CN114467578B - 一种植物线虫的防治方法 - Google Patents
一种植物线虫的防治方法 Download PDFInfo
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- CN114467578B CN114467578B CN202011269923.9A CN202011269923A CN114467578B CN 114467578 B CN114467578 B CN 114467578B CN 202011269923 A CN202011269923 A CN 202011269923A CN 114467578 B CN114467578 B CN 114467578B
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Abstract
本发明提供了一种植物线虫的防治方法。该方法为：采用植物甾醇喂养植物线虫，植物甾醇不包含胆固醇和谷固醇。本发明提供的依赖于DHCR24的线虫防控方法，完全不同于已知的轮作、化学杀线虫剂或针对线虫特定蛋白酶的基因工程策略，具有更强的靶点特异性和更高的物种广谱性。本发明通过干扰线虫的甾体代谢或通过调控植物的甾体代谢实现线虫防治，可针对不同线虫种或不同作物株系，开发特异的防治策略。也为在其它物种中(如家畜)通过调控甾醇代谢实现寄生性线虫防控提供了理论支撑和技术平台。
Description
技术领域
本发明属于生物技术领域，具体涉及一种植物线虫的防治方法。
背景技术
植物线虫病害是一种严重制约农作物产量的重要病害，其危害超过细菌和病毒，仅次于真菌病害，每年造成世界农业生产损失约1250亿美元(Chitwood，Biochemistry andfunction of nematode steroids，Critical Reviews inBiochemistry and MolecularBiology，1999，34(4)273-284)。常见的线虫防治方法有轮作和化学杀线虫剂等。轮作主要是通过易感病作物与感病轻作物的轮作，可以一定程度降低虫口密度。但部分孢囊线虫在土壤中存活时间长达10多年；且有些线虫(如根结线虫)具有宽泛的寄主范围，限制了作物轮作在线虫防治中的应用。化学杀线虫剂可以有效降低根结线虫对作物的危害，但多次使用后线虫易产生抗药性，同时也增加了农业生产成本，而且长期使用易引起严重的土壤污染，破坏土壤微环境，不符合农业可持续发展的要求。推广抗线虫品种是一种有效防治线虫病害的策略，但植物中抗性种质资源非常匮乏。
植物寄生线虫是生活在潮湿表面或液体环境，包括土壤中的水膜和其它生物中的潮湿组织中的活跃的、柔软的长形生物体。存在多种植物寄生线虫物种，包括多种孢囊线虫(如异皮线虫，Heterodera sp.)、根瘤线虫(如根结线虫，Meloidogyne sp.)、根腐线虫(如短体线虫，Pratylenchus sp.)、剑线虫(如剑线虫，Xiphinema sp)以及茎和球茎线虫(如茎线虫，Ditylenchus sp.)等。垫刃线虫(Tylenchid nematode；垫刃线虫目的成员)，包括异皮线虫科、根结线虫科和短体线虫科，是植物寄生线虫的最大和经济上最重要的组。其它重要的植物寄生线虫包括矛线虫(如剑线虫)等。线虫物种经过一系列生命周期阶段和蜕皮而生长。典型地，存在5个阶段和4次蜕皮：卵阶段；J1(即第一幼虫阶段)；M2(即第一次蜕皮)；J2(第二幼虫阶段；有时从卵孵出)；M2；J3；M3；J4；M4；A(成虫)。幼虫(“J”)阶段有时称作幼虫(“L”)阶段。
固醇(英文：sterol)又称甾醇，是类固醇的一类，是含有羟基的类固醇。它们均以环戊烷多氢菲为基本结构，并含有羟基，故称为固醇类化合物。甾醇是广泛存在于生物体内的一种重要的天然活性物质，按其原料来源分为动物性甾醇、植物性甾醇和菌类甾醇等三大类。动物性甾醇以胆固醇为主，植物性甾醇主要为谷固醇、豆甾醇和菜油甾醇等，而麦角甾醇则属于菌类甾醇。植物甾醇广泛存在于植物的根、茎、叶、果实和种子中，是植物细胞膜的组成部分，在所有来源于植物种子的油脂中都含有甾醇。
