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间作套种对小白菜田节肢动物群落及主要害虫的影响
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摘要
在化学农药产生之前,间作套种是害虫防治的主要手段之一。由于化学农药所产生的一系列弊端,人类开始寻找安全有效的控制措施。近年来,许多昆虫学家和生态学家根据他们的研究工作,认为通过采用合理的耕作制度和栽培措施,可以保护和强化农田生物多样性,恢复和重组农田生态系统的动态平衡,有利于农业害虫的生态控制和综合治理(Andow,1991;Altieri,1987,1994;Douglas等,2000)。因此,利用生物多样性控制作物虫害已成为当前的一个研究热点。本文从群落生态学的角度出发,研究不同作物间作套种对节肢动物群落天敌功能团的影响,旨在揭示多样化生境对保护和利用自然天敌生态学机理,为菜田害虫的持续控制提供理论依据。同时观察测定间作套种下对小白菜新陈代谢、对小菜蛾生长发育和三种农药相关酶系的影响。
     1.间作套种对小白菜田节肢动物群落的影响
     2004年于福建省琅歧经济开发区调查间作套种对菜田节肢动物群落的影响,间作套种系统分别为大蒜间作小白菜(CG1),大蒜套种小白菜(CG2),春菜间作小白菜(CL1),春菜套种小白菜(CL2),包菜间作小白菜(CB1)和包菜套种小白菜(CB2),以单一种植小白菜为对照(CK)。调查结果显示:
     (1)共采集到53392头个体,包括昆虫纲、蛛形纲和多足纲,经鉴定分别属于98科,111属,175个物种。整体上,单作田和多作田之间目、科、属之间差异不显著,但是科总数,多样性和均匀性指数,单作田明显低于多作田,差异显著。CL1与CL2群落相似性最高(0.71),相似性最低为CB2与CK(0.10)。
     (2)根据各个功能团划分,捕食性功能团物种数最高(59种),占总物种数的33.71%,而中性昆虫物种数最低(27种,15.43%)。各功能团丰盛度依次为:中性昆虫(占总个体数的62.13%)>植食者(27.24%)>捕食者(6.79%)>寄生者(3.84%)。其中,单作田中性昆虫数量明显高于多作田。
     (3)整体上多作田植食者的物种丰富度高于单作田,丰盛度最高为CG1和CB1。多样性指数,除了CG2,其它各个多作田均明显高于单作田。CL1,CB1和CK优势度指数没有显著差异。CB1,CB2和CK田内小菜蛾种群动态相似;黄曲条跳甲种群数量多作田低于单作田:蚜虫在生长前期多作田低于单作田,但生长后期高于单作田。
     (4)寄生性昆虫物种丰富度依次为CL1(33种)>CG1(32种)>CG2(27种)>CL1(26种)>CK(23种)>CB2(21种)>CB1(20种)。寄生者数量最高为CG1,CG2和L2,明显高于CK。多样性指数多作田明显高于单作田;但是均匀性和优势集中性指数多作田和单作田之间没有显著差异。其中,从第16天到第32天多作田蚜小蜂数量明显高于单作田。多作系统比单作系统支持更多的菜蚜茧蜂。
     (5)捕食功能团,物种丰富度最高为CG1,最低为CK,个体数量最高的为CG1(141.67头/小区),最低为(97.67头/小区)。多作田除CL1多样性指数均显著高于单作田。CL1均匀性指数显著低于其他各田,多作田的优势集中指数明显高于单作田。所采集的捕食者中,大部分属于蜘蛛类,其中球腹蛛科占34.04%,狼蛛占30.57%。CG1,CL1和CB1的狼蛛数量在各调查时间内均高于单作田。
     (6)单作田的中性昆虫数量,及优势集中性指数显著高于多作田;单作田物种数、多样性、均匀性指数显著低于多作田。丰盛度最高的物种为跳虫科,占87.35%,紧接着为双翅目昆虫。CK和CB2田的双翅目昆虫较少。
     本实验说明小白菜与非十字花科作物间作套种有利于提高天敌(包括捕食性和寄生性天敌)的多样性和丰盛度。多样化种植对天敌的影响比对植食性昆虫的影响更大。
     2.间作套种对小白菜新陈代谢的影响
     研究小白菜分别与春菜(CL)和大蒜(CG)间作套种下对小白菜各个生理生化指标的影响,包括总蛋白,可溶性糖,还原糖、硝态氮、叶绿素含量,并测定了4种酶活性,分别为硝酸还原酶(NR)、过氧化物酶(POD)、过氧化氢酶(CAT)和超氧化物歧化酶(SOD)。结果显示间作套种显著提高可溶性蛋白和硝态氮含量,春菜与小白菜(CL)套种明显提高小白菜生长后期叶绿素a的含量。总体上,间作套种对可溶性含量没有影响;还原性糖含量随着生长期的不同而变化。大蒜和小白菜套种显著提高NR、SOD和CAT酶活力。春菜与小白菜套种显著提高SOD,POD和CAT活力。本实验为小白菜与非十字花科作物间作套种提高植物营养水平和植物代谢水平提供了理论依据。
     3.间作套种对小菜蛾生长发育的影响
     研究小白菜分别与春菜(CL)或大蒜(CG)间作套种下对小菜蛾5个世代的生长发育历期、蛹重、成虫羽化率、产卵量以及卵孵化率的影响,以小白菜单作田(CK)饲养下的小菜蛾为对照。结果显示:春菜与小白菜间作套种显著延长小菜蛾各个世代幼虫生长发育历期,大蒜的影响达3个世代(F1,F4和F5)。化蛹率(88%-91%)三者之间没有显著差异,但是CG明显提高了小菜蛾3个世代(F3,F4和F5)的蛹重(平均5.20 mg/蛹),而单作田为(平均4.60 mg/蛹)。CG和CL均显著降低成虫羽化率,分别为92%和77%,对照为96%。各个世代的产卵期和产卵量表现出不稳定变化。CG和CK卵历期没有显著差异,为2.4—3.5天,但是CL比CK显著延长了小菜蛾3个世代的卵历期(F2,F3和F5)。因此,合适的间作套种可通过延长小菜蛾发育历期,降低羽化率、卵孵化率或产卵量来降低每年的世代种群。
     4.间作套种对小菜蛾三种解毒酶活性的测定
     测定小白菜分别与大蒜和春菜间作条件下对小菜蛾3龄幼虫的三种解毒酶乙酰胆碱酯酶(AChE),羧酸酯酶(CarE)和谷光甘肽S转移酶(GST)的活力影响。结果表明:与单一种植田小白菜比较,小白菜间作明显降低了小菜蛾三种酶活力,说明间作套种可推迟小菜蛾对有机磷、氨基甲酸酯类和拟除虫菊酯类农药的抗性产生速度。
