外源茉莉酸类化合物对两种落叶松的诱导抗虫性研究
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摘要
落叶松(Larch pine)是东北地区主要造林树种,20世纪90年代后期落叶松毛虫Dendrolimus superans发生猖撅,危害严重。本文以兴安落叶松Larix gemelinii、长白落叶松Larix olgensis为对象,运用茉莉酸、茉莉酸甲酯和茉莉酮3种化学信号物质喷施处理2年生落叶松苗,研究其体内化学物质的时序变化,对落叶松毛虫及其天敌—平腹小蜂寄主选择行为的影响。从营养、防御蛋白、次生代谢物质等角度探讨外源茉莉酸类化合物对2种落叶松诱导抗虫性作用,以及落叶松毛虫、平腹小蜂对植物防御物质相应的行为反应机制,为提高林木健康水平和有效控制落叶松毛虫种群密度提供理论依据。通过化学物质分析和生物测定,得到如下结论:
1外源茉莉酸类化合物诱导两种落叶松针叶内营养物质的时序变化
外源茉莉酸类化合物处理后,长白、兴安落叶松针叶内可溶性糖含量在第1、3、10、15、20d含量降低,第5d增加,呈现波动变化,但诱导2种落叶松针叶内可溶性糖含量变化不显著。
长白落叶松经外源茉莉酸类化合物处理后,用高效液相色谱仪共鉴定出17种游离氨基酸,其总量在第1、5、10d含量极显著增加,第3、15、20d含量极显著降低;在不同诱导时间内,各处理游离氨基酸总量变化差异显著。不同种游离氨基酸组分百分含量在第1、5、10d显著降低,第3、15、20d明显上升,与游离氨基酸酸总含量变化趋势相反;不同处理诱导游离氨基酸组分百分含量的变化差异显著,但氨基酸变异系数小于1,表明其诱导氨基酸组分百分含量的波动变化是正常的。
2外源茉莉酸类化合物诱导两种落叶松针叶内次生物质的时序变化
外源茉莉酸类化合物诱导处理后落叶松针叶内单宁含量变化显著。兴安落叶松在处理后第1、10d针叶内单宁含量显著降低,第3、5d含量明显升高。长白落叶松在诱导后3、5、15d针叶内单宁含量明显增加,1、10、20d含量降低。
外源茉莉酸类化合物处理后,长白落叶松针叶内酚酸总含量在第1、3、5、10d降低,第15d显著升高,第20d显著降低的波动变化;兴安落叶松针叶内酚酸总含量变化总体呈现降低的趋势,增加不显著。兴安落叶松针叶用高效液相色谱仪共鉴定出11种酚酸;咖啡酸、阿魏酸、香豆酸、苯乙酸、水杨酸、香草酸变化趋势相似,呈现诱导前期含量显著降低,后期逐渐升高的动态变化;肉桂酸和原儿茶酸则呈现诱导前期升高,后期降低的变化规律。长白落叶松针叶共检测到12种酚酸;咖啡酸、苯乙酸、水杨酸含量呈现先降后升的动态变化;绿原酸、肉桂酸、原儿茶酸、香草酸含量是先增加后降低的波动变化。
3外源茉莉酸类化合物诱导两种落叶松防御蛋白活性的时序变化
外源茉莉酸类化合物诱导后,2种落叶松针叶内多酚氧化酶和苯丙氨酸解氨酶、蛋白酶抑制剂活性明显增加,呈现先升高后降低的波动性变化;不同处理时间各处理诱导作用显著,表明2种落叶松的诱导抗虫性产生。外源茉莉酸类化合物对2种落叶松胰凝乳蛋白酶抑制剂诱导作用较强;对兴安落叶松针叶内胰凝乳蛋白酶抑制剂诱导作用更明显。
4外源茉莉酸诱导两种落叶松对落叶松毛虫寄主选择行为的影响
落叶松毛虫在不同浓度茉莉酸处理的2种落叶松松苗上产卵量差异显著;对照树苗上的落卵量显著高于茉莉酸处理树苗上的卵量;3个不同浓度茉莉酸处理的松树苗上的落卵量差异不显著;茉莉酸处理树苗对落叶松毛虫产卵有很强驱避作用。
不同处理2种落叶松对落叶松毛虫的嗅觉反应影响显著;随茉莉酸处理时间的延长,落叶松对落叶松毛虫驱避性增强;随着茉莉酸处理浓度的增加,驱避性加大。与对照相比,2种落叶松处理后1d对落叶松毛虫没有明显的驱避作用;处理后3、5d驱避性显著。
5外源茉莉酸诱导两种落叶松对平腹小蜂寄主选择行为的影响
平腹小蜂对不同处理的落叶松上落叶松毛虫卵的寄生率,随处理时间的延长逐渐降低;但不同诱导时间内的各处理间寄生率差异均不显著。随着茉莉酸处理浓度的加大,平腹小蜂对落叶松毛虫卵的寄生率呈波动变化。
平腹小蜂对不同处理落叶松的嗅觉反应明显;随着茉莉酸处理浓度的增加,趋性增强;随着处理时间的延长,趋性呈波动变化。平腹小蜂对茉莉酸处理的第1、5d的松苗趋性显著。
本文较系统地研究了应用茉莉酸、茉莉酸甲酯和茉莉酮3种外源化合物诱导两种落叶松体内化学物质的时序变化,结果表明,外源化合物诱导落叶松体内营养物质可溶性糖和游离氨基酸含量降低,次生物质单宁和酚酸含量增加,蛋白酶抑制剂等防御蛋白活性增强,落叶松直接诱导抗虫性产生;且茉莉酸诱导落叶松对落叶松毛虫搜寻寄主产生明显驱避性,对平腹小蜂的寄主选择有显著引诱力,表明茉莉酸诱导落叶松产生间接抗虫性,从而增强落叶松自身的防御能力,减少植食昆虫的危害。茉莉酸、茉莉酸甲酯和茉莉酮能够在保持植物完好的情况下,模拟剪叶损伤和昆虫取食危害,诱导营养物质和次生物质的变化,对取食的昆虫产生毒害作用,又能诱导产生吸引害虫天敌的挥发性物质,其作为植物抗虫诱导剂,不污染环境,对天敌安全,保持生态平衡,有利于害虫的可持续控制。
