">104<\/a>) \u3068\u3044\u3046\u3053\u3068\u3067\u3042\u308b\uff0e\u3053\u308c\u3089\u306e\u30b7\u30e3\u30da\u30ed\u30f3\u306e\u3044\u304f\u3064\u304b\uff0c\u4f8b\u3048\u3070 &alpha;B-\u30af\u30ea\u30b9\u30bf\u30ea\u30f3\u306f\uff0c\u6298\u308a\u305f\u305f\u307e\u308c\u305f Ig \u30c9\u30e1\u30a4\u30f3\u306b\u76f4\u63a5\u7d50\u5408\u3059\u308b\u3053\u3068\u304c\u53ef\u80fd\u3067\u3042\u308a\uff0c\u305d\u308c\u3089\u3092\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306b\u5bfe\u3057\u3066\u3088\u308a\u62b5\u6297\u6027\u306b\u3059\u308b (<strong>Figure 4B<\/strong>, \u8d64\u66f2\u7dda) (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/14676215\" rel=\"noopener noreferrer\" target=\"_blank\">103<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/19282282\" rel=\"noopener noreferrer\" target=\"_blank\">105<\/a>). \u5358\u5206\u5b50\u539f\u5b50\u9593\u529b\u5206\u5149\u6cd5\u306f\u6700\u8fd1\u5206\u5b50\u30b7\u30e3\u30da\u30ed\u30f3\u306e\u53ef\u80fd\u6027\u3092\u660e\u3089\u304b\u306b\u3057\u59cb\u3081\u3066\u304a\u308a\uff0c\u51dd\u96c6\u3092\u9632\u3050\u3060\u3051\u3067\u306a\u304f\uff0c\u3042\u308b\u7a2e\u306e\u86cb\u767d\u8cea\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53cd\u5fdc\u3092\u52a0\u901f\u3057\u3066\u3044\u308b\u3068\u3055\u308c\u308b (Haldar, \u672a\u516c\u958b\u30c7\u30fc\u30bf). \u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u529b\u5b66\u306b\u304a\u3051\u308b\u5897\u52a0\u306f\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53cd\u5fdc\u304b\u3089\u306e\u51fa\u529b\u5897\u52a0\u3068\u540c\u7fa9\u3067\u3042\u308a\uff0c\u3086\u3048\u306b 3 \u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u30e2\u30c7\u30eb\u306b\u5f93\u3063\u3066\u7b4b\u53ce\u7e2e\u306b\u76f4\u63a5\u5f71\u97ff\u3092\u4e0e\u3048\u308b\u3060\u308d\u3046\uff0e<\/p>\n<h3>A rechargeable gravitational battery<\/h3>\n<p>\u3000\u3053\u3053\u3067\u7b4b\u53ce\u7e2e\u306e\u8996\u70b9\u3092\uff0c\u5b89\u9759\u6642\u306e\u5358\u72ec\u30e2\u30fc\u30bf\u30fc\u6d3b\u6027\u304b\u3089\uff0c\u4f38\u5c55\u529b\u306b\u62b5\u6297\u3057\u306a\u304c\u3089\u6a5f\u68b0\u7684\u306b\u4f38\u5c55\u3055\u308c\u305f\u86cb\u767d\u8cea\u304c\u6298\u308a\u305f\u305f\u307e\u308c\u3066\u77ed\u7e2e\u3059\u308b\u53ef\u80fd\u6027\u306b\u79fb\u305d\u3046\uff0e\u98df\u7269\u6442\u53d6\u3092\u901a\u3058\u305f\u592a\u967d\u5149\u304b\u3089 ATP \u3078\u306e\u30a8\u30cd\u30eb\u30ae\u30fc\u5909\u63db\u3068\u52d5\u7269\u7d30\u80de\u306e\u4ee3\u8b1d\u6a5f\u69cb\u3068\u306f\uff0c\u7b4b\u53ce\u7e2e\u6642\u306b\u5229\u7528\u3055\u308c\u308b\u5316\u5b66\u30a8\u30cd\u30eb\u30ae\u30fc\u306e\u6e90\u3067\u3042\u308b\uff0e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3057\u305f\u86cb\u767d\u8cea\u306b\u3088\u308a\u63d0\u4f9b\u3055\u308c\u308b\u30a8\u30cd\u30eb\u30ae\u30fc\u3068\u306f\u306a\u3093\u3060\u308d\u3046\uff1f\u77ed\u8ddd\u96e2\u306e\u758e\u6c34\u6027\u5d29\u58ca\u3068\u7d44\u307f\u5408\u308f\u3055\u308c\u305f\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc\u53cd\u52d5\u304c\u86cb\u767d\u8cea\u6298\u308a\u305f\u305f\u307f\u306e\u529b\u3092\u4f9b\u4e0e\u3059\u308b\u3068\u6211\u3005\u306f\u77e5\u3063\u3066\u3044\u308b\uff0e\u3060\u304c\uff0c\u86cb\u767d\u8cea\u304c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u4ed5\u4e8b\u3092\u306a\u3059\u3053\u3068\u306f\u660e\u3089\u304b\u3060\u304c\uff0c\u305d\u306e\u524d\u306b\u6a5f\u68b0\u7684\u306b\u4f38\u5c55\u3055\u308c\u3066\u3044\u306a\u3051\u308c\u3070\u306a\u3089\u306a\u3044\uff0e\u591a\u304f\u306e\u6298\u308a\u305f\u305f\u307f\u53ef\u80fd\u306a Ig \u30c9\u30e1\u30a4\u30f3\u3078\u306e\u30bf\u30a4\u30c1\u30f3\u30dd\u30ea\u30da\u30d7\u30c1\u30c9\u306e\u5206\u7bc0\u5316\u306f\u3086\u3048\u306b\uff0c\u305f\u3060\u306e\u5f3e\u6027\u53ce\u7e2e\u529b\u306e\u307f\u3092\u6709\u3059\u308b\u5358\u7d14\u306a\u3070\u306d\u304b\u3089 Ig \u30c9\u30e1\u30a4\u30f3\u3092\u5206\u5316\u3055\u305b\u308b\uff0e\u3053\u306e\u65b0\u3057\u3044\u6a5f\u69cb\u306f\u9b45\u529b\u7684\u3067\u3042\u308b\uff0e\u3068\u3044\u3046\u306e\u3082\uff0c\u5316\u5b66\u30a8\u30cd\u30eb\u30ae\u30fc\u306b\u52a0\u3048\u3066\uff0c\u7b4b\u8089\u306f\u4ed6\u306e\u57fa\u672c\u7684\u306a\u529b\u3092\u5229\u7528\u3059\u308b\u304b\u3089\u3067\u3042\u308b\uff0e\u91cd\u529b\u3068\u6163\u6027\u306f\uff0c\u52d5\u7269\u304c\u30e9\u30f3\u30cb\u30f3\u30b0\u306a\u3069\u306e\u904b\u52d5\u4e2d\u306b\u7b4b\u8089\u306e\u30bf\u30a4\u30c1\u30f3\u3092\u4f38\u5c55\u3057\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3055\u305b\uff0c\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc\u529b\u3068\u758e\u6c34\u529b\u306f\u8ca0\u8377\u6e1b\u5c11\u4e2d\u306b\u4f38\u5c55\u3057\u305f\u86cb\u767d\u8cea\u306e\u5d29\u58ca\u3068\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3092\u99c6\u52d5\u3059\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/15017000\" rel=\"noopener noreferrer\" target=\"_blank\">34<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC3785139\/\" rel=\"noopener noreferrer\" target=\"_blank\">106<\/a>). \u7b4b\u30b5\u30eb\u30b3\u30e1\u30a2\u306f\u3086\u3048\u306b\uff0c\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u304c\u4f38\u5c55\u3057\u305f\u5f62\u614b\u3067\u74b0\u5883\u304b\u3089\u306e\u6a5f\u68b0\u7684\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u5438\u53ce\u3057\u84c4\u7a4d\u3059\u308b\u306e\u306b\u6700\u9069\u306b\u8a2d\u8a08\u3055\u308c\u3066\u304a\u308a\uff0c\u3053\u306e\u6a5f\u68b0\u7684\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u5316\u5b66\u7684\u306b\u6d3b\u6027\u5316\u3057\u305f\u30df\u30aa\u30b7\u30f3 II \u578b\u30e2\u30fc\u30bf\u30fc\u306b\u3088\u308a\u30c8\u30ea\u30ac\u30fc\u3055\u308c\u305f\u6642\u306b\u653e\u51fa\u3059\u308b (<strong>Figure 3B<\/strong>). \u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u304c\u524d\u9032\u3067\u304d\u306a\u3044\u307b\u3069\u9ad8\u3044\u8ca0\u8377\u304c\u304b\u304b\u3063\u3066\u3044\u308b\u6642 (6 pN) \u3067\u3082\u30bf\u30a4\u30c1\u30f3\u306f\u5bb9\u6613\u306b\u6298\u308a\u305f\u305f\u307e\u308c\u308b\uff0e\u3086\u3048\u306b\uff0c\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306f\u5931\u901f\u3057\u305f\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u3092\u89e3\u653e\u3057\uff0c\u305d\u308c\u3089\u304c\u524d\u9032\u3059\u308b\u901f\u5ea6\u3092\u8a2d\u5b9a\u3057\uff0c\u52b9\u7387\u7684\u306a\u7b4b\u53ce\u7e2e\u3092\u8a31\u3059\uff0e<\/p>\n<p>\u3000\u7b4b\u53ce\u7e2e\u306b\u304a\u3051\u308b\u30bf\u30a4\u30c1\u30f3\u306e\u3053\u306e\u6d3b\u52d5\u7684\u306a\u5f79\u5272\u306f\uff0c\u30d0\u30c3\u30c6\u30ea\u30fc\u306e\u30a2\u30ca\u30ed\u30b8\u30fc\u3067\u7406\u89e3\u3067\u304d\u308b (<strong>Figure 5A<\/strong> \u53c2\u7167). \u4f38\u5c55\u30d5\u30a7\u30fc\u30ba\u3067\u306e Ig \u30c9\u30e1\u30a4\u30f3\u306e\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306f\u5916\u529b\u304c 10 pN \u3092\u8d85\u3048\u308b\u6642\u306b\u300c\u30d0\u30c3\u30c6\u30ea\u30fc\u300d\u306b\u53ce\u7e2e\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u5145\u96fb\u3059\u308b\uff0e\u30e2\u30fc\u30bf\u30fc\u304c\u53c2\u52a0\u3057\u305f\u6642\u306b\u5916\u529b\u304c\u5f31\u307e\u308b\u306b\u3064\u308c\u3066 Ig \u30c9\u30e1\u30a4\u30f3\u518d\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306f\u84c4\u7a4d\u3057\u305f\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u653e\u51fa\u3057\uff0c\u5f15\u304d\u7d9a\u3044\u3066\u7b4b\u53ce\u7e2e\u304c\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306b\u3088\u308a\u306a\u3055\u308c\u308b\u4ed5\u4e8b\u3068\u5171\u306b\u8d77\u304d\u308b\uff0e\u305d\u308c\u3086\u3048\uff0c\u30bf\u30a4\u30c1\u30f3\u3068\u30df\u30aa\u30b7\u30f3\u306f\u4f38\u5c55\u30d5\u30a7\u30fc\u30ba\u3067\u30d0\u30c3\u30c6\u30ea\u30fc\u304c\u5145\u96fb\u3055\u308c\u308b (<strong>Figure 5A<\/strong>, \u9ed2\u7dda) \u30cf\u30a4\u30d6\u30ea\u30c3\u30c9\u30b7\u30b9\u30c6\u30e0\u306e\u4e8c\u3064\u306e\u8981\u7d20\u3067\u3042\u308a, ATP \u5316\u5b66\u30a8\u30cd\u30eb\u30ae\u30fc\u306b\u3088\u308a\u99c6\u52d5\u3055\u308c\u308b\u30df\u30aa\u30b7\u30f3\u982d\u90e8\u304c\u53c2\u52a0\u3057\u305f\u5f8c\u306b\u306e\u307f\uff0c\u3053\u306e\u84c4\u7a4d\u30a8\u30cd\u30eb\u30ae\u30fc\u306f\u653e\u51fa\u3055\u308c\u308b (<strong>Figure 5A<\/strong>, \u8d64\u7dda). \u306b\u3082\u304b\u304b\u308f\u3089\u305a\uff0c\u53ce\u7e2e\u30a8\u30cd\u30eb\u30ae\u30fc\u304c\u84c4\u7a4d\u3055\u308c\u653e\u51fa\u3055\u308c\u308b\u78ba\u7387\u306f\u3080\u3057\u308d\u7570\u306a\u308a, Figure 5A \u306b\u793a\u3059\u3088\u3046\u306b\u7570\u306a\u308b\u659c\u7dda\u306b\u53cd\u6620\u3055\u308c\u308b. 10 pN \u3067\u306e\u30bf\u30a4\u30c1\u30f3\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306f 1.5 &times; 10<sup>-4<\/sup> \u307e\u3067\u306e\u78ba\u7387\u3067\u767a\u751f\u3057 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/28394039\" rel=\"noopener noreferrer\" target=\"_blank\">65<\/a>), \u3086\u3048\u306b\u30bf\u30a4\u30c1\u30f3\u306f\u5c11\u306a\u3044\u529b\u3067\u53ce\u7e2e\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u84c4\u7a4d\u3057\uff0c\u305d\u306e\u529b\u306f 0.02 zW \u3068\u63a8\u5b9a\u3055\u308c\u3066\u3044\u308b\uff0810 pN (140 zJ) \u3067\u306e Ig \u30c9\u30e1\u30a4\u30f3\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u4ed5\u4e8b\u3092\u89b3\u5bdf\u3059\u308b\u3068 2 \u500d\u306e\u78ba\u7387\u3067\u3042\u308b\uff09. \u3057\u304b\u3057\uff0c\u518d\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306f\u66f4\u306b\u9ad8\u3044\u78ba\u7387\u3067\u767a\u751f\u3057\uff0c\u30c9\u30e1\u30a4\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306f 7 \u79d2\u9593\u3067 6 pN \u3067\u3042\u308a\uff0c\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u518d\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3067\u653e\u51fa\u3055\u308c\u308b\u305d\u306e\u529b\u306f\u7d04 100 \u500d\u3082\u5927\u304d\u304f, 6.5 zW \u306b\u3082\u9054\u3059\u308b\u3068\u63a8\u5b9a\u3055\u308c\u3066\u3044\u308b\uff0e\u305d\u306e\u529b\u306e\u95a2\u4fc2\u306f\u7570\u306a\u308b\u30c9\u30e1\u30a4\u30f3\u306b\u3088\u308b\u3082\u306e\u3067\u3042\u308d\u3046\u304c\uff0c\u751f\u4f53\u5185\u3067\u306e\u529b\u306e\u653e\u51fa\u306f\u3088\u308a\u8907\u96d1\u3067\u3042\u308b\u306b\u9055\u3044\u306a\u3044\uff0e\u52d5\u7269\u306e\u904b\u52d5\u306b\u304a\u3044\u3066\u7b4b\u8089\u304c\u3069\u3046\u3084\u3063\u3066\u52d5\u4f5c\u3059\u308b\u304b\u3092\u6975\u7aef\u306b\u5358\u7d14\u5316\u3059\u308b\u3068\uff0c\u30d2\u30e9\u30e1\u7b4b\u306e\u3088\u3046\u306a\u811a\u306e\u7b4b\u8089\u306f\u305d\u306e\u9759\u6b62\u72b6\u614b\u304b\u3089\u4f38\u5c55\u4f4d\u7f6e\u306b\u79fb\u52d5\u3057\uff0c\u9ad8\u3044\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u30bf\u30a4\u30c1\u30f3\u306b\u5909\u63db\u3059\u308b (<strong>Figure 5A<\/strong>, \u4e0b\u90e8\u30d1\u30cd\u30eb 1 \u304b\u3089 2). \u5f8c\u306b\uff0c\u5206\u5b50\u30e2\u30fc\u30bf\u30fc\u3068 ATP \u52a0\u6c34\u5206\u89e3\u304c\u53c2\u52a0\u3057\u305f\u5f8c\u306b\u30bf\u30a4\u30c1\u30f3\u306b\u5909\u63db\u3055\u308c\u308b\u529b\u306f\u6e1b\u5c11\u3057\uff0c\u30bf\u30a4\u30c1\u30f3\u304b\u3089\u6a5f\u68b0\u7684\u4ed5\u4e8b\u304c\u653e\u51fa\u3055\u308c\u3066\u7b4b\u8089\u304c\u53ce\u7e2e\u3059\u308b (<strong>Figure 5A<\/strong>, \u4e0b\u90e8\u30d1\u30cd\u30eb 2 \u304b\u3089 3). \u30bf\u30a4\u30c1\u30f3\u30c9\u30e1\u30a4\u30f3\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u30fb\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u4e0a\u306e\u30a8\u30cd\u30eb\u30ae\u30fc\u5909\u63db\u306e\u610f\u5473\u304b\u3089\u306f\uff0c\u4e0a\u8a18\u3067\u63d0\u6848\u3057\u305f\u84c4\u7a4d\u30fb\u653e\u51fa\u30b5\u30a4\u30af\u30eb\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u306e\u81ea\u7531\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u80cc\u666f\u306b\u3057\u305f\u878d\u5408\u30d7\u30ed\u30bb\u30b9\u3068\u3057\u3066\u63cf\u304f\u3053\u3068\u304c\u3067\u304d\u308b (<strong>Figure 5B<\/strong>). 12 pN \u3067\u306e\uff08\u7dd1\u8272\u80cc\u666f\uff09\u84c4\u7a4d\u30d5\u30a7\u30fc\u30ba\u4e2d\u305d\u306e\u30a8\u30cd\u30eb\u30ae\u30fc\u80cc\u666f\u306f\u4f38\u5c55\u72b6\u614b\u306b\u50be\u3044\u3066\u304a\u308a\uff08\u5927\u4f38\u5c55\uff09\uff0c\u305d\u306e\u305f\u3081\u30bf\u30a4\u30c1\u30f3\u30c9\u30e1\u30a4\u30f3\u304c\u4f38\u5c55\u3057\uff0c\u9806\u6b21\u81ea\u7531\u30a8\u30cd\u30eb\u30ae\u30fc\u969c\u58c1\u3092\u8d8a\u3048\u308b\u306e\u3092\u9858\u3063\u3066\u3044\u308b\uff0e\u5916\u529b\u304c 6 pN \u306b\u307e\u3067\u6e1b\u5c11\u3059\u308b\u3068\uff08\u8d64\u8272\u80cc\u666f\uff09\uff0c\u305d\u306e\u5206\u5b50\u306f\u518d\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306b\u3088\u308a\u77ed\u7e2e\u3057\uff0c\u305d\u306e\u30c9\u30e1\u30a4\u30f3\u306f\u53cd\u5bfe\u65b9\u5411\u306b\u969c\u58c1\u3092\u9858\u3044\uff0c\u3086\u3048\u306b\u53ce\u7e2e\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u653e\u51fa\u3059\u308b\uff0e<\/p>\n<figure id=\"attachment_3692\" aria-describedby=\"caption-attachment-3692\" style=\"width: 525px\" class=\"wp-caption aligncenter\"><a href=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure5.jpg\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure5-971x1024.jpg\" alt=\"\" width=\"525\" height=\"554\" class=\"size-large wp-image-3692\" srcset=\"https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure5-971x1024.jpg 971w, https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure5-285x300.jpg 285w, https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure5-768x810.jpg 768w, https:\/\/www.muscle-hypertrophy.com\/wp-content\/uploads\/2018\/08\/TitinFoldingFigure5.jpg 1415w\" sizes=\"auto, (max-width: 525px) 100vw, 525px\" \/><\/a><figcaption id=\"caption-attachment-3692\" class=\"wp-caption-text\"><strong>Figure 5. \u30cf\u30a4\u30d6\u30ea\u30c3\u30c9\u30e1\u30ab\u30cb\u30ba\u30e0\u3068\u3057\u3066\u306e\u7b4b\u53ce\u7e2e\u30e2\u30c7\u30eb<\/strong><br \/> <strong>(A)<\/strong> \u7b4b\u53ce\u7e2e\u306b\u304a\u3051\u308b\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u30fb\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u5f79\u5272\uff08\u30d1\u30cd\u30eb\u4e0a\u90e8\uff09\uff0e \u30bf\u30a4\u30c1\u30f3\u30c9\u30e1\u30a4\u30f3\u306f\u300c\u30d0\u30c3\u30c6\u30ea\u30fc\u300d\u3068\u3057\u3066\u6a5f\u80fd\u3057\uff0c\u53ce\u7e2e\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306b\u3088\u308a\u84c4\u3048\uff0c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u77ed\u7e2e\u6642\u306b\u4f9b\u7d66\u3059\u308b\uff0e\u7b4b\u8089\u304c\u4f38\u5c55\u3059\u308b\u6642\uff0c\u6a5f\u68b0\u7684\u5f35\u529b\u304c\u5897\u52a0\u3057\uff0c\u30bf\u30a4\u30c1\u30f3\u30c9\u30e1\u30a4\u30f3\u304c\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c9\u3059\u308b\uff08\u84c4\u7a4d\u30d5\u30a7\u30fc\u30ba (1)-(2)). \u7b4b\u8089\u304c\u53ce\u7e2e\u3059\u308b\u3068\u3001\u30df\u30aa\u30b7\u30f3\u982d\u90e8\u304c\u5f93\u4e8b\u3057\u3066\u30bf\u30a4\u30c1\u30f3\u306b\u304b\u304b\u308b\u529b\u304c 6 pN \u306b\u4f4e\u4e0b\u3057, Ig \u30c9\u30e1\u30a4\u30f3\u304c\u518d\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3057\u3066\u77ed\u7e2e\u3057\uff0c\u53ce\u7e2e\u30a8\u30cd\u30eb\u30ae\u30fc\u304c\u4f9b\u7d66\u3055\u308c\u3066\u7b4b\u53ce\u7e2e\u3092\u88dc\u52a9\u3059\u308b (3)\uff08\u4e0b\u90e8\u30d1\u30cd\u30eb\uff09\uff0e\u904b\u52d5\u4e2d\u306e\u52d5\u7269\u306e\u7b4b\u8089\u306e\u4f38\u5f35-\u53ce\u7e2e\u306e\u305f\u3081\u306e\u7c21\u7565\u5316\u3055\u308c\u305f\u30e2\u30c7\u30eb\uff0c\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304a\u3088\u3073\u518d\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u6642\u306e\u4e09\u76f8\u306e\u30bf\u30a4\u30c1\u30f3\u3092\u793a\u3059\uff0e<strong>(B)<\/strong> \u30bf\u30a4\u30c1\u30f3\u88dc\u52a9\u306e\u7b4b\u53ce\u7e2e\u306e\u81ea\u7531\u30a8\u30cd\u30eb\u30ae\u30fc\u666f\u89b3\u89e3\u91c8\uff0e\u4f38\u5c55\u30d5\u30a7\u30fc\u30ba\u306b\u304a\u3044\u3066\u306f\u305d\u306e\u666f\u89b3\u306f\u5897\u52a0\u3059\u308b\u529b\u306b\u3088\u308a\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c9\u72b6\u614b\uff08\u3088\u308a\u9577\u3044\u9577\u3055\uff09\u306b\u50be\u304d (12 pN, \u9752\u8272),  \u30bf\u30a4\u30c1\u30f3\u306f\u53ce\u7e2e\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u305d\u306e\u30c9\u30e1\u30a4\u30f3\u306e\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306b\u3088\u308a\u84c4\u7a4d\u3059\u308b\uff0e\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u304c\u53c2\u52a0\u3059\u308b\u6642\uff0c\u30bf\u30a4\u30c1\u30f3\u306b\u304b\u304b\u308b\u8ca0\u8377\u306f 6 pN \u306b\u6e1b\u5c11\u3059\u308b\u305f\u3081\uff0c\u305d\u306e\u666f\u89b3\u306f\u6298\u308a\u305f\u305f\u307e\u308c\u305f\u72b6\u614b\u306b\u6709\u5229\u3068\u306a\u308a\uff0c\u81ea\u7531\u30a8\u30cd\u30eb\u30ae\u30fc\u969c\u58c1\u3092\u4e57\u308a\u8d8a\u3048\u3066\u30c9\u30e1\u30a4\u30f3\u3092\u6298\u308a\u305f\u305f\u3080\u3053\u3068\u304c\u53ef\u80fd\u3068\u306a\u308a\uff0c\u5206\u5b50\u5168\u9577\u3092\u77ed\u7e2e\u3057\u3066\u53ce\u7e2e\u306e\u4ed5\u4e8b\u3092\u63d0\u4f9b\u3059\u308b\u3053\u3068\u304c\u53ef\u80fd\u3068\u306a\u308b\uff0e<strong>(C)<\/strong>\u3053\u306e\u4eee\u8aac\u3092\u8a66\u9a13\u3059\u308b\u306b\u306f\uff0c\u907a\u4f1d\u7684\u306b\u30b3\u30fc\u30c9\u5316\u3055\u308c\u305f\u529b\u5b66\u30bb\u30f3\u30b5\u30fc\u3092\u30bf\u30a4\u30c1\u30f3\u914d\u5217\u306b\u7d44\u307f\u8fbc\u3080\u3053\u3068\u304c\u5fc5\u8981\u3067\u3042\u308b\uff0e\u3053\u306e\u3088\u3046\u306a\u8a2d\u8a08\u306e\u4e00\u3064\u306f\uff0c\u30a8\u30ad\u30bd\u30f3 I86 \u3068 I87 \u3068\u306e\u9593\u306b\u5c0e\u5165\u3055\u308c\u305f HaloTag \u304a\u3088\u3073 SNAP \u30bf\u30b0\u5171\u6709\u6a19\u8b58\u6280\u8853\u3092\u5229\u7528\u3057\u3066\u63d0\u4f9b\u3055\u308c\u308b\uff0e\u4eee\u8aac\u7684\u306a\u7dd1\u8272\u4f9b\u4e0e\u4f53\u86cd\u5149\u56e3\u304a\u3088\u3073\u8d64\u8272\u30a2\u30af\u30bb\u30d7\u30bf\u30fc\u86cd\u5149\u56e3\u3092\u7528\u3044\u3066\uff0c\u30d1\u30cd\u30eb (A) \u306b\u793a\u3055\u308c\u308b\u6b69\u307f\u3092\u901a\u3057\u3066\uff0c\u7b4b\u8089\u306e FRET \u6bd4\u306f\u6700\u5927\u304b\u3089\u6700\u5c0f\u306b\u5909\u5316\u3059\u308b\uff0e\u3053\u308c\u306f\u751f\u4f53\u5185\u3067\u306e\u30bf\u30a4\u30c1\u30f3 1 \u5206\u5b50\u3042\u305f\u308a\u306b\u304b\u304b\u308b\u529b\u3092\u76f4\u63a5\u8aad\u307f\u51fa\u3059\u3053\u3068\u3092\u53ef\u80fd\u3068\u3059\u308b\uff0e<\/figcaption><\/figure>\n<p>\u3000\u6211\u3005\u306e\u73fe\u5728\u306e\u89b3\u5bdf\u306e\u9650\u754c\u306f\u78c1\u6c17\u30d4\u30f3\u30bb\u30c3\u30c8\u3067\u63a2\u7d22\u3057\u305f 8 \u56de\u7e70\u308a\u8fd4\u3057 Ig \u30c9\u30e1\u30a4\u30f3\u306e 6 pN \u3067\u306e\u9045\u3044\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u7387\u3067\u3042\u308b (<strong>Figure 1B<\/strong>). \u5b9f\u969b\uff0c\u5358\u5206\u5b50\u306e\u89b3\u5bdf\u3092\u751f\u4f53\u7d44\u7e54\u306b\u5916\u633f\u3059\u308b\u6642\u306b\u306f\u6211\u3005\u306f\u614e\u91cd\u3067\u306a\u304f\u3066\u306f\u306a\u3089\u306a\u3044\uff0e\u3053\u308c\u3089\u306e\u89b3\u5bdf\u304c\u8907\u96d1\u306a\u30b5\u30eb\u30b3\u30e1\u30a2\u306e\u74b0\u5883\u306b\u304a\u3044\u3066\u3069\u306e\u3088\u3046\u306b\u518d\u751f\u3055\u308c\u308b\u306e\u304b\uff0c\u6211\u3005\u306f\u307e\u3060\u5b8c\u5168\u306b\u7406\u89e3\u3057\u3066\u3044\u308b\u308f\u3051\u3067\u306f\u306a\u3044\u304b\u3089\u3067\u3042\u308b\uff0e\u3057\u304b\u3057\uff0c\u3044\u304f\u3064\u304b\u306e\u5358\u7d14\u306a\u89b3\u5bdf\u306f\u4fdd\u8a3c\u3055\u308c\u3066\u3044\u308b\uff0e\u4f38\u5c55\u3057\u305f\u30c9\u30e1\u30a4\u30f3\u306b\u52a0\u3048\u3066\uff0c\u91ce\u751f\u578b\u306e\u30bf\u30a4\u30c1\u30f3\u306f\u5de8\u5927\u306a\u975e\u69cb\u9020\u30bb\u30b0\u30e1\u30f3\u30c8\u3092\u6709\u3057\u3066\u304a\u308a\uff0c\u4f8b\u3048\u3070 PEVK \u3084 N2B \u306a\u3069\u3060\u304c\uff0c\u3053\u308c\u3089\u306f\u30bf\u30a4\u30c1\u30f3\u304c\u30dd\u30ea\u30de\u30fc\u62e1\u6563\u306e\u9650\u754c\u901f\u5ea6\u3067\u30b5\u30eb\u30b3\u30e1\u30a2\u5185\u306e\u3044\u304b\u306a\u308b\u5916\u529b\u306b\u3082\u5e73\u8861\u3059\u308b\u3053\u3068\u3092\u8a31\u3057\u3066\u304a\u308a\uff0c\u305d\u306e\u901f\u5ea6\u306f\u3069\u3093\u306a\u6700\u901f\u306e\u7b4b\u53ce\u7e2e\u901f\u5ea6\u3088\u308a\u3082\u901f\u3044 