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母代效应(Maternal effects)在自然生态系统中是普遍存在的[1-2]。亲代的取食和生活经历等因素可影响生物母体对资源的分配从而影响其后代的发育、表型分化、繁殖等生活史特征[3-5],这种由于母体介导所带来的表观遗传信息跨代传递往往提高种群适合度[6-7]。昆虫天敌亦会受到母代效应的影响[4, 8-10],林间寄生性天敌昆虫母代在成虫寿命期内完成首次寄生后可再次搜索寄生从而产生多个姊妹窝子代[11-14]。这些母代调控下的各个姊妹窝子代发育策略及其在林间的活动将影响其种群的定殖和扩散,有关寄生蜂不同姊妹窝子代生长发育情况以及表型如何分化还未见报道。
白蜡吉丁肿腿蜂(Sclerodermus pupariae Yang et Yao)是近年来在白蜡窄吉丁(Agrilus planipennis)幼虫和蛹上发现的一种卵育型抑性外寄生蜂(synovigenic idiobiont ectoprasitoid)[15],该蜂对寄主有较强的搜索和主动攻击能力,现已成为我国天牛类和吉丁甲类幼虫的优势天敌[16-17]。对白蜡吉丁肿腿蜂生态学和生物学特性的研究显示,该蜂主要营两性生殖,雄蜂先羽化,咬破雌蜂茧壳后钻入交尾。未经交配的雌蜂也能孤雌生殖,但后代全部为雄性。雌雄蜂均可多次交配,仅交配一次的雌蜂补充营养后仍继续产出雌性后代[18];多寄生,1头寄主幼虫常常繁育出数十头,有时高达100多头子代蜂[19-20]。此外,武辉等[18]在室内变温条件下观察到该蜂每代能产出3~5个姊妹窝子代,但是母代产生的各姊妹窝子代发育历期和表型分化等生物学参数尚无详细记载。因此,本研究以白蜡吉丁肿腿蜂为对象,在室内恒温条件下母蜂连续依次寄生多头寄主幼虫,观察其在寿命期内所产生的姊妹窝个数,并比较随着姊妹窝子代产出次数的增加,母蜂对寄主幼虫的寄生能力以及各姊妹窝子代的发育历期、翅型分化和性比情况,研究母蜂连续寄生所产各个姊妹窝子代的生物学参数变化,为阐明该蜂在林间定殖和种群延续机制提供理论依据。
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白蜡吉丁肿腿蜂建群蜂采集于天津市大港区官港森林公园(38°56′N,117°29′E)内绒毛白蜡(Fraxinus velutina)林受害树上的白蜡窄吉丁蛀道内。寄主麻竖毛天牛(Thyestilla gebleri)幼虫购自市售。野外肿腿蜂种群用麻竖毛天牛幼虫作为替代寄主于室内人工气候箱中(宁波海曙赛福PRX-450C)进行传代饲养建立实验种群,饲养条件为30±1℃,光周期14L:10D,相对湿度为65%±5%。室内种群F32中挑选羽化10 d且体型中等的无翅型母蜂为供试肿腿蜂,选取质量230.0±10.0 mg麻竖毛天牛幼虫作为供试寄主。
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待白蜡吉丁肿腿蜂雌雄蜂交配后,从羽化10 d的寄生蜂种群中随机选取50头,无需补充营养,直接用毛笔单头挑出并置入有1头麻竖毛天牛的指形管(1.0 cm×5.0 cm)内,用脱脂棉塞紧管口,所有供试昆虫均置于30±1℃,14L︰10D,65%±5%RH的人工气候箱培育。当母蜂首次产下的子代开始吐丝结茧时将母蜂移出,不经过交配直接将母蜂单头接入有1头麻竖毛天牛幼虫的指形管内让其寄生。依次类推,在母蜂整个寿命期内观察母蜂姊妹窝子代的产出情况,直到所有母蜂死亡。所有观察均在Zeiss体视显微镜(Zeiss, Stemi 2000-c)下完成。
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接蜂后,每天定时观察并记录白蜡吉丁肿腿蜂在连续产出姊妹窝子代后在麻竖毛天牛幼虫体上的产卵和羽化情况。以母蜂在寄主幼虫体上产出卵粒记为寄生,以母蜂子代在寄主体上成功羽化出成蜂记为寄生成功。统计母蜂对寄主幼虫的寄生率和寄生成功率,以及各姊妹窝子代中的雌蜂数量。寄生率(bi)和寄生成功率(ri)是衡量寄主适合度的重要指标,计算方法参考田军等[21]。
$ {{b}_{i~}}=\text{ }{{R}_{i}}/{{S}_{i}}\text{, }{{r}_{i}}~={{A}_{i}}/{{S}_{i}} $
式中Ri为被寄生(以麻竖毛天牛幼虫被白蜡吉丁肿腿蜂蛰刺、麻痹、停止发育并在寄主体上产卵为标志)的寄主数量,Ai为白蜡吉丁肿腿蜂在麻竖毛天牛幼虫上寄生并完成生长发育、羽化为成蜂的寄主数目,Si为供试的麻竖毛天牛幼虫总数。
b∈[0, 1], r∈[0, 1], b, r越接近1,则说明寄主越适合被白蜡吉丁肿腿蜂寄生,同时也反映出该蜂对麻竖毛天牛幼虫寄生作用。白蜡吉丁肿腿蜂雌蜂作为卵育型寄生蜂,在首次产生子代后,如果能搜索到新的寄主幼虫,会再次取食寄主补充营养从而繁育出新的姊妹窝子代。母蜂连续寄生所产的后代均为姊妹窝子代,分别记为第1个姊妹窝子代(sister broodⅠ),第2个姊妹窝子代(sister broodⅡ),第3个姊妹窝子代(sister brood Ⅲ),…,依此类推。