寄生性植物线虫没有从头合成甾醇的能力。它们摄取宿主植物甾醇并将其通过脱烷基转化为胆固醇，以满足结构和激素需求。采用化学方法阻断24-脱氢胆固醇还原酶DHCR24导致植物甾醇脱烷基化中间体的积累、胆固醇合成的终止和昆虫幼虫的发育迟缓。这表明昆虫DHCR24在植物甾醇脱烷基中的重要作用，而DHCR24可能是农药设计的潜在目标。但目前，尚不知道线虫中是否存在DHCR24介导的脱烷基作用，或者它们在这种古老生物中可能具有何种功能。因此，亟待解析线虫DHCR24的功能，并探索可否通过调控DHCR24用于线虫防治。
发明内容
有鉴于此，本发明提供了一种植物线虫的防治方法。相比轮作、化学杀线虫剂或针对线虫特定蛋白酶的基因工程策略，该方法具有更强的靶点特异性和更高的物种广谱性。
为了实现上述发明目的，本发明提供以下技术方案：
本发明提供了一种植物线虫的防治方法，该方法为：采用植物甾醇喂养植物线虫，植物甾醇不包含胆固醇和谷固醇。
通过解析秀丽隐杆线虫Caenorhabditis elegans DHCR24(CeDHCR24)突变体的固醇代谢，发明人发现CeDHCR24是线虫利用植物固醇所必需的酶，其中以谷固醇作为唯一固醇源时，CeDHCR24的缺陷导致线虫的生长受阻。采用胆固醇部分替代谷固醇，CeDHCR24缺陷线虫的生长表现出剂量依赖性。这表明通过调控DHCR24可用于线虫防治。为了证明这一发现的实用性，发明人设计了具备特定结构特征的植物甾醇。自然界植物中天然存在此类植物甾醇，且此类植物甾醇在植物体中的积累并不影响植物生长。喂养此类植物固醇的线虫无法通过脱烷基作用合成胆固醇，从而导致线虫生长发育停滞。这些结果有助于理解线虫固醇代谢的分子机制，并为防控植物寄生线虫和一般植物食性虫害提供了可行的方法。
本发明的植物线虫的防治方法包括通过阻断线虫内源的植物甾醇转化途径导致其发育停滞或通过调控植物中甾体化合物谱导致食用相应植物甾醇后的线虫生长停滞。
本发明主要涉及两方面的内容：一是利用遗传与生化方法鉴定了线虫利用植物甾醇合成自身胆固醇的酶；二是通过化学生物学手段，采用具备特异结构特征的植物甾醇喂养线虫，可阻断线虫发育。
作为优选，植物甾醇为7-去氢胆固醇、7-胆甾烯醇、4α-methylcholst-8(14)enol、Lophanol、4-甲基-7-烯胆甾烷醇、胆甾醇5,7,24-三烯醇、胆甾醇7,24-二烯醇、4α-甲基胆甾8(14),24-二烯醇、24-乙基胆甾醇5,25(27)-二烯醇、24-乙基胆甾-5,7,25(27)-三烯酚、24-乙基胆甾7,25(27)-二烯酚、4-甲基,24-乙基-8(14),25(27)-二烯醇中的一种或几种。
在本发明种，植物线虫包括但不限于孢囊线虫、根瘤线虫、根腐线虫、剑线虫、矛线虫、垫刃线虫，以及茎、球茎线虫。
在本发明种，孢囊线虫包括但不限于异皮线虫。
在本发明种，根瘤线虫包括但不限于根结线虫。
在本发明种，根腐线虫包括但不限于短体线虫。
在本发明种，剑线虫包括但不限于剑线虫。
在本发明种，矛线虫包括但不限于剑线虫。
在本发明种，垫刃线虫包括但不限于异皮线虫科、根结线虫科和短体线虫科。
在本发明种，茎、球茎线虫包括但不限于茎线虫。
作为优选，植物甾醇存在于植物体中。
本发明提供了一种植物线虫的防治方法。该方法为：采用植物甾醇喂养植物线虫，植物甾醇不包含胆固醇和谷固醇。本发明与已有技术相比具有以下显著特点和积极效果：
(1)目前为止，线虫防治的策略中尚不存在通过调控甾体代谢抑制线虫发育的应用。本发明提供的依赖于DHCR24的线虫防控方法，完全不同于已知的轮作、化学杀线虫剂或针对线虫特定蛋白酶的基因工程策略(RNAi敲低特定蛋白酶)，具有更强的靶点特异性和更高的物种广谱性。
(2)本发明提供了两种实现线虫防治的方法：通过干扰线虫的甾体代谢或通过调控植物的甾体代谢。针对不同线虫种或不同作物株系，开发特异的防治策略。也为在其它物种中(如家畜)通过调控甾醇代谢实现寄生性线虫防控提供了理论支撑和技术平台。
附图说明
图1本发明涉及的代表性甾体化合物的化学结构式；
图2喂养胆固醇的秀丽隐杆线虫突变体tm2141的甾体代谢谱，数字代表鉴定到的甾体化合物(甾体结构和质谱图参见图1)；