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/22895787\" rel=\"noopener noreferrer\" target=\"_blank\">73<\/a>). \u5b9f\u969b\uff0c\u4f38\u5c55\u9577\u304b\u3089\u306e\u53d7\u52d5\u7684\u30b5\u30eb\u30b3\u30e1\u30a2\u77ed\u7e2e\u306f\u6700\u521d\u306f\u6025\u901f\u3067\u3042\u308a (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/11927678\" rel=\"noopener noreferrer\" target=\"_blank\">107<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/14563922\" rel=\"noopener noreferrer\" target=\"_blank\">108<\/a>), \u6700\u8fd1\u306e\u751f\u4f53\u5185\u7b4b\u9855\u5fae\u93e1\u3067\u306f\uff0c\u30b5\u30eb\u30b3\u30e1\u30a2\u306f\u30bf\u30a4\u30c1\u30f3\u304c\u6570 pN \u306e\u69cb\u6210\u7684\u306a\u529b\u306e\u4e0b\u3067\u306e\u9577\u3055\u306b\u304a\u3044\u3066\u52d5\u4f5c\u3059\u308b\u3088\u3046\u3067\u3042\u308b\uff0e\u3053\u308c\u306e\u3055\u3089\u306b\u793a\u5506\u3059\u308b\u3068\u3053\u308d\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u306f\u5f1b\u7de9\u3059\u308b\u306e\u3067\u306f\u306a\u3044\u3068\u3044\u3046\u3053\u3068\u3067\u3042\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/18600262\" rel=\"noopener noreferrer\" target=\"_blank\">90<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC5858182\/\" rel=\"noopener noreferrer\" target=\"_blank\">92<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC5920519\/\" rel=\"noopener noreferrer\" target=\"_blank\">93<\/a>). \u3088\u308a\u91cd\u8981\u306a\u3053\u3068\u306f\uff0c\u77ed\u3044\uff0c\u5358\u5206\u5b50\u5b9f\u9a13\u3067\u3088\u304f\u7528\u3044\u3089\u308c\u308b 8 \u56de\u7e70\u308a\u8fd4\u3057 Ig \u306e\u64cd\u4f5c\u3055\u308c\u305f\u86cb\u767d\u8cea\u3068\u306f\u5bfe\u7167\u7684\u306b\uff0c\u91ce\u751f\u578b\u30bf\u30a4\u30c1\u30f3\u306e\u62e1\u5f35\u53ef\u80fd\u306a\u30bb\u30b0\u30e1\u30f3\u30c8\u306f 50-100 \u306e\u30a2\u30a4\u30bd\u30d5\u30a9\u30fc\u30e0\u4f9d\u5b58\u6027 Ig \u30c9\u30e1\u30a4\u30f3\u3092\u542b\u6709\u3057\u3066\u304a\u308a\uff0c\u5916\u529b\u6e1b\u5c11\u306b\u4f34\u3044\u521d\u671f\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u7387\u304c\u6025\u6fc0\u306b\u52a0\u901f\u3059\u308b\uff0e\u751f\u4f53\u5185\u3067\u89b3\u6e2c\u3055\u308c\u308b\u76f8\u5bfe\u7684\u306b\u77ed\u3044\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u306e\u5909\u5316\u3092\u8d85\u3048\uff0c\u4f8b\u3048\u3070\u5fc3\u7b4b\u7d44\u7e54\u306b\u304a\u3051\u308b\u30b5\u30eb\u30b3\u30e1\u30a2\u534a\u5206\u3042\u305f\u308a 50 nm \u307e\u3067\u306a\u3069\uff0c\u305d\u306e\u521d\u671f\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u7387\u306f\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u751f\u7406\u5b66\u7684\u306b\u95a2\u9023\u3059\u308b\u51fa\u529b\u306e\u3088\u308a\u826f\u3044\u8a08\u6e2c\u3092\u53ef\u80fd\u306b\u3057\u3066\u304f\u308c\u308b\uff0e\u3053\u308c\u3089\u306e\u52b9\u679c\u3092\u57a3\u9593\u898b\u308b\u3053\u3068\u306f\uff0c\u8a66\u9a13\u7ba1\u5185\u3067\u306e\u30bf\u30a4\u30c1\u30f3\u529b\u5b66\u3092\u8a08\u6e2c\u53ef\u80fd\u3068\u3059\u308b\u3082\u306e\u3067\u3042\u308a\uff0c\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u4e8b\u8c61\u304c 100 \u30df\u30ea\u79d2\u4ee5\u4e0b\u306e\u6642\u9593\u8ef8\u3067\u767a\u751f\u3059\u308b\u3053\u3068\u3092\u793a\u3057\u3066\u3044\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/26854230\" rel=\"noopener noreferrer\" target=\"_blank\">10<\/a>). \u6700\u5f8c\u306b\uff0c\u30b5\u30eb\u30b3\u30e1\u30a2\u306f\u30bf\u30a4\u30c1\u30f3\u7279\u7570\u7684\u30b7\u30e3\u30da\u30ed\u30f3\u3067\u6df7\u96d1\u3057\u305f\u74b0\u5883\u3067\u3042\u308a\uff0c\u305d\u308c\u304c\u6211\u3005\u306e\u6e2c\u5b9a\u3088\u308a\u3082\u3055\u3089\u306b\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u7387\u3092\u5897\u52a0\u3055\u305b\u3066\u3044\u308b\u306b\u9055\u3044\u306a\u3044 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC3897184\/\" rel=\"noopener noreferrer\" target=\"_blank\">102<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/14676215\" rel=\"noopener noreferrer\" target=\"_blank\">103<\/a>). \u3053\u308c\u3089\u306e\u52b9\u679c\u306e\u5168\u3066\u304c, <strong>Figure 1B<\/strong> \u306b\u793a\u3057\u305f\u5b9f\u9a13\u3067\u306f\u6b20\u3051\u3066\u3044\u308b\u304c\uff0c\u30b5\u30eb\u30b3\u30e1\u30a2\u5185\u90e8\u3067\u306e\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u51fa\u529b\u3092\u5f37\u5316\u3059\u308b\u3060\u308d\u3046\uff0e<\/p>\n<h3>Synchronization of titin folding with actomyosin contraction<\/h3>\n<p>\u3000\u30bf\u30a4\u30c1\u30f3\u306e\u305f\u3081\u306e\u6211\u3005\u306e\u4eee\u8aac\u306f\u7b4b\u53ce\u7e2e\u72b6\u614b\u3092\u5f37\u5316\u3059\u308b\u3082\u306e\u3067\u3042\u308b\uff0e\u3064\u307e\u308a\uff0c\u30bf\u30a4\u30c1\u30f3\u3068\u30df\u30aa\u30b7\u30f3\u306f\u5354\u529b\u3057\u3066\u50cd\u304f\uff0e\u305d\u308c\u306f\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304c\u67b6\u6a4b\u306e\u53c2\u4e0e\u306b\u3088\u308a\u5f15\u304d\u8d77\u3053\u3055\u308c\u308b\u5916\u529b\u306e\u6e1b\u5c11\u306b\u4f9d\u5b58\u3057\u3066\u3044\u308b\u305f\u3081\u3067\u3042\u308b\uff0e\u3053\u306e\u30e1\u30ab\u30cb\u30ba\u30e0\u306e\u793a\u5506\u3059\u308b\u3068\u3053\u308d\u306f\uff0c\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306b\u4ed8\u7740\u3057\u3066\u3044\u308b 6 \u3064\u306e\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u306f\u540c\u6642\u306b\u6298\u308a\u305f\u305f\u307e\u308c\u308b\u3060\u308d\u3046\u3068\u3044\u3046\u3053\u3068\u3067\u3042\u308b\uff0e\u5358\u5206\u5b50\u539f\u5b50\u9593\u529b\u5206\u5149\u6cd5\u306e\u8a66\u9a13\u306f\u5e73\u884c\u306a\u30dd\u30ea\u86cb\u767d\u8cea\u306e\u30b9\u30b1\u30fc\u30ea\u30f3\u30b0\u306e\u6cd5\u5247\u3092\u8a3c\u660e\u3057\u305f\uff0e\u86cb\u767d\u8cea\u30c9\u30e1\u30a4\u30f3\u304c\u5e73\u884c\u306b\u4f38\u5c55\u3055\u308c\u308b\u6642\uff0c\u305d\u308c\u3089\u306f\u540c\u6642\u306b\u7834\u88c2\u3057\uff0c\u529b\u306f\u4e8c\u500d\u3068\u306a\u308a\uff0c\u6c38\u7d9a\u9577\u306f\u534a\u5206\u3068\u306a\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC1366990\/\" rel=\"noopener noreferrer\" target=\"_blank\">109<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC1783889\/\" rel=\"noopener noreferrer\" target=\"_blank\">110<\/a>). \u305d\u308c\u3086\u3048\uff0c\u6211\u3005\u306e\u793a\u5506\u3059\u308b\u3068\u3053\u308d\u306f\uff0c6 \u672c\u306e\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u3082\u540c\u6642\u306b\u6298\u308a\u305f\u305f\u307e\u308c\uff0c\u305d\u308c\u306f\u592a\u3044\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u5468\u56f2\u306e\u5e73\u884c\u306b\u914d\u5217\u3055\u308c\u3066\u3044\u308b\u3053\u3068\u306b\u3088\u308b\uff0c\u3068\u3044\u3046\u3082\u306e\u3067\u3042\u308b. 6 \u672c\u306e\u5e73\u884c\u306a\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u306e\u6700\u521d\u306e\u4e00\u672c\u304c\u78ba\u7387\u7684\u306b\u6298\u308a\u305f\u305f\u307e\u308c\u308b\u6642\uff0c\u4ed6\u306e 5 \u672c\u306b\u304b\u304b\u308b\u529b\u306f\u5373\u5ea7\u306b\u6e1b\u5c11\u3059\u308b\uff0e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3084\u30a2\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u7387\u306f\u5916\u529b\u306b\u6307\u6570\u95a2\u6570\u7684\u306b\u4f9d\u5b58\u3057\uff0c\u5916\u529b\u306e\u307b\u3093\u306e\u5c11\u3057\u306e\u6e1b\u5c11\u3067\u3055\u3048\uff0c\u4ed6\u306e\u5e73\u884c\u306a\u30c9\u30e1\u30a4\u30f3\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3092\u52a0\u901f\u3059\u308b\u3060\u308d\u3046\uff0e\u3086\u3048\u306b\uff0c\u30bf\u30a4\u30c1\u30f3 Ig \u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304b\u3089\u306e\u51fa\u529b\u306f\u9ad8\u5ea6\u306b\u5354\u8abf\u7684\u3067\u3042\u308b\uff0e\u3055\u3089\u306b\uff0c\u30a2\u30af\u30c1\u30f3-\u30df\u30aa\u30b7\u30f3\u53ce\u7e2e\u304c\u540c\u671f\u3057\u3066\u3044\u308b\u306e\u306b\u30bf\u30a4\u30c1\u30f3\u306f\u305d\u3046\u3067\u306f\u306a\u3044\uff0c\u3068\u8a00\u3046\u306e\u306f\u8aa4\u89e3\u3092\u62db\u304f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/27572873\" rel=\"noopener noreferrer\" target=\"_blank\">111<\/a>). \u6b21\u306e\u70b9\u306f\u4e00\u822c\u306b\u53d7\u3051\u5165\u308c\u3089\u308c\u3066\u3044\u308b\uff0e\u3064\u307e\u308a\uff0c\u7b4b\u8089\u77ed\u7e2e\u306b\u99c6\u52d5\u3055\u308c\u308b\u30df\u30aa\u30b7\u30f3\u306f &#8220;\u30b5\u30a4\u30af\u30ea\u30c3\u30af\u975e\u540c\u671f ATP \u99c6\u52d5\u5f62\u30a2\u30af\u30c1\u30f3-\u30df\u30aa\u30b7\u30f3\u53ce\u7e2e&#8221; \u306b\u3088\u308a\u767a\u751f\u3059\u308b\u3068 (112). \u67b6\u6a4b\u5f62\u6210\u306f\u30df\u30aa\u30b7\u30f3\u982d\u90e8\u3068\u30a2\u30af\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e\u62e1\u6563\u906d\u9047\u306b\u4f9d\u5b58\u3057\u3066\u304a\u308a\uff0c\u78ba\u7387\u7684\u30d7\u30ed\u30bb\u30b9\u3067\u3042\u308b\u3068\u5b9a\u7fa9\u3055\u308c\u308b\uff0e\u52a0\u3048\u3066, ATP \u52a0\u6c34\u5206\u89e3\u306b\u3088\u308a\u653e\u51fa\u3055\u308c\u308b\u30a8\u30cd\u30eb\u30ae\u30fc\u306f kT \u3088\u308a\u5927\u304d\u306a\u30aa\u30fc\u30c0\u30fc\u306e\u5927\u304d\u3055\u306b\u904e\u304e\u306a\u3044\u305f\u3081\uff0c\u305d\u306e\u5f8c\u306e\u30df\u30aa\u30b7\u30f3\u306e\u69cb\u9020\u5909\u5316\u306f\uff0c\u5c11\u306a\u304f\u3068\u3082\u90e8\u5206\u7684\u306b\u306f\uff0c\u30d6\u30e9\u30a6\u30f3\u904b\u52d5\u306b\u4f9d\u5b58\u3059\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/20385833\" rel=\"noopener noreferrer\" target=\"_blank\">113<\/a>). \u3053\u306e\u8996\u70b9\u304b\u3089\uff0c\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306e\u30b5\u30a4\u30af\u30ea\u30f3\u30b0\u3068\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u3068\u306f\u5171\u306b\u78ba\u7387\u7684\u3067\u3042\u308b\u304c\uff0c\u6211\u3005\u306e\u30e2\u30c7\u30eb\u3067\u306f\uff0c\u7b4b\u8089\u306e\u5316\u5b66\u7684\u6d3b\u6027\u5316\u306b\u304a\u3044\u3066\u540c\u6642\u306b\u767a\u751f\u3059\u308b\uff0e<\/p>\n<h3>Testable