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洛潴 铕汔龛玎鲨: 软耱栩篁 祛脲牦痦铋 徼铍钽梃 桁. ..蓓沐朦汔痄蜞 欣

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棱蝾疱翦疣 疣珙耠囗 揰___ __________ 2008 .

喻睇 皴牮弪囵 滂耨屦蜞鲨铐眍泐 耦忮蜞, 漕牝铕 徼铍钽梓羼觇 磬箨 .. 灭箸溴 1

吾 踵疣牝屦桉蜩赅 疣犷螓

朽犷蜞 镱疋磬 桉耠邃钼囗棹 戾踵龛珈钼 溴蝈痨桧圉梃 皴沆屙蝾 祛痿钽屙弪梓羼觐 镱脲 皴沆屙蜞鲨 镫钿钼铋 祗 漯铉铘桦.

填痿钽屙弪梓羼觐 镱脲 怆弪 铖眍忭 屐屙蝾 铐蝾沐龛, , 赅 翳玷麇耜 钺铖钺脲眄 钺豚耱, 纛痨桊箦蝰 疱珞朦蜞蝈 耠铈睇 觐铕滂龛痤忄眄 忡噼祛溴轳蜮栝 耦耱噔桴 邈 觐祜铐屙 - 沐眍, 蝠囗耜痂矧钼 徨腙钼. 蒡 忡噼祛溴轳蜮 镳桠钿 蝾祗, 黩 赅驿 觌弪赅 镱 耱囗钼栩 溴蝈痨桧桊钼囗眍, 恹徼疣 钿桧 桤 祉钽桴 忸珈铈睇 矬蝈 疣玮栩 (铃腩篑钼, 2005). 腻蝈痨桧圉 皴沆屙蝾 镳铊聃钿栩 祛痿钽屙弪梓羼觐 镱脲, 镳邃耱噔屐 耦犷 蝮 鬣耱 狃桀磬, 桤 觐蝾痤 狍潴 疣珙恻弪 玎痤潲忄 镱腩耜 (暑痤麝桧, 2002; Gilbert, 2003), 镳桠钿栩 铒疱溴脲龛 镱腩驽龛 沭囗桷 镟疣皴沆屙蝾 (Martinez-Arias and Lawrence, 1985). 澡玷麇耜桁 觐祜铐屙蜞扈 祛痿钽屙弪梓羼觐泐 镱 皴沆屙蜞鲨 怆 沐睇 皴沆屙蜞鲨 桴 镳钿箨螓 (耢. 疣玟咫 2.1).

蔓镱腠屙睇 觐祜脲犟 桉耠邃钼囗栝 猁 铕桢眚桊钼囗 磬 耦玟囗桢 铕桡桧嚯 戾蝾漕 镱塍麇龛 觐腓麇耱忮眄 溧眄 桤 桤钺疣驽龛 赅痱桧 耧疱耨梃 沐眍, 桤篦屙桢 铖钺屙眍耱彘 滂磬扈觇 耧疱耨梃 沐眍 皴沆屙蜞鲨, 爨蝈爨蜩麇耜铄 祛溴腓痤忄龛 耧疱耨梃 沐眍 gap 鲨觌圊 13 14, 蜞赕 耦玟囗桢 囹豚襦 镳铖蝠囗耱忮眄-怵屐屙眍 耧疱耨梃 沐眍.

1.1

狸蝮嚯铖螯 蝈禧 湾耢铗 磬 犷朦 钺 桧纛痨圉梃 疱泱蝾痦 忡噼祛溴轳蜮 沐眍 皴沆屙蜞鲨 (Akam, 1987; Ingham, 1988; 暑痤麝桧, 2002), 珥囗 戾踵龛珈圊 疱泱鲨 桴 沐眍, 铖钺屙眍耱, 戾踵龛珈圊 耧疱耨梃 沐眍 gap, 怆 溧脲觐 礤 镱腠. 蒡, 钿眍 耱铕铐 钺耥弪 礤镱腠铗铋 耧屦桁屙蜞朦睇 溧眄 疱泱蝾痦 忡噼祛溴轳蜮, 漯筱铋 - 戾蝾漕腩汨麇耜桁 蝠箐眍耱扈, 觐蝾瘥 忸珥桕帼, 觐沅 恹忸 疱泱蝾痦铎 忡噼祛溴轳蜮梃 祉钽铌脲蝾黜钽 铕汔龛珈 溴豚弪 磬 铖眍忮 赅麇耱忮眄钽 囗嚯桤 耧疱耨梃 沐磬 祗蜞眚钼. 署囹觐  蝠箐眍耱 祛骓 钺铉磬麒螯 赅 镳钺脲禧 礤镳铗桠铕鬻桠铖蜩, 箜桕嚯铖蜩 镱腠铗 镳邃镱豚汔屐钽 疱泱蝾痦钽 戾踵龛珈.

碾 漕赅玎蝈朦耱忄 礤镳铗桠铕鬻桠铖蜩 汨镱蝈蜩麇耜钽 疱泱蝾痦钽 戾踵龛珈 蝠遽箦蝰 篦弪 怅豚溧 怦艴 疱泱蝾痤 桉耠邃箦祛泐 沐磬. 杨怵屐屙睇 耧屦桁屙蜞朦睇 戾蝾潲, 钿磬觐, 桁妣 钽疣龛麇眄簋 忸珈铈眍耱 钿眍怵屐屙眍泐 囗嚯桤 怆龛 怦艴 疱泱蝾痤 磬 溧眄 沐, 镱耜铍 铐 铖眍忄睇 磬 囗嚯桤 赅痱桧 耧疱耨梃 祗蜞眚钼, 镱塍麇龛 铕汔龛珈钼 祗蜞鲨扈 犷脲 麇 蝠艴 疱泱蝾疣 鬣耱 怆弪 礤镳铖蝾 玎溧麇. 令脲 蝾泐, 沐眄 皴螯 祗蜞眚钼 镱 铒疱溴脲龛 耦耱铊 桤 礤镱腠钽, 桦 溴翦牝眍泐 磬犷疣 疱泱蝾痦 忡噼祛溴轳蜮栝 沐眍. 亦觇 钺疣珙, 恹忸潲 耱痼牝箴 沐眄铋 皴蜩 滂觐泐 蜩镟 镳桴钿栩 溴豚螯 磬 铖眍忄龛 溧眄 祉钽桴 耧屦桁屙蝾 祗蜞眚囔. 湾镳铗桠铕鬻桠铖螯 驽 恹怆屙睇 忡噼祛溴轳蜮栝 祛驽 猁螯 镳钿屐铐耱痂痤忄磬 蝾朦觐 镳 镳钼屦赍 桴 桧蜞牝眍 疣玮桠帼祚 铕汔龛珈, 耦溴疰帙屐 镱腠 磬犷 疱泱蝾痤.

酿筱 镳钺脲爨, 忸珥桕帼 镳 桧蝈痫疱蜞鲨 赅痱桧 耧疱耨梃 祗蜞眚钼, 耦耱铊 漕赅玎蝈朦耱忮 箜桕嚯铖蜩 戾踵龛珈 忡噼祛溴轳蜮, .. 漕赅玎蝈朦耱忮 蝾泐, 黩 溧眄铄 疱泱蝾痦铄 忡噼祛溴轳蜮桢 怆弪 镳禧, 礤 铒铖疱漕忄眄. 亦觇 漕赅玎蝈朦耱忄 蝠遽簋 镳钼邃屙 漕镱腠栩咫 耧屦桁屙蝾.

磬觐礤, 耋耱怏弪 趔礓囔屙蜞朦磬 镳钺脲爨 溴祛眈蝠圉梃 镱腠铗 恹怆屙睇 疱泱蝾痦 忡噼祛溴轳蜮栝. 腻轳蜮栩咫, 礤钺躅滂祛耱 爨蝈痂眈觇 沐眍 沐眍 gap - 翳玷麇耜桴 觐祜铐屙 祛痿钽屙弪梓羼觐泐 镱 皴沆屙蜞鲨 - 潆 镳噔桦铋 耧疱耨梃 沐眍 gap 礤 铉磬鬣弪 漕耱囹铟眍耱 钽 磬犷疣 沐眍. 镳桧鲨镥 漕赅玎蝈朦耱忸 漕耱嘌邈戾眚 - 镱耠邃钼囹咫 麇疱潴桢 祛痿铍钽梓羼觇 耱痼牝箴 蝈豚 漯铉铘桦 蝾黜铖蜩 恹怆屙眍泐 疱泱蝾痦钽 戾踵龛珈 礤忸珈铈眍 徨 疱觐眈蝠箨鲨 耔耱屐 ab initio 桤 铗溴朦睇, 躅痤 觐眚痤腓痼屐 觐祜铐屙蝾. 西邂桎眍, 黩 耦怵屐屙睇 耧屦桁屙蜞朦睇 戾蝾潲 礤 镱玮铍 镳钼羼蜩 蜞牦 疱觐眈蝠箨鲨, , 耠邃钼囹咫, 铐 漕腈磬 猁螯 镳钼邃屙 in silico 镱祛 爨蝈爨蜩麇耜钽 祛溴腓痤忄龛 麒耠屙睇 疣聍弪钼 (朽蝮 et al., 2005).

酿筱铋 忸镳铖, 钿桧 桤 襦禧 忄骓 狃桀腩汨, 耦耱铊 恹耥屙梃 戾踵龛珈 溴蝈痨桧圉梃 觌弪铌 鬣耱彘 玎痤潲. 铖眍忮 钽 戾踵龛珈 脲骅 嚓蜩忄鲨 蝈 桦 桧 沐眍 疣珥 觌弪赅, 黩 镳桠钿栩 忸珥桕眍忮龛 镳铖蝠囗耱忮眄 礤钿眍痤漤铋 赅痱桧 耧疱耨梃 沐眍 ("箸铕"桦 蜞 磬琨忄屐钽 镟蝌屦磬). 漯铉铘桦 溴蝈痨桧圉 皴沆屙蝾 铒疱溴弪 镱腩驽龛 镟疣皴沆屙蝽, 礤 皴沆屙蝽 沭囗桷. 吾疣珙忄龛 镟疣皴沆屙蝾 镳邃耱怏弪 纛痨桊钼囗棹 皴沆屙蝾 磬 犷脲 镱玟龛 耱噤 疣玮栩.

祛痿钽屙弪梓羼觐 镱脲 皴沆屙蜞鲨 皴沆屙蝽 镳屣囹蝈痦 钺疣珞 镱腩覃 耧疱耨梃 沐眍 segment polarity (Lawrence and Johnston, 1989; Martinez-Arias and Lawrence, 1985; Ingham and Martinez-Arias, 1992; 畦祗脲, 2003).

孰囫耔麇耜铄 钺耥屙桢 戾踵龛珈钼 溴蝈痨桧圉梃 猁腩 镳邃腩驽眍 1969 泐潴 .骂腼屦蝾, 耵铕祗腓痤忄怿桁 蝈铕棹 镱玷鲨铐眍 桧纛痨圉梃 (Wolpert, 1969; Wolpert, 1989).

杨汶囫眍 铋 蝈铕梃 耋潼徉 觌弪觇 铒疱溴弪 邋 镱腩驽龛屐 铒疱溴脲眄铎 镳铖蝠囗耱忮眄铎 镱脲 玎痤潲, 觐蝾痤 耋耱怏弪 沭噤桢眚 觐眦屙蝠圉梃 礤觐 耔沩嚯铋 祛脲牦臌, 磬琨忄屐铋 祛痿钽屙铎. 痒栩囗桢 桧纛痨圉梃 沭噤桢眚 祛痿钽屙 邋 桧蝈痫疱蜞鲨 镳桠钿 滂趑屦屙鲨圉梃 觌弪铌 蝾 桦 桧铎 磬镳噔脲龛 玎忤耔祛耱 铗 箴钼 觐眦屙蝠圉梃 祛痿钽屙.

漯铉铘桦 镳钿箨 爨蝈痂眈觐泐 觐铕滂磬蝽钽 沐磬 bcd 怆弪 觌囫耔麇耜桁 镳桁屦铎 祛痿钽屙 (Driever and Nsslein-Volhard, 1988b; Driever and Nsslein-Volhard, 1988a;

Ephrussi and St Johnston, 2004). 橡钼邃屙睇 镱耠邃礤 怵屐 沐礤蜩麇耜桢 蝈铕弪梓羼觇 桉耠邃钼囗 箨噻帼 磬 蝾, 黩 溴轳蜮桢 钿眍泐 腓 祛痿钽屙 Bcd 礤漕耱囹铟眍 潆 忸珥桕眍忮龛 镳铖蝠忭耱忮眄 礤铑漤铕钿眍 赅痱桧 耧疱耨梃 沐眍 犭囫蝾溴痨 漯铉铘桦. 磬 疣犷蝈 桉耠邃钼囗 滂磬扈赅 镱玷鲨铐桊钼囗 钺豚耱彘 耧疱耨梃 沐眍 皴沆屙蜞鲨, 磬 铖眍忮 镱塍麇眄 溧眄 钺耋驿噱蝰 噤尻忄蝽铖螯 觐眦屣蝮嚯铋 祛溴腓 .骂腼屦蜞 潆 钺耥屙 戾踵龛珈钼 纛痨桊钼囗 皴沆屙蝽钽 镳屣囹蝈痦 犭囫蝾溴痨 漯铉铘桦.

项龛爨龛 镳桧鲨镱 铕汔龛玎鲨 趔黻鲨铐桊钼囗 祛痿钽屙弪梓羼觐泐 镱 皴沆屙蜞鲨 蝠遽箦 溴蜞朦眍泐 觐腓麇耱忮眄钽 铒桉囗 滂磬扈觇 赅驿铋 桤 邈 觐祜铐屙.

湾耢铗 磬 恹耦牦 疣琊屮帼 耧铖钺眍耱, 戾蝾 耐 麒镱 蜞 驽, 赅 祉钽桢 漯筱桢 戾蝾潲 觐腓麇耱忮眄铋 铞屙觇 耧疱耨梃 沐眍 (觐腓麇耱忮眄 PCR, CAT assays), 桁邋 钽疣龛麇眄铄 镳桁屙屙桢 潆 疱龛 铋 玎溧麒. 腻腩 蝾, 黩 怦  戾蝾潲 桉镱朦珞 泐祛沐磬螓 觌弪铌 , 蜞觇 钺疣珙, 蝈 桧纛痨圉棹 钺 耧疱耨梃 沐眍 镳铖蝠囗耱忮. 襄瘃镥牝桠睇 怆弪 桉镱朦珙忄龛 桁祗眍綦箢疱聆屙蝽钽 爨痍桊钼囗 徼铍钽梓羼觇 爨牮铎铍尻箅 耦麇蜞龛 豚珏痦铋 觐眙铌嚯铋 扈牮铖觐镨彘, 觐蝾疣 镱玮铍弪 镱塍鬣螯 赅麇耱忮眄 鲨麴钼 桤钺疣驽龛 赅痱桧 耧疱耨梃 沐眍, 泐蝾恹 潆 桤怆鬻屙 觐腓麇耱忮眄 溧眄 钺 耧疱耨梃 矬蝈 觐祜蝈痦铋 钺疣犷蜿.

西邂桎眍, 黩 赅麇耱忸 爨蝈爨蜩麇耜钽 祛溴腓痤忄龛 玎忤耔 铗 赅麇耱忄 耧屦桁屙蜞朦睇 溧眄. 枢 猁腩 铗戾麇眍 .疏蜞眍 (Kitano, 2002) 桎遴朦睇 磬犷 溧眄 漕腈屙 猁螯 桉麇痫帼 镱 镱腠铗 铞屙桠噱禧 觐祜铐屙, 桤戾屐 镟疣戾蝠钼 镱 铛忄蝮 怵屐屙眍 滂磬扈觇, 蝾黜 觐腓麇耱忮眄铎 铗眍龛, 蜞赕 耔耱屐囹梓羼觇.

项耠邃礤 铉磬鬣弪, 黩 耧铖钺 镱塍麇龛 溧眄 疣珥 蜩镱 漕腈屙 漕矬耜囹 桴 耦汶囫钼囗眢 桧蝈沭圉棹. 疹  蝠遽钼囗 铟邂桎睇, 磬耱 怵屐 腓 礤祉钽桢 磬犷瘥 溧眄 桁 箐钼脲蜮铕 , 镱铎, 镱塍麇龛 蜞觐泐 磬犷疣 溧眄 镱 耧疱耨梃 沐眍 皴沆屙蜞鲨 桁邋 忄骓铄 珥圜屙桢.

蒴翦牝桠眍 桉镱朦珙忄龛 觐腓麇耱忮眄 溧眄 镱 耧疱耨梃 沐眍 皴沆屙蜞鲨 蝠遽箦 铕汔龛玎鲨 磬犷疣 溧眄 忤溴 徉琨 溧眄, 黩 钺羼镥麒 翦牝桠眍 躔囗屙桢 恹犷痍 桧纛痨圉梃, 蜞赕 钺脲泖栩 囗嚯桤 溧眄, 磬鲥脲眄 磬 恹怆屙桢 眍恹 徼铍钽梓羼觇 玎觐眍戾痦铖蝈 桤篦噱祛泐 镳铞羼襦 纛痨箅桊钼牦 眍恹 汨镱蝈 潆 桴 磬镳噔脲眄铋 镳钼屦觇 耧屦桁屙蜞. 悟戾蜩, 黩 镱 疋铄 耋蜩 蜞赅 徉玎 溧眄 狍溴 镳铖蝠囗耱忮眄-怵屐屙睇 囹豚耦 耧疱耨梃 沐眍, 镱耜铍 铐 躔囗栩 觐腓麇耱忮眄簋 桧纛痨圉棹 钺 耧疱耨梃 疣珥 蝾麝圊 祛痿钽屙弪梓羼觐泐 镱 疣珥 祛戾眚 怵屐屙.

1.2 皱腓 疣犷螓 皱腓 滂耨屦蜞鲨铐眍 疣犷螓 耦耱腓 蝾, 黩钺:

1. 朽琊噌铗囹 眍恹 觐礅彘屦睇 戾蝾 觐腓麇耱忮眄铋 铞屙觇 箴钼 耧疱耨梃 汶噔睇 觐祜铐屙 祛痿钽屙弪梓羼觐泐 镱 皴沆屙蜞鲨 漯铉铘桦 - 沐眍 皴沆屙蜞鲨 - 磬 铖眍忮 桤钺疣驽龛, 镱塍麇眄 镱祛 觐眙铌嚯 扈牮铖觐镱. 蒡铗 戾蝾 漕腈屙 怅膻鬣螯 皴沆屙蜞鲨 桤钺疣驽龛, 箐嚯屙桢 纛眍忸泐 耔沩嚯, 铒疱溴脲龛 忸琊囫蜞 狃桀磬, 镳铖蝠囗耱忮眄簋 疱汨耱疣鲨 赅痱桧 耧疱耨梃 桧蝈沭圉棹 溧眄.

2. 项塍麒螯 桉麇痫帼 镱 镱腠铗, 蝾黜 觐腓麇耱忮眄铎 铗眍龛 耔耱屐囹梓羼觇 磬犷 觐腓麇耱忮眄 溧眄 钺 耧疱耨梃 沐眍 皴沆屙蜞鲨 赅驿铎 漯 赅驿钽 桧滂忤潴嚯钽 狃桀磬, 蜞赕 嚯铐睇, 桧蝈沭桊钼囗睇 溧眄 钺 耧疱耨梃 赅驿钽 桤 14 沐眍 皴沆屙蜞鲨 赅驿铋 钺豚耱 狃桀磬 疣珥 祛戾眚 怵屐屙;

3. 如篦栩 滂磬扈牦 纛痨桊钼囗 钺豚耱彘 耧疱耨梃 沐眍 祛痿钽屙弪梓羼觐 镱脲 皴沆屙蜞鲨 矬蝈 铞屙觇 镱腩驽龛 赅驿铋 钺豚耱 疣珥 祛戾眚 怵屐屙;

4. 杨玟囹 爨蝈爨蜩麇耜簋 祛溴朦 戾踵龛珈钼 疱泱鲨 耧疱耨梃 沐眍 gap 鲨觌 溴脲龛 溴 14. 橡桁屙屙栩  祛溴朦 潆 in silico 疱觐眈蝠箨鲨 皴蜩 沐眍 ga 恹怆屙 戾踵龛珈钼, 钺羼镥麒忄桴 皲忤汨 沭囗桷 钺豚耱彘 耧疱耨梃 桴 沐眍;

5. 碾 钺脲泖屙 疣犷螓 蝈铕弪桕钼 徼铍钽钼 溧眄 耦玟囹 镳铖蝠囗耱忮眄钼疱戾眄铋 囹豚 耧疱耨梃 沐眍 皴沆屙蜞鲨 忤溴 疱鲨铐眍 徉琨 溧眄, 漕耱箫眍 皴蜩 软蝈痦弪.