图3喂养谷固醇的野生型秀丽隐杆线虫(N2)的甾体代谢谱；
图4喂养谷固醇的秀丽隐杆线虫突变体tm2141的甾体代谢谱；
图5饲喂不同固醇的秀丽隐杆线虫突变体tm2141的发育分析；其中，
1：用胆固醇喂养(重复r＝8，被测动物总数n＝186)；
2：喂食谷固醇(r＝8，n＝117)；
3：连续两次喂食谷固醇，每次持续48h(r＝5，n＝101)；
4：谷固醇喂养48小时后，以谷固醇/胆固醇(1∶1)配方喂养(r＝10，n＝375)；
5：谷固醇喂养48小时后，以谷固醇/胆固醇(9：1)配方喂养(r＝10，n＝353)；
图6喂养赤桐甾醇(clerosterol,CLE)的秀丽线虫N2蠕虫的甾体化合物谱；数字(11、12、13和14)为鉴定到的甾体化合物，其结构和质谱参见图1；
图7喂养谷固醇(sitosterol)和赤桐甾醇(clerosterol,CLE)的秀丽线虫N2蠕虫；结构式显示了谷固醇侧链结构中C24和C25氢的位置和CLE侧链结构中C25氢的缺失。
具体实施方式
本发明公开了一种植物线虫的防治方法，本领域技术人员可以借鉴本文内容，适当改进工艺参数实现。特别需要指出的是，所有类似的替换和改动对本领域技术人员来说是显而易见的，它们都被视为包括在本发明。本发明的方法及应用已经通过较佳实施例进行了描述，相关人员明显能在不脱离本发明内容、精神和范围内对本文所述的方法和应用进行改动或适当变更与组合，来实现和应用本发明技术。
化学物名称中英文对照：
1、Cholesterol，胆固醇
2、7-dehydrocholesterol，7-去氢胆固醇
3、Lathosterol,7-烯胆甾烷醇
4、4α-methylcholst-8(14)enol
5、Lophanol
6、Lophenol
7、Cholesta-5,7,24-trienol，胆甾醇5,7,24-三烯醇
8、sitosterol,谷固醇/谷甾醇
9、Cholesta-7,24-dienol，胆甾醇7,24-二烯醇
10、4α-methylcholesta-8(14),24-dienol，4α-甲基胆甾8(14),24-二烯醇
11、24-ethylcholesta-5,25(27)-dienol,24-乙基胆甾醇5,25(27)-二烯醇
12、24-ethylcholesta-5,7,25(27)-trienol,24-乙基胆甾-5,7,25(27)-三烯酚
13、24-ethylcholesta-7,25(27)-dienol,24-乙基胆甾7,25(27)-二烯酚
14、4α-methyl,24-ethyl-8(14),25(27)-dienol,4-甲基,24-乙基-8(14),25(27)-二烯醇。
本发明提供的植物线虫的防治方法中所用试剂或仪器均可由市场购得。
下面结合实施例，进一步阐述本发明：
实施例1DHCR24阻断前后线虫甾体代谢谱的对比
本发明获得了秀丽隐杆线虫CeDHCR24敲除突变体tm2141，该突变体是F52H2.6/DHCR24等位基因，缺失范围为29539至30298，长度为760bp。该突变体等位基因丢失了外显子4和部分外显子5。tm2141突变体能够在含有5mg/L胆固醇的培养基(NGM)上正常生长。被突变的同化胆固醇(1；图1和图2)被代谢为7-脱氢胆固醇(2；图2)，7-烯胆甾烷醇(3；Lathosterol，图2)，4α-methylcholst-8(14)enol(4；图2)，lophanol(5；图2)和lophenol(6；图2)。其固醇特征与饲喂胆固醇的野生秀丽隐杆线虫(Chitwood,D.J.,Lusby,W.R.,Lozano,R.,Thompson,M.J.&Svoboda,J.A.Sterol metabolism in the nematodeCaenorhabditis elegans.Lipids，1984，19,500-506)非常相似，表明DHCR24参与胆固醇生物合成上游的固醇代谢途径。
