用于植物改良的方法与流程

本发明涉及产生具有增加的根生物量的植物的方法。
背景技术：
：维管植物的根具有多种功能。这些功能包括：1)吸收水和无机养分，2)将植物固着至地面并支持植物，3)储存食物和养分，4)无性繁殖。在对养分的浓度的反应中，根也合成细胞分裂素，其充当影响芽生长快慢的信号。具有增加的根生物量的植物也潜在地具有多种优势，包括更好的固着、更有效的水摄取、更有效的养分摄取和提高的耐旱性。这些特征的组合也可以导致产量提高，包括谷物或果实生物量增加和/或叶生物量增加。目前，对控制植物中的根生物量的遗传机制的理解有限。因此，获得控制植物中的根生物量的可用替代方法是有利的。因此，本发明的目的是提供增加植物中的根生物量的产生的方法和材料，和/或至少为公众提供有用的选择。技术实现要素：之前，white(2006)在拟南芥属(arabidopsis)中发现了两个邻近的同源基因(命名为peapod，ppd1和ppd2)，其在叶和种荚(seedpod)发育的后期阶段调控拟分生组织的细胞增殖。在拟南芥属中这些基因的缺失导致叶肥大和种荚变宽，而ppd1的过表达导致叶和长角果的大小减小(white，2006)。本申请人现已令人惊讶地证明peapod基因在植物中的过表达导致根生物量产生的增加。因此，本申请人的发明涉及通过异位表达peapod以增加植物中的根生物量的方法。具体地，本发明涉及异位表达peapod蛋白，其特征在于存在对于由广泛的植物物种表现出的所有peapod蛋白均常见的共有氨基酸基序。方法第一方面，本发明提供了增加植物中根生物量的方法，该方法包括增加植物中至少一种peapod蛋白的表达的步骤。在一种实施方式中，根生物量相对于同一物种或品种的对照植物中的根生物量增加。在一种实施方式中，至少一种peapod蛋白的表达增加是用编码peapod蛋白的多核苷酸转化植物、或其祖先植物或植物细胞的结果。在其他实施方式中，所述植物为用编码和表达peapod蛋白的至少一种多核苷酸进行转基因的。在其他方面，本发明提供了产生具有增加的根生物量的植物的方法，所述方法包括增加所述植物中至少一种peapod蛋白的表达的步骤。在一种实施方式中，使用编码peapod蛋白的至少一种多核苷酸转化所述植物。在其他实施方式中，所述方法包括用编码peapod蛋白的多核苷酸转化植物，或转化再生为植物的植物细胞的步骤。在一种实施方式中，所述方法包括测试或评估所述植物的增加的根生物量的额外步骤。在其他实施方式中，所述方法包括通过无性或有性繁殖被测试其增加的生物量的植物以进一步产生具有增加的根生物量的植物的步骤。peapod蛋白在一种实施方式中，所述peapod蛋白是包含seqidno:28、29、31、32、34和35中的至少一种序列的多肽。在其他实施方式中，所述peapod蛋白包含seqidno:28的序列。在其他实施方式中，所述peapod蛋白包含seqidno:29的序列。在其他实施方式中，所述peapod蛋白包含seqidno:31的序列。在其他实施方式中，所述peapod蛋白包含seqidno:32的序列。在其他实施方式中，所述peapod蛋白包含seqidno:34的序列。在其他实施方式中，所述peapod蛋白包含seqidno:35的序列。在其他实施方式中，所述peapod蛋白是包含与seqidno:1-26中任意一项具有至少70％同一性的序列的多肽。在其他实施方式中，所述peapod蛋白是包含选自seqidno:1-26中任意一项序列的多肽。在其他实施方式中，所述peapod蛋白是包含与seqidno:1具有至少70％同一性的序列的多肽。在其他实施方式中，所述peapod蛋白是包含seqidno:1的序列的多肽。表达peapod在植物中表达蛋白的方法是本领域技术人员公知的，并在本文中进行描述。所有这些方法都包括在本发明的范围内。通过引入多核苷酸增加peapod的表达在一种实施方式中，通过将至少一种多核苷酸引入植物细胞或植物中以增加表达。在一个优选的实施方式中，所述多核苷酸编码如本文所定义的peapod蛋白。在其他实施方式中，所述多核苷酸包含与seqidno:83-107中任意一项的编码序列具有至少70％同一性的序列。在其他实施方式中，所述多核苷酸包含与seqidno:83-107中任意一项的序列具有至少70％同一性的序列。在其他实施方式中，所述多核苷酸包含seqidno:83-107中任意一项的编码序列。在其他实施方式中，所述多核苷酸包含seqidno:83-107中任意一项的序列。在其他实施方式中，所述多核苷酸包含以上所述的序列的片段，其能够编码与peapod蛋白具有相同功能的多肽。在一种实施方式中，所述片段编码能够增加根生物量的多肽。通过表达构建体表达peapod在一个优选的实施方式中，将所述多核苷酸作为表达构建体的一部分被引入植物。在一个优选的实施方式中，所述表达构建体包含与所述多核苷酸可操作地连接的启动子。用于增加peapod表达的启动子在一种实施方式中，所述启动子能够驱动或驱动可操作地连接的多核苷酸在植物的所有组织中组成型表达。在其他实施方式中，所述启动子是组织优选的启动子。在其他实施方式中，所述启动子能够驱动或驱动可操作地连接的多核苷酸在植物的地下组织中表达。在一种实施方式中，所述启动子是地下组织优选的启动子。在其他实施方式中，所述启动子是地下组织特异性启动子。在一种实施方式中，所述启动子是光阻遏型启动子(light-repressedpromoter)。在其他实施方式中，所述启动子能够驱动或驱动可操作地连接的多核苷酸在植物的根中表达。在一种实施方式中，所述启动子是根优选的启动子。在其他实施方式中，所述启动子是根特异性启动子。多核苷酸和多肽的来源本发明的或本发明方法中使用的多核苷酸和多核苷酸的变体可以衍生自任何物种。所述多核苷酸和变体也可以是合成产生的或重组产生的，并且也可以是“基因改组(geneshuffling)”方法的产物。本发明的或本发明方法中使用的多肽和多肽的变体可衍生自任何物种。所述多肽和变体也可以是重组产生的，并且还可以由“基因改组”方法的产物表达。在一种实施方式中，所述多核苷酸、多肽或变体衍生自植物物种。在其他实施方式中，所述多核苷酸、多肽或变体衍生自裸子植物(gymnospermplant)物种。在其他实施方式中，所述多核苷酸、多肽或变体衍生自被子植物(angiospermplant)物种。在其他实施方式中，所述多核苷酸、多肽或变体衍生自双子叶植物(dicotyledonous)物种。在一个优选的实施方式中，所述多核苷酸、多肽或变体衍生自真双子叶植物(eudicot)物种。在其他实施方式中，所述多核苷酸、多肽或变体衍生自真双子叶植物物种。在其他实施方式中，所述多核苷酸、多肽或变体衍生自单子叶植物物种。优选的单子叶植物包括：棕榈、香蕉、浮萍和兰花物种。待转化的植物细胞和植物本发明的，或在本发明的方法中使用的植物细胞和植物来自于任何植物物种。在一种实施方式中，所述植物细胞或植物来自于裸子植物物种。在其他实施方式中，所述植物细胞或植物来自于被子植物物种。在其他实施方式中，所述植物细胞或植物来自于双子叶植物物种。优选的单子叶植物属包括：冰草属(agropyron)、葱属(allium)、看麦娘属(alopecurus)、须芒草属(andropogon)、燕麦草属(arrhenatherum)、天门冬属(asparagus)、燕麦属(avena)、刺竹属(bambusa)、孔颖草属(bothrichloa)、格兰马草属(bouteloua)、雀麦属(bromus)、沙茅属(calamovilfa)、蒺藜草属(cenchrus)、虎尾草属(chloris)、香茅属(cymbopogon)、狗牙根属(cynodon)、鸭茅属(dactylis)、双花草属(dichanthium)、马唐属(digitaria)、椮属(eleusine)、画眉草属(eragrostis)、荞麦属(fagopyrum)、羊茅属(festuca)、向日葵属(helianthus)、大麦属(hordeum)、黑麦草属(lolium)、芒属(miscanthis)、芒属(miscanthusxgiganteus)、稻属(oryza)、黍属(panicum)、雀稗属(paspalum)、狼尾草属(pennisetum)、虉草属(phalaris)、梯牧草属(phleum)、早熟禾属(poa)、甘蔗属(saccharum)、黑麦属(secale)、狗尾草属(setaria)、印第安草属(sorgahastum)、高粱属(sorghum)、小麦属(triticum)、香荚兰属(vanilla)、小黑麦属(xtriticosecaletriticale)和玉蜀黍属(zea)。优选的单子叶植物物种包括：冰草(agropyroncristatum)、冰生冰草(agropyrondesertorum)、长穗偃麦草(agropyronelongatum)、中间偃麦草(agropyronintermedium)、蓝莲冰草(agropyronsmithii)、丛生小麦草(agropyronspicatum)、细莲冰草(agropyrontrachycaulum)、毛冰草(agropyrontrichophorum)、古月葱(alliumascalonicum)、洋葱(alliumcepa)、藠头(alliumchinense)、韭葱(alliumporrum)、北葱(alliumschoenoprasum)、大葱(alliumfistulosum)、大蒜(alliumsativum)、大看麦娘(alopecuruspratensis)、须芒草(andropogongerardi)、大须芒草(andropogongerardii)、小须芒草(andropogonscoparious)、燕麦草(arrhenatherumelatius)、石刁柏(asparagusofficinalis)、裸燕麦(avenanuda)、燕麦(avenasativa)、大佛肚竹(bambusavulgaris)、藤茎须芒草(bothrichloabarbinodis)、白羊草(bothrichloaischaemum)、针草(bothrichloasaccharoides)、垂穗草(boutelouacuripendula)、黑格兰马草(boutelouaeriopoda)、格兰马草(boutelouagracilis)、直立雀麦(bromuserectus)、无芒雀麦(bromusinermis)、沙拂子茅(calamovilfalongifilia)、纤毛蒺藜草(cenchrusciliaris)、非洲虎尾草(chlorisgayana)、香茅(cymbopogonnardus)、狗牙根(cynodondactylon)、鸭茅(dactylisglomerata)、双花草(dichanthiumannulatum)、毛梗双花草(dichanthiumaristatum)、昆士兰蓝草(dichanthiumsericeum)、俯仰马唐(digitariadecumbens)、南非马唐(digitariasmutsii)、圆形薯蓣(dioscorearotundata)、参薯(dicsoreaalata)、麻山药(dicscoreaopposita)、黄药(dicscoreabulbifera)、甘薯(dioscoreaesculenta)、三浅裂薯蓣(dioscoreatrifida)、穇子(eleusinecoracan)、窄颖新麦草(elymusangustus)、俄罗斯新麦草(elymusjunceus)、弯叶画眉草(eragrostiscurvula)、埃塞俄比亚画眉草(eragrostistef)、荞麦(fagopyrumesculentum)、苦荞麦(fagopyrumtataricum)、苇状羊茅(festucaarundinacea)、羊茅(festucaovina)、草甸羊茅(festucapratensis)、紫羊茅(festucarubra)、向日葵(helianthusannuussunflower)、二列大麦(hordeumdistichum)、大麦(hordeumvulgare)、多花黑麦草(loliummultiflorum)、多年生黑麦草(loliumperenne)、中国银色草(miscanthissinensis)、奇岗(miscanthusxgiganteus)、水稻(oryzasativa)、秫米(panicumitalicium)、大黍(panicummaximum)、稷(panicummiliaceum)、紫黍草(panicumpurpurascens)、柳枝稷(panicumvirgatum)、柳枝稷(panicumvirgatum)、毛花雀稗(paspalumdilatatum)、百喜草(paspalumnotatum)、狼尾草(pennisetumclandestinum)、珍珠狼尾草(pennisetumglaucum)、象草(pennisetumpurpureum)、御谷(pennisetumspicatum)、虉草(phalarisarundinacea)、梯牧草(phleumbertolinii)、猫尾草(phleumpratense)、羊肉早熟禾(poafendleriana)、草地早熟禾(poapratensis)、林地早熟禾(poanemoralis)、甘蔗(saccharumofficinarum)、大茎野生蔗(saccharumrobustum)、竹蔗(saccharumsinense)、割手密(saccharumspontaneum)、黑麦(secalecereale)、非洲狗尾草(setariasphacelata)、印第安草(sorghastrumnutans)、印第安草(sorghastrumnutans)、甜高粱(sorghumdochna)、假高粱(sorghumhalepense)、苏丹草(sorghumsudanense)、高粱(sorghumvulgare)、高粱(sorghumvulgare)、小麦(triticumaestivum)、二粒小麦(triticumdicoccum)、硬粒小麦(triticumdurum)、一粒小麦(triticummonococcum)、香荚兰(vanillafragrans)、小黑麦(xtriticosecale)和玉米(zeamays)。单子叶植物的优选科系是禾本科。优选的禾本科亚科包括：柊叶竺亚科(anomochlooideae)、服叶竺亚科(pharoideae)、姜叶竺亚科(puelioideae)、竹亚科(bambusoideae)、早熟禾亚科(pooideae)、稻亚科(ehrhartoideae)、三芒草亚科(aristidoideae)、芦竹亚科(arundinoideae)、虎尾草亚科(chloridoideae)、黍亚科(panicoideae)、扁芒草亚科(danthonioideae)和百生草亚科(micrairoideae)。优选的禾本科亚科为早熟禾亚科。优选的早熟禾亚科植物包括小麦、大麦、燕麦、雀麦草和芦苇草。另一个优选的禾本科亚科为稻亚科。优选的稻亚科植物包括水稻。另一个优选的禾本科亚科为黍亚科。优选的黍亚科植物包括稗草、玉米、高粱、甘蔗(sugarcane)、能源甘蔗(energycane)、小米、福尼奥米(fonio)和须芒草。另一个优选的禾本科亚科为芦竹亚科。优选的芦竹亚科植物包括芦竹(arundodonax)。另一个优选的禾本科亚科为竹亚科。优选的竹亚科植物包括竹子。优选的禾本科物种包括来自黑麦草属(lolium)的那些物种。优选的黑麦草属物种包括长花黑麦草(loliumlongiflorum)、多花黑麦草(loliummultiflorum)、多年生黑麦草、西方黑麦草(loliumwesterwoldicum)、毒麦(loliumtemulentum)和杂种黑麦草(loliumhybridum)。其他优选的禾本科物种包括来自羊茅属(festuca)的那些物种。优选的羊茅属物种包括高羊茅(festucaarundinacea)、硬羊茅(festucaovina)、草甸羊茅(festucapratensis)和紫羊茅(festucarubra)。优选地，所述植物细胞或植物来自双子叶植物物种。优选的双子叶植物属包括：桃属(amygdalus)、槚如树属(anacardium)、花生属(arachis)、芸苔属(brassica)、木豆属(cajanus)、大麻属(cannabis)、红花属(carthamus)、山核桃属(carya)、吉贝属(ceiba)、鹰嘴豆属(cicer)、椰子属(cocos)、芫荽属(coriandrum)、小冠花属(coronilla)、棉属(cossypium)、猪屎豆属(crotalaria)、扁豆属(dolichos)、油棕属(elaeis)、大豆属(lycine)、棉花属(gossypium)、向日葵属(helianthus)、香豌豆属(lathyrus)、兵豆属(lens)、胡枝子属(lespedeza)、亚麻属(linum)、莲属(lotus)、羽扇豆属(lupinus)、澳洲坚果属(macadamia)、苜蓿属(medicago)、草木犀属(melilotus)、黎豆属(mucuna)、木犀榄属(olea)、驴食草属(onobrychis)、料豆属(ornithopus)、罂粟属(papaver)、菜豆属(phaseolus)、刺葵属(phoenix)、黄连木属(pistacia)、豌豆属(pisum)、樱桃属(prunus)、葛属(pueraria)、茶藨子属(ribes)、蓖麻属(ricinus)、胡麻属(sesamum)、可可(theobroma)、三叶草属(trifolium)、胡芦巴属(trigonella)、蚕豆属(vicia)和豇豆属(vigna)。优选的双子叶植物物种包括：：扁桃(amygdaluscommunis)、腰果(anacardiumoccidentale)、花生(arachishypogaea)、花生(arachishypogea)、油菜(brassicanapusrape)、黑芥(brassicanigra)、白菜(brassicacampestris)、木豆(cajanuscajan)、木豆(cajanusindicus)、大麻(cannabissativa)、红花(carthamustinctorius)、美国山核桃(caryaillinoinensis)、吉贝(ceibapentandra)、鹰嘴豆(cicerarietinum)、可可椰子(cocosnucifera)、芫荽(coriandrumsativum)、绣球小冠花(coronillavaria)、陆地棉(cossypiumhirsutum)、菽麻(crotalariajuncea)、扁豆(dolichoslablab)、油棕(elaeisguineensis)、亚洲棉(gossypiumarboreum)、中棉(gossypiumnanking)、海岛棉(gossypiumbarbadense)、草棉(gossypiumherbaceum)、陆地棉(gossypiumhirsutum)、大豆(glycinemax)、野大豆(glycineussuriensis)、宽叶蔓豆(glycinegracilis)、向日葵(helianthusannus)、jatrophacuneata、麻疯树(jatrophacurcas)、窄叶羽扇豆(lupinusangustifolius)、黄羽扇豆(lupinusluteus)、珍珠羽扇豆(lupinusmutabilis)、铁扫帚(lespedezasericea)、鸡眼草(lespedezastriata)、大百脉根(lotusuliginosus)、山黧豆(lathyrussativus)、兵豆(lensculinaris)、长萼妁眼草(lespedezastipulacea)、亚麻(linumusitatissimum)、百脉根(lotuscorniculatus)、白羽扇豆(lupinusalbus)、窄叶羽扇豆(lupinusangustifolius)、黄羽扇豆(lupinusluteus)、木本苜蓿(medicagoarborea)、黄花苜蓿(medicagofalcate)、金花菜(medicagohispida)、黄香草木犀(medicagoofficinalis)、紫花苜蓿(medicagosativa(alfalfa))、硬果苜蓿(medicagotribuloides)、澳洲坚果(macadamiaintegrifolia)、褐斑苜蓿(medicagoarabica)、白花草木樨(melilotusalbus)、刺毛黎豆(mucunapruriens)、油橄榄(oleaeuropaea)、红豆草(onobrychisviciifolia)、锯齿草(ornithopussativus)、绿豆(phaseolusaureus)、红叶李(prunuscerasifera)、欧洲酸樱桃(prunuscerasus)、荷包豆(phaseoluscoccineus)、欧洲李(prunusdomestica)、棉豆(phaseoluslunatus)、马哈利樱桃(prunu.maheleb)、绿豆(phaseolusmungo)、桃(prunus.persica)、樱桃(prunus.pseudocerasus)、菜豆(phaseolusvulgaris)、罂粟(papaversomniferum)、宽叶菜豆(phaseolusacutifolius)、海枣(phoenixdactylifera)、阿月浑子(pistaciavera)、豌豆(pisumsativum)、甜扁桃(prunusamygdalus)、杏(prunusarmeniaca)、野葛(puerariathunbergiana)、黑茶镳子(ribesnigrum)、红茶藨子(ribesrubrum)、鹅莓(ribesgrossularia)、蓖麻(ricinuscommunis)、芝麻(sesamumindicum)、马铃薯(solanumtuberosum)、狭叶三叶草(trifoliumaugustifolium)、分散三叶草(trifoliumdiffusum)、杂种车轴草(trifoliumhybridum)、降车轴草(trifoliumincarnatum)、球花车轴草(trifoliumingrescens)、红车轴草(trifoliumpratense)、白三叶草(trifoliumrepens)、波斯三叶草(trifoliumresupinatum)、地三叶草(trifoliumsubterraneum)、可可(theobromacacao)、埃及车轴草(trifoliumalexandrinum)、胡芦巴(trigonellafoenumgraecum)、油桐(verniciafordii)、窄叶野豌豆(viciaangustifolia)、深紫花野豌豆(viciaatropurpurea)、距花豌豆(viciacalcarata)、广布野豌豆(viciadasycarpa)、苦野豌豆(viciaervilia)、红莓苔子(vacciniumoxycoccos)、褐毛野豌豆(viciapannonica)、长豇豆(vignasesquipedalis)、豇豆(vignasinensis)、长柔毛野豌豆(viciavillosa)、蚕豆(viciafaba)、野碗豆(viciasative)和赤豆(vignaangularis)。植物和植物部分在其他方面，本发明提供了由于具有增加的peapod蛋白或其片段的表达而具有增加的根生物量的植物。在一种实施方式中，所述peapod蛋白或其片段的表达增加是用编码peapod蛋白或其片段的多核苷酸转化植物、或其祖先植物或植物细胞的结果。在其他实施方式中，所述植物为用表达peapod蛋白或其片段的多核苷酸进行转基因的。在其他实施方式中，所述多核苷酸或其片段是与组织优选的启动子可操作地连接的多核苷酸。在其他实施方式中，所述启动子是根优选的启动子。在其他实施方式中，所述启动子是根特异性启动子。在其他实施方式中，所述peapod蛋白如本文所定义。在其他实施方式中，所述编码peapod蛋白的多核苷酸如本文所定义。在其他方面，本发明提供了植物的细胞、部分、繁殖体或子代，其为用以下中的至少一种进行转基因的：a)多核苷酸，和b)多核苷酸和可操作地连接的启动子。具体实施方式在本说明书中，提及专利说明书、其他外部文献或其他信息来源时，通常是为了提供用于讨论本发明特征的上下文。除非另有具体说明，对这些外部文献的参考并不被解释为承认这些文献或这些信息来源以任何权限作为现有技术，或形成本领域的公知常识的一部分。本说明书中使用的术语“包含(comprising)”意指“至少部分由…组成”。当解释本说明书中的每个包括术语“包含”的语句时，除了以该术语开始的特征以外的特征也可能存在。相关的术语如“包含(comprise)”将以相同的方式解释。增加的根生物量具有“增加的根生物量”的植物比同一类型和年龄的对照植物产生更多的根生物量。因此，“增加的”意指相对于同一类型和年龄的对照植物的增加。优选地，具有“增加的根生物量”的植物比同一类型和年龄的对照植物产生的根生物量多至少10％、优选至少20％、更优选至少30％、更优选至少40％、更优选至少50％、更优选至少60％、更优选至少70％、更优选至少80％、更优选至少90％、更优选至少100％、更优选至少150％、更优选至少200％、更优选至少300％、更优选至少400％。在一种实施方式中，具有“增加的根生物量”的植物相比于对照植物具有以下中的至少一种：更大的根、更多的根或更延伸的根系。根生物量术语根生物量是指由植物产生的根组织的总质量。这可以由干重或湿重来评估。根本文使用的术语根意指与该术语的标准用法相同。术语根包含初生根、次生根、不定根、根分枝和根毛。根通常在地下，但是该术语也包含气生根。