1.3 吾牝 戾蝾潲 桉耠邃钼囗 务眍忭 钺牝铎 桉耠邃钼囗 怆腓顸 1580 狃桀眍 祗 滂觐泐 蜩镟 (Oregon-R), 铌疣眄 戾蝾漕 礤镳祛泐 桁祗眍綦箢疱聆屙蝽钽 爨痍桊钼囗 潆 恹怆屙 徨腙钼 镳钿箨蝾 爨蝈痂眈觇 觐铕滂磬蝽 沐眍 bicoid (bcd) caudal (cad), 沐眍 gap Krppel (Kr), knirps (kni), giant (gt), hunchback (hb), tailless (tll) 沐眍 pair-rule even-skipped (eve), fushitarazu (ftz), hairy (h), runt (run), odd-skipped (odd), sloppy-paired (slp) paired (prd). 橡桁屦眍 镱腩忤磬 桤 桴 狃桀眍 猁豚 蜞赕 镳铌疣磬 磬 汨耱铐钼 徨腙 潆 镱漶忮麒忄龛 钺豚耱彘 桤钺疣驽龛, 铗眍躐 漯囔. 4490 桤钺疣驽龛 赅痱桧 耧疱耨梃 恹 镥疱麒耠屙睇 沐眍 皴沆屙蜞鲨 狃桀眍 猁腓 镱塍麇睇 豚犷疣蝾痂 镳铘. 朽轫桷 (禹桠屦耔蝈 羊铐 琉箨, 沿) 镱祛 觐眙铌嚯 扈牮铖觐镱 Leica TCS4D Leica TS-SP2. 蒡 桤钺疣驽龛 桁妣 疣珈屦 1024 x 1024 镨犟咫彘; 潆 玎镨耔 耔沩嚯 桉镱朦珞弪 忸顸扈徼蝾恹 纛痨囹 磬耱痤殛 扈牮铖觐镟, 箪屙帼 箴钼屙 爨 疱汨耱痂痼屐铎 耔沩嚯, 蜞赕 镳铞邃箴 耜囗桊钼囗, 镱玮铍 镱塍麒螯 耦汶囫钼囗睇 溧眄 镱 耧疱耨梃 钿眍泐 沐磬 桧滂忤潴嚯 狃桀磬, 耜囗桊钼囗睇 疣珥 耧屦桁屙蜞 (Kosman et al., 1997; Kosman et al., 1998). 碾 耜囗桊钼囗 恹徼疣腓顸 狃桀睇 疣珥钽 忸琊囫蜞, 磬麒磬 10 鲨觌 漯钺脲龛 溴 觐眵 汔耱痼鲨彘.

碾 桤怆鬻屙 觐腓麇耱忮眄 溧眄 桤 桤钺疣驽龛 磬潴 铕桡桧嚯 戾蝾溧扈, 疣琊噌铗囗睇扈 溧眄铋 疣犷蝈, 桉镱朦珙忄腓 耱囗溧痱睇 戾蝾潲 钺疣犷蜿 桤钺疣驽龛 (Gonzalez and Woods, 2002).

项耜铍 耧疱耨 沐眍 皴沆屙蜞鲨, 汶噔睇 钺疣珙, 怆弪 趔黻鲨彘 镱腩驽龛 漯 怃铍 镥疱漤-玎漤彘 (A-P) 铖, 邋 疣耧疱溴脲龛 镱 蝈塍 狃桀磬 漕耱囹铟眍 躅痤 祛驽 猁螯 镳邃耱噔脲眍 钿眍 桤戾疱龛. 骡桎 钽, 漕耱囹铟睇 镳邃耱噔弪 囗嚯桤 觐腓麇耱忮眄 溧眄 桤 鲥眚疣朦眍 10% (镱 铖 y) 镱腩耜, 恹疱玎眄铋 磬镳噔脲龛 A-P, .. 怃铍 铖 x. 亦觇 钺疣珙, 赅驿铎 漯 耦镱耱噔弪 腓 邈 x-觐铕滂磬蜞, 赅痱桧 耧疱耨梃 桁妣 忤 沭圄桕钼, 铗疣驵桴 桤戾礤龛 耧疱耨梃 沐眍 怃铍 铖 x. 碾 铒桉囗 钺豚耱彘 钿眍戾痦铋 赅痱桧 耧疱耨梃 沐眍 桉镱朦珙忄腓 囡镳铌耔爨鲨 赈噤疣蜩黜 耧豚轫囔 桦 猁耱痤 桤猁蝾黜铄 溻铊黜铄 忮殁脲-镳孱狃噻钼囗桢 噔蝾爨蜩麇耜桁 铒疱溴脲龛屐 蝠遽箦祛泐 箴钼 溴觐祜铉桷梃 赅痱桧 耧疱耨梃 忤珞嚯 觐眚痤脲 赅麇耱忄 噔蝾爨蜩麇耜钽 恹溴脲龛 镳桤磬觐. 碾 箪屙屙 疣珈屦眍耱 忮牝铕 镳桤磬觐 镳桁屙腓 羿牝铕睇 囗嚯桤.

唁忤汨 钺豚耱彘 耧疱耨梃 沐眍 铞屙桠嚯 镱祛 滂耧屦耔铐眍泐 囗嚯桤. 碾 恹耥屙 戾踵龛珈钼 忡噼祛溴轳蜮 沐眍 皴蜩 沐眍 gap, 蜞赕 潆 钺耥屙 戾踵龛珈钼 皲忤泐 钺豚耱彘 耧疱耨梃 桴 沐眍 镳钼钿桦 爨蝈爨蜩麇耜铄 祛溴腓痤忄龛; 镳邃耜噻囗睇 疱泱蝾痦 忡噼祛溴轳蜮 汨镱蝈琨 耦镱耱噔腓 腓蝈疣蝮痦 溧眄 溧眄 耧屦桁屙蜞. 碾 耦玟囗 镳铖蝠囗耱忮眄-怵屐屙眍泐 囹豚襦 耧疱耨梃 沐眍 桉镱朦珙忄腓 疱鲨铐眢 延聊 IBM DB2, 潆 疱嚯桤圉梃 恺-桧蝈痿彘襦 镱朦珙忄蝈 镳桁屙腓 蜞觇 蝈蹴铍钽梃, 赅 HTML, JSP, Java-皴疴脲螓, Java-囡镫弪 Java-耜痂矧.

1.4 袜篦磬 眍忤珥 疣犷螓 磬耱 疣犷蝈 怙屦恹 疣琊噌铗囗 觐礅彘屦睇 戾蝾 镱塍麇龛 觐腓麇耱忮眄 溧眄 镱 耧疱耨梃 沐眍 皴沆屙蜞鲨 桤 觐眙铌嚯 桤钺疣驽龛 赅痱桧 耧疱耨梃 桴 沐眍. 体蝾 怅膻鬣弪 5 镳铞邃箴, 桁屙眍, 皴沆屙蜞鲨 桤钺疣驽龛, 箐嚯屙桢 纛眍忸泐 耔沩嚯, 铒疱溴脲龛 忸琊囫蜞 狃桀磬, 镳铖蝠囗耱忮眄簋 疱汨耱疣鲨 赅痱桧 耧疱耨梃 桧蝈沭圉棹 溧眄, 觐蝾瘥 祛骓 镳桁屙螯 镱耠邃钼囹咫 镱 铗溴朦眍耱;

镱塍麇 镱腠, 蝾黜 耔耱屐囹梓羼觇 磬犷 觐腓麇耱忮眄 溧眄 钺 耧疱耨梃 沐眍 皴沆屙蜞鲨 赅驿铎 漯 赅驿钽 桤 1580 桧滂忤潴嚯 狃桀眍, 蜞赕 桧蝈沭桊钼囗睇 溧眄 钺 耧疱耨梃 赅驿钽 桤 14 沐眍 皴沆屙蜞鲨;

镱赅玎眍, 黩 钺豚耱 耧疱耨梃 沐眍 皴沆屙蜞鲨, 腩赅腓珙忄眄 狍潴 玎痤潲忸 镱腩耜, 镱 戾疱 疋铄泐 纛痨桊钼囗 鲨觌 14 耢妁帼蝰 镥疱漤屐 镱膻耋 狃桀磬;

镳邃腩驽磬 爨蝈爨蜩麇耜 祛溴朦 潆 镳邃耜噻囗 戾踵龛珈钼 疱泱鲨 耧疱耨梃 沐眍 gap 鲨觌 14, 镳噔桦 忸耧痤桤忸 怵屐屙眢 滂磬扈牦 耧疱耨梃 桴 沐眍, 耱屣屙 镥疱牮囗 耦皴漤桴 钺豚耱彘 耧疱耨梃, 蜞赕 忸耧痤桤忸 皲忤汨 沭囗桷 钺豚耱彘 耧疱耨梃 沐眍 gap 躅溴 鲨觌 14;

桉耠邃钼囗 戾踵龛珈 皲忤泐 鲥眚疣朦眍 钺豚耱 耧疱耨梃 Kr, 蜞赕 玎漤桴 钺豚耱彘 耧疱耨梃 kni gt 镱 磬镳噔脲龛 镥疱漤屐 觐眦 狃桀磬;

耦玟囗 镳铖蝠囗耱忮眄-怵屐屙眍 囹豚 耧疱耨梃 沐眍 皴沆屙蜞鲨 忤溴 疱鲨铐眍 徉琨 溧眄 FlyEx, 漕耱箫眍 镱 皴蜩 软蝈痦弪.

1.5 务眍忭 镱腩驽龛, 恹眍耔禧 磬 玎蝮 1. 暑祜蝈痦 钺疣犷蜿 鲨麴钼 桤钺疣驽龛 赅痱桧 耧疱耨梃 沐眍, 镱塍麇眄 镱祛 觐眙铌嚯钽 扈牮铖觐镟 桁祗眍綦箢疱聆屙蝽钽 爨痍桊钼囗 徼铍钽梓羼觇 爨牮铎铍尻箅, 祛驽 猁螯 桉镱朦珙忄磬 潆 镱塍麇龛 觐腓麇耱忮眄 溧眄 镱 耧疱耨梃 沐眍 in situ.

2. 暑腓麇耱忮眄铄 铒桉囗桢 镳铖蝠囗耱忮眄-怵屐屙眍 滂磬扈觇 觐祜铐屙 祛痿钽屙弪梓羼觐泐 镱 皴沆屙蜞鲨 礤钺躅滂祛 潆 镱龛爨龛 戾踵龛珈钼 邈 趔黻鲨铐桊钼囗.

3. 唁忤汨 钺豚耱彘 耧疱耨梃 沐眍 皴沆屙蜞鲨 忄骓 潆 镱玷鲨铐桊钼囗 钺豚耱彘 耧疱耨梃 沐眍-扈礤 桡疣 忄骓簋 痤朦 纛痨桊钼囗梃 皴沆屙蝽钽 镳屣囹蝈痦.

4. 锑蝈痂眈觇 沐睇 bcd,cad 沐睇 gap Kr, kni, gt, hb, tll 礤 蝾朦觐 礤钺躅滂禧, 眍 漕耱囹铟睇 潆 镳噔桦铋 耧疱耨梃 沐眍 gap 狍潴 玎痤潲忸 镱腩耜.

5. 项玷鲨铐磬 桧纛痨圉 犭囫蝾溴痨 漯铉铘桦 玎溧弪 滂磬扈麇耜 戾彘 忸 怵屐屙 觐灬桧圉桢 觐眦屙蝠圉栝 镳钿箨蝾 爨蝈痂眈觇 玷泐蜩麇耜桴 沐眍. 赅驿 祛戾眚 怵屐屙  觐灬桧圉 铒疱溴弪 礤 蝾朦觐 爨蝈痂眈觇扈 祛痿钽屙囔, 眍 皲忤汔扈 沭囗桷 钺豚耱彘 耧疱耨梃 沐眍 皴沆屙蜞鲨 桤-玎 疱泱蝾痦 忡噼祛溴轳蜮栝. 蒡 蝾腙钼囗桢 镱漯噻箪邂噱 嚓蜩忭, 礤 镟耨桠睇 耧铖钺 桧蝈痫疱蜞鲨 沭噤桢眚 祛痿钽屙 疣珈噱 沭囗桷 戾驿 纛痨桊钼囗桢 桧蝈痫疱蜞鲨彘 镱玷鲨铐眍 桧纛痨圉梃.

1.6 橡嚓蜩麇耜铄 珥圜屙桢 疣犷螓 袜篦眍-镳嚓蜩麇耜 珥圜桁铖螯 疣犷螓 耦耱铊 蝾, 黩 礤 怙屦恹 疣琊噌铗囗 篑镥 镳桁屙屙 戾蝾 觐礅彘屦眍 钺疣犷蜿 赅痱桧 耧疱耨梃 沐眍 皴沆屙蜞鲨 鲥朦 镱塍麇龛 觐腓麇耱忮眄 溧眄 镱 耧疱耨梃 沐眍. 体蝾 猁 篑镥 噤囡蜩痤忄 潆 钺疣犷蜿 溧眄 镱 耧疱耨梃 沐眍 皴沆屙蜞鲨 磬 箴钼礤 煨褪 (Janssens et al., 2006), 潆 钺疣犷蜿 铒蜩麇耜桴 耩彗钼 桤钺疣驽龛 赅痱桧 耧疱耨梃 沐眍 漯圊 狃桀磬 漯铉铘桦, 蜞赕 潆 爨耜桊钼囗 耧疱耨桊簋 钺豚耱彘 镱塍麇龛 觐腓麇耱忮眄 溧眄 镱 耧疱耨梃 沐眍 疣眄屐 疣玮栩梃 觐疣腚 Acropora millepora 祛瘃觐泐 囗屐铐 Nematostella vectensis (Kozlov et al., 2007). 埋  镱玮铍弪 聍栩囹 疣琊噌铗囗睇 戾蝾 忄骓 桧耱痼戾眚铎 桤怆鬻屙 觐腓麇耱忮眄铋 桧纛痨圉梃 桤 桤钺疣驽龛 赅痱桧 耧疱耨梃 沐眍. 悟戾蜩 蜞赕, 黩 耔塍 箜桠屦襦朦眍耱 犷朦眈蜮 镳铞邃箴 疣琊噌铗囗睇 戾蝾 祛驽 礤犷朦扈 祛滂翳赅鲨扈 镳桁屙螯 潆 钺疣犷蜿 痤觐泐 耧尻蝠 徼铍钽梓羼觇 桤钺疣驽龛 , 蜞觇 钺疣珙, 镳邃耱噔弪 桧蝈疱 潆 痤觐泐 牮筱 篦屙 钺豚耱 祛脲牦痦铋 徼铍钽梃.

赅麇耱忮 钿眍 桤 镳铞邃箴 觐礅彘屦眍泐 戾蝾溧 镳邃腩驽 眍恹 戾蝾 铒疱溴脲龛 忸琊囫蜞 狃桀磬 鲨觌 疣玮栩 14, 铖眍忄眄 磬 囗嚯桤 滂磬扈黜 赅痱桧 耧疱耨梃 沐磬 eve, 铌疣眄钽 怦艴 狃桀眍, 耱囗溧痱桤圉梃 桴 赅痱桧 耧疱耨梃 铗眍耔蝈朦眍 忸琊囫蜞 狃桀磬, 铒疱溴脲眄钽 耧屦桁屙蝈. 蒡铗 戾蝾 镱玮铍弪 噔蝾爨蜩玷痤忄螯 镳铞邃箴 镳邃耜噻囗 忸琊囫蜞, 溴豚弪 礤眢骓 蝠箐铄礻铄 铒疱溴脲龛 忸琊囫蜞 耧屦桁屙蜞朦睇 矬蝈 , 蜞觇 钺疣珙, 怆弪 忄骓 篑钼屦眈蜮钼囗桢 戾蝾溧 铒疱溴脲龛 忸琊囫蜞 狃桀磬 漯铉铘桦 疣眄屐 狃桀沐礤珏.

暑腓麇耱忮眄 溧眄 镱 耧疱耨梃 沐眍 皴沆屙蜞鲨, 镱塍麇眄 溧眄铋 疣犷蝈, 箜桕嚯 镱 铛忄蝮 怵屐屙眍 滂磬扈觇, 蝾黜, 桁妣 觌弪铟眍 疣琊屮屙桢 镱塍麇睇 疱珞朦蜞蝈 耔耱屐囹梓羼觇 爨聒蜞犴 耧屦桁屙蝾, 镳钼钿桁 钿眍 豚犷疣蝾痂 桉镱朦珙忄龛屐 钿龛 蝈 驽 耱囗溧痱桤钼囗睇 戾蝾漕. 蒡 铖钺屙眍耱 皲咫嚯 镱塍麇眄 溧眄 桉觌栩咫 忸耱疱犷忄眄 扈痤恹 耦钺耱忸, 桉镱朦珞桁 桴 赅 蝈铕弪梓羼觇 桉耠邃钼囗, 蜞 潆 桤篦屙 戾踵龛珈钼 皴沆屙蜞鲨, 耢. 磬镳桁屦, Holloway et al., 2003; Pereanu and Hartenstein, 2004; Diambra and da Costa, 2005;

Aegerter-Wilmsen et al., 2005; Isalan et al., 2005; Ludwig et al., 2005; Holloway et al., 2006; Krishna et al., 2005; Ochoa-Espinosa et al., 2005; Perkins et al., 2006; Yucel and Small, 2006; Zinzen and Papatsenko, 2007; Bergmann et al., 2007.

杨玟囗睇 镳铖蝠囗耱忮眄-怵屐屙眍 囹豚 耧疱耨梃 沐眍 皴沆屙蜞鲨 FlyEx 怆弪 铗牮 疱耋瘃铎, 痤觐 桉镱朦珞屐 扈痤恹 耦钺耱忸 徼铍钽钼 徼铊眙铕爨蜩觐. 亦 磬镳桁屦, 2006 泐潴 钺 觐腓麇耱忸 钺疣龛 FlyEx 耦耱噔桦 犷脲 260000.

疣犷蝈 桉耠邃箦蝰 鲥眚疣朦睇 忸镳铖 狃桀腩汨 - 戾踵龛珈 溴蝈痨桧圉梃 觌弪铌 鬣耱彘 玎痤潲 祛痿钽屙弪梓羼觇 镱. 项扈祛 钽, 恹镱腠屙睇 桉耠邃钼囗 桁妣 忄骓铄 珥圜屙桢 潆 塍鼬邈 镱龛爨龛 戾滂鲨眈觇 囫镥牝钼 疣玮栩, 耦忸牦镯铖螯 疣琊噌铗囗睇 戾蝾漕 祛溴脲 纛痨桊箦, 觐礤黜铎 栩钽, 戾蝾漕腩汨麇耜簋 徉珞 潆 疱觐眈蝠箨鲨 沐眄铋 皴蜩 铕汔龛珈 镳 铗耋蝰蜮梃 桦 钽疣龛麇眄铎 桉镱朦珙忄龛 祗蜞沐礤玎.

1.7 里痤徉鲨 疣犷螓 绣珞朦蜞螓 滂耨屦蜞鲨铐眍 疣犷螓 猁腓 漕腩驽睇 磬 觐眙屦屙鲨: Computational Cell Biology Workshop [CSHL 2007] (Cold Spring Harbor, USA, 2007); 燕黻-襄蝈疳箴泷觐 戾驿箜囵钿眍 觐眙屦屙鲨 镱 磬眍徼铗艴眍腩汨, 镫屙囵睇 漕觌噤, (燕黻-襄蝈疳箴, 2006);

3 Workshop on Data Integration for the Life Sciences [DILS2006] (Hinxton, UK); 1, 3, 4 International Conference on Bioinformatics of Genome Regulation and Structure [BGRS1998, 2002, 2004, 2006], 2006 . - 镳桡豚眄 漕觌噤 (皖忸耔徼瘃, 1998, 2002, 2004 2006); 3 4 TICSP Workshop on Computational Systems Biology, 镳桡豚眄 漕觌噤 (Tampere, Finland, 2005 2006); 2, 3 4 International Symposium on Networks in Bioinformatics, 2005 2006 .. - 镳桡豚眄 漕觌噤, [ISNB 2005, 2006, 2007] (Amsterdam, the Netherlands, 2005, 2006 2007); NETTAB 2005 workshop, Workflows management: new abilities for the biological information overflow (Naples, Italy, 2005); 2d Integrative Bioinformatics Workshop, 镳桡豚眄 漕觌噤, (Bielefeld, Germany, 2005); 42, 44, 45 46 Annual Drosophila Research Conferences, (Washington DC, San Diego, Washington DC Chicago, USA, 2001, 2003, 2004 2005); Intl. Moscow Conference on Computational Molecular Biology (Moscow, Russia, 2003); 7, 11 Int. Conference on Intelligent Systems for Molecular Biology [ISMB99, ISMB02, ISMB03], 1999 . - 镳屐 SGI 玎 塍鼬栝 漕觌噤 磬 觐眙屦屙鲨, (Heidelberg, Germany, 1999;

Edmonton, Canada, 2002; Brisbane, Australia, 2003); Computation in Cells: EPSRC Emerging Computing Paradigms Workshop, 镳桡豚眄 漕觌噤, (Hertfordshire, UK, 2000) ..

桉镱朦珙忄龛屐 爨蝈痂嚯钼 滂耨屦蜞鲨 噔蝾痤 皲咫囗 2 镳桡豚眄 漕觌噤 Lawrence Berkeley National Laboratory, USA (2002 2006 沣.), 漕觌噤 磬 填耜钼耜铎 皴扈磬疱 镱 徼铊眙铕爨蜩赍 (2006), 镳桡豚眄 漕觌噤 2003 . Genetics Department, Cambridge University (UK) Bioinformatics Research Centre, University of Glasgow (UK), 镳桡豚眄 漕觌噤 2005 . 磬 耰oinformati駍 裲lloquium, Georg-August-Universitat Gottingen (Germany) 磬 Waterman Seminars, the Leibniz Institute of Plant Genetics and Crop Plant Research (IPK, Germany), 镳桡豚眄 漕觌噤 Laurence H. Baker Center for Bioinformatics and Biological Statistics Seminar Series, Iowa University, USA (2001), 蜞赕 2 镳桡豚眄 脲牿梃 磬 the Les Houches Summer School "Multiple Aspects of DNA and RNA: from biophysics to bioinformatics"(2004). 亦赕 皲咫囗 3 镳桡豚眄 漕觌噤 磬 皴扈磬疣 Dagstuhl: 3d Dagstuhl Seminar for Information and Simulation Systems for the Analysis of Gene Regulation and Metabolic Pathways (2001); Dagstuhl Seminar 04281 "Integrative Bioinformatics - Aspects of the Virtual Cell (2004), Dagstuhl Seminar 03051, "Information and Process Integration: A Life Science Perspective"(2003) 祉钽桴 漯筱桴 皴扈磬疣. 唁咫囗 蜞赕 漕觌噤 磬 皴扈磬疱, 铕汔龛珙忄眄铎 翳痨铋 Leica, "杨怵屐屙睇 觐眙铌嚯 扈牮铖觐稃 翳痨 Leica 桴 镳桁屙屙桢 徼铍钽梃"(燕黻-襄蝈疳箴, 2006).

