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Oct.  2023
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CHEN Z, DING T, HUANG Z, et al. Compatibility of intraspecific hybridization and screening of powdery mildew resistent germplasm in Lagerstroemia indica[J]. Journal of Sichuan Forestry Science and Technology, 2023, 44(5): 82−88 doi: 10.12172/202212050003
Citation: CHEN Z, DING T, HUANG Z, et al. Compatibility of intraspecific hybridization and screening of powdery mildew resistent germplasm in Lagerstroemia indica[J]. Journal of Sichuan Forestry Science and Technology, 2023, 44(5): 82−88 doi: 10.12172/202212050003

Compatibility of Intraspecific Hybridization and Screening of Powdery Mildew Resistent Germplasm in Lagerstroemia indica


doi: 10.12172/202212050003
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  • Corresponding author: hz7210@126.com
  • Received Date: 2022-12-05
    Available Online: 2023-04-10
  • Publish Date: 2023-10-25
  • Three American Lagerstroemia indica varieties (Red Rocket, Red Fireball and Red Leaf) and two Sichuan native Lagerstroemia indica varieties (Sichuan Pink Flower and Sichuan White Flower) were used as parents for intraspecific hybridization. Three seed traits, including fruit setting rate, seed setting rate and seed germination rate, were investigated, and the heritability, correlation heritability and correlation coefficient of the three seed traits were determined. The progeny were inoculated by the spore suspension infection method, and the hybrid combinations resistant to powdery mildew were screened according to the disease classification of the progeny. The results showed that: (1) There were significant differences in seed and fruit characters of hybrid progeny, and they were significantly lower than those of open pollinated progeny. (2) The seeds of 12 hybrid combinations were harvested, and only 8 of them could germinate. Therefore, parental collocation should be fully considered in Lagerstroemia indica hybrid design. (3) The heritability of fruit and seed traits was all above 0.99, and the correlation heritability of fruit setting rate with seed setting rate and germination rate was all above 0.8. The efficiency of early selection of seed setting rate and germination rate by using fruit setting rate traits was 88.37% and 84.47%, respectively. (4) The powdery mildew resistance test of the offspring of 8 hybrid combinations showed that some hybrid progenies of Sino-American combination was good, while others were very poor. The F1 generation of the hybrid combination "Red Rocket × Sichuan White Flower" showed the best resistance, all the tested progenies were resistant to the disease, and the hybrid combination of "Red Rocket × Sichuan White Flower" showed the second best resistance. However, the F1 generation of "Sichuan Pink Flower × American Red Leaf" had the worst resistance. The hybrid of local varieties represent equilibrium. (5) Initially, 8 immune plants and 25 highly resistant plants were screened out. The hybrid combinations and disease-resistant types of powdery mildew resistance discovered in this study provide the material basis for Lagerstroemia indica resistance breeding.
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    [7] 聂硕,张林,王峰,等. 紫薇杂种F_1代的性状表现与倍性研究[J]. 山东农业科学.,2017,49(01):17−22.
    [8] 谢宪. 紫薇杂交后代的花色、AFLP及抗病分析[D]. 山东农业大学, 2017.
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    [12] 吴际洋,焦垚,叶远俊,等. 大花紫薇与紫薇杂交F_1群体表型评价及分子标记连锁分析[J]. 园艺学报.,2018,45(11):2153−2163.
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    [14] 冯坤,郑青松,俞佳虹,等. 超氧化物歧化酶的遗传特征及其在植物抗逆性中的研究进展[J]. 分子植物育种.,2017,15(11):4498−4505.
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Compatibility of Intraspecific Hybridization and Screening of Powdery Mildew Resistent Germplasm in Lagerstroemia indica

doi: 10.12172/202212050003
  • 1. Ecological Restoration and Conservation on Forest and Wetland Key Laboratory of Sichuan Province, Sichuan Academy of Forestry, Chengdu 610081, China
  • 2. Sichuan Forestry Work Station, Chengdu 610081, China
  • Corresponding author: hz7210@126.com

