西北植物学报2003,23(5):822—829Acta Bot.Boreal.-Occident.Sin.文章编号:1000-4025(2003)05-0822-08小麦抗白粉病基因解超杰,杨作民,孙其信*(中国农业大学农学与生物技术学院,北京100094)摘 要:到目前为止,小麦中已经鉴定出31个主效抗白粉病基因位点(Pm1-P m31),对这些小麦抗白粉病基因位点的来源、染色体定位、遗传特点以及载体品种等方面进行了概括性综述.关键词:小麦;白粉病;抗病基因;染色体定位中图分类号:Q945.8;S512.1 文献标识码:AResistance genes to powdery mildew in wheatXIE Chao-jie,YANG Zuo-m in,SU N Qi-x in*(College of Agronomy and Bio-technology,China Agricultural U nivers ity,Beijing100094,C hina)Abstract:Pow dery mildew is o ne o f the m ost serious w heat diseases in China.Breeding fo r resistant culti-vars has pro ved the effective and an env ir onm entally safe w ay to co ntrol this disease.Up to now,31m ajor w heat pow dery m ildew r esistance g enes(P m1—Pm31)had been repor ted.This paper briefly review ed the so urces,chrom oso mal location,g enetics,and the carr ying cultivars o f these P m genes.Key words:wheat;pow dery mildew;resistance g enes;chromo som al locatio n 小麦白粉病是由小麦白粉菌Blumeria grami-nis f.sp.tritici(=Ery sip he gr aminis f.sp.tritici)引起的一种气传病害,是威胁我国小麦生产的重要常见病害之一[1].培育抗病品种是防治小麦白粉病的一项既安全又经济有效的措施,而抗病育种的基础是多样化的抗源.深入研究抗病基因的抗性表现及遗传特点,将有助于对这些抗源进行有效利用.目前小麦白粉病抗性基因的研究中有关主效基因的研究较多,其正式命名的基因符号为Pm(po w-dery m ildew),未正式命名的临时符号或早期符号为Ml.曾有相关文献对小麦抗白粉病基因研究进行综述报道[2～8].近年来研究工作进展很快,陆续有新基因鉴定出来,其所在的染色体位置也基本明确(表1).P m1:该基因最早在澳大利亚春小麦品种T hew、法国品种Nor mandie以及加拿大品种Ax minster中发现.Sears和Briggle[9]利用中国春端体系将Pm1基因定位于7A染色体长臂上. Hsam等[10]发现该基因位点包括4个复等位基因(P m1a,Pm1b,P m1c,Pm1d),其载体品种分别为Ax minster/8*Cc、含有一粒小麦(T riticum mono-coccum,2n=2x=14,AA)抗白粉病基因的抗病衍生系M ocZlatka、德国抗病新品系Weihenstephan Stamm M1N(M1N)(原定名为P m18)和来自罗马尼亚的斯卑尔脱小麦(T.sp elta v ar.d uhamelianum(TSD),2n=6x=42,AABBDD)抗病系T RI2258.收稿日期:2002-04-12;修改稿收到日期:2002-07-04基金项目:中以农业研究基金(SIARF)项目、国家自然科学基金项目(30200174)、北京市科委项目(951501800)作者简介:解超杰(1969-),男(汉族),副教授,博士,主要从事小麦抗病育种研究.*通讯联系人.C or res pon dence to:SUN Qi-xin.