署铎 蝾泐 疱珞朦蜞螓 疣犷螓 钺耋驿嚯桉 磬 燕黻-襄蝈疳箴泷觐 皴扈磬疱 镱 觐祜蝈痦铋 徼铍钽梃, 磬 皴扈磬疣 赅翦漯 狃桀腩汨 沐礤蜩觇 燕黻-襄蝈疳箴泷觐泐 泐耋溧瘃蜮屙眍泐 箜桠屦耔蝈蜞 猁腓 怅膻麇睇 脲牿棹 磬 体驿箜囵钿眍 铍-皴扈磬疱 BGRS "葩铍, 耔耱屐磬 徼铍钽 耋镥痍铎稂屦睇 恹麒耠屙 徼铊眙铕爨蜩赍" 2005 . (皖忸耔徼瘃, 蓄耨).

1.8 象犭桕圉梃 项 爨蝈痂嚯囔 滂耨屦蜞鲨 铒筢腓觐忄眍 45 磬篦睇 疣犷 (怦 耦噔蝾瘃蜮), 蝾 麒耠 23 耱囹 疱翦痂痼屐 磬篦睇 骟痦嚯圊.

2 羊痼牝箴 滂耨屦蜞鲨铐眍 疣犷螓 蔫耨屦蜞鲨 耦耱铊 桤 忖邃屙, 7 汶噔, 恹忸漕 徼犭桀沭圄梃 (229 磬桁屙钼囗). 佩 镱腠 钺 耦耱噔弪 165 耱疣龛, 觐腓麇耱忸 痂耋黻钼 44.

2.1 杨溴疰囗桢 疣犷螓 骂 骡邃屙梃 钺耥 嚓蝮嚯铖螯 鲥腓 疣犷螓, 邋 磬篦磬 眍忤珥, 镳嚓蜩麇耜 珥圜桁铖螯. 阳铕祗腓痤忄睇 镱腩驽龛, 恹眍耔禧 磬 玎蝮. 橡桠邃屙 桧纛痨圉 钺 囡痤徉鲨 疣犷螓, 觐腓麇耱忮 矬犭桕圉栝 噔蝾疣, 铒桉囗 耱痼牝箴 滂耨屦蜞鲨.

襄疴 汶噔 揅钼疱戾眄 镳邃耱噔脲龛 溴蝈痨桧圉梃 皴沆屙蝾 漯铉铘桦麛 怆弪 钺珙痤 腓蝈疣蝮瘥.

漯铉铘桦 祛痿铍钽梓羼觇 疣珉梓桁 沭囗桷 皴沆屙蝾 纛痨桊簋蝰 泐疣玟 镱玟礤 蝾泐 怵屐屙, 觐沅 镳铊聃钿栩 溴蝈痨桧圉 皴沆屙蝾. 灶痨桊钼囗棹 沭囗桷 皴沆屙蝾 镳邃耱怏弪 牮囹觐怵屐屙眍 镱怆屙桢 沭囗桷 镟疣皴沆屙蝾, 觐蝾瘥 忸珥桕帼 忸 怵屐 箐腓礤龛 玎痤潲忸 镱腩耜 镱皴疱滂礤 赅驿钽 狍潴泐 皴沆屙蜞 (Campos-Ortega and Hartenstein, 1985). 腻蝈痨桧圉 皴沆屙蝾 镳桠钿栩 铒疱溴脲龛 镱腩驽龛 沭囗桷 镟疣皴沆屙蝾 (Martinez-Arias and Lawrence, 1985) 镳铊聃钿栩 祛痿钽屙弪梓羼觐 镱脲, 镳邃耱噔屐 耦犷 磬 耱噤梃 耔眦栩栲朦眍 犭囫蝾溴痨 鲨觌 溴脲龛 溴 14 蝮 鬣耱 狃桀磬, 桤 觐蝾痤 疣珙恻弪 玎痤潲忄 镱腩耜 (Gilbert, 2003).

朽眄栝 囡 疣玮栩 漯铉铘桦 怅膻鬣弪 耱噤棹 漯钺脲龛 耔眦栩栲朦眢 犭囫蝾溴痨. 羊噤 耔眦栩栲朦眍 犭囫蝾溴痨 镳钿铍驵弪 鲨觌 溴脲龛 溴 10 镱 鲨觌 溴脲龛 13.  怵屐 漯, 扈沭桊钼噔 磬 镥痂翦痂, 钺疣珞 箫铕漕麇眄 耠铋 屦汨 (屦汨溧 - 漯, 囫耦鲨桊钼囗眍 篦囫蜿铎 鲨蝾镫噻禧), 镳桦邈帼 镱忮瘐眍耱 狃桀磬. 骤觌 溴脲龛 溴 14 - 镥疴 鲨觌 镱耠邃簋 耱噤梃, 磬琨忄屐铋 觌弪铟眍 犭囫蝾溴痨铋. 奏耱 鲨觌 14, 镳邃耱怏 汔耱痼鲨, 磬琨忄弪 鲨觌铎 溴脲龛 14. 骂 怵屐 鲨觌 14 漯 箐腓 怃铍 耦狁蜮屙眍 徉玎朦眍-囡桕嚯铋 铖 , 怦觐疱 镱耠 钽, 磬麒磬弪 桧忄汨磬鲨 戾灬疣, 觐蝾疣 钺铖钺弪 漯 铗溴朦睇 觌弪觇. 锣骓, 黩 怙腩螯 漕 镱腠钽 钺疣珙忄龛 觌弪铟眍 戾灬疣睇 赅驿 纛痨桊簋 觌弪赅 耦钺弪 龛驽 疣耧铍铈屙睇 驽腧觐 镱耩邃耱忸 鲨蝾镫噻爨蜩麇耜桴 赅磬腩. 蒡 溴豚弪 忸珈铈睇 钺戾 徨腙囔 戾驿 耦皴漤桁 屦汨溧扈 蝈麇龛 怦彘 耱噤梃 耔眦栩栲朦眍 犭囫蝾溴痨 鲨觌 溴脲龛 14.

秒噔睇 翳玷麇耜桢 觐祜铐屙 祛痿钽屙弪梓羼觐泐 镱 皴沆屙蜞鲨 耋螯 沐睇 皴沆屙蜞鲨 - 桎屙蜩翳鲨痤忄睇, 觌铐桊钼囗 (Akam, 1987; Ingham, 1988) 镱漯噻溴脲睇 磬 麇螓疱 沭箫稃. 锑蝈痂眈觇 觐铕滂磬蝽 沐睇 (Nsslein-Volhard et al., 1987) 耧疱耨桊簋蝰 爨蝈痂眈觇 沐眍祛 怆 磬 疣玮栩桢 镥疱漤邈, 玎漤邈 桦 蝈痨桧嚯 疣轭眍 狃桀磬. 缅睇 漯筱桴 沭箫 gap, pair-rule segment polarity 怆 玷泐蜩麇耜桁, ..

耧疱耨桊簋蝰 沐眍祛 狃桀磬 (Nsslein-Volhard and Wieschaus, 1980).

令朦眈蜮 爨蝈痂眈觇 觐铕滂磬蝽 玷泐蜩麇耜桴 沐眍 皴沆屙蜞鲨 觐滂痼 蝠囗耜痂秭桀眄 羿牝铕, 觐蝾瘥 疱泱腓痼 耧疱耨棹 玷泐蜩麇耜桴 沐眍 皴沆屙蜞鲨. 亦觇 钺疣珙, 沐睇 皴沆屙蜞鲨 钺疣珞 皴螯 忡噼祛溴轳蜮簋 漯筱 漯筱铎 沐眍怵邈箅蝾痤. 理嚯桤 翦眍蜩镱 溻铋睇 祗蜞眚钼 恹忤 桢疣瘐棹 疱泱蝾痦 忡噼祛溴轳蜮栝 铋 沐眄铋 皴蜩 (耢. 钺珙瘥 Akam, 1987; Ingham, 1988), 耦铗忮蝰蜮梃 觐蝾痤 沐睇 犷脲 恹耦觇 箴钼礤 (磬镳桁屦, 爨蝈痂眈觇 觐铕滂磬蝽 沐睇) 疱泱腓痼 沐睇 犷脲 龛珀桴 箴钼礤 (磬镳桁屦, 沐睇 gap), 礤 磬钺铕铗. 项扈祛 钽 恹怆屙 耋耱忮眄 疱泱蝾痦 忡噼祛溴轳蜮 戾驿 沐磬扈, 镳桧噤脲驵扈 钿眍祗 箴钼睨 桢疣瘐梃.

骂 怛铕铋 汶噔 "锑蝈痂嚯 戾蝾潲" 镳桠邃屙 桧纛痨圉 钺 钺牝 戾蝾溧 桉耠邃钼囗.

蝠弪 汶噔 "体蝾 镱塍麇龛 觐腓麇耱忮眄 溧眄 镱 耧疱耨梃 沐眍 皴沆屙蜞鲨" 疣耨祛蝠屙 铕桡桧嚯 戾蝾, 蜞赕 脲驵 铖眍忮 礤泐 嚯泐痂蜢, 疣琊噌铗囗睇 潆 觐腓麇耱忮眄铋 铞屙觇 箴钼 耧疱耨梃 沐眍 皴沆屙蜞鲨 磬 铖眍忮 桤钺疣驽龛, 镱塍麇眄 镱祛 觐眙铌嚯钽 扈牮铖觐镟. 橡邃腩驽眄 戾蝾 钺疣犷蜿 怆弪 觐礅彘屦睇 怅膻鬣弪 5 徉珙恹 镳铞邃箴: 皴沆屙蜞鲨 桤钺疣驽龛, 箐嚯屙桢 纛眍忸泐 耔沩嚯, 铒疱溴脲龛 忸琊囫蜞 狃桀磬, 镳铖蝠囗耱忮眄簋 疱汨耱疣鲨 赅痱桧 耧疱耨梃 桧蝈沭圉棹 溧眄. 物桉囗 蝾黜铖螯 赅驿铋 桤 疣琊噌铗囗睇 镳铞邃箴.

绣珞朦蜞螓 镳邃耱噔脲睇 耱囹 A3, A5, A6, A8, A9, A15, A16, A17, A19, A20, A21, A22, A25, A29, A31, A36, A38, A40, A42, A43, 45.

彦沆屙蜞鲨 桤钺疣驽龛 怅膻鬣弪 麇螓疱 囡: (1) 镳桠邃屙桢 耧屦桁屙蜞朦眍泐 桤钺疣驽龛 耱囗溧痱眍 铕桢眚圉梃, 蜞 黩钺 镥疱漤栝 觐礤 狃桀磬 磬躅滂腭 耠邂, 耧桧磬 鬣耱 疋屦躞; (2) 恹溴脲龛 钺豚耱, 玎龛爨屐铋 钺牝铎 磬 桤钺疣驽龛, 铗皴麇龛 矬耱铋 钺豚耱 镱 牮; (3) 镱耱痤屙桢 "溴痦铋 爨耜"狃桀磬, 蝾 羼螯 徼磬痦钽 桤钺疣驽龛, 觐蝾痤 蝾朦觐 镨犟咫 沭囗桷圊 溴 桁妣 礤眢脲怏 桧蝈眈桠眍耱; (4) 恹麒耠屙桢 觐铕滂磬 鲥眚痤桎钼 溴 耩邃礤 桧蝈眈桠眍耱 綦箢疱聆屙鲨 赅驿铎 漯 潆 赅驿钽 赅磬豚 扈牮铖觐镟, .. 潆 赅驿钽 镳钿箨蜞 耜囗桊钼囗眍泐 沐磬. 佯邃睨 桧蝈眈桠眍耱 綦箢疱聆屙鲨 聍栩噱 镳铒铕鲨铐嚯铋 箴钼睨 耧疱耨梃 沐磬 (Kosman et al., 1997).

蒡囡 镳桠邃屙 桤钺疣驽龛 耱囗溧痱眢 铕桢眚圉棹 磬麒磬弪 镱耱痤屙 钿眍泐 桤钺疣驽龛 磬 铖眍忮 篑疱漤屙 溻篚 铒蜩麇耜桴 耩彗钼 潆 赅驿钽 桤 蝠艴 沐眄 镳钿箨蝾, 疱汨耱痂痼屐 钿眍泐 狃桀磬. 碾 桤钺疣驽龛, 镳铌疣眄 磬 汨耱铐钼 徨腙, 桉镱朦珞屐 赅麇耱忮 溴痦 爨痍屦钼, 磬 铖眍忮 桴 溻篚 耩彗钼 耱痤桁 镨犟咫 爨犟桁箪. 泥脲 潆 赅驿钽 狃桀磬 耦玟噱 沭筢簋 爨耜 矬蝈 觐眈蝠箬痤忄龛 镱镨犟咫钽 爨犟桁嚯钽 桤钺疣驽龛 桤 麇螓疱 桤钺疣驽龛, 镱塍麇眄 疣珥 赅磬豚 扈牮铖觐镟, 镳桁屙屙 铎 桤钺疣驽龛 镱痤泐忸泐 戾滂囗睇 翳朦蝠钼, 蜞赕 礤耜铍桴 鲨觌钼 铉梃 滂豚蜞鲨 (Gonzalez and Woods, 2002). 锑耜 狃桀磬 桉镱朦珞弪 潆 恹麒耠屙 筱豚, 磬 觐蝾瘥 磬漕 镱忮痦篁 狃桀, 潆 箐嚯屙 耠篦嚅睇 镨犟咫彘 磬 沭囗桷圊.

袜 怛铕铎 囡 耱痤栩 汶噤赅 爨耜 狃桀磬, 觐蝾疣 蝾黜铖蜩 镱怛铕弪 邈 纛痨. 碾 钽 耦玟噱 眍忸 桤钺疣驽龛 镨犟咫钽 爨犟桁箪 磬 铖眍忮 镳邃忄痂蝈朦眍 镱忮痦篁 钺疱玎眄 桤钺疣驽龛 镱塍麇眄铎 桤钺疣驽龛 镳桁屙屐 镱耠邃钼囹咫 恹疣忭桠囗桢 汨耱钽疣祆, 戾滂囗睇 镱痤泐恹 翳朦蝠. 镱塍麇眄铎 徼磬痦铎 桤钺疣驽龛 镳桁屙屐 蝠囗耵铕爨鲨 觌桎钼 疣耨蝾龛, 戾滂囗睇 镱痤泐恹 翳朦蝠, 黩 镳桠钿栩 耦玟囗棹 爨耜 铟屦蜞龛扈, 犷脲 耦铗忮蝰蜮簋扈 磬蝮疣朦睇 觐眚箴囔 狃桀磬. 镱祛 嚯泐痂蜢 劐磬-枢耱囗 (Shen and Castan, 1986), 玎镱腠屙 铉梃 镱塍鬣屐 眍怏 汶噤牦 爨耜 怦邈 狃桀磬 桉镱朦珞屐 邋 潆 钺疱玎龛 桤钺疣驽龛, 镱塍麇眄 忸 怦艴 赅磬豚 扈牮铖觐镟.

袜 蝠弪 麇蜮屦蝾 囡圊 耱痤栩 蜞 磬琨忄屐 溴痦 爨耜 磬 铖眍忮 桤钺疣驽龛, 镳铌疣眄钽 磬 汨耱铐钼 徨腙, 羼腓 铐 桁邋蝰, 桦 驽 磬 铖眍忮 镨犟咫钽 爨犟桁箪 耧疱耨梃 蝠艴 铖蜞朦睇 沐眍. 秧圜嚯 潆 篑桦屙 觐眚疣耱 篁铟礤龛 沭囗桷 溴 镳桁屙屐 腩赅朦眍 恹疣忭桠囗桢 汨耱钽疣祆, 箐嚯屐 沭囗箅桊钼囗睇  嚯泐痂蜢铎 署桁扈眈 镳钼钿桁 礤耜铍 鲨觌钼 戾滂囗眍 翳朦蝠圉梃. 泥脲, 镱耠 桧忮痱桊钼囗 珥圜屙栝 怦艴 镨犟咫彘, 耦玟噱蝰 桤钺疣驽龛 忸漕疣玟咫. 吾豚耱 忸漕疣玟咫 铒疱溴弪 赅 蝈痧栩铕, 玎蜞 钿龛 漯铎 钽疣龛麇眄 腓龛彘 痂眍 钿桧 镨犟咫. 枢驿 钺豚耱 忸漕疣玟咫 踵疣牝屦桤箦蝰 箜桕嚯 珥圜屙桢 沭噤圉梃 皴痤泐 鲡弪. 疱珞朦蜞蝈 镳桁屙屙 铉梃 镱痤泐忸泐 翳朦蝠  桤钺疣驽龛 镳孱狃噻箦蝰 徼磬痦铄, 沅 沭囗桷 忸漕疣玟咫 桁妣 珥圜屙桢 0. 项耠 箪眍驽龛 钽 徼磬痦钽 桤钺疣驽龛 磬 桤钺疣驽龛 镨犟咫钽 爨犟桁箪 桦 桤钺疣驽龛 汨耱铐钼 徨腙钼, 赅驿铄 漯 铗溴弪 铗 耦皴漤桴 溴 沭囗桷彘 桤 眢脲恹 镨犟咫彘.

蒡 铒屦圉 镱玮铍弪 疣玟咫栩 礤觐蝾瘥 漯, 耠桠羼 磬 桤钺疣驽龛. 玎忮瘌屙桢 铉 镱耠邃簋 镳孱狃噻钼囗桢 疣耨蝾龛 (Vincent et al., 1997) 镱痤泐恹 翳朦蝠铎 镱玮铍 箐嚯栩 桤 爨耜 镱耱铕铐龛 蝽, 礤 怆桢 漯囔 狃桀磬.

项塍麇眄 徼磬痦 爨耜 桉镱朦珞弪 潆 桤怆鬻屙 觐腓麇耱忮眄 溧眄 钺 耧疱耨梃 沐眍. 暑铕滂磬螓 鲥眚痤桎 赅驿钽 漯 恹麒耠 镱祛 桧忄痂囗蝾 祛戾眚钼 (Hu, 1962). 袜腩驽龛 爨耜 磬 桤钺疣驽龛, 镱塍麇眄 赅驿铎 赅磬脲 扈牮铖觐镟, 镱玮铍弪 恹麒耠栩 耩邃龛 觐眦屙蝠圉梃 镳钿箨蝾 怦艴 耜囗桊钼囗睇 沐眍 赅驿铎 漯.

暑礤黜 疱珞朦蜞 镳邃耱噔弪 耦犷 蜞犭桷, 耦溴疰帙簋 (x,y) 觐铕滂磬螓 赅驿钽 漯 镳铞屙蜞 铗 潆桧 痂睇 狃桀磬, 蜞赕 篑疱漤屙眢 桧蝈眈桠眍耱 綦箢疱聆屙鲨, 桦 铗眍耔蝈朦睇 箴钼屙 耧疱耨梃 潆 赅驿钽 桤 耜囗桊钼囗睇 溧眄钽 狃桀磬 沐眍. 朽琊噌铗囗磬 镳铞邃箴 桤怆鬻屙 觐腓麇耱忮眄铋 桧纛痨圉梃 桤 徼铍钽梓羼觇 桤钺疣驽龛 疱嚯桤钼囗 铕桡桧嚯铎 镟赍蝈 ProStack (Processing of Stacks) 潆 忤珞嚯钽 镱耱痤屙 耠铈睇 镳铞邃箴 钺疣犷蜿 溧眄 桤钺疣驽龛.

枢麇耱忸 镱耱痤屙 溴痦铋 爨耜 钺眍 觐眚痤腓痼弪 忤珞嚯 矬蝈 邋 磬腩驽龛 磬 桤钺疣驽龛, 镱塍麇眄铄 镳 耜囗桊钼囗梃 汨耱铐钼, 桦 磬 镨犟咫 爨犟桁箪 桤钺疣驽龛, 镱塍麇眄 漯筱桴 赅磬豚 扈牮铖觐镟. 碾 噔蝾爨蜩玎鲨 磬扈 镳邃腩驽 麒耠屙睇 戾蝾 觐眚痤 赅麇耱忄 溴痦铋 爨耜, 铖眍忄眄 磬 蝾, 黩 疣玑痤 桧蝈眈桠眍耱彘 潆 镨犟咫彘 忭篁痂 漯 漕腈屙 猁螯 戾睃, 麇 蜞觐忸 戾驿 镨犟咫扈 忭篁痂 漯 忭 邈.

愉嚯屙桢 纛眍忸泐 耔沩嚯. 疹痤 桤忮耱眍, 黩 磬潴 耦 耧弼梏梓羼觇 铌疣忄龛屐, 戾蝾潲 桁祗眍綦箢疱聆屙蝽钽 爨痍桊钼囗 徼铍钽梓羼觇 钺疣琏钼 in situ 镳桠钿 镱怆屙棹 礤耧弼梏梓羼觐泐 耔沩嚯, 磬琨忄屐钽 纛眍. 骂珥桕眍忮龛 钽 耔沩嚯 恹玮囗 祉钽桁 羿牝铕囔, 桤 觐蝾瘥 忄骓 痤朦 镳桧噤脲骅 镥疴梓睇 怛铕梓睇 囗蜩蝈豚. 湾耧弼梏梓羼觇 纛 祛驽 猁螯 筢疣 怵篦眢 潆 赅驿钽 觐黻疱蝽钽 耠嚅溧 矬蝈 篑蜞眍怅 珥圜屙 铘赳弪. 武磬觐, 铟邂桎眍, 黩 镳铖蝾 篑蜞眍怅铋 铘赳弪 礤忸珈铈眍 漕徼螯 箐嚯屙 纛磬 桤 怦艴 赅痱桧 耧疱耨梃 怦艴 沐眍 忖桎 蝾泐, 黩 潆 赅驿铋 赅痱桧 蝠遽簋蝰 疋铊 篑蜞眍怅 潆 箐嚯屙 纛磬.

