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虫害挥发物对邻近枣树直接防御反应的作用
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摘要
枣镰翅小卷蛾(Ancylis Sativa.Liu,又名枣粘虫),鳞翅目卷蛾科,是危害枣树的重要害虫,在枣树萌芽期和成熟期严重危害,造成二次发芽或严重影响果实品质。本文以枣树(Ziziphus jujuba Mill.)—枣蠊翅小卷蛾为材料,研究枣镰翅小卷蛾幼虫取食后,虫害枣树挥发物(herbivore-induced plant volatiles,HIPVs)对邻近枣树直接防御反应的影响,探讨损伤信号在枣树间的传递及直接防御作用。研究结论如下:
     虫害3 d后,与虫害枝通气的株内和株间邻近枝蛋白酶抑制剂(PI)、多酚氧化酶(PPO)和脂氧合酶(LOX)活性都显著高于未通气的邻近枝条,且与虫害枝不通气的邻近枝3种酶活性和对照差异不显著。与虫害枝通气条件下,株内邻近枝3种酶活性略高于株间邻近枝。这些结果表明,虫害枣树挥发物作为损伤信号主要通过空气在株间进行传递。
     虫害24 h后,虫害株与邻近株通气5 d内,枣树PI、PPO和LOX活性大小依次是虫害株>邻近株>对照株;通气的邻近株内3种防御酶活性显著高于对照,并且PI和LOX活性先上升后下降,而PPO活性一直保持相对较高的水平。茉莉酸甲酯(MeJA)处理与枣粘虫取食试验结果基本一致,而绿叶挥发物(GLVs)处理的枣树3种酶活性增高迅速,2 h后就达到最大值,但高活性持续时间只有48 h。这表明虫害挥发物、MeJA诱导挥发物(MeJA-VOCs)和GLVs都可作为损伤信号通过空气传递到邻近枣树,激发邻近枣树的防御反应,且HIPVs和MeJA-VOCs激发的直接防御反应可持续5 d以上,而GLVs诱导的防御反应持续时间仅为2 d。
     虫害3 d后,在自然条件下的虫害株和邻近株3种酶活性均显著高于对照株,且邻近株PI活性呈曲折下降型变化,PPO活性呈“W”型变化,LOX活性则呈“N”形变化。从防御反应持续时间上看,邻近株PI高活性(与对照株相比)可持续5 d,PPO和LOX高活性可持续5 d以上;而虫害株3种酶的高活性持续时间更长,强度更大。
     综上所述,本文探明了田间条件下,枣树株间信号的传递途径和作用方式,探讨了虫害挥发物引起邻近枣树防御反应的作用机制。
Jujube armyworm (Ancylis Sativa. Liu), Lepidoptera, Tortricidae, is an important infested pest against jujube. Larvae can seriously feed leaves in bud stage and fruit mature stage, causing the second germination or fruit quality drop. Taking jujube and A. Sativa as materials, we researched direct defensive effects of neibouring jujube exposed by herbivore-induced plant volatiles (HIPVs) released from A. Sativa infested-jujube, explored transmission of wound-signal and direct defensive function in jujubes. The results as follow:
     After three days' feeding, PI, PPO and LOX activities in adjacent intra-shoot and inter-shoot aerating with herbivored-shoot were significantly higher than those adjacent shoots with no aeration. Three enzyme activities in no aerating shoot had no significant difference with uninfested control shoot. Moreover, in the condition of aeration, three enzyme activities in intra-shoot slightly higher than inter-shoot. These results indicate that HIPVs are an vital airborne wound-signal, primarily transmit by atmosphere among plants.
     After 24 hours' attack, PI, PPO and LOX amounts were greastest in A. Sativa feed plant among five days ventilation. In aerating neighboring jujube, three defensive enzymes significantly higher than the undamaged control jujube, and PI and LOX activities increased and then decreased, while PPO activity remained at a high level. Methyl jasmonate (MeJA) treatment had the same effect with A. Sativa feeding, however, though three enzyme activities increased rapidly to maximumafter 2 hour's GLVs exposedness, the duration of high activities were only 48 hours in contrast to 5 days exposed by HIPVs and MeJA induced volatiles (MeJA-VOCs). This illustrated that HIPVs, MeJA-VOCs and GLVs were important damage-signals, which could transmit to nearby plant by airborne, priming defense response in nearby plants.
     After 3 days' infestation, under natural conditions, three enzyme activities in infested-plant and adjacent plant all significantly higher than undamaged control jujube, moreover, PI activity in nearby plant was reflexional change, PPO activity was "W"-type change, LOX activity was presented "N" shape change. Judging from the defense response duration, the high PI activity (compared with the control) in adjacent plant sustained 5 days, PPO and LOX lasted over 5 days, but in herbivored-plant, the duration of three kind of enzymes continued much longer and stronger.
     In summary, this study proved signal function and transmission pathway in inter-jujube in field conditions, explored defense response mechanism in neighbouring jujube exposed by HIPVs.
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