表1 小麦抗白粉病基因T able1 Resistance g enes to wheat po wdery mildew基因Gene 位点Location来源Sour ce代表品系Cultivar sPm1a7AL普通小麦Axm inster/8*CcPm1b7AL一粒小麦M ocZlatkaPm1c7AL一粒小麦M1NPm1d7AL斯卑尔脱小麦T.sp elta var d uhamelianum T RI2258 Pm25DS未确定Ulka/8*CcPm3a1AS普通小麦As os anPm3b1AS普通小麦Ch ulPm3c1AS普通小麦S on or aPm3d1AS普通小麦Kolibr iPm3e1AS普通小麦AUS6449Pm3f1AS普通小麦M ichigan Amber/8*CcPm3g1AS普通小麦Aris tidePm3h1AS普通小麦Abess iPm3I1AS普通小麦N324Pm3j1AS普通小麦GUS122Pm4a2AL二粒小麦Khapli/8*CcPm4b2AL波斯小麦ArmadaPm5a7BL二粒小麦HopePm5b7BL未确定IbisPm5c7BL印度园粒小麦KolandiPm5d7BL普通小麦IGV1-455Pm5e7BL普通小麦复壮30Pm6T2B/2G提莫菲维小麦Coker747Pm7T4BS.4BL-2R#1L黑麦(Ros en)T ran secPm8T1BL/1RS黑麦(Petku s)Amb as sadorPm97AL普通小麦Normandie(Pm1,P m2)Pm101D普通小麦Norin4Pm116BS普通小麦Ch ines e Sprin gPm12T6BS-6SS.6S L拟斯卑尔脱山羊草Line#31(W embley*6/Ae.sp eltoid es) Pm13T3BL.3BS-3S l#1S高大山羊草R1AT3DL.3DS-3Sl#1S R1DPm146B普通小麦Norin10(P m15)Pm157DS普通小麦Norin10(P m14)Pm164A野生二粒小麦T.d icoccoides CL1060025Pm17T1AL/1R#2S黑麦(Insave F.A.)AmigoT1BL/1R#2S T AM107Pm197D粗山羊草XX186Pm20T6BS/6RL黑麦(Prolific)PI583795Pm21T6AL/6VS簇毛麦R137,R55等Pm221D普通小麦VirestPm235A普通小麦Line81-7241Pm241DS普通小麦Ch iyacaoPm251A野生一粒小麦NC96BGTA5Pm262BS野生二粒小麦T TD140Pm27T6B/6G提莫菲维小麦146-155-TPm281B未确定M eriPm297DL卵穗山羊草PovaPm305BS野生二粒小麦87-1/C20//2*8866Pm31(M lG)6AL野生二粒小麦G-305-M/781//Jing411*3 Pm2:Pug sley和Carter[11]发现前苏联地方品种Ulka中带有不同于P m1的抗病基因,后来U lka 中的抗病基因被定名为P m2[12].M cIntosh和Bak-er[13]将P m2定位于5D染色体短臂上.Nyquist[14]发现在提莫菲维小麦(T.timo p hee-vii,2n=4x=28,AAGG)的六倍体小麦衍生系 CI12633 中至少有两个抗白粉病基因,其中一个显性基因为Mlx.Brig gle[15]根据U lka和CI12632 (CI12632为CI12633的姊妹系)测交分析及Ulka/ 8*Cc和CI12632/8*Cc的抗性比较结果,认为P m2和Mlx是等位基因,而其中另一个基因后来证明是P m6.关于P m2的来源有不同的报道.Bennett[7]分析了CI12632和CI12633的系谱(T.timo p heevii//8235期解超杰,等:小麦抗白粉病基因Illino is No.1/Chinese Spring),认为P m2的来源未知.因为Illinois No.1和Chinese Spring不含有Pm2,而提莫菲维小麦无D组染色体,其中的抗病基因不可能位于5D染色体上.谢皓,陈孝[4]和贾继增[2]认为P m2来自于CI12632和CI12633系谱中的提莫菲维小麦,经易位到染色体5DS上.齐莉莉等[16]根据与已知白粉病抗性基因比较的抗谱分析表明,阿拉拉特小麦(T. ar aratum,2n=4x=28,AAGG)携有主效抗病基因Pm2.阿拉拉特小麦和提莫菲维小麦具有相同染色体组,该结论似乎与谢皓,陈孝[4]和贾继增[2]的推论相符.