Study on the cytoplasmic incompatibility in some natural population of Culex pipiens complex in China and Japan

The cytoplasmic incompatibility in the field population of Culex pipiens complex from China and Japan was studied. The results of the crossing experiments between 11 different strains suggest that some Japanese strains might be possible to be used for control Chinese strains by using male sterile technique, and the heavy infection of wolbachiae in mosquitoes may have some relations with the cytoplasmic incompatibility in the field population of Cx. pipiens complex.     

Taxonomy of oxalotrophic <Emphasis Type="Italic">Methylobacterium</Emphasis> strains

Most of the oxalotrophic bacteria are facultative methylotrophs and play important ecological roles in soil fertility and cycling of elements. This study gives a detailed picture of the taxonomy and diversity of these bacteria and provides new information about the taxonomical variability within the genus Methylobacterium. Twelve mesophilic, pink-pigmented, and facultatively methylotrophic oxalate-oxidizing strains were included in this work that had been previously isolated from the soil and some plant tissues by the potassium oxalate enrichment method. The isolates were characterized using biochemical tests, cellular lipid profiles, spectral characteristics of carotenoid pigments, G+C content of the DNA, and 16S rDNA sequencing. The taxonomic similarities among the strains were analyzed using the simple matching (S _SM) and Jaccard (S _J) coefficients, and the UPGMA clustering algorithm. The phylogenetic position of the strains was inferred by the neighbor-joining method on the basis of the 16S rDNA sequences. All isolates were Gram-negative, facultatively methylotrophic, oxidase and catalase positive, and required no growth factors. Based on the results of numerical taxonomy, the strains formed four closely related clusters sharing ≥85% similarity. Analysis of the 16S rDNA sequences demonstrated that oxalotrophic, pink-pigmented, and facultatively methylotrophic strains could be identified as members of the genus Methylobacterium. Except for M. variabile and M. aquaticum, all of the Methylobacterium type strains tested had the ability of oxalate utilization. Our results indicate that the capability of oxalate utilization seems to be an uncommon trait and could be used as a valuable taxonomic criterion for differentiation of Methylobacterium species.     

Does a parthenogenesis-inducing <Emphasis Type="Italic">Wolbachia</Emphasis> induce vestigial cytoplasmic incompatibility?

Wolbachia is a maternally inherited bacterium that manipulates the reproduction of its host. Recent studies have shown that male-killing strains can induce cytoplasmic incompatibility (CI) when introgressed into a resistant host. Phylogenetic studies suggest that transitions between CI and other Wolbachia phenotypes have also occurred frequently, raising the possibility that latent CI may be widespread among Wolbachia. Here, we investigate whether a parthenogenesis-inducing Wolbachia strain can also induce CI. Parthenogenetic females of the parasitoid wasp Asobara japonica regularly produce a small number of males that may be either infected or not. Uninfected males were further obtained through removal of the Wolbachia using antibiotics and from a naturally uninfected strain. Uninfected females that had mated with infected males produced a slightly, but significantly more male-biased sex ratio than uninfected females that had mated with uninfected males. This effect was strongest in females that mated with males that had a relatively high Wolbachia titer. Quantitative PCR indicated that infected males did not show higher ratios of nuclear versus mitochondrial DNA content. Wolbachia therefore does not cause diploidization of cells in infected males. While these results are consistent with CI, other alternatives such as production of abnormal sperm by infected males cannot be completely ruled out. Overall, the effect was very small (9%), suggesting that if CI is involved it may have degenerated through the accumulation of mutations.     

