猕猴桃是猕猴桃科(Actinidiaceae)猕猴桃属(Actinidia Lindl.)落叶藤本果树,该属有54 个种和21 个变种,共75 个分类单元[1]。中国是猕猴桃属植物的原产地,野生资源丰富,地域分布广泛。自20世纪被人工驯化以来,该产业在世界范围内的栽培面积不断扩大[2],因其果实风味独特、营养丰富而深受人们喜爱。随着产业的发展,不同区域间引种频率增加,生产中的细菌性溃疡病也越发严重,对产业造成了严重危害[3]。自1984年日本首次出现该病以来[4],随后短短几年内,迅速成为威胁世界猕猴桃产业的毁灭性病害[5]。中国、新西兰、意大利、韩国等世界各大猕猴桃产区深受其害[6-9]。中国作为猕猴桃生产大国,在四川、安徽、陕西等主产省份也早已出现,并逐渐向其他栽培地区蔓延[10]。因此,开展溃疡病相关研究,了解其致病机制和作用机制、探索病害防控技术,选育抗病新品种,对助力产业高质量发展具有重要意义。随着现代测序技术的发展,分子生物学技术为猕猴桃溃疡病研究提供了一种有效途径。本文旨在综述猕猴桃溃疡病抗性研究方面最新进展,为该产业抗性育种和高效防控技术研发提供理论基础。
猕猴桃细菌性溃疡病在一年内有冬末春初和秋季两个发病高峰[11]。主要危害叶片、花、果实和枝蔓,病菌侵染多从伤口、皮孔、落叶痕、枝条分叉等部位开始[12]。枝蔓发病初期呈水渍状,后病斑扩大、颜色加深;早期病斑部位流白色黏液,不久转为铁锈红色,剥开病斑皮层后,可见韧皮部腐烂,木质部变黑[13-14]。叶片染病后会形成不规则的黑色斑点,并带有黄色晕圈。溃疡病具有隐蔽性、传染性、暴发性和毁灭性等特点,一旦发生,轻则减产、重则毁园。2010年新西兰首次发现溃疡病之后,感病果园数量迅速增加,到2012 年占全部果园的37%[15],对该国猕猴桃产业造成严重危害。研究发现,不同栽培品种对溃疡病的抗性不一,一些商品性很好的品种抗病能力却很弱,如新西兰的黄肉品种Hort16A 以及中国的红心品种红阳[16]。
猕猴桃溃疡病病菌主要借助风、雨等在果园内和果园间迅速散播,低温潮湿的环境可极大地促进病原菌繁殖,一些不规范的农艺操作以及低温、冷害、冻害等极端气候现象也有助于病菌的进一步传播和流行[17]。苗木、接穗、工具、人员等都可能是携带病菌的重要载体,此外,昆虫[18]、花粉[19]、非猕猴桃属植物[20]等也可以作为中间媒介或中间寄主来实现病菌的传播和侵染。
丁香假单胞杆菌猕猴桃致病变种(Pseudomonas syringae pv. actinidiae,Psa)是引起猕猴桃细菌性溃疡病的致病菌[4],其关键致病因子是存在T3SS(type Ⅲsecretion system)的蛋白分泌系统[10]。该分泌系统能分泌多种有毒效应因子(如HopZ5、Avr-Rpm1等)来破坏植物的免疫防御反应,使病原体快速适应宿主环境[21-22]。Psa 的整合共轭元件(integrative and conjugative elements,ICEs)是可移动元件,赋予其新的表型,且常被认为是细菌病原体产生耐药性进化的机制[22-23]。研究人员通过对目前收集到的Psa 株系进行基因组比较、系统进化和起源分析等,可将目前已发现的Psa 株系分为6 类[24]:第一类是在日本和意大利海沃德品种上采集到的病原菌Psa1[25];第二类是在韩国发现的病原菌Psa2[26];第三类是于2008年首次在意大利发现,并对世界各国产业造成毁灭性伤害的Psa3[27];第四类是在新西兰发现的病原菌Psa4,该类病菌致病能力较弱,仅引起叶斑[9],与前三类存在明显不同,虽然Psa4是从猕猴桃属植物上分离出来的,但由于其表型、遗传和系统发育不同,后将其更名为Pseudomonas syringae pv.actinidifoliorum(Pfm)[28];第五类和第六类是在日本发现的Psa5[29]和Psa6[30]。在已有的研究报道中,并没有发现对溃疡病有效的治愈方法,生产上频繁使用的药物防治使得Psa菌株对铜和链霉素产生了抗药性[31]。因此,选育抗性强的品种来增强自身的抗病性,是解决溃疡病危害最直接的方法。