然后将来自胆固醇平板的tm2141子代饲养在仅包含谷固醇的平板上，同时将野生型蠕虫用作对照。野生型线虫可以高效的将谷固醇(8；有关结构和质谱图参见图1)代谢为胆固醇(1；有关结构和质谱图参见图1)和随后的终产物(图3)。同时，观察到存在少量的胆甾醇5,7,24-三烯醇(cholesta-5,7,24-trienol，7；有关结构和质谱图，请参见图1)，这可能是通过核修饰衍生自cholesta-5,24-dienol。相反，在tm2141突变体中未检测到胆固醇(图4)。相反，cholestea-5,7,24-trienol(7；图1)是主要固醇(图4)，约占突变体中总固醇的60％，这表明tm2141中固醇脱烷基步骤被阻断，而DHCR24缺失后，24-脱氢胆固醇(desmosterol)的核修饰成为主要的代谢流。胆甾醇5,7,24-三烯醇(cholesta-5,7,24-trienol，7)，胆甾醇7,24-二烯醇(cholesta-7,24-dienol，9；有关结构和质谱图参见图1；通过核修饰衍生自胆甾醇5,7,24-三烯酚)和4α-甲基胆甾8(14)，24-二烯醇(4α-methylcholesta-8(14),24-dienol，10；有关结构和质谱图，请参见图1；通过核修饰从cholesta-7,24-二烯醇中衍生)(图4)。
甾体化合物提取和鉴定方法参照Lu等，Regulation of the cholesterolbiosynthetic pathway and its integration with fatty acid biosynthesis in theoleaginous microalga Nannochloropsis oceanic,2014,Biotechnology for Biofuels,7:81和Clade-specific sterol metabolites in dinoflagellate endosymbionts areassociated with coral bleaching in response to environmental cues,2020,mSystems,doi:10.1128/mSystems.00765-20。采用Agilent6890N气相色谱仪检测，FFAP柱子，分流比：20∶1；进样口温度：250℃；柱温：150℃-230℃程序升温；检测器温度：250℃；气化室温度：350℃；尾吹：40mL/min；氢气流速：45mL/min；空气流速：450mL/min；检测器：氢火焰离子化检测器(FID)；进样量：1μL，具体操作参照按仪器操作说明书进行操作。
实施例2通过阻断线虫内源的植物甾醇转化途径导致其发育停滞
当用足够的胆固醇培养时，tm2141能够完成整个生命周期，与野生型秀丽隐杆线虫(N2)动物没有区别。胆固醇平板上同步化后L1蠕虫(参阅秀丽隐杆线虫生命周期：Fielenbach,N.&Antebi,A.C.elegans dauer formation and the molecular basis ofplasticity.Genes&Development,2008,22,2149-2165)被转移到仅含谷固醇的平板中。25℃下48h后，94％的野生型蠕虫生长到成虫(图5中为1)，而tm2141的发育明显阻滞，仅20％的蠕虫达到成年阶段(图5中为2)。使用谷固醇饲养的tm2141的后代产生了同步的L1蠕虫。这些蠕虫体内被去除了胆固醇，之后转移至新鲜的谷固醇平板上培养。在25℃下孵育48小时后，100％蠕虫未达到成年期(图5中的3)。可见，固醇脱烷基化对于仅依赖植物固醇的秀丽隐杆线虫的生长是必不可少的。
由于仅依赖谷固醇的tm2141无法完成其生命周期，因此用胆固醇替代了10％或50％的谷固醇。相同数量的tm2141蠕虫接种在这两种不同固醇配方(谷固醇/胆固醇比为9：1或1：1)的新鲜平板上。48小时后，74％的tm2141蠕虫在谷固醇/胆固醇(1：1)平板上达到L4阶段(参阅秀丽隐杆线虫生命周期：Fielenbach,N.&Antebi,A.C.elegans dauerformation and the molecular basis of plasticity.Genes&Development,2008,22,2149-2165)(图5中的4)。而喂养谷固醇/胆固醇(9：1)配方的突变蠕虫中，仅有26％进入L4阶段(图5中的5)。因此，胆固醇的添加对tm2141表型的回复表现出剂量依赖性。
