仲夏棉蚜种群崩溃机制研究进展
2015-12-09高桂珍吕昭智
高桂珍,吕昭智
(1.中国科学院新疆生态与地理研究所,生物地理与生物资源重点实验室,乌鲁木齐830011 2.新疆农业大学林学与园艺学院,乌鲁木齐 830052)
棉蚜Aphis gossypii Glover 是世界性分布的棉花害虫,也是我国长江、黄河和西北棉区的重要害虫。棉蚜主要以刺吸叶片汁液和分泌蜜露的方式影响棉花的生长和品质,由于棉蚜危害时间长、危害重、繁殖加强、种群蔓延速度快等特点(马祁,2002),给棉花生产带来了巨大的损失。每年的7月中旬左右棉蚜种群数量迅速下降,这为棉蚜的预测和防治工作提供了切入点,生产中若能预知棉蚜种群崩溃现象的发生,可有效减少不必要的农药使用,有效降低成本投入和环境损害,但何种因素导致棉蚜种群崩溃迄今尚不清楚。本文对可能导致仲夏棉蚜种群崩溃的主要影响因素:植物次生代谢物、暴风雨、高温、种内竞争和天敌分别进行论述,探讨其对棉蚜存活、繁殖和有翅蚜形成的影响,旨在为探明仲夏棉蚜种群崩溃机制提供参考。
1 棉蚜种群过程及其崩溃
棉蚜种群经历快速增长到达高峰期,然后在5-10 d 内棉蚜种群从高峰值迅速消亡,这在不同国家不同地区均有发生,是普遍存在的现象(见表1)。由表1 可见,不同寄主、不同地区的棉蚜种群崩溃时间不同,这是由于不同寄主植物的物候、营养以及不同地区的气候不同所致。新疆属于西北内陆棉区,播种时间较黄河流域、长江流域棉区早,且光照热量充足,棉蚜始发期和高峰期提前,因此新疆棉区较中国其他地区棉蚜种群崩溃时间早。
表1 不同寄主不同地区棉蚜种群崩溃参数Tab.1 The parameters of population crash of Aphis gossypii on different host plants and different areas
2 棉蚜种群崩溃影响因素分析
蚜虫种群崩溃的研究主要涉及生物因素(植物营养恶化、天敌数量增加以及蚜虫种内竞争)和非生物因素(剧烈天气变化),这些因素引起蚜虫繁殖率降低、死亡率升高和有翅蚜迁飞,从而导致蚜虫种群崩溃 (Jones,1979;Ba-Angood and Stewart,1980;Karley et al.,2004)。桃 蚜、麦蚜等多种蚜虫均存在种群崩溃的现象。植物营养恶化和天敌增加导致桃蚜种群繁殖率降低、死亡率增加,使其种群崩溃 (Kift et al.,1998;Karley et al.,2003);高温引起麦双尾蚜死亡和迁移从而导致其种群崩溃(Basky,1993);小麦营养恶化降低麦长管蚜繁殖率(Watt,1979)、促进有翅蚜形成并迁移(Watt and Dixon,1981)导致其种群崩溃。由此可见,为害不同作物的不同蚜虫种类,其种群崩溃的影响因素和种群崩溃过程均存在差异,而关于棉蚜种群崩溃的关键影响因素及其作用机制未见系统报道。
2.1 植物次生代谢物
害虫的取食为害能引起植物的生理应激反应,大量产生并累积对昆虫有防御效应的次生性抗虫物质。植物次生代谢物在一定程度上影响蚜虫的行为和数量(Douglas,2003),植物组织中可溶性糖和氨基酸的种类和含量明显影响植物对昆虫的抗性或防御能力(Slosser et al.,2004)。单宁是广谱的防御物质,兼有拒虫和抗生的双重性质。刘旭明研究表明,蚜虫为害棉花后诱导棉株单宁的加快合成并积累 (刘旭明,1989);刘泽辉等(2008)研究表明棉酚含量与抗虫性呈正相关;陆宴辉等(2005)研究表明单宁和棉酚对棉蚜种群增长有抑制作用。棉花在棉蚜为害胁迫条件下,体内棉酚、单宁、脯氨酸和可溶性糖含量以及抗蚜性相关酶类如苯丙氨酸解氨酶、多酚氧化酶、过氧化氢酶和超氧化物歧化酶活性都有不同程度的上升(刘旭明和杨奇华,1991,1993;董应才等,1996;李进步等,2008);棉蚜为害后的棉株使得棉蚜种群数量下降,增长率降低,发育历期延长(董应才等,1996)。