不同种类和品种的猕猴桃对Psa的抗病性有所不同,研究人员分别从形态结构、生理和分子水平等方面对其抗病能力进行了研究分析。对24 个不同种类或品种的猕猴桃进行抗性评价,不同种类抗性由强到弱的顺序依次为:毛花猕猴桃(A.eriantha)、美味猕猴桃(A.chinensis var.delicious)、中华猕猴桃(A.chinensis)[32]。利用29个猕猴桃种或品种(系)进行抗性评价,在离体枝条接种Psa 病原菌6 d 后,中华猕猴桃红阳、6-65、2-72就开始发病并溢出少量白色黏液,而接种21 d 后软枣猕猴桃(A.arguta)和毛花猕猴桃才开始出现病斑且病斑直径明显小于中华猕猴桃,体现出抗性相对较强[33]。通过采用离体枝条和叶片进行人工接种Psa病原菌的方法对51份软枣猕猴桃进行抗性鉴定,结果显示51份资源中高抗33 份、中抗18 份,无高感、中感和感病种质,体现了软枣猕猴桃具有较好的抗性[34]。采用室内和田间接种方法分别对12个和23个品种进行抗性鉴定,总体抗性趋势为软枣猕猴桃和毛花猕猴桃强于美味猕猴桃,而中华猕猴桃表现最差[35]。近年来,研究人员选育出了多个抗病新品种,如先沃五号[36]、华金3号[37]、金塘一号[38]等。
猕猴桃的组织结构及活性成分与抗病性有一定关系。通过比较不同抗性资源的叶片和枝条结构发现,叶片气孔密度和长度、枝条皮孔密度和长度与病情指数呈显著正相关[39-40]。接种病原菌后,抗病品种金魁的枝条、叶片中可溶性糖和木质素含量也显著高于感病品种金丰[41]。在不同品种中,抗病性越强,叶片中可溶性蛋白质和酚类物质含量越高[42]。韧皮部蔗糖代谢的增加可能引起免疫应答相关酶的增加,但不同品种的抗病能力与蔗糖的含量呈负相关[43]。还有研究发现,抗性砧木也能够有效增强接穗的抗病性[44]。
综上所述,不同猕猴桃种类、品种(系)的抗性情况存在差异,但总体抗性趋势为软枣猕猴桃和毛花猕猴桃强于美味猕猴桃,且强于中华猕猴桃。这种抗性差异可能与生理生化特性及遗传背景等因素有关。因此,准确评价和筛选出高抗猕猴桃细菌性溃疡病的品种(系)至关重要。
分子标记(Molecular Markers)是以个体间核苷酸序列变异为基础的遗传标记,能在DNA水平上直接反映遗传的多态性[45]。与其他生物遗传标记如生化标记、细胞学标记和形态学标记相比,具有不受个体发育时期和外界环境影响、多态性高、易于检测等显著的优点[46]。分子标记技术的发展主要经历了3个阶段,第1 个阶段是以Southern 杂交为基础的限制性片段长度多态性(Restriction fragment length polymorphism,RFLP)标记;第2 个阶段是以PCR 反应为基础的随机扩增多态性(Random amplified polymorphic DNA,RAPD)、扩增片段长度多态性(Amplified fragment length polymorphism,AFLP)、简单重复序列(Simple sequence repeat,SSR)标记;第3个阶段则是以测序为基础的单核苷酸标记(Single nucleotide polymorphism,SNP)[47]。目前,SNP技术已被广泛应用于物种亲缘关系进化分析、种质资源鉴定保存、分子遗传图谱构建、植物抗病基因定位及遗传育种等方面。