在一种实施方式中，术语根包含植物的非含叶、非含节部分。增加的耐旱性在一种实施方式中，具有“增加的根生物量”的植物也具有增加的耐旱性。同样地，“增加的”意指相对于同一类型和年龄的对照植物的增加。术语“增加的耐旱性”旨在描述在次优的水化条件下其在生长和发育的任一方面都比在相同条件下的适宜的对照植物表现更好的植物。对照植物在一种实施方式中，所述对照植物是野生型植物。在其他实施方式中，所述对照植物是未转化的植物。在其他实施方式中，所述对照植物是尚未用peapod多核苷酸转化的植物。在其他实施方式中，所述对照植物是尚未用构建体转化的植物。在其他实施方式中，所述对照植物是用对照构建体转化的植物。在一种实施方式中，所述构建体是空载体构建体。异位表达术语“异位表达”旨在被广义地解释。该术语是指多核苷酸或多肽在以下的任意一项中的表达：·在其中所述多核苷酸或多肽未被正常表达的细胞、器官、组织或植物，·在所述多核苷酸或多肽未被正常表达时的时间或发育阶段的细胞、器官、组织或植物，和·在其中所述多核苷酸或多肽以比其正常表达时更高的水平表达的细胞、器官、组织或植物。组织优选的启动子在某些实施方式中，所述编码多核苷酸的peapod蛋白在组织优选的启动子的控制下表达。关于组织优选的启动子的术语“优选的”意指启动子首先在该组织中驱动表达。因此，例如，根优选的启动子在根组织中比在植物的其他组织或器官中驱动更高水平的可操作地连接的多核苷酸的表达。根优选的启动子根优选的启动子在根组织中比在植物的其他组织或器官中驱动更高水平的可操作地连接的多核苷酸的表达。根优选的启动子可以包括非光合组织优选的启动子和光阻遏调节型启动子。非光合组织优选的启动子非光合组织优选的启动子包括优选在植物的非光合组织/器官中表达的启动子。非光合组织优选的启动子也可包括光阻遏型启动子。光阻遏型启动子光阻遏型启动子的实例可见于us5,639,952和us5,656,496。根特异性启动子根特异性启动子的实例可见于us5,837,848和us2004/0067506以及us2001/0047525。关于优选在某一组织中表达的启动子的术语“优选表达”意指在该组织中比在植物的其他组织中以更高水平表达的启动子。关于启动子的术语“组织特异性的”意指所述启动子基本上仅在该组织而非植物的其他组织中表达。在一种实施方式中，所述根优选的启动子是根特异性启动子。本文使用的术语“基因”意指包括编码多肽或蛋白的编码序列的内源基因组序列。所述编码序列被一个或多个内含子间隔。基因通常还包括启动子序列、5’非翻译序列、3’非翻译序列和终止子序列。调节蛋白表达的基因组序列也可被认为是基因的一部分。多核苷酸和片段本文使用的术语“多核苷酸”意指任意长度但优选至少15个核苷酸的单链或双链脱氧核糖核苷酸或核糖核苷酸聚合物，并且包括以下非限制性实例：基因的编码和非编码序列、正义和反义序列互补物、外显子、内含子、基因组dna、cdna、前mrna、mrna、rrna、sirna、mirna、trna、核酶、重组多肽、分离的和纯化的天然存在的dna或rna序列、合成的rna和dna序列、核酸探针、引物和片段。多核苷酸的“片段”是指较长的多核苷酸序列的连续子序列。优选地，所述片段为所公开的多核苷酸的至少15个核苷酸、优选至少16个核苷酸、更优选至少17个核苷酸、更优选至少18个核苷酸、更优选至少19个核苷酸、更优选至少20个核苷酸、更优选至少21个核苷酸、更优选至少22个核苷酸、更优选至少23个核苷酸、更优选至少24个核苷酸、更优选至少25个核苷酸、更优选至少26个核苷酸、更优选至少27个核苷酸、更优选至少28个核苷酸、更优选至少29个核苷酸、更优选至少30个核苷酸、更优选至少31个核苷酸、更优选至少32个核苷酸、更优选至少33个核苷酸、更优选至少34个核苷酸、更优选至少35个核苷酸、更优选至少36个核苷酸、更优选至少37个核苷酸、更优选至少38个核苷酸、更优选至少39个核苷酸、更优选至少40个核苷酸、更优选至少41个核苷酸、更优选至少42个核苷酸、更优选至少43个核苷酸、更优选至少44个核苷酸、更优选至少45个核苷酸、更优选至少46个核苷酸、更优选至少47个核苷酸、更优选至少48个核苷酸、更优选至少49个核苷酸、更优选至少50个核苷酸、更优选至少51个核苷酸、更优选至少52个核苷酸、更优选至少53个核苷酸、更优选至少54个核苷酸、更优选至少55个核苷酸、更优选至少56个核苷酸、更优选至少57个核苷酸、更优选至少58个核苷酸、更优选至少59个核苷酸、更优选至少60个核苷酸、更优选至少61个核苷酸、更优选至少62个核苷酸、更优选至少63个核苷酸、更优选至少64个核苷酸、更优选至少65个核苷酸、更优选至少66个核苷酸、更优选至少67个核苷酸、更优选至少68个核苷酸、更优选至少69个核苷酸、更优选至少70个核苷酸、更优选至少71个核苷酸、更优选至少72个核苷酸、更优选至少73个核苷酸、更优选至少74个核苷酸、更优选至少75个核苷酸、更优选至少76个核苷酸、更优选至少77个核苷酸、更优选至少78个核苷酸、更优选至少79个核苷酸、更优选至少80个核苷酸、更优选至少81个核苷酸、更优选至少82个核苷酸、更优选至少83个核苷酸、更优选至少84个核苷酸、更优选至少85个核苷酸、更优选至少86个核苷酸、更优选至少87个核苷酸、更优选至少88个核苷酸、更优选至少89个核苷酸、更优选至少90个核苷酸、更优选至少91个核苷酸、更优选至少92个核苷酸、更优选至少93个核苷酸、更优选至少94个核苷酸、更优选至少95个核苷酸、更优选至少96个核苷酸、更优选至少97个核苷酸、更优选至少98个核苷酸、更优选至少99个核苷酸、更优选至少100个核苷酸、更优选至少150个核苷酸、更优选至少200个核苷酸、更优选至少250个核苷酸、更优选至少300个核苷酸、更优选至少350个核苷酸、更优选至少400个核苷酸和最优选至少500个核苷酸的连续核苷酸。多核苷酸序列的片段可用于反义rna干扰(rnai)、基因沉默、三螺旋或核酶技术中，或作为包括在微阵列中的引物、探针，或用于本发明的基于多核苷酸的可选方法中。在一种实施方式中，所述片段编码执行或能够执行与由该片段是其一部分的更长的多核苷酸编码的多肽相同的功能的多肽。术语“引物”是指通常具有游离的3’oh基团的短多核苷酸，其与或能够与模板杂交，并用于引发与靶标互补的多核苷酸的聚合。术语“探针”是指在基于杂交的测定中，用于或可用于检测与探针互补的多核苷酸序列的短多核苷酸。所述探针可由如本文所定义的多核苷酸的“片段”组成。多肽和片段本文使用的术语“多肽”包含任意长度但优选至少5个氨基酸的氨基酸链，包括其中氨基酸残基通过共价肽键连接的全长蛋白。本发明的或用于本发明方法中的多肽可以是纯化的天然产物，或者可以部分或全部使用重组或合成技术产生。该术语可是指多肽、多肽的聚合体，例如，二聚体或其他多聚体、融合多肽、多肽片段、多肽变体或其衍生物。多核苷酸的“片段”是指较长的多核苷酸序列的连续子序列。优选所述片段多肽的“片段”是指较长的多肽的连续子序列。优选所述片段为至少5个、更优选至少10个、更优选至少20个、更优选至少30个、更优选至少40个、更优选至少50个、更优选至少100个、更优选至少120个、更优选至少150个、更优选至少200个、更优选至少250个、更优选至少300个、更优选至少350个、更优选至少400个。在一种实施方式中，所述片段执行或能够执行与该片段是其一部分的多肽相同的功能。优选地，所述片段执行生物活性所必需的功能和/或提供多肽的三维结构。应用于本文公开的多核苷酸或多肽序列的术语“分离的”用于指从其天然细胞环境中去除的序列。在一种实施方式中，所述序列与其天然存在的侧翼序列相分离。分离的分子可以通过包括生化、重组和合成技术的任意方法或方法的组合来获得。术语“重组体(recombinant)”是指经合成产生的或从在其天然环境中围绕它的序列中去除的多核苷酸序列。所述重组序列可以与其天然环境中不存在的序列重新组合。“重组体”多肽序列通过从“重组体”多核苷酸序列的翻译而产生。关于衍生自特定属或物种的本发明的多核苷酸或多肽的术语“衍生自”意指所述多核苷酸或多肽与天然存在于该属或物种中的多核苷酸或多肽具有相同的序列。衍生自特定属或物种的多核苷酸或多肽因此可以合成产生或重组产生。变体本文使用的术语“变体”是指不同于特定鉴定的序列的多核苷酸或多肽序列，其中一个或多个核苷酸或氨基酸残基被缺失、置换或添加。变体可以是天然存在的等位基因变体或非天然存在的变体。变体可以来自同一物种或来自其他物种并且可以包含同源物(homologue)、旁系同源物(paralogue)和直系同源物(orthologue)。在某些实施方式中，本文公开的多肽和多核苷酸的变体具有与所公开的多肽或多核苷酸的生物活性相同或相似的生物活性。关于多肽和多核苷酸的术语“变体”包含如本文所定义的多肽和多核苷酸的所有形式。多核苷酸变体变体核苷酸序列优选显示出与本发明的序列至少50％、更优选至少51％、更优选至少52％、更优选至少53％、更优选至少54％、更优选至少55％、更优选至少56％、更优选至少57％、更优选至少58％、更优选至少59％、更优选至少60％、更优选至少61％、更优选至少62％、更优选至少63％、更优选至少64％、更优选至少65％、更优选至少66％、更优选至少67％、更优选至少68％、更优选至少69％、更优选至少70％、更优选至少71％、更优选至少72％、更优选至少73％、更优选至少74％、更优选至少75％、更优选至少76％、更优选至少77％、更优选至少78％、更优选至少79％、更优选至少80％、更优选至少81％、更优选至少82％、更优选至少83％、更优选至少84％、更优选至少85％、更优选至少86％、更优选至少87％、更优选至少88％、更优选至少89％、更优选至少90％,更优选至少91％、更优选至少92％、更优选至少93％、更优选至少94％、更优选至少95％、更优选至少96％、更优选至少97％、更优选至少98％和最优选至少99％的同一性。同一性通过至少20个核苷酸位置、优选至少50个核苷酸位置、更优选至少100个核苷酸位置以及最优选本发明的全长多核苷酸的比较窗口而发现。多核苷酸序列同一性可以通过以下方式测定。使用bl2seq中的blastn(来自blast程序套件，2.2.5版本[2002年11月])将所述目标多核苷酸序列与候选多核苷酸序列进行比较(tatianaa.tatusova,thomasl.madden(1999),"blast2sequences-anewtoolforcomparingproteinandnucleotidesequences",femsmicrobiollett.174:247-250)，该程序可从ncbi公开获得(ftp:jjftp.ncbi.nih.govjblastj)。在一种实施方式中，使用bl2seq的默认参数。在其他实施方式中，使用bl2seq的默认参数，除了应该关闭对低复杂度部分的过滤以外。也可以使用全局序列比对程序(例如，needleman,s.b.andwunsch,c.d.(1970)j.mol.biol.48,443-453)在候选多核苷酸序列和目标多核苷酸序列之间的整个重叠长度上计算多核苷酸序列同一性。needleman-wunsch全局比对算法的完整实施方式可见于emboss程序包中的needle程序(rice,p.longden,l.andbleasby,a.emboss:theeuropeanmolecularbiologyopensoftwaresuite,trendsingeneticsjune2000,第16卷,第6期，第276-277页)，其可从http://www.hgmp.mrc.ac.uk/software/emboss/获得。欧洲生物信息学研究所服务器也提供了在线上(在http:/www.ebi.ac.uk/emboss/align/上)进行两条序列之间的emboss-needle全局比对的工具。或者，可以使用gap程序，其计算两条序列的最佳全局比对，而不会对末端空位进行罚分。gap记载于以下文章中：huang,x.(1994)onglobalsequencealignment.computerapplicationsinthebiosciences10,227-235。计算多核苷酸的序列同一性％的优选方法基于使用clustalx比对待比较的序列(jeanmouginetal.,1998,trendsbiochem.sci.23,403-5)。本发明的多核苷酸变体还包含表现出与一个或多个特定鉴定的序列具有相似性的那些，其可能保留这些序列的功能等效性并且无法被合理预期已随机发生。所述关于多肽的序列相似性可以使用可公开获得的ncbi的blast程序套件的bl2seq程序(2.2.5版本[2002，11月])(ftp://ftp.ncbi.nih.gov/blast/)来确定。或者，本发明的变体多核苷酸在严格条件下与特定的多核苷酸序列或其互补物杂交。术语“在严格条件下杂交”及其语法上的同义语是指多核苷酸分子与靶多核苷酸分子(如固定在dna或rna印迹(如southern印迹或northern印迹)上的靶多核苷酸分子)在确定的温度和盐浓度的条件下杂交的能力。在严格杂交条件下杂交的能力可以通过在较不严格条件下初始杂交然后增加严格性至预期严格性的条件下杂交来测定。对于长度大于约100个碱基的多核苷酸分子，典型的严格杂交条件是不超过25至30℃(例如，10℃)，低于天然双链的解链温度(tm)(通常参见，sambrooketal.,eds,1987,molecularcloning,alaboratorymanual,第2版，coldspringharborpress；ausubeletal.,1987,currentprotocolsinmolecularbiology,greenepublishing)。大于约100个碱基的多核苷酸分子的tm可以通过公式tm＝81.5+0.41％(g+c-log(na+)来计算(sambrooketal.,eds,1987,molecularcloning,alaboratorymanual,第2版，coldspringharborpress；boltonandmccarthy,1962,pnas84:1390)。长度大于约100个碱基的多核苷酸的典型严格条件为杂交条件，如在6xssc、0.2％sds的溶液中预洗涤；在65℃、6xssc、0.2％sds中过夜杂交；然后在65℃下在1xssc、0.1％sds中洗涤两次，每次30分钟，以及在65℃下在0.2xssc、0.1％sds中洗涤两次，每次30分钟。对于长度小于100个碱基的多核苷酸分子，示例性的严格杂交条件为低于tm的5至10℃。平均而言，长度小于100bp的多核苷酸分子的tm降低约(500/寡核苷酸长度)℃。对于已知作为肽核酸(pna)的dna模拟物(nielsenetal.,science.1991dec6；254(5037):1497-500)，tm值比dna-dna杂交体或dna-rna杂交体的tm值高，并且可以使用giesenetal.,nucleicacidsres.1998nov1；26(21):5004-6中记载的公式计算。具有长度小于100个碱基的dna-pna杂交体的示例性严格杂交条件为低于tm的5至10℃。本发明的变体多核苷酸还包含这样的多核苷酸，其不同于本发明的序列，但是因遗传密码的简并性而编码与由本发明的多核苷酸编码的多肽具有相似活性的多肽。不改变多肽的氨基酸序列的序列改变是“沉默变异”。除了atg(甲硫氨酸)和tgg(色氨酸)，相同氨基酸的其他密码子可以通过本领域公知的技术改变，例如，在特定的宿主生物体中以优化密码子的表达。本发明也包括这样的多核苷酸序列改变，其导致所编码的多肽序列的一个或几个氨基酸的保守置换，而不明显改变其生物活性。本领域技术人员将会知晓制备表型沉默氨基酸置换的方法(参见，例如，bowieetal.,1990,science247,1306)。由于所编码的多肽序列中的沉默变异和保守置换而导致的变体多核苷酸可以通过如前所述的tblastx算法使用可公开获得的ncbi的blast程序套件的bl2seq程序(2.2.5版本[2002，11月])(ftp://ftp.ncbi.nih.gov/blast/)来测定。多肽变体关于多肽的术语“变体”包含天然存在的、重组产生的和合成产生的多肽。变体多肽序列优选表现出与本发明的序列至少50％、更优选至少51％、更优选至少52％、更优选至少53％、更优选至少54％、更优选至少55％、更优选至少56％、更优选至少57％、更优选至少58％、更优选至少59％、更优选至少60％、更优选至少61％、更优选至少62％、更优选至少63％、更优选至少64％、更优选至少65％、更优选至少66％、更优选至少67％、更优选至少68％、更优选至少69％、更优选至少70％、更优选至少71％、更优选至少72％、更优选至少73％、更优选至少74％、更优选至少75％、更优选至少76％、更优选至少77％、更优选至少78％、更优选至少79％、更优选至少80％、更优选至少81％、更优选至少82％、更优选至少83％、更优选至少84％、更优选至少85％、更优选至少86％、更优选至少87％、更优选至少88％、更优选至少89％、更优选至少90％,更优选至少91％、更优选至少92％、更优选至少93％、更优选至少94％、更优选至少95％、更优选至少96％、更优选至少97％、更优选至少98％和最优选至少99％的同一性。同一性通过至少20个氨基酸位置、优选至少50个氨基酸位置、更优选至少100个氨基酸位置以及最优选本发明的全长多肽的比较窗口而发现。多肽序列同一性可以通过以下方式确定。使用bl2seq中的blastn(来自blast程序套件，2.2.5版本[2002年11月]，其可从ncbi公开获得(ftp://ftp.ncbi.nih.gov/blast/))将所述目标多肽序列与候选多肽序列进行比较。在一种实施方式中，使用bl2seq的默认参数。在其他实施方式中，使用bl2seq的默认参数，除了应关闭对低复杂度部分的过滤。也可以使用全局序列比对程序在候选多肽序列和目标多肽序列之间的整个重叠长度上计算多肽序列同一性。如上所讨论的emboss-needle(可从http:/www.ebi.ac.uk/emboss/align/上获得)和gap(huang,x.(1994)onglobalsequencealignment.computerapplicationsinthebiosciences10,227-235)也是用于计算多肽序列同一性的合适的全局序列比对程序。计算多肽的序列同一性％的优选方法基于使用clustalx比对待比较的序列(jeanmouginetal.,1998,trendsbiochem.sci.23,403-5)。变体多肽包括这样的多肽，其中氨基酸序列与本文的多肽的区别在于一个或多个不影响肽的生物活性的保守氨基酸置换、缺失、添加或插入。保守置换通常包括一个氨基酸置换为具有相似特征的另一个，例如，以下组中的置换：缬氨酸、甘氨酸；甘氨酸、丙氨酸；缬氨酸、异亮氨酸、亮氨酸；天冬氨酸、谷氨酸；天冬酰胺、谷氨酰胺；丝氨酸、苏氨酸；赖氨酸、精氨酸；和苯丙氨酸、酪氨酸。非保守置换需要将这些类别之一的一个成员换成另一类别的成员。对进化的生物序列的分析已证明并不是所有的序列改变都具有同样可能性，反映了保守置换和非保守置换在生物水平上的至少部分差异。例如，某些氨基酸置换可以频繁发生，而其他则非常少见。氨基酸残基的进化性改变或置换可以通过评分矩阵(也称为置换矩阵)来模拟。将所述矩阵用于生物信息学分析中以鉴定序列之间的关系，一个实例为以下所示的blosum62矩阵(表1)。表1：含有所有可能的置换分数的blosum62矩阵[henikoffandhenikoff,1992]使用所示的blosum62矩阵生成相应的行和列的交汇点处存在的各比对的氨基酸对的分数。例如，从谷氨酸残基(e)至天冬氨酸残基(d)的置换分数为2。对角线示出了还未改变的氨基酸的分数。大部分置换改变具有负分数。所述矩阵只包含整数。确定合适的评分矩阵以产生对给定序列组最佳的比对被认为是在本领域技术的范围内。表1中的blosum62矩阵也可以用作blast检索中的缺省矩阵，尽管并不局限于此。其他变体包括具有影响肽稳定性的修饰的肽。这种类似物可以包含，例如，肽序列中的一个或多个非-肽键(其替换肽键)。也包括这样的类似物，其包括非天然存在的l-氨基酸的残基，例如，d-氨基酸或非天然存在的合成氨基酸，例如，β或γ氨基酸和环状类似物。构建体、载体和其组件术语“遗传构建体”是指多核苷酸分子，通常为双链dna，其可能已经被插入至另一个多核苷酸分子(插入多核苷酸分子)(例如，但不限于cdna分子)中。遗传构建体可以包含必要的元件，其允许转录插入多核苷酸分子，并且，任选地，将转录物翻译为多肽。所述插入多核苷酸分子可衍生自宿主细胞，或可衍生自不同的细胞或生物体和/或可以是重组多核苷酸。一旦在宿主细胞内，所述遗传构建体可以整合在宿主染色体dna中。所述遗传构建体可与载体连接。术语“载体”是指多核苷酸分子，通常为双链dna，其被用于将所述遗传构建体转运到宿主细胞中。所述载体能够在至少一种其他宿主系统(如大肠杆菌)中复制。术语“表达构建体”是指包括必要元件的遗传构建体，所述元件允许转录所述插入多核苷酸分子，并且，任选地，将转录物翻译为多肽的必要的元件。表达构建体通常在5’至3’方向上包含：a)在所述构建体将转化至其中的宿主细胞中具有功能的启动子；b)待表达的多核苷酸；c)在所述构建体将转化至其中的宿主细胞中具有功能的终止子。术语“编码区域”或“开放阅读框”(orf)是指能够在适当的调控序列的控制下产生转录产物和/或多肽的基因组dna序列的正义链或cdna序列。所述编码序列通过5’翻译起始密码子和3’翻译终止密码子的存在而鉴定。当插入至遗传构建体中时，“编码序列”在其与启动子序列和终止子序列可操作地连接时能够被表达。“可操作地连接”意指待表达的序列被置于调控元件的控制下，所述调控元件包括启动子、组织特异性调控元件、时间调控元件、增强子、抑制子和终止子。术语“非编码区域”是指非翻译序列，其为翻译起始位点的上游和翻译终止密码子的下游。这些序列也分别称为5’utr和3’utr。这些区域包括转录起始和终止以及调控翻译效率所必需的元件。终止子是终止转录的序列，并存在于所翻译序列下游的基因的3’非翻译末端。终止子是mrna稳定性的重要决定因素，并且在一些情况下被发现具有空间调控功能。术语“启动子”是指编码区域上游的调控基因转录的非转录顺式调控元件。启动子包含指定转录起始位点和保守框(如tata框)以及被转录因子结合的基序的顺式起始元件。启动子可以与待表达的多核苷酸是同源的。这意指所述启动子和多核苷酸天然以可操作地连接的形式存在。或者，所述启动子可以与待表达的多核苷酸是异源的。这意指所述启动子和多核苷酸天然不以可操作地连接的形式存在。“转基因”是通过转化被引入生物体中的多聚核苷酸。所述转基因可以衍生自相同物种或与所述转基因被引入至其中的生物体的物种不同的物种。所述转基因也可以是合成的并且天然不存在于任何物种中。“转基因植物”是指含有作为遗传操作或转化结果的新遗传物质的植物。所述新遗传物质可衍生自与所得转基因植物的物种相同或不同的植物，或可是合成的。优选地，所述“转基因植物”因转基因的存在而与天然存在的任何植物不同。“反向重复”是重复的序列，其中，重复序列的后半段是互补链，例如：(5’)gatcta.......tagatc(3’)(3’)ctagat.......atctag(5’)只要在重复区域间具有3-5个碱基对的间隔区，通读转录将产生经过互补碱基配对来形成发卡结构的转录物。所述间隔区可以是通常长度至少为3个碱基对的任意多核苷酸序列。宿主细胞宿主细胞可以衍生自，例如，细菌、真菌、昆虫、哺乳动物或植物生物体。分离或产生多核苷酸的方法本发明的多核苷酸分子可以使用本领域普通技术人员已知的多种技术分离。例如，可以通过使用mullisetal.,eds.1994thepolymerasechainreaction,birkhauser(以引用的方式纳入本文)中记载的聚合酶链式反应(pcr)分离所述多肽。本发明的多肽可以使用如本文所定义的衍生自本发明的多核苷酸序列的引物来扩增。分离本发明的多核苷酸的其他方法包括使用具有如本文所示的杂交探针的序列的多肽的全部或部分。将标记的多核苷酸探针与固定在固体支持物(如硝化纤维素滤膜或尼龙膜)上的多核苷酸杂交的技术可以被用于筛选基因组文库或cdna文库。示例性的杂交和洗涤条件为：在65℃下在5.0xssc、0.5％十二烷基硫酸钠、1xdenhardt溶液中杂交20小时；在1.0xssc,1％(w/v)十二烷基硫酸钠中洗涤(在55℃下洗涤3次，每次20分钟)，以及任选地，在60℃下在0.5xssc、1％(w/v)十二烷基硫酸钠中洗涤1次(20分钟)。在60℃、0.1xssc、1％(w/v)十二烷基硫酸钠的条件下进行任选的进一步洗涤(20分钟)。本发明的多核苷酸片段可以通过本领域公知的技术(例如，限制性内切酶消化、寡核苷酸合成和pcr扩增)产生。部分多核苷酸序列可以在本领域公知的方法中使用，以鉴定相应的全长多核苷酸序列。这种方法包括基于pcr的方法，5’race(frohmanma,1993,methodsenzymol.218:340-56)和基于杂交的方法、基于计算机/数据库的方法。此外，例如，使用以基于已知区域的引物为起始的反向pcr允许捕获位于本文公开的多核苷酸序列侧翼的未知序列(trigliaetal.,1998,nucleicacidsres16,8186,以引用的方式纳入本文)。该方法使用若干个限制性酶来以在基因的已知区域中产生合适的片段。所述片段然后通过分子内连接进行环化并用作pcr模板。发散引物(divergentprimer)从已知区域设计而来。为了完全组装全长的克隆，可使用标准分子生物学方法(sambrooketal.,molecularcloning:alaboratorymanual,第2版coldspringharborpress,1987)。