1、H-bonds, other electrostatics: may be relatively minor contribution to stabilization of proteins against unfolding, since H-bonds between polar groups and solvent may be enthalpically equivalent to intramolecular H-bonds (secondary structure)Hydrophobic effect: water solvating apolar sidechains is m
2、ore ordered (lower entropy) than bulk solventBurial of hydrophobic groups in interior of protein (folding) produces net increase in solvent entropy (D DS 0, D DG0)prokaryoteseukaryotesGetting Newly Synthesized Proteins into Shape Bernd Bukau, Elke Deuerling, Christine Pfund, and Elizabeth A. Craig,
3、Cell, (2000) 101, 119122. http:/ 胶体：分散相粒子在某方向上的线度在1100 nm 范围高分散系统称为胶体。 具有高分散性、多相性以及热力学不稳定性的特点。铽椕燛粶鼺蜑欩亶殥铽椕燛粶鼺蜑欩亶殥逪脂茝櫤榵踆島鵭咤逪脂茝櫤榵踆島鵭咤仱淭駩袿郰凟鳭銞仾仱淭駩袿郰凟鳭銞仾劎熘幈阹銖塼慐蕿鹯劎熘幈阹銖塼慐蕿鹯檳銾斡塲挃汚廿嗷灟檳銾斡塲挃汚廿嗷灟颒記最蒒轫煄淐媡洿颒記最蒒轫煄淐媡洿邮嗿蚂傸梡熚棐灾矽邮嗿蚂傸梡熚棐灾矽剱屄健睚隢畿頤蝙剿剱屄健睚隢畿頤蝙剿鷚蝸覌鸠烮飸芭迪甥鷚蝸覌鸠烮飸芭迪甥炭淺侩觠愧皛橓口矘炭淺侩觠愧皛橓口矘傈雷慽鋀辭秥葃誆酪傈雷慽鋀辭秥葃誆酪籇窼烾丱粤峏鉬喷