Non-specific association between filamentous bacteria and fungus-growing ants

Fungus-growing ants and their fungal cultivar form a highly evolved mutualism that is negatively affected by the specialized parasitic fungus Escovopsis. Filamentous Pseudonocardia bacteria occurring on the cuticle of attine ants have been proposed to form a mutualistic interaction with these ants in which they are vertically transmitted (i.e. from parent to offspring colonies). Given a strictly vertical transmission of Pseudonocardia, the evolutionary theory predicts a reduced genetic variability of symbionts among ant lineages. The aim of this study was to verify whether actinomycetes, which occur on Acromyrmex octospinosus leaf-cutting ants, meet this expectation by comparing their genotypic variability with restriction fragment length polymorphisms. Multiple actinomycete strains could be isolated from both individual ant workers and colonies (one to seven strains per colony). The colony specificity of actinomycete communities was high: Only 15% of all strains were isolated from more than one colony, and just 5% were present in both populations investigated. Partial sequencing of 16S ribosomal deoxyribonucleic acid of two of the isolated strains assigned both of them to the genus Streptomyces. Actinomycetes could also be isolated from workers of the two non-attine ant species Myrmica rugulosa and Lasius flavus. Sixty-two percent of the strains derived from attine ants and 80% of the strains isolated from non-attine ants inhibited the growth of Escovopsis. Our data suggest that the association between attine ants and their actinomycete symbionts is less specific then previously thought. Soil-dwelling actinomycetes may have been dynamically recruited from the environment (horizontal transmission), probably reflecting an adaptation to a diverse community of microbial pathogens.     

金属离子诱导Trichoderma sp.WL-Go合成纳米金特性研究

生物法合成纳米金是一种环境友好且经济高效的合成途径,受到广泛关注.普遍认为微生物合成纳米金是通过胞外分泌生物大分子而实现的一种自发脱毒过程.同时,相关研究表明低浓度的重金属离子对菌株胞外酶的活性会产生一定的影响,进而会对菌株合成纳米金的能力产生影响.基于此,本研究选择一株前期筛选得到的真菌Trichoderma sp. WL-Go,探究不同金属离子诱导菌株对其合成的纳米金特性的影响.结果表明,经Co~(2+)、Al~(3+)、Zn~(2+)、Sn~(2+)、Ni~(2+)等金属离子诱导后,菌株WL-Go合成纳米金的能力均有所提升,而Pb~(2+)、Cu~(2+)、Fe~(3+)与对照组相比,其合成的纳米金浓度及转化率都无明显变化.此外,Co~(2+)诱导菌株合成的纳米金呈现肉眼可见的团簇状,发生明显的团聚现象.本实验还考察了生物合成纳米金对4-硝基苯酚还原的催化特性,结果表明,Sn~(2+)和Pb~(2+)的诱导使菌株WL-Go合成的纳米金催化速率得到明显提升,而其他金属离子均有所抑制.最后选取革兰氏阳性菌Arthrobacter sp. W1和革兰氏阴性菌Escherichia coli BL21 (DE3)验证了不同金属离子诱导后合成的纳米金均具有良好的生物相容性.综上,本研究为对于拓宽纳米金的工业化应用前景有着重要意义.     

Accelerated biodegradation of nitrophenols in the rhizosphere of Spirodela polyrrhiza

We investigated the biodegradation of 2-nitrophenol (2-NP), 4-nitrophenol (4-NP), and 2,4-dinitrophenol (2,4-DNP) in the rhizosphere of Spirodela polyrrhiza plants by conducting degradation experiments with three river water samples supplemented with each nitrophenol (NP). We then isolated NP-degrading bacteria both from the S. polyrrhiza roots and from the river water. In the river water samples, removal of the three NP was accelerated in the presence of S. polyrrhiza plants. The three NPs persisted in an autoclaved solution with sterile plants suggests that NP removal was accelerated largely by bacterial NP biodegradation rather than by adsorption and uptake by the plants. We isolated 8 strains of NP-degrading bacteria: 6 strains from the S. polyrrhiza roots and 2 strains from river water without the plants. The 2-NP- and 2,4-DNP-degrading bacteria were isolated only from the S. polyrrhiza roots. The 4-NPdegrading bacteria different from those isolated from the river water samples were also found on S. polyrrhiza roots. The 2-NP- and 4-NP-degrading strains isolated from the roots utilized the corresponding NP (0.5 mmol/L) as the sole carbon and energy source. The 2,4-DNP-degrading strains isolated from the roots showed substantial 2,4-DNP-degrading activity, but the presence of other carbon and energy sources was required for their growth. The isolated NP-degrading bacteria from the roots must have contributed to the accelerated degradation of the three NPs in the rhizosphere of S. polyrrhiza. Our results suggested that rhizoremediation with S. polyrrhiza may be effective for NP-contaminated surface water.     

