亚洲玉米螟对气候变暖的响应及其对温度胁迫的适应机制研究
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摘要
亚洲玉米螟Ostrinia furnacalis (Guenee)是我国玉米生产上最重要的害虫,其分布广泛、为害严重且生态适应性强。昆虫是变温动物,气候变暖必然对其产生深刻影响。为明确亚洲玉米螟在气候变暖条件下种群动态响应性变化及其适应机制,本文结合历史资料,通过田间调查研究了气候变暖条件下亚洲玉米螟及其天敌发生规律的变化；研究了气候变暖引发的频率和烈度不断增加的干旱和洪涝灾害对亚洲玉米螟种群动态的影响；以亚洲玉米螟对低温的耐受性和适应性为切入点,运用过冷却点(supercooling points, SCP)测定和低温暴露试验研究了亚洲玉米螟的耐寒策略、耐寒能力和地理种群间耐寒力变化；采用体内生化物质测定、mRNA差异显示(mRNA differential display, mRNA DD-PCR)和蛋白质组学方法,从生理生化、基因表达和蛋白调控层面上研究了亚洲玉米螟对低温胁迫的响应和调控机制；主要结果如下：
1.田间落卵系统调查结果显示,近10a沈阳地区亚洲玉米螟全年落卵时间延长,1代卵末期和田间终见卵时间均延后10d左右,1代卵期延长,1,2代落卵间隔期缩短；沈阳地区亚洲玉米螟发现3代卵,3代卵可发育到老熟幼虫并安全越冬,将导致沈阳地区由2代区变为2,3代混生区。
2.卵期和越冬期调查结果显示,沈阳地区亚洲玉米螟田间落卵量高,1代和2代分别达2992.67粒/百株和8746.33粒/百株,近10年2代卵量有显著下降趋势；赤眼蜂是卵期主要天敌,1代卵寄生率在2%～15%之间,随1代卵期延长有上升趋势,2代卵寄生率介于40%～70%之间,后期达90%以上；沈阳地区亚洲玉米螟越冬死亡率高,介于45%-65%之间,寄蝇、茧蜂和白僵菌三种主要天敌的控制力在22.50%-34.11%之间,连续四年未发现寄生性线虫。
3.对辽宁干旱和洪涝灾区亚洲玉米螟发生动态监测显示,受6-9月持续干旱影响,受灾当年亚洲玉米螟种群数量大幅下降(穗期虫量16.67头／百株,为害率34.67%),第二年亚洲玉米螟心叶期为害较轻(为害率16.67%),但穗期为害上升(虫量73.11头／百株,为害率47.78%),第三年种群数量恢复至较高水平(心叶期为害率37.33%,穗期虫量94头／百株,为害率80.44%)；在7月下旬洪涝灾害发生条件下,受灾年份亚洲玉米螟发生为害维持在较高水平(穗期虫量172头／百株,为害率98.33%),但不能直接影响第二年亚洲玉米螟的发生情况(心叶期为害率8.33%)。
4.对东北地区7个地理种群(佳木斯、德惠、公主岭、沈阳、瓦房店、普兰店和旅顺)亚洲玉米螟野生滞育幼虫耐寒性评价显示：试虫SCP介于-6.18～-29.98℃间,个体间差异大；试虫-30℃以上低温暴露12h无死亡,-30℃以下处理后死亡率随温度降低而升高,-80℃冷冻处理12h后仍有部分能够复苏,表明东北地区亚洲玉米螟耐寒能力极强,以耐结冰的耐寒性策略越冬,SCP不能评价其耐寒性强弱,低温存活率和LT50可以作为界定其耐寒性强弱的指标；7种群12h低温暴露死亡率随纬度升高而降低,差异显著(P<0.05),其致死中温(LT50)介于-52.19～-68.19℃间,LT50亦随纬度升高而降低,种群间差异显著(P<0.05)。
5.对亚洲玉米螟越冬幼虫耐寒能力和抗寒物质含量测定显示,虫体干物质中脂肪含量最高,12月下旬达62.51%,其次为蛋白质、甘露醇和糖类,血淋巴中甘油含量最高,2月达170.96mg/mL,其次为蛋白质、甘露醇和糖类；随时间延长,越冬期亚洲玉米螟含水量和蛋白质含量逐渐降低；随温度变化,越冬幼虫耐寒能力呈“倒U型”变化,糖类和甘露醇呈“U型”变化,脂肪和甘油呈“倒U型”变化；致死中温和死亡温度变化相关性极显著(P<0.01)；死亡温度变化与游离脂肪含量变动显著负相关(P<0.05),与血淋巴中甘油含量变化趋势负相关,但未达到显著相关水平(r=-0.802,P=0.055),致死中温变化与甘油含量变化致显著负相关(P<0.05)；脂肪和甘油为亚洲玉米螟抵御低温胁迫积累的主要物质,蛋白质、甘露醇和糖类在低温胁迫响应中提供能量和物质转换原料。
6.采用mRNA DD-PCR技术对亚洲玉米螟4℃和0℃低温胁迫下的基因表达谱进行分析显示,4℃和0℃胁迫下分别发现154和170条mRNA差异带,回收条带后二次扩增分别得到121和149个单一条带,经反向Northern杂交证实,这些片段均是组成型基因表达。认为亚洲玉米螟通过调节某些组成型基因的表达量或激活某些蛋白的生物学活性、调节某些物质含量、改变细胞膜的构象等方式来响应短暂低温胁迫。
7.对亚洲玉米螟幼虫血淋巴中耐低温胁迫相关蛋白质组进行分析,发现胁迫与否表达量稳定差异2倍以上蛋白点24个,将差异蛋白点做质谱分析后进行生物信息学比对,成功鉴定21个点,其中18个上调表达,8个下调表达；其中包括热激蛋白、肌动蛋白、精氨酸激酶等酶系、真核翻译起始因子、离子通道和免疫球蛋白等多个类群,这些蛋白在防御反应、能量和物质代谢、物质运输、基因和离子调控等方面发挥着重要作用。
Asian corn borer (ACB)(Ostrinia furnacalis(Guenee)), distributed widely due to its strong ecological adaptation, is the most important pests which cause enormous damage to maize production. Insect, as a poikilotherm, would been effected deeply by global climate warming.