猕猴桃属植物种类繁多,地域分布范围广泛,因此对其种质资源进行准确鉴定是合理利用的基础。利用分子标记技术筛选抗性优异种质资源已广泛应用于大豆[48]、水稻[49]、番茄[50]、黄瓜[51]等植物的抗性育种中,猕猴桃中也有少量应用。通过利用ISSR标记进行遗传多样性分析,发现不同品种对溃疡病的抗病能力与遗传有关,且各品种抗性强弱分组与ISSR聚类组有明显相关性,表明了该技术可用于辅助选育抗溃疡病品种[52]。采用SSR技术结合BSA分析方法对杂交F1代群体及40 份资源进行抗病基因(PR)分子标记筛选,获得了与抗病基因连锁的SSR 分子标记UDK97-428116[53]。利用SCoT分子标记确定了9个中华猕猴桃红肉品种的亲缘关系并筛选出较抗溃疡病品种[16]。对6 个品系进行抗病相关的RAPD 分析,结果发现,抗病品系都有一条1458 bp的DNA片段,而感病品系均无该条带[54]。通过构建表达序列标签(EST)文库,并基于同源序列设计EST引物,成功鉴定出部分参与基础防御途径的基因,为猕猴桃抗性育种提供了基础[55]。
遗传连锁图谱是通过遗传重组交换结果进行连锁分析所得到的基因或分子标记在染色体上相对位置的线性排列图[56],是数量性状定位(quantitative trait locus,QTL)、分子标记辅助选择育种等研究的理论依据。因此,构建高密度的遗传连锁图谱是植物进化过程、遗传育种及功能基因组学等研究的重要环节。
利用果树基因组中的多种分子标记技术构建高精度的遗传图谱已逐渐成为该学科研究的重要内容。近年来,已在枣[57]、苹果[58]、葡萄[59]、柑橘[60]、樱桃[61]、梨[62]等果树作物中构建了分子遗传图谱。目前,猕猴桃相关分子遗传图谱的构建也在逐步完善。自2001年Testolin等[63]利用AFLP和SSR标记,分别构建中华猕猴桃和硬齿猕猴桃的遗传图谱以来,目前已报道构建的遗传图谱共有11 组(表1)。这些遗传图谱对猕猴桃相关性状定位研究具有重要意义。在目前已构建的图谱中,应用于溃疡病研究的只有两组,分别是2019年以二倍体中华猕猴桃为亲本和2020 年以四倍体中华猕猴桃为亲本构建。在得到的二倍体中华猕猴桃高密度遗传图谱中,通过QTL 定位技术在Hort16A 的27 号染色体上定位到了一个主效QTL,在P1 中定位到了6 个微效QTLs(3、14、15、22、24 和28 号染色体),并验证27、14、22、28之间的互作效应,结果表明,F1代对溃疡病抗性的增强是Hort16A 和P1 上的QTL 加性效应引起的[64]。随后,在四倍体中华猕猴桃遗传图谱中定位到了4个抗Psa的QTLs(LG1、LG2、LG4和LG7);其中,高抗亲本包含3 个QTLs(LG1、LG4 和LG7),耐病亲本包含1 个QTL(LG2)[65]。主要抗性基因与数量抗性因子的结合,可以保持抗性品种的持久性,QTL的加性效应增强了子代对病原菌的抗性[66-67]。
表1 猕猴桃遗传图谱
Table 1 Genetic linkage map of kiwifruit
年份Year 2001标记类型Markers mapping SSR、AFLP参考文献Reference[63]2009 SSR [68]2013 SNP [69]2015 SSR、SNP [70]2015 SNP [71-72]2017 SNP [73]2019 SNP [64]2020 SNP [65]2021 SNP应用Application性别性状Sex traits性别性状Sex traits辅助基因组拼接Assisted genome splicing辅助基因组拼接Assisted genome splicing性别、果实性状Sex and fruit