Isolation of five Rubrobacter strains from biodeteriorated monuments

In the last few years, the microbial colonisation of mural paintings in ancient monuments has been attracting the attention of microbiologists and conservators. The genus Rubrobacter is commonly found in biodeteriorated monuments, where it has been reported to cause rosy discolouration. However, to date, only three species of this genus have been isolated, all from thermophilic environments. In this paper, we studied three monuments: the Servilia and Postumio tombs in the Roman Necropolis of Carmona (Spain), and Vilar de Frades church (Portugal), in search of Rubrobacter strains. In all cases, biodeterioration and the formation of efflorescences were observed, and five Rubrobacter strains were isolated. These isolates showed different physiology and migration in denaturing gradient gel electrophoresis, suggesting they might represent new species within this genus. The isolates reproduced some biodeterioration processes in the laboratory and revealed their biomediation in crystal formation.     

2株苯胺降解的分离鉴定及其降解特性研究

从处理印染废水的活性污泥中分离得到2株苯胺降解菌,从菌落、细胞形态、生理生化及16S rRNA基因扩增测序等方面对2株菌进行了鉴定,并比较分析2株菌在好氧与缺氧条件下的苯胺降解、偶氮染料脱色及苯胺脱氨氧化酶基因tdnQ和黄素还原酶基因(fre)的携带情况.结果表明,2株菌属于Pseudomonas属和Shewanella属,分别命名为Pseudomonas sp.AN30和Shewanella sp.DN425.AN30菌株在振荡好氧条件下72h内对250mg/L苯胺的降解率为96.1%,DN425菌株的降解率为13.8%;在静置缺氧条件下AN30菌株的苯胺降解率为39.6%,DN425菌株的降解率仅为8.6%.DN425菌株在静置缺氧条件下4h内可将初始浓度为50 mg/L的偶氮染料酸性大红彻底脱色,而AN30菌株对酸性大红不具有脱色能力.以总DNA为模板,分别用tdnQ基因和fre基因特异性引物进行扩增,2株菌均能扩增出大小分别为380bp和630bp左右的目标条带,显示2菌株均携带有苯胺脱氨氧化酶基因和黄素还原酶基因.     

Shinella zoogloeoides BC026对吡啶的降解特性研究

从首钢焦化厂的污水处理系统中分离1株能以吡啶为唯一碳、氮源的细菌BC026,它具有自絮凝特性,对卡那霉素、氨苄青霉素和壮观霉素具有抗性,可在阿须贝无氮培养基中良好生长.通过16S rRNA序列分析和Biolog微生物鉴定系统鉴定,确定该菌为Shinella zoogloeoides.纯菌对单基质的降解实验表明,在30℃、180 r/min和pH为7的条件下,当投菌量为0.1 g/L时,BC026可在17 h内将400 mg/L吡啶完全降解;在吡啶初始浓度为99～1 806 mg/L的无机盐培养基中,BC026均能保持降解活性,较高初始浓度的吡啶对BC026的生长产生一定抑制,但BC026在适应后对吡啶的降解速率较快;降解最适温度为30～35℃,最适pH为8.BC026对吡啶的代谢途径研究表明：降解的第一步是断开吡啶的2条C—N链,生成氨氮和戊二醛,随后戊二醛被氧化为戊二酸,并最终转化为二氧化碳和水;吡啶中的氮有59.5%转化成氨氮.     

Atlantic sturgeons (<Emphasis Type="Italic">Acipenser sturio</Emphasis>, <Emphasis Type="Italic">Acipenser oxyrinchus</Emphasis>): American females successful in Europe

Recent molecular data on the maternally inherited mitochondrial (mt) DNA have challenged the traditional view that the now extinct Baltic sturgeon population belonged to the European sturgeon Acipenser sturio. Instead, there is evidence that American sea sturgeon Acipenser oxyrinchus historically immigrated into the Baltic Sea. In this study, we test the hypothesis that A. oxyrinchus introgressed into, rather than replaced, the A. sturio population in the Baltic. We established four single nucleotide polymorphisms (SNPs) in the nuclear MHC II antigen gene with a species-specific SNP pattern. Using an ancient DNA approach and two independent lines of molecular evidence (sequencing of allele-specific clones, SNaPshot), we detected both A. sturio and A. oxyrinchus alleles in the available museum material of the now extinct Baltic sturgeon population. The hybrid nature of the Baltic population was further confirmed by very high levels of heterozygosity. It had been previously postulated that the immigration of the cold-adapted A. oxyrinchus into the Baltic occurred during the Medieval Little Ice Age, when temperature likely dropped below the degree inducing spawning in A. sturio. Under this scenario, our new findings suggest that the genetic mosaic pattern in the Baltic sturgeon population (oxyrinchus mtDNA, sturio and oxyrinchus MHC alleles) is possibly caused by sex-biased introgression where spawning was largely restricted to immigrating American females, while fertilization was predominantly achieved by abundant local European males. The hybrid nature of the former Baltic sturgeon population should be taken into account in the current reintroduction measures. Electronic supplementary material  Supplementary material is available in the online version of this article at and is accessible for authorized users.     