Field survey and analytics of historical date, population fluctuation both ACB and its natural enemy and the effects of drought and flood to ACB were carried out to clarify the influence on population fluctuation of ACB under the global climate warming.
The cold-resistance strategy, freeze resistance ability and the diversity of different geographical populations of ACB by measuring the supercooling points (SCP) and by exposing to low-temperature were also studied. Response and adaptation mechanism of ACB to extreme temperature stress were conducted by biochemical, mRNA differential display (mRNA DD-PCR) and proteomics techniques. The results are as follows:
1. Systematic survey result showed that annual egg-laying period of ACB had been prolonged over the past decade in Shenyang region. The egg-laying end date of1st generation was prolonged about lOds. The egg stage period of1st generation ACB increased and the oviposition interval between1st and2nd generation decreased in Shenyang. The eggs of3rd generation ACB were founded and it could grew partially to mature larva and overwintered safely, It could be concluded that Shenyang region was transformed from the two-generation zone into the two and three mixed generations zone of ACB.
2. An amount of ACB oviposition was very high in field in Shenyang. The oviposition amount of1st and2nd generation was2992.67eggs/centgener plot and8746.33eggs/centgener plot respectively. The oviposition amount of2nd generation decreased significantly over the past decade. The major natural enemy of ACB during the egg-laying period was Trichogramma and parasitism rate was2%-15%in1st generation egg occur period. The parasitism rate increased with the prolonging of ACB1st generation egg-laying period. The parasitism rate was40%-70%in2nd generation egg-laying period, and reached90percentage above at the later period of2nd generation egg-laying period. The ACB overwintering mortality was high in Shenyang (45%-65%).
3. It was found that the ACB population decreased sharply due to lasting drought during June to September in2009in Beipiao county (Liaoning province) and ACB1st generation ACB population is lower next year, however2nd generation ACB population at that year restored to normal amount. No significant ACB population fluctuation was observed under the condition of flood disaster in Tieling city (Liaoning province) at the end of July in2010.