traits基因组评价Genomic evaluation溃疡病性状KBC traits溃疡病性状KBC traits果实性状Fruit traits[74]2021 SNP [75]2023作图群体Mapping parents中华猕猴桃(2x)×硬齿猕猴桃(2x)A.chinensis(2x)×A.callosa(2x)中华猕猴桃(2x)×中华猕猴桃(2x)A.chinensis(2x)×A.chinensis(2x)中华猕猴桃(2x)×毛花猕猴桃(2x)A.chinensis(2x)×A.eriantha(2x)中华猕猴桃(2x)×中华猕猴桃(2x)A.chinensis(2x)×A.chinensis(2x)山梨猕猴桃(2x)×中华猕猴桃(2x)A.rufa(2x)×A.chinensis(2x)中华猕猴桃(2x)×中华猕猴桃(2x)A.chinensis(2x)×A.chinensis(2x)中华猕猴桃(2x)×中华猕猴桃(2x)A.chinensis(2x)×A.chinensis(2x)中华猕猴桃(4x)×中华猕猴桃(4x)A.chinensis(4x)×A.chinensis(4x)美味猕猴桃(6x)×美味猕猴桃(6x)A.chinensis var.delicious(6x)×A.chinensis var.delicious(6x)中华猕猴桃(4x)×黑蕊猕猴桃(4x)A.chinensis(4x)×A.melanandra(4x)软枣猕猴桃(4x)×软枣猕猴桃(4x)A.arguta(4x)×A.arguta(4x)SNP果实性状Fruit traits性别性状Sex traits[76]
溃疡病可能是由数量性状引起[64],而数量性状通常会被多个基因共同作用,且易受到环境因素的影响,遗传情况复杂,因而数量性状的研究非常困难。随着分子标记技术的发展,利用遗传标记和QTL间的遗传连锁现象,可以确定QTL在染色体上的位置和效应,从而提高育种效率、加快新品种选育。
当植物受到病原菌的感染时,会激发植物的天然防御系统,使植物产生抗病反应。植物的天然免疫系统可分为两个水平,第一是通过植物细胞表面的模式识别受体(Pattern Recognition Receptors,PRRs)识别病原微生物保守成分(Pathogen Associated Molecular Patterns,PAMPs),从而激活与病原相关分子模式成分触发的免疫反应(PAMP-Triggered-Immunity,PTI)[77],第二是病原微生物释放的效应因子触发的免疫反应(Effector-Triggered-Immunity,ETI)[78]。植物强大的免疫系统能抵御大多数病原微生物的侵染,但是丁香假单胞杆菌等部分病菌能向寄主植物细胞内注入毒力蛋白,抑制植物免疫力,从而引起毁灭性病害[79]。核苷酸结合富亮氨酸重复受体(nucleotide-binding and leucine-rich repeat receptors,NLRs)是植物最大的免疫受体家族[80],来自拟南芥和本氏烟草的NLR 蛋白ZAR1 能识别HopZ5 并触发细胞死亡[81]。目前,已鉴定出多个识别Psa效应因子的NLR 蛋白,如RPA1[82]、NbPTR1[83]等,这些蛋白在植物抗病反应中具有关键作用。
NHL(NDR1/HIN1-like)基因家族成员在抵御丁香假单胞杆菌的侵染时具有积极作用。如拟南芥NHL 基因家族的成员NHL2 和NHL3 在抵御丁香假单胞杆菌侵染中发挥重要作用[84];在采用不同丁香假单胞杆菌处理拟南芥时,发现NHL3 基因的表达量均显著增加,且过表达该基因增强了植株对Pseudomonas syringae pv.tomato DC3000(Pst)病菌的抗性[85]。NHL基因在其他病原菌防御反应中也发挥了重要作用。如拟南芥NHL10 基因能调控黄瓜对花叶病毒(CMV)的超敏反应[86];在马铃薯中过表达NHL 基因成员StPOTHR1 增强了植株对疫霉病(Phytophthora infestans)的抗性[87]。
