不同茶树品种抗假眼小绿叶蝉机理研究
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摘要
假眼小绿叶蝉Empoasca vitis G the（同翅目，叶蝉科）是我国重要的茶树害虫，其为害严重影响到茶叶的质量和产量。由于成虫将卵产于植物组织内部，化学防治很难达到理想的效果。本文从品种抗虫性的角度出发，根据假眼小绿叶蝉在茶树品种上的田间虫口密度、生命周期、成虫繁殖力等指标，对供试品种的抗假眼小绿叶蝉水平进行分级。其次，利用电子刺探图谱（Electrical penetration graph，EPG）技术研究了假眼小绿叶蝉在不同抗虫水平茶树品种上的电生理反应，实现茶树品种抗性因子的定位。以此为基础，调查了不同抗虫水平茶树品种的外观形态、物理结构和生化特征的差异，并对相关生化成分进行生物测定，最终确定茶树对假眼小绿叶蝉的抗性因子和抗性机理。主要结果如下：
1.通过对天台(TT)等12个茶树品种的田间茶枝幼虫孵化数、笼内接虫产卵若虫孵化数和田间虫口密度的调查，筛选出恩标(EB)、竹山一号(ZS)、蓝天(LT)、斑竹园(BZY)、长兴紫笋(CX)、举岩(JY)、建德(JD)和德清(DQ)8个品种做进一步研究。
2.调查假眼小绿叶蝉在8个初筛品种上的生命周期、每雌产若量和若虫成活率，根据结果将其分为两大类。第Ⅰ类：举岩、德清、建德和长兴紫笋，抗假眼小绿叶蝉品种；第Ⅱ类：蓝天、斑竹园、竹山一号和恩标，感假眼小绿叶蝉品种。其中，举岩对假眼小绿叶蝉抗性最强，恩标对假眼小绿叶蝉抗性最弱。
3.运用EPG、TEM（Transmission electronic microscope）等昆虫行为学和植物组织学手段，明确假眼小绿叶蝉在茶树上的取食过程，首次证实假眼小绿叶蝉是破损细胞取食者(Cell rupture feeder)。假眼小绿叶蝉取食茶树过程中产生7种EPG波形。其中，主要波形Np (non-probing)、E1、E2和E3分别代表口针的非刺探、渠道开通和多细胞刺探、叶肉主动取食和维管束主动取食行为。假眼小绿叶蝉在茶树上一次完整的刺探行为通常为Np-E1-E2-E3-E1-Np。
4. EPG非连续型参数的分析结果显示，假眼小绿叶蝉在感虫品种上的取食时间长于抗虫品种，恩标和竹山一号与抗虫品种之间的差异达到显著水平；另一方面，EPG连续型参数的分析发现，长兴紫笋、德清和建德表面均存在抗性，而蓝天显然缺少这种抗性；举岩、建德和德清的叶片下表皮-叶肉层存在抗性，但感虫品种在这个层面上均没有抗性；举岩、长兴紫笋和建德的叶肉组织都表现出对假眼小绿叶蝉的抗性，斑竹园和恩标却明显缺乏这种抗性。
5.外观形态上，抗虫品种与感虫品种的当年生新梢色泽（抗虫品种深于感虫品种）、一芽三叶长度（抗虫品种＜感虫品种）、叶片大小（抗虫品种＜感虫品种）、叶片硬度（抗虫品种＞感虫品种）和叶身伸展状态都存在差异。物理结构上，抗虫品种的茸毛长度和密度都显著大于感虫品种；抗虫品种的蜡质含量也显著高于除恩标以外的其它感虫品种；此外，抗虫品种的下表皮厚度均大于感虫品种。生化成分上，抗虫品种的茶多酚和绿原酸含量显著低于感虫品种，相反，γ-氨基丁酸含量却明显高于感虫品种。
6.茶树品种理化指标与假眼小绿叶蝉每雌产若量和生命周期的相关性分析显示，茸毛长度、茸毛密度、叶表皮蜡质含量和γ-氨基丁酸含量与假眼小绿叶蝉每雌产若量呈负相关，相关性均达到极显著；下表皮厚度与每雌产若量也呈显著负相关；茶多酚与假眼小绿叶蝉的每雌产若量呈极显著正相关。另一方面，茸毛长度和茸毛密度与假眼小绿叶蝉生命周期呈极显著正相关；下表皮厚度与假眼小绿叶蝉的生命周期呈显著正相关；茶多酚与假眼小绿叶蝉的生命周期呈显著负相关。绿原酸与假眼小绿叶蝉每雌产若量和生命周期均没有相关关系。
7.配制全纯人工饲料进行γ-氨基丁酸、茶多酚、绿原酸、天冬氨酸和茶氨酸的生物测定。高浓度茶多酚对假眼小绿叶蝉若虫的生长和存活表现抑制作用，与茶树品种中茶多酚含量和假眼小绿叶蝉生命周期的相关性分析结果不一致；高浓度γ-氨基丁酸对假眼小绿叶蝉若虫的生长发育和成活率有负面影响；虽然假眼小绿叶蝉的成活率不受绿原酸浓度的影响，但第1、2龄历期随其浓度的逐步升高呈现出递增趋势。另外，天冬氨酸与茶树抗虫性没有相关关系，但假眼小绿叶蝉1、2龄历期随其浓度的增加而缩短；茶氨酸对假眼小绿叶蝉的存活和龄期没有影响。
8.茶树不同抗虫品种具有不同的抗虫机理。蜡质含量、茸毛长度和密度决定表面抗性，其中，蜡质含量是关键因子；下表皮厚度是形成叶片下表皮-叶肉层抗性的重要原因；生化成分可能是叶肉组织抗性的主要决定因素。而且，叶肉层面的化学抗性是决定茶树抗假眼小绿叶蝉水平的关键作用因子，γ-氨基丁酸可能正是关键因子之一。
Tea green leafhopper Empoasca vitis G the (Hemiptera: Cicadellidae), is an importantpest of tea plant in China, and seriously affect the tea quality and yield. Because the eggs werelaid in plant tissues, chemical pesticide applications usually fail to achieve the desired result.Standing on the piont of view of variety resistance, the tea cultivars were grouped basing onthe results of leafhopper’s field population density, life cycle duration, adult reproduction, etc..Secondly, resistance factors location of tea cultivars were achieved through the study ofleafhopper’s feeding behavior on different cultivars, which