predictions and the techniques to examine them<\/h3>\n<p>\u3000\u7b4b\u8089\u306f\u8907\u96d1\u306a\u30b7\u30b9\u30c6\u30e0\u3067\u3042\u308a\uff0c\u968e\u5c64\u69cb\u9020\u306b\u914d\u5217\u3055\u308c\u305f\u86cb\u767d\u8cea\u306e\u3044\u304f\u3064\u304b\u306e\u76f8\u4e92\u4f5c\u7528\u3092\u542b\u3093\u3067\u3044\u308b\uff0e\u5206\u5b50\u30ec\u30d9\u30eb\u304b\u3089\u3053\u308c\u3089\u306e\u5404\u86cb\u767d\u8cea\u306e\u7279\u5b9a\u306e\u5f79\u5272\u3092\u5b9a\u7fa9\u3059\u308b\u3053\u3068\u306f\u56f0\u96e3\u306a\u4e8b\u696d\u3067\u3042\u308b\uff0e\u3057\u304b\u3057\uff0c\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u306e\u30bf\u30f3\u30c7\u30e0\u30e2\u30b8\u30e5\u30fc\u30eb\u5f0f\u914d\u7f6e\u306f\u7b4b\u8089\u306e\u53d7\u52d5\u7684\u5f35\u529b\u306e\u5f79\u5272\u306b\u5358\u306b\u5c02\u5ff5\u3059\u308b\u3060\u3051\u3067\u306f\u306a\u304f\uff0c\u5b9f\u969b\u306b\u3053\u308c\u3089\u306e\u30c9\u30e1\u30a4\u30f3\u304c\u5916\u529b\u306e\u7bc4\u56f2\u5185\u3067\u4f38\u5c55\u3057\u518d\u5ea6\u6298\u308a\u305f\u305f\u307e\u308c\u3066\u52d5\u4f5c\u3059\u308b\u3053\u3068\u306f\uff0c\u4eca\u3084\u660e\u3089\u304b\u306b\u306a\u3063\u305f\u3088\u3046\u306b\u898b\u3048\u308b\uff0e\u7b4b\u8089\u306e\u77ed\u7e2e\u6642\u306b\u3042\u305f\u308a\uff0c\u3053\u308c\u306f\u53ce\u7e2e\u306e\u4ed5\u4e8b\u3092\u767a\u751f\u3055\u305b\u308b\u30bf\u30a4\u30c1\u30f3\u306e\u6d3b\u52d5\u6a5f\u80fd\u3092\u4f34\u3046\uff0e\u3057\u304b\u3057\uff0c\u7b4b\u53ce\u7e2e\u306b\u304a\u3051\u308b\u30bf\u30a4\u30c1\u30f3\u306e\u7cbe\u5bc6\u306a\u5f79\u5272\u304a\u3088\u3073\uff0c\u3044\u304b\u306b\u3057\u3066\u30c9\u30e1\u30a4\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u51fa\u529b\u304c\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306e\u51fa\u529b\u3068\u76f8\u4e92\u4f5c\u7528\u3059\u308b\u304b\u306e\u8a73\u7d30\u306a\u30d3\u30b8\u30e7\u30f3\u3092\u63d0\u4f9b\u3059\u308b\u305f\u3081\u306b\uff0c\u591a\u304f\u306e\u5b9f\u9a13\u306e\u4ed5\u4e8b\u304c\u6b8b\u3055\u308c\u3066\u3044\u308b\uff0e\u5168\u4f53\u3092\u901a\u3058\u3066\u8b70\u8ad6\u3055\u308c\u305f\u4eee\u8aac\u306f\u691c\u8a3c\u53ef\u80fd\u306a\u4e88\u6e2c\u306e\u9375\u3092\u4f5c\u308b\uff0e\u3064\u307e\u308a\uff0c\u7b4b\u8089\u306e\u6d3b\u6027\u5316\u306f\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306e\u53c2\u52a0\u3078\u3068\u7e4b\u304c\u308a\uff0c\u30bf\u30a4\u30c1\u30f3\u3078\u306e\u8ca0\u8377\u3092\u6e1b\u5c11\u3055\u305b\u3066\uff0c\u4ee5\u524d\u306b\u4f38\u5c55\u72b6\u614b\u306b\u52d5\u54e1\u3055\u308c\u305f Ig \u30c9\u30e1\u30a4\u30f3\u3092\u6298\u308a\u305f\u305f\u3080\u30c8\u30ea\u30ac\u30fc\u3068\u306a\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/26854230\" rel=\"noopener noreferrer\" target=\"_blank\">10<\/a>). \u6211\u3005\u306f\u81ea\u5206\u81ea\u8eab\u306b\u554f\u308f\u306a\u3051\u308c\u3070\u306a\u3089\u306a\u3044\uff0e\u30bf\u30a4\u30c1\u30f3\u306e\u3053\u306e\u30a8\u30cd\u30eb\u30ae\u30c3\u30b7\u30e5\u306a\u8ca2\u732e\u306f\u3044\u304b\u306b\u3057\u3066\u5206\u96e2\u3057\u305f\u306e\u304b\uff1f\u307e\u305f\uff0c\u73fe\u5728\u306e\u751f\u7269\u7269\u7406\u5b66\u7684\u30c4\u30fc\u30eb\u306f\u5341\u5206\u306a\u306e\u304b\uff1f\u3068\uff0e<\/p>\n<p>\u3000\u539f\u5b50\u9593\u529b\u5206\u5149\u5b66\u8005\u305f\u3061\u306f\uff0c\u5168\u7d44\u7e54\u30ec\u30d9\u30eb\u3067\u306e\u5358\u5206\u5b50\u6280\u8853\u306e\u7cbe\u5ea6\u3092\u3082\u305f\u3089\u3057\u3066\u304f\u308c\u308b\u30c4\u30fc\u30eb\u3092\u6301\u3064\u3053\u3068\u3092\u5fc3\u304b\u3089\u671b\u3093\u3067\u3044\u308b\uff0e\u3042\u308b\u7a0b\u5ea6\uff0c\u3053\u308c\u306f\u4e00\u822c\u306b\u306f\u30b3\u30fc\u30c9\u5316\u3055\u308c\u305f\u529b\u5b66\u30bb\u30f3\u30b5\u30fc\u3068\u5171\u306b\u8a8d\u8b58\u3055\u308c\u3066\u304d\u3066\u304a\u308a\uff0c\u57f9\u990a\u7d30\u80de\u306e\u529b\u3092\u652f\u3048\u308b\u7d30\u80de\u9aa8\u683c\u6210\u5206\u5185\u90e8\u3067\u767a\u73fe\u3057\u3066\u3044\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/27544876\" rel=\"noopener noreferrer\" target=\"_blank\">114<\/a>). \u3053\u308c\u3089\u306e\u529b\u5b66\u30bb\u30f3\u30b5\u30fc\u306f\u4e8c\u7a2e\u306e\u86cd\u5149\u86cb\u767d\u8cea\u3092\u7528\u3044\uff0c\u975e\u653e\u5c04\u6027\u79fb\u52d5\u3092\u901a\u3058\u3066\u30a8\u30cd\u30eb\u30ae\u30fc\u3092\u5909\u63db\u3059\u308b\u3053\u3068\u304c\u3067\u304d (FRET), \u77ed\u3044\u975e\u69cb\u9020\u5316\u307e\u305f\u306f\u69cb\u9020\u5316\u30da\u30d7\u30c1\u30c9\u306b\u5206\u96e2\u3057\u3066\u3044\u308b\uff0e\u3053\u308c\u3089\u306e\u4e00\u822c\u306b\u30b3\u30fc\u30c9\u5316\u3055\u308c\u305f\u529b\u5b66\u30bb\u30f3\u30b5\u30fc\u306f\u30b9\u30da\u30af\u30c8\u30ea\u30f3\u86cb\u767d (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/22389408\" rel=\"noopener noreferrer\" target=\"_blank\">115<\/a>), \u30d3\u30f3\u30ad\u30e5\u30ea\u30f3\u304a\u3088\u3073\u30bf\u30ea\u30f3 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/26523364\" rel=\"noopener noreferrer\" target=\"_blank\">116<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC2901888\/\" rel=\"noopener noreferrer\" target=\"_blank\">117<\/a>), \u305d\u3057\u3066\u6838\u30cd\u30b9\u30d7\u30ea\u30f3 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/26745407\" rel=\"noopener noreferrer\" target=\"_blank\">118<\/a>) \u306b\u7d44\u307f\u8fbc\u307e\u308c\uff0c\u3053\u308c\u3089\u306f\u5168\u3066\u7d30\u80de\u5185\u30b9\u30c8\u30ec\u30b9\u7dda\u7dad\u304b\u3089\u306e\u5f35\u529b\u306e\u6750\u6599\u3067\u3042\u308b\uff0e\u529b\u5b66\u30bb\u30f3\u30b5\u30fc\u3092\u6709\u3059\u308b\u86cb\u767d\u8cea\u306b\u304b\u304b\u308b\u5f35\u529b\u304c\u5897\u52a0\u3059\u308b\u306b\u3064\u308c, FRET \u30c9\u30ca\u30fc\u304a\u3088\u3073\u30a2\u30af\u30bb\u30d7\u30bf\u30fc\u306f\u5206\u96e2\u3057\u3066 FRET \u7387\u304c\u4f4e\u4e0b\u3059\u308b\uff0e\u3053\u308c\u3089\u306e\u529b\u5b66\u30bb\u30f3\u30b5\u30fc\u306f FRET \u7387\u3092\u8a08\u6e2c\u3059\u308b\u3053\u3068\u3067\u30ad\u30e3\u30ea\u30d6\u30ec\u30fc\u30b7\u30e7\u30f3\u3055\u308c\uff0c\u305d\u308c\u306f 0-10 pN \u306e\u7bc4\u56f2\u3092\u8d85\u3048\u308b\u5358\u5206\u5b50\u539f\u5b50\u9593\u529b\u5206\u5149\u6cd5\u3092\u7528\u3044\u3066\u9069\u7528\u3055\u308c\u305f\u5916\u529b\u306e\u4e0b\u3067\u306e FRET \u7387\u3067\u3042\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/26824190\" rel=\"noopener noreferrer\" target=\"_blank\">119<\/a>). \u30bf\u30a4\u30c1\u30f3 I-band \u306f\u6280\u8853\u8005\u306b\u3068\u3063\u3066 <strong>Figure 5<\/strong> \u306b\u63cf\u304b\u308c\u305f\u3088\u3046\u306b\u529b\u5b66\u30bb\u30f3\u30b5\u30fc\u306e\u7406\u60f3\u7684\u306a\u90e8\u4f4d\u306b\u306a\u308a\u3046\u308b\uff0e\u7d30\u80de\u57f9\u990a\u5b9f\u9a13\u3068\u7570\u306a\u308a\uff0c\u5236\u5fa1\u3055\u308c\u3066\u3044\u306a\u3044\u65b9\u6cd5\u3067\u7d30\u80de\u63a5\u7740\u5f8c\u306e\u7d30\u80de\u9aa8\u683c\u5f35\u529b\u304c\u5897\u52a0\u3059\u308b\u3068\u3053\u308d\u3067\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u306b\u304b\u304b\u308b\u305d\u306e\u5f35\u529b\u306f\u7b4b\u7dda\u7dad\u727d\u5f15\u88c5\u7f6e\u306b\u3088\u308a\u7d76\u5999\u306b\u8abf\u7bc0\u3055\u308c\u308b\u53ef\u80fd\u6027\u304c\u3042\u308b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/4939977\" rel=\"noopener noreferrer\" target=\"_blank\">9<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/14563922\" rel=\"noopener noreferrer\" target=\"_blank\">108<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/21903813\" rel=\"noopener noreferrer\" target=\"_blank\">120<\/a>). \u30bf\u30a4\u30c1\u30f3\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u306e\u30b5\u30eb\u30b3\u30e1\u30a2\u5185\u90e8\u3067\u306e\u7d71\u4e00\u3055\u308c\u305f\u914d\u7f6e\u306f\uff0c\u767a\u8272\u56e3\u304c\u9ad8\u5ea6\u306b\u6574\u5217\u3059\u308b\u3053\u3068\u3092\u78ba\u4fdd\u3057\u3066\u304a\u308a\uff0c\u7121\u50b7\u306e\u7b4b\u8089\u5185\u306e\u86cd\u5149\u8aad\u307f\u53d6\u308a\u306e\u512a\u308c\u305f\u30b7\u30b0\u30ca\u30eb\u30ce\u30a4\u30ba\u6bd4\u3092\u63d0\u4f9b\u3057\u3066\u304f\u308c\u308b\uff0e\u6700\u3082\u91cd\u8981\u306a\u3053\u3068\u3060\u304c\uff0c\u30bf\u30a4\u30c1\u30f3\u306b\u304b\u304b\u308b\u5916\u529b\u306f\u30d0\u30b9\u5185\u306e\u30ab\u30eb\u30b7\u30a6\u30e0\u6fc3\u5ea6\u4e0a\u6607\u306b\u3088\u308a\u8abf\u7bc0\u3055\u308c\u308b\u53ef\u80fd\u6027\u304c\u3042\u308a\uff0c\u305d\u306e\u7d50\u679c\u67b6\u6a4b\u5f62\u6210\u304c\u30bf\u30a4\u30c1\u30f3\u304b\u3089\u529b\u3092\u77ed\u7d61\u3055\u305b\u308b\u7d50\u679c\u3068\u306a\u308b\u3053\u3068\u3092\u6211\u3005\u306f\u793a\u5506\u3057\u305f\uff0e\u7b4b\u53ce\u7e2e\u306b\u304a\u3051\u308b\u6211\u3005\u306e 3 \u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u30e2\u30c7\u30eb\u306b\u5f93\u3046\u3068\uff0c\u751f\u7406\u7684\u30b9\u30c8\u30ec\u30c3\u30c1\u306e\u4e0b\u3067\u306e\u7b4b\u8089\u5185\u306e\u67b6\u6a4b\u5f62\u6210\u306f\u5404\u30bf\u30a4\u30c1\u30f3\u306b\u304b\u304b\u308b\u529b\u3092 10 pN \u4ee5\u4e0b\u306b\u6e1b\u5c11\u3055\u305b\uff0c\u305d\u3053\u3067\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u304c\u30d4\u30fc\u30af\u51fa\u529b\u3092\u653e\u51fa\u3059\u308b\uff0e\u4e00\u822c\u7684\u306b\u30b3\u30fc\u30c9\u5316\u3055\u308c\u305f\u529b\u5b66\u30bb\u30f3\u30b5\u30fc\u304c\u30bf\u30a4\u30c1\u30f3\u3054\u3068\u306e\u529b\u3092\u6c7a\u5b9a\u3059\u308b\u3053\u3068\u3092\u53ef\u80fd\u3068\u3059\u308b\u3060\u3051\u3067\u306f\u306a\u304f\uff0c\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53ce\u7e2e\u3067\u306e\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u3082\u307e\u305f\u30c8\u30ea\u30ac\u30fc\u3055\u308c\u308b\uff0e\u5404\u7b4b\u7fa4\u306f\u7570\u306a\u308b\u9577\u3055\u306e\u30bf\u30a4\u30c1\u30f3\u30a2\u30a4\u30bd\u30d5\u30a9\u30fc\u30e0\u3092\u6709\u3057\u3066\u3044\u308b\uff0e\u4f8b\u3048\u3070\u7570\u306a\u308b 40 \u30bb\u30c3\u30c8\u306e\u30a6\u30b5\u30ae\u306e\u7b4b\u8089\u306e\u3088\u3046\u306b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/16230467\" rel=\"noopener noreferrer\" target=\"_blank\">87<\/a>). \u6211\u3005\u306e\u793a\u5506\u3059\u308b\u3068\u3053\u308d\u3067\u306f\uff0c\u3053\u306e\u9577\u3055\u306f\u6b21\u306e\u3088\u3046\u306b\u30d7\u30ed\u30b0\u30e9\u30e0\u3055\u308c\u3066\u3044\u308b\uff0e\u3064\u307e\u308a\uff0c\u30bf\u30a4\u30c1\u30f3\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u53ce\u7e2e\u306f\uff0c\u7279\u5b9a\u306e\u7b4b\u8089\u7fa4\u306e\u6700\u9069\u306a\u4ed5\u4e8b\u3092\u3059\u308b\u30b5\u30eb\u30b3\u30e1\u30a2\u9577\u306e\u4e2d\u3067\u767a\u751f\u3059\u308b\u3068\uff0e\u305d\u306e\u3088\u3046\u306a\u30c4\u30fc\u30eb\u306f\u7b4b\u8089\u306e\u52d5\u4f5c\u306e\u7814\u7a76\u306b\u3068\u3063\u3066\uff0c\u751f\u4f53\u5185\u3084\u8a66\u9a13\u7ba1\u5185\u3044\u305a\u308c\u306b\u304a\u3044\u3066\u3082\u975e\u5e38\u306b\u8cb4\u91cd\u3067\u3042\u308a\uff0c\u30bf\u30a4\u30c1\u30f3\u304a\u3088\u3073\u30bf\u30a4\u30c1\u30f3\u306b\u304b\u304b\u308b\u8ca0\u8377\u3092\u6e1b\u5c11\u3055\u305b\u308b\u306e\u306b\u5fc5\u8981\u306a\u67b6\u6a4b\u306e\u6570\u306b\u3088\u308a\u7d4c\u9a13\u3059\u308b\u751f\u7406\u5b66\u7684\u306a\u529b\u3092\u3088\u308a\u826f\u304f\u7406\u89e3\u3059\u308b\u305f\u3081\u306e\u5206\u91ce\u3078\u306e\u79fb\u52d5\u306e\u52a9\u3051\u3068\u306a\u308b\u3060\u308d\u3046\uff0e\u3053\u308c\u3089\u306e\u4e8c\u3064\u306e\u529b\u306f\uff0c\u30bf\u30a4\u30c1\u30f3\u306e\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u901f\u5ea6\u3068\u4f75\u305b\u3066\uff0c\u7b4b\u53ce\u7e2e\u6642\u306e\u30bf\u30a4\u30c1\u30f3\u3068\u30df\u30aa\u30b7\u30f3\u306e\u76f8\u5bfe\u7684\u306a\u51fa\u529b\u3092\u6c7a\u5b9a\u3059\u308b\u306e\u306b\u91cd\u8981\u3067\u3042\u308b\uff0e<\/p>\n<p>\u3000\u305d\u306e\u3088\u3046\u306a\u751f\u4f53\u5185\u30c4\u30fc\u30eb\u306e\u767a\u5c55\u3078\u306e\u6700\u521d\u306e\u6bb5\u968e\u306f\u9054\u6210\u3055\u308c\u3066\u3044\u308b. I-band \u9060\u4f4d\u90e8\u5206\u3078\u306e HaloTag \u9175\u7d20\u306e\u30ce\u30c3\u30af\u30a4\u30f3\u3092\u6709\u3059\u308b\u30e2\u30c7\u30eb\u30de\u30a6\u30b9\u306f\u30bf\u30a4\u30c1\u30f3\u306e\u751f\u7269\u7269\u7406\u5b66\u7684\u7279\u6027\u306e\u305f\u3081\u306e\u591a\u76ee\u7684\u30c4\u30fc\u30eb\u3068\u3057\u3066\u5949\u4ed5\u3057\u305f (<a href=\"https:\/\/www.cell.com\/biophysj\/abstract\/S0006-3495(16)31949-X?code=cell-site\" rel=\"noopener noreferrer\" target=\"_blank\">121<\/a>). HaloTag \u9175\u7d20\u306e\u4f7f\u7528\u306f\uff0c\u30ac\u30e9\u30b9\u88fd\u30ab\u30d0\u30fc\u30b9\u30ea\u30c3\u30d7\u306b\u5358\u5206\u5b50\u3092\u727d\u5f15\u3059\u308b\u5b9f\u9a13\u306e\u305f\u3081\u306e\u5171\u6709\u7d50\u5408\u30a2\u30f3\u30ab\u30fc\u86cb\u767d\u8cea\u3068\u3057\u3066\uff0c\u6211\u3005\u306e\u30e9\u30dc\u3067\u958b\u767a\u3055\u308c\u305f\u3082\u306e\u3067\u3042\u308a (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/23909704\" rel=\"noopener noreferrer\" target=\"_blank\">41<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/27409974\" rel=\"noopener noreferrer\" target=\"_blank\">49<\/a>), \u86cd\u5149\u30a4\u30e1\u30fc\u30b8\u30f3\u30b0\u306e\u5e83\u7bc4\u56f2\u306e\u53d6\u5f97\u3092\u53ef\u80fd\u3068\u3057\u305f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/25157152\" rel=\"noopener noreferrer\" target=\"_blank\">122<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC3808294\/\" rel=\"noopener noreferrer\" target=\"_blank\">123<\/a>). HaloTag \u306f\u30af\u30ed\u30ed\u30a2\u30eb\u30ab\u30f3\u57fa\u8cea\u3092\u8a8d\u8b58\u3057\u3066\u5171\u6709\u7d50\u5408\u3057\uff0c\u69d8\u3005\u306a\u6709\u6a5f\u767a\u8272\u56e3\u3068\u4ed8\u7740\u3057\u3066\u512a\u308c\u305f\u30a4\u30e1\u30fc\u30b8\u30f3\u30b0\u7279\u6027\u3092\u6301\u3064\uff0e\u6297\u4f53\u3068\u9055\u3063\u3066\uff0c\u30af\u30ed\u30ed\u30a2\u30eb\u30ab\u30f3\u30ea\u30ac\u30f3\u30c9\u306f\u975e\u5e38\u306b\u5c0f\u3055\u304f\uff0c\u5bb9\u6613\u306b\u30b5\u30eb\u30b3\u30e1\u30a2\u306b\u6d78\u900f\u3057\u3066\u9ad8\u6fc3\u5ea6\u306e\u6a19\u8b58\u306b\u9054\u3059\u308b\u3053\u3068\u304c\u3067\u304d\u308b. SNAP-Tag, CLIP-Tag \u3084 TMP-tag (122) \u306a\u3069\u306e\u76f4\u4ea4\u6a19\u8b58\u90e8\u4f4d\u3092 HaloTag \u306b\u7d9a\u3051\u3066\u914d\u7f6e\u3059\u308b\u3053\u3068\u3067\uff0c\u4e0a\u8ff0\u3057\u305f\u30bb\u30f3\u30b5\u30fc\u306b\u8fd1\u4f3c\u3057\u305f\u4e00\u822c\u7684\u306b\u30b3\u30fc\u30c9\u5316\u3055\u308c\u305f\u30bb\u30f3\u30b5\u30fc\u304c\u9054\u6210\u3055\u308c\u308b\u53ef\u80fd\u6027\u304c\u3042\u308b (<strong>Figure 5C<\/strong>). \u3053\u308c\u306f\u5148\u8ff0\u3057\u305f\u6709\u6a5f\u67d3\u6599\u3092\u7528\u3044\u308b\u3053\u3068\u304c\u3067\u304d\u308b\u70b9\u3067\u4e00\u822c\u7684\u306b\u30b3\u30fc\u30c9\u5316\u3055\u308c\u305f\u529b\u5b66\u30bb\u30f3\u30b5\u30fc\u306b\u5bfe\u3057\u3066\u512a\u4f4d\u6027\u304c\u3042\u308a\uff0c\u86cd\u5149\u86cb\u767d\u8cea\u306e\u30bb\u30f3\u30b5\u30fc\u3088\u308a\u512a\u308c\u305f\u7279\u6027\u3092\u6301\u3064\uff0e\u4f7f\u7528\u3057\u305f\u86cd\u5149\u767a\u8272\u56e3\u306e\u30bb\u30c3\u30c8\u306f\u624b\u5143\u306e\u5b9f\u9a13\u3067\u3082\u9069\u5fdc\u3059\u308b\u53ef\u80fd\u6027\u304c\u3042\u308a\uff0c\u5358\u4e00\u7b4b\u7dda\u7dad\u4f38\u5c55\u5b9f\u9a13\u3084\u751f\u4f53\u5185\u30a4\u30e1\u30fc\u30b8\u30f3\u30b0\u306b\u3082\u9069\u5fdc\u3059\u308b\u53ef\u80fd\u6027\u304c\u3042\u308b. HaloTag \u306b\u52a0\u3048\u3066\uff0c\u73fe\u5728\u306e\u30de\u30a6\u30b9\u30e2\u30c7\u30eb\u306f TEV \uff08\u30a6\u30a4\u30eb\u30b9\uff09\u30d7\u30ed\u30c6\u30a2\u30fc\u30bc\u90e8\u4f4d\u3092\u3082\u6709\u3057\u3066\u3044\u308b\uff0e\u3053\u308c\u307e\u3067\u306e\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u3092\u5207\u65ad\u3057\u3088\u3046\u3068\u3059\u308b\u8a66\u307f (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/11927678\" rel=\"noopener noreferrer\" target=\"_blank\">107<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/3755803\" rel=\"noopener noreferrer\" target=\"_blank\">124<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/1587789\" rel=\"noopener noreferrer\" target=\"_blank\">125<\/a>) \u3068\u9055\u3063\u3066\uff0c\u3053\u308c\u306f\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\u306e\u534a\u5206\uff08\u30ce\u30c3\u30af\u30a4\u30f3\u306e\u30d8\u30c6\u30ed\u63a5\u5408\u4f53\u30de\u30a6\u30b9\uff09\u304b\uff0c\u5168\u30bf\u30a4\u30c1\u30f3\u5206\u5b50\uff08\u30db\u30e2\u63a5\u5408\u4f53\uff09\u3092\u9ad8\u5ea6\u306b\u7279\u7570\u7684\u306b\u5207\u65ad\u3059\u308b\u3053\u3068\u304c\u53ef\u80fd\u3067\u3042\u308a\uff0c\u305d\u308c\u306b\u3088\u308a\u7b4b\u5f3e\u6027\u306b\u304a\u3051\u308b\u30bf\u30a4\u30c1\u30f3\u306e\u5f79\u5272\u3068\u529b\u306e\u767a\u751f\u3092\u3088\u308a\u3088\u304f\u7406\u89e3\u3059\u308b\u3053\u3068\u304c\u3067\u304d\u308b\uff0e<\/p>\n<p>\u3000\u7b4b\u53ce\u7e2e\u306e\u8907\u96d1\u3055\u3092\u3088\u308a\u6df1\u304f\u63a2\u6c42\u3059\u308b\u3053\u3068\u306f\u53e4\u5178\u7684\u306a\u7b4b\u53ce\u7e2e\u5b9f\u9a13\u306e\u518d\u8a55\u4fa1\u3092\u8981\u3059\u308b\uff0e\u305d\u308c\u306b\u306f\u7b4b\u8089\u306e\u53d7\u52d5\u7684\u304a\u3088\u3073\u80fd\u52d5\u7684\u30e1\u30ab\u30cb\u30ba\u30e0\u306e\u4e21\u8005\u306b\u304a\u3044\u3066\u30bf\u30a4\u30c1\u30f3\u304c\u4e3b\u8981\u306a\u5f79\u5272\u3092\u679c\u305f\u3059\u3053\u3068\u3092\u5ff5\u982d\u306b\u7f6e\u304f\u5fc5\u8981\u304c\u3042\u308b\uff0e\u30df\u30aa\u30b7\u30f3\u981a\u90e8\u306e\u8ffd\u5f93\u306b\u304a\u3051\u308b\u4e0b\u65b9\u50be\u5411\u3092\u9451\u307f\u308b\u306b (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/18022371\" rel=\"noopener noreferrer\" target=\"_blank\">89<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/9845077\" rel=\"noopener noreferrer\" target=\"_blank\">126<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/11832949\" rel=\"noopener noreferrer\" target=\"_blank\">127<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/15058307\" rel=\"noopener noreferrer\" target=\"_blank\">128<\/a>), \u53ce\u7e2e\u3059\u308b\u30b5\u30eb\u30b3\u30e1\u30a2\u306e\u4e3b\u8981\u306a\u30a8\u30cd\u30eb\u30ae\u30fc\u84c4\u7a4d\u6e90\u306f\u518d\u8003\u3055\u308c\u306a\u304f\u3066\u306f\u306a\u3089\u306a\u3044\uff0e\u304a\u305d\u3089\u304f\u4e00\u822c\u306b\u30b3\u30fc\u30c9\u5316\u3055\u308c\u305f\u529b\u5b66\u30bb\u30f3\u30b5\u30fc\u306f, Huxley \u3068 Simmons \u306e\u53e4\u5178\u7684\u306a\u30af\u30a4\u30c3\u30af\u30ea\u30ea\u30fc\u30b9\u5b9f\u9a13 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/4939977\" rel=\"noopener noreferrer\" target=\"_blank\">9<\/a>) \u306e\u30d0\u30fc\u30b8\u30e7\u30f3\u3092\u518d\u8003\u3059\u308b\u3053\u3068\u3067\uff0c\u30bf\u30a4\u30c1\u30f3\u304c\u4e3b\u8981\u306a\u30a8\u30cd\u30eb\u30ae\u30fc\u84c4\u7a4d\u6e90\u3067\u3042\u308b\u3053\u3068\u3092\u8a3c\u660e\u3059\u308b\u3053\u3068\u306b\u306a\u308b\u3060\u308d\u3046\uff0e\u305d\u3057\u3066\u518d\u69cb\u7bc9\u3055\u308c\u305f\u30a2\u30af\u30c1\u30f3\u30df\u30aa\u30b7\u30f3\u8907\u5408\u4f53\u306e\u5358\u5206\u5b50\u5b9f\u9a13\u304c\u6539\u5584\u3057\u3064\u3064\u3042\u308a (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/8139653\" rel=\"noopener noreferrer\" target=\"_blank\">26<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/20689017\" rel=\"noopener noreferrer\" target=\"_blank\">28<\/a>,<a href=\"https:\/\/www.nature.com\/articles\/ncomms16036\" rel=\"noopener noreferrer\" target=\"_blank\">29<\/a>), \u3055\u3089\u306b\u30df\u30aa\u30b7\u30f3\u306b\u3088\u308b\u529b\u306e\u767a\u751f\u30e1\u30ab\u30cb\u30ba\u30e0\u3078\u306e\u6d1e\u5bdf\u3092\u3082\u305f\u3089\u3057\u3064\u3064\u3042\u308b\u306b\u3082\u304b\u304b\u308f\u3089\u305a\uff0c\u7121\u50b7\u306e\u30b5\u30eb\u30b3\u30e1\u30a2\u5185\u90e8\u3067\u5358\u4e00\u306e\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306e\u901f\u5ea6\u3084\u30b9\u30c8\u30ed\u30fc\u30af\u30b5\u30a4\u30ba\u3092\u76f4\u63a5\u89b3\u5bdf\u3057\u305f\u5b9f\u9a13\u306f\u307e\u3060\u5b58\u5728\u3057\u306a\u3044\uff0e\u3053\u306e\u89b3\u70b9\u304b\u3089\u306f\uff0c\u305d\u306e\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306e\u5206\u91ce\u306f\u30bf\u30a4\u30c1\u30f3\u3088\u308a\u5f8c\u306e\u6bb5\u968e\u3067\u3042\u308b\uff0e\u3055\u3089\u306b\uff0c\u71b1\u529b\u5b66\u30a8\u30f3\u30b8\u30f3\u306e\u3088\u3046\u306b\u7b4b\u8089\u3092\u6271\u3046\u3053\u3068\u306f, Fenn \u3084 Wilkie \u305d\u306e\u4ed6\u306e\u7814\u7a76\u8005\u3089\u304c\u71b1\u96fb\u5806\u3068\u71b1\u91cf\u8a08\u3092\u7528\u3044\u3066\u7b4b\u8089\u30a8\u30f3\u30bf\u30eb\u30d4\u30fc\u3092\u8a08\u6e2c\u3057\u305f\u7d50\u679c\uff0c\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u5358\u72ec\u306b\u71b1\u7523\u751f\u3092\u5272\u308a\u5f53\u3066\u3066\u3057\u307e\u3063\u305f\u3088\u3046\u306b\uff0c\u554f\u984c\u3092\u904e\u5ea6\u306b\u5358\u7d14\u5316\u3057\u3066\u3057\u307e\u3046 (<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/5639798\" rel=\"noopener noreferrer\" target=\"_blank\">27<\/a>,<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC1405508\/\" rel=\"noopener noreferrer\" target=\"_blank\">129<\/a>). \u5358\u7d14\u306a\u7c98\u5f3e\u6027\u30dd\u30ea\u30de\u30fc\u306f\u529b\u306e\u6e1b\u5c11\u306b\u4e57\u3058\u3066\u5de8\u5927\u306a\u30a8\u30f3\u30c8\u30ed\u30d4\u30fc\u5897\u5927\u3092\u7d4c\u3066\uff0c\u5468\u8fba\u74b0\u5883\u304b\u3089\u71b1\u3092\u5438\u53ce\u3059\u308b\uff0e\u6298\u308a\u305f\u305f\u307e\u308c\u305f Ig 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Fenn \u3089\u306e\u7d50\u679c\u306f\u4eca\u3084\u518d\u8003\u3055\u308c\u308b\u5fc5\u8981\u304c\u3042\u308a\uff0c\u6211\u3005\u306f\u30bf\u30a4\u30c1\u30f3\u86cb\u767d\u8cea\u30d5\u30a9\u30fc\u30eb\u30c7\u30a3\u30f3\u30b0\u306e\u71b1\u529b\u5b66\u304c\u7c98\u5f3e\u6027\u30dd\u30ea\u30de\u30fc\u307b\u3069\u5358\u7d14\u3067\u306f\u306a\u3044\u3053\u3068\u3092\u77e5\u3063\u3066\u3044\u308b\uff0e<\/p>\n<p>\u3000\u7d50\u8ad6\u3068\u3057\u3066\uff0c\u7b4b\u53ce\u7e2e\u306b\u304a\u3051\u308b\u6211\u3005\u306e\u8996\u70b9\u306f\uff0c\u5b89\u9759\u5358\u72ec\u306e\u30e2\u30fc\u30bf\u30fc\u6d3b\u6027\u304b\u3089\uff0c\u4f38\u5c55\u529b\u306b\u6297\u3057\u3066\u6298\u308a\u305f\u305f\u307e\u308c\u308b\u6a5f\u68b0\u7684\u306b\u5c55\u958b\u3057\u305f\u86cb\u767d\u8cea\u306e\u53ef\u80fd\u6027\u306b\u7126\u70b9\u3092\u79fb\u3057\u305f\uff0e\u30bf\u30a4\u30c1\u30f3\u306e\u3053\u306e\u30e6\u30cb\u30fc\u30af\u306a\u7d20\u6750\u7279\u6027\u306f\uff0c\u305d\u306e\u6298\u308a\u305f\u305f\u307f\u53ef\u80fd\u306a Ig \u30c9\u30e1\u30a4\u30f3\u306b\u3088\u308a\u4e0e\u3048\u3089\u308c, Hookean \u3070\u306d\u3068\u3082\u975e\u69cb\u9020\u30dd\u30ea\u30de\u30fc\u3068\u3082\u7570\u306a\u308a\uff0c\u552f\u4e00\uff0c\u5f3e\u6027\u53ce\u7e2e\u3092\u7d4c\u9a13\u3059\u308b\u53d7\u52d5\u7684\u30a8\u30e9\u30b9\u30c8\u30de\u30fc\u3067\u3042\u308b\u3053\u3068\u3060\u308d\u3046\uff0e\u5358\u5206\u5b50\u30c7\u30fc\u30bf\u306f\u660e\u3089\u304b\u306b\u793a\u3057\u3066\u3044\u308b\u304c, 6 pN \u4ee5\u4e0a\u3067\u306f\uff0c\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u306f\u5931\u901f\u3057\u3066\u524d\u9032\u3067\u304d\u305a\uff0c\u6a5f\u68b0\u7684\u4ed5\u4e8b\u3092\u306a\u3059\u3053\u3068\u304c\u3067\u304d\u306a\u3044\uff0e\u305d\u308c\u306b\u5fdc\u3058\u3066\uff0c\u30bf\u30a4\u30c1\u30f3 Ig \u30c9\u30e1\u30a4\u30f3\u306f\u305d\u306e\u5916\u529b\u306e\u7bc4\u56f2\u3067\u5bb9\u6613\u306b\u518d\u6298\u308a\u305f\u305f\u307f\uff0c\u53ce\u7e2e\u306e\u4ed5\u4e8b\u3092\u63d0\u4f9b\u3057\u3066\u5931\u901f\u3057\u305f\u30df\u30aa\u30b7\u30f3\u30e2\u30fc\u30bf\u30fc\u3092\u89e3\u653e\u3057\uff0c\u52b9\u679c\u7684\u306a\u7b4b\u53ce\u7e2e\u3092\u8a31\u3059\uff0e\u3086\u3048\u306b\uff0c\u3053\u306e\u65b0\u3057\u3044\u6839\u62e0\u306e\u793a\u3059\u3068\u3053\u308d\u306f\uff0c\u7b2c\u4e09\u306e\u30d5\u30a3\u30e9\u30e1\u30f3\u30c8\u3068\u3057\u3066\u8a8d\u8b58\u3055\u308c\u305f\u306e\u3068\u540c\u69d8\u306b\uff0c\u7b4b\u53ce\u7e2e\u306b\u304a\u3044\u3066\u30a2\u30af\u30c1\u30f3\u3068\u30df\u30aa\u30b7\u30f3\u3068\u5171\u306b\u50cd\u304f\u4e0d\u53ef\u5206\u306e\u30d1\u30fc\u30c8\u30ca\u30fc\u3068\u3057\u3066\uff0c\u30bf\u30a4\u30c1\u30f3\u3092\u30b5\u30eb\u30b3\u30e1\u30a2\u306e\u6d3b\u6027\u8981\u7d20\u3068\u3057\u3066\u8003\u616e\u3059\u308b\u5fc5\u8981\u304c\u3042\u308b\uff0e<\/p>\n<h3>References<\/h3>\n<p>1. Maruyama K, Matsubara S, Natori R, Nonomura Y, Kimura S, et al. Connectin, an Elastic Protein of Muscle Characterization and Function. J Biochem (Tokyo). 1977; 82(2):317\u201337. [PubMed: 914784]<\/p>\n<p>2. Polissar MJ. Physical chemistry of contractile process in muscle. I. A physicochemical model of contractile mechanism. Am J Physiol. 1952; 168(3):766\u201381. [PubMed: 14914890]<\/p>\n<p>3. Polissar MJ. Physical chemistry of contractile process in muscle. II. Analysis of other mechanochemical properties of muscle. Am J Physiol. 1952; 168(3):782\u201392. [PubMed: 14914891]<\/p>\n<p>4. Polissar MJ. Physical chemistry of contractile process in muscle. III. Interpretation of thermal behavior of stimulated muscle. Am J Physiol. 1952; 168(3):793\u2013804. [PubMed: 14914892]<\/p>\n<p>5. Polissar MJ. Physical chemistry of contractile process in muscle. IV. Estimates of size of contractile unit. Am J Physiol. 1952; 168(3):805\u201311. [PubMed: 14914893]<\/p>\n<p>6. Gordon AM, Huxley AF, Julian FJ. Tension development in highly stretched vertebrate muscle fibres. J Physiol. 1966; 184(1):143\u201369. [PubMed: 5921535]<\/p>\n<p>7. Gordon AM, Huxley AF, Julian FJ. The variation in isometric tension with sarcomere length in vertebrate muscle fibres. J Physiol. 1966; 184(1):170\u201392. [PubMed: 5921536]<\/p>\n<p>8. Huxley AF, Niedergerke R. Structural Changes in Muscle During Contraction: Interference Microscopy of Living Muscle Fibres. Nature. 1954; 173(4412):971\u201373. [PubMed: 13165697]<\/p>\n<p>9. Huxley AF, Simmons RM. Proposed Mechanism of Force Generation in Striated Muscle. Nature. 1971; 233(5321):533\u201338. [PubMed: 4939977]<\/p>\n<p>10. Rivas-Pardo JA, Eckels EC, Popa I, Kosuri P, Linke WA, Fern\u00e1ndez JM. Work Done by Titin Protein Folding Assists Muscle Contraction. Cell Rep. 2016; 14(6):1339\u201347. [PubMed: 26854230]<\/p>\n<p>11. Huxley HE. Structural Arrangements and the Contraction Mechanism in Striated Muscle. Proc R Soc Lond B Biol Sci. 1964; 160(981):442\u201348. [PubMed: 14214772]<\/p>\n<p>12. Maruyama K, Natori R, Nonomura Y. New elastic protein from muscle. Nature. 1976; 262(5563): 58\u201360. [PubMed: 934326]<\/p>\n<p>13. Maruyama K, Kimura S, Ohashi K, Kuwano Y. Connectin, an Elastic Protein of Muscle. Identification of \u201cTitin\u201d with Connectin. J Biochem (Tokyo). 1981; 89(3):701\u20139. [PubMed: 7287636]<\/p>\n<p>14. Stull JT. Myosin Minireview Series. J Biol Chem. 1996; 271(27):15849\u201315849. [PubMed: 8663555]<\/p>\n<p>15. Bang M-L, Centner T, Fornoff F, Geach AJ, Gotthardt M, et al. The Complete Gene Sequence of Titin, Expression of an Unusual \u2248700-kDa Titin Isoform, and Its Interaction With Obscurin Identify a Novel Z-Line to I-Band Linking System. Circ Res. 2001; 89(11):1065\u201372. [PubMed: 11717165]<\/p>\n<p>16. Burkholder TJ, Lieber RL. Sarcomere length operating range of vertebrate muscles during movement. J Exp Biol. 2001; 204(9):1529\u201336. [PubMed: 11296141]<\/p>\n<p>17. Millman BM. The Filament Lattice of Striated Muscle. Physiol Rev. 1998; 78(2):359\u201391. [PubMed: 9562033]<\/p>\n<p>18. Cazorla O, Freiburg A, Helmes M, Centner T, McNabb M, et al. Differential Expression of Cardiac Titin Isoforms and Modulation of Cellular Stiffness. Circ Res. 2000; 86(1):59\u201367. [PubMed: 10625306]<\/p>\n<p>19. Bogomolovas J, Gasch A, Simkovic F, Rigden DJ, Labeit S, Mayans O. Titin kinase is an inactive pseudokinase scaffold that supports MuRF1 recruitment to the sarcomeric M-line. Open Biol. 2014; 4(5):140041. [PubMed: 24850911]<\/p>\n<p>20. Lange S, Xiang F, Yakovenko A, Vihola A, Hackman P, et al. The Kinase Domain of Titin Controls Muscle Gene Expression and Protein Turnover. Science. 2005; 308(5728):1599\u20131603. [PubMed: 15802564]<\/p>\n<p>21. Linke WA, Hamdani N. Gigantic Business Titin Properties and Function Through Thick and Thin. Circ Res. 2014; 114(6):1052\u201368. [PubMed: 24625729]<\/p>\n<p>22. Hessel AL, Lindstedt SL, Nishikawa KC. Physiological Mechanisms of Eccentric Contraction and Its Applications: A Role for the Giant Titin Protein. Front Physiol. 2017; 8 <\/p>\n<p>23. Tskhovrebova L, Trinick J. Titin: properties and family relationships. Nat Rev Mol Cell Biol. 2003; 4(9):679\u201389. [PubMed: 14506471]<\/p>\n<p>24. Linke WA, Kr\u00fcger M. The Giant Protein Titin as an Integrator of Myocyte Signaling Pathways. Physiology. 2010; 25(3):186\u201398. [PubMed: 20551232]<\/p>\n<p>25. Spudich JA. Hypertrophic and Dilated Cardiomyopathy: Four Decades of Basic Research on Muscle Lead to Potential Therapeutic Approaches to These Devastating Genetic Diseases. Biophys J. 2014; 106(6):1236\u201349. [PubMed: 24655499]<\/p>\n<p>26. Finer JT, Simmons RM, Spudich JA. Single myosin molecule mechanics: piconewton forces and nanometre steps. Nature. 1994; 368(6467):113\u201319. [PubMed: 8139653]<\/p>\n<p>27. Wilkie DR. Heat work and phosphorylcreatine break-down in muscle. J Physiol. 1968; 195(1):157\u2013 83. [PubMed: 5639798]<\/p>\n<p>28. Kaya M, Higuchi H. Nonlinear Elasticity and an 8-nm Working Stroke of Single Myosin Molecules in Myofilaments. Science. 2010; 329(5992):686\u201389. [PubMed: 20689017]<\/p>\n<p>29. Kaya M, Tani Y, Washio T, Hisada T, Higuchi H. Coordinated force generation of skeletal myosins in myofilaments through motor coupling. Nat Commun. 2017; 8 ncomms16036. <\/p>\n<p>30. Herzog W, Schappacher G, DuVall M, Leonard TR, Herzog JA. Residual Force Enhancement Following Eccentric Contractions: A New Mechanism Involving Titin. Physiology. 