Silverman Scott K
Department of Chemistry, University of Illinois at Urbana-Champaign, 600 South Mathews Avenue, Urbana, Illinois 61801, United States.
Acc Chem Res. 2015 May 19;48(5):1369-79. doi: 10.1021/acs.accounts.5b00090. Epub 2015 May 5.
Catalysis is a fundamental chemical concept, and many kinds of catalysts have considerable practical value. Developing entirely new catalysts is an exciting challenge. Rational design and screening have provided many new small-molecule catalysts, and directed evolution has been used to optimize or redefine the function of many protein enzymes. However, these approaches have inherent limitations that prompt the pursuit of different kinds of catalysts using other experimental methods. Nature evolved RNA enzymes, or ribozymes, for key catalytic roles that in modern biology are limited to phosphodiester cleavage/ligation and amide bond formation. Artificial DNA enzymes, or deoxyribozymes, have great promise for a broad range of catalytic activities. They can be identified from unbiased (random) sequence populations as long as the appropriate in vitro selection strategies can be implemented for their identification. Notably, in vitro selection is different in key conceptual and practical ways from rational design, screening, and directed evolution. This Account describes the development by in vitro selection of DNA catalysts for many different kinds of covalent modification reactions of peptide and protein substrates, inspired in part by our earlier work with DNA-catalyzed RNA ligation reactions. In one set of studies, we have sought DNA-catalyzed peptide backbone cleavage, with the long-term goal of artificial DNA-based proteases. We originally anticipated that amide hydrolysis should be readily achieved, but in vitro selection instead surprisingly led to deoxyribozymes for DNA phosphodiester hydrolysis; this was unexpected because uncatalyzed amide bond hydrolysis is 10(5)-fold faster. After developing a suitable selection approach that actively avoids DNA hydrolysis, we were able to identify deoxyribozymes for hydrolysis of esters and aromatic amides (anilides). Aliphatic amide cleavage remains an ongoing focus, including via inclusion of chemically modified DNA nucleotides in the catalyst, which we have recently found to enable this cleavage reaction. In numerous other efforts, we have investigated DNA-catalyzed peptide side chain modification reactions. Key successes include nucleopeptide formation (attachment of oligonucleotides to peptide side chains) and phosphatase and kinase activities (removal and attachment of phosphoryl groups to side chains). Through all of these efforts, we have learned the importance of careful selection design, including the frequent need to develop specific "capture" reactions that enable the selection process to provide only those DNA sequences that have the desired catalytic functions. We have established strategies for identifying deoxyribozymes that accept discrete peptide and protein substrates, and we have obtained data to inform the key choice of random region length at the outset of selection experiments. Finally, we have demonstrated the viability of modular deoxyribozymes that include a small-molecule-binding aptamer domain, although the value of such modularity is found to be minimal, with implications for many selection endeavors. Advances such as those summarized in this Account reveal that DNA has considerable catalytic abilities for biochemically relevant reactions, specifically including covalent protein modifications. Moreover, DNA has substantially different, and in many ways better, characteristics than do small molecules or proteins for a catalyst that is obtained "from scratch" without demanding any existing information on catalyst structure or mechanism. Therefore, prospects are very strong for continued development and eventual practical applications of deoxyribozymes for peptide and protein modification.
催化作用是一个基本的化学概念,许多种类的催化剂都具有相当大的实用价值。开发全新的催化剂是一项令人兴奋的挑战。合理设计和筛选已经提供了许多新型小分子催化剂,定向进化已被用于优化或重新定义许多蛋白质酶的功能。然而,这些方法存在固有的局限性,这促使人们使用其他实验方法来寻找不同类型的催化剂。自然界进化出了RNA酶(即核酶),用于关键的催化作用,在现代生物学中,这些作用仅限于磷酸二酯键的切割/连接和酰胺键的形成。人工DNA酶(即脱氧核酶)在广泛的催化活性方面具有巨大潜力。只要能够实施合适的体外筛选策略来鉴定它们,就可以从无偏向性(随机)的序列群体中识别出脱氧核酶。值得注意的是,体外筛选在关键的概念和实践方面与合理设计、筛选及定向进化不同。本综述描述了通过体外筛选开发用于肽和蛋白质底物多种不同类型共价修饰反应的DNA催化剂的过程,这部分受到我们早期DNA催化RNA连接反应工作的启发。在一组研究中,我们致力于寻找DNA催化的肽主链切割方法,其长期目标是开发基于人工DNA的蛋白酶。我们最初预计酰胺水解应该很容易实现,但体外筛选却意外地得到了用于DNA磷酸二酯水解的脱氧核酶;这是出乎意料的,因为未催化的酰胺键水解速度要快10^5倍。在开发出一种能够有效避免DNA水解的合适筛选方法后,我们成功鉴定出了用于酯和芳香酰胺(酰苯胺)水解的脱氧核酶。脂肪族酰胺的切割仍是一个持续关注的重点,包括通过在催化剂中引入化学修饰的DNA核苷酸来实现,我们最近发现这能够使这种切割反应发生。在许多其他研究中,我们研究了DNA催化的肽侧链修饰反应。主要成果包括核肽的形成(寡核苷酸与肽侧链的连接)以及磷酸酶和激酶活性(磷酸基团从侧链上去除和连接到侧链上)。通过所有这些努力,我们认识到精心设计筛选的重要性,包括经常需要开发特定的“捕获”反应,以使筛选过程仅提供那些具有所需催化功能的DNA序列。我们已经建立了识别能够接受离散肽和蛋白质底物的脱氧核酶的策略,并且我们已经获得了数据,以便在筛选实验开始时为随机区域长度的关键选择提供参考。最后,我们证明了包含小分子结合适体结构域的模块化脱氧核酶的可行性,尽管发现这种模块化的价值很小,但这对许多筛选工作具有启示意义。本综述中总结的这些进展表明,DNA在生物化学相关反应中具有相当大的催化能力,特别是包括共价蛋白质修饰。此外,对于一种“从头开始”获得且不依赖任何关于催化剂结构或机制的现有信息的催化剂而言,DNA具有与小分子或蛋白质截然不同且在许多方面更优的特性。因此,脱氧核酶在肽和蛋白质修饰方面持续发展并最终实现实际应用的前景非常广阔。