当从特定物种产生转基因植物时，使用衍生自该物种的序列转化这样的植物可能是有益的。益处可能是改善公众对在产生转基因生物中的物种间转化的关注。此外，当基因的下调是期望的结果时，利用与所述植物中的序列相同(或至少是高度相似的)的序列可能是必需的，对该基因而言，降低的表达是期望的。因为这些以及其他原因，能够鉴定和分离在几个不同植物物种中鉴定和分离出特定基因的直系同源物是可期望的。变体(包括直系同源物)可以通过所描述的方法进行鉴定。用于鉴定变体的方法物理方法变体多肽可以通过使用基于pcr的方法进行鉴定(mullisetal.,eds.1994thepolymerasechainreaction,birkhauser)。通常，可用于通过pcr扩增本发明的多聚核苷酸分子变体的引物的多聚核苷酸序列可以基于编码相应氨基酸序列的保守区域的序列。或者，可以使用本领域技术人员所公知的文库筛选方法(sambrooketal.,molecularcloning:alaboratorymanual,第2版，coldspringharborpress,1987)。当鉴定所述探针序列的变体时，相对于当寻找精确的序列匹配时，其杂交和/或洗涤的严格性通常将被降低。多肽变体也可以通过物理方法进行鉴定，例如，通过使用抗本发明的多肽的抗体筛选表达文库(sambrooketal.,molecularcloning:alaboratorymanual,第2版,coldspringharborpress,1987)或在这样的抗体的帮助下将多肽从天然来源中鉴定出。基于计算机的方法包括多核苷酸和多肽变体的本发明的变体序列也可以通过本领域技术人员所公知的基于计算机的方法使用公开的域序列比对算法和序列相似性搜索工具来搜索序列数据库(公开的域数据库包括genbank、embl、swiss-prot、pir等)而进行鉴定。参见，例如，nucleicacidsres.29:1-10and11-16,2001中在线资源的实例。相似性搜索检索和比对用于与待分析序列相比较的靶序列(即查询序列)。序列比较算法使用评分矩阵来为每一条比对序列赋予总分数。用于鉴定序列数据库中变体的示例性程序族是包括blastn、blastp、blastx、tblastn和tblastx的blast程序套件(2.2.5版本[2002年11月])，它们可以从(ftp://ftp.ncbi.nih.gov/blast/)或从美国马里兰州贝塞斯达市(邮编20894)38a栋8n805房间的国家医学图书馆国家生物技术信息中心(ncbi)公开获得。ncbi服务器也提供了使用这些程序来筛选大量可公开获得的序列数据库的工具。blastn比较核苷酸查询序列和核苷酸序列数据库。blastp比较氨基酸查询序列和蛋白序列数据库。blastx比较在所有阅读框中被翻译的核苷酸查询序列和蛋白序列数据库。tblastn比较蛋白查询序列和在所有阅读框中动态翻译的核苷酸序列数据库。tblastx比较核苷酸查询序列的六读码框翻译和核苷酸序列数据库的六读码框翻译。blast程序可以以默认参数使用或者可以根据需要改变参数以改进筛选。包括blastn、blastp和blastx的blast算法族的使用在出版物altschuletal.,nucleicacidsres.25:3389-3402,1997中进行了描述。由blastn、blastp、blastx、tblastn、tblastx或相似算法产生的被查询序列产生的对于一个或多个数据库序列的“击中序列数”(hit)比对和鉴定了序列的相似部分。击中序列数以相似性的程度和序列重叠的长度的顺序排列。对数据库序列的击中序列数通常代表了仅是被查询序列的序列长度的部分的重叠。blastn、blastp、blastx、tblastn和tblastx算法也针对比对产生“期望”值。期望值(e)表明当搜索含有随机连续序列的相同大小的数据库时，人们能“期望”随机看到的击中的数目。期望值用作决定对于数据库的击中是否表明真的相似性的显著性阈值。例如，指定到多核苷酸击中数的e值为0.1被解释为意味着，在被筛选的数据库大小的数据库中，人们可能期望看到具有简单随机地相似分数的序列的被比对部分的0.1匹配。对于被比对和匹配部分具有0.01或更低的e值的序列，通过使用所述blastn、blastp、blastx、tblastn或tblastx算法在那个数据库中发现随机匹配的可能性是1％或更低。一组相关序列的多序列比对可以使用clustalw(thompson,j.d.,higgins,d.g.andgibson,t.j.(1994)clustalw:improvingthesensitivityofprogressivemultiplesequencealignmentthroughsequenceweighting,positions-specificgappenaltiesandweightmatrixchoice.nucleicacidsresearch,22:4673-4680,http://www-igbmc.u-strasbg.fr/bioinfo/clustalw/top.html)或t-coffee(cedricnotredame,desmondg.higgins,jaapheringa,t-coffee:anovelmethodforfastandaccuratemultiplesequencealignment,j.mol.biol.(2000)302:205-217))或pileup进行，这些算法使用渐进配对比对。(fenganddoolittle,1987,j.mol.evol.25,351)。模式识别软件应用可用于发现基序或标记序列。例如，meme(用于基序挖掘的多重em)在一组序列中发现基序标记序列，和mast(基序比对和搜索工具)使用这些基序来鉴定查询序列中的相似或相同基序。mast结果以具有适当统计数据和发现的基序的可视化概况的系列比对的形式提供。meme和mast在圣地亚哥的加利福尼亚大学开发。prosite(bairochandbucher,1994,nucleicacidsres.22,3583；hofmannetal.,1999,nucleicacidsres.27,215)是鉴定从基因组或cdna序列翻译而来的未表征的蛋白的功能的方法。prosite数据库(www.expasy.org/prosite)含有生物学上重要的模式和配置文件并被设计以使其能与适当的计算机工具一起使用，以指定新的序列至已知的蛋白家族或测定哪些已知的结构域存在于所述序列中(falquetetal.,2002,nucleicacidsres.30,235)。prosearch是能够使用给定序列模式或标记搜索swiss-prot和embl数据库的工具。分离多肽的方法包括变体多肽的本发明的，或用于本发明的方法中使用的多肽可以通过使用本领域公知的肽合成方法进行制备，例如使用固相合成技术的定向肽合成(例如stewartetal.,1969,insolid-phasepeptidesynthesis,whfreemanco,加利福尼亚圣弗朗西斯科)或自动化合成，例如使用应用appliedbiosystems431a肽合成仪(加利福尼亚，福斯特市)。所述多肽的突变形式也可以在该合成过程中产生。本发明的或在本发明方法中使用的多肽和变体多肽也可以通过使用本领域公知的各种技术从天然来源中纯化而来(例如，deutscher,1990,ed,methodsinenzymology，vol.182,guidetoproteinpurification)。或者，本发明的或在本发明的方法中使用的多肽和变体多肽可能在合适的宿主细胞中重组表达并按照如下所述从细胞中分离而来。产生构建体和载体的方法本发明的遗传构建体包含一条或多条本发明的多核苷酸序列和/或编码本发明的多肽的多核苷酸，并可用于转化例如细菌、真菌、昆虫、哺乳动物或植物生物体。本发明的遗传构建体旨在包括如本文所定义的表达构建体。产生和使用遗传构建体和载体的方法在本领域中是公知的并通常记载于sambrooketal.,molecularcloning:alaboratorymanual，第2版，coldspringharborpress,1987；ausubeletal.,currentprotocolsinmolecularbiology，greenepublishing,1987)中。产生包含多核苷酸、构建体或载体的宿主细胞的方法本发明提供了包含本发明的遗传构建体或载体的宿主细胞。包含本发明的遗传构建体(如表达构建体)的宿主细胞可以用于本领域公知的方法中(例如，sambrooketal.,molecularcloning:alaboratorymanual,第2版，coldspringharborpress,1987；ausubeletal.,currentprotocolsinmolecularbiology，greenepublishing,1987)以用于重组产生本发明的多肽。该方法可以涉及在适当的培养基中在适于或有助于本发明的多肽的表达的条件下培养宿主细胞。可任选地分泌至培养物中的经表达的重组多肽然后可以被从培养基、宿主细胞或培养物培养基中通过本领域公知的方法分离(例如，deutscher,ed,1990,methodsinenzymology，第182卷,guidetoproteinpurification)。产生包含构建体和载体的植物细胞和植物的方法本发明还提供了包含本发明的遗传构建体的植物细胞，以及被修饰以改变本发明的多核苷酸或多肽的表达的植物细胞。包含该细胞的植物也形成了本发明的一个方面。用多肽转化植物细胞、植物和其部分的方法在下述文献中进行了描述：draperetal.,1988,plantgenetictransformationandgeneexpression.alaboratorymanualblackwellsci.pub.oxford,p.365；potrykusandspangenburg,1995,genetransfertoplants.springer-verlag,berlin.；和gelvinetal.,1993,plantmolecularbiol.manual.kluweracad.pub.dordrecht。包括转化技术的转基因植物的综述在galunandbreiman,1997,transgenicplants.imperialcollegepress,london中提供。植物遗传操纵的方法大量植物转化策略是可获得的(例如，birch,1997,annrevplantphysplantmolbiol,48,297，hellensrp,etal(2000)plantmolbiol42:819-32,hellensretalplantmeth1:13)。例如，在多核苷酸/多肽被正常表达时，可以设计策略以增加多核苷酸/多肽在植物细胞、器官中和/或在特定发育阶段的表达，或在多核苷酸/多肽不能正常被表达时，设计策略以在细胞、组织、器官中和/或在特定发育阶段异位表达多核苷酸/多肽。所表达的多核苷酸/多肽可以衍生自待转化的植物物种或可衍生自不同的植物物种。在转基因植物中用于表达基因的遗传构建体通常包括驱动一个或多个克隆多核苷酸表达的启动子、终止子和用于检测遗传构建体在被转化的植物中存在的选择性标记序列。适用于遗传构建体的启动子在单子叶植物或双子叶植物的细胞、组织或器官中具有功能并包括细胞、组织和器官特异性启动子，细胞循环特异性启动子，时间启动子，诱导型启动子，在大多数植物组织中有活性的组成型启动子和重组启动子。启动子的选择将依赖于所期望的克隆的多核苷酸在时间和空间上的表达。所述启动子可能是与目的转基因正常相关的启动子，或衍生自其他植物、病毒和植物病原性细菌和真菌的基因的启动子。无需过多的实验，本领域技术人员将能够通过使用包含本发明的多核苷酸序列的遗传构建体选择适用于修饰和调控植物特性的启动子。组成型植物启动子的实例包括camv35s启动子、胆脂碱合成酶启动子和章鱼碱合成酶启动子和来自于玉米的ubi1启动子。在特定组织中有活性并对内部发育信号或外部非生物胁迫或生物胁迫产生应答的植物启动子记载于科学文献中。示例性的启动子记载于例如wo02/00894中，其以引用的方式纳入本文。在植物转化遗传构建体中普遍使用的示例性的终止子包括，例如，花椰菜花叶病毒(camv)35s终止子、根癌农杆菌(agrobacteriumtumefaciens)胭脂碱合成酶或章鱼肉碱合成酶终止子、玉米(zeamays)醇溶蛋白基因终止子、水稻(oryzasativa)adp葡糖焦磷酸化酶终止子和马铃薯pi-ii终止子。在植物转化中普遍使用的选择性标志物包括赋予卡那霉素抗性的新霉素磷酸转移酶ii基因(nptii)、赋予壮观霉素和链霉素抗性的aada基因、赋予ignite(agrevo)和basta(hoechst)抗性的膦丝菌素乙酰基转移酶(bar基因)和赋予潮霉素抗性的潮霉素磷酸转移酶基因(hpt)。包含可能用于植物和植物组织中启动子表达分析的报告基因(表达对于宿主而言是外来的、通常是酶活性和/或可视信号(例如荧光素酶、gus、gfp)的活性的编码序列)的遗传构建体的使用也是被考虑的。报告基因的文献在herrera-estrellaetal.,1993,nature303,209，和schrott,1995,in:genetransfertoplants(potrykus,t.,spangenberg.eds)springerverlag.berline,pp.325-336中进行了综述。基因沉默在多核苷酸/多肽被正常表达时，可以设计转化策略以降低多核苷酸/多肽在植物细胞、组织、器官中和/或在特定发育阶段的表达。基因沉默策略可能会关注基因本身或影响所编码的多肽的表达的调控元件。“调控元件”在本文中以其最广泛可能的意义使用并包括与目的基因相互作用的其他基因。被设计用于降低或沉默本发明的多核苷酸/多肽的表达的遗传构建体可以包括本发明多核苷酸的反义拷贝。在这样的构建体中，所述多核苷酸被以相对于启动子和终止子的反义方向进行放置。“反义”多核苷酸通过将多核苷酸或多核苷酸或片段反向获得，以使得所产生的转录物将与该基因的mrna转录物互补，例如，5’gatcta3’(编码链)3’ctagat5’(反义链)3’cuagau5’mrna5’gaucucg3’反义rna被设计用于基因沉默的遗传构建体也可以包括反向重复序列。“反向重复序列”是重复的序列，其中，重复序列的后半段是互补链，例如，5’-gatcta………tagatc-3’3’-ctagat………atctag-5’所形成的转录物可能经历互补碱基配对以形成发卡结构。通常在重复区域之间的至少3-5个碱基对的间隔区是允许发卡形成所必需的。包括这种反向重复序列的构建体可用于rna干扰(rnai)中，并因此可称为rnai构建体。另一种沉默方法涉及靶向于mirna的转录等价物的小反义rna的使用(llaveetal.,2002,science297,2053)。这样的对应于本发明的多核苷酸的小反义rna的使用是特别考虑的。本文所用的术语遗传构建体也包括小反义rna和其他影响基因沉默的这样的多肽。如本文所定义的，用表达构建体进行的转化也可以通过被称为有义抑制的过程产生基因沉默(例如，napolietal.,1990,plantcell2,279；decarvalhoniebeletal.,1995,plantcell,7,347)。在一些情况下，有义抑制可涉及全部或部分编码序列的过表达但也可涉及所述基因的非编码区(如内含子或5’或3’非翻译区(utr))的表达。嵌合的部分正义构建体能用于协调沉默多个基因(abbottetal.,2002,plantphysiol.128(3):844-53；jonesetal.,1998,planta204:499-505)。这样的用于沉默本发明的多核苷酸的表达的有义抑制的使用也是被考虑的。被设计用于基因沉默的遗传构建体中多核苷酸插入物可能对应于相应基因的编码序列和/或非编码序列(如启动子和/或内含子和/或5’或3’utr序列)。其他基因沉默策略包括显性负向方法和核酶构建体的使用(mcintyre,1996,transgenicres,5,257)。转录前沉默可能通过基因本身或其调控元件的突变而实现。这样的突变可以包括点突变、移码、插入、缺失和置换。转化方案以下是公开的能够被用于遗传转化下述植物物种的遗传转化方案的代表性出版物：水稻(alametal.,1999,plantcellrep.18,572)、苹果(yaoetal.,1995,plantcellreports14,407-412)、玉米(美国专利序列no.5,177,010和5,981,840)、小麦(ortizetal.,1996,plantcellrep.15,1996,877)、番茄(美国专利序列no.5,159,135)、马铃薯(kumaretal.,1996plantj.9,:821)、木薯(lietal.,1996nat.biotechnology14,736)、莴苣(michelmoreetal.,1987,plantcellrep.6,439)、烟草(horschetal.,1985,science227,1229)、棉花(美国专利序列no.5,846,797和5,004,863)、草(美国专利no.5,187,073和6,020,539)、薄荷(niuetal.,1998,plantcellrep.17,165)、柑橘植物(penaetal.,1995,plantsci.104,183)；葛缕子(krenetal.,1997,plantcellrep,17,39)、香蕉(美国专利序列no.5,792,935)、大豆(美国专利no.5,416,011、5,569,834、5,824,877、5,563,04455和5,968,830)、菠萝(美国专利序列no.5,952,543)、白杨(美国专利no.4,795,855)、普通单子叶植物(美国专利no.5,591,616和6,037,522)、芸苔(美国专利no.5,188,958、5,463,174和5,750,871)、谷物(美国专利no.6,074,877)、梨(matsudaetal.,2005,plantcellrep.24(1):45-51)、李子(rameshetal.,2006、song和sink2005、gonzalezpadillaetal.,2003)、草莓(oosumietal.,2006、oosumietal.,2006planta.223(6):1219-30；foltaetal.,2006plantaapr14；pmid:16614818foltaetal.,2006)、玫瑰(lietal.,2003)、悬钩子(grahametal.,1995methodsmolbiol.1995；44:129-33)、番茄(danetal.,2006,plantcellreportsv25:432-441)、苹果(yaoetal.,1995,plantcellrep.14,407-412)和毛花猕猴桃(actinidiaeriantha)(wangetal.,2006,plantcellrep.25,5:425-31)、欧洲桦(silverbirch)(keinonen-mettalaetal.,1998,plantcellrep.17:356-361.)和山杨(nilssono,etal.,1992,transgenicresearch.1:209-220)。其他物种的转化也被本发明所考虑。合适的方法和方案在科学文献中可以获得。本领域已知的多个其他方法可能被用于改变本发明的核苷酸和/或多肽的活性的表达。这样的方法包括但不限于tilling(tilletal.,2003,methodsmolbiol,2％,205)、所谓的“deletagene”技术(lietal.,2001,plantjournal27(3),235)和人工转录因子(如合成的锌指转录因子)的使用(例如，jouvenotetal.,2003,genetherapy10,513)。此外，靶向于特定多肽的抗体或其片段也可以在植物中表达以调控该多肽的活性(joblingetal.,2003,nat.biotechnol,21(1),35)。转座子标签方法也可以被使用。此外，与本发明的多肽相互作用的肽可以通过如相位展示的技术(dyaxcorporation)进行鉴定。这样的相互作用的多肽可以在植物中表达或应用到植物以影响本发明的多肽的活性。上述改变本发明的核苷酸和/或多肽的表达的每一种方法的使用都是被特别考虑的。术语“改变本发明的或在本发明的方法中使用的多核苷酸或多肽的表达”或“改变的本发明的或在本发明的方法中使用的多核苷酸或多肽的表达”旨在包括对应于本发明的多核苷酸的基因组dna被修饰从而导致本发明的多核苷酸或多肽的改变的表达的情况。所述基因组dna的修饰可能是通过遗传转化或其他的本领域已知的诱导突变的方法而进行。所述“改变的表达”可能与产生的信使rna和/或多肽的量的增加或减少有关，并可能由于产生的多核苷酸和多肽的序列的改变而导致多肽的活性改变。选择植物的方法还提供了选择具有增加的根生物量的植物的方法。该方法包括测试植物中是否存在至少一种peapod多核苷酸或多肽的表达改变，包括本文所定义的或所公开的那些。当增加的根生物量特征不必然易于测量时，该方法可以在幼年或早期发育阶段使用。多核苷酸(如信使rna)的表达经常被用作相应多肽表达的指标。测量多核苷酸的表达的示例性的方法包括但不限于northern分析、rt-pcr和斑点印记(dot-blot)分析(sambrooketal.,molecularcloning:alaboratorymanual,第2版，coldspringharborpress,1987)。因此，如本文所定义的，本发明的多核苷酸或其部分在鉴定具有增加的根生物量的植物的方法中可用作探针或引物。本发明或本文公开的多肽可以在被设计以鉴定这样的植物的杂交实验中用作探针或基于pcr的实验中用作引物。或者，可以获得抗本文所述或所公开的peapod多肽的抗体。获得和使用抗体的方法是本领域中的标准方法(参见例如：antibodies,alaboratorymanual,harlowalane,eds,coldspringharbourlaboratory，1998)。这样的抗体可以被用在检测所述多肽的表达改变的方法中。这样的方法可以包括elisa(kemeny,1991,apracticalguidetoelisa,nypergamonpress)和western分析(towbin&gordon,1994,jimmunolmethods,72,313)。这些分析多核苷酸或多肽表达和选择具有增加的根生物量的植物的方法在被设计以产生具有所述改变的特征的品种的传统育种程序中是有用的。植物术语“植物”旨在包括完整的植物或植物的任意部分、植物的繁殖体和子代。术语“繁殖体”意指可用在有性或无性生殖或繁殖中的植物的任意部分，包括种子和插穗。本发明的植物可能被种植，并且进行自交或与不同的植物植株进行杂交且所产生的具有期望的表型特征的杂交体可以被鉴定。两代或更多代植物可以被种植以确保目标表型特征被稳定维持和遗传。从这样的标准育种方法产生的植物也形成本发明的一个方面。通过赤霉素(ga)、油菜素内酯(br)和其他植物激素控制植物生长和发育赤霉素(ga)和油菜素内酯(br)是两类植物激素，它们参与植物形态发生和生长的很多方面，包括：种子萌发、细胞伸长、维管组织发育、种子大小、叶直立、开花、叶和果实衰老(mathewetal2009,nzjar52,213-225；houetal2010,developmentalcell19,884-894；jiangandlin2013,plantsignalingandbehaviour8:10,e25928)。考虑到ga和br在植物发育中的作用，调控ga和br的水平或它们下游靶标的能力在提高很多植物物种的产量和质量方面是高度期望的。实际上，有一些这两种激素中的一种的外源施用被用于提高农艺价值的商业实例。ga可被应用于黑麦草以刺激过季生长以及促进开花(mathewetal2009,nzjar52,213-225)，其也可以被用于抵消较低温度对甘蔗(热带c4草)的副作用。ga也可以用于增大无籽葡萄和樱桃的果实大小，以促进苹果和梨的结实并延迟特别是柑橘属作物的果皮老化(sun2011,currentbiology21,r338-r345)。类似地，建议将br制品用于提高马铃薯、番茄、黄瓜、辣椒和大麦、水稻、玉米、小麦、棉花和烟草的作物产量和质量(prusakovaetal1999,agrarianrussia,41-44；khripachetal2000annalsofbotany86,441-447；anjumetal2011j.agronomycropsci.197,177-185；vardhini2012j.phytology4,1-3)。但是，商业上应用的油菜素内酯的低采用率可以反映出成本和在类固醇被外源施用时植物不能对其进行有效吸收的事实。此外，需要严格控制时机和外源施用的ga和br的浓度限制了它们的应用。在大多数情况下，被子植物中的ga和br生物合成和分解代谢途径已经被表征并包括负调控因子和下游转录因子靶标。ga或br一旦结合至其各自的受体，复杂的信号通路便接续发生，在这两种情况中，调控的中心点均涉及改变负调控因子della(在ga的情况下)和转录调控因子(在br的情况下)的泛素-蛋白酶体通路。della蛋白的去除导致对生长抑制的去除和对ga应答的生长和发育的促进。相反地，对br的检测导致未被磷酸化的bzr1蛋白在细胞核中的累积。bzr1的去磷酸化阻止了其被蛋白酶体降解并反而使得bzr1能与其他dna结合转录因子结合并与转录辅因子相互作用。