然而,Lutz等[17]通过抗谱分析、染色体定位以及等位性测验证明合成六倍体小麦 XX194 (T. durum'M oro ccos183'/A e.squarrosa'AE457/78')含有Pm2基因,抗病基因来自于方穗山羊草(A e. squar rosa,2n=2x=14,DD).Lutz等[17]还介绍Pugsley[18]发现澳大利亚小麦选系 Javelin-325 带有Mlx(即P m2)基因,该基因来自 Sears 合成六倍体小麦(T.dur um cv'Ium illo'/A e.squar rosa)中的方穗山羊草,因为Ium illo是感病品种.这些结果表明P m2很有可能是来自方穗山羊草.尽管如此,仍不能排除P m2可能是自发突变产生的[17].Pm3:Br ig gle[12]在对23个小麦抗病品种(系)的研究中发现并命名了该基因.以后Br ig gle[15]又进一步证明该基因位点具有3个等位基因,即P m3a, Pm3b,P m3c,载体品种分别为Asosan,Chul和Sonora.单体及端体分析结果表明,这些抗病基因位于1AS上[19].Heun和Fischbeck[20]发现小麦品种Ko libri中含有抗病基因Mlk,Zeller等[21]将其定位到1A,并证明是P m3的复等位基因,定名为Pm3d.目前Pm3位点已经发现了10个等位基因[8],其载体品种分别是澳大利亚品种A US6449(P m3e)、美国品种Michig an Amber/8*Cc(P m3f)、法国品种Aris-tide(P m3g,即M lar)、埃塞俄比亚硬粒小麦和德国春小麦品种Amor杂交组合的六倍体小麦衍生系Abessi(P m3h),来自尼泊尔的品系N324(Pm3i)和来自俄罗斯的品系GUS122(Pm3j).Pm4:该位点目前有2个等位基因(P m4a和Pm4b).据Bennett[7]介绍,Briggle[22]首先发现在印度广泛用作育种亲本的二粒小麦(T.dicoccum,2n =4x=28,AABB)品系Khapli的抗病性.后来Br ig gle[23]将Khapli中的抗病基因转移到普通小麦品种Chancellor中,命名为P m4,以后发现了另一个等位基因,改名为P m4a[24].P m4b(原名Mle)来源于四倍体波斯小麦(T. carthlicum,2n=4x=28,AABB),带有Pm4b的品种有ELS,T P229,Arm ada等.The T T等[24]发现Pm4a和Mle(即Pm4b)都位于2AL上,表现出等位基因或紧密连锁的遗传特性.遗传研究表明Khapli带有3个抗病基因,而Khapli/8*Cc只含有P m4a,说明从Khapli只转过来一个抗病基因[7].这种现象在从近缘物种向小麦中导入抗病基因的过程中很普遍.P m5:Lebsock和Brigg le[25]将小麦品种H ope 具有的一个隐性抗白粉基因命名为P m5,该基因苗期不能充分表达,4～5叶期接种则抗性表现明显.因为Hope是感病普通小麦品种Mar quis和二粒小麦Yaroslav的杂交后代,因此推断P m5来源于二粒小麦Yaroslav[7].Hope的抗白粉病基因定位在染色体7BL上[26].Ho pe染色体7BL上除有Pm5以外,还有一个隐性抗秆锈基因Sr17和一个不完全显性抗叶锈基因Lr14a.Hope曾在小麦育种中广泛应用.抗白粉病基因M li最初在法国冬小麦品种Flanders等中发现[7].后来Heun和Fischbeck[27]证明Mli抗性方式与Pm5相似,并将Mli定名为P m5.但是M cIntosh[28]指出P m5是由二粒小麦转移到普通小麦Hope,而M li却没有这种明显的起源关系.Hsam等[29]证明P m5为一复等位基因位点,包括Hope中的Pm5a、Ibis等中的P m5b(即M li)、印度圆粒小麦(T.sp haerococcum v ar. R otund atum)品系Kolandi中的P m5c和普通小麦品系IGV1-455中的P m5d,而P m5d的来源是中国的普通小麦.最近,Huang等[30]在中国小麦品系复壮30中又鉴定出Pm5e.P m6:来源于提莫菲维小麦,曾被认为是最成功和广泛应用的抗白粉病基因[7],由Jorg ensen和Jensen[31]正式命名.