利用转录组学的方法对Psa侵染后猕猴桃中的基因表达进行分析,发现免疫系统PTI、ETI、HR 中多个抗性基因的表达受到诱导,它们可能通过调整代谢过程,并改变次级代谢产物的产生以抑制Psa的生长[88]。研究发现,在受到Psa侵染后,高抗品种华特中编码Pti1 和RPS2 的效应受体及参与水杨酸信号通路的NPR1、TGA 和PR-1 基因均显著上调表达[89]。PR-1基因能在溃疡病菌诱导下显著表达,且其过表达能增强烟草对溃疡病的抗性[90]。AcTGA07基因的过表达显著增强了猕猴桃的抗性,且TGA转录因子能特异性地结合PR 基因的启动子区域并与NPR 蛋白相互作用[91]。NPR1 同源基因AeNPR1a在烟草中过表达后,相比与野生型株系其Psa 和Pst感染症状明显减轻,转基因烟草抗性显著增强;与接种Pst 的拟南芥npr1-1 突变体植株相比,NPR1a回补株系抗性显著增强[92]。使用壳聚糖处理可诱导猕猴桃防御相关基因(PR1、PR5)表达,使其产生系统获得性抗性(systemic acquired resistance,SAR)[93]。
自1984 年在日本首次发现猕猴桃细菌性溃疡病以来,世界各产区深受其害。经过多年的研究积累,对该病害的发病规律与传播途径、致病菌Psa的致病机制、不同种类或品种猕猴桃的抗病特性等研究均已取得一定成果,但一直未有有效的防治方法。因此,在今后的生产实践中应选择抗性强的品种、加强苗木或花粉等媒介物的检疫监测、提高果园栽培管理农艺措施,这对早期病害预防和减少种植者的经济损失具有重要作用。
中国猕猴桃种质资源丰富,倍性多样,充分利用这些宝贵资源进行抗病新品种培育具有重要意义,但传统实生选种或杂交育种不仅耗时长而且目标性状聚合困难,后代易出现性状分离。随着现代测序技术的发展,通过QTL定位、转录组等方法,在猕猴桃中已经挖掘出许多与抗溃疡病相关的基因和转录因子等,这不仅为抗病材料的选育提供了重要的基因资源和分子标记,还可在早期进行目标性状的预判和选择,极大地提高育种效率。但是由于猕猴桃遗传背景复杂,分子标记技术还无法很好的应用于育种实践,今后还需加大稳定性较好的分子标记开发力度。
近年来QTL 定位技术在许多果树重要性状鉴定方面的应用取得重大进展并已在猕猴桃树种中实施,猕猴桃定向高效精准育种策略成为可能。但该树种染色体基数大、倍性复杂且杂合度高,因此构建高密度的遗传图谱较为困难,QTL 精细定位实施还很少。今后应针对不同研究目的,将图谱构建与杂交育种工作相结合,构建高层次和高实用价值的遗传连锁图。
随着基因组、转录组、代谢组等各类技术的发展以及猕猴桃多倍体基因组信息的不断完善,可以建立不同倍性组合的远缘杂交新基因导入群体开展QTL 定位及抗性基因发掘工作,从而开发出溃疡病抗性相关分子标记,为猕猴桃抗性新品种培育提供新途径。虽然已报道许多与抗性相关基因,但整体而言对猕猴桃抗病分子机制及调控网络研究存在不足,在抗病关键基因挖掘方面的研究也有待进一步深入。今后应注重抗性基因的功能验证,以及关键基因在调控网络中的作用,结合免疫途径、代谢途径等相关反应,解析其抗病分子机制。
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Research advances on reisistance to kiwifruit bacterial canker