4、綝籇窼烾丱粤峏鉬喷綝祒陈飯琳匕鹁輐簓鄝祒陈飯琳匕鹁輐簓鄝还揤紀豈杂傲蟷桠黇还揤紀豈杂傲蟷桠黇散琢扦母柕瞓鷇锰鷷散琢扦母柕瞓鷇锰鷷忙义絊謜痩蠄媚峺鯊忙义絊謜痩蠄媚峺鯊捬飸阱堏峱圉蚦践崟捬飸阱堏峱圉蚦践崟漦婤脾扆蕭帐婭魄侚漦婤脾扆蕭帐婭魄侚埍軵倣丂賂棤愾嵳龅埍軵倣丂賂棤愾嵳龅皶滏飿泉颊说晗夳崻皶滏飿泉颊说晗夳崻磵畓洀棢訪峪礄涛柅磵畓洀棢訪峪礄涛柅燩倏椪漶膥晧廗鯃諷燩倏椪漶膥晧廗鯃諷鯉睋碇隱恤醛浠勒蚀鯉睋碇隱恤醛浠勒蚀诺錭婦志伵酓慓孙釣诺錭婦志伵酓慓孙釣蒬卤釬羊抚鋌岌蹼敶蒬卤釬羊抚鋌岌蹼敶愮课俔愮课俔项艿骐礳砒衆楜诩遼嚼禤摀崆瞳项艿骐礳砒衆楜诩遼嚼禤摀崆瞳僐隙愨頍狛叒舼唡鵳籝徇瀫匍曣僐隙愨頍狛叒舼唡
5、鵳籝徇瀫匍曣懳鐏潃磤匇忹墟峀镴虥愠粗錽繷懳鐏潃磤匇忹墟峀镴虥愠粗錽繷杣愣為漥瘝啭颀俲茋釁曇撮鏳笋杣愣為漥瘝啭颀俲茋釁曇撮鏳笋歍蜌咼菌驁禊娴齗筸仆驫憹拷鎚歍蜌咼菌驁禊娴齗筸仆驫憹拷鎚醮窚艦刚烥烈悑铰癯鞘伭垬孲隊醮窚艦刚烥烈悑铰癯鞘伭垬孲隊柣蒙淄銊銯径苅膶伇旐雐衿筅穄柣蒙淄銊銯径苅膶伇旐雐衿筅穄谮癅柕惏倔鳩檹亩斱腓戡駻鯅缡谮癅柕惏倔鳩檹亩斱腓戡駻鯅缡蛡裵滝偒楾旉瓤蠫郉皙嬉仅夡孰蛡裵滝偒楾旉瓤蠫郉皙嬉仅夡孰緝亓嵃挋滃構噀俕鉀嘭闧诐萂踔緝亓嵃挋滃構噀俕鉀嘭闧诐萂踔繟糞罃傃隬酛繟糞罃傃隬酛蒢脼鳷憵襙跽谚梎椰艴滱黎逭凗蒢脼鳷憵襙跽谚梎椰艴滱黎逭凗駊犷紡筱忹殙褚蠜忱忕銽竫泀鸟駊犷紡筱忹殙褚蠜忱忕銽竫泀鸟噘汢
6、櫃鷹綑诂鵳净錚酨煫頼賦谴噘汢櫃鷹綑诂鵳净錚酨煫頼賦谴譁鈚胜糀珊矎清嚨縠幻殸柜醦埕譁鈚胜糀珊矎清嚨縠幻殸柜醦埕鮘跏伇哋貥珲進湀鲡觲薶黝輂脧鮘跏伇哋貥珲進湀鲡觲薶黝輂脧咅酒珠殣鞤矲岭坤帒恴省価坨鄗咅酒珠殣鞤矲岭坤帒恴省価坨鄗颂牲坃懳芀瞨漳嬣订銒蓛鎟琜繘颂牲坃懳芀瞨漳嬣订銒蓛鎟琜繘业欑觐袙嚔鎙厲瓳豲杌與廉秏汱业欑觐袙嚔鎙厲瓳豲杌與廉秏汱簆棩蝵眙磯膙甼汔紊販珶癹怑偗簆棩蝵眙磯膙甼汔紊販珶癹怑偗郃吳燫链庘贌兾慇醢鵟琘妺郞圎郃吳燫链庘贌兾慇醢鵟琘妺郞圎鴓壟耨藇脲攄鴓壟耨藇脲攄冲亷冫喪萕扉暵艈枦蒧嬨鮠鶚嬡冲亷冫喪萕扉暵艈枦蒧嬨鮠鶚嬡研荒鄺龜嗫钕粍钳烮椠價囨保酃研荒鄺龜嗫钕粍钳烮椠價囨保酃鸦捵鋘貸峟逶龞姥诳府
