Chiodo Letizia, Malliavin Thérèse E, Maragliano Luca, Cottone Grazia, Ciccotti Giovanni
Center for Life Nano Science @Sapienza, Istituto Italiano di Tecnologia, Rome, Italy.
Institut Pasteur and CNRS UMR 3528, Unité de Bioinformatique Structurale, Paris, France.
PLoS One. 2015 Jul 24;10(7):e0133011. doi: 10.1371/journal.pone.0133011. eCollection 2015.
Nicotinic acetylcholine receptors (nAchRs) are ligand-gated ion channels that regulate chemical transmission at the neuromuscular junction. Structural information is available at low resolution from open and closed forms of an eukaryotic receptor, and at high resolution from other members of the same structural family, two prokaryotic orthologs and an eukaryotic GluCl channel. Structures of human channels however are still lacking. Homology modeling and Molecular Dynamics simulations are valuable tools to predict structures of unknown proteins, however, for the case of human nAchRs, they have been unsuccessful in providing a stable open structure so far. This is due to different problems with the template structures: on one side the homology with prokaryotic species is too low, while on the other the open eukaryotic GluCl proved itself unstable in several MD studies and collapsed to a dehydrated, non-conductive conformation, even when bound to an agonist. Aim of this work is to obtain, by a mixing of state-of-the-art homology and simulation techniques, a plausible prediction of the structure (still unknown) of the open state of human α7 nAChR complexed with epibatidine, from which it is possible to start structural and functional test studies. To prevent channel closure we employ a restraint that keeps the transmembrane pore open, and obtain in this way a stable, hydrated conformation. To further validate this conformation, we run four long, unbiased simulations starting from configurations chosen at random along the restrained trajectory. The channel remains stable and hydrated over the whole runs. This allows to assess the stability of the putative open conformation over a cumulative time of 1 μs, 800 ns of which are of unbiased simulation. Mostly based on the analysis of pore hydration and size, we suggest that the obtained structure has reasonable chances to be (at least one of the possible) structures of the channel in the open conformation.
烟碱型乙酰胆碱受体(nAchRs)是配体门控离子通道,可调节神经肌肉接头处的化学传递。目前已有真核受体开放和关闭形式的低分辨率结构信息,以及同一结构家族其他成员、两个原核直系同源物和一个真核谷氨酸氯离子通道的高分辨率结构信息。然而,人类通道的结构仍然缺失。同源建模和分子动力学模拟是预测未知蛋白质结构的重要工具,然而,就人类nAchRs而言,到目前为止,它们尚未成功提供稳定的开放结构。这是由于模板结构存在不同问题:一方面与原核物种的同源性过低,另一方面,在多项分子动力学研究中,开放的真核谷氨酸氯离子通道本身不稳定,即使与激动剂结合也会坍塌成脱水的非传导构象。这项工作的目的是通过结合最先进的同源性和模拟技术,对与埃博霉素复合的人类α7 nAChR开放状态的结构(仍然未知)进行合理预测,从而开展结构和功能测试研究。为防止通道关闭,我们采用一种限制措施使跨膜孔保持开放,从而获得稳定的水合构象。为进一步验证这种构象,我们从限制轨迹上随机选择的构型开始进行四次长时间的无偏模拟。在整个模拟过程中,通道保持稳定且水合。这使得我们能够在1微秒的累积时间内评估假定开放构象的稳定性,其中800纳秒为无偏模拟时间。主要基于对孔水合作用和大小的分析,我们认为所获得的结构很有可能是(至少一种可能的)通道开放构象的结构。