Larch pine is main species of forest in northeast of China.Dendrolimus superans(Butler) boomed since 90s in the twentieth century and serious harmful result was seen.In this research, Larix gmelinii and Larix olgensis were selected as objects for research.Temporal variation of chemical compounds in two larch species,and effects of host plant selection of Dendrolimus superans and of Anastatus japonicus,which were performed to analysis through treatment of jasmonic acid(JA),methyl jasnonate(MJA) and cis-jasmone.The influence of exogenous jasmonates induced resistance of two larch species to insects and relative biochemical mechanism to Dendrolimus superan and Anastatus japonicus were analyzed about the factors, such as nutrient compounds,defensive proteins and secondary metabolites.It was important for improveing the healthy level of forest and control of population density of Dendrolimus superan.By using analysis of chemical compounds and biometrics,results of the research were listed below:
1.Changing of nutrient compounds in the needles of Larix gmelinii and L.olgensis induced with exogenous jasmonates
The contents of soluble sugar in the needles of Larix gmelinii and L.olgensis was fluctuant variation,and which decreased at the 1~(st),3~(rd),10~(th),15~(th),20~(th) day,and which increased at the 5~(th) day after exogenous jasmonates application.The variation of the contents of soluble sugar was no significant difference.
17 species of free amino acid were identified in the needles of Larix olgensis by HPLC.In contrasted with CK,the total contents of free amino acids increased at the 1~(st),5~(th),10~(th) day,and which decreased at the 3~(rd),15~(th),20~(th) day in the needles of L.olgensis after exogenous jasmonates application.The variation of the total contents of free amino acids was significantly different in the period of the experimentation.The component percents of free amino acids dropped at the 1~(st),5~(th),10~(th) day,and which raised at the 3~(rd),15~(th),20~(th) day in the needles of L. olgensis treated with jasmonates.But variational coefficient of free amino acids was smaller than one.Which sharp reversed with the variation of the total contents of free amino acids.The result indicated the variation of the component percents of free amino acids was normal fluctuation.
2.Changing of secondary metabolites in the needles of Larix gmelinii and L.olgensis induced with exogenous jasmonates
The contents of tannins changed significantly in the needles of Larix gmelinii and L. olgensis after exogenous jasmonates application.The contents of tannins in the needles of Larix gmelinii dropped at the 1~(st),10~(th) day,and which raised at the 3~(rd),5~(th) day.While which of L. olgensis decreased at the 1~(st),10~(th),20~(th) day,and which increased at the 3~(rd),5~(th),15~(th) day after application exogenous jasmonates.