4. Freeze-resistance capability and tactics of wild diapause ACB larvae were compared among seven ACB geographical populations, which were collected from Jiamusi, Dehui, Gongzhuling, Shenyang, Wafangdian, Pulandian, and Lushun in Northeast China by measuring the supercooling point (SCPs) and by surveying the low temperature survival rates. The results showed that the range of SCPs was-6.18--29.98℃and had significant difference(P<0.05) among the seven geographic populations. No strict correlation between the SCPs and the geographic latitudes of population was observed. The mortality had significant difference(P<0.05) among the seven geographic populations and the mortality decreased with the increasing latitudes after12h cold exposed, the Median lethal temperatures (LT5o) decreased with the increasing latitudes and had significant difference(P<0.05)(-52.19～68.19℃). No death was observed the-30℃above after12h cold exposed. The mortality increased with the temperatures decreased below the-30℃. However, part of ACB individual could resistant-80℃. after12h frozen. So the cold-resistance standard of ACB was not SCP but survival rate of low temperature and LT50.
5. The freeze-resistance capability and relative body content of diapaused ACB overwintering larvae were studied during October2012to March2013. The results showed that the larvae freeze-resistance capability changed with temperature alteration. The water and protein contents within the ACB body descended gradually in winter. The fat which made up the highest percent of body dry matter in that time, reached62.51%at end of December. Glycerin, reached170.96mg/ml at February, took the highest hemolymph content. The correlation between Median lethal temperatures and the lethal temperatures alteration was significant (P<0.01). The alteration of lethal temperatures was significant negative correlated with the alteration of dissociative fat content (P<0.05), and was positive correlated (not significant, r=-0.802, P=0.055) with the alteration of glycerine content possessing in hemolymph. The alteration of lethal temperatures was significant correlated with alteration of glycerine content (P<0.05). In conclusion, the fat and the glycerin are the two main freeze-resistance substances of ACB. Moreover that plenty of mannitol accumulated was found in the hemolymph of diapause ACB larvae.
6. The gene expression of ACB at0℃and4℃under rapid cold acclimatization test was carried out by using the mRNA DD-PCR technology. The results showed that there were154and170difference zones through reverse transcription of total mRNA after test, and121and141single stripe were obtained through secondary amplification respectively. No specific gene transcription was found after rapid cooling domesticated of ACB through the reverse Northern blot. In conclusion, ACB could response to rapid cooling acclimatization by activating specific protein, adjusting substances content, changing the conformation of the membranes and regulating some specific gene expression.
7. The ACB haemolymph protein group of freezing and resistance to freezing were identified by two-dimensional electrophoresis. The results showed there were24proteins with two fold expression differences.21points were identified successfully after mass spectrometry analysis, including17up-regulation expressions and4down-regulation expressions. These protein play an important role in metabolism, energy supply and defense reaction of insects