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记录母蜂接入寄主幼虫的时间(a′)、第1粒卵粒的产出时间(a)、卵粒开始孵化时间(b)、寄生蜂幼虫吐丝时间(c)、第1头雄蜂羽化时间(d)以及第1头雌蜂羽化时间(e),将a-a′、b-a、c-b、d-c、e-c、d-a、e-a分别记作产卵前期、卵期、幼虫期、雄蜂茧蛹期、雌蜂茧蛹期、雄蜂发育历期、雌蜂发育历期;统计各姊妹窝子代雌蜂有翅率、雄蜂数量和性比。以有翅雌蜂个体占子代雌蜂总量的百分比作为雌蜂有翅率,以雄性个体占子代数量的百分比作为寄生蜂性比。
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采用单因素方差分析比较产出不同姊妹窝次数的母蜂的产卵前期、不同姊妹窝子代卵期、幼虫期、茧蛹期、雄蜂发育历期和雌蜂发育历期的差异,用LSD法检验各参数均值之间的差异显著性。采用卡方检验评价不同姊妹窝子代有翅率、性比的差异,组内比较的显著水平经过布氏矫正后为0.008 3。所有数据使用SPSS 20.0软件分析完成。
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研究发现,白蜡吉丁肿腿蜂在麻竖毛天牛寄主幼虫上可以产生4个姊妹窝子代,产出第2个姊妹窝后母蜂存活个体数开始降低。第3个姊妹窝产出少量雌蜂,产生第4个姊妹窝后的母蜂继续接入寄主幼虫则不能成功寄生。母蜂连续3次的寄生行为均对寄主幼虫有较高的寄生率和寄生成功率(>75%),随着产出姊妹窝次数的增加,母蜂对寄主幼虫的寄生能力和子代雌蜂数量逐渐降低(表 1)。
表 1 连续产出姊妹窝子代的白蜡吉丁肿腿蜂寄生能力和子代雌蜂数量
Table 1. Parasitic ability and number of female progeny of S. pupariae produced successive sister broods
母蜂所产姊妹窝次数
Number of sister brood produced by female重复数
Replicates寄生率
Parasitism rate/%寄生成功
Successful parasitism/%平均雌蜂产出量
Average female progeny1 50 98.0 a 92.0 a 70.54±16.03A 2 50 82.0 a 76.0 a 52.65±14.07A 3 37 83.8 a 81.1 a 21.97±17.31B 4 20 10.0 b 10.0 b 4.00±2.83B 5 1 0 0 0 表中数据为平均值±标准差。同列数据后不同的小写字母表示在0.008 3水平上差异显著(卡方检验);同列数据后不同大写字母表示在0.05水平上差异显著(单因素方差分析,LSD检验)。Data in the table are presented as mean ± SD. Those in the same column followed by different small letters are significantly different at the 0.008 3 level, by Chi-square test. Those in the same column followed by different capital letters are significantly different at the 0.05 level, by one-way ANOVA, LSD test. -
随着白蜡吉丁肿腿蜂产出姊妹窝次数的增多,母蜂产卵前期逐渐延长,当寄生蜂已经繁育出3个姊妹窝子代后,再次寄生新的寄主幼虫时产卵前期显著延长(df = 3, 112, F = 3.816, P = 0.012)。各个虫态的发育历期虽然在各个姊妹窝之间存在差异,但其历期并无逐代延长或者缩短的规律(表 2)。
表 2 连续产出姊妹窝子代的白蜡吉丁肿腿蜂产卵前期和子代发育历期
Table 2. Pre-oviposition duration and progeny developmental duration of S. pupariae produced successive sister brood
母蜂所产姊妹窝次数
Number of sister broodproduced by female产卵前期
Pre-ovipositionperiod卵期
Egg duration幼虫期
Larval duration雄蜂茧蛹期
Male pupalduration雌蜂茧蛹期
Female pupalduration雄蜂发育历期
Developmental duration of male雌蜂发育历期
Developmental duration of female1 5.37±0.57 b 2.65±0.85c 6.89±0.71a 12.53±0.72 a 12.74±0.68a 19.41±1.05 ab 19.63±1.14 b 2 5.45±0.67 b 3.63±0.54a 7.34±2.64a 11.58±1.08 b 11.89±1.01b 18.92±1.94 b 19.24±1.92 b 3 5.60±1.40 b 3.03±0.81b 5.73±1.17b 11.27±1.02 b 11.77±0.77 b 20.03±1.59 a 20.53±1.43 a 4 7.50±0.70 a 3.00±0.00abc 6.00±0.00ab 11.00±0.00 b 11.00±0.00b 20.00±0.00 ab 20.00±0.00ab 表中数据为平均值±标准差,同列数据后不同小写字母表示在0.05水平上差异显著(单因素方差分析,LSD检验)。Data in the table are presented as means followed ± SD, those in the same column, by different small letters are significantly different at the 0.05 level, by one-way ANOVA, LSD test. -