这导致对参与生长和其他细胞过程的成千上万个基因的调控，包括抑制br生物合成基因的表达(heetal2005,science307,1634-1638；guoetal2013,currentopinionplantbiol.16,545-553)。存在很多外源信号和环境提示，它们影响ga-gid1-della调控模块，在该模块中della通过与多种来自其他通路的关键调控蛋白的直接蛋白-蛋白相互作用整合了不同的信号传导活性。因此，della蛋白是主要的生长抑制子，其通过整合来自其他激素通路的内部信号(植物生长素、脱落酸、茉莉酸和乙烯)和外部生物刺激(病原体)和非生物刺激(光照条件、冷和盐胁迫)来控制植物生长和发育(sun2011,currentbiology21,r338-r345)。干旱是限制植物生长和农业生产力的最重要的环境限制因素之一。不出所料地，在提高的耐旱性和更延伸的根系之间存在正相关性，所述更延伸的根系包括更深的根和更多的侧根，它们都能够进行土壤勘探和地下资源获取(yuetal2008,plantcell20,1134-1151；werneretal2010,plantcell22,3905-3920)。因此，由此可见，共同的农业目标是对根系结构的优化以有助于克服由水或养分缺乏造成的作物植物的产量限制。但是，在所有缩减作物产量的非生物胁迫中，干旱是最具破坏性的一种并且是对抗育种者工作的最顽固胁迫。因为性状受很多基因控制，传统的育种方法是困难的并且难以评估(werneretal2010,plantcell22,3905-3920)。当标记辅助选择(mas)、数量性状基因座(qtl)和其他基因组学方法被广泛用于辅助育种工作以产生抵抗干旱的栽培种时(tuberosaandsalvi,2006,trendsinplantscience,11:405-412)，所述系统受限于筛选群体中存在的变异。有趣地是，水稻仅具有一个della蛋白(slr1)，玉米具有两个(d8和d9)(lawitetal2010,plantcellphysiol51,1854-1868)，而拟南芥属具有5个(ga1、rga、rgl1、rgl2和rgl3)(achardandgenschik2009,j.exp.bot.60,1085-1092)。此外，在最近的系统进化分析中，chenetal2013发现了它们所分析的六个禾本科物种有5个仅具有单个della，而18个双子叶植物物种中有14个具有两个或更多个della蛋白。相比之下，水稻中存在bzr家族中的6个成员，玉米中存在10个成员(www.grassius.org)以及拟南芥属中存在6个成员(wangetal2002,developmentalcell2,505-513)。植物的生长和发育依赖于提供其固着生活习性所需的高发育可塑性的信号传导通路之间的多种连接(gallego-bartolomeetal2012,pnas109,13446-13451)。因此，目前的证据，而不是以绝缘形式存在的各激素信号传导通路表明不同的通路之间存在高程度的相互作用，以及给定的激素通常调控由其余激素引起的输出。例如，近来已证明ga和br信号传导通路之间的相互干预包括della和bzr1/bes1之间的直接相互作用，由此della蛋白不仅影响蛋白稳定性而且抑制bzr1的转录活性(liandhe2013,plantsignalingandbehaviour8:7,e24686及其中的文献)。因此，由ga促进细胞伸长部分是通过去除della介导的对bzr1的抑制。近来已证明植物生长和发育可以通过直接操控主生长调节子della(lawit,kundu,raoandtomes,2007,isolatedpolynucleotidemoleculescorrespondingtomutantandwild-typeallelesofthemaized9geneandmethodsofuse,wo2007124312a2)和bzr1(choryandwang,2005,genesinvolvedinbrassinosteroidhormoneactioninplants,us6,921,848b2)来改变。类固醇激素在协调植物和动物的广泛的发育和生理过程中起着重要作用(thummelandchory2002,genesdev.16,3113-3129)。在植物中，类固醇激素油菜素内酯(br)在生长、发育和对生物和非生物胁迫的应答中起着广泛的作用(zhuetal2013,development140,1615-1620；clouse2011,plantcell23,1219-1230)。相比于通过核受体起作用的动物类固醇激素信号传导，在植物中，br结合于细胞表面受体激酶brassinosteroidinsensitive1(bri1)的胞外结构域并激活调控基因表达的细胞内信号转导级联(clouse2011,plantcell23,1219-1230；kinoshitaetal2005,nature433,167-171)。有包括中间体的激活和失活的多个步骤，所述中间体导致两个转录因子brassinazoleresistant1(bzr1)和bzr2(也称为bes1)的磷酸化。因此，信号转导bzr转录因子是将信号传导转换为br应答基因表达的靶组分。在植物激素信号传导中存在一种新兴模式，其中，由激素激活的靶转录因子也被特定的阻遏因子复合物负调控。例如，在茉莉酸(ja)、植物生长素、脱落酸(aba)和独角金内酯(sl)信号传导通路中，所述靶转录因子被利用作为由激素通路特异性抑制因子招募的常用协同阻遏物的topless(tpl)的抑制因子复合物负调控(pauwelsetal2010,nature464,788-791)。在ja转导通路中，jasmonatezimdomain(jaz)家族的转录阻遏因子直接地或通过衔接蛋白novelinteractorofjaz(ninja)与靶ja-应答转录激活子myc2相互作用并招募tpl(pauwelsetal2010,nature464,788-791)。因此，调控影响很多不同的目的农业性状的ga和br通路的能力对于商业性农业具有相当大的价值。本申请人的发明如上所述，本发明涉及通过异位表达peapod以增加植物中的根生物量的方法。不希望被理论所束缚，本申请人已经证明peapod(ppd)似乎通过与主生长调节子della和bzr的直接或间接相互作用来调控ga和br通路。对ppd蛋白的一级氨基酸结构的分析表明高度保守的新的植物特异性结构域仅存在于这些蛋白中。在广泛的双子叶植物、松柏类植物和一些单子叶植物(棕榈、香蕉、兰花和浮萍)而非禾本科植物中发现了这些基因的同源物。拟南芥属的ppd基因编码植物特异性tify家族(其以称为zim的结构域内发现的核心tif[f/y]xg基序命名)成员的蛋白(vanholmeetal2007,trendsplantsci.12,239-244)。这两个拟南芥属ppd蛋白ppd1和ppd2包括在与12个已经充分表征的充当茉莉酮酸酯应答的阻遏因子的jaz蛋白相同的ii类tify组中。但是，ppd蛋白和该组中的另一个非-jaz蛋白似乎并不参与对茉莉酮酸酯激素信号传导的应答(pauwelsetal2010,nature464,788-791)。同样地，不希望被理论所束缚，本申请人提出根据本发明，根生物量的增加由新的机制介导，该机制使用ppd基因调控ga和br通路。以下实施例3和4支持该提议。本发明也可以广义地说是由本申请的说明书中单独或共同提及或指出的部分、元素和特征，以及所述部分、元素或特征中的任意两个或更多个的任意或全部组合组成，并且其中在本文中提到的特定整体在本发明涉及的领域中具有已知的等同性，这种已知的等同性被认为如同单独提出一样被纳入本文。附图说明参照以下附图可以更好地理解本发明，其中：图1示出了包含来自一些植物物种的peapod区域的46个氨基酸残基，相同的残基用星号表示。图2示出了来自一些植物物种的peapod区域内的内部27个氨基酸残基，相同的残基用星号表示。图3示出了来自一些植物物种的peapod蛋白的tify结构域的6个氨基酸残基，相同的残基用星号表示。图4示出了ppd蛋白和保守的ppd、tify和jas*区域的大致位置的示意图。图5示出了在y2h测定中ppd蛋白的二聚化以及tpl和ninja之间的相互作用。图6示出了在y2h测定中ppd和ninja之间的相互作用以及tpl和bzr1之间的相互作用。图7示出了使用bifc测定证明的在嫩叶(a和b)和老叶(c)中的ppd、ninja、tpl和bzr1之间的相互作用。图8示出了ppd-ninja-tpl-bzr1复合物的示意图。图9示出了在y1h测定中ppd和bzr1之间的相互作用。图10示出了野生型、δppd突变体和peapod过表达子(ppd-ox)下胚轴长度对外源ga和pac施用的应答。图11示出了与野生型和载体对照相比过表达来自拟南芥(arabidopsisthaliana)的ppd(ppd-ox)或来自无油樟(ambroellatrichopoda)的peapod的黑麦草植物的根生长的增加。图12示出了与对照植物相比过表达ppd(ppd-ox)的苜蓿植物的根长度的增加。图13示出了来自拟南芥、北美云杉(piceasitchensis)、无油樟、小果野蕉(musaacuminate)、白三叶草(trifoliumrepens)和江南卷柏(selaginellamoellendorffii)的peapod蛋白是功能等效的。来自每一个的优化的peapod编码序列被用于互补peapod缺失突变体δppd拟南芥(生态型兰兹贝格(landsbergerecta))。在等同的发育阶段拍摄幼苗图片。实施例现在参照以下非限制性实施例对本发明进行说明。实施例1：对多个植物物种的peapod基因的表征为了鉴定其他植物物种中的ppd基因直系同源物，将来自拟南芥属ppd1基因(seqidno:27)的保守的ppd区域(46个氨基酸)用于使用搜索程序tblastn和blastp进行公共植物基因序列数据库搜索(altschuletal1990)。从多种植物物种(包括苔藓植物、松柏类植物、所检查的双子叶植物的所有目和一些单子叶植物目，包括棕榈、香蕉、兰花和浮萍)中鉴定peapod序列。peapod序列不存在于禾本科植物中。代表性的peapod蛋白和核酸序列分别示于seqidno:1-26和seqidno:83-107中。来自拟南芥ppd1的46个氨基酸peapod区域示于seqidno:27中。如图1所示，用vectornti(vnti)比对来自多肽seqidno:1-26的该区域。seqidno:28示出了这46个氨基酸ppd区域的共有序列。seqidno:29示出了相同的共有区域，但是示出了哪些氨基酸可存在于各可变位置处。来自拟南芥ppd1的46个氨基酸peapod区域内的27个氨基酸子序列示于seqidno:30中。对图1中的每个相同序列的这27个氨基酸子序列的比对示于图2中。seqidno:30示出了这27个氨基酸ppd区域的共有序列。seqidno:32示出了相同的共有区域，但是示出了哪些氨基酸可存在于各可变位置处。在每一个seqidno:1-26的ppd肽序列中，也有保守的tify基序，其位于所述46个氨基酸ppd区域之后。分隔开ppd区域的c-末端和tify基序的n-末端的氨基酸残基的数目取决于ppd的来源；例如，对于seqidno:1-26，所述数目在46至140个氨基酸之间变化。seqidno:33示出了拟南芥属ppd1序列的tify基序。对来自seqidno:1-26的tify基序的比对(如vanholmeetal2007,trendsplantsci.12,239-244中所述)示于图3中。seqidno:34示出了这6个氨基酸tify基序的共有序列。seqidno:35示出了相同的共有区域，但是示出了哪些氨基酸可存在于各可变位置。ppd和tify结构域中完全保守的残基在图1-3中用星号突出显示。本申请人认为以上所述的这些区域和基序存在于所有鉴定的peapod蛋白中并且可用于诊断这些peapod蛋白。实施例2：证明来自多个植物物种的peapod序列的peapod功能可以通过拟南芥属δppd突变体叶表型的互补作用来确认任意peapod序列的功能。首先将拟南芥属δppd突变体叶表型的互补作用用于鉴定拟南芥属ppd基因(white2006)。这可以通过将与圆顶叶和玫瑰花结的扭曲相反的野生型扁平叶表型和正常玫瑰花结形恢复为“螺旋桨”表型而看到。peapod序列，如seqinno:1-26中的那些(包括棕榈、松柏类植物、苔藓、兰花和其他双子叶植物物种)，或任意其他待测试的peapod序列可以通过本领域技术人员公知的方法被转化到拟南芥属δppd突变体中。这类方法中的一个实例在下文描述。克隆和基因构建体用于在拟南芥属δppd突变体中过表达拟南芥属ppd1的camv35s::拟南芥ppd1构建体的产生合成表达构建体以使得在拟南芥属δppd突变体中拟南芥ppd1在camv35s启动子(seqidno:129)下过表达。优化ppdorf以用于在拟南芥属中表达，这包括修饰的joshi序列(joshi1997,nucleicacidresearch15,6643-6653)、密码子的优化、mrna不稳定性序列的去除、多聚a信号序列的去除、隐性剪接位点的去除、bamhi可去除的c-末端v5表位和his标签尾部(编码seqidno:37)的添加和双终止密码子的添加。然后将该构建体(有或没有所述尾部)通过lr反应置于camv35s启动子和ocs终止子之间，其分别编码为seqidno:105和seqidno:111。用于在拟南芥属δppd突变体中过表达白三叶草ppd1的camv35s::白三叶草ppd构建体的产生合成表达构建体以使得在拟南芥属δppd突变体中白三叶草ppd在camv35s启动子(seqidno:129)下过表达。优化ppdorf以用于在拟南芥属中表达，这包括修饰的joshi序列(joshi1997,nucleicacidresearch15,6643-6653)、密码子的优化、mrna不稳定性序列的去除、多聚a信号序列的去除、隐性剪接位点的去除、bamhi可去除的c-末端v5表位和his标签尾部(编码seqidno:37)的添加和双终止密码子的添加。然后将该构建体(有或没有所述尾部)通过lr反应置于camv35s启动子和ocs终止子之间，其分别编码为seqidno:106和seqidno:112。用于在拟南芥属δppd突变体中过表达无油樟ppd的camv35s::无油樟ppd构建体的产生合成表达构建体以使得在拟南芥属δppd突变体中无油樟ppd在camv35s启动子(seqidno:129)下过表达。优化ppdorf以用于在拟南芥属中表达，这包括修饰的joshi序列(joshi1997,nucleicacidresearch15,6643-6653)、密码子的优化、mrna不稳定性序列的去除、多聚a信号序列的去除、隐性剪接位点的去除、bamhi可去除的c-末端v5表位和his标签尾部(编码seqidno:37)的添加和双终止密码子的添加。然后将该构建体(有或没有所述尾部)通过lr反应置于camv35s启动子和ocs终止子之间，其分别编码为seqidno:107和seqidno:113。用于在拟南芥属δppd突变体中过表达小果野蕉ppd的camv35s::小果野蕉ppd构建体的产生合成表达构建体以使得在拟南芥属δppd突变体中小果野蕉ppd在camv35s启动子(seqidno:129)下过表达。优化ppdorf以用于在拟南芥属中表达，这包括修饰的joshi序列(joshi1997,nucleicacidresearch15,6643-6653)、密码子的优化、mrna不稳定性序列的去除、多聚a信号序列的去除、隐性剪接位点的去除、bamhi可去除的c-末端v5表位和his标签尾部(编码seqidno:37)的添加和双终止密码子的添加。然后将该构建体(有或没有所述尾部)通过lr反应置于camv35s启动子和ocs终止子之间，其分别编码为seqidno:108和seqidno:114。用于在拟南芥属δppd突变体中过表达北美云杉ppd的camv35s::北美云杉ppd1构建体的产生合成表达构建体以使得在拟南芥属δppd突变体中北美云杉ppd在camv35s启动子(seqidno:129)下过表达。优化ppdorf以用于在拟南芥属中表达，这包括修饰的joshi序列(joshi1997,nucleicacidresearch15,6643-6653)、密码子的优化、mrna不稳定性序列的去除、多聚a信号序列的去除、隐性剪接位点的去除、bamhi可去除的c-末端v5表位和his标签尾部(编码seqidno:37)的添加和双终止密码子的添加。然后将该构建体(有或没有所述尾部)通过lr反应置于camv35s启动子和ocs终止子之间，其分别编码为seqidno:109和seqidno:115。用于在拟南芥属δppd突变体中过表达江南卷柏ppd1的camv35s::江南卷柏ppd构建体的产生合成表达构建体以使得在拟南芥属δppd突变体中江南卷柏ppd在camv35s启动子(seqidno:129)下过表达。优化ppdorf以用于在拟南芥属中表达，这包括修饰的joshi序列(joshi1997,nucleicacidresearch15,6643-6653)、密码子的优化、mrna不稳定性序列的去除、多聚a信号序列的去除、隐性剪接位点的去除、bamhi可去除的c-末端v5表位和his标签尾部(编码seqidno:37)的添加和双终止密码子的添加。然后将该构建体(有或没有所述尾部)通过lr反应置于camv35s启动子和ocs终止子之间，其分别编码为seqidno:110和seqidno:116。植物材料和生长条件拟南芥(l.)heynh生态型ler可以被用作野生型(wt)。功能缺失突变体(缺失ppd1和ppd2)的δppd缺失如先前white2006,pnas103,13238-13243中所述。使植物生长于持续为21℃的温度可控的温室中或在23℃下在16h光照_8h黑暗循环下的可控环境室中。拟南芥属转化可以通过蘸花(floraldip)浸润法将上述构建体转化到拟南芥属中(cloughandbent,1998,plantj16,735-43)。转化δppd株系以通过标准技术表达ppd多肽。通过标准pcr分析技术并结合转基因特异性和t-dna引物确认转基因植物。在t1幼苗(经浸润的植物的子代)中产生野生型叶和玫瑰花结表型的δppd株系的互补作用确认了所引入的基因的peapod功能，如照片所示。该方法可用于确认本申请人指定的任意基因的peapod功能，表明其适用于本发明。通过peapod缺失突变体δppd拟南芥生态型landsbergerecta的互补作用证明来自拟南芥、北美云杉、无油樟、小果野蕉和江南卷柏的peapod蛋白是功能等效的(图13)。实施例3：peapod可能参与调控油菜素类固醇信号传导通路本申请人使用酵母双杂交(y2h)测定、双分子荧光(bifc)来研究ppd、ninja、tpl和bzr1之间的相互作用。克隆和构建体如下产生用于y2h和bifc测定的构建体。合成编码以下的开放阅读框的拟南芥属dna序列：at4g14713(ppd1)和ppd1的截短和缺失衍生物、ppd1、ppd1δppd(编码氨基酸1-61的序列的n-末端截短)、ppd1δtify(编码氨基酸154-186的序列的内部缺失)、ppd1δjas*(编码氨基酸229-313的序列的c-末端截短)(图4)、at4g28910(ninja)、at1g15750(tpl)、at1g75080(bzr1)、合成的puas-35s启动子和编码gal4dbd和c-myc融合蛋白的序列，并通过geneart验证序列。提供大部分序列作为备用于gateway克隆到酵母和植物表达载体中的pentr221中的克隆。例外的是，整合了5’xho1和3’nco1限制性酶切位点的用于植物内转录激活测定的启动子序列被提供在pma-rq中进行克隆。用于瞬时luc报告测定的质粒：将具有-105bpcamv35s启动子上游的5xuasgal4dna结合位点的合成启动子克隆至包含含内含子的萤火虫荧光素酶基因(luc)和35spro::海肾荧光素酶(ren)作为内标的双荧光素酶构建体pnwa62内的xhoi-ncoi位点，以构建paml7。对于gal4dbd融合蛋白的过表达，使用gateway技术将编码gal4dna结合结构域(gal4dbd氨基酸1-147)以及n-末端gal4dbd与2xvp16激活子结构域(gal4dbd-vp16)或ppd1(gal4dbd-ppd1)的融合体(使用编码ggggs的接头)的dna序列克隆至prsh1(winichayakuletal2008)中以构建载体prsh1-gal4dbd、prsh1-gal4dbd-vp16和prsh1-gal4dbd-ppd1，用于在植物内表达该融合蛋白。用于酵母双杂交分析的质粒将bzr1、ninja、tpl和ppd1，以及ppd1的截短或缺失衍生物(ppd1δppd、ppd1δtify和ppd1δjas*)的全长编码序列gateway亚克隆至pdest32(n-末端gal4dbd)或pdest22(n-末端gal4ad)，以构建作为诱饵载体的pdest32-ppd1、pdest32-ppd1δppd、pdest32-ppd1δtify、pdest32-ppd1δjas*、pdest32-tpl以及作为猎物载体的pdest22-ppd1、pdest22-bzr1和pdest22-ninja。当被表达时，这些构建体产生序列56-67和70-72中所列的蛋白，包括dna结合结构域(dbd)、激活结构域(ad)、与dbd融合的ppd1(ppd1-dbd)、与ad融合的ppd1(ppd1-ad)、未与ad融合的tify结构域的ppd1(ppd1-tify-ad)、未与ad融合的jas结构域的ppd1(ppd1-jas*-ad)、topless(tpl)、与dbd融合的tpl(tpl-dbd)、ninja、与ad融合的ninja(ninja-ad)、与ad融合的bzr1(bzr1-ad)、不含与dbd融合的ppd结构域的ppd1(ppd1-δppd-dbd)、不含与dbd融合的tify结构域的ppd1(ppd1-tify-dbd)、不含与dbd融合的jas结构域的ppd1(ppd1-jas*-dbd)。用于双分子荧光互补的质粒使用具有n-末端融合体的双元bifc-gatewayyfp载体pdest-vyne(r)gw(venus氨基酸1-173)和pdest-vyce(r)gw(venus氨基酸156-239)构建以下载体：pdestnyfp-bzr1、pdestnyfp-ninja、pdestnyfp-ppd1、pdestcyfp-bzr1、pdestcyfp-ppd1、pdestcyfp-ppd1δppd、pdestcyfp-ppd1δtify和pdestcyfp-ppd1δjas*。对于与ppd1或bzr相互作用的蛋白在植物中的瞬时表达，将ninja和tpl亚克隆至prsh1以构建prsh1-ninja和prsh1-tpl。用于免疫共沉淀的质粒：将编码ppd1和3xc-mycc-末端融合体的合成的dna构建体亚克隆至prsh1以产生prsh1-ppd1-3xc-myc，同时将ninjacdna序列亚克隆至pb7fwg2,0(karimietal2002,trendsplantsci.7,193-195)以构建pb7fwg2-ninja-gfp。当被表达时，这些构建体产生序列63、65、73-82中所列的蛋白，包括topless(tpl)、ninja、双分子荧光(bifc)nyfp、bifccyfp、bifcnyfp-ninja、bifcnyfp-bzr1、bifccyfp-ppd1、bifccyfp-ninja、bifccyfp-bzr1、bifccyfp-ppd1-ppd、bifccyfp-ppd1-tify、bifccyfp-ppd1-jas*。使用proquest双杂交体系(invitrogen)分析ppd1、ninja、tpl和bzr1之间的相互作用。使用诱饵构建体和猎物构建体的组合共转化酵母菌株mav203(invitrogen)，在合成缺陷型(sd)sd/-leu/-trp琼脂平板上进行选择。在具有不同浓度的3-氨基三唑(3-at)(sigma)的sd/-leu/-trp/-his平板上使用10μl1:10和1:100稀释度的液滴测试转化菌株的相互作用。使用pdestnyfp和pdestcyfp质粒的组合，使用或不使用用于表达ninja(prsh1-ninja)或tpl(prsh1-tpl)和用于本氏烟(nicotianabenthamiana)叶的农杆菌-浸润的质粒进行瞬时bifc实验。对于浸润，从平板上重悬含有该双元载体的根癌农杆菌gv3101菌株并制备以用于如上所述用于luc测定的转化。将所有的yfp和表达菌株以1:1(体积/体积)的比例混合，并添加1/10体积的菌株p19。40h后，从各经浸润的叶中取5个叶盘样品。在取样用于显微镜荧光观察之前2小时，将叶用1μg/mldapi溶液浸润以染色细胞核。使用olympusfluoviewfv10i共聚焦激光扫描显微镜检测yfp荧光和dapi染色。每个实验重复两次。使用ppd1作为诱饵蛋白的y2h筛选鉴定ninja为与ppd1的直接相互作用因子。bifc测定的结果表明ppd1在植物中与ninja相互作用，并且表明tify基序对于该相互作用也是必需的(图7)。可能ninja作为tpl和ppd1之间的桥梁而发挥功能。使用y2h，没有观察到ppd1和bzr1之间的直接相互作用(图5)。但是，近来的串联亲和纯化(tap)实验已表明tpl可能与bzr1相互作用(wangetal2013,mol.cell.proteomics12,3653-3665)，并且在此，y2h结果确认发生了直接的相互作用(图6)。为了确定ppd蛋白的分子功能，在植物内研究了ppd1、ninja、tpl和bzr1的相互作用。