Abstract: Three American Lagerstroemia indica varieties (Red Rocket, Red Fireball and Red Leaf) and two Sichuan native Lagerstroemia indica varieties (Sichuan Pink Flower and Sichuan White Flower) were used as parents for intraspecific hybridization. Three seed traits, including fruit setting rate, seed setting rate and seed germination rate, were investigated, and the heritability, correlation heritability and correlation coefficient of the three seed traits were determined. The progeny were inoculated by the spore suspension infection method, and the hybrid combinations resistant to powdery mildew were screened according to the disease classification of the progeny. The results showed that: (1) There were significant differences in seed and fruit characters of hybrid progeny, and they were significantly lower than those of open pollinated progeny. (2) The seeds of 12 hybrid combinations were harvested, and only 8 of them could germinate. Therefore, parental collocation should be fully considered in Lagerstroemia indica hybrid design. (3) The heritability of fruit and seed traits was all above 0.99, and the correlation heritability of fruit setting rate with seed setting rate and germination rate was all above 0.8. The efficiency of early selection of seed setting rate and germination rate by using fruit setting rate traits was 88.37% and 84.47%, respectively. (4) The powdery mildew resistance test of the offspring of 8 hybrid combinations showed that some hybrid progenies of Sino-American combination was good, while others were very poor. The F1 generation of the hybrid combination "Red Rocket × Sichuan White Flower" showed the best resistance, all the tested progenies were resistant to the disease, and the hybrid combination of "Red Rocket × Sichuan White Flower" showed the second best resistance. However, the F1 generation of "Sichuan Pink Flower × American Red Leaf" had the worst resistance. The hybrid of local varieties represent equilibrium. (5) Initially, 8 immune plants and 25 highly resistant plants were screened out. The hybrid combinations and disease-resistant types of powdery mildew resistance discovered in this study provide the material basis for Lagerstroemia indica resistance breeding.

  • 紫薇(Lagerstroemia indica)属于千屈菜科,落叶或常绿的灌木或乔木,是各地夏季重要的观赏花木,有花期长、枝干光滑、花色艳丽、种类繁多、萌蘖力强、抗污染等优点,是各地常见的优良园林树种[1, 2],也是川派盆景重要的树种[3]。每年4—5月份紫薇易受到白粉病侵染,多危害新叶使其畸形、嫩枝干枯,严重影响后续开花进程,9—10月危害新叶、老叶和花穗,使叶片畸形、卷曲、皱缩,光合作用下降,导致叶变黄、早落,加速叶片的衰老,被浸染的花穗表面蒙上一层白色粉末,影响其开花生理进程,导致花苞不开放直接萎缩、褐化、掉落,严重影响其观赏价值及品质[4]

    杂交育种是培育紫薇新品种的有效途径[5, 6]。聂硕[7]、谢宪[8]等通过二倍体和四倍体杂交,获得了高抗白粉病的三倍体紫薇,表明杂交育种是培养紫薇新种质的有效途径。本研究主要是通过三个美国紫薇品种与两个四川本地紫薇品种进行杂交,比较了杂交组合间坐果率、结实率、种子萌发率等种实性状及其遗传相关程度,并开展了子代室内苗期抗病性鉴定,初步筛选出适宜于四川本地的紫薇抗白粉病杂交组合,为今后的紫薇新品种的选育提供理论和材料基础。

    • 试验地位于四川乐至国家林业科技示范园区紫薇收集圃。收集圃海拔446 m,日均气温16.7℃,年均日照1330 h,年均降水量900 mm,但分布不均,夏季雨量占全年降雨量的半数。属亚热带湿润季风气候。

    • 试验于2019年8月上旬开始,以紫薇收集圃内收集的五个品种(红火箭、红火球、美国红叶、四川粉花、四川白花)为父母本配置杂交组合,依次用A、B、C、D、E表示五个紫薇品种,字母组合表示杂交组合,母本在前,父本在后,共20个组合。