白蜡吉丁肿腿蜂产出的4个姊妹窝子代之间雌蜂有翅率差异显著(df = 3, χ2 = 76.104, P<0.001),在最先产出的两个姊妹窝子代中,寄生蜂种群均保持着一定比例的有翅型和无翅型个体,随着姊妹窝数量的增加,雌蜂有翅率递减;第1个和第2个姊妹窝子代的有翅率在40%左右,第3个姊妹窝子代开始降至23.3%,第4个姊妹窝子代所产少量子代雌蜂均为无翅型(图 1)。
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白蜡吉丁肿腿蜂各姊妹窝子代雄蜂产出量有显著差异(df=3, 112, F=12.2, P<0.001)(图 2),随着母蜂所产姊妹窝数增加,雄蜂数量逐渐增加,第3个姊妹窝子代产出后有所回落;各姊妹窝子代之间性比也有显著差异(df=3, χ2 =1 091.142, P<0.001),第3个姊妹窝子代性比接近1:1。
白蜡吉丁肿腿蜂不同姊妹窝子代翅型分化和性比变化
Wing Dimorphism and Sex Ratio Changes in Progeny of Various Sister Broods in Parasitoid Sclerodermus pupariae (Hymenoptera: Bethylidae)
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摘要:
目的 探究具有多次产卵习性的寄生蜂不同姊妹窝子代种群参数的变化,从母代效应角度明确寄生蜂在自然栖境中增殖和延续种群的机制。 方法 室内30℃±1℃,光周期14L︰10D,相对湿度65%±5%的条件下测定了白蜡吉丁肿腿蜂在其成虫寿命期内所有姊妹窝子代产出过程中对麻竖毛天牛幼虫的寄生作用,比较各姊妹窝子代中雌蜂数量、发育历期、有翅型雌蜂比例和性比等参数的变化。 结果 白蜡吉丁肿腿蜂雌蜂对寄主幼虫的寄生能力和自身的产雌数量逐渐减低;各姊妹窝子代中雌性有翅率逐次递减,其中第3姊妹窝子代降幅最大,直至为零;各姊妹窝子代的雌雄性比呈抛物线趋势,第3姊妹窝子代雌雄性比接近1:1。 结论 白蜡吉丁肿腿蜂姊妹窝子代间翅的非遗传多型性可能是该蜂在应对林间异质环境的一种适应性进化。白蜡吉丁肿腿蜂第3姊妹窝子代的产出可能是其成虫寿命期内调控翅型和性比达到种群适合度最大化的关键节点。 Abstract:Objective This study aims to explore the changes of population parameters among different sister broods in parasitoids with multiple host attacking habit, so as to explain the mechanisms of expanding and sustaining population from the perspective of maternal effects. Method The parasitism on Thyestilla gebleri larva of Sclerodermus pupariae was studied during the production of successive sister broods in its lifetime. The number of female progeny, developmental duration, percentage of winged female, sex ratio and the other parameters of all the sister broods were recorded in the laboratory under the conditions of 30℃, photoperiod of 14L:0D, and RH 65% ±5%. Result The parasitism ability on host larvae and brood sizes reduced gradually. The percentage of winged females decreased with the number of produced sister broods till to zero, with the most dramatic decrease in the third sister brood. The sex ratio of progeny showed a parabolic trend at various successive sister broods and it was close to 1:1 for the third sister brood. Conclusion Wing polyphenism in various sister broods of S. pupariae may reflect adaptive evolution of this parasitoid coping with heterogeneous environment of forest. The production of the third sister brood might be a key point for regulating wing morph and sex ratio of progeny to maximize the fitness of the population in the lifetime in S. pupariae. -
