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缨翅目 (Thysanoptera) 和膜翅目 (Hymenoptera)昆虫的繁殖采用单倍−二倍体性别决定系统(Haplodiploid sex-determinationsystem)。雌虫行两性生殖时,受精卵发育成二倍体雌虫,未受精卵发育成单倍体雄虫;而产雄孤雌生殖(Arrhenotoky)的后代全为单倍体雄虫[1-3]。它们的性别分配可用多种理论模型来表征[4],其中,常用的模型有局域配偶竞争(Local mate competition)理论[5]和获精受限雌虫模式(Constrained females model)[6]。这2种模型的背后隐含着一个重要的假设前提:雌虫能感知和评价周边环境的信息,然后对子代性比进行调节[4]。普通大蓟马(Megalurothrips usitatus) 是一种植食性蓟马,主要为害豆科植物的嫩芽、花朵和嫩荚等器官[7-11]。在我国海南,该虫已成为豇豆(Vigna unguiculata)的头号害虫[12]。跟很多植食性蓟马一样,普通大蓟马采用两性生殖和产雄孤雌生殖方式来繁衍后代[7,9,12]。蓟马的性比受很多因素的影响,包括蓟马的生物学和生态学特性、温度、食物和防控因子等[3,13-15]。雌虫通过控制受精囊内精子的输入和释放来调控子代性比[16]。蓟马性比的调节可分为2个部分,即交配期间的性比调节和交配后的性比调节[12]。迄今,有关同种成虫对蓟马交配期间性比调节的影响尚未见报道。本研究将评价同种成虫的气味对普通大蓟马雌虫交配和子代性比的干扰作用,旨在明确该虫在交配期间能否利用气味来判别周边同种成虫的性别和交配状况,然后调控雌虫获得精子的能力,进而对子代性比进行调节。
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用不同气味源处理配对的普通大蓟马24 h后,被测雌虫所产子代的雄虫数、雌虫数和成虫数都有不同程度的变化(表1)。
表 1 同种成虫的气味处理配对期间的普通大蓟马对子代成虫数量的影响
气味源 子代成虫数/头 雄虫 雌虫 成虫 对照 16.55±2.36ab 38.09±3.09a 54.64±3.29ab 已交配雌虫 21.04±1.52a 22.80±1.67b 43.84±2.48b 未交配雌虫 13.32±1.63b 46.95±4.97a 60.26±6.49a 已交配雄虫 14.39±1.71b 45.35±4.56a 59.74±5.39a 注:表中数值为平均数±标准误,同一列数据后标有不同字母表示在P<0.05上差异显著。 子代雄虫数:对照、已交配雌虫气味处理、未交配雌虫气味处理和已交配雄虫气味处理的子代雄虫数分别为16.55、21.04、13.32和14.39头,其中,已交配雌虫气味处理与未交配雌虫气味处理和已交配雄虫气味处理之间的差异达到了显著水平(P<0.05)。
子代雌虫数:已交配雌虫气味处理的子代雌虫数最少,仅为22.80头,跟其他处理均有显著差异(P<0.05)。另3个处理的子代雌虫数介于38.09~46.95头,但彼此之间无显著差异。
子代成虫数:子代成虫数最少的处理仍是已交配雌虫气味处理(43.84头),与未交配雌虫气味处理和已交配雄虫气味处理之间的差异均达到了显著水平(P<0.05),但与对照(54.64头)相比差异不显著。
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子代性比:普通大蓟马子代性比的响应随成虫气味源而异(图1)。已交配雌虫气味处理的子代性比高达0.48,同其他3个处理之间的差异均达到显著水平(P<0.05)。未交配雌虫气味处理的子代性比最低,仅为0.22,跟对照(0.31)差异显著(P<0.05)。已交配雄虫气味处理的子代性比(0.26)比对照的子代性比略低,两者之间没有显著差异。
子代性比日变化:在不同气味处理条件下,普通大蓟马子代的日性比均呈现波动性(图2),其中,已交配雌虫气味处理的子代性比日变化别具特色。从图2可知,在雌虫产卵观测期的第1~13天,已交配雌虫气味处理的日性比介于0.37~0.67,多数接近0.50,但均大于对照;随后日性比变化趋势出现反转,从产卵第14天至第20天,对照的日性比大于已交配雌虫气味处理;至产卵第21天时,已交配雌虫气味处理的日性比达到1.00,即所产子代全为雄虫,意味着雌虫受精囊内的精子已消耗殆尽。对照、未交配雌虫气味处理和已交配雄虫气味处理的日性比波动范围分别为0.15~0.7、0.00~0.33和0.10~0.44。在产卵第1~11天,未交配雌虫气味处理和已交配雄虫气味处理的日性比波动类似于对照。但是,从在产卵第12天起,这3个处理的日性比波动呈现出不一样的态势。
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各种气味处理条件下,被测雌虫的产卵天数平均值介于11.52~12.96 d,彼此之间没有显著差异,表明同种雌虫和已交配雄虫的气味对雌虫产卵期都没有明显影响。
Offspring sex ratio response to odors from conspecific adults in Megalurothrips usitatus (Bagrall)(Thysanoptera:Thripidae)