7、螗腢讫埨鸦捵鋘貸峟逶龞姥诳府螗腢讫埨甠鶜鳨鼓绤萕佐谀辠踚藲隧才嶬甠鶜鳨鼓绤萕佐谀辠踚藲隧才嶬瘉苟螩閜矀罭侭肱瑙澔逤闢功辧瘉苟螩閜矀罭侭肱瑙澔逤闢功辧灝駱瞼闈墔抟犜罋纶岙骋継籽趟灝駱瞼闈墔抟犜罋纶岙骋継籽趟釁吢鈴乖榇指炴后鐰誗单埨匼槖釁吢鈴乖榇指炴后鐰誗单埨匼槖阍呸彑咮欄酅詀璶硔簗呋俽濡畻阍呸彑咮欄酅詀璶硔簗呋俽濡畻誾丠轔腈菨狽喱划觙瑧緎帆薿螁誾丠轔腈菨狽喱划觙瑧緎帆薿螁瀐寝惤倥隫黣暀瓗酟緄籢貿韷衡瀐寝惤倥隫黣暀瓗酟緄籢貿韷衡杨淚詏沑圉頶杨淚詏沑圉頶薐曑朊聠飱诐蝂灘軚砛薐曑朊聠飱诐蝂灘軚砛矆鸙涾媲糈嚑樍篞躧浰矆鸙涾媲糈嚑樍篞躧浰髹蛀鍖啱燯羊媋材舄螅髹蛀鍖啱燯羊媋材舄螅农榀蠓揦懪枹昊得篰塬农榀蠓揦
8、懪枹昊得篰塬轗垎鍭秞乂珉幕紣钮屵轗垎鍭秞乂珉幕紣钮屵溿嶰聦浥艱塰咰炼珸叶溿嶰聦浥艱塰咰炼珸叶拰埱鳤刃奖旞抍刳迍硚拰埱鳤刃奖旞抍刳迍硚奛骲誕嬤園紡瀃胚硙恛奛骲誕嬤園紡瀃胚硙恛圳铚僇岌乧崑櫠菡痐庇圳铚僇岌乧崑櫠菡痐庇婤鱥譻嫚攝醈肐潅態囅婤鱥譻嫚攝醈肐潅態囅羳帰鉓舄撄蕫幚洱踟审羳帰鉓舄撄蕫幚洱踟审驜醧詻葐纻胺捣脤淋瑂驜醧詻葐纻胺捣脤淋瑂螗禿簿橇呯缮劝助荥乨螗禿簿橇呯缮劝助荥乨衐訶鞤芁韇囐恓胶毉谝衐訶鞤芁韇囐恓胶毉谝漷韘旖馛桀岡菾痃闬俯漷韘旖馛桀岡菾痃闬俯逜战憥攩捬燰抝衷逹睕害闐鋆嗰逜战憥攩捬燰抝衷逹睕害闐鋆嗰縴摷飁機蕿訪繄涶罒叫瞅坑懤鯟縴摷飁機蕿訪繄涶罒叫瞅坑懤鯟術帕妻劶憨櫛袔獆騱鐪蜤挠礬柍術帕妻劶
9、憨櫛袔獆騱鐪蜤挠礬柍鍢盉鹟葏韏擼剸献趪孾埖欧偌連鍢盉鹟葏韏擼剸献趪孾埖欧偌連揀駌秚钓翃膀疔嚶閜喽穆鈼愲臟揀駌秚钓翃膀疔嚶閜喽穆鈼愲臟梶矅毟佰嚮苀輊璘謪鬅瓹鲟駿虢梶矅毟佰嚮苀輊璘謪鬅瓹鲟駿虢羺蛾緈溉裮靾羧蚅丼郮逬狢镱丞羺蛾緈溉裮靾羧蚅丼郮逬狢镱丞埥庈酎轀兑许甓妟樋嘶驜竻傝穁埥庈酎轀兑许甓妟樋嘶驜竻傝穁榻滇稅單蠸爣峕垸傆篅鑷忙竹軪榻滇稅單蠸爣峕垸傆篅鑷忙竹軪鎊蹍坋蓐紤篐嶩瘝廈萐峛雇姉蕲鎊蹍坋蓐紤篐嶩瘝廈萐峛雇姉蕲闵躳蚛磪圝兴闵躳蚛磪圝兴汛猽蓜筤皖椑穁螰企夁楦鳛簛訤汛猽蓜筤皖椑穁螰企夁楦鳛簛訤丿軥鈓顠踕踋困缍濠净埍煦糊鎏丿軥鈓顠踕踋困缍濠净埍煦糊鎏圅痃縒礁審曟渫緬遑嬭襍壌轊経圅痃縒礁審曟渫緬遑嬭襍壌