使用双分子荧光(bifc)证明在未成熟的本氏烟叶的扁平细胞中，ppd1似乎在细胞核中与bzr1相互作用(图7a、b)。ppd1中的ninja-结合tify基序对于该相互作用是必需的。此外，在完全展开的叶中当nyfp-ppd1和cyfp-bzr1共表达时，未观察到相互作用(图7c)。有趣地是，ppd1和bzr1之间的相互作用在共表达ninja而非单独的tpl时得到恢复，表明在成熟的叶中缺少相互作用是由于内源ninja的限制。对于未成熟的叶，当ppd1ninja-结合tify基序缺失时，甚至在ninja和tpl共表达的存在下，也观察不到ppd1和bzr1之间的相互作用(图7c)。这些结果表明ppd1、ninja、tpl和bzr1在植物内作为复合物存在，并且表明ninja是招募ppd1以通过tpl与bzr1相互作用所必需的。ppd1似乎并不直接与靶bzr1转录因子相互作用。相反地，ppd1蛋白相互作用实验结果表明了这样一种模式，其中ppd蛋白使用ninja作为衔接体来招募tpl转录共表达子，该ppd-ninja-tpl复合物与bzr转录因子的ear基序相互作用(图8)。因此，在该模式中，拟南芥的peapod1(ppd1)蛋白会充当br信号传导通路的阻遏物并与ninja和tpl组合来负调控bzr1。实施例4：peapod可能参与调控赤霉素信号传导通路已知赤霉酸(ga)处理降低ga阻遏物della蛋白(包括rga1)的水平；为了确定ppd、della和ga信号传导通路之间的关系，本申请人进行ppd和della(rga1)之间的酵母双杂交(y2h)分析，并将赤霉酸(ga)激素和ga生物合成抑制剂(多效唑，pac)施用至野生型、δppd突变体和δppd突变体ppd过表达子(ppd-ox)。将proquest双杂交系统(invitrogen)用于分析ppd1和rga1之间的相互作用。将ppd1以及ppd1的截短或缺失衍生物(ppd1δppd、ppd1δtify和ppd1δjas*)的全长编码序列(图4)gateway亚克隆至pdest32(n-末端gal4dbd)或pdest22(n-末端gal4ad)。当被翻译时，这些产生了以下肽序列：seqidno:58、68、69、70、71、72，它们分别为ppd1-dbd、rga1、rga1-ad、ppd1-ppd-dbd、ppd1-tify-dbd、pp1-jas*-dbd。使用诱饵构建体和猎物构建体的组合共转化酵母菌株mav203(invitrogen)，在合成缺陷型(sd)sd/-leu/-trp琼脂平板上进行选择。在具有不同浓度的3-氨基三唑(3-at)(sigma)的sd/-leu/-trp/-his平板上使用10μl1:10和1:100稀释度的液滴测试转化菌株的相互作用。使用ppd1-dbd作为诱饵测试ppd1-rga1相互作用。将转化的酵母以10倍稀释液点样在对照培养基(-2)或具有15mm3at的选择培养基上。对照为空载体、dbd、gal4dna结合结构域、ad、gal4激活结构域(图4)。y2h结果表明ppd可以直接与della结合(图9)。对于外源施用ga或pac，用70％乙醇、0.01％tritonx-100对种子进行表面消毒10min，然后用100％乙醇消毒5min，在无菌的滤纸上空气干燥，转移到含有半强度的ms盐、1％蔗糖和0.8％琼脂的培养基平板上。将平板在黑暗中于4℃孵育5天，然后转移到24℃下的14h光照/10h黑暗的日循环中。通过荧光灯管(philipstld58w/865)以100μmm-2s-1的强度提供光照。使野生型(col-0)δppd突变体和转基因ppd-ox幼苗在含有不同浓度的ga(图10a)或pac(图10b)的培养基上生长5天。将ga(acros有机物)和pac(sigma-aldrich)分别溶解于乙醇和丙酮中，过滤消毒并添加到培养基平板中。将乙醇或丙酮(0.5％)用于对照处理。使幼苗在24℃下的14h光照/10h黑暗的日循环中生长5天，然后分析下胚轴长度(n＝35)。每次处理重复2次，误差棒＝平均值的标准误差。della的降低导致促进细胞扩增的della靶基因的转录的增加并可以通过测量生长在含不同水平ga的培养基上的幼苗的下胚轴长度来定量。最低浓度的ga(1μm)不能促进野生型(wt)下胚轴的伸长，而功能丧失的ppd突变体(δppd)和过表达苗的转基因ppd(ppd-ox)都显示出增加的下胚轴长度(图10a)。在更高的ga浓度(5-50μm)下，野生型下胚轴的伸长以剂量依赖的方式发生。在比较中，δppd和ppd-ox幼苗分别显示出对最高达5和10μmga的超敏感性伸长，其中它们大约达到相同的长度(图10a)。通过施用外源性多效唑(pac)抑制ga生物合成；这导致della阻遏蛋白的增加和细胞扩增的相应降低。野生型幼苗显示出下胚轴伸长的从0至10μmpac呈剂量依赖性降低(图10b)。再次，δppd幼苗显示出高敏感性应答，观察到下胚轴伸长在相同范围的pac施用下降低更大。但是，ppd-ox幼苗是相对不敏感的，直到pac浓度增加超过0.1μm，这之后，它们也显示出下胚轴长度的降低(图10b)δppd幼苗对ga的超敏感性应答潜在地反映了不存在通过ppd的转录因子阻遏时，用于降解的增加的della的靶标的组合。类似地，在δppd背景中添加pac可能导致与wt相比下胚轴伸长的降低更大，因为这种情况在不存在della天然拮抗剂之一-ppd时发生的，表明ppd和ga竞争与della结合。可以预测的是ppd的过表达将导致更高水平的对della的拮抗作用，因此，这些植物的下胚轴伸长应该对ga超敏感；实际上，这是本发明人在ppd-ox幼苗中所观察到的结果。在相反的情况下，当ga水平降低(通过施用pac)时，ppd-ox幼苗无应答直到pac浓度大于0.1μm。这可能反映了在该点内源ga水平具有足够的降低以观察到未被过表达的ppd拮抗的della蛋白的影响。实施例5：通过增加peapod在植物中的表达来增加根生物量构建体以下描述的是几种用于在不同启动子的控制下表达来自从不同物种的peapod序列的构建体，用于在双子叶植物和单子叶植物中表达。用于拟南芥属ppd在双子叶植物中的(根优选的)过表达的ptobrb7δ1.3::ppd双子叶植物构建体的产生合成含有用于在双子叶植物的根中表达的ppd1的编码序列的构建体，核酸序列示于seqidno:40和126中。ppdorf来自于拟南芥ppd1cdna(登录号nm_202819)并被修饰以包括修饰的joshi序列(joshi1997,nucleicacidresearch15,6643-6653)、隐性剪接位点的去除，在seqidno:40的情况中来自多年生黑麦草dgat1的第3个内含子的加入(seqidno:39)、bamhi可去除的c-末端v5表位和his标签尾部(编码seqidno:37)的添加和双终止密码子的添加。优化内含子的位置以用于剪接位点预测，通过netgene2进行(http://www.cbs.dtu.dk/services/netgene2/)。然后通过lr反应将seqidno40(具有或不具有所述尾部)的构建体置于tobrb7δ1.3启动子(yamamotoetal1991,plantcell,3:371-382)和nos终止子之间以产生seqidno:42和seqidno:44，其分别编码为seqidno:36和seqidno:38。在seqidno126所示的构建体(具有或不具有所述c-末端尾部)上进行类似的亚克隆步骤，其分别编码为seqidno:108和seqidno:114。用于拟南芥属ppd在双子叶植物中的(根优选的)过表达的ptobrb7δ0.6::ppd双子叶植物构建体的产生合成含有用于在双子叶植物的根中表达的ppd1的编码序列的构建体，核酸序列示于seqidno:40和126中。ppdorf来自于拟南芥ppd1cdna(登录号nm_202819)并被修饰以包括修饰的joshi序列(joshi1997,nucleicacidresearch15,6643-6653)、隐性剪接位点的去除，在seqidno:40的情况中来自多年生黑麦草dgat1的第3个内含子的加入(seqidno:39)、bamhi可去除的c-末端v5表位和his标签尾部(编码seqidno:37)的添加和双终止密码子的添加。优化内含子的位置以用于剪接位点预测，通过netgene2进行(http://www.cbs.dtu.dk/services/netgene2/)。然后通过lr反应将(具有或不具有所述尾部)的构建体置于tobrb7δ0.6启动子(yamamotoetal1991,plantcell,3:371-382)和nos终止子之间以产生seqidno:43和seqidno:45，其分别编码为seqidno:36和seqidno:38。在seqidno126所示的构建体(具有或不具有所述c-末端尾部)上进行类似的亚克隆步骤，其分别编码为seqidno:108和seqidno:114。用于拟南芥属ppd在双子叶植物中的(根优选的)过表达的patwrky6::ppd双子叶植物构建体的产生合成含有用于在双子叶植物的根中表达的ppd1的编码序列的构建体，核酸序列示于seqidno:41和126中。ppdorf来自于拟南芥ppd1cdna(登录号nm_202819)并被修饰以包括修饰的joshi序列(joshi1997,nucleicacidresearch15,6643-6653)、隐性剪接位点的去除，在seqidno:41的情况中来自多年生黑麦草dgat1的第3个内含子的加入(seqidno:39)、bamhi可去除的c-末端v5表位和his标签尾部(编码seqidno:37)的添加和双终止密码子的添加。优化内含子的位置以用于剪接位点预测，通过netgene2进行(http://www.cbs.dtu.dk/services/netgene2/)。然后通过lr反应将(具有或不具有所述尾部)的构建体置于烟草atwrky6启动子(robatzekandsomssich2001,theplantjournal,28:123-133)和nos终止子之间以产生seqidno:44和seqidno:47，其分别编码为seqidno:36和seqidno:38。在seqidno126所示的构建体(具有或不具有所述c-末端尾部)上进行类似的亚克隆步骤，其分别编码为seqidno:108和seqidno:114。用于拟南芥属ppd在单子叶植物中的(根优选的)过表达的ptobrb7δ1.3::ppd单子叶植物构建体的产生合成含有用于在单子叶植物的根中表达的ppd1的编码序列的构建体，核酸序列示于seqidno:49和120中。优化ppdorf以用于在单子叶植物中表达，这包括修饰的joshi序列(joshi1997,nucleicacidresearch15,6643-6653)、密码子的优化、mrna不稳定性序列的去除、多聚a信号序列的去除、隐性剪接位点的去除、在seqidno:49的情况中来自多年生黑麦草dgat1的第3个内含子的加入(seqidno:48)、bamhi可去除的c-末端v5表位和his标签尾部(编码seqidno:37)的添加和双终止密码子的添加。优化内含子的位置以用于剪接位点预测，通过deepc2进行(http://www.cbs.dtu.dk/services/netgene2/)。然后通过lr反应将(具有或不具有所述尾部)的构建体置于ptobrb7δ1.3启动子(yamamotoetal1991,plantcell,3:371-382)和nos终止子之间以产生seqidno:50和seqidno:53，其分别编码为seqidno:36和seqidno:38。在seqidno120所示的构建体(具有或不具有所述c-末端尾部)上进行类似的亚克隆步骤，其分别编码为seqidno:108和seqidno:114。用于拟南芥属ppd在双子叶植物中的(根优选的)过表达的ptobrb7δ0.6::ppd单子叶植物构建体的产生合成含有用于在单子叶植物的根中表达的ppd1的编码序列的构建体，核酸序列示于seqidno:49和120中。优化ppdorf以用于在单子叶植物中表达，这包括修饰的joshi序列(joshi1997,nucleicacidresearch15,6643-6653)、密码子的优化、mrna不稳定性序列的去除、多聚a信号序列的去除、隐性剪接位点的去除、在seqidno:49的情况中来自多年生黑麦草dgat1的第3个内含子的加入(seqidno:48)、bamhi可去除的c-末端v5表位和his标签尾部(编码seqidno:37)的添加和双终止密码子的添加。优化内含子的位置以用于剪接位点预测，通过deepc2进行(http://www.cbs.dtu.dk/services/netgene2/)。然后通过lr反应将(具有或不具有所述尾部)的构建体置于ptobrb7δ0.6启动子(yamamotoetal1991,plantcell,3:371-382)和nos终止子之间以产生seqidno:51和seqidno:54，其分别编码为seqidno:36和seqidno:38。在seqidno120所示的构建体(具有或不具有所述c-末端尾部)上进行类似的亚克隆步骤，其分别编码为seqidno:108和seqidno:114。用于拟南芥属ppd在单子叶植物中的(根优选的)过表达的patwrky6::ppd单子叶植物构建体的产生合成含有用于在单子叶植物的根中表达的ppd1的编码序列的构建体，核酸序列示于seqidno:49和120中。优化ppdorf以用于在单子叶植物中表达，这包括修饰的joshi序列(joshi1997,nucleicacidresearch15,6643-6653)、密码子的优化、mrna不稳定性序列的去除、多聚a信号序列的去除、隐性剪接位点的去除、在seqidno:49的情况中来自多年生黑麦草dgat1的第3个内含子的加入(seqidno:48)、bamhi可去除的c-末端v5表位和his标签尾部(编码seqidno:37)的添加和双终止密码子的添加。优化内含子的位置以用于剪接位点预测，通过deepc2进行(http://www.cbs.dtu.dk/services/netgene2/)。然后通过lr反应将(具有或不具有所述尾部)的构建体置于patwrky6启动子(robatzekandsomssich2001theplantjournal,28:123-133)和nos终止子之间以产生seqidno:52和seqidno:55，其分别编码为seqidno:36和seqidno:38。在seqidno120所示的构建体(具有或不具有所述c-末端尾部)上进行类似的亚克隆步骤，其分别编码为seqidno:108和seqidno:114。含有来自其他植物物种的ppd的表达构建体的产生为了证明可使用来自非拟南芥属的物种的ppd，我们合成了这样的构建体，其在被置于合适的启动子下时会表达来自以下物种的ppd：拟南芥(ppd对照)、豆科植物白三叶草、原始被子植物无油樟、单子叶植物小果野蕉、松柏类植物挪威云杉和北美云杉以及石松门植物江南卷柏。优化这些构建体以用于在水稻(包括单子叶植物内含子)或拟南芥属(无内含子)中表达。对这些构建体进行改造以包含可去除的尾部，该尾部由编码v5-his标签的核酸序列的任一侧上的两个bamhi位点(编码甘氨酸-丝氨酸)组成。所述肽序列(有和没有c-末端v5-his标签尾部)示于序列108-119中。被优化用于在水稻中表达的相应的核酸序列示于序列120-125中，被优化用于在拟南芥属中表达的相应的核酸序列示于序列126-131中。通过gatewaylr反应将拟南芥属优化版本置于双元载体的适当的启动子下游，并通过蘸花方法转化到δppd拟南芥中。通过gatewaylr反应将水稻优化版本置于适当的启动子下游，并通过生物裂解的方式(biolystically)转化到黑麦草中。植物转化拟南芥属的转化按照实施例2所述的方法转化拟南芥属。苜蓿的转化使用改良自samacetal.2000(methodsinmolecularbiology343:301-311)的方法通过农杆菌介导的转化产生过表达peapod构建体的苜蓿植物。植物组织对regen-sy的高度可再生基因型(bingham1991cropsci.31,1098)进行鉴定，其具有在体外条件下从体细胞胚胎形成完整植株的能力。使用该方案选择单一基因型用于植物转化实验。农杆菌菌株和载体使用双元载体prsh1::ppd-op和prsh1::ppd-op-v5用于所有的转化实验。载体prsh1是part27(gleave1992plantmolbiol20:1203-1207)的衍生物并且包含由camv35s启动子表达的bar选择基因和也在camv35s启动子的表达下的禾本科植物优化的peapod基因。将质粒转移到根癌农杆菌菌株gv310中，并在含有200mgl-1壮观霉素的ym平板上选择转化体。在从细菌培养物中制备质粒dna之后，通过限制性图谱确认质粒成功转移到农杆菌中。将双元质粒的培养物加入到含有100mg/l壮观霉素的25mlmannitolglutamateluria(mgl)培养基中。使细菌培养物于28℃在旋转振荡器上生长过夜(16个小时)。通过离心(3000xg)收集培养物，去除上清液并将细胞重悬于溶于用于植物转化的制品中的5ml10mmmgso4溶液中。植物转化收集来自regen-sy的组织培养物相应基因型的第二和第三完全展开叶，并在50％乙醇溶液中冲洗30秒，然后在无菌蒸馏水中洗涤，接着在次氯酸钠溶液(3％可用氯)中进行表面消毒。然后将无菌叶切成叶蝶，接着将它们浮于sh1.5d愈伤组织诱导培养基的溶液上。对于叶蝶的转化，将约3ml农杆菌培养液加入到12mlsh1.5d中，并孵育15分钟，偶尔搅拌以将叶用农杆菌全部覆盖。然后去除细菌溶液，将叶小片在无菌滤纸上稍微印干，然后使近轴侧朝上置于用0.8％植物用琼脂凝胶化的sh1.5d上。接着将平板于25℃在黑暗中孵育5天。接种后，从平板上去除叶小片，并在无菌sh1.5d液体培养基中冲洗3-4次，在无菌滤纸上印干，放置于补充有用于筛选的5mg/l草铵膦和100mg/l头孢噻肟和特美汀的sh1.5d平板上，并于25℃在黑暗中孵育15天。然后将在叶小片上形成的愈伤组织传代至新鲜的sh1.5d选择培养基中，再持续15天。然后将转化的愈伤组织转移到补充有草甘膦、头孢噻肟和特美汀的bl0培养基中用于形成体细胞胚胎。一旦胚胎形成，将它们解剖并转移到用于发芽筛选的半强度msn培养基中。将整个生根植株转移并使其在温室中定植(establish)。pcr分析进行pcr分析以确认t-dna稳定整合至从转化实验中回收的植物的基因组中。使用基因组dnamini试剂盒(geneaid)从约50mg的体外生长的叶中提取基因组dna。使用特异性针对nos3’聚腺苷酸化信号的引物对(ocs3’-1f,5’-gatatgcgagacgcctatga-3’[seqidno:132]；ocs3’-1r,5’-gagttcccttcagtgaacgt-3’[seqidno:133])产生439bp的扩增产物。也包括仅包含质粒dna模板、未转化的植物dna或水的对照反应。pcr反应的方案由如下组成：94℃下起始变性5分钟，30个循环的95℃30s、55℃15s、72℃1min，和72℃延伸10min。在tae缓冲液中通过凝胶电泳将扩增产物在1.0％琼脂糖凝胶上进行分析并使用bio-radgeldoc成像系统进行显影。用于苜蓿转化的培养基的一般组成a.mglb.sh1.5dc.bl0(blaydes培养基)·blaydes改良的基础培养基(phytotechnologylaboratories)d.半msn培养基·ms大量和微量营养元素，半强度(duchefa)黑麦草的转化使用改良自altpeteretal.2000(molecularbreeding6:519-528)的方法通过微粒轰击产生过表达peapod构建体的黑麦草植物。从高达7mm的不成熟花序诱导用于转化的愈伤组织。收获花分蘖，在次氯酸钠溶液(4％可用氯)中进行表面消毒，解剖，然后在黑暗中于25℃下培养4至6周，接着在含大量营养物、微量营养物和维生素的murashige和skoog(ms)基础培养基(1962physiolplant.15:473-497)上进行转化，该培养基补充有30g/l麦芽糖、5mg/l2,4-d，ph被调节至5.8，并用6g/l琼脂糖固化。使用invitrogenpurelinkhipureplasmidmaxiprep试剂盒制备质粒，浓度调节至1μg/μl。将质粒pach1用于选择，该质粒包含含有由具有第一个内含子的水稻肌动蛋白启动子表达并终止于nos3’聚腺苷酸化信号的嵌合潮霉素转移酶(hph)基因(bilangetal.1991gene100:247-250)的表达盒。将含有ppd表达盒的质粒以1:1的摩尔比与pach1混合。使用sanfordetal.1993的方法(meth.enzymol.217:483-509.)将质粒dna包被于m17钨颗粒(1.4μm平均直径分布)上，并使用dupontpds-1000/氦生物弹射粒子输送系统转化至靶组织中。在转化前高达6小时，将愈伤组织传代培养至含有64g/l甘露醇的愈伤组织起始培养基上。转化后(约16小时)，然后将转化的愈伤组织转移到补充有2mg/l2,4-d、不含甘露醇的ms基础培养基中。2天后，将愈伤组织转移至含有200mg/l潮霉素的相同培养基中，并在黑暗中培养4小时用于转基因株系的选择。在补充有0.2mg/l激动素、30g/l蔗糖、和50mg/l潮霉素，被调节至ph5.8并用8g/l植物用琼脂固化的ms基础培养基上在光照下发生来自体细胞胚胎的整株植物的再生。将转化的植物转移到含有温室的环境中用于分析。转化体的pcr分析进行pcr分析以确认hph和ppd转基因稳定整合至从转化实验中回收的植物的基因组中。使用基因组dnamini试剂盒(geneaid)从约50mg的体外生长的叶中提取基因组dna。分别使用特异性针对hphgene的引物对(hpt-1,5’-gctggggcgtcggtttccactatccg-3’[seqidno:134]；hpt-2,5’-cgcataacagcggtcattgactggagc-3’)[seqidno:135]；和特异性针对nos3’聚腺苷酸化信号的引物对(nos3’-1f,5’-ctgttgccggtcttgcgatg-3’–[seqidno:136]；nos3’-1r,5’-gtcacatagatgacaccgcg-3’–[seqidno:137])产生375bp和202bp的扩增产物。也包括仅包含质粒dna模板、未转化的植物dna或水的对照反应。pcr反应的方案由如下组成：94℃下起始变性5分钟，30个循环的95℃30s、55℃15s、72℃1min，和72℃延伸10min。在tae缓冲液中通过凝胶电泳将扩增产物在1.0％琼脂糖凝胶上进行分析并使用bio-radgeldoc成像系统进行显影。禾本科植物转化体的dna印迹分析使用dna印迹杂交评估每个株系的转基因拷贝数。使用doylej和doylej1990的方法(focus,12:13-15)从温室生长的植物的叶材料提取基因组dna用于dna印迹杂交。将dna(20μg)消化，并在0.8％琼脂糖凝胶上进行分离并使用0.4nnaoh通过毛细管转移将其转移至尼龙膜(roche)上。当探测hph和ppd转基因时，分别用xbai或hindiii消化基因组dna。使用digpcr合成试剂盒制备探针。分别使用特异性针对hph基因的引物对(rgh1,5’-ctcgtgctttcagcttcgatgtag-3’[seqidno:138]；rgh5,5’-gctggggcgtcggtttccactatcgg-3’[seqidno:139])和特异性针对ppd的引物对(grppd1f,5’-cacaggatggattctccaagg-3’[seqidno:140]；grppd1r,5’-taaggtccacggagaggttc-3’[seqidno:141])产生906bp和586bp的扩增产物用于探针。使用标准缓冲液(roche)在45℃下进行预杂交(1小时)和杂交(12小时)。使用cdp-star作为化学发光底物根据制造商的方案使用非放射性方法实现检测。使用bio-radchemidocmp系统和软件检测光信号。抗ppd1蛋白的多克隆抗体的产生和免疫印迹使用全长的拟南芥ppd1蛋白通过genscript产生定制的抗ppd1亲和纯化的兔多克隆抗体。在1:5000的稀释度下所述抗体能够通过免疫印迹检测低于10ng的经纯化的ppd蛋白。将植物组织冷冻在液氮中，并研磨成细粉。将冷冻的组织粉末以1.0/1.5(重量/体积)的比例加入到含有50mmtrisph7.5、150mmnacl、1mmedta、10％(体积/体积)甘油、5mmdtt、1％(体积/体积)完全蛋白酶抑制剂混合物(sigma)和1％(体积/体积)tritonx-100的提取缓冲液中，均质化直到解冻，然后以16,300g在4℃下离心12min。通过bradford测定(coomassieplus,thermoscientific)对上清液中的总的溶解蛋白定量，进行调节以得到等同的总蛋白浓度/样品(通常在10-40μg之间)，在1xnupagelds样品缓冲液(invitrogen)中变性并在4-12％bis-trissds/page凝胶(novex)上运行。在使用iblot装置(invitrogen)印迹到pvdf膜上后，使用1:5,000稀释度的10抗ppd1多克隆抗体，然后使用1:5,000稀释度的20山羊抗兔hrp抗体(sigma)，用westernbrightecl试剂(advansta)检测蛋白，并使用chemidoctm仪器(biorad)捕获图像。