西邂桎眍, 黩 溧驽 礤珥圜栩咫 箴钼屙 纛磬 桉赅玷 麒耠屙眍 珥圜屙桢 箴钼 耧疱耨梃 沐磬. 令脲 蝾泐, 箴钼屙 纛磬 忄瘘桊箦 铗 狃桀磬 狃桀眢 铗 耧屦桁屙蜞 耧屦桁屙蝮, 黩 礤 镱玮铍弪 镳钼羼蜩 耩噔礤龛 溧眄, 镱塍麇眄 疣珥 耧屦桁屙蜞.

朽琊噌铗囗睇 磬扈 戾蝾 箐嚯屙 纛眍忸泐 耔沩嚯 铖眍忄 磬 磬犭屙梃, 黩 箴钼屙 綦箢疱聆屙鲨 眢朦-祗蜞眚圊, 铌疣眄 磬 铗耋蝰蜮簋 徨腩, 躅痤 囡镳铌耔扈痼弪 溻箪屦睇 镟疣犷腩桎铎 (桦, 犷脲 钺 耠篦噱, 恹矬觌铋 镱忮瘐眍耱 怛铕钽 镱潢). 蒡铗 镟疣犷腩桎 铟屙 犭桤铌 耔祆弪痂黜铎 铗眍耔蝈朦眍 铖彘 x y.

务眍忭 桎 戾蝾溧 箐嚯屙 纛磬 耦耱铊 铒疱溴脲龛 礤镳羼耔痼桴 钺豚耱彘 狃桀磬, 觐蝾瘥 桉镱朦珞 潆 囡镳铌耔爨鲨 纛眍忸泐 耔沩嚯, 黩钺 玎蝈 箐嚯栩 邈 爨聒蜞徼痤忄龛屐 赅痱桧 耧疱耨梃.

湾耧疱耨桊簋 钺豚耱 - 篦囫蜿 狃桀磬, 觐蝾瘥 溧眄 沐 礤 耧疱耨桊箦蝰 犷朦眈蜮 溴 - 潆 赅驿钽 沐磬 镥疴铐圜嚯 铒疱溴 磬 铖眍忮 蝙囹咫钽 忤珞嚯钽 桤篦屙 赅痱桧 耧疱耨梃 忸 怦艴 狃桀磬. 青蝈 礤耧疱耨桊簋 钺豚耱 篁铟 磬 溻箪屦眍 赅痱桧 耧疱耨梃 赅驿钽 狃桀磬. 橡 铎, 耔塍 桉牮桠脲龛 镱腩 耧疱耨梃 怃铍 铖 y, 桴 礤钺躅滂祛 恹镳扈螯.

灶眍恹 耔沩嚯 囡镳铌耔扈痼弪 赈噤疣蜩黜 镟疣犷腩桎铎 镱 铒铕睇 蝾麝囔, 恹溴脲眄 桤 礤耧疱耨桊簋 钺豚耱彘 狃桀磬 恹镳祀屙睇扈 镱腩襦扈. 橡铞邃箴 镳桁屙弪 镱耠邃钼囹咫 觐 怦屐 桤钺疣驽龛 赅痱桧 耧疱耨梃, 镱塍麇眄 钿眍泐 狃桀磬. 令朦 礤耧疱耨桊簋 钺豚耱 耠邃箦 疣玑桠囹 磬 礤耜铍 钺豚耱彘 礤耋耱忮眄铋 忄痂圉桢 纛磬. 物铕睇扈 蝾麝囔 聍栩帼蝰 蝾麝, 觐蝾瘥 箴钼屙 桧蝈眈桠眍耱 綦箢疱聆屙鲨 磬 镳邂噱 玎溧眄钽 镱痤泐忸泐 珥圜屙, 镳桁屦眍 疣忭钽 2/3 疣玑痤襦 珥圜屙栝 耧疱耨梃 钺豚耱.

相疣犷腩桎 S(x, y) = 11x2 +22y2 +12xy +1x+2y +0, 囡镳铌耔扈痼栝 纛, 耱痤栩 镳 镱祛 栩屦囹桠眍 铒蜩扈玎鲨铐眍 镳铞邃箴. 侮铐鬣蝈朦眍 纛 箐嚯弪 桤 怦邈 狃桀磬 腓礤轫 铗钺疣驽龛屐 桧蝈眈桠眍耱, 觐蝾痤 镳孱狃噻箦 珥圜屙 綦箢疱聆屙鲨 疣忭 桦 戾睃 箴钼 纛磬 眍朦, 爨犟桁嚯 忸珈铈眢 綦箢疱聆屙鲨 (255) 襦祗 皴. 橡孱狃噻钼囗桢 桁邋 忤 ai - S(xi, yi) a(n) = max 255, 0, i = 1... N, (1) i 255 - S(xi, yi) 疱珞朦蜞蝈 镱塍鬣屐 眍痨桊钼囗眢 赅痱桧 耧疱耨梃, 觐蝾痤 赅驿铄 漯 踵疣牝(n) 痂珞弪 忮牝铕铎 vi = {xi, yi, a(n)}, i = 1,... N.

i 锣骓 镱澉屦觏篁, 黩 爨聒蜞徼痤忄龛 眍痨嚯桤钼囗睇 桧蝈眈桠眍耱彘 滂囡噻铐 [0,255] 礤钺躅滂祛 耔塍 玎忤耔祛耱 纛磬 铗 恹犷疣 镥疴梓睇 怛铕梓睇 囗蜩蝈 铗耋蝰蜮 蜞觐 玎忤耔祛耱 箴钼 耧疱耨梃 沐磬.

橡铞邃箴 箐嚯屙 纛磬 桤 痤觇 赅痱桧 耧疱耨梃 沐眍 hb cad, 蜞赕 沐眍 pair-rule 磬 疣眄桴 耱噤 溧弪 塍鼬栝 疱珞朦蜞 镳 漕矬龛 耔祆弪痂黜铖蜩 镱 铖 x 磬鬣朦眍泐 镳栳腓驽龛 镟疣犷腩桎. 缅 bcd 镳邃耱噔弪 耦犷 耧弼栲朦睇 耠篦嚅, 镱耜铍 潆 礤泐 玎漤 沭囗桷 耧疱耨梃 礤 铒疱溴脲磬. 疹痤 桤忮耱眍, 黩 觐眦屙蝠圉 徨腙 Bcd 疣耧疱溴脲磬 镱 耧铐屙鲨嚯铎 沭噤桢眚 爨犟桁箪铎 囗蝈痂铕眍 觐眦 漕耱桡噱 纛眍忸泐 箴钼 镱耱屦桀痦铋 蝠弪 狃桀磬 (Driever and NssleinVolhard, 1988b; Driever and Nsslein-Volhard, 1988a). 帖 眍痨桊箦 赅痱桧 耧疱耨梃 bcd, 漕徼忄顸 耧铐屙鲨嚯铖蜩 眍痨桊钼囗眍 赅痱桧.

项潋忮疰溴龛 钺铖眍忄眄铖蜩 镳邃腩驽眄钽 戾蝾溧 箐嚯屙 纛眍忸泐 耔沩嚯 祛骓 镱塍麒螯 矬蝈 忤珞嚯铋 桧耧尻鲨 赅痱桧 耧疱耨梃. 央噌 戾耱铎 忤珞嚯铋 铞屙觇 怆弪 礤忭铄 镳邃镱腩驽龛 蝾, 黩 龛 钿桧 桤 沐眍 皴沆屙蜞鲨 礤 桁邋 疣忭铎屦眍 龛珀钽 箴钼 耧疱耨梃 疣轭磬, 脲驵 忭 镳铖蝠囗耱忮眄 铕汔龛麇眄 钺豚耱彘 耧疱耨梃. 武磬觐, 镳 囗嚯桤 溧眄, 镱塍麇眄 镳 铌疣忄龛 钿眍泐 蝾泐 驽 徨腙 疣珥 镥疴梓睇扈 怛铕梓睇扈 囗蜩蝈豚扈, 耱囗钼栩 铟邂桎眍, 黩 沭箫稃 囗蜩覃忸痤蝾 疣珥钽 赅麇耱忄 溧 钿龛 蝈 驽 镳铖蝠囗耱忮眄 铕汔龛麇眄 钺豚耱 耧疱耨梃, 眍 疣珥 箴钼龛 纛磬. 令脲 蝾泐, 戾蝾 箐嚯屙 纛眍忸泐 耔沩嚯 猁 蝙囹咫 蝈耱桊钼囗 磬 祗蜞眚睇 狃桀磬, 泐祛玷泐蝽 镱 眢朦-祗蜞鲨 赅觐-腓犷 桤 沐眍 eve, gt, kni 桦 Kr 铌疣眄 潆 恹怆屙 镳钿箨蜞 祗蜞眚眍泐 沐磬. 枢痱桧 耧疱耨梃 蜞觇 祗蜞眚钼 镱耠 箐嚯屙 纛磬 蝠囗耵铕扈痼 眢脲忸 箴钼屙 耧疱耨梃 忸 怦屐 狃桀礤.

妈珞嚯 铞屙赅 赅痱桧 耧疱耨梃 镱赅琨忄弪, 黩 戾蝾 箐嚯屙 纛磬 溧弪 躅痤 疱珞朦蜞螓 潆 狃桀眍, 铗眍躐 鲨觌 14. 诣镨黜 疱珞朦蜞螓 箐嚯屙 纛磬 需. 1: 绣珞朦蜞螓 箐嚯屙 纛磬 桤 桧滂忤潴嚯 赅痱桧 耧疱耨梃 溧 沐眍. 青 桉觌屙桢 耠篦噱, 觐沅 鲨觌 溴脲龛 溴 箨噻囗, 怦 铖蜞朦睇 赅痱桧 耧疱耨梃 镱塍麇睇 狃桀磬, 镳桧噤脲驵 鲨觌 14. 湾眍痨嚯桤钼囗睇 赅痱桧 耧疱耨梃 桤钺疣驽睇 徨臌扈 牮箧赅扈; 赅痱桧 耧疱耨梃 镱耠 箐嚯屙 纛眍忸泐 耔沩嚯 镱赅玎睇 麇痦 牮箧赅扈; 镟疣犷腩桎, 囡镳铌耔扈痼栝 纛, 桤钺疣驽 耧腩铋 腓龛彘.

桤 赅痱桧 耧疱耨梃 bcd, hb, Kr, eve, run h 磬 铋 耱噤梃 镳桠邃屙 磬 痂. 1. 碾 赅痱桧 耧疱耨梃 沐眍 gap pair-rule 磬 疣眄桴 耱噤 蝾黜铖螯 戾蝾溧 珥圜栩咫 龛驽.

鲨觌圊 溴脲龛 1012 赅痱桧 耧疱耨梃 桴 沐眍 踵疣牝屦桤簋蝰 铟屙 龛珀桁 箴钼礤 桧蝈眈桠眍耱 綦箢疱聆屙鲨, 镳桉篁耱怏彘 镳嚓蜩麇耜 忸 怦艴 鬣耱 狃桀磬.

袜 痂. 1 镳桠邃屙 蜩镨黜 赅痱桧 耧疱耨梃 eve kni 鲨觌 12 疱珞朦蜞 箐嚯屙 纛磬. 软蝈痫疱蜞鲨 蜞觇 赅痱桧 礤钿眍珥圜磬: 忸珈铈眍, 黩 忮顸 耔沩嚯 钺篑腩怆屙 礤耧弼梏梓羼觐 綦箢疱聆屙鲨彘 玷泐蜩麇耜 耧疱耨 沐眍 铗耋蝰蜮箦.

物疱溴脲龛 忸琊囫蜞 狃桀磬. 项耜铍 铌疣忄龛 耜囗桊钼囗桢 狃桀眍 铖簌羼蜮腩顸 徨 篦弪 桴 忸琊囫蜞, 礤钺躅滂禧 耧弼栲朦睇 戾蝾潲 潆 铒疱溴脲龛 怵屐屙 疣玮栩. 耔塍 蝾泐, 黩 鲨觌 溴脲龛 10 镱 13 怆 礤镳钿铍骅蝈朦睇扈 镱 怵屐屙, 潆 铒疱溴脲龛 忸琊囫蜞 狃桀磬 磬 铎 镳铎彐篁赍 怵屐屙 漕耱囹铟屙 镱漶麇 觐腓麇耱忄 溴. 武磬觐, 鲨觌 溴脲龛 14 玎龛爨弪 铌铍 50 扈眢 , 镱铎 镳铖蝾 镱漶麇 觐腓麇耱忄 溴 礤漕耱囹铟屙. 橡邃腩驽眄 潆 疱龛 铋 玎溧麒 耧屦桁屙蜞朦睇 戾蝾 蝠遽箦 桉镱朦珙忄龛 钿眍泐 桤 赅磬腩 扈牮铖觐镟 潆 镱塍麇龛 桤钺疣驽龛 戾灬疣. 雾 铖眍忄 磬 桤戾疱龛 耱屣屙 桧忄汨磬鲨 戾灬疣睇 翳犟桊钼囗睇 狃桀眍 桉镱朦珙忄龛 耱囗溧痱眍 牮桠铋, 铒桉帼 怵屐屙眢 滂磬扈牦 桧忄汨磬鲨 戾灬疣 in vivo, 潆 铒疱溴脲龛 忸琊囫蜞 狃桀磬 (Merrill et al., 1988).

帖 疣琊噌铗嚯 眍恹 戾蝾 铒疱溴脲龛 忸琊囫蜞 狃桀磬 鲨觌 溴脲龛 溴 14, 铖眍忄眄 磬 囗嚯桤 恹耦觐 滂磬扈黜 赅痱桧 耧疱耨梃 沐磬 eve, 铌疣眄钽 怦艴 狃桀眍, 耱囗溧痱桤圉梃 桴 赅痱桧 耧疱耨梃 铗眍耔蝈朦眍 忸琊囫蜞 狃桀磬, 铒疱溴脲眄钽 耧屦桁屙蝈. 蒡铗 戾蝾 镱玮铍弪 噔蝾爨蜩玷痤忄螯 镳铞邃箴 镳邃耜噻囗 忸琊囫蜞, 桉觌噱 蝠箐铄礻铄 铒疱溴脲龛 忸琊囫蜞 耧屦桁屙蜞朦睇 矬蝈 , 蜞觇 钺疣珙, 珥圜栩咫 箅篦弪 戾蝾 铒疱溴脲龛 忸琊囫蜞 狃桀磬 漯铉铘桦 疣眄屐 狃桀沐礤珏.

赅麇耱忮 镥疴钽 汔 疣琊噌铗赍 戾蝾溧 怦 狃桀睇, 镳桧噤脲驵 鲨觌 溴脲龛 14, 猁腓 疣耧疱溴脲睇 矬蝈 忤珞嚯钽 囗嚯桤 赅痱桧 耧疱耨梃 eve 镱 忸顸扈 怵屐屙睇 觌囫襦. 项耜铍 狃桀睇 猁腓 耜囗桊钼囗 徨 篦弪 桴 忸琊囫蜞, 疣珞祉 镳邃镱腩骅螯 疣忭铎屦眍 疣耧疱溴脲龛 忸琊囫蜞 恹犷痍 狃桀眍, 镳桧噤脲驵 鲨觌 14. 亦 赅 怦 忸皴禳 怵屐屙睇 觌囫耦 耦溴疰囹 镳桁屦眍 钿桧嚓钼铄 觐腓麇耱忸 狃桀眍 鲨觌 溴脲龛 14 玎龛爨弪 铌铍 50 扈眢 疣玮栩, 祛骓 聍栩囹, 黩 赅驿 觌囫 镱 潆栩咫铖蜩 耦铗忮蝰蜮箦 镳桁屦眍 6.5 扈眢蜞. 孰囫耔翳赅鲨 狃桀眍, 铖眍忄眄 磬 囗嚯桤 滂磬扈觇 赅痱桧 耧疱耨梃 沐磬 eve, 躅痤 耦汶囫箦蝰 耦 耱屣屙 桧忄汨磬鲨 戾灬疣.

碾 镳邃耜噻囗 忸琊囫蜞 狃桀磬 镱 赅痱桧 耧疱耨梃 桉镱朦珙忄眍 钺钺龛 戾蝾溧 SVM (Support Vector machine) 磬 耠篦嚅 疱沭羼耔铐眍泐 铞屙桠囗 (Vapnik, 1995; Smola and Scholkopf, 1998). 体蝾 镳邃耜噻囗 忸琊囫蜞 怅膻鬣弪 蝠 囡: (1) 纛痨桊钼囗桢 钺篦帼 恹犷痍 狃桀眍, 赅驿 屐屙 觐蝾痤 踵疣牝屦桤箦蝰 磬犷痤 踵疣牝屦睇 镳桤磬觐, 铒桉帼 赅痱桧 耧疱耨梃, 耧屦桁屙蜞朦眍 铒疱溴脲眄 忸琊囫蝾;

(2) 镱耱痤屙桢 疱沭羼耔铐眍 趔黻鲨 镱 钺篦帼 恹犷痍; (3) 镳邃耜噻囗桢 忸琊囫蜞 潆 狃桀眍, 礤 怅膻麇眄 钺篦帼 恹犷痍 礤 桁妣 耧屦桁屙蜞朦睇 溧眄 耱屣屙 桧忄汨磬鲨 戾灬疣睇.

赅麇耱忮 钺篦帼 恹犷痍 猁豚 桉镱朦珙忄磬 沭箫镟 桤 103 狃桀眍, 镳桧噤脲驵 怵屐屙睇 觌囫襦 3 - 8. 枢驿 狃桀 钺篦帼 恹犷痍 猁 铛囵嚓蝈痂珙忄 祉钽铎屦睇 忮牝铕铎, 觐祜铐屙蜞扈 觐蝾痤泐 怆 忸琊囫 狃桀磬 13 镟疣戾蝠钼, 疣忭 耱疱祗爨 耧疱耨梃 沐磬 eve. 骂琊囫 猁 铒疱溴脲 矬蝈 桤戾疱龛 耱屣屙 桧忄磴桧圉梃 戾灬疣 磬 漕瘃嚯铋 耱铕铐 赅驿钽 桤 狃桀眍 桉镱朦珙忄龛 耱囗溧痱眍 牮桠铋 (Merrill et al., 1988).

锑臌 疣珈屦 钺篦帼 恹犷痍 (103 狃桀磬) 镱 耩噔礤龛 疣珈屦铎 忮牝铕 镳桤磬觐 (13) 祛驽 镱怆鬻 礤漕耱钼屦眍 镳邃耜噻囗桢 铗耋蝰蜮桢 篑蝾轺桠铖蜩 镳 桤戾礤龛 镟疣戾蝠钼 嚯泐痂蜢. 碾 箪屙屙 疣珈屦眍耱 忮牝铕 镳桤磬觐 猁 桉镱朦珙忄 羿牝铕睇 囗嚯桤. 项耠 镳桁屙屙 羿牝铕眍泐 囗嚯桤 恹犷痍 桤 501 狃桀磬, 铗眍躐 鲨觌 14, 猁豚 镱塍麇磬 耦忸牦镯铖螯 桤 13 眍恹 羿牝铕钼. 锣骓, 黩 潆 1狃桀眍 钺篦帼 恹犷痍, 镥疴 溻 羿牝铕 桁妣 珥圜栩咫簋 觐痧咫鲨 忸琊囫蝾, 92% 25% 耦铗忮蝰蜮屙眍. 蒡 镱玮铍桦 磬溴骓 镳邃耜噻囹 忸琊囫, 篦栩 忮牝铕 麇痱 蝾朦觐 镥疴 蝠 羿牝铕, 筲咫梓桦 耜铕铖螯 聃钿桁铖蜩.

疱珞朦蜞蝈 镱溽铕 镟疣戾蝠钼, 溧桴 磬桦篦 耦怙噤屙桢 疱沭羼耔铐眍 趔黻鲨 耧屦桁屙蜞朦睇扈 溧眄, 猁腩 篑蜞眍怆屙, 黩 祛溴腓 耦 耜嚯痦 漯铎 镱腓眍扈嚯 漯铎 怛铕钽 镱潢 塍鼬 怦邈 耦汶囫簋蝰 溧眄.

绣珞朦蜞螓 铞屙觇 蝾黜铖蜩 镳邃耜噻囗 忸琊囫蜞, 铖簌羼蜮脲眄铋 矬蝈 镱耠邃钼囹咫钽 桉觌屙 钿眍泐 狃桀磬 桤 钺篦帼 恹犷痍 桉镱朦珙忄龛 铖蜞怿桴 狃桀眍 赅麇耱忮 疣犷麇 恹犷痍 潆 镳邃耜噻囗 忸琊囫蜞 桉觌屙眍泐 狃桀磬, 镱赅玎腓, 黩 钺 祛溴腓 桁妣 镳桁屦眍 钿桧嚓钼簋 蝾黜铖螯 耠篦噱 桉镱朦珙忄龛 忮牝铕 麇痱 钿眍泐, 溻篚 桦 蝠艴 羿牝铕钼. 硒栳赅 镳邃耜噻囗 忸琊囫蜞 耦耱噔弪 铌铍 溻篚 扈眢 疣玮栩. 蒡 躅痤 疱珞朦蜞, 羼腓 镳桧螯 忸 忭桁囗桢, 黩 忸琊囫 狃桀眍 钺篦帼 恹犷痍 磬躅滂蝰 镳邃咫圊 铗 20 漕 50 扈眢 磬鬣豚 鲨觌 溴脲龛 14.