但棉花体内单宁、棉酚等次生代谢物含量很低(平均0.2%左右),其对棉蚜的抑制作用不足以导致棉蚜种群数量的迅速下降 (陆宴辉等,2005),且7月中旬是棉花生长的旺盛期,营养丰富充足。所以植物次生代谢物在一定程度上抑制棉蚜的种群数量,但其不是仲夏棉蚜种群崩溃的关键影响因子。
2.2 暴风雨
剧烈的天气变化尤其是强烈的暴雨和强风,会导致蚜虫的快速大量死亡(Walker et al.,1984;Legg and Brewer,1995;Jas'kiewicz,1997),但是这些异常的事件无法解释连续多年和多个地点发生的种群崩溃现象(Parker et al.,2000)。新疆棉区几乎每年的7月中旬均发生棉蚜种群崩溃现象,并连续发生于南北疆多个地点,且新疆干燥少雨,年平均降水量仅147 mm,南疆地区甚至不到60 mm (郑鸿雁等,1999),因此暴风雨不能成为仲夏棉蚜种群崩溃的主要原因。
2.3 温度
温度是影响昆虫生长、发育、繁殖及存活等生命活动最重要的因素(Hoffmann,1985)。棉蚜在高温条件下比培养在适宜条件下发育速率慢(Stinner et al.,1974;Curry et al.,1978;Kersting et al.,1999)、存活率低(Ma et al.,2004)。在30℃时棉蚜繁殖力普遍明显下降 (赵静雅等,2001)。在26℃-30℃范围内,棉蚜第二代产仔量明显低于第一代(吴孔明和刘芹轩,1992)。在35℃时,棉蚜不能够完成生命循环,也不能产仔(Zamani et al.,2006),35℃的持续高温对若蚜是致命的 (Aldyhim and Khalil,1993;Satar et al.,2005)。但中国南部蚜虫在35℃下能够生存并且产仔(Xia et al.,1999b),即不同地理区域的棉蚜种群之间耐高温能力存在一定差异 (刘健等,2003;Parajulee,2007)。同时值得注意的是,恒温试验环境与田间正午的高温相差甚远,潘飞等(2014)提出了开展日变温模式下昆虫生物学特性研究的重要性,且研究发现变温与恒温对麦蚜(Ma et al.,2004a,b)、桃 蚜 (Davis et al.,2006)、棉蚜(李晶晶等,2011)等存活和繁殖的影响存在差异,高桂珍等(2012)研究表明波动性高温胁迫显著降低棉蚜的存活和繁殖,38℃是棉蚜耐高温能力的拐点。而仲夏季节正午的温度往往到达35℃甚至40℃以上,因此极端高温很可能是仲夏棉蚜种群崩溃的关键影响因素。
温度高低对有翅蚜形成的影响存在争论,许多试验表明,较低温度适合有翅蚜产生,较高温度抑制其产生 (Johnson,1966;Schaefers and Judge,1971;王佩玲,2014),但 Muller(2001)认为高温增加了蚜虫个体的活性,相对增加了种群拥挤度,促使有翅蚜形成。温度对有翅蚜形成的影响有待进一步研究,从而明确促使和抑制有翅蚜形成的温度范围。
2.4 种内竞争
由于蚜虫主要营孤雌繁殖并且世代重叠,蚜虫种群很可能是密度制约调节过程(Sequeira and Dixon,1997)。目前国内外已报道了在许多蚜虫种类上存在着密度制约效应(Maudsley et al.,1996;Alyokhin et al.,2005;Rhainds and Messing,2005;Bommarco et al.,2007)。Mashanova 等(2008)报道密度制约分布也许可以解释仲夏蚜虫种群崩溃的现象,并认为母代效应是仲夏蚜虫种群崩溃的关键因子。蚜虫的种内竞争可引起其死亡率升高、繁殖力降低和扩散加快 (Dixon,1971;Day,1986;Kindlmann and Dixon,1996)。种内竞争对有翅蚜的形成具有重要作用,密度越大,蚜虫间接触越多,有翅蚜比例越高 (Lees,1961;Johnson,1965)。孟玲和李保平(2000)研究发现棉蚜有翅蚜的数量随棉蚜种群密度的增大而增加。有翅蚜迁飞在一定程度上加速了棉蚜种群的崩溃(Hughes and Gilbert,1968)。高桂珍等 (2009)研究结果表明,随着种群密度增加,棉蚜种群增长率降低;初始密度越大,棉蚜种群崩溃越提前。同时,种内密度效应受到温度影响,温度较高时蚜虫种内密度效应显著(Agrawal et al.,2004)。高桂珍等(2013)研究表明在不同的温度区间和密度范围内,温度和密度对棉蚜的作用大小、主次发生了改变。因此,种内竞争显著影响棉蚜存活、繁殖和有翅蚜比例,且暴发年份高峰期单叶蚜量甚至可以达千头以上,激烈的种内竞争是否启动了棉蚜种群的崩溃需要进一步验证。同时,温度和密度效应对棉蚜种群的联合作用需要更多关注。