据Bennett[7]介绍,Nyquist[14]发现在CI12633中除含有Mlx(即P m2)外,还有一个独立的基因,与M lx共同控制成株抗性,并与抗秆锈病基因S r36(原定名为Sr9c)紧密连锁,Jor-g ensen和Jensen[32]证明这两个基因分别是Pm2和P m6.P m6与S r36紧密连锁,而S r36定位于2B 上,因此P m6也位于2B上.Friebe等[33]根据C-分带证明P m6位于易位染色体T2B/2G上,易位部分824西 北 植 物 学 报23卷涉及2B的两个臂.陶文静等[34]也通过分子标记证明了这一结论.Pm7:该基因是通过辐射易位由黑麦(Secale cer eale,2n=2x=14,RR)品种Rosen导入普通小麦T ransec中[35],其中黑麦染色体2RL的一段易位到小麦染色体4BL上,形成T4BS.4BL-2RL易位系[33].该易位片段还带有抗叶锈基因L r25,这两个基因表现为紧密连锁.Pm7并没有得到广泛应用.Pm8:来源于黑麦,位于黑麦品种Petkus染色体1RS上.通过1R(1B)代换系(如洛夫林13等)或1BL/1R#1S易位系(如洛夫林10等)转入普通小麦中.在这个黑麦染色体片段上除有P m8外,还有抗条锈基因Yr9,抗叶锈基因L r26,和抗秆锈基因Sr31[28,33].转入小麦的黑麦Pm基因中,P m8的应用最为广泛.Pm8最初推广时表现高抗,但随着与之相对应的致病性小种频率迅速上升,其抗性很快丧失[7]. Pm8在我国曾经广泛地被应用,现在也已丧失抗性[1].Pm8基因的表达受到其它基因的抑制,包括位于小麦1A S染色体上的抑制基因Su P m8[36]和位于小麦品种Caribo染色体7D上的Pm8与Pm17的抑制基因[37].Pm9:法国品种No rmandie中除含有Pm1和Pm2外,还有P m9.Schneider[38]对Pm9与P m1和Pm2相对遗传关系的研究发现,Pm9呈单基因隐性遗传,与P m2互相独立,而与Pm1连锁,遗传距离为8.5cM.已知Pm1位于7AL,因此判断P m9也在7AL上.含有该基因的品种还有Mephisto,An-field,Pom pe和Ring.Pm10和Pm11:To sa等[39,40]利用小麦白粉菌(E.graminis f.sp.tritici)和冰草白粉菌(E. graminis f.sp.agr o p y ri)的杂交后代,在No rin4, Norin26等品种中检测出P m10基因,在中国春中检测出P m11,并分别定位到染色体1D和6BS上.Pm12:来源于拟斯卑尔脱山羊草(Ae.sp el-toides,2n=2x=14,SS),通过易位转移到小麦品种Wembley中,获得抗病材料品系31号(line# 31).P m12最初定位于小麦染色体6A,但Jia等[41]利用分子标记将该基因重新定位到易位染色体T6BS-6SS.6SL上.Pm13:来源于高大山羊草(Ae.longissim a(= T.longissimum),2n=2x=14,SlSl).Ceoloni等[42]通过诱导部分同源染色体配对重组方法将高大山羊草3Sl短臂携带的显性抗白粉病基因Pm13转移到中国春中,获得了R1A,R1D等抗病易位系.单体分析和C分带研究表明,R1A等易位系中Pm13位于易位染色体T3BL.3BS-3S l#1S上,而R1D等易位系中该基因位于易位染色体T3DL.3DS-3 S l#1S上[33].P m14和Pm15:T osa等[43]用小麦白粉菌和冰草白粉菌的杂交种,发现在Akabozu,Norin10等品种中含有P m14和P m15基因,分别定位于6B和7D染色体上.P m10,Pm11,P m14和Pm15只对冰草白粉菌表现抗性,不抗小麦白粉菌,在小麦抗病育种中无实际利用价值.P m16:Reader和M iller[44]将野生二粒小麦(T.d icoccoides,2n=4x=28,AABB)材料CL1060025中的抗白粉病基因Pm16转入普通小麦Norman 中,并将该基因定位于4A染色体上.P m17:来源于黑麦品种Insave F.A..经用X 射线处理普通小麦与八倍体小黑麦杂交种,获得了抗麦二叉蚜的品种Amigo[45].在Am igo中,黑麦染色体1RS易位到1A上,形成T1AL.1R#2S易位系.在这段1RS片段上不仅含有抗麦二叉蚜基因G b2,还含有一个与Pm8不同的抗白粉病基因,命名为P m17[46,47].