was investgited by electricalpenetration graph (EPG) technique. Based on these results, the morphology, anatomicalstucture and biochemical compositions of different tea cultivars were analyzed, as for thecompositions with significantly different content between resistant and susceptible cultivars,the bioassay experiment were then carried out to determine the resistance factors and theresistance mechanism in resistant tea cultivars. The main results are as follows:
1. The eight tea cultivars including En biao (EB), Zhu shan1(ZS), Lan tian (LT), Banzhu yuan (BZY), Chang xing zi sun (CX), Ju yan (JY), Jian de (JD) and De qing (DQ) wereselected for the further study according to the results of hatched nymph number from the fieldtea shoots, hatched nymph number from the artificially infested tea shoots and fieldpopulation density on12tea cultivars such as TT etc.
2. Life cycle duration, nymph number from per female and nymph survival rate of E.vitis on the eight tea cultivars were investigated, and two groups were classified according towhich results. Ⅰ: resistant group including JY, DQ, JD and CX, which were resistant to E.vitis; Ⅱ: susceptible group including LT, BZY, ZS and EB, which were susceptible to E. vitis.JY is the most resistant to E. vitis and EB is the most susceptible to E. vitis in the eight teacultivars.
3. The feeding behaviors of E. vitis on tea plant were identified by using planthistological and insect behavioral mothods, which including EPG, transmission electronicmicroscope and other techniques. Thus the result firstly comfirmed that E. vitis blongs to cellrupture feeder. Seven EPG waveforms were produced by E. vitis on tea plant, the mainwaveforms Np, E1, E2and E3were correlated with stylet non-probing, channel-cutting and multiple-cell laceration, mesophyll active ingestion and vascular active ingestion, respectively.The whole process of a probe was used to performing as Np-E1-E2-E3-E1-Np.
4. The result of EPG non-sequential parameters analysis showed that the ingestiondurations of E. vitis on susceptible cultivars are longer than resistant cultivars, although onlyEB and ZS were significantly longer than all of resistant cultivars. On the other hand, EPGsequential parameters caculation showed that the surface resistance was existed on CX, DQand JD, but this resistance was absent from LT; the epidermis-mesophyll of JY, JD and DQshowed resistace to E. vitis, but all of the four susceptible cultivars were lack of the resistance;the mesophyll tissue of JY, CX and JD were resistant to E. vitis, but this resistance have notbeen discovered in BZY and EB.