2.5 天敌
天敌是生态控制蚜虫种群数量的重要因素(Steenis,1992;Zhang,1992;Rutledge et al.,2004;吕昭智等,2005;冯宏祖等,2007)。高孝华等(2001)和朱三荣等(2006)研究了龟纹瓢虫对棉蚜的捕食功能作用和搜索行为,结果表明其捕食量随猎物的密度增加而上升,当猎物增加到一定水平,捕食量趋向稳定。丁瑞丰等(2015)研究表明人工释放草蛉对棉蚜具有一定的防治效果。然而,种内干扰、空间大小和异质性对捕食过程产生了重要影响,天敌在田间捕食量远不及室内较小空间的捕食量(Butler and May,1971;苏建伟等,2000)。天敌对棉蚜的作用同时受到温度的影响(Romo and Tylianakis,2013)。夏敬源等(1999a)研究表明七星瓢虫对棉蚜的搜索率随温度增加直线上升,而处置率则呈曲线上升;Yu等(2013)研究表明小十三星瓢虫对棉蚜的捕食率随着温度的升高呈先上升后下降的趋势。
天敌对蚜虫的影响除了直接的捕食或寄生外,还具有间接的非致死作用(Andrew et al.,2012;李姣等,2014),从而影响蚜虫的发育、繁殖(李姣等,2013)和有翅蚜形成 (Swapna et al.,2014)。Byers (2005)研究表明天敌的存在影响若蚜的生长发育;Mondor 等(2005)研究表明增加捕食风险,棉蚜将产生更多有翅蚜后代。因此,评估天敌作用时,除了研究天敌对害虫的直接捕食或寄生的效率,还应考虑到天敌存在对害虫的非直接致死效应。
天敌对棉蚜具有控制作用,棉蚜发生早期,天敌可以控制棉蚜,但棉蚜数量快速增长及高峰期阶段,天敌不足以控制棉蚜 (Slosser et al.,1998)。这同时在马铃薯蚜虫 (Walker et al.,1984)、麦蚜(Kindlmann et al.,2010)上均有报道。棉蚜和瓢虫的种群数量具有明显的跟随效应(朱世华等,2000;王林霞等,2004),棉蚜种群下降期,瓢虫种群数量刚好达到高峰期,所以天敌很可能在棉蚜种群崩溃后期发挥重要作用。
3 总结与展望
仲夏棉蚜种群崩溃是自然控制棉蚜的重要途径,高温、种内竞争以及天敌均对棉蚜存活、繁殖和有翅蚜形成产生了重要的影响。温度在棉蚜种群崩溃甚至整个种群过程中发挥了重要作用;初始密度影响棉蚜种群崩溃发生的早晚,即初始密度越大,崩溃发生越提前;天敌对加速棉蚜种群后期崩溃发挥了重要作用。但仍存在以下问题尚待明确:极端高温以及高温出现的频率对棉蚜如何影响?其对种群崩溃发生的早晚及下降速率如何影响?温度对有翅蚜的形成存在争论,温度胁迫刺激棉蚜产生有翅蚜的阈值范围有待明确。同时,各因素之间对棉蚜的影响不是独立存在的,温度、种内竞争以及天敌对棉蚜的联合作用是未来研究的重点。
数学模型是研究种群动态的有力手段,它在蚜虫自然种群研究中具有巨大的潜力。由于影响因素众多,对棉蚜整个种群过程的预测其准确性、有效性和推广性较差。种群崩溃现象为棉蚜预测和防治提供了新思路与切入点,目标性强,重要影响因素范围缩小,提高了预测可行性。Mashanova 等(2008)首次用模型解决蚜虫种群崩溃问题,通过假设模型得出密度制约可以解释仲夏蚜虫种群崩溃现象,但缺乏有力的室内实验数据和田间实际种群动态的支持。在明确以上因素对棉蚜影响的基础上,需进一步结合田间实际棉蚜、天敌及气象监测数据,建立仲夏棉蚜种群崩溃数学模型,从而有效预测仲夏棉蚜种群崩溃发生时间,减少农药盲目使用,有效降低成本投入和环境损害,对开发防治新策略具有重要意义。
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