热纤维梭菌不同菌株的纤维素降解和酒精生产能力

以高温厌氧细菌热纤维梭菌(Clostridium thermocellum LQRI和VPI菌株)和嗜热厌氧乙醇菌(Thermoanaerobacterethanolicus X514和39E菌株)为对象,以不同浓度纤维素为底物,分析了热纤维梭菌不同菌株的纤维素降解和酒精生产能力.结果表明,在热纤维梭菌纯培养体系中,LQRI对纤维素的降解能力明显高于VPI,前者约为后者的1.2倍,但两者的酒精生产能力没有显著差异,LQRI在1%纤维素条件下酒精终浓度约为12 mmol/L,VPI约为10 mmol/L.随着底物纤维素浓度的增加(1%、2%、5%),两者对纤维素降解和酒精生产能力均呈明显下降趋势.在热纤维梭菌和嗜热厌氧乙醇菌混合培养体系中,LQRI对纤维素的降解能力明显高于VPI,前者约为后者的1.28～1.58倍.随着底物浓度的增加,两者的纤维素降解率均逐渐下降,但热纤维梭菌对纤维素降解的绝对数值还受到其它混合培养菌种的影响.此外,LQRI+Thermoanaerobacter混合培养体系的酒精生产能力明显高于VPI+Thermoanaerobacter,前者约为后者的1.27～1.77倍.随着底物浓度的增加,混合培养体系酒精生产能力没有明显下降趋势.     

拟柱孢藻生长及碱性磷酸酶活性对不同磷浓度和磷形态响应的株系间差异

拟柱孢藻(Cylindrospermopsis raciborskii)是热带地区普遍存在的蓝藻种类,近年来广泛扩张到温带地区水体,耐受低磷环境和多生态型的存在被认为是该藻能成功入侵的重要原因.为进一步了解不同藻株对磷波动的生理响应是否存在差异,本研究以广东省镇海水库分离的4株拟柱孢藻(N1、 N8、 N9和N10)为材料,观测它们的生长和碱性磷酸酶(ALP)活性在不同无机磷(Pi)浓度(HP=7.13 mg·L^-1、 MP=0.64 mg·L^-1、 LP=0.03 mg·L^-1)和磷形态[磷酸氢二钾(K₂HPO₄)、焦磷酸钾(K₄P₂O₇)、三聚磷酸钾(K₅P₃O₁₀)、 D-葡萄糖-6-磷酸(D-G-6-P)、三磷酸腺苷(ATP)、环磷酸腺苷(cAMP)]的变化.结果表明, 4株拟柱孢藻的生长对Pi浓度变化的响应基本一致,即它们的生物量都随Pi...     

Moss-associated methylobacteria as phytosymbionts: an experimental study

Methylotrophic bacteria inhabit the surface of plant organs, but the interaction between these microbes and their host cells is largely unknown. Protonemata (gametophytes) of the moss Funaria hygrometrica were cultivated in vitro under axenic conditions and the growth of the protonemal filaments recorded. In the presence of methylobacteria (different strains of Methylobacterium), average cell length and the number of cells per filament were both enhanced. We tested the hypothesis that auxin (indole-3-acetic acid, IAA), secreted by the epiphytic bacteria and taken up by the plant cells, may in part be responsible for this promotion of protonema development. The antiauxin parachlorophenoxyisobutyric acid (PCIB) was used as a tool to analyze the role of IAA and methylobacteria in the regulation of cell growth. In the presence of PCIB, cell elongation and protonema differentiation were both inhibited. This effect was compensated for by the addition of different Methylobacterium strains to the culture medium. Biosynthesis and secretion of IAA by methylobacteria maintained in liquid culture was documented via a colorimetric assay and thin layer chromatography. Our results support the hypothesis that the development of Funaria protonemata is promoted by beneficial phytohormone-producing methylobacteria, which can be classified as phytosymbionts.     