1.田间落卵系统调查结果显示,近10a沈阳地区亚洲玉米螟全年落卵时间延长,1代卵末期和田间终见卵时间均延后10d左右,1代卵期延长,1,2代落卵间隔期缩短；沈阳地区亚洲玉米螟发现3代卵,3代卵可发育到老熟幼虫并安全越冬,将导致沈阳地区由2代区变为2,3代混生区。
2.卵期和越冬期调查结果显示,沈阳地区亚洲玉米螟田间落卵量高,1代和2代分别达2992.67粒/百株和8746.33粒/百株,近10年2代卵量有显著下降趋势；赤眼蜂是卵期主要天敌,1代卵寄生率在2%～15%之间,随1代卵期延长有上升趋势,2代卵寄生率介于40%～70%之间,后期达90%以上；沈阳地区亚洲玉米螟越冬死亡率高,介于45%-65%之间,寄蝇、茧蜂和白僵菌三种主要天敌的控制力在22.50%-34.11%之间,连续四年未发现寄生性线虫。
3.对辽宁干旱和洪涝灾区亚洲玉米螟发生动态监测显示,受6-9月持续干旱影响,受灾当年亚洲玉米螟种群数量大幅下降(穗期虫量16.67头／百株,为害率34.67%),第二年亚洲玉米螟心叶期为害较轻(为害率16.67%),但穗期为害上升(虫量73.11头／百株,为害率47.78%),第三年种群数量恢复至较高水平(心叶期为害率37.33%,穗期虫量94头／百株,为害率80.44%)；在7月下旬洪涝灾害发生条件下,受灾年份亚洲玉米螟发生为害维持在较高水平(穗期虫量172头／百株,为害率98.33%),但不能直接影响第二年亚洲玉米螟的发生情况(心叶期为害率8.33%)。
4.对东北地区7个地理种群(佳木斯、德惠、公主岭、沈阳、瓦房店、普兰店和旅顺)亚洲玉米螟野生滞育幼虫耐寒性评价显示：试虫SCP介于-6.18～-29.98℃间,个体间差异大；试虫-30℃以上低温暴露12h无死亡,-30℃以下处理后死亡率随温度降低而升高,-80℃冷冻处理12h后仍有部分能够复苏,表明东北地区亚洲玉米螟耐寒能力极强,以耐结冰的耐寒性策略越冬,SCP不能评价其耐寒性强弱,低温存活率和LT50可以作为界定其耐寒性强弱的指标；7种群12h低温暴露死亡率随纬度升高而降低,差异显著(P<0.05),其致死中温(LT50)介于-52.19～-68.19℃间,LT50亦随纬度升高而降低,种群间差异显著(P<0.05)。
5.对亚洲玉米螟越冬幼虫耐寒能力和抗寒物质含量测定显示,虫体干物质中脂肪含量最高,12月下旬达62.51%,其次为蛋白质、甘露醇和糖类,血淋巴中甘油含量最高,2月达170.96mg/mL,其次为蛋白质、甘露醇和糖类；随时间延长,越冬期亚洲玉米螟含水量和蛋白质含量逐渐降低；随温度变化,越冬幼虫耐寒能力呈“倒U型”变化,糖类和甘露醇呈“U型”变化,脂肪和甘油呈“倒U型”变化；致死中温和死亡温度变化相关性极显著(P<0.01)；死亡温度变化与游离脂肪含量变动显著负相关(P<0.05),与血淋巴中甘油含量变化趋势负相关,但未达到显著相关水平(r=-0.802,P=0.055),致死中温变化与甘油含量变化致显著负相关(P<0.05)；脂肪和甘油为亚洲玉米螟抵御低温胁迫积累的主要物质,蛋白质、甘露醇和糖类在低温胁迫响应中提供能量和物质转换原料。
6.采用mRNA DD-PCR技术对亚洲玉米螟4℃和0℃低温胁迫下的基因表达谱进行分析显示,4℃和0℃胁迫下分别发现154和170条mRNA差异带,回收条带后二次扩增分别得到121和149个单一条带,经反向Northern杂交证实,这些片段均是组成型基因表达。认为亚洲玉米螟通过调节某些组成型基因的表达量或激活某些蛋白的生物学活性、调节某些物质含量、改变细胞膜的构象等方式来响应短暂低温胁迫。
7.对亚洲玉米螟幼虫血淋巴中耐低温胁迫相关蛋白质组进行分析,发现胁迫与否表达量稳定差异2倍以上蛋白点24个,将差异蛋白点做质谱分析后进行生物信息学比对,成功鉴定21个点,其中18个上调表达,8个下调表达；其中包括热激蛋白、肌动蛋白、精氨酸激酶等酶系、真核翻译起始因子、离子通道和免疫球蛋白等多个类群,这些蛋白在防御反应、能量和物质代谢、物质运输、基因和离子调控等方面发挥着重要作用。
Asian corn borer (ACB)(Ostrinia furnacalis(Guenee)), distributed widely due to its strong ecological adaptation, is the most important pests which cause enormous damage to maize production. Insect, as a poikilotherm, would been effected deeply by global climate warming.
Field survey and analytics of historical date, population fluctuation both ACB and its natural enemy and the effects of drought and flood to ACB were carried out to clarify the influence on population fluctuation of ACB under the global climate warming.
The cold-resistance strategy, freeze resistance ability and the diversity of different geographical populations of ACB by measuring the supercooling points (SCP) and by exposing to low-temperature were also studied. Response and adaptation mechanism of ACB to extreme temperature stress were conducted by biochemical, mRNA differential display (mRNA DD-PCR) and proteomics techniques. The results are as follows:
1. Systematic survey result showed that annual egg-laying period of ACB had been prolonged over the past decade in Shenyang region. The egg-laying end date of1st generation was prolonged about lOds. The egg stage period of1st generation ACB increased and the oviposition interval between1st and2nd generation decreased in Shenyang. The eggs of3rd generation ACB were founded and it could grew partially to mature larva and overwintered safely, It could be concluded that Shenyang region was transformed from the two-generation zone into the two and three mixed generations zone of ACB.