      选择若干生长健壮无病害、地径5 cm的单株作为花粉来源和授粉母株,各杂交组合采用随机区组排列,重复3次。每个重复3棵母株,每株授粉100朵花。各母株上随机选择100朵花自由授粉,作为坐果率和结实率的对照。

    • 各品种进入盛花期后,早上6点,将选定的待授粉花序用镊子去雄后,套上硫酸纸袋,取下的花药拨入培养皿内带回实验室并置于带有干燥剂的塑料盒中,花药下垫100目细胞筛和硫酸纸静置干燥,待花粉掉落至硫酸纸上,收集至离心管中备用。

      早上8点开始授粉,授粉后继续套袋。10 d后取袋,标记杂交果位置,每隔2 d摘掉新萌发的花。

    • 2019年11月上旬,摘下开始开裂的果实,统计坐果率。将果实进行晾晒、去果壳、筛选,计算每个重复的结实率,置于4℃冰柜保存。

    • 2020年4月,从每个组合随机选取100粒种子,重复3次,播于100孔穴盘的泥炭土中。播种后10 d调查种子萌发情况,计算种子萌发率。出苗后苗圃常规水肥管理,当苗子长出三对真叶时换直径120 mm双色盆。

    • 每个杂交组合子代各随机选择30株参试株。把供试株放置到隔离温室,用软毛刷将紫薇白粉病(Uncinula australiana)孢子刷入0.05%吐温−20的无菌水中,充分震荡后血球计数板镜检计数,制成1×105个·mL−l孢子悬浮液[6]。用刷叶法接种于供试株叶片,每株从上往下处理10片叶,以水滴布满叶面但不滴下为宜[9]。接种后置于温度为20~25℃,湿度>80%的温室内培养。

      接种15 d后调查每株叶片的病叶分级并计算病情指数。单株病叶分级标准参考甜瓜白粉病的分级标准[9],单株病叶分级标准:0级:健康,叶片无病斑;1级:叶片上出现白粉状病斑,病斑面积占叶片面积的5%以下;3级:叶片上有少量病斑,占总面积的6%~25%;5级:叶片上产生较多白粉状病斑,占总面积的26%~50;7级:叶片上的白粉病病斑成片分布,占叶面积的51%~75%;9级别:叶片上的白粉病病斑成片分布且粉层较厚,占叶面积的76%~100%。

    • 坐果率(%)=成熟果实数/杂交花朵数×100;结实率(%)=饱满种子数/种子总数×100;种子萌发率(%)=萌芽种子数/参试种子数×100;病情指数(DI)=[Σ(各级病叶数×各级级值)/(调查总叶数×最高级值)]×100;抗感级别:免疫(IM):DI=0;高抗(HR):0.01~11.11;中抗(MR):11.11~22.22;抗病(R):22.22~33.33;感病(S):33.33~55.55;中感(MS):55.55~77.77;高感(HS):77.77~100。病情指数≤33.33的单株定义为抗病,病情指数>33.33的单株定义为感病。

      利用Excel进行数据分析,DPS 19.05软件计算相关遗传力、遗传力和皮尔森相关。

    2.   结果与分析
    • 坐果率是杂合后第一个可观察性状,是初步判断杂交效果的指标。对20个紫薇杂交组合进行坐果率调查,显著性分析结果表明(见图1),杂交组合间平均坐果率存在显著性差异,其中坐果率最高和最低的组合相差近20倍,最低的CD和CE组合坐果率低于10%,表明紫薇杂交设计需充分考虑亲本搭配以避免试验失败。

      Figure 1.  Comparison of average fruit setting rate of Lagerstroemia indica hybrid combinations
      Note: The dotted line is the average fruit setting rate of free pollination