Intercropping is a widely employed conventional practice throughout the world. It can enhance the nutrient cycle, soil fertility utilization, land and sunlight use efficiency, crop productivity and pest damage reduction. This dissertation presents the results on the study of the effects of intercropping garlic (Allium sativum), lettuce (Lactuca sativa) or green cabbage (Brassica oleracea) with Chinese cabbage (B. chinensis) on the arthropod community, metabolism of the Chinese cabbage, development and growth of diamondback moth (Plutella xylostella) (DBM), as well as the enzyme activities in the. DBM's larvae .
     1.Effects of intercropping on the structure of arthropod communities in Chinese cabbage fields
     Field trials were carried out at Langqi, Fujian, P. R. China in 2004, to determine the effects of intercropping Chinese cabbage with another crucifer species, green cabbage, and two non-cruciferous crops, i.e., garlic and lettuce, on the arthropod communities. Two intercropping plot designs were used in this study. Design 1 planted 2 ridges of the Chinese cabbages with 1 ridge of accompanying plants. The accompanying crops were garlic (CG1) , lettuce (CL1) or green cabbage (CB1) . Design 2 under-sowed the Chinese cabbages in the center (100 cm wide) with garlic (CG2) , lettuce (CL2) or green cabbage (CB2) on both edges (25 cm wide) of a same ridge. The monoculture of the Chinese cabbages (CK) was the control for comparisons in both design 1 and 2.The results obtained from the experiments showed that:
     (1) A total of 53,392 arthropod individuals belonging to 3 classes, 98 families, with 111 genera and 175 species were collected. In general, no significant differences were found in the number of orders (11 or 12 orders), families or genera between CK and the various intercropped treatments on the experimental lot; but significantly lower number of species was observed in CK than the intercropping treatments;
     (2) The greatest number of the species collected belonged to the predator guild (i.e., 59 species and 33.71% of the total). The neutral insect guild had the least number of species (i.e., 27 species at 15.43%). The abundance of each guild was as followed: neutral insects (i.e., 62.13% of the total number of individual insects)>herbivores (i.e., 27.24%)> predators (i.e., 6.79%)>parasitoid (i.e., 3.84%);
     (3) Species richness of herbivorous guild was higher in the intercropping treatments than CK. There were significant differences in diversity index between CK and the intercropping treatments, except CG2.No significant differences on dominance indices were found in CL1, CB1 and CK.