后来Hsam等[48]通过 Am ig o/Helios(T1BL. 1RS) 杂交组合将P m17转移到易位系T1BL.1RS 上,并证明P m17和P m8之间是等位基因关系[49].近来发现小麦品种Car ibo的7D染色体上带有P m8与P m17的抑制基因[37].P m18:据Hsam等[10]介绍,德国普通小麦品系W eihenstephan Stam m M1(M1)携带有P m4b基因,而在1986年得到一个M1种子样品,经鉴定这个样品的抗谱与原M1不同,为一新抗病系,取名为W eihenstephan Stamm M1N(M1N),其中含有的抗病基因被定名为Pm18[50].Hsam等[10]通过单体分析发现该基因位于7A染色体上,并与Ax minis-ter/8*Cc中的P m1a基因等位,据此将其重新命名为Pm1c.Hsam等[10]认为Pm1c可能来自于一粒小麦.P m19:Lutz等[17]根据对小麦白粉菌不同生理小种的抗性反应,在合成六倍体小麦XX186中发现了一个显性抗病基因,并利用单体分析证明该基因位于7D染色体上.由于在7D上尚未有已知P m基因,因此命名该基因为P m19.XX186的组合为8255期解超杰,等:小麦抗白粉病基因T.durum v ar Santa M arta'/A e.squarrosa BGRC 1458',由于硬粒小麦'Santa M ar ta'是感病品种,所以判断P m19来自方穗山羊草.Pm20:来自黑麦品种Prolific.Friebe等[51]用小麦-黑麦6RL(6D)单体代换系M S6RL(6D)与小麦-黑麦T6BS.6R#2L纯合易位系品种T AM104杂交,通过同源重组将抗白粉病基因Pm20转移到易位染色体T6BS.6R#2L上,所获得的抗病材料命名为 KS93WGRC28 [52].据Fr iebe等[33]报道,在 KS93WGRC28 中的易位染色体形成T6BS.6R# 2L-6R#3L.C分带结果表明,Pm20位于Pro lific的6R#3染色体长臂的端粒附近.Pm21:这是中国学者发现的第一个小麦抗白粉病基因[53].该基因位于簇毛麦(H ay naldia villosa, 2n=2x=14,VV)6V染色体短臂,通过染色体添加、代换和易位转入普通小麦中[53,54].目前应用的多为T6AL.6VS易位系,李辉等[54]还获得了T6DL.6VS易位系.Pm22:Peusha等[55]通过抗谱分析发现小麦品种Virest带有新的抗白粉病基因,单体分析表明该基因为一显性主效基因,位于1D染色体上,命名为Pm22.含有该基因的还有意大利品种Elia,Est M ottin,Ov est和T udest等.Pm23:四川农业大学杨足君(YANG Z J)和任正隆(REN ZH L)报道在小麦品系81-7241中发现新的抗白粉病基因,并定位于染色体5A上,命名为Pm23[28].Pm24:Huang等[56]发现中国小麦地方品种Chiy acao中含有一个与已知P m基因抗谱不同的新基因,通过单体分析将Chiyacao中的这个显性主效抗病基因定位于6D上,并命名为P m24.但是后来通过分子标记证明该基因应该是位于1DS[57].Pm25:Shi等[58]从野生一粒小麦(T.m onococ-cum var.boeoticum,2n=2x=14,AA)向普通小麦转移了一个显性抗白粉病基因,命名为P m25.遗传分析表明该基因与Pm3a连锁,据此将该基因定位于1A上.Pm26:来自野生二粒小麦.Rong等[59]将以色列野生二粒小麦T TD140中的一个抗白粉病基因转移到普通小麦Bethlehem中.该基因为隐性,命名为P m26,通过对一系列染色体臂代换系(chro mo-some-arm substitution lines CASLs)进行RFLP标记被定位于2BS上.P m27:Ja¨r ve等[60]通过杂交和回交从提莫菲维小麦材料K-38555向普通小麦146-155中转移了一个显性抗白粉病基因,获得纯合抗病系146-155-T.单体分析和分子标记研究表明该基因来自6G并易位到6B染色体上,命名为P m27.P m28:Peusha等[61]根据单体分析结果证明芬兰-爱沙尼亚春小麦品种M eri带有一个显性抗白粉基因,定位于1B.该基因与原来1B上来自黑麦的P m8和P m17的抗性表现不同,命名为P m28.