实施例3通过调控植物中甾醇谱导致食用相应植物甾醇后的线虫生长停滞
同位素标记实验表明，昆虫C24和C25氢对于固醇脱烷基至关重要(Fujimoto,Y.,Kimura,M.,Takasu,A.,Khalifa,F.A.M.&Ikekawa,N.Mechanism of stigmasteroldealkylation in insect.Tetrahedron Letters,1984,25,1501–1504；Ikekawa,N.,Morisaki,M.&Fujimoto,Y.Sterol metabolism in insects:dealkylation ofphytosterol to cholesterol.Accounts of Chemical Research,1993,26,139-146)。基于这一点，设计选取了24-乙基胆甾醇5,25(27)-二烯醇(clerosterol，CLE；11，图1)。CLE在植物体内天然存在，但缺少C25氢(结构和质谱图参见图1)。
外源固醇及其经核修饰的中间体是蠕虫的主要成分(图3)。虽然在动物体内未检测到胆固醇，但CLE被吸收并通过核修饰转变为24-乙基胆甾-5,7,25(27)-三烯酚(24-ethylcholesta-5,7,25(27)-trienol，12)，24-乙基胆甾7,25(27)-二烯酚(24-ethylcholesta-7,25(27)-dienol，13)和4-甲基,24-乙基-8(14),25(27)-二烯醇(4-methyl,24-ethyl-8(14),25(27)-dienol，14)(图6)(结构和质谱图参见图1)。因此，即使野生型线虫中存在有效的24-脱烷基途径，其蠕虫仍无法将人工设计的植物甾醇转化为胆固醇。将谷固醇上的野生型秀丽隐杆线虫的L1子代分别在含有谷固醇(对照)或CLE的平板上培养。与对照相比，在CLE平板上的动物的发育被严重阻滞(图7)。因此，人工设计植物甾醇可以抑制野生型线虫的发育。
以上所述仅是本发明的优选实施方式，应当指出，对于本技术领域的普通技术人员来说，在不脱离本发明原理的前提下，还可以做出若干改进和润饰，这些改进和润饰也应视为本发明的保护范围。
Claims (9)
1.一种植物线虫的防治方法，其特征在于，采用植物甾醇喂养植物线虫，所述植物甾醇不包含胆固醇和谷固醇；所述植物甾醇为7-去氢胆固醇、7-胆甾烯醇、4α-methylcholst-8(14)enol、Lophanol、4-甲基-7-烯胆甾烷醇、胆甾醇5,7,24-三烯醇、胆甾醇7,24-二烯醇、4α-甲基胆甾8(14),24-二烯醇、24-乙基胆甾醇5,25(27)-二烯醇、24-乙基胆甾-5,7,25(27)-三烯酚、24-乙基胆甾7,25(27)-二烯酚、4-甲基,24-乙基-8(14),25(27)-二烯醇中的一种或几种。
2.根据权利要求1所述的防治方法，其特征在于，所述植物线虫包括但不限于孢囊线虫、根瘤线虫、根腐线虫、剑线虫、矛线虫、垫刃线虫，以及茎、球茎线虫。
3.根据权利要求2所述的防治方法，其特征在于，所述孢囊线虫包括但不限于异皮线虫。
4.根据权利要求2所述的防治方法，其特征在于，所述根瘤线虫包括但不限于根结线虫。
5.根据权利要求2所述的防治方法，其特征在于，所述根腐线虫包括但不限于短体线虫。
6.根据权利要求2所述的防治方法，其特征在于，所述剑线虫包括但不限于剑线虫。
7.根据权利要求2所述的防治方法，其特征在于，所述矛线虫包括但不限于剑线虫。
8.根据权利要求2所述的防治方法，其特征在于，所述垫刃线虫包括但不限于异皮线虫科、根结线虫科和短体线虫科；
所述茎、球茎线虫包括但不限于茎线虫。
9.根据权利要求1至8中任一项所述的防治方法，其特征在于，所述植物甾醇存在于植物体中。
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