5. In morphology, the difference between resistant and susceptible cultivars wasdiscovered in new tea shoots color (resistant cultivars showed darker than susceptiblecultivars), length of new tea shoots with one bud and three leaves (resistant cultivars＜susceptible cultivars), leaves size (resistant cultivars＜susceptible cultivars), leaveshardness (resistant cultivars＞susceptible cultivars) and extension of leaf body. As forphysical structure, the trichome length and density of leaves of resistant cultivars weresignificantly higher than of susceptible ones; all resistant cultivals had significantly higherwax content than susceptible cultivars except EB. Additionally, lower epidermal thickness ofresistant cultivars were higher than that of susceptible ones. For the biochemical compositions,the content of tea polyphenols and chlorogenic acid in resistant cultivars were significantlylower than susceptible cultivars. On the contrary, the content of γ-aminobutyric acid inresistant cultivars was obviously higher than that of susceptible cultivars; theanine content inDQ, JY and CX was significantly lower than all susceptible cultivars; moreover, the contentof aspartic acid of resistant cultivars was lower than all susceptible cultivars, although thevalues of resistant cultivars and two susceptible cultivars ZS and BZY reached significantlevel.
6. Correlations of physical and chemical data of tea cultivars with life cycle duration andnymph number of each female of E. vitis were analyzed. The result showed that trichomelength and density, wax content and γ-aminobutyric acid content were negatively correlatedwith the nymph number from per female, this correlation reached highly significant level; leaflower epidermal thickness showed significantly negative correlation with the nymph numberfrom per female, but content of tea polyphenols was positively correlated to it. On the otherhand, trichome length and density and leaf lower epidermal thickness showed significantlypositive correlation with life cycle duration of E. vitis. Contrarily, the significantly negativecorrelation was discovered between the content of tea polyphenols and life cycle duration of E. vitis.
7. Artificial diets were made for the biaassay analysis of γ-aminobutyric acid. Theinhibition role was discovered in high concentration of tea polyphenols on growth andsurvival rate of E. vitis, which was inconsistent with the previous result of correlation analysis.High concentration of γ-aminobutyric acid had a negative impact on the growth and thesurvival rate of E. vitis. Although chlorogenic acid has not been found any certain correlationwith the survival rate of E. vitis, the duration of1stand2ndinstar of E. vitis were graduallyincreased with the increase of the content of chlorogenic acid in diet. Additionally, althoughaspartic acid was not related to tea resistance, the duration of1stand2ndinstar of E. vitis wasshortened with the concentration increasing; the survive and development of E. vitis have notbeen influenced by the changes of concentration of theanine.
8. Different tea cultivars have different resistance mechanisms to E.vitis. The surfaceresistance was determined by wax content, trichome length and density. Thickness of thelower epidermis was the important reason for epidermis-mesophyll layer resistance.Biochemical components may be the main determinant of the mesophyll resistance. Moreover,the chemical resistance was the key role for determining the resistance level of tea cultivars toE. vitis, γ-aminobutyric acid may be one of the key factors of the chemical resistance.