2016; 31(4): 300\u2013312. [PubMed: 27252165]<\/p>\n<p>31. Herzog W, Leonard TR. Force enhancement following stretching of skeletal muscle. J Exp Biol. 2002; 205(9):1275\u201383. [PubMed: 11948204]<\/p>\n<p>32. Nishikawa KC, Monroy JA, Uyeno TE, Yeo SH, Pai DK, Lindstedt SL. Is titin a \u201cwinding filament\u201d? A new twist on muscle contraction. Proc R Soc Lond B Biol Sci. 2012; 279(1730):981\u2013 90. <\/p>\n<p>33. Carrion-Vazquez M, Oberhauser AF, Fowler SB, Marszalek PE, Broedel SE, et al. Mechanical and chemical unfolding of a single protein: A comparison. Proc Natl Acad Sci. 1999; 96(7):3694\u201399. [PubMed: 10097099]<\/p>\n<p>34. Fernandez JM, Li H. Force-Clamp Spectroscopy Monitors the Folding Trajectory of a Single Protein. Science. 2004; 303(5664):1674\u201378. [PubMed: 15017000]<\/p>\n<p>35. Kellermayer MSZ, Smith SB, Granzier HL, Bustamante C. Folding-Unfolding Transitions in Single Titin Molecules Characterized with Laser Tweezers. Science. 1997; 276(5315):1112\u201316. [PubMed: 9148805]<\/p>\n<p>36. Li H, Linke WA, Oberhauser AF, Carrion-Vazquez M, Kerkvliet JG, et al. Reverse engineering of the giant muscle protein titin. Nature. 2002; 418(6901):998\u20131002. [PubMed: 12198551]<\/p>\n<p>37. Rief M, Gautel M, Oesterhelt F, Fernandez JM, Gaub HE. Reversible Unfolding of Individual Titin Immunoglobulin Domains by AFM. Science. 1997; 276(5315):1109\u201312. [PubMed: 9148804]<\/p>\n<p>38. Schlierf M, Li H, Fernandez JM. The unfolding kinetics of ubiquitin captured with single-molecule force-clamp techniques. Proc Natl Acad Sci U S A. 2004; 101(19):7299\u20137304. [PubMed: 15123816]<\/p>\n<p>39. Tskhovrebova L, Trinick J, Sleep JA, Simmons RM. Elasticity and unfolding of single molecules of the giant muscle protein titin. Nature. 1997; 387(6630):308\u201312. [PubMed: 9153398]<\/p>\n<p>40. Popa I, Kosuri P, Alegre-Cebollada J, Garcia-Manyes S, Fernandez JM. Force dependency of biochemical reactions measured by single-molecule force-clamp spectroscopy. Nat Protoc. 2013; 8(7):1261\u201376. [PubMed: 23744288]<\/p>\n<p>41. Popa I, Berkovich R, Alegre-Cebollada J, Badilla CL, Rivas-Pardo JA, et al. Nanomechanics of HaloTag Tethers. J Am Chem Soc. 2013; 135(34):12762\u201371. [PubMed: 23909704]<\/p>\n<p>42. Chen H, Fu H, Zhu X, Cong P, Nakamura F, Yan J. Improved High-Force Magnetic Tweezers for Stretching and Refolding of Proteins and Short DNA. Biophys J. 2011; 100(2):517\u201323. [PubMed: 21244848]<\/p>\n<p>43. Chen H, Yuan G, Winardhi RS, Yao M, Popa I, et al. Dynamics of Equilibrium Folding and Unfolding Transitions of Titin Immunoglobulin Domain under Constant Forces. J Am Chem Soc. 2015; 137(10):3540\u201346. [PubMed: 25726700]<\/p>\n<p>44. Svoboda K, Schmidt CF, Schnapp BJ, Block SM. Direct observation of kinesin stepping by optical trapping interferometry. Nature. 1993; 365(6448):721\u201327. [PubMed: 8413650]<\/p>\n<p>45. Martonfalvi Z, Bianco P, Linari M, Caremani M, Nagy A, et al. Low-force transitions in single titin molecules reflect a memory of contractile history. J Cell Sci. 2014; 127(4):858\u201370. [PubMed: 24357719]<\/p>\n<p>46. Neuman KC, Nagy A. Single-molecule force spectroscopy: optical tweezers, magnetic tweezers and atomic force microscopy. Nat Methods. 2008; 5(6):491\u2013505. [PubMed: 18511917]<\/p>\n<p>47. Huhle A, Klaue D, Brutzer H, Daldrop P, Joo S, et al. Camera-based three-dimensional real-time particle tracking at kHz rates and \u00c5ngstr\u00f6m accuracy. Nat Commun. 2015; 6 <\/p>\n<p>48. Dulin D, Cui TJ, Cnossen J, Docter MW, Lipfert J, Dekker NH. High Spatiotemporal-Resolution Magnetic Tweezers: Calibration and Applications for DNA Dynamics. Biophys J. 2015; 109(10): 2113\u201325. [PubMed: 26588570]<\/p>\n<p>49. Popa I, Rivas-Pardo JA, Eckels EC, Echelman DJ, Badilla CL, et al. A HaloTag Anchored Ruler for Week-Long Studies of Protein Dynamics. J Am Chem Soc. 2016; 138(33):10546\u201353. [PubMed: 27409974]<\/p>\n<p>50. Bustamante C, Marko JF, Siggia ED, Smith S. Entropic elasticity of lambda-phage DNA. Science. 1994; 265(5178):1599\u20131600. [PubMed: 8079175]<\/p>\n<p>51. Rief M, Clausen-Schaumann H, Gaub HE. Sequence-dependent mechanics of single DNA molecules. Nat Struct Mol Biol. 1999; 6(4):346\u201349. <\/p>\n<p>52. Rief M, Oesterhelt F, Heymann B, Gaub HE. Single Molecule Force Spectroscopy on Polysaccharides by Atomic Force Microscopy. Science. 1997; 275(5304):1295\u201397. [PubMed: 9036852]<\/p>\n<p>53. Marszalek PE, Oberhauser AF, Pang Y-P, Fernandez JM. Polysaccharide elasticity governed by chair\u2013boat transitions of the glucopyranose ring. Nature. 1998; 396(6712):661\u201364. [PubMed: 9872313]<\/p>\n<p>54. Florin EL, Moy VT, Gaub HE. Adhesion forces between individual ligand-receptor pairs. Science. 1994; 264(5157):415\u201317. [PubMed: 8153628]<\/p>\n<p>55. Radmacher M, Fritz M, Hansma HG, Hansma PK. Direct observation of enzyme activity with the atomic force microscope. Science. 1994; 265(5178):1577\u201379. [PubMed: 8079171]<\/p>\n<p>56. Carrion-Vazquez M, Marszalek PE, Oberhauser AF, Fernandez JM. Atomic force microscopy captures length phenotypes in single proteins. Proc Natl Acad Sci. 1999; 96(20):11288\u201392. [PubMed: 10500169]<\/p>\n<p>57. Valle-Orero J, Rivas-Pardo JA, Tapia-Rojo R, Popa I, Echelman DJ, et al. Mechanical Deformation Accelerates Protein Ageing. Angew Chem Int Ed. n\/a-n\/a. <\/p>\n<p>58. Oberhauser AF, Hansma PK, Carrion-Vazquez M, Fernandez JM. Stepwise unfolding of titin under force-clamp atomic force microscopy. Proc Natl Acad Sci. 2001; 98(2):468\u201372. [PubMed: 11149943]<\/p>\n<p>59. Wiita AP, Ainavarapu SRK, Huang HH, Fernandez JM. Force-dependent chemical kinetics of disulfide bond reduction observed with single-molecule techniques. Proc Natl Acad Sci. 2006; 103(19):7222\u201327. [PubMed: 16645035]<\/p>\n<p>60. Wiita AP, Perez-Jimenez R, Walther KA, Gr\u00e4ter F, Berne BJ, et al. Probing the chemistry of thioredoxin catalysis with force. Nature. 2007; 450(7166):124\u201327. [PubMed: 17972886]<\/p>\n<p>61. Ainavarapu SRK, Bruji\u0107 J, Huang HH, Wiita AP, Lu H, et al. Contour Length and Refolding Rate of a Small Protein Controlled by Engineered Disulfide Bonds. Biophys J. 2007; 92(1):225\u201333. [PubMed: 17028145]<\/p>\n<p>62. Sotomayor M, Schulten K. Single-Molecule Experiments in Vitro and in Silico. Science. 2007; 316(5828):1144\u201348. [PubMed: 17525328]<\/p>\n<p>63. Flory, PJ. Principles of Polymer Chemistry. Cornell University Press; 1953. p. 696 <\/p>\n<p>64. Stacklies W, Vega MC, Wilmanns M, Gr\u00e4ter F. Mechanical Network in Titin Immunoglobulin from Force Distribution Analysis. PLOS Comput Biol. 2009; 5(3):e1000306. [PubMed: 19282960]<\/p>\n<p>65. Yuan G, Le S, Yao M, Qian H, Zhou X, et al. Elasticity of the Transition State Leading to an Unexpected Mechanical Stabilization of Titin Immunoglobulin Domains. Angew Chem Int Ed. 2017 n\/a-n\/a. <\/p>\n<p>66. Berkovich R, Garcia-Manyes S, Klafter J, Urbakh M, Fern\u00e1ndez JM. Hopping around an entropic barrier created by force. Biochem Biophys Res Commun. 2010; 403(1):133\u201337. [PubMed: 21050839]<\/p>\n<p>67. Garcia-Manyes S, Dougan L, Badilla CL, Bruji\u0107 J, Fern\u00e1ndez JM. Direct observation of an ensemble of stable collapsed states in the mechanical folding of ubiquitin. Proc Natl Acad Sci. 2009; 106(26):10534\u201339. [PubMed: 19541635]<\/p>\n<p>68. Redfield C, Smith RAG, Dobson CM. Structural characterization of a highly\u2013ordered \u201cmolten globule\u201d at low pH. Nat Struct Mol Biol. 1994; 1(1):23\u201329. <\/p>\n<p>69. M\u00e1rtonfalvi Z, Bianco P, Naftz K, Ferenczy GG, Kellermayer M. Force generation by titin folding. Protein Sci. 2017 n\/a-n\/a. <\/p>\n<p>70. Eckels EC, Rivas-Pardo JA, Valle-Orero J, Popa I, Fernandez JM. The Science of Stretching: Mechanical Anisotropy in Titin Ig Domains. Biophys J. 2016; 110(3):393a. <\/p>\n<p>71. Li H, Wang H-C, Cao Y, Sharma D, Wang M. Configurational Entropy Modulates the Mechanical Stability of Protein GB1. J Mol Biol. 2008; 379(4):871\u201380. [PubMed: 18472109]<\/p>\n<p>72. Li H, Carrion-Vazquez M, Oberhauser AF, Marszalek PE, Fernandez JM. Point mutations alter the mechanical stability of immunoglobulin modules. Nat Struct Mol Biol. 2000; 7(12):1117\u201320. <\/p>\n<p>73. Berkovich R, Hermans RI, Popa I, Stirnemann G, Garcia-Manyes S, et al. Rate limit of protein elastic response is tether dependent. Proc Natl Acad Sci. 2012; 109(36):14416\u201321. [PubMed: 22895787]<\/p>\n<p>74. Berkovich R, Garcia-Manyes S, Urbakh M, Klafter J, Fernandez JM. Collapse Dynamics of Single Proteins Extended by Force. Biophys J. 2010; 98(11):2692\u20132701. [PubMed: 20513414]<\/p>\n<p>75. Neupane K, Woodside MT. Quantifying Instrumental Artifacts in Folding Kinetics Measured by Single-Molecule Force Spectroscopy. Biophys J. 2016; 111(2):283\u201386. [PubMed: 27369870]<\/p>\n<p>76. Brenner H. The slow motion of a sphere through a viscous fluid towards a plane surface. Chem Eng Sci. 1961; 16(3\u20134):242\u201351. <\/p>\n<p>77. Valle-Orero J, Eckels EC, Stirnemann G, Popa I, Berkovich R, Fernandez JM. The elastic free energy of a tandem modular protein under force. Biochem Biophys Res Commun. 2015; 460(2): 434\u201338. [PubMed: 25796331]<\/p>\n<p>78. Alegre-Cebollada J, Kosuri P, Giganti D, Eckels E, Rivas-Pardo JA, et al. S-Glutathionylation of Cryptic Cysteines Enhances Titin Elasticity by Blocking Protein Folding. Cell. 2014; 156(6): 1235\u201346. [PubMed: 24630725]<\/p>\n<p>79. Li H, Fernandez JM. Mechanical Design of the First Proximal Ig Domain of Human Cardiac Titin Revealed by Single Molecule Force Spectroscopy. J Mol Biol. 2003; 334(1):75\u201386. [PubMed: 14596801]<\/p>\n<p>80. Pardo JAR, Eckels E, Valle-Orero J, Fernadez JM. Disulfide Bonding in the Contractile Work of Titin. Biophys J. 2017; 112(3):456a. <\/p>\n<p>81. Hill AV. The Heat of Shortening and the Dynamic Constants of Muscle. Proc R Soc Lond B Biol Sci. 1938; 126(843):136\u201395. <\/p>\n<p>82. Debold EP, Patlak JB, Warshaw DM. Slip Sliding Away: Load-Dependence of Velocity Generated by Skeletal Muscle Myosin Molecules in the Laser Trap. Biophys J. 2005; 89(5):L34\u201336. [PubMed: 16169988]<\/p>\n<p>83. Visscher K, Schnitzer MJ, Block SM. Single kinesin molecules studied with a molecular force clamp. Nature. 