转化体的根生物量分析从野生型和用在组成型启动子下的拟南芥属ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物的塑料生长袋中并修剪至相同的高度。使植物在温室中生长约6周，通过观察生长袋外的根生长可以看出，与野生型植物相比，用在组成型启动子下的拟南芥属ppd转化的黑麦草植物的根生物量/生长/长度/分支增加(图11)。可以通过在流水下冲洗根、去除所连接的地上部分(叶和茎)并在65℃下干燥根48小时，然后对干重进行称重来对根生物量的增加进行定量(表2)。表2从野生型和用在三个根启动子之一下的拟南芥属ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物的塑料生长袋中并修剪至相同的高度。使植物在温室中生长约6周，通过观察生长袋外的根生长可以看出，与野生型植物相比，用在组成型启动子下的拟南芥属ppd转化的黑麦草植物的根生物量/生长/长度/分支增加(图12)。可以通过在流水下冲洗根、去除所连接的地上部分(叶和芽)并在65℃下干燥根48小时，然后对干重进行称重来对根生物量的增加进行定量(表3和表4)。表3表4从野生型和用在组成型启动子下的拟南芥属ppd转化的苜蓿植物(品种usd5)中取约相等大小的插条。将插条种植到含有盆栽混合物的塑料生长袋中。使植物在具有x小时日长的生长室中生长约6周；通过观察生长袋外的根生长可以看出，与野生型植物相比，用在组成型启动子下的拟南芥属ppd转化的苜蓿植物的根生物量/生长/长度/分支增加。可以通过在流水下冲洗根(对这些进行拍照，图12)，然后去除所连接的地上部分(叶和茎)并在65℃下干燥根48小时，然后进行称重来对根生物量的增加进行定量(表5)。表5植物株系合并的来自10个插条的干根重(mg)未转化的对照43转化的株系1110转化的株系254转化的株系3160转化的株系459转化的株系553转化的株系685转化的株系7148从野生型和用在组成型启动子下的白三叶草ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物的塑料生长袋中并修剪至相同的高度。使植物在温室中生长约6周，通过观察生长袋外的根生长可以看出，与野生型植物相比，用在组成型启动子下的白三叶草ppd转化的黑麦草植物的根生物量/生长/长度/分支增加。可以通过在流水下冲洗根、去除所连接的地上部分(叶和茎)并在65℃下干燥根48小时，然后对干重进行称重来对根生物量的增加进行定量。从野生型和用在三个根启动子之一下的白三叶草ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物的塑料生长袋中并修剪至相同的高度。使植物在温室中生长约6周，通过观察生长袋外的根生长可以看出，与野生型植物相比，用在组成型启动子下的白三叶草ppd转化的黑麦草植物的根生物量/生长/长度/分支增加。可以通过在流水下冲洗根、去除所连接的地上部分(叶和茎)并在65℃下干燥根48小时，然后对干重进行称重来对根生物量的增加进行定量。从野生型和用在组成型启动子下的无油樟ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物的塑料生长袋中并修剪至相同的高度。使植物在温室中生长约6周，通过观察生长袋外的根生长可以看出，与野生型植物相比，用在组成型启动子下的无油樟ppd转化的黑麦草植物的根生物量/生长/长度/分支增加(图11)。可以通过在流水下冲洗根、去除所连接的地上部分(叶和茎)并在65℃下干燥根48小时，然后对干重进行称重来对根生物量的增加进行定量。从野生型和用在三个根启动子之一下的无油樟ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物的塑料生长袋中并修剪至相同的高度。使植物在温室中生长约6周，通过观察生长袋外的根生长可以看出，与野生型植物相比，用在组成型启动子下的无油樟ppd转化的黑麦草植物的根生物量/生长/长度/分支增加。可以通过在流水下冲洗根、去除所连接的地上部分(叶和茎)并在65℃下干燥根48小时，然后对干重进行称重来对根生物量的增加进行定量。从野生型和用在组成型启动子下的小果野蕉ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物的塑料生长袋中并修剪至相同的高度。使植物在温室中生长约6周，通过观察生长袋外的根生长可以看出，与野生型植物相比，用在组成型启动子下的小果野蕉ppd转化的黑麦草植物的根生物量/生长/长度/分支增加。可以通过在流水下冲洗根、去除所连接的地上部分(叶和茎)并在65℃下干燥根48小时，然后对干重进行称重来对根生物量的增加进行定量。从野生型和用在三个根启动子之一下的小果野蕉ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物的塑料生长袋中并修剪至相同的高度。使植物在温室中生长约6周，通过观察生长袋外的根生长可以看出，与野生型植物相比，用在组成型启动子下的小果野蕉ppd转化的黑麦草植物的根生物量/生长/长度/分支增加。可以通过在流水下冲洗根、去除所连接的地上部分(叶和茎)并在65℃下干燥根48小时，然后对干重进行称重来对根生物量的增加进行定量。从野生型和用在组成型启动子下的北美云杉ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物的塑料生长袋中并修剪至相同的高度。使植物在温室中生长约6周，通过观察生长袋外的根生长可以看出，与野生型植物相比，用在组成型启动子下的北美云杉ppd转化的黑麦草植物的根生物量/生长/长度/分支增加。可以通过在流水下冲洗根、去除所连接的地上部分(叶和茎)并在65℃下干燥根48小时，然后对干重进行称重来对根生物量的增加进行定量。从野生型和用在三个根启动子之一下的北美云杉ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物的塑料生长袋中并修剪至相同的高度。使植物在温室中生长约6周，通过观察生长袋外的根生长可以看出，与野生型植物相比，用在组成型启动子下的北美云杉ppd转化的黑麦草植物的根生物量/生长/长度/分支增加。可以通过在流水下冲洗根、去除所连接的地上部分(叶和茎)并在65℃下干燥根48小时，然后对干重进行称重来对根生物量的增加进行定量。从野生型和用在组成型启动子下的江南卷柏ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物的塑料生长袋中并修剪至相同的高度。使植物在温室中生长约6周，通过观察生长袋外的根生长可以看出，与野生型植物相比，用在组成型启动子下的江南卷柏ppd转化的黑麦草植物的根生物量/生长/长度/分支增加。可以通过在流水下冲洗根、去除所连接的地上部分(叶和茎)并在65℃下干燥根48小时，然后对干重进行称重来对根生物量的增加进行定量。从野生型和用在三个根启动子之一下的江南卷柏ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物的塑料生长袋中并修剪至相同的高度。使植物在温室中生长约6周，通过观察生长袋外的根生长可以看出，与野生型植物相比，用在组成型启动子下的江南卷柏ppd转化的黑麦草植物的根生物量/生长/长度/分支增加。可以通过在流水下冲洗根、去除所连接的地上部分(叶和茎)并在65℃下干燥根48小时，然后对干重进行称重来对根生物量的增加进行定量。将来自野生型和用在组成型启动子下的白三叶草ppd转化的δppd拟南芥属植物的种子划破并播种于盆栽混合物中。使植物在具有x小时日长的生长室中生长约6周；通过观察盆外的根生长可以看出，与野生型植物相比，用在组成型启动子下的白三叶草ppd转化的拟南芥属植物的根生物量/生长/长度/分支增加。可以通过在流水下冲洗根、然后去除所连接的地上部分(叶和茎)并在65℃下干燥根48小时，然后进行称重来对根生物量的增加进行定量。将来自野生型和用在组成型启动子下的小果野蕉ppd转化的δppd拟南芥属植物的种子划破并播种于盆栽混合物中。使植物在具有x小时日长的生长室中生长约6周；通过观察盆外的根生长可以看出，与野生型植物相比，用在组成型启动子下的小果野蕉ppd转化的拟南芥属植物的根生物量/生长/长度/分支增加。可以通过在流水下冲洗根、然后去除所连接的地上部分(叶和茎)并在65℃下干燥根48小时，然后进行称重来对根生物量的增加进行定量。将来自野生型和用在组成型启动子下的无油樟ppd转化的δppd拟南芥属植物的种子划破并播种于盆栽混合物中。使植物在具有x小时日长的生长室中生长约6周；通过观察盆外的根生长可以看出，与野生型植物相比，用在组成型启动子下的无油樟ppd转化的拟南芥属植物的根生物量/生长/长度/分支增加。可以通过在流水下冲洗根、然后去除所连接的地上部分(叶和茎)并在65℃下干燥根48小时，然后进行称重来对根生物量的增加进行定量。将来自野生型和用在组成型启动子下的挪威云杉ppd转化的δppd拟南芥属植物的种子划破并播种于盆栽混合物中。使植物在具有x小时日长的生长室中生长约6周；通过观察盆外的根生长可以看出，与野生型植物相比，用在组成型启动子下的挪威云杉ppd转化的拟南芥属植物的根生物量/生长/长度/分支增加。可以通过在流水下冲洗根、然后去除所连接的地上部分(叶和茎)并在65℃下干燥根48小时，然后进行称重来对根生物量的增加进行定量。将来自野生型和用在组成型启动子下的江南卷柏ppd转化的δppd拟南芥属植物的种子划破并播种于盆栽混合物中。使植物在具有x小时日长的生长室中生长约6周；通过观察盆外的根生长可以看出，与野生型植物相比，用在组成型启动子下的江南卷柏ppd转化的拟南芥属植物的根生物量/生长/长度/分支增加。可以通过在流水下冲洗根、然后去除所连接的地上部分(叶和茎)并在65℃下干燥根48小时，然后进行称重来对根生物量的增加进行定量。对转化体的耐旱性分析从野生型和用在组成型启动子下的拟南芥属ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。从野生型和用在组成型启动子下的白三叶草ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。从野生型和用在组成型启动子下的小果野蕉ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。从野生型和用在组成型启动子下的无油樟ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。从野生型和用在组成型启动子下的北美云杉ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。从野生型和用在组成型启动子下的江南卷柏ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。从野生型和用在三个根启动子之一下的拟南芥属ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。从野生型和用在三个根启动子之一下的白三叶草ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。从野生型和用在三个根启动子之一下的小果野蕉ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。从野生型和用在三个根启动子之一下的无油樟ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。从野生型和用在三个根启动子之一下的北美云杉ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。从野生型和用在三个根启动子之一下的江南卷柏ppd转化的黑麦草植物中取等数量(通常为4-10个)的分蘖。将分蘖种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。将相同数目(通常为4-10)的野生型和用在组成型启动子下的拟南芥属ppd转化的δppd拟南芥属幼苗种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。将相同数目(通常为4-10)的野生型和用在组成型启动子下的白三叶草ppd转化的δppd拟南芥属幼苗种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。将相同数目(通常为4-10)的野生型和用在组成型启动子下的小果野蕉ppd转化的δppd拟南芥属幼苗种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。将相同数目(通常为4-10)的野生型和用在组成型启动子下的无油樟ppd转化的δppd拟南芥属幼苗种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。将相同数目(通常为4-10)的野生型和用在组成型启动子下的挪威云杉ppd转化的δppd拟南芥属幼苗种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。将相同数目(通常为4-10)的野生型和用在组成型启动子下的江南卷柏ppd转化的δppd拟南芥属幼苗种植到含有盆栽混合物和土壤的大盆中。使植物在温室中生长，然后将一半的各类型植物进行水胁迫(通常为12％的重量含水量，正好在永久凋萎点之上)，同时使另一半保持含水(通常为22％的重量含水量，约为田间持水量)。通过与wt植物比较根和芽生物量可以对过表达ppd的植物的增加的干旱胁迫耐受性进行定量。文献achard，p.genschikp.(2009).releasingthebrakesofplantgrowth：howgasshutdowndellaproteins.j.exp.bot.，60：1985-1092.altschul，s.f.，gish，w.，miller，w.，myers，e.w.，lipman，d.j.(1990).basiclocalalignmentsearchtool.j.mol.biol.215：403-410.anjum，sa.，wang，lc.，farooq，m.，hussain，m.，zou，cm.(2011)brassinolideapplicationimprovesthedroughttoleranceinmaizethroughmodulationofenzymaticantioxidantsandleafgasexchange.j.agronomyandcropsci.197：177-185.choryandwang，2005，genesinvolvedinbrassinosteroidhormoneactioninplants，us6,921,848b2cloughs.j.andbenta.f.(1998)floraldip：asimplifiedmethodforagrobacterium-mediatedtransformationofarabidopsisthaliana.plantj.16：735-743.clouse，s.d.brassinosteroidsignaltransduction：fromreceptorkinaseactivationtotranscriptionalnetworksregulatingplantdevelopment.plantcell23，1219-1230(2011).clouse，s.d.＆sasse，j.m.brassinosteroids：essentialregulatorsofplantgrowthanddevelopment.annu.rev.plantphysiol.plantmol.biol.16，427-451(1998).gallego-bartolome，j.，minguet，eg.，grau-enguix，f.，abbas，m.，locascio，a.，thomas，sg.，alabadi，d.，blazquez，ma.(2012).molecularmechanismfortheinteractionbetweengibberellinandbrassionosteroidsignallingpathwaysinarabidopsis.pnas，109：13446-13451.garcia，m.e.，lynch，t.，peeters，j.，snowden，c.＆finkelstein，r.asmallplant-specificproteinfamilyofabifivebindingproteins(afps)regulatesstressresponseingerminatingarabidopsisseedsandseedlings.plantmol.biol.67，643-658(2008).guo，h.，li，l.，aluru，m.，sluru，s.，yin，y.(2013).mechanismsandnetworkesforbrassinosteroidregulatedgeneexpression.currentopinioninplantbiology16：545-553.he，j-x.，gendron，jm.，sun，y.，gampaia，sl.，gendron，n.，sun，cq.，wang，z-y.(2005).bzr1isatranscriptionalrepressorwithdualrolesinbrassinosteroidhomeostasisandgrowthresponses.science307：1634-1638.hothorn，m.，belkhadir，y.，dreux，m.，dabi，t.，noel，j.p.，wilson，i.a.＆chory，j.structuralbasisofsteroidhormoneperceptionbythereceptorkinasebri1.nature474，467-471(2011).hou，x.，leelyc，xia，k.，yan，y.，yuh.(2010).dellasmodulatejasmonatesignallingviacompetitivebindingtojazs.devel.cell19：884-894.jiang，w-b.，lin，w-h.(2013).brassinosteroidfunctionsinarabidopsisseeddevelopment.plantsignaling&behavior8：10，e25928；october2013.jiang，l.，liu，x.，x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valglyval130135140serproalalysserileleualalysproleulysleuleuthrglu145150155160gluaspileserglnleuthrarggluaspcysarglyspheleulys165170175asplysglymetargargprosertrpasnlysserglnalailegln180185190glnvalleuserleulysalaleutyrgluproglyaspaspsergly195200205alaglyilephearglysileleuvalserglnprovalasnpropro210215220argvalthrthrthrleuilegluproserasngluleuglualacys225230235240glyargvalsertyrprogluaspasnglyalacyshisargmetasp245250255serproargseralaglupheserglyglyserglyhisphevalser260265270glulysaspglyhislysthrthrileserproargserproalaglu275280285thrsergluleuvalglyglnmetthrilephetyrserglylysval290295300asnvaltyraspglyileproproglulysalaargserilemethis305310315320phealaalaasnproileaspleuprogluasnglyilephealaser325330335serargmetileserlysleuileserlysglulysmetmetgluleu340345350proglnlysglyleuglulysalaasnserserargaspserglymet355360365gluglyglnalaasnarglysvalserleuglnargtyrargglulys370375380arglysasparglyspheserlysalalyslyscysproglyvalala385390395400serserserleuglumetpheleuasncysglnproargmetlysala405410415alatyrserglnasnleuglycysthrglyserproleuhissergln420425430serprogluserglnthrlysserproasnleuservalaspleuasn435440445sergluglyile450<210>60<211>391<212>prt<213>artificialsequence<220><223>consensussequence<400>60metprolyslyslysarglysvalserserglyalaasnpheasngln151015serglyasnilealaaspserserleuserphethrphethrasnser202530serasnglyproasnleuilethrthrglnthrasnserglnalaleu354045serglnproilealaserserasnvalhisaspasnphemetasnasn505560gluilethralaserlysileaspaspglyasnasnserlysproleu65707580serproglytrpthraspglnthralatyrasnalapheglyilethr859095thrglymetpheasnthrthrthrmetaspaspvaltyrasntyrleu100105110pheaspaspgluaspthrproproasnprolyslysgluglyglyser115120125asnglnthrserleutyrlyslysalaglysertyrgluproglyasp130135140aspserglyalaglyilephearglysileleuvalserglnproval145150155160asnproproargvalthrthrthrleuilegluproserasngluleu165170175glualacysglyargvalsertyrprogluaspasnglyalacyshis180185190argmetaspserproargseralaglupheserglyglyserglyhis195200205phevalserglulysaspglyhislysthrthrileserproargser210215220proalagluthrsergluleuvalglyglnmetthrilephetyrser225230235240glylysvalasnvaltyraspglyileproproglulysalaargser245250255ilemethisphealaalaasnproileaspleuprogluasnglyile260265270phealaserserargmetileserlysleuileserlysglulysmet275280285metgluleuproglnlysglyleuglulysalaasnserserargasp290295300serglymetgluglyglnalaasnarglysvalserleuglnargtyr305310315320argglulysarglysasparglyspheserlysalalyslyscyspro325330335glyvalalaserserserleuglumetpheleuasncysglnproarg340345350metlysalaalatyrserglnasnleuglycysthrglyserproleu355360365hisserglnserprogluserglnthrlysserproasnleuserval370375380aspleuasnsergluglyile385390<210>61<211>427<212>prt<213>artificialsequence<220><223>consensussequence<400>61metprolyslyslysarglysvalserserglyalaasnpheasngln151015serglyasnilealaaspserserleuserphethrphethrasnser202530serasnglyproasnleuilethrthrglnthrasnserglnalaleu354045serglnproilealaserserasnvalhisaspasnphemetasnasn505560gluilethralaserlysileaspaspglyasnasnserlysproleu65707580serproglytrpthraspglnthralatyrasnalapheglyilethr859095