     (4) Species richness of parasitoid guild decreased in the order of CL1 (33 species)>CG1 (32 species)>CG2 (27species)> CL1 (26 species)> CK (23 species)>CB2 (21 species)>CB1(20 species). Significantly higher abundance of parasitoids was found in CG1, CL1 and CG2 than CK. Although significantly higher diversity index of parasitoid species was found in intercropping treatments than CK, no significant difference of evenness and dominant indices were found between them.
     (5) In regard to the predatory guild. the highest species richness was found in CG1 and the lowest in CK. The highest abundance was found in CL1 (i.e.. 141.67 predators plot). whereas the lowest in CB1 (i.e.. 97.67 predators plot). Except CL1.significantly higher diversity indices were found in intercropping treatments than CK. The majority of spiders found in the fields belonged tothe Theridiidae family (i.e.. 34.04%) and Lycosida family (i.e.. 30.57%).
     (6) Higher abundance and dominance indices in neutral insect guild and lower species richness, diversity and evenness indices were found in CK than the intercropping treatments. The most abundant species in the neutral insect guild were springtails (i.e., 87.35%), followed by dipters of the Ephyridae family. Overall, fewer dipters were caught in CK and CB2.
     Therefore, it was suggested that intercropping the Chinese cabbages with non-cruciferous crops might increase the abundance and diversity of the predators and parasitoids. and that the effects brought about by intercropping might be more pronounced on the natural enemies than on the herbivores.
     2.Effects of intercropping with non-crucifers on the metabolism of Chinese cabbage
     The physiological and biochemical indices of the Chinese cabbage consisted of protein, sugar, reducing sugar. nitrate-N and chlorophyll contents, as well as four plant metabolic indicators, including nitrate reductase (NR). superoxide dismutases (SOD). peroxidases (POD) and catalase (CAT) activities, were compared among the Chinese cabbages grown under different intercropping treatments. In general, higher average protein and nitrate-N contents were found in the intercropped Chinese cabbages than in CK. Significantly higher chlorophyll-a content was found in the cabbages from CL than CK. during the later half of the growing season. No significant differences in sugar concentration were found in CG and CL. as compared to CK. Reducing sugar content varied over the growing period of the Chinese cabbages in CG and CL. In terms of enzyme activity, NR, SOD and CAT activities increased in CG, and SOD, POD and CAT activities increased in CL, as compared to CK. The results suggested that the Chinese cabbages intercropped with non-crucifer plants not only increased plant nutritional quality, but also affected the plant's metabolism.
     3.Effects of intercropping garlic or lettuce with Chinese cabbage on the growth and development of Plutella xylostella (Lepidoptera: Plutellidae)
     During 2005, the effects of intercropping on the growth and development of the diamondback moth over 5 generations were examined in Fuzhou. China. The duration of the larval development was generally significantly longer in both intercropping treatments than CK. Intercropping did not affect the larvae pupating rate. However. it did affect the weight of pupa. Pupae in the intercropping lots tended to be 10%-15% larger for three of the five generations. Despite the size increase, pupa's mortality rate was 2-5 times greater in the intercropping treatments than CK. While female DBM laid similar numbers of eggs in all treatments, the egg stage in 3 of the 5 generations was significantly longer in the lettuce intercropping. This suggested that intercropping with different accompanying crop might cause a reduction in the annual number of DBM generations due to prolonged developmental stage and decreased adult emergence rate.
     4.DBM enzyme activity in relation to intercropping
     Activities of three insecticide related enzymes, acetylcholinesterase (AChE), carboxylesterase (CarE) and glutathione S-transferase (GST), of DBM were compared among third instar larvae for 5 successive generations that had been fed on the Chinese cabbages from one of the 3 treatments: CL, CG or CK. Enzyme activities in 30 larvae from each treatment were measured by spectrophotometry. The AChE, CarE and GST activities in the larvae, which were fed with treatment cabbage foliage, decreased significantly, as compared to CK. Therefore, potentially, intercropping could increase delay DBM's insecticide sensitivity on organophosphate. carbamate and pyrethroid insecticides.
     In conclusion, our study suggested that intercropping the Chinese cabbage with garlic or lettuce could increase the efficiency of nature enemies and decrease that of pest insects. Proper intercropping could not only reduce DBM population but also increase the insect's susceptibility to insecticides by disrupting its metabolism.
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