2株芽孢杆菌利用不同碳源生产絮凝剂的研究

探讨了在纯培养和混合培养的条件下,2株芽孢杆菌F2和F6生产高效絮凝剂过程中对碳源的利用情况.研究表明,多种分子量在200以下的有机物能够作为碳源用于高效絮凝剂的生产,包括:单糖、双糖、醇、有机酸和酯类等,具有一定的普适性.发现利用废糖蜜、生物制氢废液和纤维素糖化液等廉价的生物质废料能够生产出性能良好的生物絮凝剂.     

Contribution of anadromous fish to the diet of European catfish in a large river system

Many anadromous fish species, when migrating from the sea to spawn in fresh waters, can potentially be a valuable prey for larger predatory fish, thereby efficiently linking these two ecosystems. Here, we assess the contribution of anadromous fish to the diet of European catfish (Silurus glanis) in a large river system (Garonne, southwestern France) using stable isotope analysis and allis shad (Alosa alosa) as an example of anadromous fish. Allis shad caught in the Garonne had a very distinct marine δ¹³C value, over 8‰ higher after lipid extraction compared to the mean δ¹³C value of all other potential freshwater prey fish. The δ¹³C values of European catfish varied considerably between these two extremes and some individuals were clearly specializing on freshwater prey, whereas others specialized on anadromous fish. The mean contribution of anadromous fish to the entire European catfish population was estimated to be between 53% and 65%, depending on the fractionation factor used for δ¹³C.     

南方冬季条件下根际促生菌与肥料配施对巨菌草迁移累积镉砷的影响

优良菌种是微生物肥料的基础,研究微生物肥料中低温适应型菌株在低温条件下的促生性,对于提高南方冬闲时植物修复重金属污染农田具有重要意义.通过形态观察及18S rRNA基因序列分析,对菌株进行了鉴定,在此基础上进行植物盆栽试验,分析了3株不同根际促生菌单独施加及其与肥料（鸡粪+微量元素）配施对镉砷复合污染土壤中巨菌草迁移累积Cd、As的影响.分离菌株的初步鉴定结果表明,3株根际促生菌（M1、M2、M3）分别为云南木霉（Trichoderma yunnanense）、卵形孢球托霉（Gongronella butleri）、越橘间座壳菌（Diaporthe vaccinii）.纯培养试验表明,M1、M2、M3均可在南方冬季低温条件下生长.盆栽试验结果表明,与CK处理相比,单独施菌或肥料及菌株与肥料配施均能提高巨菌草茎叶生物量,分别提高14.12%~84.35%、13.36%、57.25%~143.13%.在所有处理中,M3与肥料配施后,植株茎叶生物量达到最大.不同处理可影响巨菌草对Cd和As的吸收累积,但对巨菌草各部位影响不同.单独施菌时,M3对巨菌草茎叶Cd、As积累量提升最多,分别提高145.83%、64.03%;当菌株与肥料配施时,M3与肥料配施对巨菌草茎叶Cd、As积累量提升最多,分别提高94.17%、142.45%.可见,3株根际促生菌在南方冬季均能提高巨菌草生物量,其中,菌株M3与肥料配施具有开发为有效治理镉砷复合污染土壤的微生物肥料的良好前景.     

电磁环境胁迫对铜绿微囊藻氧化应激和光合固碳酶的影响作用研究

研究了3种典型电磁环境(射频电磁辐射(RF-EMF)、极低频电磁场(ELF-EMF)、交变电场(AEF))胁迫对铜绿微囊藻(Microcystis aeruginosa)细胞氧化应激(超氧化物歧化酶(SOD)、丙二醛(MDA))和光合固碳酶(核酮糖-1,5-二磷酸羧化酶/加氧酶(Rubisco)、果糖-1,6-二磷酸醛缩酶(FBA))的影响作用.结果表明,短期和长期RF-EMF、ELF-EMF及AEF暴露胁迫铜绿微囊藻细胞,总体能极显著提高藻细胞的SOD活性(p0.01).铜绿微囊藻经RF-EMF、ELF-EMF及AEF短期暴露处理后,均极显著提高了藻细胞内MDA含量(p0.01);铜绿微囊藻经RFEMF、AEF长期暴露处理后,藻细胞内MDA含量极显著降低(p0.01).RF-EMF暴露处理铜绿微囊藻,总体能显著提高藻细胞Rubisco酶活性(p0.05).长期RF-EMF、AEF和ELF-EMF暴露下的铜绿微囊藻,总体表现出FBA酶活性极显著降低(p0.01).以上结果说明,RF-EMF、ELFEMF、AEF均能够诱导铜绿微囊藻发生氧化应激反应.而RF-EMF能够对铜绿微囊藻细胞参与光合作用的关键酶(Rubisco和FBA)产生一定的调控作用,表明射频电磁波能够在一定程度上影响铜绿微囊藻细胞的光合作用.     