The contents of phenolic acid in the needles of Larix olgensis dropped at the 1~(st),3~(rd),5~(th), 10~(th) day,and which raised at the 15~(th) day,then which dropped at the 20~(th) day after treated with exogenous jasmonates.The contents of phenolic acid in the needles of Larix gmelinii was decreasing in the period of the experimentation.11 species of phenolic acid were identified in the needles of Larix gmelinii by HPLC.There was similar variation for the contents of caffeic acid,ferulic acid,p-coumaric acid,phenylacetic acid,salicylic acid and vanillic acid.Which raised formerly,dropped later in needles of L.gmelinii induced with exogenous jasmonates. While which of cinnamic acid and chlorogenic acid sharp reversed with those.12 species of phenolic acid were identified in the needles of Larix olgensis by HPLC.The variation of the contents of caffeic acid,phenylacetic acid and salicylic acid dropped formerly,raised later in needles of L.olgensis treated with exogenous jasmonates.And which of cinnamic acid, chlorogenic acid,chlorogenic acid and vanillic acid sharp reversed with those.
3.Changing of the activity of defendable enzymes in the needles of Larix gmelinii and L.olgensis induced by exogenous jasmonates
The activities of PPO,PAL,trypsin inhibitor and chymotrypsin inhibitor increased significantly in the needles of Larix gmelinii and L.olgensis induced with exogenous jasmonates.The variation of their activity was waved in the period of the experimentation. While the induced function of the different concentration jasmonates was significantly different. Which indicated 2 species of larch pines formed the induced resistance to insect.
4.Effect of two larch species induced with exogenous jasmonic acid on the behavior of host plant selection of Dendrolimus superans
The number of oviposition of Dendrolimus superans was significantly different in the seedlings treated with different concentration JA.The number of egg in control seedlings were much more than which in the seedlings treated with JA.There was not different for the number of eggs in the seedlings after different concentration JA application.There was strongly drivable for the seedlings treated with JA to the oviposition behavior of Dendrolimus superans.
The drivable function of two larch species treated with JA to Dendrolimus superans was increased with prolonging days and increasing concentration treated with JA.Compared with CK,the drivable function of larch pine to Dendrolimus superans was not different at the 1~(st) day after jasmonic acid application,and which was significantly different at the 3~(rd),5~(th) day following jasmonic acid application.
5.Effect of two larch species induced with exogenous jasmonic acid on the behavior of host plant selection of Anastatus japonicus
The parasitism rate of Anastatus japonicus to Dendrolimus superans eggs was dropped with prolonging days treated with JA,and which was not different in the different treatments in the period of the experimentation.Which was waving with increasing concentration treated with JA.
The attractive function of two larch species treated with JA to Anastatus japonicus was increased with prolonging days,and which was waving with increasing concentration treated with JA.Which was significantly different at the 1~(st),5~(th) day following JA application.