亦觇 钺疣珙, 潆 嚓牦疣蝽钽 镳邃耜噻囗 忸琊囫蜞 狃桀眍, 礤 怅膻麇眄 钺篦帼 恹犷痍, 漕耱囹铟眍 蝾朦觐 蝠艴 羿牝铕钼. 昨钺 镳钼羼蜩 疣珉梓桢 戾驿 祛溴扈, 镳桁屐 忸 忭桁囗桢, 黩 疣聍弪 玎溧麇 耦 耜嚯痦 漯铎 聃钿栩 泐疣玟 猁耱疱 (磬镳桁屦, 潆 蝠艴 羿牝铕钼 玎 59 栩屦圉梃 镳铗桠 197) , 耠邃钼囹咫, 耜嚯痦 祛溴朦 铌噻噱蝰 犷脲 镳邃镱黩栩咫铋.

填溴腓 耦 耜嚯痦 漯铎 镱腓眍扈嚯 漯铎 怛铕钽 镱潢 猁腓 桉镱朦珙忄睇 潆 镳邃耜噻囗 忸琊囫蜞 398 狃桀眍, 礤 镳桧噤脲驵 钺篦帼 恹犷痍, 镳梓屐 忮牝铕 镳桤磬觐 猁 耦耱噔脲 桤 蝠艴 羿牝铕钼. 橡邃耜噻囗睇 忸琊囫螓 躅痤 觐痧咫桊钼嚯 觌囫耔翳赅鲨彘 狃桀眍 镱 怵屐屙睇 觌囫襦.

绣汨耱疣鲨 赅痱桧 耧疱耨梃 沐眍 皴沆屙蜞鲨. 锐镱朦珙忄眄 潆 镱塍麇龛 溧眄 觐眙铌嚯 扈牮铖觐稃 镱玮铍 镱塍鬣螯 钿眍泐 狃桀磬 赅痱桧 耧疱耨梃 蝾朦觐 蝠艴 沐眍. 袜 驽 桧蝈疱耋弪 镳铖蝠囗耱忮眄-怵屐屙磬 滂磬扈赅 耧疱耨梃 怦艴 沐眍, 觐眚痤腓痼桴 纛痨桊钼囗桢 镳屣囹蝈痦 皴沆屙蜞鲨. 耔塍 桧滂忤潴嚯铋 忄痂噌咫铖蜩 疣珈屦钼 狃桀眍, 桧纛痨圉棹 钺 铗眍耔蝈朦眍 疣耧铍铈屙梃 赅痱桧 耧疱耨梃 疣珥 沐眍 镳铖蝠囗耱忮 礤忸珈铈眍 镱塍麒螯 镳铖螓 耦忪妁屙桢 赅痱桧 耧疱耨梃 桧滂忤潴嚯 狃桀眍, 铌疣眄 囗蜩蝈豚扈 疣珥 徨腙囔. 昨钺 疱螯  玎溧黧, 溧眄, 镱塍麇眄 潆 桧滂忤潴嚯 狃桀眍, 漕腈睇 猁螯 镳桠邃屙 钺 耔耱屐 觐铕滂磬 镱祛 镳铞邃箴 疱汨耱疣鲨.

溧眄铋 疣犷蝈 猁 疣琊噌铗囗 戾蝾 疱汨耱疣鲨, 铖眍忄眄 磬 恹溴脲龛 觐眚痤朦睇 蝾麇 桤钺疣驽龛 磬 镳孱狃噻钼囗梃 觐铕滂磬 潆 爨犟桁嚯 镱腠钽 耦忪妁屙 桴 蝾麇 疣珥 桤钺疣驽龛 (Brown, 1992). 赅麇耱忮 觐眚痤朦睇 蝾麇 钺眍 桉镱朦珞 赅觇-腓犷 踵疣牝屦睇 镳桤磬觇 桤钺疣驽龛. 项耜铍 磬 桉耠邃钼囗梃 怦 狃桀睇 猁腓 耜囗桊钼囗 潆 恹怆屙 耧疱耨梃 沐磬 eve 赅痱桧 耧疱耨梃 钽 沐磬 耔朦眍 戾弪 忸 怵屐屙, 赅麇耱忮 觐眚痤朦睇 蝾麇 桉镱朦珙忄腓 踵疣牝屦睇 镳桤磬觇 铋 赅痱桧 - 耱疱祗禧 钿眍戾痦铋 赅痱桧 耧疱耨梃, 镱塍麇眄铋 矬蝈 耱疣牿梃 溧眄 桤 10% 鲥眚疣朦眍 镱腩覃 狃桀磬.

绣汨耱疣鲨 桤钺疣驽龛 镳铊玮钿栩 镱耩邃耱忸 爨聒蜞徼痤忄龛 溻箪屦睇 赅痱桧 耧疱耨梃 怃铍 铖 噌聆桉 镱祛 圄翳眄钽 镳孱狃噻钼囗, 扈龛扈玷痼邈 耋祆囵眍 疣耨蝾龛 戾驿 x-觐铕滂磬蜞扈 怦艴 耱疱祗祛 磬 疣珥 赅痱桧圊 耩邃龛 镱腩驽龛屐 耦铗忮蝰蜮簋 耱疱祗祛 镱 怦屐 疱汨耱痂痼屐 桤钺疣驽龛. 阻耠 耱疱祗祛 忄瘘桊箦 玎忤耔祛耱 铗 觌囫襦. 赅麇耱忮 趔黻鲨铐嚯 赅麇耱忄 桉镱朦珙忄磬 戾蝠桕, 铖眍忄眄 磬 恹麒耠屙梃 觌桎钼 疣耨蝾龛.

翌黜铖螯 疱汨耱疣鲨 铞屙桠嚯 镱 忮腓麒礤 耱囗溧痱睇 铗觌铐屙栝 镱腩驽龛 耱疱祗祛 铗 耩邃礤泐 珥圜屙 腩赅腓玎鲨 耦铗忮蝰蜮簋泐 耱疱祗爨 狃桀眍 钿眍泐 蝾泐 驽 怵屐屙眍泐 觌囫襦. 翌黜铖螯 戾蝾溧 疱汨耱疣鲨 恹 狃桀眍 桤 镱玟龛 怵屐屙睇 觌囫耦 箪屙噱蝰 铗 鲥眚疣 狃桀磬 镥痂翦痂. 狃桀眍 怵屐屙眍泐 觌囫襦 8 蝾黜铖螯 疱汨耱疣鲨 鲥眚疣朦眍 10% 镱腩皴 磬躅滂蝰 镳邃咫圊 0.220.47%.

漯筱桴 篦囫蜿圊 狃桀磬  珥圜屙 觐脲犭 铗 0.29 漕 4.42% 忸 祉钽桴 耠篦 猁忄 戾睃 1%. 项耜铍 滂囔弪 钿眍泐 漯 鲥眚疱 狃桀磬 镳栳腓玷蝈朦眍 疣忮 1% 潆桧 狃桀磬 (妮), 磬 镥痂翦痂 镳桁屦眍 1.5 疣玎 犷朦, 镱塍麇眄 珥圜屙 疋桎弪咫蜮簋 恹耦觐 蝾黜铖蜩 疱汨耱疣鲨.

暑眈蝠箬痤忄龛 桧蝈沭桊钼囗睇 赅痱桧 耧疱耨梃 沐眍 皴沆屙蜞鲨. 秒噔眍 鲥朦 镳铖蝠囗耱忮眄铋 疱汨耱疣鲨 箐嚯屙 纛磬 怆弪 镱耱痤屙桢 赅痱 忡噼祉钽 疣耧铍铈屙 钺豚耱彘 耧疱耨梃 怦艴 沐眍 皴沆屙蜞鲨 潆 祉铈羼蜮 狃桀眍 钿桧嚓钼钽 忸琊囫蜞. 吾豚耱 耧疱耨梃 赅驿钽 沐磬 磬 赅痱 狍潴 镳邃耱噔脲睇 疋铄 擙蜞腩眄铋 桦 撹眚邈痂痤忄眄铋 赅痱桧铋 耧疱耨梃, 觐蝾疣 踵疣牝屦桤箦蝰 耩邃龛扈 潆 怦邈 怵屐屙眍泐 觌囫襦 珥圜屙扈 桧蝈眈桠眍耱 綦箢疱聆屙鲨. 蠕遴朦磬 桧蝈沭桊钼囗磬 赅痱桧 耧疱耨梃 溧眄钽 沐磬 祛汶 猁 猁螯 镱耱痤屙 汨镱蝈蜩麇耜铋 耔蝮圉梃, 觐沅 怦 狃桀睇 桁咫 猁 钿桧嚓钼簋 溴痦簋 耱痼牝箴, .. 耦耱腓 猁 桤 祉铈羼蜮 溴 钿桧嚓钼 潆 怦艴 狃桀眍 觐铕滂磬蜞扈. 蜞觐 耠篦噱 耩邃龛 桧蝈眈桠眍耱 綦箢疱聆屙鲨 祛汶 猁 猁螯 恹麒耠屙 礤玎忤耔祛 赅驿铎 漯. 武磬觐, 磬 襦祛 溴脲 觐腓麇耱忸 镱腩驽龛 溴 疣珥 狃桀磬, 蜞赕 镫铗眍耱 疣耧疱溴脲龛 溴 忭篁痂 赅驿钽 桧滂忤潴嚯钽 狃桀磬 礤镱耱眄. 项铎 漕腈磬 猁螯 恹麒耠屙 篑疱漤屙磬 溴痦 耱痼牝箴 赅痱桧 耧疱耨梃, 篦栩帼  铖钺屙眍耱. 蒡 玎溧鬣 疱磬 矬蝈 铒疱溴脲龛 镳铖蝠囗耱忮眄铋 镫铗眍耱 疣耧疱溴脲龛 溴, 黩 镱玮铍桦 恹麒耠栩 耩邃礤 珥圜屙桢 滂囔弪疣 漯 赅驿铋 蝾麝 篑疱漤屙眍泐 狃桀磬.

暑眈蝠箬痤忄龛 溻箪屦睇 桧蝈沭桊钼囗睇 赅痱桧 耧疱耨梃 禧 恹镱腠桦, 桉镱朦珞 疱汨耱痂痤忄眄 溧眄, 潆 狃桀眍 赅驿钽 怵屐屙眍泐 觌囫襦. 碾 钽 赅驿铎 漯 赅驿钽 桧滂忤潴嚯钽 狃桀磬 耱噔桦 耦铗忮蝰蜮桢 漯 篑疱漤屙眍 溴痦铋 耱痼牝箴 赅痱桧 耧疱耨梃, 桁妣 磬栳铍邋 犭桤觇 觐铕滂磬螓, 镱耠 麇泐 恹麒耠腓 耩邃睨 桧蝈眈桠眍耱 綦箢疱聆屙鲨 镱 怦屐 桧滂忤潴嚯 漯囔, 铗礤皴眄 溧眄铎 耩邃礤祗 漯.

软蝈沭桊钼囗磬 赅痱桧 耧疱耨梃 软蝈沭桊钼囗磬 赅痱桧 耧疱耨梃 gt, Kr, eve 软滂忤潴嚯 狃桀 gt, Kr, kni gt, Kr, eve 需. 2: 莉豚 耧疱耨梃 沐眍 忸 怵屐屙眍 觌囫皴 8. . 软蝈沭桊钼囗睇 赅痱桧 耧疱耨梃 gt, Kr eve; .赅痱桧 耧疱耨梃 蝈 驽 沐眍 桧滂忤潴嚯钽 狃桀磬 蝾泐 驽 觌囫襦;

. 桧蝈沭桊钼囗磬 赅痱桧 耧疱耨梃 Kr, kni gt 忤痱筻朦眍 狃桀礤.

橡桁屦 桧蝈沭桊钼囗眍 溻箪屦眍 赅痱桧 耧疱耨梃 沐眍 gt, Kr eve 狃桀眍 忸顸祛泐 怵屐屙眍泐 觌囫襦 镳邃耱噔脲 磬 痂. 2. 佯噔礤龛 铋 赅痱桧 赅痱桧铋 耧疱耨梃 蝈 驽 沐眍 桧滂忤潴嚯钽 狃桀磬 觌囫襦 8 (痂. 2) 镱赅琨忄弪, 黩 桧蝈沭桊钼囗磬 赅痱桧 鲥腩 镳噔桦 镥疱溧弪 铖眍忭 铖钺屙眍耱 耧疱耨梃 桧滂忤潴嚯 沐眍, 怅膻鬣 纛痨 疣珈屦 钺豚耱彘 耧疱耨梃. 橡邃耱噔脲眄 磬 铎 驽 痂耋黻 赅痱桧 耧疱耨梃 沐眍 Kr, kni gt (痂. 2) 桧蝈疱耥 蝈, 黩 耩邃 桉镱朦珙忄眄 溧眄铋 疣犷蝈 狃桀眍 礤 猁腩 狃桀眍, 钿眍怵屐屙眍 铌疣眄 潆 恹怆屙 耧疱耨梃 桴 沐眍. 枢痱桊 桧蝈沭桊钼囗睇 赅痱桧 耧疱耨梃 磬 篑疱漤屙眢 溴痦簋 耱痼牝箴, 蝈镥瘘 祛骓 耦玟噔囹 蜞 磬琨忄屐 "忤痱筻朦睇 狃桀睇" 眍恹扈 觐灬桧圉扈 沐眍 潆 忤珞嚯桤圉梃 忡噼祉钽 疣耧铍铈屙 桴 钺豚耱彘 耧疱耨梃.

碾 觐眈蝠箬痤忄龛 钿眍戾痦 桧蝈沭桊钼囗睇 赅痱桧 耧疱耨梃 沐眍 皴沆屙蜞鲨 觐铕滂磬螓 溴 赅驿铋 铗疱汨耱痂痤忄眄铋 钿眍戾痦铋 赅痱桧 耧疱耨梃 沭箫镨痤忄腓顸 怃铍 铖 x 镱 R 桧蝈疴嚯囔. 青蝈 忭篁痂 赅驿钽 桧蝈疴嚯 恹麒耠腩顸 耩邃礤 珥圜屙桢 桧蝈眈桠眍耱 綦箢疱聆屙鲨 镱 怦屐 狃桀磬 溧眄钽 怵屐屙眍泐 觌囫襦. 阻耠 R 恹徼疣腩顸 桤 蝈 耦钺疣驽龛, 黩 鲥眚疣朦眍 鬣耱 狃桀磬 滂囔弪 钿眍泐 漯 耦耱噔弪 镳桁屦眍 1% 铗 邈 潆桧, , 耠邃钼囹咫, R 漕腈眍 猁螯 疣忭 100 潆 蝾泐, 黩钺 镳噔桦 耢钿咫桊钼囹 溴痦簋 耱痼牝箴 赅痱桧 耧疱耨梃. 武眍戾痦 桧蝈沭桊钼囗睇 赅痱桧 耧疱耨梃 爨蝈痂眈觇 沐眍 忸 怵屐屙眍 觌囫皴 3, 沐眍 gap 沐眍 pair-rule 忸 怵屐屙眍 觌囫皴 8 桤钺疣驽睇 磬 痂. 3.

朽琊噌铗囗睇 磬扈 戾蝾潲 猁腓 镳桁屙屙 潆 镱塍麇龛 觐腓麇耱忮眄 溧眄 钺 耧疱耨梃 沐眍 皴沆屙蜞鲨 赅驿铎 漯 赅驿钽 桤 1580 桧滂忤潴嚯 狃桀眍 (痂. 4 痂. 5), 蜞赕 嚯铐睇, 桧蝈疸痂痤忄眄 溧眄 钺 耧疱耨梃 赅驿钽 桤 14 沐眍 皴沆屙蜞鲨 赅驿铋 钺豚耱 狃桀磬 疣珥 祛戾眚 怵屐屙. 项塍麇眄 溧眄 桁妣 疣琊屮屙桢 镱 镳铖蝠囗耱怏 钿眍 漯 疣琊屮屙桢 镱 怵屐屙 铌铍 6.扈眢 疣玮栩.

麇蜮屦蝾 汶噔 "蔫磬扈麇耜 镳桊钿 镱玷鲨铐眍 桧纛痨圉梃" 钺耋驿帼蝰 戾踵龛珈 纛痨桊钼囗 皴沆屙蝽钽 镳屣囹蝈痦 犭囫蝾溴痨 漯铉铘桦 疋弪 眍恹, 镱塍麇眄 磬扈 溧眄 滂磬扈赍 纛痨桊钼囗 钺豚耱彘 耧疱耨梃 沐眍 皴沆屙蜞鲨. 蒡 疱珞朦蜞螓 镳邃耱噔脲睇 耱囹 A1, A2, A10, A30, A33.

2bcd cad 2110 10 20 30 40 50 60 70 80 90 12Kr hb kni tll 2gt 110 10 20 30 40 50 60 70 80 90 12h eve 2run ftz 110 10 20 30 40 50 60 70 80 90 1 2slp prd 2odd 1 1 0 10 20 30 40 50 60 70 80 90 1碾桧 狃桀磬, % 需. 3: 武眍戾痦 桧蝈沭桊钼囗睇 赅痱桧 耧疱耨梃 爨蝈痂眈觇 沐眍 忸 怵屐屙眍 觌囫皴 3 (), 沐眍 gap () 沐眍 pair-rule (,) 忸 怵屐屙眍 觌囫皴 杨汶囫眍 觌囫耔麇耜铋 蝈铕梃, 镳邃腩驽眄铋 1969 泐潴 .骂腼屦蝾 (Wolpert, 1969;

Wolpert, 1989), 耋潼徉 觌弪觇 铒疱溴弪 邋 镱腩驽龛屐 铒疱溴脲眄铎 镳铖蝠囗耱忮眄铎 镱脲 玎痤潲, 觐蝾痤 耋耱怏弪 沭噤桢眚 觐眦屙蝠圉梃 礤觐 耔沩嚯铋 祛脲牦臌, 磬琨忄屐铋 祛痿钽屙铎. 痒栩囗桢 桧纛痨圉梃 沭噤桢眚 祛痿钽屙 邋 桧蝈痫疱蜞鲨 镳桠钿 滂趑屦屙鲨圉梃 觌弪铌 蝾 桦 桧铎 磬镳噔脲龛 玎忤耔祛耱 铗 箴钼 觐眦屙蝠圉梃 祛痿钽屙. 耦铗忮蝰蜮梃 眍忸, 篁铟礤眄铋 觐眦屣鲨彘, 潆 觌囫耔翳赅鲨 祛脲牦臌 赅麇耱忮 祛痿钽屙 (Wolpert, 1989; Gurdon and Bourillot, 2001) 漕腈眍 恹镱腠螯 礤耜铍 篑腩忤: 祛痿钽屙 漕腈屙 猁螯 疣耧疱溴脲 镱 镳铖蝠囗耱怏 纛痨 镱耱屣屙眍 戾邈铖 沭噤桢眚, 镥疱戾螯 溴轳蜮钼囹 磬 犷朦 疣耨蝾龛, 蜞赕 桉觌栩咫 礤镱耩邃耱忮眄 钺篑豚怆桠囹 玎忤 铗 镱痤汔 桧潴牿棹 镱 牮嚅礤 戾疱 溻篚 疣珥 耦耱龛 耧疱耨梃 沐眍 觌弪赅-扈.

软蝈眈桠眍耱 綦箢疱聆屙鲨 锣骓, 黩 篁铟礤眄 觐眦屣鲨 祛痿钽屙 镱蜩 镱玷鲨铐眍 桧纛痨圉梃 铖眍忄睇 磬 麇蜿铎 疣玢疣龛麇龛 戾驿 纛痨桊钼囗桢 桧蝈痫疱蜞鲨彘 镱玷鲨铐眍 桧纛痨圉梃. 铖眍怏 桴 觐眦屣鲨 玎腩驽磬 耱囹梓羼赅 耔耱屐 觐铕滂磬, 磬豚汔屐 磬 噌耦膻蝽 镟耨桠眢 蜿囗-扈睃 赅驿 祛戾眚 疣玮栩 镱耠 漕耱桄屙 沭噤桢眚铎 耱圉桀磬痦钽 耦耱龛. 绣珞朦蜞螓 囗嚯桤 滂磬扈觇 纛痨桊钼囗 钺豚耱彘 耧疱耨梃 沐眍 皴沆屙蜞鲨, 镱塍麇眄 溧眄铋 疣犷蝈, 溴祛眈蝠桊簋 噌耦膻蝽簋 礤噤尻忄蝽铖螯 蜞觐 觐眦屣蝮嚯铋 祛溴腓 潆 铒桉囗 戾踵龛珈钼 纛痨桊钼囗 镟蝌屦磬 犭囫蝾溴痨 漯铉铘桦.

需. 4: 枢痱桧 耧疱耨梃 爨蝈痂眈觇 沐眍 沐眍 gap 磬鬣脲 (怵屐屙眍 觌囫 1 (1)) 觐眦 (怵屐屙睇 觌囫覃 7 桦 8 (7 桦 8)) 鲨觌 14. 碾 赅驿钽 沐磬 怵屐屙眍泐 觌囫襦 镳桠邃屙 溻箪屦磬 钿眍戾痦 赅痱桧 耧疱耨梃 蜩镨黜钽 桧滂忤潴嚯钽 狃桀磬.

碾 桤篦屙 滂磬扈觇 纛痨桊钼囗 钺豚耱彘 耧疱耨梃 沐眍 祛痿钽屙弪梓羼觐 镱脲 皴沆屙蜞鲨 赅驿 钺豚耱 猁豚 铛囵嚓蝈痂珙忄磬 礤犷朦 麒耠铎 踵疣牝屦睇 镳桤磬觐 (耢. 疣玟咫 1.3). 赅麇耱忮 蜞觇 镳桤磬觐 恹徼疣腓 镱腩驽龛 爨犟桁箪钼 耧疱耨梃, 蜞赕 x-觐铕滂磬螓 蝾麇, 觐蝾瘥 箴钼屙 耧疱耨梃 漕耱桡嚯 礤觐泐 镱痤泐忸泐 珥圜屙, 磬镳桁屦 50% 铗 爨犟桁嚯钽 箴钼.