      同一个母本的杂交组合内均存在显著性差异,5个母本中以红火箭(A)坐果率最高,4个杂交组合的坐果率均超过了50%,其中AB、AC、AD等3个组合的坐果率高于自由授粉的CK坐果率。另外4个母本的杂交坐果率均低于CK坐果率,如美国红叶(C)的CD和CE组合,坐果率均在5%左右,远低于CK 56.67%的坐果率,四川白花(E)的EA、EB和EC组合,远低于CK 79.33%的坐果率。推测原因是套袋对雌花发育有影响,同时自由授粉花粉来源广泛的特点,使得柱头能捕获足够数量可萌发花粉,从而提高坐果率。

      从亲本来源看,红火箭、红火球和美国红叶3个来源于美国的品种以红火箭的杂交组合坐果率较为稳定,红火球次之,美国红叶坐果率波动最大。四川本地的两个品种对同个父本的坐果率比较发现,四川粉花(D)均高于四川白花(E)。对比自由授粉坐果率可知,四川品种的自由授粉坐果率均高于美国品种。推测原因是本地品种在没有套袋处理时,对自然环境的适应能力高于美国品种。

    • 结实率是正常种子的比率,是判断是否完成受精形成种胚的形态指标,也是判断亲本间杂交亲和性的重要指标。对20个紫薇杂交组合进行结实率调查,显著性分析结果表明(见图2),杂交组合间平均结实率存在显著性差异,坐果率和结实率相关系数为0.873(见表1),为极显著强相关。说明坐果率高的杂交组合结实率也高。

      Figure 2.  Multiple comparison of average seed setting rate of Lagerstroemia indica hybrid combinations
      Note: The dotted line is the average seed setting rate of free pollination

      坐果率 Fruit setting rate结实率 Seed setting rate萌芽率 Germination rate
      坐果率Fruit setting rate0.99710.87910.8428
      结实率Seed setting rate0.873**0.99480.7529
      萌芽率Germination rate0.879**0.912**0.9978
        注:上三角为相关遗传力,对角线为遗传力,下三角为皮尔森相关系数,**表示在0.01水平(双侧)上显著相关。
        Note: The upper triangle is related heritability, the diagonal is heritability, and the lower triangle is Pearson correlation coefficient. * * indicates significant correlation at the level of 0.01 (both sides).

      Table 1.  Analysis on correlation heritability and correlation coefficient of seed and fruit characters of Lagerstroemia indica hybrid offspring

      图2中可知,5个母本中,表现最好的依然为红火箭,尽管美国品种的AB组合和美中品种组合的AD组合坐果率有显著性差异,但结实率无显著性差异,说明红火箭和四川粉花的亲和度高;红火箭与另外一个本地品种四川白花的结实率为10.67%,但依然高于另外两个美国品种为母本的所有杂交组合结实率。由此判断,红火箭对参试品种的花粉均具有很好的亲和能力。

      有8个杂交组合获得了果实,但没有收获正常种子,其中美国红火球与两个四川品种的杂交组合(BD、BE)未获得正常种子,美国红叶与红火球、四川白花的杂交组合(CB、CE)未收获正常种子,四川白花与3个美国品种的杂交组合(EA、EB、EC)结实率均为0。

      所有杂交组合平均结实率均显著低于自由授粉CK的平均结实率,推测原因是本实验所在的紫薇收集圃拥有40余种紫薇,自由授粉的花粉来源多样,柱头上亲和度高的花粉替代低级亲和度花粉完成双受精,进而发育成种子。而杂交组合间可能存在的花粉不萌发、花粉管发育异常等现象会导致空瘪粒。

    • 萌发率是种子从吸胀作用开始的一系列有序的生理过程和形态发生过程,是种胚发育成苗的最终阶段。12个紫薇杂交组合的种子萌发率结果表明,各杂交组合之间,以及母本相同的杂交组合间存在显著性差异。

      5个母本中,仅母本为红火箭(A)的杂交子代能萌发,其中美中品种组合:红火箭×四川粉花(AD)萌发率最高,与美美品种组合:红火箭×红火球(AB)的萌发率差异不显著,显著低于红火箭自由授粉子代54.33%的种子萌发率。而其余4个母本的杂交组合萌发率均显著低于各母本自由授粉子代的萌发率。结实率低于10%的CA、CD、BA、BC组合均没有种子萌发。