Therefore,this paper expounded exogenous jasmonates induced two larch species to form direct and indirect resitance to insects.Which prevented or disturbed pest insects feeding,and protected natural enemies.While induced resistance to insect maintained the balance of forest ecosystem,which would play a key role in pest integrated control.
1外源茉莉酸类化合物诱导两种落叶松针叶内营养物质的时序变化
外源茉莉酸类化合物处理后,长白、兴安落叶松针叶内可溶性糖含量在第1、3、10、15、20d含量降低,第5d增加,呈现波动变化,但诱导2种落叶松针叶内可溶性糖含量变化不显著。
长白落叶松经外源茉莉酸类化合物处理后,用高效液相色谱仪共鉴定出17种游离氨基酸,其总量在第1、5、10d含量极显著增加,第3、15、20d含量极显著降低;在不同诱导时间内,各处理游离氨基酸总量变化差异显著。不同种游离氨基酸组分百分含量在第1、5、10d显著降低,第3、15、20d明显上升,与游离氨基酸酸总含量变化趋势相反;不同处理诱导游离氨基酸组分百分含量的变化差异显著,但氨基酸变异系数小于1,表明其诱导氨基酸组分百分含量的波动变化是正常的。
2外源茉莉酸类化合物诱导两种落叶松针叶内次生物质的时序变化
外源茉莉酸类化合物诱导处理后落叶松针叶内单宁含量变化显著。兴安落叶松在处理后第1、10d针叶内单宁含量显著降低,第3、5d含量明显升高。长白落叶松在诱导后3、5、15d针叶内单宁含量明显增加,1、10、20d含量降低。
外源茉莉酸类化合物处理后,长白落叶松针叶内酚酸总含量在第1、3、5、10d降低,第15d显著升高,第20d显著降低的波动变化;兴安落叶松针叶内酚酸总含量变化总体呈现降低的趋势,增加不显著。兴安落叶松针叶用高效液相色谱仪共鉴定出11种酚酸;咖啡酸、阿魏酸、香豆酸、苯乙酸、水杨酸、香草酸变化趋势相似,呈现诱导前期含量显著降低,后期逐渐升高的动态变化;肉桂酸和原儿茶酸则呈现诱导前期升高,后期降低的变化规律。长白落叶松针叶共检测到12种酚酸;咖啡酸、苯乙酸、水杨酸含量呈现先降后升的动态变化;绿原酸、肉桂酸、原儿茶酸、香草酸含量是先增加后降低的波动变化。
3外源茉莉酸类化合物诱导两种落叶松防御蛋白活性的时序变化
外源茉莉酸类化合物诱导后,2种落叶松针叶内多酚氧化酶和苯丙氨酸解氨酶、蛋白酶抑制剂活性明显增加,呈现先升高后降低的波动性变化;不同处理时间各处理诱导作用显著,表明2种落叶松的诱导抗虫性产生。外源茉莉酸类化合物对2种落叶松胰凝乳蛋白酶抑制剂诱导作用较强;对兴安落叶松针叶内胰凝乳蛋白酶抑制剂诱导作用更明显。
4外源茉莉酸诱导两种落叶松对落叶松毛虫寄主选择行为的影响
落叶松毛虫在不同浓度茉莉酸处理的2种落叶松松苗上产卵量差异显著;对照树苗上的落卵量显著高于茉莉酸处理树苗上的卵量;3个不同浓度茉莉酸处理的松树苗上的落卵量差异不显著;茉莉酸处理树苗对落叶松毛虫产卵有很强驱避作用。
不同处理2种落叶松对落叶松毛虫的嗅觉反应影响显著;随茉莉酸处理时间的延长,落叶松对落叶松毛虫驱避性增强;随着茉莉酸处理浓度的增加,驱避性加大。与对照相比,2种落叶松处理后1d对落叶松毛虫没有明显的驱避作用;处理后3、5d驱避性显著。
5外源茉莉酸诱导两种落叶松对平腹小蜂寄主选择行为的影响
平腹小蜂对不同处理的落叶松上落叶松毛虫卵的寄生率,随处理时间的延长逐渐降低;但不同诱导时间内的各处理间寄生率差异均不显著。随着茉莉酸处理浓度的加大,平腹小蜂对落叶松毛虫卵的寄生率呈波动变化。
平腹小蜂对不同处理落叶松的嗅觉反应明显;随着茉莉酸处理浓度的增加,趋性增强;随着处理时间的延长,趋性呈波动变化。平腹小蜂对茉莉酸处理的第1、5d的松苗趋性显著。
本文较系统地研究了应用茉莉酸、茉莉酸甲酯和茉莉酮3种外源化合物诱导两种落叶松体内化学物质的时序变化,结果表明,外源化合物诱导落叶松体内营养物质可溶性糖和游离氨基酸含量降低,次生物质单宁和酚酸含量增加,蛋白酶抑制剂等防御蛋白活性增强,落叶松直接诱导抗虫性产生;且茉莉酸诱导落叶松对落叶松毛虫搜寻寄主产生明显驱避性,对平腹小蜂的寄主选择有显著引诱力,表明茉莉酸诱导落叶松产生间接抗虫性,从而增强落叶松自身的防御能力,减少植食昆虫的危害。茉莉酸、茉莉酸甲酯和茉莉酮能够在保持植物完好的情况下,模拟剪叶损伤和昆虫取食危害,诱导营养物质和次生物质的变化,对取食的昆虫产生毒害作用,又能诱导产生吸引害虫天敌的挥发性物质,其作为植物抗虫诱导剂,不污染环境,对天敌安全,保持生态平衡,有利于害虫的可持续控制。
Larch pine is main species of forest in northeast of China.Dendrolimus superans(Butler) boomed since 90s in the twentieth century and serious harmful result was seen.In this research, Larix gmelinii and Larix olgensis were selected as objects for research.Temporal variation of chemical compounds in two larch species,and effects of host plant selection of Dendrolimus superans and of Anastatus japonicus,which were performed to analysis through treatment of jasmonic acid(JA),methyl jasnonate(MJA) and cis-jasmone.The influence of exogenous jasmonates induced resistance of two larch species to insects and relative biochemical mechanism to Dendrolimus superan and Anastatus japonicus were analyzed about the factors, such as nutrient compounds,defensive proteins and secondary metabolites.It was important for improveing the healthy level of forest and control of population density of Dendrolimus superan.By using analysis of chemical compounds and biometrics,results of the research were listed below:
1.Changing of nutrient compounds in the needles of Larix gmelinii and L.olgensis induced with exogenous jasmonates
The contents of soluble sugar in the needles of Larix gmelinii and L.olgensis was fluctuant variation,and which decreased at the 1~(st),3~(rd),10~(th),15~(th),20~(th) day,and which increased at the 5~(th) day after exogenous jasmonates application.The variation of the contents of soluble sugar was no significant difference.