目前尚未确定P m28的来源.P m29:Zeller等[62]从普通小麦Poros的卵穗山羊草(A e.ov ata,2n=4x=28,UU MM)二体异附加系POS的后代中,选出了具有42条染色体的抗白粉病品系Po va,遗传分析表明Pova带有一个显性抗白粉病基因.该基因位于小麦染色体7D上,但与7D上的P m19基因独立分离,进一步的分子标记分析将其定位于7D长臂,正式命名为P m29.P m30:中国农业大学小麦组通过杂交和回交将以色列野生二粒小麦材料C20的抗白粉病基因导入普通小麦遗传背景中,并通过分子标记将该基因定位在染色体5BS上,现在该基因被正式命名为P m30[63].这是由中国学者发现和定名的第3个小麦抗白粉病基因.P m31(MlG):来源于以色列野生二粒小麦材料G-305-M.这是中国农业大学小麦组得到的第二个来自以色列野生二粒小麦的抗白粉病基因,微卫星分子标记表明该基因位于小麦染色体6A长臂,临时命名为M lG[64],现被定名为P m31.抗白粉病基因在小麦染色体上广泛分布,目前,除2D,3A,4D,6D染色体外,其余染色体上都有抗白粉病基因位点存在.这些抗白粉病基因大多表现显性遗传,少数为隐性(如Pm5,P m9,P m26等).许多Pm基因来源于小麦近缘种属,因此利用外源抗病基因可以极大地丰富普通小麦抗病基因资源,同时也有助于扩大小麦育种的遗传基础.参考文献:[1] YANG Z M(杨作民),T ANG B R(唐伯让),SHEN K Q(沈克全),XIA X C(夏先春).A strategic problem in w heat resis tance b reeding-building and utilization of sources of second-line res istance 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olecular marker s[J].Journal of T ritic eae Cr ops (麦类作物学报),1999,19(3):11-14(in Ch ines e).[7] BENNET T F G A.Resis tance to pow dery mild ew in w heat:a review of its us e in agriculture and b reeding program s[J].Plant P athol ,1984,33:279-300.[8] ZE LLER F J ,HSAM S L K .Progress in breeding for resistance to p ow dery m ildew in common wh eat (T riticum aestiv um L .)[A ].In :Slinkard AE (ed )Proc 9th Int Wheat Genet S ymp [C ].Sas katoon ,Sask .,Canada ,U nivers ity Ex tention Pr ess ,1998,Vol .1:178-180.[9] SEARS E R,BRIGGL E L W.M apping the gene P m 1for res istance to Erysiph e gram inis f.sp.tritici on ch romosome 7A of w heat[J].Crop Sc i ,1969,9:96-97.[10] HSAM S L K,HU ANG X Q,ERNST F,HARTL L,ZELL ER F J.Chromos om al location of genes for res istance to pow der y mildewin common w heat (Tr iticum ae stiv um L.em.Th ell.)5.Alleles at the P m 1locus [J].T heor Ap pl Genet ,1998,96:1129-1134.[11] PULGSLEY A T ,CART ER M V.Resis tance of tw elve varieties of T riticum v ulgar 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