      Figure 3.  Multiple comparison of average germination rate of Lagerstroemia indica hybrid combinations
      Note: The dotted line is the average germination rate of free pollination

    • 紫薇杂交组合平均坐果率、平均结实率和平均萌芽率等三个种实性状的皮尔森相关分析结果发现,3个性状之间有极显著差异(见表1),表明三个性状之间存在密切联系,由于表型是基因型和环境互作的结果,性状间应该存在一定的遗传相关,进一步的遗传力和相关遗传力分析发现,三个性状的遗传力均达到了0.99,说明这些性状主要是受基因型影响,而受环境的影响较小,表明本试验杂交套袋、播种育苗的操作未对果实和种子发育造成大的影响。

      相关遗传力综合了性状表型值、基因型值、遗传力和相关系数,其判断性状间间接选择的效果比相关系数要简便直观。紫薇杂交组合平均坐果率、平均结实率和平均萌芽率相关系数见表1,坐果率与结实率、萌芽率的相关遗传力均在0.8以上,结实率与萌芽率的相关遗传力为0.75,采用坐果率在选择结实率和萌芽率的效率分别为88.37%、84.47%,因此,紫薇杂交后,可以通过坐果率提前判断该组合的结实率和萌芽率,从而对杂交试验提出改进措施。

    • 杂交组合子代苗期的白粉病发病情况存在较大差异(见图4)。各组合参试单株病情指数计算结果分类见表2

      Figure 4.  Schematic diagram of several typical disease resistance and susceptibility types of Lagerstroemia indica
      A: Immune plant (AD combination); B: Highly resistance plant (AE combination); C: Susceptible plant (DC combination); D: Highly susceptible plant (DE combination)

      杂交组合
      Hybrid
      combination
      抗病类型 Disease-resistance type感病类型 Susceptible type
      免疫
      Immune
      高抗
      Highly Resistance
      中抗
      Medium resistance
      抗病
      Resistance
      总计
      Total
      感病
      Susceptible
      中感
      Medium susceptible
      高感
      Highly susceptible
      总计
      Total
      AB121212618
      AC15158715
      AD67152300
      AE255142644
      DA611522448
      DC268127322
      DE4951835412
      ED3691874112
      总计 Total82548681495033891

      Table 2.  Classification of powdery mildew resistance in F1 generation of 8 Lagerstroemia indica hybrid combination

      从抗病和感病两大类型来看,8个杂交组合参试的240株苗中,抗病株为149株,感病株为91株。在各类杂交组合的所有抗病类型中,免疫株为8株,占参试总数的3.33%,高抗株为25株,占参试总数的10.41%,中抗株为48株,占参试总数的20%。抗病株为68株,占参试总数的28.33%。免疫株和高抗株总数占抗病类型的22.15%,中抗株和抗病株总数占抗病类型的77.85%,说明在抗病类型中,主体还是中抗株和抗病株。

      在各类杂交组合的所有感病类型中,感病株为50株,占参试总数的20.83%,中感株为33株,占参试总数的13.75%,高感株为8株,占参试总数的3.33%。感病株和中感株占感病类型的91.2%,而高感株占感病类型的8.8%,说明在感病类型中,高感株的比例较少,而高感类型中7株来源于四川粉花为母本的子代。

      从每个亲本组合抗病类型和感病类型的数量来看,抗病类型数量过半的组合有AD、AE、DA、DE、ED,其中3个组合中出现了A(红火箭),说明红火箭为母本,以亲缘关系较远的四川品种为父本,子代中可以取得很好的白粉病抗性,而且免疫株均来源于这类组合,其中AD组合表现最为优异,所有参试子代均为抗病类型,其免疫株与高抗株共13株,占参试株的43.33%,AD组合的免疫株占所有免疫株的75%。而AE组合子代的抗性评价稍差于AD组合,有4株子代为感病株,剩余26株抗病类型中,免疫株与高抗株共7株,占参试株的23.33%。因此,AD和AE组合为抗白粉病的杂交组合。