腾格里沙漠东南缘不同生境白刺(Nitraria)灌丛沙堆的空间分布格局

运用地统计学和数量生态学方法,从维持天然白刺灌丛稳定性和持久生态效益角度出发,对腾格里沙漠东南缘不同生境白刺灌丛沙堆的空间异质性进行了研究,分析了不同生境白刺固沙生态效益.结果表明,生境地下水埋深和白刺植株发育状况的差异直接影响了白刺耐沙埋的能力,间接地影响了白刺沙堆的固沙效应.湖盆低地白刺种群的植株盖度、密度和生物量显著高于山前冲积扇区（p<0.01）,植株高度则相反,且无显著差异（p>0.05）.湖盆低地的半固定生境沙堆高度平均为1.20 m,单个沙堆积沙体积平均为88.19 m³,固定生境的山前冲积扇沙堆高度平均为1.14 m,单个沙堆积沙体积平均为33.16 m³.不同生境条件下,白刺沙堆的斑块大小和分布存在明显差异（p<0.01）.湖盆低地的白刺灌丛沙堆趋向于大斑块、小密度,空间自相关距离较长,而山前冲积扇区相反.湖盆低地由自相关因素引起的沙堆大小和积沙体积空间异质性占主要部分,分别体现在1.2～84 m和1.2～505 m尺度;而山前冲积扇的随机因素引起的沙堆大小和积沙体积空间异质性占到100％,在100 m以下尺度该区白刺灌丛沙堆大小和积沙体积具有恒定的变异,其空间分布格局在这个尺度无规律体现呈随机分布.     

2株海洋石油降解细菌的降解能力

为实施海洋石油污染的生物治理,从厦门储油码头油污水中分离得到了2株石油降解菌.它们能够在以柴油、萘或芘为唯一碳源的培养基中生长,并适应于海洋环境的温度、pH和盐度.它们对萘都有很强的降解能力,3d内降解率可达87.53%和84.01%;7 d内对芘降解率分别为8.35%、5.37%.经16S rDNA同源性分析表明2菌株之间及其与施氏假单胞菌的同源性皆为99%;Biolog生化鉴定也表明它们为不同的菌株.通过兼并PCR扩增,序列分析发现2种菌编码完全相同的萘双加氧酶基因,该基因与其它假单胞菌的萘双加氧酶大亚基有98%同源性.     

Bacteria on housefly eggs, Musca domestica, suppress fungal growth in chicken manure through nutrient depletion or antifungal metabolites

Female houseflies, Musca domestica (Diptera: Muscidae), lay their eggs in ephemeral resources such as animal manure. Hatching larvae compete for essential nutrients with fungi that also colonize such resources. Both the well-known antagonistic relationship between bacteria and fungi and the consistent presence of the bacterium Klebsiella oxytoca on housefly eggs led us to hypothesize (1) that K. oxytoca, and possibly other bacteria on housefly eggs, help curtail the growth of fungal resource competitors and (2) that such fungi indeed adversely affect the development of housefly larvae. Bacteria washed from housefly eggs significantly reduced the growth of fungi in chicken manure. Nineteen bacterial strains and ten fungal strains were isolated from housefly eggs or chicken manure, respectively. Co-culturing each of all the possible bacterium–fungus pairs revealed that the bacteria as a group, but no single bacterium, significantly suppressed the growth of all fungal strains tested. The bacteria's adverse effect on fungi is due to resource nutrient depletion and/or the release of antifungal chemicals. Well-established fungi in resources significantly reduced the number of larval offspring that completed development to adult flies. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.