1.通过对天台(TT)等12个茶树品种的田间茶枝幼虫孵化数、笼内接虫产卵若虫孵化数和田间虫口密度的调查，筛选出恩标(EB)、竹山一号(ZS)、蓝天(LT)、斑竹园(BZY)、长兴紫笋(CX)、举岩(JY)、建德(JD)和德清(DQ)8个品种做进一步研究。
2.调查假眼小绿叶蝉在8个初筛品种上的生命周期、每雌产若量和若虫成活率，根据结果将其分为两大类。第Ⅰ类：举岩、德清、建德和长兴紫笋，抗假眼小绿叶蝉品种；第Ⅱ类：蓝天、斑竹园、竹山一号和恩标，感假眼小绿叶蝉品种。其中，举岩对假眼小绿叶蝉抗性最强，恩标对假眼小绿叶蝉抗性最弱。
3.运用EPG、TEM（Transmission electronic microscope）等昆虫行为学和植物组织学手段，明确假眼小绿叶蝉在茶树上的取食过程，首次证实假眼小绿叶蝉是破损细胞取食者(Cell rupture feeder)。假眼小绿叶蝉取食茶树过程中产生7种EPG波形。其中，主要波形Np (non-probing)、E1、E2和E3分别代表口针的非刺探、渠道开通和多细胞刺探、叶肉主动取食和维管束主动取食行为。假眼小绿叶蝉在茶树上一次完整的刺探行为通常为Np-E1-E2-E3-E1-Np。
4. EPG非连续型参数的分析结果显示，假眼小绿叶蝉在感虫品种上的取食时间长于抗虫品种，恩标和竹山一号与抗虫品种之间的差异达到显著水平；另一方面，EPG连续型参数的分析发现，长兴紫笋、德清和建德表面均存在抗性，而蓝天显然缺少这种抗性；举岩、建德和德清的叶片下表皮-叶肉层存在抗性，但感虫品种在这个层面上均没有抗性；举岩、长兴紫笋和建德的叶肉组织都表现出对假眼小绿叶蝉的抗性，斑竹园和恩标却明显缺乏这种抗性。
5.外观形态上，抗虫品种与感虫品种的当年生新梢色泽（抗虫品种深于感虫品种）、一芽三叶长度（抗虫品种＜感虫品种）、叶片大小（抗虫品种＜感虫品种）、叶片硬度（抗虫品种＞感虫品种）和叶身伸展状态都存在差异。物理结构上，抗虫品种的茸毛长度和密度都显著大于感虫品种；抗虫品种的蜡质含量也显著高于除恩标以外的其它感虫品种；此外，抗虫品种的下表皮厚度均大于感虫品种。生化成分上，抗虫品种的茶多酚和绿原酸含量显著低于感虫品种，相反，γ-氨基丁酸含量却明显高于感虫品种。
6.茶树品种理化指标与假眼小绿叶蝉每雌产若量和生命周期的相关性分析显示，茸毛长度、茸毛密度、叶表皮蜡质含量和γ-氨基丁酸含量与假眼小绿叶蝉每雌产若量呈负相关，相关性均达到极显著；下表皮厚度与每雌产若量也呈显著负相关；茶多酚与假眼小绿叶蝉的每雌产若量呈极显著正相关。另一方面，茸毛长度和茸毛密度与假眼小绿叶蝉生命周期呈极显著正相关；下表皮厚度与假眼小绿叶蝉的生命周期呈显著正相关；茶多酚与假眼小绿叶蝉的生命周期呈显著负相关。绿原酸与假眼小绿叶蝉每雌产若量和生命周期均没有相关关系。
7.配制全纯人工饲料进行γ-氨基丁酸、茶多酚、绿原酸、天冬氨酸和茶氨酸的生物测定。高浓度茶多酚对假眼小绿叶蝉若虫的生长和存活表现抑制作用，与茶树品种中茶多酚含量和假眼小绿叶蝉生命周期的相关性分析结果不一致；高浓度γ-氨基丁酸对假眼小绿叶蝉若虫的生长发育和成活率有负面影响；虽然假眼小绿叶蝉的成活率不受绿原酸浓度的影响，但第1、2龄历期随其浓度的逐步升高呈现出递增趋势。另外，天冬氨酸与茶树抗虫性没有相关关系，但假眼小绿叶蝉1、2龄历期随其浓度的增加而缩短；茶氨酸对假眼小绿叶蝉的存活和龄期没有影响。
8.茶树不同抗虫品种具有不同的抗虫机理。蜡质含量、茸毛长度和密度决定表面抗性，其中，蜡质含量是关键因子；下表皮厚度是形成叶片下表皮-叶肉层抗性的重要原因；生化成分可能是叶肉组织抗性的主要决定因素。而且，叶肉层面的化学抗性是决定茶树抗假眼小绿叶蝉水平的关键作用因子，γ-氨基丁酸可能正是关键因子之一。
Tea green leafhopper Empoasca vitis G the (Hemiptera: Cicadellidae), is an importantpest of tea plant in China, and seriously affect the tea quality and yield. Because the eggs werelaid in plant tissues, chemical pesticide applications usually fail to achieve the desired result.Standing on the piont of view of variety resistance, the tea cultivars were grouped basing onthe results of leafhopper’s field population density, life cycle duration, adult reproduction, etc..Secondly, resistance factors location of tea cultivars were achieved through the study ofleafhopper’s feeding behavior on different cultivars, which was investgited by electricalpenetration graph (EPG) technique. Based on these results, the morphology, anatomicalstucture and biochemical compositions of different tea cultivars were analyzed, as for thecompositions with significantly different content between resistant and susceptible cultivars,the bioassay experiment were then carried out to determine the resistance factors and theresistance mechanism in resistant tea cultivars. The main results are as follows:
1. The eight tea cultivars including En biao (EB), Zhu shan1(ZS), Lan tian (LT), Banzhu yuan (BZY), Chang xing zi sun (CX), Ju yan (JY), Jian de (JD) and De qing (DQ) wereselected for the further study according to the results of hatched nymph number from the fieldtea shoots, hatched nymph number from the artificially infested tea shoots and fieldpopulation density on12tea cultivars such as TT etc.
