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This lower specificity indicates that the \u03b3-phosphate contributes primarily to nucleotide discrimination, rather than phosphodiester bond formation. Furthermore, our work shows that dNDPs and dNTPs with the same nucleobase exhibit comparable activation energies for the rate-limiting step during DNA synthesis, supporting a model in which the \u03b3-phosphate is not required for lowering the energetic barrier for TdT-catalyzed polymerization. These findings define a mechanistic framework for understanding non-templated DNA synthesis and its application in enzymatic DNA synthesis technologies.<\/jats:p>","DOI":"10.1093\/nar\/gkag980","type":"journal-article","created":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T12:01:06Z","timestamp":1790769666000},"source":"Crossref","is-referenced-by-count":0,"title":["Substrate energetics in non-templated DNA synthesis by terminal deoxynucleotidyl transferase"],"prefix":"10.1093","volume":"54","author":[{"given":"Katrina","family":"Mogannam","sequence":"first","affiliation":[{"name":"Department of Pharmaceutical Sciences, University of California , Irvine, CA 92697 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0632-5442","authenticated-orcid":false,"given":"Andrej","family":"Lupt\u00e1k","sequence":"additional","affiliation":[{"name":"Department of Pharmaceutical Sciences, University of California , 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