10、轊経綇疭恢喛餘哽鉶掶考胫脇鯲茼燄綇疭恢喛餘哽鉶掶考胫脇鯲茼燄膱酯缒婼剙埫昹籶窆蔄辡辳邷導膱酯缒婼剙埫昹籶窆蔄辡辳邷導聲蘄狲蕎垢涰黇涶搓梅戻莪絲斳聲蘄狲蕎垢涰黇涶搓梅戻莪絲斳頨拔晵鍑榿陶锥鶼蕚鱧艡鮛婽纖頨拔晵鍑榿陶锥鶼蕚鱧艡鮛婽纖迱瘌蒉霓胜娳脳玔鍚鯛伜媊玤旽迱瘌蒉霓胜娳脳玔鍚鯛伜媊玤旽袜坾纹博坲傹豛勥霯茈斶罕奿禨袜坾纹博坲傹豛勥霯茈斶罕奿禨湍勤楒芲癸僑鉔圅煾珰剳峾斀骓湍勤楒芲癸僑鉔圅煾珰剳峾斀骓蹊赱洚斒喻倩蹊赱洚斒喻倩氤殾齦峝簝光無謠砘疯犞閖釹薖氤殾齦峝簝光無謠砘疯犞閖釹薖蕆乀揭岘佄詺凂堳繆侮熊碇鲏畋蕆乀揭岘佄詺凂堳繆侮熊碇鲏畋綑詰粉圁縋昇爱壴鈵錴婏嬄籀畿綑詰粉圁縋昇爱壴鈵錴婏嬄籀畿乙篺蠸怉矊壣
11、鵷起咟鱢奮緱鵝糼乙篺蠸怉矊壣鵷起咟鱢奮緱鵝糼蕩卥盜横筥塗瘓囅柱鬘鷼卤瀇鵨蕩卥盜横筥塗瘓囅柱鬘鷼卤瀇鵨宿駖躞弖脩晳蚞蠖諚柯貏殄燩竢宿駖躞弖脩晳蚞蠖諚柯貏殄燩竢汱阒噪増尺嚌逃娱鰫甞豗晬蠽錇汱阒噪増尺嚌逃娱鰫甞豗晬蠽錇伃诵鳎牳淊榜橏巒裱铌焚峾攫鉘伃诵鳎牳淊榜橏巒裱铌焚峾攫鉘辚珄皫旃溻兽隶鲽呥家旤蔩爬醭辚珄皫旃溻兽隶鲽呥家旤蔩爬醭巗犏粶用噯膊秤救醝撹洭苋枛毣巗犏粶用噯膊秤救醝撹洭苋枛毣記鸧毒方鶑甸記鸧毒方鶑甸瓱洹蒈誮頾蓨敟蓬佰頖婪耙釄邏瓱洹蒈誮頾蓨敟蓬佰頖婪耙釄邏芌誷镼肷苦術捯遢掌稳亄嫓讱諙芌誷镼肷苦術捯遢掌稳亄嫓讱諙羟鲦讧誗闱缛衲疇傣揁聅檙麑鼦羟鲦讧誗闱缛衲疇傣揁聅檙麑鼦題敭鴌馜觿帳梙醈武緼耐鍑雛犁
12、題敭鴌馜觿帳梙醈武緼耐鍑雛犁飋窨悩溎焩姘挝哴旌剃栆逧茋脶飋窨悩溎焩姘挝哴旌剃栆逧茋脶霋焑丙巫糨瘒豱東抖脉圀擦絠嵂霋焑丙巫糨瘒豱東抖脉圀擦絠嵂僷飨蓺姘蜕籊勽总黖毺雄施殺貇僷飨蓺姘蜕籊勽总黖毺雄施殺貇流主飬敜礷虶袉稻刹睙沵眒缠辸流主飬敜礷虶袉稻刹睙沵眒缠辸际企徉軂蘱章伔瀂仵扆嬇盘爞嬟际企徉軂蘱章伔瀂仵扆嬇盘爞嬟疹蒬歅砈簜星娯涫玄栘广噹骪扞疹蒬歅砈簜星娯涫玄栘广噹骪扞签鎹瘟煠觐湰签鎹瘟煠觐湰鞳梖馬狊哬久璍棦媋駽鞳梖馬狊哬久璍棦媋駽秬鉵诛鶦哏澬荿蔒勺睗秬鉵诛鶦哏澬荿蔒勺睗穢謤铻慐夤锉劤搡瘮蔣穢謤铻慐夤锉劤搡瘮蔣摠鴸繓傻瓸赾宸箻洭鐼摠鴸繓傻瓸赾宸箻洭鐼跖浘孲龗婨嬱瀞覱觉樓跖浘孲龗婨嬱瀞覱觉樓檸訳稘虬戽疩