2. An amount of ACB oviposition was very high in field in Shenyang. The oviposition amount of1st and2nd generation was2992.67eggs/centgener plot and8746.33eggs/centgener plot respectively. The oviposition amount of2nd generation decreased significantly over the past decade. The major natural enemy of ACB during the egg-laying period was Trichogramma and parasitism rate was2%-15%in1st generation egg occur period. The parasitism rate increased with the prolonging of ACB1st generation egg-laying period. The parasitism rate was40%-70%in2nd generation egg-laying period, and reached90percentage above at the later period of2nd generation egg-laying period. The ACB overwintering mortality was high in Shenyang (45%-65%).
3. It was found that the ACB population decreased sharply due to lasting drought during June to September in2009in Beipiao county (Liaoning province) and ACB1st generation ACB population is lower next year, however2nd generation ACB population at that year restored to normal amount. No significant ACB population fluctuation was observed under the condition of flood disaster in Tieling city (Liaoning province) at the end of July in2010.
4. Freeze-resistance capability and tactics of wild diapause ACB larvae were compared among seven ACB geographical populations, which were collected from Jiamusi, Dehui, Gongzhuling, Shenyang, Wafangdian, Pulandian, and Lushun in Northeast China by measuring the supercooling point (SCPs) and by surveying the low temperature survival rates. The results showed that the range of SCPs was-6.18--29.98℃and had significant difference(P<0.05) among the seven geographic populations. No strict correlation between the SCPs and the geographic latitudes of population was observed. The mortality had significant difference(P<0.05) among the seven geographic populations and the mortality decreased with the increasing latitudes after12h cold exposed, the Median lethal temperatures (LT5o) decreased with the increasing latitudes and had significant difference(P<0.05)(-52.19～68.19℃). No death was observed the-30℃above after12h cold exposed. The mortality increased with the temperatures decreased below the-30℃. However, part of ACB individual could resistant-80℃. after12h frozen. So the cold-resistance standard of ACB was not SCP but survival rate of low temperature and LT50.
5. The freeze-resistance capability and relative body content of diapaused ACB overwintering larvae were studied during October2012to March2013. The results showed that the larvae freeze-resistance capability changed with temperature alteration. The water and protein contents within the ACB body descended gradually in winter. The fat which made up the highest percent of body dry matter in that time, reached62.51%at end of December. Glycerin, reached170.96mg/ml at February, took the highest hemolymph content. The correlation between Median lethal temperatures and the lethal temperatures alteration was significant (P<0.01). The alteration of lethal temperatures was significant negative correlated with the alteration of dissociative fat content (P<0.05), and was positive correlated (not significant, r=-0.802, P=0.055) with the alteration of glycerine content possessing in hemolymph. The alteration of lethal temperatures was significant correlated with alteration of glycerine content (P<0.05). In conclusion, the fat and the glycerin are the two main freeze-resistance substances of ACB. Moreover that plenty of mannitol accumulated was found in the hemolymph of diapause ACB larvae.
6. The gene expression of ACB at0℃and4℃under rapid cold acclimatization test was carried out by using the mRNA DD-PCR technology. The results showed that there were154and170difference zones through reverse transcription of total mRNA after test, and121and141single stripe were obtained through secondary amplification respectively. No specific gene transcription was found after rapid cooling domesticated of ACB through the reverse Northern blot. In conclusion, ACB could response to rapid cooling acclimatization by activating specific protein, adjusting substances content, changing the conformation of the membranes and regulating some specific gene expression.
7. The ACB haemolymph protein group of freezing and resistance to freezing were identified by two-dimensional electrophoresis. The results showed there were24proteins with two fold expression differences.21points were identified successfully after mass spectrometry analysis, including17up-regulation expressions and4down-regulation expressions. These protein play an important role in metabolism, energy supply and defense reaction of insects
引文
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