17 species of free amino acid were identified in the needles of Larix olgensis by HPLC.In contrasted with CK,the total contents of free amino acids increased at the 1~(st),5~(th),10~(th) day,and which decreased at the 3~(rd),15~(th),20~(th) day in the needles of L.olgensis after exogenous jasmonates application.The variation of the total contents of free amino acids was significantly different in the period of the experimentation.The component percents of free amino acids dropped at the 1~(st),5~(th),10~(th) day,and which raised at the 3~(rd),15~(th),20~(th) day in the needles of L. olgensis treated with jasmonates.But variational coefficient of free amino acids was smaller than one.Which sharp reversed with the variation of the total contents of free amino acids.The result indicated the variation of the component percents of free amino acids was normal fluctuation.
2.Changing of secondary metabolites in the needles of Larix gmelinii and L.olgensis induced with exogenous jasmonates
The contents of tannins changed significantly in the needles of Larix gmelinii and L. olgensis after exogenous jasmonates application.The contents of tannins in the needles of Larix gmelinii dropped at the 1~(st),10~(th) day,and which raised at the 3~(rd),5~(th) day.While which of L. olgensis decreased at the 1~(st),10~(th),20~(th) day,and which increased at the 3~(rd),5~(th),15~(th) day after application exogenous jasmonates.
The contents of phenolic acid in the needles of Larix olgensis dropped at the 1~(st),3~(rd),5~(th), 10~(th) day,and which raised at the 15~(th) day,then which dropped at the 20~(th) day after treated with exogenous jasmonates.The contents of phenolic acid in the needles of Larix gmelinii was decreasing in the period of the experimentation.11 species of phenolic acid were identified in the needles of Larix gmelinii by HPLC.There was similar variation for the contents of caffeic acid,ferulic acid,p-coumaric acid,phenylacetic acid,salicylic acid and vanillic acid.Which raised formerly,dropped later in needles of L.gmelinii induced with exogenous jasmonates. While which of cinnamic acid and chlorogenic acid sharp reversed with those.12 species of phenolic acid were identified in the needles of Larix olgensis by HPLC.The variation of the contents of caffeic acid,phenylacetic acid and salicylic acid dropped formerly,raised later in needles of L.olgensis treated with exogenous jasmonates.And which of cinnamic acid, chlorogenic acid,chlorogenic acid and vanillic acid sharp reversed with those.
3.Changing of the activity of defendable enzymes in the needles of Larix gmelinii and L.olgensis induced by exogenous jasmonates
The activities of PPO,PAL,trypsin inhibitor and chymotrypsin inhibitor increased significantly in the needles of Larix gmelinii and L.olgensis induced with exogenous jasmonates.The variation of their activity was waved in the period of the experimentation. While the induced function of the different concentration jasmonates was significantly different. Which indicated 2 species of larch pines formed the induced resistance to insect.
4.Effect of two larch species induced with exogenous jasmonic acid on the behavior of host plant selection of Dendrolimus superans
The number of oviposition of Dendrolimus superans was significantly different in the seedlings treated with different concentration JA.The number of egg in control seedlings were much more than which in the seedlings treated with JA.There was not different for the number of eggs in the seedlings after different concentration JA application.There was strongly drivable for the seedlings treated with JA to the oviposition behavior of Dendrolimus superans.
The drivable function of two larch species treated with JA to Dendrolimus superans was increased with prolonging days and increasing concentration treated with JA.Compared with CK,the drivable function of larch pine to Dendrolimus superans was not different at the 1~(st) day after jasmonic acid application,and which was significantly different at the 3~(rd),5~(th) day following jasmonic acid application.
5.Effect of two larch species induced with exogenous jasmonic acid on the behavior of host plant selection of Anastatus japonicus
The parasitism rate of Anastatus japonicus to Dendrolimus superans eggs was dropped with prolonging days treated with JA,and which was not different in the different treatments in the period of the experimentation.Which was waving with increasing concentration treated with JA.
The attractive function of two larch species treated with JA to Anastatus japonicus was increased with prolonging days,and which was waving with increasing concentration treated with JA.Which was significantly different at the 1~(st),5~(th) day following JA application.
Therefore,this paper expounded exogenous jasmonates induced two larch species to form direct and indirect resitance to insects.Which prevented or disturbed pest insects feeding,and protected natural enemies.While induced resistance to insect maintained the balance of forest ecosystem,which would play a key role in pest integrated control.
引文
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