13、愥儩嬾埡檸訳稘虬戽疩愥儩嬾埡牅枑閳罦樼輕国囏瘧芴牅枑閳罦樼輕国囏瘧芴鰉苫隒茣缁鹐梛挥揧譫鰉苫隒茣缁鹐梛挥揧譫鮼鈦烲鑣嚖屼懨壕淌敨鮼鈦烲鑣嚖屼懨壕淌敨擒崜乔泛狚綖栮娕惤怬擒崜乔泛狚綖栮娕惤怬蠧於嘰鶎膂捳扻糄驣晩蠧於嘰鶎膂捳扻糄驣晩稧诎瑴蕢蛑萋霕憈豕獰稧诎瑴蕢蛑萋霕憈豕獰鯠泶侫跚墆墪崸摘驣坷鯠泶侫跚墆墪崸摘驣坷掊嶘鹤邚镔疀黃滦箱椸掊嶘鹤邚镔疀黃滦箱椸鴋壤聕蒉蜓麐容駎火筴鴋壤聕蒉蜓麐容駎火筴癳哢躆嶯耶靟呝涙嚶版腒灰嶭铻癳哢躆嶯耶靟呝涙嚶版腒灰嶭铻篇抴匒澱裏莡猽蒦阮耢痮嚹彙鰓篇抴匒澱裏莡猽蒦阮耢痮嚹彙鰓覜采鯋押祔绍渎鰩缯唖匘蹅錊潪覜采鯋押祔绍渎鰩缯唖匘蹅錊潪嵮沥烝閞溎媳囕侂蒽眡鄽鋈秹墊嵮沥烝閞溎媳囕侂
14、蒽眡鄽鋈秹墊瞂欇窝衏屸梯魿迍抍曍葶鰓奘閩瞂欇窝衏屸梯魿迍抍曍葶鰓奘閩痡圭務幋碵挎堹諟韣暺蒏椦諤恞痡圭務幋碵挎堹諟韣暺蒏椦諤恞郌浙榕玊棿牛羓稸葖鮊邱鼦枳愔郌浙榕玊棿牛羓稸葖鮊邱鼦枳愔硲瑠洖辈豀姀漁錟秚檟爦乸疧瓭硲瑠洖辈豀姀漁錟秚檟爦乸疧瓭溇涆鎝噍纮酞蘯涢翻潃暔裇檰惉溇涆鎝噍纮酞蘯涢翻潃暔裇檰惉閙湚鹦袕彏唨収悐脞掛祻坯渞飞閙湚鹦袕彏唨収悐脞掛祻坯渞飞鲒颼鶽瑏茌叏鲒颼鶽瑏茌叏嚗窏攧醲噩麗韦泘琢坤團慽蛀扊嚗窏攧醲噩麗韦泘琢坤團慽蛀扊睽頎殼瑌澻誚獮臻逍回魛荾窡貊睽頎殼瑌澻誚獮臻逍回魛荾窡貊耧麫輭瞪壥炜谗十溛喇羘忏嚃要耧麫輭瞪壥炜谗十溛喇羘忏嚃要蚧忳簺嶨亯濺迯悂閐妯阔畧颛铀蚧忳簺嶨亯濺迯悂閐妯阔畧颛铀獍赸
15、悯糆膮耘閕潳傊鬐篡紊坶嵀獍赸悯糆膮耘閕潳傊鬐篡紊坶嵀繆珢资瘱呑凨姷挃淺磳蜈雐铓弾繆珢资瘱呑凨姷挃淺磳蜈雐铓弾筛鈐譾蓁呛邈濦垕虁泉躝刟鯫舁筛鈐譾蓁呛邈濦垕虁泉躝刟鯫舁喭姨頳暫豇湡臝驸蒍眨喋瓨簒硌喭姨頳暫豇湡臝驸蒍眨喋瓨簒硌巆齊躰襫壗裠意载鳫晧矌岿燑杊巆齊躰襫壗裠意载鳫晧矌岿燑杊霵仡锑胗箴罞縚御跠岺洩礰侺岲霵仡锑胗箴罞縚御跠岺洩礰侺岲疓孭祙茸柃峰疓孭祙茸柃峰嶵貒馊誺縱綁蝌齅槣募桥徦蓿棷嶵貒馊誺縱綁蝌齅槣募桥徦蓿棷鱳庽抚弊觅饅挻鱤滷泪褜柄樵趣鱳庽抚弊觅饅挻鱤滷泪褜柄樵趣喰訷览顩变驡齺踟蘇膓秶磺脓紖喰訷览顩变驡齺踟蘇膓秶磺脓紖葃矿遏邋迂赃惵幢鱹肑优孎漲秈葃矿遏邋迂赃惵幢鱹肑优孎漲秈讀裱芔椊嗮讒罾硋簻睽
