Beiles Avigdor, Raz Shmuel, Ben-Abu Yuval, Nevo Eviatar
Institute of Evolution, University of Haifa, Haifa, 31905, Israel.
Department of Computational Biology, University of Haifa, Haifa, 31905, Israel.
Biol Direct. 2015 Oct 14;10:58. doi: 10.1186/s13062-015-0074-5.
The current analysis of transposon elements (TE) in Drosophila melanogaster at Evolution Canyon, (EC), Israel, is based on data and analysis done by our collaborators (Drs. J. Gonzalez, J. Martinez and W. Makalowski, this issue). They estimated the frequencies of 28 TEs (transposon elements) in fruit flies (D. melanogaster) from the ecologically tropic, hot, and dry south-facing slope (SFS) or "African" slope (AS) of EC and compared it with the TE frequencies on the temperate-cool and humid north-facing slope (NFS) or "European" slope (ES), separated, on average, by 250 m. The flies were sampled from two stations on each slope. We received their results, including the frequencies of each TE on each slope, and the probabilities of the statistical analyses (G-tests) of each TE separately. We continued the analysis of the inter-slope differences of the frequencies of the TEs, and based our different conclusions on that analysis and on the difference between micro (=EC) and macro (2000 km.) comparisons [Gonzalez et al. 2015 doi: 10.1186/s13062-015-0075-4 ].
Our collaborators based all their conclusions on the non-significant results of each of the individual tests of the 28 TEs. We analysed also the distribution of the TE differences between the slopes, based on their results. Thirteen TEs were more frequent on the SFS, 11 were more frequent on the NFS, and four had equal frequencies. Because of the equalizing effect of the ongoing migration, only small and temporary differences between the slopes (0 - 0.06) were regarded by us as random fluctuations (drift). Three TEs were intermediate (0.08-0.09) and await additional research. The 11 TEs with large frequency differences (0.12 - 0.22) were regarded by us as putative adaptive TEs, because the equalizing power of ongoing migration will eliminate random large differences. Five of them were higher on the SFS and six were higher on the NFS. Gaps in the distribution of the differences distinguished between the large and small differences. The large gap among the 11 TEs favored on the NFS was significant and supports our rejection of drift as the only explanation of the distribution of the slope differences. The gaps in the distribution of the differences separated the putative TEs with strong enough selection from those TEs that couldn't overrule the migration. The results are compared and contrasted with the directional effect of the frequencies of the same TEs in the study of global climatic comparisons across thousands of kilometers. From the 11 putative adaptive TEs in the local "Evolution Canyon," six differentiate in the same direction as in the continental comparisons and four in the opposite direction. One TE, FBti0019144, differentiated in EC in the same direction as in Australia and in the opposite direction to that of North America.
We presume that the major divergent evolutionary driving force at the local EC microsite is natural selection overruling gene flow. Therefore, after we rejected drift as an explanation of all the large slope differences, we regarded them as putatively adaptive. In order to substantiate the individual TE adaptation, we need to increase the sample sizes and reveal the significant adaptive TEs. The comparison of local and global studies show only partial similarity in the adaptation of the TEs, because of the dryness of the ecologically tropical climate in EC, in contrast to the wet tropical climate in the global compared climates. Moreover, adaptation of a TE may be expressed only in part of the time and specific localities.
目前对以色列进化峡谷(EC)中黑腹果蝇转座子元件(TE)的分析,是基于我们的合作者(J. 冈萨雷斯博士、J. 马丁内斯博士和W. 马卡洛维斯基博士,本期)所做的数据和分析。他们估计了来自EC生态热带、炎热干燥的朝南坡(SFS)或“非洲”坡(AS)的果蝇(黑腹果蝇)中28种TE(转座子元件)的频率,并将其与平均相距250米的温带凉爽湿润的朝北坡(NFS)或“欧洲”坡(ES)上的TE频率进行了比较。果蝇是从每个坡的两个站点采集的。我们收到了他们的结果,包括每个坡上每种TE的频率,以及每种TE单独进行统计分析(G检验)的概率。我们继续分析了TE频率的坡间差异,并基于该分析以及微观(=EC)和宏观(2000公里)比较之间的差异得出了不同结论[冈萨雷斯等人,2015年,doi:10.1186/s13062 - 015 - 0075 - 4]。
我们的合作者所有结论均基于对28种TE各自单独测试的不显著结果。我们也根据他们的结果分析了坡间TE差异的分布。13种TE在SFS上更常见,11种在NFS上更常见,4种频率相等。由于持续迁移的均衡作用,我们认为坡间只有微小且暂时的差异(0 - 0.06)是随机波动(漂变)。3种TE处于中间水平(0.08 - 0.09),有待进一步研究。我们认为11种频率差异较大(0.12 - 0.22)的TE是假定的适应性TE,因为持续迁移的均衡作用会消除随机的大差异。其中5种在SFS上频率更高,6种在NFS上频率更高。差异分布中的间隙区分了大差异和小差异。在NFS上受青睐的11种TE之间的大间隙很显著,支持我们拒绝将漂变作为坡间差异分布的唯一解释。差异分布中的间隙将具有足够强选择作用的假定TE与那些无法排除迁移影响的TE区分开来。将这些结果与在数千公里全球气候比较研究中相同TE频率的定向效应进行了比较和对比。在当地“进化峡谷”的11种假定适应性TE中,6种与大陆比较中的方向相同,4种相反。一种TE,FBti0019144,在EC中的分化方向与在澳大利亚相同,与北美相反。
我们推测在当地EC微地点主要的分化进化驱动力是自然选择压倒基因流。因此,在我们拒绝将漂变作为所有大坡间差异的解释后,我们认为它们是假定适应性的。为了证实单个TE的适应性,我们需要增加样本量并找出显著的适应性TE。局部和全球研究的比较表明,由于EC生态热带气候干燥,与全球比较气候中的湿润热带气候不同,TE的适应性仅显示出部分相似性。此外,TE的适应性可能仅在部分时间和特定地点表现出来。