帖 篑蜞眍忤腓, 黩 犷朦眈蜮 钺豚耱彘 耧疱耨梃 沐眍 皴沆屙蜞鲨 耱囹桉蜩麇耜 珥圜桁 戾 疋铄 镱腩驽龛 镱 戾疱 纛痨桊钼囗. 务钺屙眍 忮腓觇 皲忤汨 鲥眚疣朦眍 钺豚耱 耧疱耨梃 Kr, 蜞赕 玎漤桴 钺豚耱彘 耧疱耨梃 gt, kni hb (痂. 4). 蒡 钺豚耱 皲忤汔 铗 玎漤邈 觐眦 狃桀磬 镥疱漤屐. 亦 磬镳桁屦, 玎漤桢 沭囗桷 钺豚耱彘 耧疱耨梃 Kr, gt kni 耢妁帼蝰 镳桁屦眍 磬 5, 6 15 溴 耦铗忮蝰蜮屙眍 (蜞犭. 1).

滂囔弪 溴, 疣耧铍铈屙睇 怃铍 鲥眚疣朦眍 镥疱漤彗噤礤 铖, 镳栳腓玷蝈朦眍 疣忮 1% 妮 需. 5: 枢痱桧 耧疱耨梃 沐眍 pair-rule 磬鬣脲 (怵屐屙眍 觌囫 1 (1)) 觐眦 (怵屐屙眍 觌囫 8 (8)) 鲨觌 14. 碾 赅驿钽 沐磬 怵屐屙眍泐 觌囫襦 镳桠邃屙 溻箪屦磬 钿眍戾痦 赅痱桧 耧疱耨梃 蜩镨黜钽 桧滂忤潴嚯钽 狃桀磬.

亦犭桷 1: 唁忤汨 钺豚耱彘 耧疱耨梃 沐眍 gap 忸 怵屐屙.

吾豚耱 gt ant hb ant 橡桤磬 A 镨 2 镨 3 P P (141) (61) (145) (144) (173) 软蝈疴嚯 18 58 18 18 1栄溻桡 -1.43 * 0.64 * -5.53 0.06 * -0.吾豚耱 Kr cent kni post gt post hb post 橡桤磬 A P A P A P A P (261) (261) (134) (134) (137) (137) (136) (136) 软蝈疴嚯 18 18 18 18 18 18 38 3栄溻桡 1.02 5.23 1.6 6.32 6.38 15.21 2.02 4.碾 赅驿铋 钺豚耱 耧疱耨梃 镳桠邃屙 忮腓麒睇 皲忤泐 镥疱漤彘 (A) 玎漤彘 (P) 沭囗桷. 碾 镥疱漤彘 钺豚耱 耧疱耨梃 gt 镳桠邃屙 蜞赕 皲忤汨 镱腩驽龛 镨觐 钺豚耱 耧疱耨梃 2 (磬 蝾 桧蝈疴嚯, 沅 铐 耋耱怏弪) 钺豚耱 耧疱耨梃 (痂. 4). 唁忤汨 镱腩骅蝈朦睇, 羼腓 溻桄屙桢 钺豚耱彘 耧疱耨梃 镳铊聃钿栩 铗 玎漤邈 觐眦 镥疱漤屐, 皲忤汨 钺疣蝽铎 磬镳噔脲龛 桁妣 铗痂鲟蝈朦睇 珥圜屙.

软蝈疴嚯 怵屐屙 桉麒耠 忸 怵屐屙睇 觌囫襦. 碾 赅驿铋 钺豚耱 耧疱耨梃 耜钺赅 箨噻囗 钺 恹犷痍 狃桀眍, 桉镱朦珙忄眄铋 潆 恹麒耠屙 皲忤汔.

氢彗漕麝铋 铗戾麇睇 皲忤汨, 礤 怆桢 耱囹桉蜩麇耜 珥圜桁. "ant", "cent" "post"钺铉磬麇睇 钺豚耱 耧疱耨梃, 疣耧铍铈屙睇 镥疱漤彘, 鲥眚疣朦眍 玎漤彘 鬣耱 狃桀磬 耦铗忮蝰蜮屙眍.

灭囗桷 镥疱漤彘 钺豚耱 耧疱耨梃 沐磬 gt (痂. 4) 礤 皲忤汔,  钺豚耱 礤 耋驵弪 耦 怵屐屙屐. 武磬觐, 忸 怵屐屙眍 觌囫皴 4 脲忸 沭囗桷 铋 钺豚耱 磬麒磬弪 纛痨桊钼囹 眍忄 钺豚耱 (镨 2). 吾疣珙忄龛 眍忸 钺豚耱 耦镳钼铈溧弪 皲忤泐 "痤滂蝈朦耜铋 钺豚耱"(镨 3) 磬 螯 溴 磬镳噔脲龛, 镳铗桠铒铍铈眍 溻桄屙棹 铖蜞朦睇 钺豚耱彘 耧疱耨梃 沐眍 gap, .. 玎漤屐 觐眦 狃桀磬 (蜞犭. 1). 桧蝈疴嚯 镥疴钽 镱 忸顸祛 怵屐屙眍 觌囫覃 牮篁桤磬 玎漤彘 沭囗桷 镥疱漤彘 钺豚耱 耧疱耨梃 hb 筲咫梓桠噱蝰. 箨噻囗眍 桧蝈疴嚯 怵屐屙  沭囗桷 皲忤汔弪 磬镳噔脲龛 玎漤屐 觐眦 狃桀磬 镳栳腓玷蝈朦眍 磬 0.9 溴, 钿磬觐 襦爨 镥疱漤 钺豚耱 耧疱耨梃 hb 礤 戾弪 疋铄泐 镱腩驽龛 (蜞犭. 1).

令朦眈蜮 钺豚耱彘 耧疱耨梃 沐眍 pair-rule 蝾驽 桤戾 疋铄 镱腩驽龛 躅溴 鲨觌 14 耱囹桉蜩麇耜 珥圜桁 耢妁帼蝰 镥疱漤屐 镱膻耋 狃桀磬 (蜞犭. 2). 青 桉觌屙桢 h odd 镱腩襦 1 沐眍 pair-rule 腓犷 礤 溻桡噱蝰, 腓犷 皲忤汔弪 铟屙 礤珥圜栩咫. 项腩襦 1 h 皲忤汔弪 镱 磬镳噔脲龛 玎漤屐 觐眦, 镱腩襦 1 odd - 镥疱漤屐, 镳梓屐 镱耠邃龛 皲忤 觐痧咫桊箦 钺疣珙忄龛屐 镱腩覃 2. 袜栳铍邋 珥圜栩咫 皲忤汨 钺磬痼驽睇 潆 镱腩 7 eve, h run, 镳梓屐  镱腩襦 eve 耢妁噱蝰 镳桁屦眍 磬 螯 溴.

亦犭桷 2: 唁忤汨 钺豚耱彘 耧疱耨梃 沐眍 pair-rule 忸 怵屐屙.

沐 / 钺豚耱 1 2 3 4 5 6 eve .. 38 38 38 38 48 48 3(N=654) 皲忤 0.47 1.94 1.97 3.44 3.21 2.62 5.ftz .. 38 38 38 48 38 48 5(N=158) 皲忤 0.20 * 1.27 1.72 1.86 2.27 3.84 1.01 * h .. 38 38 38 48 48 48 4(N=105) 皲忤 -2.18 1.54 3.61 1.11 2.61 3.25 3.odd .. 38 48 38 58 38 38 (N=85) 皲忤 2.41 1.13 * 2.98 0.92 3.65 2.66 run .. 38 48 38 58 38 38 5(N=65) 皲忤 0.20 * 1.66 2.81 2.56 1.57 3.41 3. 赅麇耱忮 踵疣牝屦睇 镳桤磬觐 桉镱朦珙忄眍 爨犟桁嚯铄 珥圜屙桢 桧蝈眈桠眍耱 綦箢疱聆屙鲨 赅驿铋 镱腩皴. 橡铎彐篁觇 怵屐屙, 潆 觐蝾瘥 恹麒耠屙 皲忤汨, 磬麒磬 祛戾眚 镱怆屙 镱腩覃 桉麒耠 忸 怵屐屙睇 觌囫襦 (..). 吾 恹犷痍 N 镳桠邃屙 潆 赅驿钽 沐磬. 唁忤汨 潆 镱腩覃 7 odd 礤 镳桠钿蝰, 镱耜铍 铐 纛痨桊箦蝰 襦祛 觐眦 鲨觌 14A. 唁忤汨 镱腩骅蝈朦睇, 羼腓 溻桄屙桢 钺豚耱彘 镳铊聃钿栩 铗 玎漤邈 觐眦 镥疱漤屐, 皲忤汨 钺疣蝽铎 磬镳噔脲龛 桁妣 铗痂鲟蝈朦睇 珥圜屙. 氢彗漕麝铋 铗戾麇睇 皲忤汨, 礤 怆桢 耱囹桉蜩麇耜 珥圜桁.

理嚯桤 皲忤泐 钺豚耱彘 耧疱耨梃 沐磬 eve 120 狃桀眍, 觐蝾瘥 蝾黜 忸琊囫 猁 铒疱溴脲 矬蝈 桤戾疱龛 耱屣屙 桧忄汨磬鲨 戾灬疣 桉镱朦珙忄龛 耱囗溧痱眍 牮桠铋 (Merrill et al., 1988), 镱赅玎, 黩 躅溴 鲨觌 14 镱腩覃 eve 皲忤汔 镥疱漤屐 镱膻耋 狃桀磬 礤镳屦眍, 躅 耜铕铖螯 溻桄屙 镱腩 5 6 戾礤 疣忭铎屦磬, 麇 蜞觐忄 镱腩覃 7.

拟桄屙桢 礤 牮嚅龛, 疣耧铍铈屙睇 忭篁痂 赅痱桧 耧疱耨梃 镱腩 沐眍 pair-rule, 鬣 怦邈 觐痧咫桊箦 钺疣珙忄龛屐 眍忸 钺豚耱 耧疱耨梃. 亦 磬镳桁屦, 镱腩襦 odd 皲忤汔弪 铟屙 耔朦眍 忸 怵屐屙睇 觌囫襦 3 镱 5, 觐沅 铗 玎漤彘 沭囗桷 铗镱麝钼噱蝰 镱腩襦 2. 氧钿睇 钺疣珙, 钺疣珙忄龛 镱腩覃 7 odd 忸 怵屐屙睇 觌囫襦 5 - 7 耦镳钼铈溧弪 耥圜嚯 耋驽龛屐 镱腩覃 6 桧蝈疴嚯 戾驿 怵屐屙睇扈 觌囫襦扈 7, 玎 觐蝾瘥 耠邃箦 耔朦睇 皲忤 镨赅 铋 镱腩覃 忸 怵屐屙睇 觌囫襦 7 - 8.

项耜铍 礤溧忭 猁腩 镳钿屐铐耱痂痤忄眍 溻桄屙桢 溴 犭囫蝾溴痨 狃桀磬 (Keranen et al., 2006), 蜞赕 怦脲漶蜮桢 蝾泐, 黩 犷朦眈蜮 沐眍 皴沆屙蜞鲨 觐滂痼 蝠囗耜痂秭桀眄 羿牝铕, 镳邃耱噔弪 忄骓 镳铊玮羼蜩 铞屙牦 怅豚溧 祛痿钽屙弪梓羼觇 溻桄屙栝 溴 皲忤汨 钺豚耱彘 耧疱耨梃 桴 沐眍. 铋 鲥朦 骅恹 狃桀眍 铞屙桦 磬 漕瘃嚯铋 忮眚疣朦眍 耱铕铐圊 皲忤汨 蝠艴 溴, 镱腩驽龛 觐蝾瘥 镳桁屦眍 耦怙噤噱 戾耱铎 纛痨桊钼囗 泐腩忭铋 犷痤玟觇 (cf) (漯 31), 镱腩驽龛屐 爨犟桁箪 镱腩覃 3 h 祛戾眚 邋 钺疣珙忄龛 忸 怵屐屙眍 觌囫皴 3 (漯 51) 镱腩驽龛屐 玎漤彘 沭囗桷 鲥眚疣朦眍 钺豚耱 耧疱耨梃 Kr 忸 怵屐屙眍 觌囫皴 1 (漯 62).

31 T6.5 扈.

cf 玎漤 沭囗桷 Kr 31 51 T19.5 扈.

镱腩襦 3 h cf 31 47 51 57 00110000 0001111 111T52 扈.

镱腩襦 3 h 玎漤 沭囗桷 Kr cf A- 250 h 忸 怵屐屙眍 觌囫皴 run 忸 怵屐屙眍 觌囫皴 200 h 忸 怵屐屙眍 觌囫皴 run 忸 怵屐屙眍 觌囫皴 1h 爨犟桁箪 镱腩覃 1忸 怵屐屙睇 觌囫襦 3 run 扈龛祗 戾驿 镱腩襦扈 2 3 忸 怵屐屙睇 觌囫襦 3 20 25 30 35 40 45 50 55 % 碾桧 蒽狃桀磬 需. 6: 拟桄屙桢 溴 in vivo 耩噔礤龛 耦 皲忤汔扈 钺豚耱彘 耧疱耨梃. 相礤腓 (), (), () - 耦铗忮蝰蜮簋 箨噻囗睇 祛戾眚囔 怵屐屙 赅漯 翳朦爨, 溴祛眈蝠桊簋泐 漕瘃嚯簋 耱铕铐 骅忸泐 狃桀磬. () - 钿眍戾痦 沭圄桕 赅痱桧 耧疱耨梃 run h 忸 怵屐屙睇 觌囫襦 3 8. 吴嚯囔, 磬痂耦忄眄 矬黻蜩痦铋 腓龛彘, () 钺铉磬麇睇 镱腩驽龛 玎漤彘 沭囗桷 Kr 镱腩覃 3 h 忸 怵屐屙眍 觌囫皴 8. 氧钿睇 疱珞朦蜞螓 猁腓 镱塍麇睇 磬 漕瘃嚯铋 忮眚疣朦眍 耱铕铐圊 15 狃桀眍. () 镱赅玎眍, 黩 镱腩驽龛 爨犟桁箪 镱腩覃 3 h 扈龛祗爨 戾骘铍铖眍泐 镳铎彐篁赅 2/3 run 皲忤汔 铗眍耔蝈朦眍 漯筱 漯筱 桧蝈疴嚯 戾驿 怵屐屙睇扈 觌囫襦扈 3 8.

侮噻嚯铖 (痂.6), 黩 躅溴 鲨觌 14 漯 51 62 皲忤汔 镥疱漤屐 觐眦 狃桀磬 磬 疣耨蝾龛, 耦铗忮蝰蜮簋 镱腩忤礤 滂囔弪疣 漯. 泥朦礤轼桢 漕赅玎蝈朦耱忄 蝾泐, 黩 钺磬痼驽眄 溧眄铋 疣犷蝈 皲忤汨 钺豚耱彘 耧疱耨梃 礤 钺篑腩怆屙 溻桄屙桢 溴, 祛骓 镱塍麒螯, 疣耨祛蝠邂 镱忮溴龛 戾骘铍铖眍泐 镳铎彐篁赅 2/3 沐磬 run 镱腩覃 3 h. 觐眦 鲨觌 14, 忸 怵屐屙眍 觌囫皴 8, 赅痱桧 耧疱耨梃 桴 沐眍 耱痤泐 觐祜脲戾眚囵睇 , 耠邃钼囹咫 爨犟桁箪 耧疱耨梃 h 扈龛祗 耧疱耨梃 run 软蝈眈桠眍耱 綦疱聆屙鲨 腩赅腓珞 钿眍 戾耱 (漯). 武磬觐, 赅 耠邃箦 桤 痂耋黻 6, 磬 疣眄桴 耱噤 钽 鲨觌 戾骘铍铖睇 镳铎彐篁铌 2/3 run 镱腩襦 3 h 腩赅腓珞 疣珥 戾耱圊 狃桀磬. 亦觇 钺疣珙, 躅溴 鲨觌 14 戾骘铍铖睇 镳铎彐篁铌 2/3 run 镱腩襦 3 h 皲忤汔 铗眍耔蝈朦眍 漯筱 漯筱. 西邂桎眍, 黩  怆屙桢 礤 祛驽 猁螯 钺篑腩怆屙 溻桄屙桢 溴.

蒡 羿牝, 蜞赕 溧眄 钺 囫桁戾蝠梓眍 疣耧铍铈屙梃 钺豚耱彘 腩赅腓玎鲨 煨褪 沐眍 皴沆屙蜞鲨 铗眍耔蝈朦眍 钺豚耱彘 腩赅腓玎鲨 耦铗忮蝰蜮簋 徨腙钼 (c. 蝮 汶噔), 疋桎弪咫蜮簋 蝾, 黩 皲忤汨 钺豚耱彘 耧疱耨梃 狍潴 玎痤潲忸 镱腩耜 怆 耠邃耱忤屐 疱泱鲨 嚓蜩忭铖蜩 沐眍 礤 钺篑腩怆屙 溻桄屙桢 溴.

蒡 皲忤汨 镱 镱潢 忮腓麒睇 耦怙噤帼 疣珈屦囔 镱腩 耧疱耨梃 沐眍 pair-rule (35 溴) , 耠邃钼囹咫, 铟屙 忄骓 潆 镱玷鲨铐桊钼囗 钺豚耱彘 耧疱耨梃 沐眍忪桫屙彘 潆 纛痨桊钼囗 皴沆屙蝽钽 镳屣囹蝈痦.

锣骓, 黩 镳铗桠铒铍铈眍耱 漯囔, 疣耧铍铈屙睇 狍潴 玎痤潲忸 镱腩耜, 漯 31, 疣耧铍铈屙眍 戾耱 纛痨桊钼囗 泐腩忭铋 犷痤玟觇, 皲忤汔弪 玎漤屐 觐眦 狃桀磬 镳桁屦眍 磬 疣耨蝾龛 疣忭铄 2 漯囔 (痂.6). 蒡铗 皲忤 耦怙噤噱 镱 磬镳噔脲龛 耦镱耱噔桁 镱 忮腓麒礤 疣珈屦囔 皲忤泐 钺豚耱彘 耧疱耨梃 沐眍 皴沆屙蜞鲨, 腩赅腓珙忄眄 泐腩忭铎 铗溴脲 狃桀磬 (耢. 痂. 4 痂. 5). 亦觇 钺疣珙, 礤 桉觌屙, 黩 皲忤汨 钺豚耱彘 耧疱耨梃 沐眍 皴沆屙蜞鲨 泐腩忭铎 铗溴脲 祛泱 猁螯 钺篑腩怆屙 溻桄屙桢 溴.

枢 桤忮耱眍, 沭噤桢眚 Bcd 猁 镥疴 祛痿钽屙弪梓羼觇 沭噤桢眚铎, 觐蝾瘥 箐嚯铖 筲桎弪 礤镱耩邃耱忮眄 耧屦桁屙蝈 (Driever and Nsslein-Volhard, 1988b; Driever and Nsslein-Volhard, 1988a). 武磬觐, 耠邃箦 铗戾蜩螯, 黩 镱扈祛 Bcd 犭囫蝾溴痨 漯铉铘桦 耋耱怏弪 怛铕铋 爨蝈痂眈觇 沭噤桢眚 Hb, 黩 铒疱溴脲龛 镱腩驽龛 沭囗桷 钺豚耱彘 耧疱耨梃 沐眍-扈礤 镳铊聃钿栩 疱珞朦蜞蝈 耔礤疸弪梓羼觐泐 忡噼祛溴轳蜮 桴 溻篚 沭噤桢眚钼 (Driever and Nsslein-Volhard, 1988b; Simpson-Brose et al., 1994; Reinitz et al., 1995). 署铎 蝾泐, 磬 疣犷蝈 镱赅玎眍, 黩 觐眦屙蝠圉 祛痿钽屙 Bcd 礤镱耱眄 耔朦眍 戾弪 忸 怵屐屙, 钺豚耱 耧疱耨梃 沐眍 gap pair-rule, 怆桴 扈扈 溴轳蜮 爨蝈痂眈觇 沭噤桢眚钼, 戾 疋铄 镱腩驽龛 镱 戾疱 疣玮栩. 埋  铉磬鬣弪, 黩 镱玷鲨铐眢 桧纛痨圉棹 犭囫蝾溴痨 漯铉铘桦 礤朦 疣耨爨蝠桠囹 赅 礤觐 镱耱眄铄, 玎溧眄铄 觐眦屙蝠圉桢 祛痿钽屙 镱玷鲨铐眍 麒耠. 袜钺铕铗, 溧眄 溻桄屙梃 钺豚耱彘 耧疱耨梃 沐眍-扈礤 疋桎弪咫蜮簋 蝾, 黩 镱玷鲨铐磬 桧纛痨圉 礤 怆弪 耱囹梓眍, 镱耱眄 猁耱痤 戾弪. 枢 镱赅玎眍 蝾 汶噔 滂耨屦蜞鲨 (耢. 龛驽), 皲忤汨 篁铟礤龛 沭囗桷 耧疱耨梃 沐眍 gap 镳铊聃钿 玎 聍弪 忡噼祉铋 疱泱鲨 桴 沐眍 礤 玎忤 铗 溴轳蜮 爨蝈痂眈觇 沭噤桢眚钼 滂趑箸梃 镳钿箨蝾 沐眍 gap 戾驿 漯囔. 锑蝈痂眈觇 沭噤桢眚 沐睇 gap 耦忪羼蝽 铒疱溴 镱腩驽龛 沭囗桷 耧疱耨梃 沐眍 pair-rule, 觐蝾瘥 蝾驽 皲忤汔 镱 戾疱 疣玮栩. 朽珞祉 镳邃镱腩骅螯, 黩 蜞觇 皲忤汨 怆 磬 镱腩驽龛 镱腩 耧疱耨梃 沐眍 segment polarity , 蝈 襦禧, 磬 镱腩驽龛 镟疣皴沆屙蝽 沭囗桷 (Ingham and Martinez-Arias, 1992).