thrglymetpheasnthrthrthrmetaspaspvaltyrasntyrleu100105110pheaspaspgluaspthrproproasnprolyslysgluglyglyser115120125asnglnthrserleutyrlyslysalaglysermetaspvalglyval130135140serproalalysserileleualalysproleulysleuleuthrglu145150155160gluaspileserglnleuthrarggluaspcysarglyspheleulys165170175asplysglymetargargprosertrpasnlysserglnalailegln180185190glnvalleuserleulysalaleutyrgluproglyaspaspsergly195200205alaglyilephearglysileleuvalserglnprovalasnpropro210215220argvalthrthrthrleuilegluproserasngluleuglualacys225230235240glyargvalsertyrprogluaspasnglyalacyshisargmetasp245250255serproargseralaglupheserglyglyserglyhisphevalser260265270glulysaspglyhislysthrthrileserproargserproalaglu275280285thrsergluleuilemethisphealaalaasnproileaspleupro290295300gluasnglyilephealaserserargmetileserlysleuileser305310315320lysglulysmetmetgluleuproglnlysglyleuglulysalaasn325330335serserargaspserglymetgluglyglnalaasnarglysvalser340345350leuglnargtyrargglulysarglysasparglyspheserlysala355360365lyslyscysproglyvalalaserserserleuglumetpheleuasn370375380cysglnproargmetlysalaalatyrserglnasnleuglycysthr385390395400glyserproleuhisserglnserprogluserglnthrlysserpro405410415asnleuservalaspleuasnsergluglyile420425<210>62<211>368<212>prt<213>artificialsequence<220><223>consensussequence<400>62metprolyslyslysarglysvalserserglyalaasnpheasngln151015serglyasnilealaaspserserleuserphethrphethrasnser202530serasnglyproasnleuilethrthrglnthrasnserglnalaleu354045serglnproilealaserserasnvalhisaspasnphemetasnasn505560gluilethralaserlysileaspaspglyasnasnserlysproleu65707580serproglytrpthraspglnthralatyrasnalapheglyilethr859095thrglymetpheasnthrthrthrmetaspaspvaltyrasntyrleu100105110pheaspaspgluaspthrproproasnprolyslysgluglyglyser115120125asnglnthrserleutyrlyslysalaglysermetaspvalglyval130135140serproalalysserileleualalysproleulysleuleuthrglu145150155160gluaspileserglnleuthrarggluaspcysarglyspheleulys165170175asplysglymetargargprosertrpasnlysserglnalailegln180185190glnvalleuserleulysalaleutyrgluproglyaspaspsergly195200205alaglyilephearglysileleuvalserglnprovalasnpropro210215220argvalthrthrthrleuilegluproserasngluleuglualacys225230235240glyargvalsertyrprogluaspasnglyalacyshisargmetasp245250255serproargseralaglupheserglyglyserglyhisphevalser260265270glulysaspglyhislysthrthrileserproargserproalaglu275280285thrsergluleuvalglyglnmetthrilephetyrserglylysval290295300asnvaltyraspglyileproproglulysalaargserilemethis305310315320phealaalaasnproileaspleuprogluasnglyilephealaser325330335serargmetileserlysleuileserlysglulysmetmetgluleu340345350proglnlysglyleuglulysalaasnserserargaspserglymet355360365<210>63<211>1131<212>prt<213>artificialsequence<220><223>consensussequence<400>63metserserleuserarggluleuvalpheleuileleuglnpheleu151015aspgluglulysphelysgluthrvalhislysleugluglngluser202530glyphephepheasnmetlystyrphegluaspgluvalhisasngly354045asntrpaspgluvalglulystyrleuserglyphethrlysvalasp505560aspasnargtyrsermetlysilephephegluilearglysglnlys65707580tyrleuglualaleuasplyshisaspargprolysalavalaspile859095leuvallysaspleulysvalpheserthrpheasnglugluleuphe100105110lysgluilethrglnleuleuthrleugluasnphearggluasnglu115120125glnleuserlystyrglyaspthrlysseralaargalailemetleu130135140valgluleulyslysleuileglualaasnproleupheargasplys145150155160leuglnpheprothrleuargasnserargleuargthrleuileasn165170175glnserleuasntrpglnhisglnleucyslysasnproargproasn180185190proaspilelysthrleuphevalasphissercysglyproproasn195200205glyalaargalaproserprovalasnasnproleuleuglyglyile210215220prolysalaglyglypheproproleuglyalahisglyprophegln225230235240prothralaserprovalprothrproleualaglytrpmetserser245250255proserservalprohisproalavalseralaglyalailealaleu260265270glyglyproserileproalaalaleulyshisproargthrpropro275280285thrasnalaserleuasptyrproseralaaspsergluhisvalser290295300lysargthrargprometglyileseraspgluvalasnleuglyval305310315320asnmetleuprometserpheserglyglnalahisglyhisserpro325330335alaphelysalaproaspaspleuprolysthrvalalaargthrleu340345350serglnglyserserprometsermetaspphehisproilelysgln355360365thrleuleuleuvalglythrasnvalglyaspileglyleutrpglu370375380valglyserarggluargleuvalglnlysthrphelysvaltrpasp385390395400leuserlyscyssermetproleuglnalaalaleuvallysglupro405410415valvalservalasnargvaliletrpserproaspglyserleuphe420425430glyvalalatyrserarghisilevalglnleutyrsertyrhisgly435440445glygluaspmetargglnhisleugluileaspalahisvalglygly450455460valasnaspileserpheserthrproasnlysglnleucysvalile465470475480thrcysglyaspasplysthrilelysvaltrpaspalaalathrgly485490495vallysarghisthrphegluglyhisglualaprovaltyrserval500505510cysprohistyrlysgluasnileglnpheilepheserthralaleu515520525aspglylysilelysalatrpleutyraspasnmetglyserargval530535540asptyraspalaproglyargtrpcysthrthrmetalatyrserala545550555560aspglythrargleuphesercysglythrserlysaspglygluser565570575pheilevalglutrpasnglusergluglyalavallysargthrtyr580585590glnglyphehislysargserleuglyvalvalglnpheaspthrthr595600605lysasnargtyrleualaalaglyaspasppheserilelysphetrp610615620aspmetaspalavalglnleuleuthralaileaspglyaspglygly625630635640leuglnalaserproargileargpheasnlysgluglyserleuleu645650655alavalserglyasngluasnvalilelysilemetalaasnserasp660665670glyleuargleuleuhisthrphegluasnilesersergluserser675680685lysproalaileasnserilealaalaalaalaalaalaalaalathr690695700seralaglyhisalaaspargseralaasnvalvalserileglngly705710715720metasnglyaspserargasnmetvalaspvallysprovalilethr725730735glugluserasnasplysserlysiletrplysleuthrgluvalser740745750gluproserglncysargserleuargleuprogluasnleuargval755760765alalysileserargleuilephethrasnserglyasnalaileleu770775780alaleualaserasnalailehisleuleutrplystrpglnargasn785790795800gluargasnalathrglylysalathralaserleuproproglngln805810815trpglnproalaserglyileleumetthrasnaspvalalagluthr820825830asnprogluglualavalprocysphealaleuserlysasnaspser835840845tyrvalmetseralaserglyglylysileserleupheasnmetmet850855860thrphelysthrmetalathrphemetproproproproalaalathr865870875880pheleualaphehisproglnaspasnasnileilealaileglymet885890895aspaspserthrileglniletyrasnvalargvalaspgluvallys900905910serlysleulysglyhisserlysargilethrglyleualapheser915920925asnvalleuasnvalleuvalserserglyalaaspalaglnleucys930935940valtrpasnthraspglytrpglulysglnargserlysvalleupro945950955960leuproglnglyargproasnseralaproseraspthrargvalgln965970975phehisglnaspglnalahispheleuvalvalhisgluthrglnleu980985990alailetyrgluthrthrlysleuglucysmetlysglntrpalaval99510001005arggluserleualaproilethrhisalathrphesercysasp101010151020serglnleuvaltyralaserphemetaspalathrvalcysval102510301035pheserseralaasnleuargleuargcysargvalasnproser104010451050alatyrleuproalaserleuserasnserasnvalhisproleu105510601065valilealaalahisproglngluproasnmetphealavalgly107010751080leuseraspglyglyvalhisilephegluproleugluserglu108510901095glylystrpglyvalalaproproalagluasnglyseralaser110011051110glyalaprothralaproservalglyalaseralaseraspgln111511201125proglnarg1130<210>64<211>1292<212>prt<213>artificialsequence<220><223>consensussequence<400>64metlysleuleuserserilegluglualacysasnilecysargleu151015lyslysleulyscysserlysglulysprolyscysalalyscysleu202530lysasnasntrpglucysargtyrserprolysthrlysargserpro354045leuthrargalahisleuth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uence<220><223>consensussequence<400>68metlysargasphishisglnpheglnglyargleuserasnhisgly151015thrserserserserserserileserlysasplysmetmetmetval202530lyslysglugluaspglyglyglyasnmetaspaspgluleuleuala354045valleuglytyrlysvalargserserglumetalagluvalalaleu505560lysleugluglnleugluthrmetmetserasnvalglngluaspgly65707580leuserhisleualathraspthrvalhistyrasnprosergluleu859095tyrsertrpleuaspasnmetleusergluleuasnproproproleu100105110proalaserserasnglyleuaspprovalleuproserprogluile115120125cysglypheproalaserasptyraspleulysvalileproglyasn130135140alailetyrglnpheproalaileaspserserserserserasnasn145150155160glnasnlysargleulyssercysserserproaspsermetvalthr165170175serthrserthrglythrglnileglyglyvalileglythrthrval180185190thrthrthrthrthrthrthrthralaalaglygluserthrargser195200205valileleuvalaspserglngluasnglyvalargleuvalhisala210215220leumetalacysalaglualaileglnglnasnasnleuthrleuala225230235240glualaleuvallysglnileglycysleualavalserglnalagly245250255alametarglysvalalathrtyrphealaglualaleualaargarg260265270iletyrargleuserproproglnasnglnileasphiscysleuser275280285aspthrleuglnmethisphetyrgluthrcysprotyrleulysphe290295300alahisphethralaasnglnalaileleuglualaphegluglylys305310315320lysargvalhisvalileaspphesermetasnglnglyleuglntrp325330335proalaleumetglnalaleualaleuarggluglyglyproprothr340345350pheargleuthrglyileglyproproalaproaspasnserasphis355360365leuhisgluvalglycyslysleualaglnleualaglualailehis370375380valglupheglutyrargglyphevalalaasnserleualaaspleu385390395400aspalasermetleugluleuargproseraspthrglualavalala405410415valasnservalphegluleuhislysleuleuglyargproglygly420425430ileglulysvalleuglyvalvallysglnilelysprovalilephe435440445thrvalvalgluglngluserasnhisasnglyprovalpheleuasp450455460argphethrgluserleuhistyrtyrserthrleupheaspserleu465470475480gluglyvalproasnserglnasplysvalmetsergluvaltyrleu485490495glylysglnilecysasnleuvalalacysgluglyproaspargval500505510gluarghisgluthrleuserglntrpglyasnargpheglyserser515520525glyleualaproalahisleuglyserasnalaphelysglnalaser530535540metleuleuservalpheasnserglyglnglytyrargvalgluglu545550555560serasnglycysleumetleuglytrphisthrargproleuilethr565570575thrseralatrplysleuserthralaalatyr580585<210>69<211>726<212>prt<213>artificialsequence<220><223>consensussequence<400>69metprolyslyslysarglysvalserserglyalaasnpheasngln151015serglyasnilealaaspserserleuserphethrphethrasnser202530serasnglyproasnleuilethrthrglnthrasnserglnalaleu354045serglnproilealaserserasnvalhisaspasnphemetasnasn505560gluilethralaserlysileaspaspglyasnasnserlysproleu65707580serproglytrpthraspglnthralatyrasnalapheglyilethr859095thrglymetpheasnthrthrthrmetaspaspvaltyrasntyrleu100105110pheaspaspgluaspthrproproasnprolyslysgluglyglyser115120125asnglnthrserleutyrlyslysalaglysermetlysargasphis130135140hisglnpheglnglyargleuserasnhisglythrserserserser145150155160serserileserlysasplysmetmetmetvallyslysglugluasp165170175glyglyglyasnmetaspaspgluleuleualavalleuglytyrlys180185190valargserserglumetalagluvalalaleulysleugluglnleu195200205gluthrmetmetserasnvalglngluaspglyleuserhisleuala210215220thraspthrvalhistyrasnprosergluleutyrsertrpleuasp225230235240asnmetleusergluleuasnproproproleuproalaserserasn245250255glyleuaspprovalleuproserprogluilecysglypheproala260265270serasptyraspleulysvalileproglyasnalailetyrglnphe275280285proalaileaspserserserserserasnasnglnasnlysargleu290295300lyssercysserserproaspsermetvalthrserthrserthrgly305310315320thrglnileglyglyvalileglythrthrvalthrthrthrthrthr325330335thrthrthralaalaglygluserthrargservalileleuvalasp340345350serglngluasnglyvalargleuvalhisalaleumetalacysala355360365glualaileglnglnasnasnleuthrleualaglualaleuvallys370375380glnileglycysleualavalserglnalaglyalametarglysval385390395400alathrtyrphealaglualaleualaargargiletyrargleuser405410415proproglnasnglnileasphiscysleuseraspthrleuglnmet420425430hisphetyrgluthrcysprotyrleulysphealahisphethrala435440445asnglnalaileleuglualaphegluglylyslysargvalhisval450455460ileaspphesermetasnglnglyleuglntrpproalaleumetgln465470475480alaleualaleuarggluglyglyproprothrpheargleuthrgly485490495ileglyproproalaproaspasnserasphisleuhisgluvalgly500505510cyslysleualaglnleualaglualailehisvalglupheglutyr515520525argglyphevalalaasnserleualaaspleuaspalasermetleu530535540gluleuargproseraspthrglualavalalavalasnservalphe545550555560gluleuhislysleuleuglyargproglyglyileglulysvalleu565570575glyvalvallysglnilelysprovalilephethrvalvalglugln580585590gluserasnhisasnglyprovalpheleuaspargphethrgluser595600605leuhistyrtyrserthrleupheaspserleugluglyvalproasn610615620serglnasplysvalmetsergluvaltyrleuglylysglnilecys625630635640asnleuvalalacysgluglyproaspargvalgluarghisgluthr645650655leuserglntrpglyasnargpheglyserserglyleualaproala660665670hisleuglyserasnalaphelysglnalasermetleuleuserval675680685pheasnserglyglnglytyrargvalglugluserasnglycysleu690695700metleuglytrphisthrargproleuilethrthrseralatrplys705710715720leuserthralaalatyr725<210>70<211>413<212>prt<213>artificialsequence<220><223>consensussequence<400>70metlysleuleuserserilegluglualacysasnilecysargleu151015lyslysleulyscysserlysglulysprolyscysalalyscysleu202530lysasnasntrpglucysargtyrserprolysthrlysargserpro354045leuthrargalahisleuthrgluvalgluserargleugluargleu505560gluglnleupheleuleuilepheproarggluasnleuasnmetile65707580leulysmetaspserleuglnaspilelysalaleuleuthrglyleu859095phevalglnaspasnvalasnlysaspalavalthraspargleuala100105110servalgluthraspmetproleuthrleuargglnhisargileser115120125alathrserserserglugluserserasnlysglyglnargglnleu130135140thrvalserserargserasnglnthrserleutyrlyslysalagly145150155160sertyrgluproglyaspaspserglyalaglyilephearglysile165170175leuvalserglnprovalasnproproargvalthrthrthrleuile180185190gluproserasngluleuglualacysglyargvalsertyrproglu195200205aspasnglyalacyshisargmetaspserproargseralagluphe210215220serglyglyserglyhisphevalserglulysaspglyhislysthr225230235240thrileserproargserproalagluthrsergluleuvalglygln245250255metthrilephetyrserglylysvalasnvaltyraspglyilepro260265270proglulysalaargserilemethisphealaalaasnproileasp275280285leuprogluasnglyilephealaserserargmetileserlysleu290295300ileserlysglulysmetmetgluleuproglnlysglyleuglulys305310315320alaasnserserargaspserglymetgluglyglnalaasnarglys325330335valserleuglnargtyrargglulysarglysasparglyspheser340345350lysalalyslyscysproglyvalalaserserserleuglumetphe355360365leuasncysglnproargmetlysalaalatyrserglnasnleugly370375380cysthrglyserproleuhisserglnserprogluserglnthrlys385390395400serproasnleuservalaspleuasnsergluglyile405410<210>71<211>449<212>prt<213>artificialsequence<220><223>consensussequence<400>71metlysleuleuserserilegluglualacysasnilecysargleu151015lyslysleulyscysserlysglulysprolyscysalalyscysleu202530lysasnasntrpglucysargtyrserprolysthrlysargserpro354045leuthrargalahisleuthrgluvalgluserargleugluargleu505560gluglnleupheleuleuilepheproarggluasnleuasnmetile65707580leulysmetaspserleuglnaspilelysalaleuleuthrglyleu859095phevalglnaspasnvalasnlysaspalavalthraspargleuala100105110servalgluthraspmetproleuthrleuargglnhisargileser115120125alathrserserserglugluserserasnlysglyglnargglnleu130135140thrvalserserargserasnglnthrserleutyrlyslysalagly145150155160sermetaspvalglyvalserproalalysserileleualalyspro165170175leulysleuleuthrglugluaspileserglnleuthrarggluasp180185190cysarglyspheleulysasplysglymetargargprosertrpasn195200205lysserglnalaileglnglnvalleuserleulysalaleutyrglu210215220proglyaspaspserglyalaglyilephearglysileleuvalser225230235240glnprovalasnproproargvalthrthrthrleuilegluproser245250255asngluleuglualacysglyargvalsertyrprogluaspasngly260265270alacyshisargmetaspserproargseralaglupheserglygly275280285serglyhisphevalserglulysaspglyhislysthrthrileser290295300proargserproalagluthrsergluleuilemethisphealaala305310315320asnproileaspleuprogluasnglyilephealaserserargmet325330335ileserlysleuileserlysglulysmetmetgluleuproglnlys340345350glyleuglulysalaasnserserargaspserglymetgluglygln355360365alaasnarglysvalserleuglnargtyrargglulysarglysasp370375380arglyspheserlysalalyslyscysproglyvalalaserserser385390395400leuglumetpheleuasncysglnproargmetlysalaalatyrser405410415glnasnleuglycysthrglyserproleuhisserglnserproglu420425430serglnthrlysserproasnleuservalaspleuasnserglugly435440445ile<210>72<211>390<212>prt<213>artificialsequence<220><223>consensussequence<400>72metlysleuleuserserilegluglualacysasnilecysargleu151015lyslysleulyscysserlysglulysprolyscysalalyscysleu202530lysasnasntrpglucysargtyrserprolysthrlysargserpro354045leuthrargalahisleuthrgluvalgluserargleugluargleu505560gluglnleupheleuleuilepheproarggluasnleuasnmetile65707580leulysmetaspserleuglnaspilelysalaleuleuthrglyleu859095phevalglnaspasnvalasnlysaspalavalthraspargleuala100105110servalgluthraspmetproleuthrleuargglnhisargileser115120125alathrserserserglugluserserasnlysglyglnargglnleu130135140thrvalserserargserasnglnthrserleutyrlyslysalagly145150155160sermetaspvalglyvalserproalalysserileleualalyspro165170175leulysleuleuthrglugluaspileserglnleuthrarggluasp180185190cysarglyspheleulysasplysglymetargargprosertrpasn195200205lysserglnalaileglnglnvalleuserleulysalaleutyrglu210215220proglyaspaspserglyalaglyilephearglysileleuvalser225230235240glnprovalasnproproargvalthrthrthrleuilegluproser245250255asngluleuglualacysglyargvalsertyrprogluaspasngly260265270alacyshisargmetaspserproargseralaglupheserglygly275280285serglyhisphevalserglulysaspglyhislysthrthrileser290295300proargserproalagluthrsergluleuvalglyglnmetthrile305310315320phetyrserglylysvalasnvaltyraspglyileproproglulys325330335alaargserilemethisphealaalaasnproileaspleuproglu340345350asnglyilephealaserserargmetileserlysleuileserlys355360365glulysmetmetgluleuproglnlysglyleuglulysalaasnser370375380serargaspserglymet385390<210>73<211>142<212>prt<213>artificialsequence<220><223>consensussequence<400>73metvalserlysglyglugluleuphethrglyvalvalproileleu151015valgluleuaspglyaspvalasnglyhislyspheservalsergly202530gluglygluglyaspalathrtyrglylysleuthraspaspglyasn354045tyrlysthrargalagluvallysphegluglyaspthrleuvalasn505560argilegluleulysglyileaspphelysgluaspglyasnileleu65707580glyhislysleuglutyrasntyrasnserhisasnvaltyrilethr859095alaasplysglnlysasnglyilelysalaasnphelysilearghis100105110asnileglugluglnlysleuileserglugluaspleuglyproarg115120125prothrserglyservalaspleugluglythralaprogly130135140<210>74<211>111<212>prt<213>artificialsequence<220><223>consensussequence<400>74metasplysglnlysasnglyilelysalaasnphelysilearghis151015asnilegluaspglyglyvalglnleualaasphistyrglnglnasn202530thrproileglyaspglyprovalleuleuproaspasnhistyrleu354045sertyrglnseralaleuserlysaspproasnglulysargasphis505560metvalleuleugluphevalthralaalaglyilethrleuglymet65707580aspgluleutyrlystyrprotyraspvalproasptyralaglypro859095argprothrserglyservalaspleugluglythralaprogly100105110<210>75<211>568<212>prt<213>artificialsequence<220><223>consensussequence<400>75metvalserlysglyglugluleuphethrglyvalvalproileleu151015valgluleuaspglyaspvalasnglyhislyspheservalsergly202530gluglygluglyaspalathrtyrglylysleuthraspaspglyasn354045tyrlysthrargalagluvallysphegluglyaspthrleuvalasn505560argilegluleulysglyileaspphelysgluaspglyasnileleu65707580glyhislysleuglutyrasntyrasnserhisasnvaltyrilethr859095alaasplysglnlysasnglyilelysalaasnphelysilearghis100105110asnileglugluglnlysleuileserglugluaspleuglyproarg115120125prothrserglyservalaspleugluglythralaproglysermet130135140aspaspaspasnglyleugluleuserleuglyleusercysglygly145150155160serthrglylysalalysglyasnasnasnasnasnalaglyserser165170175sergluasntyrargalagluglyglyaspargseralalysvalile180185190aspaspphelysasnpheleuhisprothrserglnargproalaglu195200205proserserglyserglnargseraspserglyglnglnproprogln210215220asnphepheasnaspleuserlysalaprothrthrglualagluala225230235240serthrlysproleutrpvalgluaspgluserarglysglualagly245250255asnlysarglyspheglypheproglymetasnaspasplyslyslys260265270glulysaspserserhisvalaspmethisglulyslysthrlysala275280285serhisvalserthralathraspgluglyserthralagluasnglu290295300aspvalalaglusergluvalglyglyglyserserserasnhisala305310315320lysgluvalvalargproprothraspthrasnilevalaspasnleu325330335thrglyglnargargserasnhisglyglyserglythrglugluphe340345350thrmetargasnmetsertyrthrvalprophethrvalhisprogln355360365asnvalvalthrsermetprotyrserleuprothrlysglusergly370375380glnhisalaalaalathrserleuleuglnproasnalaasnalagly385390395400asnleuproilemetpheglytyrserprovalglnleuprometleu405410415asplysaspglyserglyglyilevalalaleuserglnserprophe420425430alaglyargvalproserasnseralathralalysglygluglylys435440445glnprovalalaglugluglysersergluaspalasergluargpro450455460thrglyaspasnserasnleuasnthralapheserpheasppheser465470475480alailelysproglymetalaalaaspvallyspheglyglysergly485490495alaargproasnleuprotrpvalserthrthrglyserglyprohis500505510glyargthrileserglyvalthrtyrargtyrasnalaasnglnile515520525lysilevalcysalacyshisglyserhismetserproglugluphe530535540valarghisalasergluglutyrvalserproglusersermetgly545550555560metthralaalaseralahisthr565<210>76<211>479<212>prt<213>artificialsequence<220><223>consensussequence<400>76metvalserlysglyglugluleuphethrglyvalvalproileleu151015valgluleuaspglyaspvalasnglyhislyspheservalsergly202530gluglygluglyaspalathrtyrglylysleuthraspaspglyasn354045tyrlysthrargalagluvallysphegluglyaspthrleuvalasn505560argilegluleulysglyileaspphelysgluaspglyasnileleu65707580glyhislysleuglutyrasntyrasnserhisasnvaltyrilethr859095alaasplysglnlysasnglyilelysalaasnphelysilearghis100105110asnileglugluglnlysleuileserglugluaspleuglyproarg115120125prothrserglyservalaspleugluglythralaproglysermet130135140thrseraspglyalathrserthrseralaalaalaalaalaalaala145150155160alaalaalaalaargarglysprosertrparggluarggluasnasn165170175argargarggluargargargargalavalalaalalysiletyrthr180185190glyleuargalaglnglyasptyrasnleuprolyshiscysaspasn195200205asngluvalleulysalaleucysvalglualaglytrpvalvalglu210215220gluaspglythrthrtyrarglysglycyslysproleuproglyglu225230235240ilealaglythrserserargvalthrprotyrserserglnasngln245250255serproleuserseralapheglnserproileprosertyrglnval260265270serproserserserserpheproserproserargglygluproasn275280285asnasnmetserserthrphephepropheleuargasnglyglyile290295300proserserleuproserleuargileserasnsercysprovalthr305310315320proprovalserserprothrserlysasnprolysproleuproasn325330335trpgluserilealalysglnsermetalailealalysglnsermet340345350alaserpheasntyrprophetyralavalseralaproalaserpro355360365thrhisarghisglnphehisthrproalathrileproglucysasp370375380gluseraspserserthrvalaspserglyhistrpileserphegln385390395400lysphealaglnglnglnpropheseralasermetvalprothrser405410415prothrpheasnleuvallysproalaproglnglnmetserproasn420425430thralaalapheglngluileglyglnsersergluphelyspheglu435440445asnserglnvallysprotrpgluglygluargilehisaspvalgly450455460metgluaspleugluleuthrleuglyasnglylysalaarggly465470475<210>77<211>425<212>prt<213>artificialsequence<220><223>consensussequence<400>77metasplysglnlysasnglyilelysalaasnphelysilearghis151015asnilegluaspglyglyvalglnleualaasphistyrglnglnasn202530thrproileglyaspglyprovalleuleuproaspasnhistyrleu354045sertyrglnseralaleuserlysaspproasnglulysargasphis505560metvalleuleugluphevalthralaalaglyilethrleuglymet65707580aspgluleutyrlystyrprotyraspvalproasptyralaglypro859095argprothrserglyservalaspleugluglythralaproglyser100105110metaspvalglyvalserproalalysserileleualalysproleu115120125lysleuleuthrglugluaspileserglnleuthrarggluaspcys130135140arglyspheleul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