2. Life cycle duration, nymph number from per female and nymph survival rate of E.vitis on the eight tea cultivars were investigated, and two groups were classified according towhich results. Ⅰ: resistant group including JY, DQ, JD and CX, which were resistant to E.vitis; Ⅱ: susceptible group including LT, BZY, ZS and EB, which were susceptible to E. vitis.JY is the most resistant to E. vitis and EB is the most susceptible to E. vitis in the eight teacultivars.
3. The feeding behaviors of E. vitis on tea plant were identified by using planthistological and insect behavioral mothods, which including EPG, transmission electronicmicroscope and other techniques. Thus the result firstly comfirmed that E. vitis blongs to cellrupture feeder. Seven EPG waveforms were produced by E. vitis on tea plant, the mainwaveforms Np, E1, E2and E3were correlated with stylet non-probing, channel-cutting and multiple-cell laceration, mesophyll active ingestion and vascular active ingestion, respectively.The whole process of a probe was used to performing as Np-E1-E2-E3-E1-Np.
4. The result of EPG non-sequential parameters analysis showed that the ingestiondurations of E. vitis on susceptible cultivars are longer than resistant cultivars, although onlyEB and ZS were significantly longer than all of resistant cultivars. On the other hand, EPGsequential parameters caculation showed that the surface resistance was existed on CX, DQand JD, but this resistance was absent from LT; the epidermis-mesophyll of JY, JD and DQshowed resistace to E. vitis, but all of the four susceptible cultivars were lack of the resistance;the mesophyll tissue of JY, CX and JD were resistant to E. vitis, but this resistance have notbeen discovered in BZY and EB.
5. In morphology, the difference between resistant and susceptible cultivars wasdiscovered in new tea shoots color (resistant cultivars showed darker than susceptiblecultivars), length of new tea shoots with one bud and three leaves (resistant cultivars＜susceptible cultivars), leaves size (resistant cultivars＜susceptible cultivars), leaveshardness (resistant cultivars＞susceptible cultivars) and extension of leaf body. As forphysical structure, the trichome length and density of leaves of resistant cultivars weresignificantly higher than of susceptible ones; all resistant cultivals had significantly higherwax content than susceptible cultivars except EB. Additionally, lower epidermal thickness ofresistant cultivars were higher than that of susceptible ones. For the biochemical compositions,the content of tea polyphenols and chlorogenic acid in resistant cultivars were significantlylower than susceptible cultivars. On the contrary, the content of γ-aminobutyric acid inresistant cultivars was obviously higher than that of susceptible cultivars; theanine content inDQ, JY and CX was significantly lower than all susceptible cultivars; moreover, the contentof aspartic acid of resistant cultivars was lower than all susceptible cultivars, although thevalues of resistant cultivars and two susceptible cultivars ZS and BZY reached significantlevel.
6. Correlations of physical and chemical data of tea cultivars with life cycle duration andnymph number of each female of E. vitis were analyzed. The result showed that trichomelength and density, wax content and γ-aminobutyric acid content were negatively correlatedwith the nymph number from per female, this correlation reached highly significant level; leaflower epidermal thickness showed significantly negative correlation with the nymph numberfrom per female, but content of tea polyphenols was positively correlated to it. On the otherhand, trichome length and density and leaf lower epidermal thickness showed significantlypositive correlation with life cycle duration of E. vitis. Contrarily, the significantly negativecorrelation was discovered between the content of tea polyphenols and life cycle duration of E. vitis.
7. Artificial diets were made for the biaassay analysis of γ-aminobutyric acid. Theinhibition role was discovered in high concentration of tea polyphenols on growth andsurvival rate of E. vitis, which was inconsistent with the previous result of correlation analysis.High concentration of γ-aminobutyric acid had a negative impact on the growth and thesurvival rate of E. vitis. Although chlorogenic acid has not been found any certain correlationwith the survival rate of E. vitis, the duration of1stand2ndinstar of E. vitis were graduallyincreased with the increase of the content of chlorogenic acid in diet. Additionally, althoughaspartic acid was not related to tea resistance, the duration of1stand2ndinstar of E. vitis wasshortened with the concentration increasing; the survive and development of E. vitis have notbeen influenced by the changes of concentration of theanine.
8. Different tea cultivars have different resistance mechanisms to E.vitis. The surfaceresistance was determined by wax content, trichome length and density. Thickness of thelower epidermis was the important reason for epidermis-mesophyll layer resistance.Biochemical components may be the main determinant of the mesophyll resistance. Moreover,the chemical resistance was the key role for determining the resistance level of tea cultivars toE. vitis, γ-aminobutyric acid may be one of the key factors of the chemical resistance.
引文
彩万志,庞雄飞,花保祯.2001.普通昆虫学.北京:中国农业大学出版社:127
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