疋弪 怦邈 恹 耜噻囗眍泐, 镱玷鲨铐眢 桧纛痨圉棹 犭囫蝾溴痨 漯铉铘桦 耠邃箦 疣耨爨蝠桠囹 赅 滂磬扈麇耜 戾簋 忸 怵屐屙 觐灬桧圉棹 觐眦屙蝠圉栝 镳钿箨蝾 爨蝈痂眈觇 玷泐蜩麇耜桴 沐眍. 赅驿 祛戾眚 怵屐屙  觐灬桧圉 铒疱溴弪 礤 蝾朦觐 爨蝈痂眈觇扈 祛痿钽屙囔, 眍 皲忤汔扈 沭囗桷 钺豚耱彘 耧疱耨梃 沐眍 皴沆屙蜞鲨 疱珞朦蜞蝈 疱泱蝾痦 忡噼祛溴轳蜮栝. 蒡 蝠嚓蝾怅 镱漯噻箪邂噱 嚓蜩忭 耧铖钺 桧蝈痫疱蜞鲨 沭噤桢眚 祛痿钽屙 疣珈噱 沭囗桷 戾驿 纛痨桊钼囗桢 桧蝈痫疱蜞鲨彘 镱玷鲨铐眍 桧纛痨圉梃.

蜞 汶噔 "绣泱蝾痦 忡噼祛溴轳蜮 皴蜩 沐眍 gap" 镱疋磬 疱珞朦蜞蜞 爨蝈爨蜩麇耜钽 祛溴腓痤忄龛 戾踵龛珈钼 疱泱鲨 耧疱耨梃 沐眍 gap 鲨觌 14. 碾 漕赅玎蝈朦耱忄 耋耱忸忄龛 狃桀礤 漯铉铘桦 镳邃耜噻囗睇 疱泱蝾痦 戾踵龛珈钼 桉镱朦珙忄睇 桤忮耱睇 桤 腓蝈疣蝮瘥 溧眄. 锐耠邃钼囗 蜞赕 戾踵龛珈, 钺羼镥麒忄栝 皲忤汨 沭囗桷 钺豚耱彘 耧疱耨梃 沐眍 gap.

蒡铗 爨蝈痂嚯 桤腩驽 疣犷蜞 A4, A10, A11, 18, A24, A28, A34, A37, A41, A44.

填溴腓痼弪 耧疱耨 沐眍 皴蜩, 怅膻鬣彘 沐睇 bcd, cad, hb, Kr, gt, kni, tll, 潆 麇泐 桉镱朦珞弪 戾蝾, 磬玮囗睇 戾蝾漕 沐眄 皴蝈 (gene circuits) 怙屦恹 镳邃腩驽眄 (Mjolsness et al., 1991; Reinitz et al., 1995). 祛溴腓 疣耨爨蝠桠噱蝰 镱忮溴龛 耔耱屐 鲥镱麝 溴, 疣耧铍铈屙眍 怃铍 鲥眚疣朦眍 铖 狃桀磬 磬镳噔脲龛 A朠 玎觌屙眍 桧蝈疴嚯 戾驿 觐铕滂磬蜞扈, 疣忭 35% 92% 妮. 项忮溴龛 溴 忸 怵屐屙 祛溴腓痼弪 镱祛 蝠艴 镱耠邃钼囹咫 镳铞羼耦: 桧蝈痿噻, 扈蝾 溴脲龛. 惕蝾 铗腓鬣弪 铗 桧蝈痿噻 蝾朦觐 铗耋蝰蜮桢 耔眚彗 徨腙. 腻脲龛 漯 怆弪 滂耜疱蝽 忸 怵屐屙.

a 马篁疱眄邋 耦耱龛 i-泐 漯 铒疱溴弪 磬犷痤 觐眦屙蝠圉栝 vi 蝠囗耜痂秭桀眄 羿牝铕钼, 觐滂痼屐 沐磬扈 皴沆屙蜞鲨 桧溴犟囔 a. 如戾礤龛 觐眦屙蝠圉栝 忸 a 怵屐屙 dvi /dt 玎忤耔 铗 桧蝈眈桠眍耱 蝠艴 镳铞羼耦, 镳铊聃钿 蝈麇龛 桧蝈痿噻 耔眚彗, 滂趑箸梃 疣耧噤 徨腙, 镳邃耱噔脲眄 耋祆铋 耠嚆噱禧 镳噔铋 鬣耱 耠邃簋泐 礤腓礤轫钽 滂耜疱蝽钽 箴噔礤龛 疱嚓鲨-滂趑箸梃:

a dvi N ab b Bcd = Rag T vi + mavi + ha b=dt a a a a (2) +Da(n) (vi-1 - vi ) + (vi+1 - vi ) a -avi, 沅 a = 1,..., N N 麒耠 玷泐蜩麇耜桴 沐眍 祛溴腓.

ab 洛腓麒睇 T (2) 镳邃耱噔 爨蝠桷 觐翳鲨屙蝾 疱泱蝾痦 忡噼祛溴轳蜮栝 Bcd 踵疣牝屦桤簋 疱泱蝾痦铄 溴轳蜮桢 镳钿箨蜞 沐磬 b 磬 耧疱耨棹 沐磬 a. vi 铒桉噱 觐眦屙蝠圉棹 祛痿钽屙 Bcd 漯 i. 绣泱鲨 徨腙铎 Bcd 嚓蜩忭铖蜩 沐磬 a 铒桉噱蝰 镟疣戾蝠铎 ma. 相疣戾蝠 ha 踵疣牝屦桤箦 疱泱蝾痦 怅豚 漯筱桴 爨蝈痂眈觇 羿牝铕钼 蝠囗耜痂秭梃, 疣忭铎屦眍 疣耧疱溴脲眄 犭囫蝾溴痨 狃桀磬. 悟眍耔蝈朦磬 桧蝈眈桠眍耱 耔眚彗 铒桉噱蝰 镱祛 耔沆铊漤铋 趔黻鲨 疱泱鲨钖耧疱耨梃 N 1 ab b Bcd g(ua) = ua/ (ua)2 + 1 + 1, 沅 ua = T vi + mavi + ha 羼螯 耋祆囵睇 怅豚 b=疱泱蝾痤 耧疱耨棹 沐磬 a. 锑犟桁嚯 桧蝈眈桠眍耱 耔眚彗 镳钿箨蜞 沐磬 a 钺铉磬麇磬 麇疱 Ra. 相疣戾蝠 a 羼螯 耜铕铖螯 疣耧噤 镳钿箨蜞 沐磬 a 耦铗眍耔蝰 镥a a 痂钿铎 1/2 镱塍疣耧噤 溧眄钽 徨腙 镱 纛痨箅 1/2 = ln 2/a. 暑翳鲨屙 滂趑箸梃 Da(n) 钺疣蝽 镳铒铕鲨铐嚯屙 赈噤疣蝮 疣耨蝾龛 戾驿 耦皴漤桁 漯囔.

暑眦屙蝠圉梃 徨腙钼, 钺篑腩怆屙睇 爨蝈痂眈觐 耧疱耨桢 沐眍 hb cad (撿囹屦桧耜桢 沭噤桢眚麛), 桉镱朦珞 赅 磬鬣朦睇 篑腩忤 潆 箴噔礤龛 鲨觌 13. 琼圜屙 怦艴 镟疣戾蝠钼 磬躅蝰 矬蝈 镱溷铐觇 疱龛 祛溴腓 耧屦桁屙蜞朦睇 溧眄 镱 耧疱耨梃 9 镱耠邃钼囹咫 祛戾眚圊 怵屐屙 - 鲨觌 13 8 怵屐屙睇 觌囫襦 鲨觌 14, .. 疱弪 钺疣蝽 玎溧鬣 爨蝈爨蜩麇耜钽 祛溴腓痤忄龛. 惕龛扈玷痼弪 趔黻鲨铐嚯 赅麇耱忄, 疣忭 耋祆 赈噤疣蝾 疣珥铖蝈 戾驿 耧屦桁屙蜞朦睇扈 溧眄 疱珞朦蜞蜞扈 疣聍弪 镱 祛溴腓:

a a E = (vi (t)model - vi (t)data)2 + Epenalty, (3) 沅 Epenalty - 疣繇 趔黻鲨. 洋祆桊钼囗桢 镳钼钿栩 镱 怦屐 桁妣祚 溧眄 Nd, 蝾 羼螯, 镱 怦屐 桤戾疱眄 觐眦屙蝠圉 徨腙 潆 怦艴 沐眍 a, 怦艴 溴 i 怦艴 怵屐屙睇 觌囫耦 t. 碾 磬躅驿屙 镟疣戾蝠钼 祛溴腓 桉镱朦珙忄豚顸 镟疣腚咫 忮瘃 耱铛囫蜩麇耜钽 戾蝾溧 铒蜩扈玎鲨 - 戾蝾溧 麒耠屙眍泐 铗骅汔 肃爨 (PLSA). 蒡铗 戾蝾 镱漯钺眍 铒桉囗 Metropolis et al., 1953; Kirkpatrick et al., 1983; Lam and Delosme, 1988a;

Lam and Delosme, 1988b; Reinitz et al., 1995; Chu, 2001. 亦 赅 PLSA 桁邋 耱铛囫蜩麇耜簋 镳桊钿, 趔黻鲨铐嚯 赅麇耱忄 桁邋 祉钽 腩赅朦睇 扈龛祗祛, 蝾 铒蜩爨朦睇 珥圜屙 镟疣戾蝠钼, 磬殇屙睇 疱珞朦蜞蝈 礤耜铍桴 镳铛钿钼 嚯泐痂蜢, 祛泱 铗腓鬣螯 溧驽 镳 聃钿睇 珥圜屙 趔黻鲨铐嚯. 磬 疣犷蝈 赅驿 麒耠屙睇 耧屦桁屙 玎觌嚯 磬躅驿屙梃 66 镟疣戾蝠钼 潆 耔耱屐 桤 348 滂趑屦屙鲨嚯 箴噔礤龛, 钿桧 鲨觌 疣犷螓 嚯泐痂蜢 玎龛爨 磬 觌囫蝈疱 桤 10 镳铞羼耦痤 Intel Xeon2.4GHz 铗 漕 160 鬣耦 玎忤耔祛耱 铗 磬鬣朦眍泐 镳栳腓驽龛.

疱珞朦蜞蝈 疣聍弪钼 猁腩 铗钺疣眍 10 皴蝈, 赅痱桧 耧疱耨梃 觐蝾瘥 磬躅滂腓顸 躅痤 耦铗忮蝰蜮梃 溧眄. 朽耧疱溴脲龛 疣珉梓睇 觐翳鲨屙蝾 爨蝠桷 疱泱蝾痦 忡噼祛溴轳蜮栝 皴 礤 钿桧嚓钼, 眍 潆 犷朦眈蜮 镟疣戾蝠钼 篑蝾轺桠 蝈礓屙鲨 铒疱溴脲眄铎 蜩矬 疱泱蝾痦钽 忡噼祛溴轳蜮.

袜 痂. 7 镳桠邃屙 溧眄 踵疣牝屦 忡噼祛溴轳蜮 爨蝈痂眈觇 沐眍 沐眍 gap.

妈漤, 黩 沐睇 bcd cad 铖眍忭铎 嚓蜩忤痼 沐睇 gap, 嚓蜩忭铖螯 hb, kni, gt, Kr 疱泱腓痼弪 镱 镳桧鲨矬 噔蝾嚓蜩忄鲨. 缅睇 gap 腓犷 礤 忡噼祛溴轳蜮簋, 腓犷 疱镳羼耔痼 漯筱 漯筱. 锐觌屙桢 怆弪 溴轳蜮桢 Gt 磬 hb. 袜栳铍邋 耔朦磬 忡噼祉 疱镳羼耔 磬犭噱蝰 戾驿 镟疣扈 沐眍 gt Kr, 蜞赕 戾驿 hb kni, 镳梓屐 钺豚耱 嚓蜩忭铖蜩 桴 沐眍 礤 镥疱牮帼蝰. 橡钿箨 蝈痨桧嚯钽 沐磬 gap tll 疱镳羼耔痼弪 怦 漯筱桢 沐睇 gap, 牮铎 hb.

bcd cad hb Kr gt kni tll hb 0/0/10 0/0/10 0/0/10 0/10/0 0/2/8 10/0/0 1/8/Kr 0/0/10 0/0/10 9/1/0 0/0/10 10/0/0 9/1/0 10/0/gt 0/0/10 0/0/10 9/0/1 10/0/0 0/3/7 1/9/0 10/0/kni 3/0/7 0/0/10 10/0/0 6/4/0 9/1/0 0/0/10 10/0/0. 0.0.-0.-0.0. 0.0.-0.-0.需. 7: 朽耧疱溴脲龛 觐翳鲨屙蝾 爨蝠桷 疱泱蝾痦 忡噼祛溴轳蜮栝. . 孰囫耔翳赅鲨 镟疣戾蝠钼 镱 蜩矬 忡噼祛溴轳蜮. 羊痤觇 - 沐睇-扈龛 a, 耱铍狯 - 沐睇疱泱蝾瘥 b (耢. 箴噔礤龛 (2)). 琼圜屙 镟疣戾蝠钼 T m -0.005 觌囫耔翳鲨痤忄腓顸 赅 疱镳羼耔 疱泱蝾痤 b 溴轳蜮 沐磬-扈龛 a, 珥圜屙 桧蝈疴嚯 铗 -0.0漕 0.005 觌囫耔翳鲨痤忄腓顸 赅 铗耋蝰蜮桢 忡噼祛溴轳蜮, 珥圜屙 镟疣戾蝠钼 0.0觌囫耔翳鲨痤忄腓顸 赅 嚓蜩忄鲨. 阻耠 耦铗忮蝰蜮簋 麒耠 皴蝈, 觐蝾瘥 溧眄铄 珥圜屙桢 镟疣戾蝠 铉磬鬣弪 疱镳羼耔/铗耋蝰蜮桢 忡噼祛溴轳蜮/嚓蜩忄鲨 耦铗忮蝰蜮屙眍.

吾睇 梏蝾 恹溴脲睇 忡噼祛溴轳蜮, 觌囫耔翳鲨痤忄眄 犷朦眈蜮 皴蝈 赅 嚓蜩忄鲨, 牦瘃桠 铉磬鬣弪 铗耋蝰蜮桢 忡噼祛溴轳蜮, 骅痦 梏蝾 铗戾麇睇 忡噼祛溴轳蜮, 觌囫耔翳鲨痤忄眄 赅 疱镳羼耔. -. 灭圄桕 疣玑痤襦 珥圜屙栝 镟疣戾蝠钼 m T 镳 疱泱鲨 耧疱耨梃 Kr (), kni (), gt () hb ().

琼圜屙桢 镟疣戾蝠 琼圜屙桢 镟疣戾蝠 m m T T T T T T T T T T T T Kr Kr Kr Kr Kr Kr Kr kni kni kni kni kni kni kni cad hb Kr gt kni tll Kr kni cad hb gt tll m m T T T T T T T T T T T T gt gt gt gt gt gt hb hb hb hb hb hb gt hb kni tll cad hb Kr gt cad hb Kr gt kni tll 项扈祛 囗嚯桤 爨蝠桷 疱泱蝾痦 忡噼祛溴轳蜮栝, 鲥朦 恹怆屙 耧弼梏梓羼觇 疱泱蝾痦 忡噼祛溴轳蜮栝, 钺篑豚怆桠帼 纛痨桊钼囗桢 沭囗桷 钺豚耱彘 耧疱耨梃 沐眍-扈礤, 猁 镳钼邃屙 沭圄梓羼觇 囗嚯桤 觐灬桧桊钼囗眍泐 怅豚溧 疣珥 ab b Bcd 疱泱蝾痤 (T vi mavi ) 疱泱鲨 沐磬-扈龛.

灭圄梓羼觇 囗嚯桤 镱赅玎, 黩 Bcd 忭铖栩 忮耦禧 怅豚 嚓蜩忄鲨 镥疱漤桴 钺豚耱彘 嚓蜩忭铖蜩 gt hb, 蜞赕 鲥眚疣朦眍 钺豚耱 耧疱耨梃 Kr. Cad 铌噻噱 嚓蜩忤痼邋 溴轳蜮桢 磬 耧疱耨棹 kni gt 玎漤彘 镱腩忤礤 狃桀磬. 令脲 蝾泐, Cad 忭铖栩 铗眍耔蝈朦眍 犷朦 怅豚 嚓蜩忄鲨 Kr 鲥眚疣朦眍 钺豚耱 狃桀磬, 蜞赕 嚓蜩忄鲨 耧疱耨梃 hb 镥疱漤彘 镱腩忤礤 狃桀磬 磬 疣眄桴 耱噤 疣玮栩.

络噼祉 疱镳羼耔痼桢 忡噼祛溴轳蜮 沐眍 gap 腩赅腓珙忄睇 镳铖蝠囗耱忮. 袜镱祉桁, 黩 Kr gt 桁妣 礤镥疱皴赅桢 钺豚耱 耧疱耨梃 犭囫蝾溴痨 (痂. 3, 8a).

骂 怦艴 皴 Gt 疱镳羼耔痼弪 Kr (痂. 7,). 蒡 忡噼祛溴轳蜮桢 铒疱溴弪 镱腩驽龛 赅 镥疱漤彘, 蜞 玎漤彘 沭囗桷 钺豚耱 嚓蜩忭铖蜩 Kr 鲨觌 14A (痂. 8). 吾豚耱 嚓蜩忭铖蜩 hb kni 镥疱皴赅 钺豚耱 嚓蜩忭铖蜩 Kr (痂. 3, 痂. 8). 犷朦眈蜮 皴蝈 Hb Kni 疱镳羼耔痼 Kr, 钿磬觐  忸玟彘耱忤 怆 犷脲 耠噌, 麇 溴轳蜮桢 Gt (痂. 7,). Kni 篦囫蜮箦 铒疱溴脲龛 镱腩驽龛 玎漤彘 沭囗桷 鲥眚疣朦眍 钺豚耱 嚓蜩忭铖蜩 Kr, Hb - 镥疱漤彘.

需. 8: 理嚯桤 溴轳蜮 疱镳羼耦痤, 铒疱溴桴 镱腩驽龛 沭囗桷 钺豚耱 耧疱耨梃 Kr. ( ) 绣龛 箴噔礤龛 祛溴腓 镱玟礤 鲨觌 14A. ( ) 橡铘桦 疱镳羼耔 潆 Kr 镱玟礤 鲨觌 14A. 洋祆囵睇 怅豚 疱泱蝾痤 uKr 镱赅玎 耧腩铋 麇痦铋 腓龛彘 赅 趔黻鲨 镱腩驽龛 怃铍 镳钿铍铋 铖 狃桀磬 镳铞屙蜞. 青牮帏屙睇 钺豚耱 镱赅琨忄 桧滂忤潴嚯 怅豚 疱泱蝾痤. 妙痂珙眚嚯 矬黻蜩痦 腓龛 钽疣龛麒忄 箴钼龛 疱泱鲨, 龛驽 觐蝾瘥 耧疱耨 戾睃 10%(龛骓 腓龛) , 恹 觐蝾痤泐 犷朦 90% (忮瘐 腓龛) 铗 爨犟桁嚯钽 箴钼. 洛痱桕嚯 矬黻蜩痦 腓龛 钽疣龛麒忄 钺豚耱 磬 镳钿铍铋 铖, 沅 ua 龛驽 10% 箴钼 嚓蜩忭铖蜩. 羊疱腙 箨噻噱 磬 礤珥圜栩咫 箴钼屙 溴疱镳羼耔 Kr 铗耋蝰蜮梃 Gt. 氢彗漕麝 镱赅琨忄 磬耜铍 皲忤礤蝰 沭囗桷 钺豚耱 耧疱耨梃 Kr 铗耋蝰蜮梃 Hb Kni.

绣珞朦蜞螓 沭圄梓羼觐泐 囗嚯桤 觐灬桧桊钼囗眍泐 怅豚溧 疣珥 疱泱蝾痤 疱泱鲨 铖蜞朦睇 沐眍 gap (hb, kni gt) 躅痤 耦汶囫簋蝰 疱珞朦蜞蜞扈 囗嚯桤 觐翳鲨屙蝾 爨蝠桷 疱泱蝾痦 忡噼祛溴轳蜮栝 (痂. 7).

亦觇 钺疣珙, 磬扈 钺磬痼驽眍 螯 徉珙恹 疱泱蝾痦 戾踵龛珈钼, 钺羼镥麒忄桴 忡噼祉铄 疣耧铍铈屙桢 钺豚耱彘 耧疱耨梃 沐眍 gap 狃桀礤 (痂. 9):

嚓蜩忄鲨 耧疱耨梃 沐眍 gap 镱 溴轳蜮桢 Bcd Cad, 疣耧痤耱疣 磬 痤觇 钺豚耱 狃桀磬;

噔蝾嚓蜩忄鲨 沐眍 gap;

耔朦磬 疱镳羼耔 镱 镳桧鲨矬 钺疣蝽铋 疋玷 戾驿 沐磬扈 gap 礤镥疱牮帼扈 钺豚耱扈 耧疱耨梃;

囫桁戾蝠梓磬 疱镳羼耔 戾驿 沐磬扈 gap, 鼽 钺豚耱 耧疱耨梃 镥疱牮帼蝰 疱镳羼耔 镱 镳桧鲨矬 镳祛 疋玷 耧疱耨梃 沐眍 gap 玎漤彘 鬣耱 狃桀磬 镱 溴轳蜮桢 蝈痨桧嚯钽 沐磬 tll.

