Pratico Elizabeth D, Wang Yangming, Silverman Scott K
Department of Chemistry, University of Illinois at Urbana-Champaign 600 South Mathews Avenue, Urbana, IL 61801, USA.
Nucleic Acids Res. 2005 Jun 20;33(11):3503-12. doi: 10.1093/nar/gki656. Print 2005.
RNA molecules with internal 2',5'-branches are intermediates in RNA splicing, and branched RNAs have recently been proposed as retrotransposition intermediates. A broadly applicable in vitro synthetic route to branched RNA that does not require self-splicing introns or spliceosomes would substantially improve our ability to study biochemical processes that involve branched RNA. We recently described 7S11, a deoxyribozyme that was identified by in vitro selection and has general RNA branch-forming ability. However, an important restriction for 7S11 is that the branch-site RNA nucleotide must be a purine (A or G), because a pyrimidine (U or C) is not tolerated. Here, we describe the compact 6CE8 deoxyribozyme (selected using a 20 nt random region) that synthesizes 2',5'-branched RNA with any nucleotide at the branch site. The Mn2+-dependent branch-forming ligation reaction is between an internal branch-site 2'-hydroxyl nucleophile on one RNA substrate with a 5'-triphosphate on another RNA substrate. The preference for the branch-site nucleotide is U > C congruent with A > G, although all four nucleotides are tolerated with useful ligation rates. Nearly all other nucleotides elsewhere in both RNA substrates allow ligation activity, except that the sequence requirement for the RNA strand with the 5'-triphosphate is 5'-pppGA, with 5'-pppGAR (R = purine) preferred. These characteristics permit 6CE8 to prepare branched RNAs of immediate practical interest, such as the proposed branched intermediate of Ty1 retrotransposition. Because this branched RNA has two strands with identical sequence that emerge from the branch site, we developed strategies to control which of the two strands bind with the deoxyribozyme during the branch-forming reaction. The ability to synthesize the proposed branched RNA of Ty1 retrotransposition will allow us to explore this important biochemical pathway in greater detail.
具有内部2',5'-分支的RNA分子是RNA剪接的中间体,并且分支RNA最近被认为是逆转录转座中间体。一条广泛适用的体外合成分支RNA的路线,该路线不需要自我剪接内含子或剪接体,将大大提高我们研究涉及分支RNA的生化过程的能力。我们最近描述了7S11,一种通过体外筛选鉴定的脱氧核酶,具有一般的RNA分支形成能力。然而,7S11的一个重要限制是分支位点的RNA核苷酸必须是嘌呤(A或G),因为嘧啶(U或C)不被接受。在这里,我们描述了紧凑的6CE8脱氧核酶(使用20个核苷酸的随机区域进行筛选),它能在分支位点合成带有任何核苷酸的2',5'-分支RNA。依赖于Mn2+的分支形成连接反应发生在一个RNA底物上的内部分支位点2'-羟基亲核试剂与另一个RNA底物上的5'-三磷酸之间。分支位点核苷酸的偏好顺序是U > C ≈ A > G,尽管所有四种核苷酸都能以有用的连接速率被接受。除了带有5'-三磷酸的RNA链的序列要求是5'-pppGA,5'-pppGAR(R = 嘌呤)更受青睐外,两个RNA底物中几乎所有其他位置的核苷酸都允许连接活性。这些特性使6CE8能够制备具有直接实际意义的分支RNA,例如所提出的Ty1逆转录转座的分支中间体。由于这种分支RNA有两条从分支位点出现的具有相同序列的链,我们开发了策略来控制在分支形成反应过程中两条链中的哪一条与脱氧核酶结合。合成所提出的Ty1逆转录转座分支RNA的能力将使我们能够更详细地探索这一重要的生化途径。