Key words:
- Sclerodermus pupariae
- / maternal effect
- / sister brood
- / parasitic effect
- / percentage of winged female
- / sex ratio
- / fitness
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表 1 连续产出姊妹窝子代的白蜡吉丁肿腿蜂寄生能力和子代雌蜂数量
Table 1. Parasitic ability and number of female progeny of S. pupariae produced successive sister broods
母蜂所产姊妹窝次数
Number of sister brood produced by female重复数
Replicates寄生率
Parasitism rate/%寄生成功
Successful parasitism/%平均雌蜂产出量
Average female progeny1 50 98.0 a 92.0 a 70.54±16.03A 2 50 82.0 a 76.0 a 52.65±14.07A 3 37 83.8 a 81.1 a 21.97±17.31B 4 20 10.0 b 10.0 b 4.00±2.83B 5 1 0 0 0 表中数据为平均值±标准差。同列数据后不同的小写字母表示在0.008 3水平上差异显著(卡方检验);同列数据后不同大写字母表示在0.05水平上差异显著(单因素方差分析,LSD检验)。Data in the table are presented as mean ± SD. Those in the same column followed by different small letters are significantly different at the 0.008 3 level, by Chi-square test. Those in the same column followed by different capital letters are significantly different at the 0.05 level, by one-way ANOVA, LSD test. 表 2 连续产出姊妹窝子代的白蜡吉丁肿腿蜂产卵前期和子代发育历期
Table 2. Pre-oviposition duration and progeny developmental duration of S. pupariae produced successive sister brood
母蜂所产姊妹窝次数
Number of sister broodproduced by female产卵前期
Pre-ovipositionperiod卵期
Egg duration幼虫期
Larval duration雄蜂茧蛹期
Male pupalduration雌蜂茧蛹期
Female pupalduration雄蜂发育历期
Developmental duration of male雌蜂发育历期
Developmental duration of female1 5.37±0.57 b 2.65±0.85c 6.89±0.71a 12.53±0.72 a 12.74±0.68a 19.41±1.05 ab 19.63±1.14 b 2 5.45±0.67 b 3.63±0.54a 7.34±2.64a 11.58±1.08 b 11.89±1.01b 18.92±1.94 b 19.24±1.92 b 3 5.60±1.40 b 3.03±0.81b 5.73±1.17b 11.27±1.02 b 11.77±0.77 b 20.03±1.59 a 20.53±1.43 a 4 7.50±0.70 a 3.00±0.00abc 6.00±0.00ab 11.00±0.00 b 11.00±0.00b 20.00±0.00 ab 20.00±0.00ab 表中数据为平均值±标准差,同列数据后不同小写字母表示在0.05水平上差异显著(单因素方差分析,LSD检验)。Data in the table are presented as means followed ± SD, those in the same column, by different small letters are significantly different at the 0.05 level, by one-way ANOVA, LSD test. -
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