需. 9: 氧屐囹梓羼觐 镳邃耱噔脲龛 蝾镱沭圄梃 皴蜩 沐眍 gap. 吾豚耱 耧疱耨梃 沐眍 hb, kni, gt, Kr tll 镱赅玎睇 忤溴 镳祛筱铍桕钼. 襄疱漤 鬣耱 狃桀磬 耠邂. 氧屐 铖眍忄磬 磬 溧眄, 镳邃耱噔脲眄 磬 痂耋黻 7a, 镳梓屐 镱赅玎睇 蝾朦觐 蝈 忡噼祛溴轳蜮, 觐蝾瘥 磬犭嚯桉 9 皴 gap 桤 10. 绣镳羼耔 戾驿 沐磬扈 镱赅玎磬 忤溴 耦邃桧栩咫 腓龛 篑桕囔. 肘弪 纛磬 钺铉磬鬣弪 钺豚耱, 桉稃螓忄桢 嚓蜩忤痼邋 溴轳蜮桢 徨腙钼 Bcd (蝈祉-皴瘥) Cad (疋弪腩-皴瘥).

理嚯桤 疱珞朦蜞蝾 麒耠屙眍泐 祛溴腓痤忄龛 滂磬扈觇 耧疱耨梃 沐眍 gap 镱玮铍弪 恹忤螯 疱泱蝾痦 戾踵龛珈, 钺篑豚怆桠帼 皲忤汨 沭囗桷 钺豚耱彘 耧疱耨梃 桴 沐眍. 悟戾蜩, 黩 麒耠屙眍 祛溴腓痤忄龛 怆弪 磬耱 怵屐 邃桧耱忮眄 漕耱箫睇 耧铖钺铎 疱龛 铋 镳钺脲禧 耔塍 礤忸珈铈眍耱 桤篦屙 邋 沐礤蜩麇耜桁 戾蝾溧扈, 镱耜铍 祗蜞眚, 觐蝾瘥 蜞觇 皲忤汨 铗耋蝰蜮簋, 镱赅 礤 镱塍麇睇.

绣龛 祛溴朦眍 玎溧麒, 耦铗忮蝰蜮簋 磬殇屙睇 铒蜩爨朦睇 珥圜屙 镟疣戾蝠钼, 恹耦觐 蝾黜铖螯 忸耧痤桤忸滂 皲忤汨 钺豚耱彘 耧疱耨梃 沐眍 gap 狍潴 玎痤潲忸 镱腩耜 狃桀磬. 令脲 蝾泐, 桉耠邃钼囗桢 镱忮溴龛 祛溴腓 镱赅玎腩, 黩 耔眚彗 徨腙 镳铊聃钿栩 桉觌栩咫 镥疱漤彘 镱腩忤礤 赅驿铋 桤 桉耠邃箦禧 钺豚耱彘 (鲥眚疣朦眍 钺豚耱 耧疱耨梃 Kr, 钺豚耱彘 耧疱耨梃 kni gt 玎漤彘 镱腩忤礤 狃桀磬), 蝾沅 赅 玎漤彘 鬣耱 钺豚耱彘 耧疱耨梃 镳孱犭噤噱 镳铞羼 疣耧噤 耦铗忮蝰蜮簋 徨腙钼. 谚眚彗 徨腙 镥疱漤彘 鬣耱 疣耧噤 徨腙 玎漤彘 鬣耱 赅驿铋 桤 钺豚耱彘, 溴轳蜮 耦忪羼蝽, 皲忤汔  钺豚耱 镱 磬镳噔脲龛 镥疱漤屐 镱膻耋 狃桀磬.

描镱蝈玎 钺 囫桁戾蝠梓眍 疣耧铍铈屙梃 珙 耔眚彗 徨腙 镱 铗眍龛 钺豚耱 耧疱耨梃, 耵铕祗腓痤忄眄 磬 铖眍忄龛 祛溴腓痤忄龛, 猁豚 镱潋忮疰溴磬 耧屦桁屙蜞朦睇扈 戾蝾溧扈. 聋腩 镱赅玎眍, 黩 鲨觌 14 钺豚耱 疣耧疱溴脲龛 觐眦屙蝠圉栝 煨褪 Kr, kni gt 礤 耔祆弪痂黜 腩赅腓珙忄睇 铗眍耔蝈朦眍 钺豚耱彘 疣耧疱溴脲龛 觐眦屙蝠圉栝 耦铗忮蝰蜮簋 徨腙钼 皲忤眢螓 铗眍耔蝈朦眍 镱耠邃龛 镱 磬镳噔脲龛 镥疱漤屐 觐眦 狃桀磬. 令脲 蝾泐, 钺豚耱 疣耧疱溴脲龛 觐眦屙蝠圉栝 煨褪 桴 沐眍 蜞 驽, 赅 钺豚耱 疣耧疱溴脲龛 觐眦屙蝠圉栝 徨腙, 耢妁帼蝰 躅溴 鲨觌 14 镥疱漤屐 镱膻耋 狃桀磬.

绣龛 箴噔礤龛 祛溴腓 徨 滂趑箸梃 (c 眢脲恹扈 珥圜屙扈 镟疣戾蝠 Da), 耦铗忮蝰蜮簋 磬殇屙睇 铒蜩爨朦睇 珥圜屙 镟疣戾蝠钼, 忸耧痤桤忸滂 忸 怵屐屙 镳铖蝠囗耱忮 皲忤汨 钺豚耱彘 耧疱耨梃. 亦觇 钺疣珙, 滂趑箸 礤 铌噻噱 耋耱忮眄钽 怆龛 磬 皲忤汨 钺豚耱彘 耧疱耨梃 沐眍 gap.

碾 钺耥屙 戾踵龛珈钼 皲忤汔 钺豚耱彘 耧疱耨梃 Kr, kni gt 猁 镳钼邃屙 沭圄梓羼觇 囗嚯桤 疱泱鲨 耧疱耨梃 桴 沐眍 钺豚耱 皲忤汔 镳桦邈帼 钺豚耱.

侮噻嚯铖, 黩 鲥眚疣朦眍 钺豚耱 耧疱耨梃 Kr 玎漤桴 钺豚耱彘 耧疱耨梃 gt kni 皲忤 玎漤桴 沭囗桷 镳铊聃钿栩 疱珞朦蜞蝈 囫桁戾蝠梓睇 疱泱蝾痦 忡噼祛溴轳蜮栝 戾驿 Kr kni, kni gt, gt hb (痂. 9). 襄疱漤桢 沭囗桷 桴 钺豚耱彘 耧疱耨梃 蝾驽 溻桡帼蝰 怦脲漶蜮桢 耢妁屙 玎漤桴 沭囗桷 耦皴漤桴 钺豚耱彘 耧疱耨梃 沐眍 gap, 疣耧铍铈屙睇 犭桄 镥疱漤屐 镱膻耋 狃桀磬, 腓犷 疱珞朦蜞蝈 蝾泐, 黩 沭囗桷 钺豚耱彘 耧疱耨梃 gt hb 镥疱漤彘 鬣耱 狃桀磬 耱囗钼蝰 戾礤 镱腩汨扈. 亦觇 钺疣珙, 皲忤汨 镥疱漤桴 沭囗桷 钺豚耱彘 耧疱耨梃 祛骓 疣耨爨蝠桠囹 赅 怛铕梓眍 怆屙桢, 镳铊聃钿 桤-玎 滂磬扈麇耜钽 镱忮溴龛 玎漤桴 沭囗桷.

劐耱 汶噔 "杨玟囗桢 囹豚襦 耧疱耨梃 沐眍 皴沆屙蜞鲨 忸 怵屐屙 镳铖蝠囗耱忮" 镱疋磬 铒桉囗棹 耱痼牝箴 趔黻鲨铐嚯 忸珈铈眍耱彘 囹豚襦 耧疱耨梃 沐眍 皴沆屙蜞鲨 忸 怵屐屙 镳铖蝠囗耱忮, FlyEx (http://urchin.spbcas.ru;

http://flyex.ams.sunysb.edu/flyex).

锑蝈痂嚯 桤腩驽 磬 耱囹 A7, A12, A13, A14, A23, A26, A27, A32, A35, A39.

FlyEx - 疱鲨铐磬 徉玎 溧眄, 耦玟囗磬 磬 铖眍忮 延聊 IBM DB2, 镱朦珙忄蝈朦耜栝 桧蝈痿彘 觐蝾痤 疋玎 徉珙 溧眄 镱祛 WWW-皴疴屦, Apache Tomcat Java皴疴屦钼 镳桦铈屙栝. 如 镱耠邃龛, 磬栳铍栝 桧蝈疱 镳邃耱噔弪 皴疴屦 桤钺疣驽龛.

蒡铗 皴疴屦 疣琊噌铗囗 潆 钺疣犷蜿 桤钺疣驽龛 赅痱桧 耧疱耨梃 沐眍. 雾 镱玮铍弪 恹镱腠螯 蜞觇 铒屦圉梃 赅 钺滂礤龛 溻篚 桦 蝠艴 桤钺疣驽龛 皴痤 嚯 钿眍 鲡弪眍 祉铈羼蜮屙眍 桤钺疣驽龛, 沐礤疣鲨 疣珥铖蝽钽 桤钺疣驽龛 戾驿 溻箪 桤钺疣驽龛扈, 爨耜桊钼囗桢 钿眍泐 桤钺疣驽龛 漯筱桁, 钺羼镥麒忄弪 赅漯桊钼囗桢 桤钺疣驽龛, 翳朦蝠圉棹 桧蝈眈桠眍耱 綦箢疱聆屙鲨 篑桦屙桢 觐眚疣耱.

袜 皴疱滂眢 2007 . FlyEx 耦溴疰栩 桧纛痨圉棹 1580 狃桀磬 4490 赅痱桧圊 耧疱耨梃 14 沐眍 皴沆屙蜞鲨. 碾 赅驿钽 狃桀磬 躔囗栩 溴痦 爨耜, 怦 狃桀睇 铗眍蝰 鲨觌囔 溴脲龛 10 镱 14, 钺 麒耠 玎镨皴 潆 磬犷痤 觐腓麇耱忮眄 溧眄 耦耱噔弪 铌铍 9.5 祀., 潆 篑疱漤屙睇 溧眄 铌铍 270 螓. FlyEx 蜞赕 躔囗栩 觐翳鲨屙螓 疱汨耱疣鲨 眍痨嚯桤圉梃, 桉镱朦珞屐 潆 镱耱痤屙 擁 脲蝮 疱汨耱痂痤忄眄 溧眄 溧眄 箐嚯屙睇 纛眍恹 耔沩嚯铎. 碾 赅驿钽 戾蝾溧 疱汨耱疣鲨 赅驿钽 狃桀磬, 镳桧噤脲驵泐 怵屐屙睇 觌囫襦 28 鲨觌 溴脲龛 溴 14, 躔囗蝰 镱 2 觐翳鲨屙蜞 疱汨耱疣鲨. 项 6 觐翳鲨屙蝾 眍痨嚯桤圉梃 躔囗栩 潆 赅驿钽 沐磬 赅驿铎 狃桀礤. 软蝈沭桊钼囗睇 溧眄 镱耱痤屙 潆 赅驿钽 怵屐屙眍泐 觌囫襦, 眍 蝾朦觐 潆 蝈 沐眍, 觐蝾瘥 耜囗桊钼囗 犷脲 麇 9 狃桀磬.

袜镱腠屙桢 徉琨 溧眄 FlyEx 镳钿铍驵弪.

务眍忭 鲥朦 徉琨 溧眄 FlyEx 耦耱铊 铗忮蜞 磬 玎镳铖 镱朦珙忄蝈脲 滂磬扈赍 纛痨桊钼囗 钺豚耱彘 耧疱耨梃 沐眍 皴沆屙蜞鲨. 灶痨 玎镳铖钼 疣玟咫 "理嚯桤 溧眄"钺羼镥麒忄 漕耱箫 疣珉梓睇 铒屦圉 磬 桤钺疣驽龛扈 觐腓麇耱忮眄 溧眄 镱 c镳羼耔 沐眍. 蒡 铒屦圉梃 镱玮铍 镱朦珙忄蝈膻 镱塍鬣螯 桧纛痨圉棹 忡噼祉铎 疣耧铍铈屙梃 钺豚耱彘 耧疱耨梃 沐眍 皴沆屙蜞鲨 疣珉梓睇 怵屐屙睇 祛戾眚, 赅麇耱忮 溧眄 镱 耧疱耨梃 沐眍, 滂磬扈赍 纛痨桊钼囗 钺豚耱彘 耧疱耨梃, 耱屣屙 忄痂噌咫铖蜩 耧疱耨梃 沐磬 疣珥 篦囫蜿圊 蝈豚 狃桀磬 疣珥 祛戾眚 怵屐屙.

FlyEx - 铗牮 疱耋瘃, 痤觐 桉镱朦珞屐 扈痤恹 耦钺耱忸 徼铍钽钼 (耢. 疣玟咫 "橡嚓蜩麇耜铄 珥圜屙桢 疣犷螓"). 碾 钺脲泖屙 疣犷螓 溧眄, 躔囗桁 忸 FlyEx, 镳邃篑祛蝠屙 忸珈铈眍耱 恹沭箸觇 膻猁 钺祛 溧眄 镱 玎镳铖 镱朦珙忄蝈.

皴潼祛 汶噔 "青觌屙桢. 理嚯桤 疣礤 疣琊噌铗囗睇 眍恹 戾蝾漕 祛溴脲" 镳钼邃屙 耩噔龛蝈朦睇 囗嚯桤 疣礤 桉镱朦珙忄眄 疣琊噌铗囗睇 溧眄铋 疣犷蝈 戾蝾漕 镱塍麇龛 溧眄 镱 耧疱耨梃 沐眍 皴沆屙蜞鲨 爨蝈爨蜩麇耜桴 祛溴脲 溴蝈痨桧圉梃 皴沆屙蝾 镳铞羼襦 皴沆屙蜞鲨, 蜞赕 觐腓麇耱忮眄 溧眄 镱 耧疱耨梃 沐眍 皴沆屙蜞鲨 溧眄, 镱塍麇眄 漯筱桁 觐腚尻蜩忄扈 桉耠邃钼囹咫彘. 项溻邃屙 栩钽 滂耨屦蜞鲨铐眍 疣犷螓.

朽犷蝮 玎忮瘌帼 蔓忸潲.

3 蔓忸潲 1. 朽琊噌铗囗 铕桡桧嚯 觐礅彘屦睇 戾蝾 觐腓麇耱忮眄铋 铞屙觇 箴钼 耧疱耨梃 沐眍 皴沆屙蜞鲨 磬 铖眍忮 桤钺疣驽龛, 镱塍麇眄 镱祛 觐眙铌嚯钽 扈牮铖觐镟. 蒡铗 戾蝾 怅膻鬣弪 蜞觇 镳铞邃箴 赅 皴沆屙蜞鲨 桤钺疣驽龛, 箐嚯屙桢 纛眍忸泐 耔沩嚯, 铒疱溴脲龛 忸琊囫蜞 狃桀磬, 镳铖蝠囗耱忮眄簋 疱汨耱疣鲨 赅痱桧 耧疱耨梃 桧蝈沭圉棹 溧眄.

2. 项赅玎眍, 黩 忸琊囫 狃桀磬 鲨觌 溴脲龛 溴 14 祛驽 猁螯 镳邃耜噻囗 蝾黜铖螯 漕 2 扈眢 疣玮栩 磬 铖眍忄龛 囗嚯桤 恹耦觐 滂磬扈黜铋 赅痱桧 耧疱耨梃 沐磬 even-skipped.

3. 项塍麇 镱腠, 蝾黜 耔耱屐囹梓羼觇 磬犷 觐腓麇耱忮眄 溧眄 钺 耧疱耨梃 沐眍 皴沆屙蜞鲨 赅驿铎 漯 赅驿钽 桤 1580 桧滂忤潴嚯 狃桀眍, 蜞赕 桧蝈沭桊钼囗睇 溧眄 钺 耧疱耨梃 赅驿钽 桤 14 沐眍 皴沆屙蜞鲨. 软蝈沭桊钼囗睇 溧眄 桁妣 镳铖蝠囗耱忮眄铄 疣琊屮屙桢 钿眍 漯 疣琊屮屙桢 镱 怵屐屙 15 扈眢 疣玮栩 潆 鲨觌 13 铌铍 6.5 扈眢 疣玮栩 潆 鲨觌 14.

4. 项赅玎眍, 黩 钺豚耱 耧疱耨梃 沐眍 皴沆屙蜞鲨, 腩赅腓珙忄眄 狍潴 玎痤潲忸 镱腩耜, 镱 戾疱 疋铄泐 纛痨桊钼囗 鲨觌 14 耢妁帼蝰 镥疱漤屐 镱膻耋 狃桀磬. 唁忤汨 钺豚耱彘 耧疱耨梃 镱 镱潢 忮腓麒睇 耦怙噤帼 疣珈屦囔 镱腩 耧疱耨梃 沐眍 pair-rule. 鸯妁屙 溴 铎 疣轭礤 狃桀磬 珥圜栩咫 戾睃, 麇 皲忤汨 钺豚耱彘 耧疱耨梃.

5. 朽琊噌铗囗 爨蝈爨蜩麇耜 祛溴朦, 觐蝾疣 镳噔桦 忸耧痤桤忸滂 怵屐屙 滂磬簋 扈牦 耧疱耨梃 沐眍 皴沆屙蜞鲨, 疣珈屦 镥疱牮囗 钺豚耱彘 耧疱耨梃 耦皴漤桴 沐眍, 蜞赕 皲忤汨 沭囗桷 钺豚耱彘 耧疱耨梃 沐眍 gap 鲨觌 14.

6. 绣觐眈蝠箬痤忄眄 疣礻圊 祛溴腓 蝾镱沭圄 皴蜩 沐眍 gap 怅膻鬣弪 皴 螯 徉珙恹 疱泱蝾痦 戾踵龛珈钼: (1) 嚓蜩忄鲨 沐眍 gap 镱 溴轳蜮桢 Bcd Cad;

(2) 噔蝾嚓蜩忄鲨 沐眍 gap; (3) 耔朦眢 疱镳羼耔 镱 镳桧鲨矬 钺疣蝽铋 疋玷 戾驿 沐磬扈 gap 礤镥疱牮帼扈 钺豚耱扈 耧疱耨梃; (4) 囫桁戾蝠梓眢 疱镳羼耔 戾驿 沐磬扈 gap, 鼽 钺豚耱 耧疱耨梃 镥疱牮帼蝰; (5) 疱镳羼耔 镱 镳桧鲨矬 镳祛 疋玷 沐眍 gap 玎漤彘 鬣耱 狃桀磬 镱 溴轳蜮桢 蝈痨桧嚯钽 沐磬 tll. 蒡 蝾镱沭圄 怆弪 忭篁疱眄 礤镳铗桠铕鬻桠铋, 漕耱囹铟眍 潆 钺耥屙 磬犭噱禧 赅痱桧 耧疱耨梃 沐眍 gap , 鲥腩, 耦汶囫箦蝰 腓蝈疣蝮痦 溧眄.

7. 项潋忮疰溴磬 汨镱蝈玎 蝾, 黩 狍潴 玎痤潲忸 镱腩耜 钺豚耱 疣耧疱溴脲龛 觐眦屙蝠圉栝 煨褪 Kr, kni gt 腩赅腓珞 囫桁戾蝠梓眍 铗眍耔蝈朦眍 钺豚耱彘 疣耧疱溴脲龛 觐眦屙蝠圉栝 耦铗忮蝰蜮簋 徨腙钼 皲忤眢螓 铗眍耔蝈朦眍 镱耠邃龛 磬镳噔脲龛 镥疱漤屐 镱膻耋 狃桀磬.

8. 项赅玎眍, 黩 麒耠屙眍 祛溴腓痤忄龛 镱玮铍弪 耦镱耱噔栩 沭箫矬 沐眄 忡噼祛溴轳蜮栝 耦 耧弼梏梓睇 疱泱蝾痦 镳铞羼耦: 镳邃耜噻囗, 黩 皲忤汨 沭囗桷 耧疱耨梃 沐眍 gap 镳铊聃钿 桤-玎 囫桁戾蝠梓眍耱 疱泱蝾痦 忡噼祛溴轳蜮栝 戾驿 桁 沐磬扈, 镳 觐蝾瘥 沐, 钺豚耱 耧疱耨梃 觐蝾痤泐 疣耧铍铈屙 犭桄 玎漤屐 觐眦 狃桀磬, 疱镳羼耔痼弪 沐 镥疱牮帼轳 钺豚耱 耧疱耨梃, 眍 疣耧铍铈屙眍 犭桄 镥疱漤屐 觐眦. 蒡 皲忤汨 礤 玎忤 铗 溴轳蜮 爨蝈痂眈觇 沭噤桢眚钼 滂趑箸梃 镳钿箨蝾 沐眍 gap 戾驿 漯囔.

9. 杨玟囗 徉玎 溧眄 FlyEx (http://urchin.spbcas.ru; http://flyex.ams.sunysb.edu/flyex), 怆 镳铖蝠囗耱忮眄-怵屐屙睇 囹豚耦 耧疱耨梃 沐眍 皴沆屙蜞鲨. 拎玎 镳邃铖蜞怆弪 镱朦珙忄蝈膻 桧纛痨圉棹 滂磬扈赍 纛痨桊钼囗 钺豚耱彘 耧疱耨梃 沐眍 皴沆屙蜞鲨 镱玮铍弪 镳铊玮羼蜩 镱塍麇龛 膻猁 钺祛 溧眄 镱 玎镳铖.

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