1999; 400(6740):184\u201389. [PubMed: 10408448]<\/p>\n<p>84. Clemen AE-M, Vilfan M, Jaud J, Zhang J, B\u00e4rmann M, Rief M. Force-Dependent Stepping Kinetics of Myosin-V. Biophys J. 2005; 88(6):4402\u201310. [PubMed: 15764664]<\/p>\n<p>85. Gennerich A, Carter AP, Reck-Peterson SL, Vale RD. Force-Induced Bidirectional Stepping of Cytoplasmic Dynein. Cell. 2007; 131(5):952\u201365. [PubMed: 18045537]<\/p>\n<p>86. Linke WA, Stockmeier MR, Ivemeyer M, Hosser H, Mundel P. Characterizing titin\u2019s I-band Ig domain region as an entropic spring. J Cell Sci. 1998; 111(11):1567\u201374. [PubMed: 9580564]<\/p>\n<p>87. Prado LG, Makarenko I, Andresen C, Kr\u00fcger M, Opitz CA, Linke WA. Isoform Diversity of Giant Proteins in Relation to Passive and Active Contractile Properties of Rabbit Skeletal Muscles. J Gen Physiol. 2005; 126(5):461\u201380. [PubMed: 16230467]<\/p>\n<p>88. Campbell KS. Dynamic coupling of regulated binding sites and cycling myosin heads in striated muscle. J Gen Physiol. 2014; 143(3):387\u201399. [PubMed: 24516189]<\/p>\n<p>89. Piazzesi G, Reconditi M, Linari M, Lucii L, Bianco P, et al. Skeletal Muscle Performance Determined by Modulation of Number of Myosin Motors Rather Than Motor Force or Stroke Size. Cell. 2007; 131(4):784\u201395. [PubMed: 18022371]<\/p>\n<p>90. Llewellyn ME, Barretto RPJ, Delp SL, Schnitzer MJ. Minimally invasive high-speed imaging of sarcomere contractile dynamics in mice and humans. Nature. 2008; 454(7205):784\u201388. [PubMed: 18600262]<\/p>\n<p>91. Rassier DE, MacIntosh BR, Herzog W. Length dependence of active force production in skeletal muscle. J Appl Physiol. 1999; 86(5):1445\u20131457. [PubMed: 10233103]<\/p>\n<p>92. Cromie MJ, Sanchez GN, Schnitzer MJ, Delp SL. Sarcomere lengths in human extensor carpi radialis brevis measured by microendoscopy. Muscle Nerve. 2013; 48(2):286\u201392. [PubMed: 23813625]<\/p>\n<p>93. Sanchez GN, Sinha S, Liske H, Chen X, Nguyen V, et al. In Vivo Imaging of Human Sarcomere Twitch Dynamics in Individual Motor Units. Neuron. 2015; 88(6):1109\u201320. [PubMed: 26687220]<\/p>\n<p>94. Gittes F, Mickey B, Nettleton J, Howard J. Flexural rigidity of microtubules and actin filaments measured from thermal fluctuations in shape. J Cell Biol. 1993; 120(4):923\u201334. [PubMed: 8432732]<\/p>\n<p>95. Fudge DS, Gardner KH, Forsyth VT, Riekel C, Gosline JM. The Mechanical Properties of Hydrated Intermediate Filaments: Insights from Hagfish Slime Threads. Biophys J. 2003; 85(3): 2015\u201327. [PubMed: 12944314]<\/p>\n<p>96. Lovelady HH, Shashidhara S, Matthews WG. Solvent specific persistence length of molecular type I collagen. Biopolymers. 2014; 101(4):329\u201335. [PubMed: 23907817]<\/p>\n<p>97. Hidalgo C, Hudson B, Bogomolovas J, Zhu Y, Anderson B, et al. PKC Phosphorylation of Titin\u2019s PEVK Element A Novel and Conserved Pathway for Modulating Myocardial Stiffness. Circ Res. 2009; 105(7):631\u201338. [PubMed: 19679839]<\/p>\n<p>98. K\u00f6tter S, Gout L, Frieling-Salewsky MV, M\u00fcller AE, Helling S, et al. Differential changes in titin domain phosphorylation increase myofilament stiffness in failing human hearts. Cardiovasc Res. 2013; 99(4):648\u201356. [PubMed: 23764881]<\/p>\n<p>99. Perkin J, Slater R, Del Favero G, Lanzicher T, Hidalgo C, et al. Phosphorylating Titin\u2019s Cardiac N2B Element by ERK2 or CaMKII\u03b4 Lowers the Single Molecule and Cardiac Muscle Force. Biophys J. 2015; 109(12):2592\u20132601. [PubMed: 26682816]<\/p>\n<p>100. Avner BS, Shioura KM, Scruggs SB, Grachoff M, Geenen DL, et al. Myocardial infarction in mice alters sarcomeric function via post-translational protein modification. Mol Cell Biochem. 2012; 363(1\u20132):203\u201315. [PubMed: 22160857]<\/p>\n<p>101. Li Y, Linke WA. Mechanically Unfolded Titin Immunoglobulin Domains Refold Faster and More Accurately in Presence of Chaperone Alpha-B-Crystallin. Biophys J. 2017; 112(3):42a. <\/p>\n<p>102. Kotter S, Unger A, Hamdani N, Lang P, Vorgerd M, et al. Human myocytes are protected from titin aggregation-induced stiffening by small heat shock proteins. J Cell Biol. 2014; 204(2):187\u2013 202. [PubMed: 24421331]<\/p>\n<p>103. Bullard B, Ferguson C, Minajeva A, Leake MC, Gautel M, et al. Association of the Chaperone \u03b1B-crystallin with Titin in Heart Muscle. J Biol Chem. 2004; 279(9):7917\u201324. [PubMed: 14676215]<\/p>\n<p>104. Brinkmeier H, Ohlendieck K. Chaperoning heat shock proteins: Proteomic analysis and relevance for normal and dystrophin-deficient muscle. PROTEOMICS \u2013 Clin Appl. 2014; 8(11\u201312):875\u2013 95. [PubMed: 24895218]<\/p>\n<p>105. Zhu Y, Bogomolovas J, Labeit S, Granzier H. Single molecule force spectroscopy of the cardiac titin N2B element: effects of the molecular chaperone alphaB-crystallin with disease-causing mutations. J Biol Chem. 2009; 284(20):13914\u201323. [PubMed: 19282282]<\/p>\n<p>106. Roach NT, Venkadesan M, Rainbow MJ, Lieberman DE. Elastic energy storage in the shoulder and the evolution of high-speed throwing in Homo. Nature. 2013; 498(7455):483\u201386. [PubMed: 23803849]<\/p>\n<p>107. Minajeva A, Neagoe C, Kulke M, Linke WA. Titin-based contribution to shortening velocity of rabbit skeletal myofibrils. J Physiol. 2002; 540(1):177\u201388. [PubMed: 11927678]<\/p>\n<p>108. Opitz CA, Kulke M, Leake MC, Neagoe C, Hinssen H, et al. Damped elastic recoil of the titin spring in myofibrils of human myocardium. Proc Natl Acad Sci. 2003; 100(22):12688\u201393. [PubMed: 14563922]<\/p>\n<p>109. Dugdale TM, Dagastine R, Chiovitti A, Mulvaney P, Wetherbee R. Single Adhesive Nanofibers from a Live Diatom Have the Signature Fingerprint of Modular Proteins. Biophys J. 2005; 89(6): 4252\u201360. [PubMed: 16169972]<\/p>\n<p>110. Sarkar A, Caamano S, Fernandez JM. The Mechanical Fingerprint of a Parallel Polyprotein Dimer. Biophys J. 2007; 92(4):L36\u201338. [PubMed: 17158577]<\/p>\n<p>111. Bianco P, Reconditi M, Piazzesi G, Lombardi V. Is muscle powered by springs or motors? J Muscle Res Cell Motil. 2016:1\u20133. [PubMed: 26842420]<\/p>\n<p>112. Caremani M, Pinzauti F, Reconditi M, Piazzesi G, Stienen GJM, et al. Size and speed of the working stroke of cardiac myosin in situ. Proc Natl Acad Sci. 2016; 113(13):3675\u201380. [PubMed: 26984499]<\/p>\n<p>113. Takano M, Terada TP, Sasai M. Unidirectional Brownian motion observed in an in silico single molecule experiment of an actomyosin motor. Proc Natl Acad Sci. 2010; 107(17):7769\u201374. [PubMed: 20385833]<\/p>\n<p>114. Freikamp A, Cost A-L, Grashoff C. The Piconewton Force Awakens: Quantifying Mechanics in Cells. Trends Cell Biol. 2016; 26(11):838\u201347. [PubMed: 27544876]<\/p>\n<p>115. Meng F, Sachs F. Orientation-based FRET sensor for real-time imaging of cellular forces. J Cell Sci. 2012; 125(3):743\u201350. [PubMed: 22389408]<\/p>\n<p>116. Austen K, Ringer P, Mehlich A, Chrostek-Grashoff A, Kluger C, et al. Extracellular rigidity sensing by talin isoform-specific mechanical linkages. Nat Cell Biol. 2015; 17(12):1597\u20131606. [PubMed: 26523364]<\/p>\n<p>117. Grashoff C, Hoffman BD, Brenner MD, Zhou R, Parsons M, et al. Measuring mechanical tension across vinculin reveals regulation of focal adhesion dynamics. Nature. 2010; 466(7303):263\u201366. [PubMed: 20613844]<\/p>\n<p>118. Arsenovic PT, Ramachandran I, Bathula K, Zhu R, Narang JD, et al. Nesprin-2G, a Component of the Nuclear LINC Complex, Is Subject to Myosin-Dependent Tension. Biophys J. 2016; 110(1): 34\u201343. [PubMed: 26745407]<\/p>\n<p>119. Brenner MD, Zhou R, Conway DE, Lanzano L, Gratton E, et al. Spider Silk Peptide Is a Compact, Linear Nanospring Ideal for Intracellular Tension Sensing. Nano Lett. 2016; 16(3): 2096\u20132102. [PubMed: 26824190]<\/p>\n<p>120. Prosser BL, Ward CW, Lederer WJ. X-ROS Signaling: Rapid Mechano-Chemo Transduction in Heart. Science. 2011; 333(6048):1440\u201345. [PubMed: 21903813]<\/p>\n<p>121. Rivas-Pardo JA, M\u00e1rtonfalvi Z, Manteca A, Eckels EC, Echelman DJ, et al. A Multi-Tool Mouse Model to Study the Elasticity of Native Titin. Biophys J. 2017; 112(3):167a. <\/p>\n<p>122. Kohl J, Ng J, Cachero S, Ciabatti E, Dolan M-J, et al. Ultrafast tissue staining with chemical tags. Proc Natl Acad Sci. 2014; 111(36):E3805\u201314. [PubMed: 25157152]<\/p>\n<p>123. Stagge F, Mitronova GY, Belov VN, Wurm CA, Jakobs S. Snap-, CLIP- and Halo-Tag Labelling of Budding Yeast Cells. PLoS ONE. 2013; 8(10):e78745. [PubMed: 24205303]<\/p>\n<p>124. Horowits R, Kempner ES, Bisher ME, Podolsky RJ. A physiological role for titin and nebulin in skeletal muscle. Nature. 1986; 323(6084):160\u201364. [PubMed: 3755803]<\/p>\n<p>125. Higuchi H. Changes in Contractile Properties with Selective Digestion of Connectin (Titin) in Skinned Fibers of Frog Skeletal Muscle. J Biochem (Tokyo). 1992; 111(3):291\u201395. [PubMed: 1587789]<\/p>\n<p>126. Dobbie I, Linari M, Piazzesi G, Reconditi M, Koubassova N, et al. Elastic bending and active tilting of myosin heads during muscle contraction. Nature. 1998; 396(6709):383\u201387. [PubMed: 9845077]<\/p>\n<p>127. Piazzesi G, Reconditi M, Linari M, Lucii L, Sun Y-B, et al. Mechanism of force generation by myosin heads in skeletal muscle. Nature. 2002; 415(6872):659\u201362. [PubMed: 11832949]<\/p>\n<p>128. Reconditi M, Linari M, Lucii L, Stewart A, Sun Y-B, et al. The myosin motor in muscle generates a smaller and slower working stroke at higher load. Nature. 2004; 428(6982):578\u201381. [PubMed: 15058307]<\/p>\n<p>129. Fenn WO. A quantitative comparison between the energy liberated and the work performed by the isolated sartorius muscle of the frog. J Physiol. 1923; 58(2\u20133):175\u2013203. 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