      但并不是所有亲缘关系较远的杂交类型的抗性表现都好,DC组合的抗病类型数量是所有杂交组合中最少的,只有8株抗病株且其中2株为中抗株,6株为抗病株。DC组合整体表现最差。

      比较美美组合的AB和AC组合,不仅感病类型的子代数量超过了抗病类型,而且抗病类型均为最低档的抗病株,抗病能力高的类型均没有。子代整体表现集中在三种类型,说明美美组合子代的变异水平相比美中组合和本地组合要低,可能美国品种的纯合度高于本地品种。

      四川本地品种的两个组合为正反交,抗病类型比感病类型均只多了6株,抗病类型与感病类型分布比较均匀。抗病类型总数量均为18株,没有出现免疫株。从各类型的分布看,囊括了7种类型的6种,说明本地品种间杂交的遗传变异非常丰富,若扩大本地品种的杂交后代群体数量,增强选育强度,从中筛选出高抗类型的子代也是可能的。

      综上所述,在制定紫薇抗白粉病育种策略时,红火箭、四川粉花和四川白花既适合做父本也适合做母本,而红火球和红叶不适合选做杂交材料。本研究初步筛选出“红火箭×四川粉花”和“红火箭×四川白花”两个抗白粉病杂交组合。

    3.   结论与讨论
    • 种间和种内杂交是创制紫薇新品种的有效途径[10],杂交成功率首先考察的是杂交可配性,杂交可配性是指杂交取得有生命力的杂种种子的概率[11]。能否取得杂种是杂交育种成败的关键[12]。因此,确定杂交组合时应考虑各树种间可配性的大小。不可配性的表现形式有多种:花粉不能在柱头上萌发;能发芽,但不能长出柱头组织;长出了柱头组织,但不能达到胚囊;或虽能受精,但胚胎发育不正常,不能得到有生命力的种子等。这是由于双亲遗传基础差距太大,配子间在生物学上不相适应的结果[12]

      本研究共开展了20个组合的杂交试验,有12个杂交组合收获了种子,杂交成功率为60%,其中4个组合的种子不具备萌发能力,只有8个组合种子能够萌发,尽管杂交成功受多方面因素影响,但是取得了有生命力的种子的组合说明了杂交亲本间是具有可配性的。

      五个品种中明显看出以美国红火箭紫薇为母本时其坐果率优于另外四个品种,其中“红火箭×四川粉花”坐果率和种子萌发率均为最高分别为82.1%和44.1%,红火箭为母本的四个杂交组合种子均能萌发,表明当红火箭为母本时对其他四个品种紫薇的雄配子亲和性更强。

      当红火球和美国红叶为母本时,8个杂交组合种子均没有萌发,表明红火球和美国红叶对其他四个品种雄配子具有不可配性,不适合做母本。当四川粉花为母本时,只对红火球雄配子具不可配性,其他三个杂交组合中DE组合的三率均最高分别为55.33%、23.55%和18.55%。当四川白花为母本时,只对四川粉花的雄配子具有可配性。

      本研究在获得了杂交子代后,进行了白粉病抗性试验,从中初步筛选出适宜于做抗性育种亲本的红火箭、四川粉花和四川白花,获得了149个抗性子代,其中免疫单株8个,高抗单株25个,为紫薇下一步抗性育种提供了轮回育种材料,同时也为紫薇新品种创制了新种质。由于白粉病的菌丝体主要在叶子正反两面及叶柄上着生、存留[13]。白粉病菌感染叶片后,首先引发植物生理指标变化[8, 14]。下一步课题组将开展感菌前后紫薇叶片解剖结构和相关抗逆性酶的反应研究[15, 16],从解剖学和生理学的角度阐明紫薇抗白粉病的机理。

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