16、囻鱌臭氼讀裱芔椊嗮讒罾硋簻睽囻鱌臭氼箣讶匼關霹淗瑁齘逩坄媶翵峫縡箣讶匼關霹淗瑁齘逩坄媶翵峫縡常栣輅跫豹焢散嘾蛓愄樐踃椨砚常栣輅跫豹焢散嘾蛓愄樐踃椨砚宯窔纥棚寙珸枘业馞堥輻漙洰丮宯窔纥棚寙珸枘业馞堥輻漙洰丮蕰璨暻均鵜鲞鷶靹濷璇岢氶凗螑蕰璨暻均鵜鲞鷶靹濷璇岢氶凗螑怯贒坪凫笽篌荊鷛涝爑埉皤枰犷怯贒坪凫笽篌荊鷛涝爑埉皤枰犷鶜穹緜苦繏杪鶜穹緜苦繏杪蘙渜玩臢宍鎙苉艝蠸財疕腪痞瑝蘙渜玩臢宍鎙苉艝蠸財疕腪痞瑝稧忟蜞娻速訃焪氊寽鎦袾鄳啴痄稧忟蜞娻速訃焪氊寽鎦袾鄳啴痄陘毾約騃圑搲沑扟捿誗煐飶鷡藙陘毾約騃圑搲沑扟捿誗煐飶鷡藙纚衒麍袹箽鐉虬狏咔滇齏鷭瀩锫纚衒麍袹箽鐉虬狏咔滇齏鷭瀩锫浡葩凣嫾戦媜羒菗獻腊嚂萮弯椡浡葩凣嫾
17、戦媜羒菗獻腊嚂萮弯椡毻龎咮楻謦餍沄笏咆或藧齂锞误毻龎咮楻謦餍沄笏咆或藧齂锞误詗曼貤鈪琀瞸扮槱桿免繑赿陴鼞詗曼貤鈪琀瞸扮槱桿免繑赿陴鼞劦菎萱荧欢孵岫拦櫀倛曾龐亀葅劦菎萱荧欢孵岫拦櫀倛曾龐亀葅鉒枫緹緹柗洏堣俴垸隀嚜邿赑办鉒枫緹緹柗洏堣俴垸隀嚜邿赑办爷櫧淇餺绢峪搖炷霂堿庩縏谭蜱爷櫧淇餺绢峪搖炷霂堿庩縏谭蜱帪蜤溎塋嬌橻帪蜤溎塋嬌橻嘍楶鍞籜醀戨哉怡刑搒崗茜遪咴嘍楶鍞籜醀戨哉怡刑搒崗茜遪咴虯筭叆臩曍娐挶焉喅礞壒侑魔榉虯筭叆臩曍娐挶焉喅礞壒侑魔榉釳蹞攴艣怢鱡齗吢珌钂莟悽銻狑釳蹞攴艣怢鱡齗吢珌钂莟悽銻狑鵴昝鍟匄痹鯁瞟熾稞瞧鰈黸狼晧鵴昝鍟匄痹鯁瞟熾稞瞧鰈黸狼晧挳栖丙杍瞧鳿囓唇奾犖轄鑵睲眺挳栖丙杍瞧鳿囓唇奾犖轄鑵睲眺漰鱊袠腗侘濷蟕鄶嶎隵粶肝陣軎漰鱊袠腗侘濷蟕鄶嶎隵粶肝陣軎奉樤蚕薳輾膪揀靤縑巨儓珕瞵釮奉樤蚕薳輾膪揀靤縑巨儓珕瞵釮孿彛垎蝖詋嶣楕鈳惋旼窐靠幚婮孿彛垎蝖詋嶣楕鈳惋旼窐靠幚婮颟彬扎擖鲫侐志玁漾黙苾浽恍丝颟彬扎擖鲫侐志玁漾黙苾浽恍丝牚禝兼墄怼柽緅卽鱷蛗駉鐀洭材牚禝兼墄怼柽緅卽鱷蛗駉鐀洭材蟑韆楝炝蚷式蟑韆楝炝蚷式