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摘要: 为了探究同种成虫的气味对普通大蓟马(Megalurothrips usitatus)交配期间性比调节的影响,设置对照(无气味)、已交配雌虫、未交配雌虫和已交配雄虫等4种气味源,分别处理单对刚羽化成虫24 h,然后用幼嫩豇豆(Vigna unguiculata)豆荚单头饲养被测雌虫,观测每日所产子代的性别和数量,计算子代性比。结果表明:已交配雌虫气味处理的子代雌虫数(22.80头)最少,跟其他处理差异显著。同对照的子代性比(0.31)相比,已交配雌虫和未交配雌虫的气味分别导致子代性比显著提高和降低,子代性比分别为0.48和0.22,而已交配雄虫气味对子代性比无显著影响。4个处理中,仅已交配雌虫气味处理的雌虫在产卵最后1天的日性比达到1.00,即雌虫体内的精子已消耗殆尽。成虫气味源对雌虫产卵天数没有明显影响。实验结果说明,普通大蓟马在交配期间能依靠嗅觉判别周边雌虫的交配状况,然后通过调控雌虫的获精数量,实现对子代性比的自调节。Abstract: To explore the effects of odors from conspecific adults on sex ratio adjustment during mating by bean flower thrips, Megalurothrips usitatus, one of main pests on Vigna unguiculata . In this paper, a pair of newly emerged male and female of the been flower thrips was exposed for 24 hours to the odors from 6 mated females, 6 virgin females, 6 mated males and no adult as control, respectively. Then the focal female was transferred to a glass tube containing a section of a young cowpea pod for her feeding and oviposition. The pod was replaced daily until the focal female died. The pod containing eggs was transferred to a new glass tube for the offsprings’ growth. Oviposition days of the focal females were recorded, and their adult offspring were sexed and counted. Sex ratios of offspring were calculated as proportion of males. The results showed that the number of female offspring per focal female for the treatment with the odor from mated females was 22.80, being significantly less than those for other treatments. The sex ratios of adult offspring produced by the thrips treated with the odors from mated and virgin females were 0.48 and 0.22, respectively, which both were significantly different from that of control (0.31). The odor from mated males, however, showed no significant effect on offspring sex ratio. The daily sex ratio at the last oviposition day for the focal females treated with the odor from mated females reached 1.00, suggesting that their sperms had been depleted. The oviposition days of the focal females did not differ among all treatments. Our research revealed that the bean flower thrips during mating can use olfactory cues to assess mating status of conspecific females, and then control the number of sperms transferred into spermathecae, thereby adjusting sex ratio of their offspring.
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Key words:
- Megalurothrips usitatus /
- cowpea pods /
- arrhenotoky /
- sex allocation /
- olfactory cues
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表 1 同种成虫的气味处理配对期间的普通大蓟马对子代成虫数量的影响
气味源 子代成虫数/头 雄虫 雌虫 成虫 对照 16.55±2.36ab 38.09±3.09a 54.64±3.29ab 已交配雌虫 21.04±1.52a 22.80±1.67b 43.84±2.48b 未交配雌虫 13.32±1.63b 46.95±4.97a 60.26±6.49a 已交配雄虫 14.39±1.71b 45.35±4.56a 59.74±5.39a 注:表中数值为平均数±标准误,同一列数据后标有不同字母表示在P<0.05上差异显著。 -
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