Center for Computational & Integrative Biology, Massachusetts General Hospital
Department of Molecular Biology, Massachusetts General Hospital
Department of Chemistry and Chemical Biology, Harvard University
Department of Genetics, Harvard Medical School
Howard Hughes Medical Institute


Prywes N, Michaels YS, Pal A, Oh SS, Szostak JW. Thiolated uridine substrates and templates improve the rate and fidelity of ribozyme-catalyzed RNA copying. Chem Commun (Camb). 2016 May. PDF.

Engelhart AE, Adamala KP, Szostak JW. A simple physical mechanism enables homeostasis in primitive cells. Nat Chem. 2016 May. PDF.

Adamala KP, Engelhart AE, Szostak JW. Collaboration between primitive cell membranes and soluble catalysts. Nat Commun. 2016 Mar. PDF.

Li L, Lelyveld VS, Prywes N, Szostak JW. Experimental and computational evidence for a loose transition state in phosphoroimidazolide hydrolysis. J Am Chem Soc. 2016 Mar. PDF.

Izgu EC, Oh SS, Szostak JW. Synthesis of activated 3'-amino-3'-deoxy-2-thio-thymidine, a superior substrate for the nonenzymatic copying of nucleic acid templates. Chem Commun (Camb). 2016 Feb. PDF.


Lelyveld VS, Björkbom A, Ransey EM, Sliz P, Szostak JW. Pinpointing RNA-protein cross-Links with site-specific stable isotope-labeled oligonucleotides. J Am Chem Soc. 2015 Dec PDF.

Björkbom A, Lelyveld VS, Zhang S, Zhang W, Tam CP, Blain JC, Szostak JW. Bidirectional direct sequencing of noncanonical RNA by two-dimensional analysis of mass chromatograms. J Am Chem Soc. 2015 Nov PDF.

Larsen AT, Fahrenbach AC, Sheng J, Pian J, Szostak JW. Thermodynamic insights into 2-thiouridine-enhanced RNA hybridization. Nucleic Acids Res. 2015 Sep PDF.

Kamat NP, Tobe S, Hill IT, Szostak JW. Electrostatic localization of RNA to protocell membranes by cationic hydrophobic peptides. Angew Chem Int Ed Engl. 2015 Sep PDF.

Adamala K, Engelhard AE, Kamat NP, Jin L, Szostak JW. Construction of a liposome dialyzer for the preparation of high-value, small-volume liposome formulations. Nat Protoc 2015 Jun. PDF.

Izgu EC, Fahrenbach AC, Zhang N, Li L, Zhang W, Larsen, AT, Blain, JC, Szostak JW. Uncovering the thermodynamics of monomer binding for RNA replication. J Am Chem Soc. 2015 Apr. PDF.

Heuberger BD, Pal A, Del Frate F, Topkar VV, Szostak JW. Replacing uridine with 2-thiouridine enhances the rate and fidelity of nonenzymatic RNA primer extension. J Am Chem Soc. 2015 Feb. PDF.

Adamala K, Engelhart AE, Szostak JW. Generation of functional RNAs from inactive oligonucleotide complexes by non-enzymatic primer extension. J Am Chem Soc. 2015 Jan. PDF.


Hentrich C, Szostak JW. Controlled growth of filamentous fatty acid vesicles under flow. Langmuir 2014 Dec. PDF.

Budin I, Prywes N, Zhang N, Szostak JW. Chain-length heterogeneity allows for the assembly of fatty acid vesicles in dilute solutions. Biophys J. 2014 Oct. PDF.

Sheng J, Larsen A, Heuberger BD, Blain JC, Szostak JW. Crystal structure studies of RNA duplexes containing s(2)U:A and s(2)U:U base pairs. J Am Chem Soc. 2014 Oct. PDF.

Blain JC, Szostak JW. Progress toward synthetic cells. Annu Rev Biochem. 2014 Mar. PDF.

Jia TZ, Hentrich C, Szostak JW. Rapid RNA exchange in aqueous two-phase system and coacervate droplets. Orig Life Evol Biosph. 2014 Feb. PDF.

Sheng J, Li L, Engelhart AE, Gan J, Wang J, Szostak JW. Structural insights into the effects of 2'-5' linkages on the RNA duplex. Proc Natl Acad Sci USA. 2014 Feb. PDF.

Li L, Szostak JW. The free energy landscape of pseudorotation in 3'-5' and 2'-5' linked nucleic acids. J Am Chem Soc. 2014 Feb. PDF.

Blain JC, Ricardo A, Szostak JW. Synthesis and nonenzymatic template-directed polymerization of 2'-amino-2'-deoxythreose nucleotides. J Am Chem Soc. 2014 Feb. PDF.


Adamala K, Szostak JW. Nonenzymatic template-Directed RNA synthesis inside model protocells. Science. 2013 Nov. PDF.

Zhu TF, Budin I, Szostak JW. Preparation of fatty acid micelles. Methods Enzymol. 2013 PDF.

Zhu TF, Budin I, Szostak JW. Vesicle extrusion through polycarbonate track-etched membranes using a hand-held mini-extruder. Methods Enzymol. 2013 PDF.

Zhu TF, Budin I, Szostak JW. Preparation of fatty acid or phospholipid vesicles by thin-film rehydration. Methods Enzymol. 2013 PDF.

Josephson K, Ricardo A, Szostak JW. mRNA display: from basic principles to macrocycle drug discovery. Drug Discov Today. 2013 Oct PDF.

Zhang S, Blain JC, Zielinska D, Gryaznov SM, Szostak JW. Fast and accurate nonenzymatic copying of an RNA-like synthetic genetic polymer. Proc Natl Acad Sci USA. 2013 Oct PDF.

Adamala K, Szostak JW. Competition between model protocells driven by an encapsulated catalyst. Nat Chem. 2013 Jun PDF.

Engelhart AE, Powner MW, Szostak JW. Functional RNAs exhibit tolerance for non-heritable 2'-5' versus 3'-5' backbone heterogeneity. Nat Chem. 2013 May. PDF.

Zhang S, Zhang N, Blain JC, Szostak JW. Synthesis of N3'-P5'-linked phosphoramidate DNA by nonenzymatic template-directed primer extension. J Am Chem Soc. 2013 Jan. PDF.


Budin I, Debnath A, Szostak JW. Concentration-driven growth of model protocell membranes. J Am Chem Soc. 2012 Dec. PDF.

Powner MW, Zheng SL, Szostak JW. Multicomponent assembly of proposed DNA precursors in water. J Am Chem Soc. 2012 Aug PDF.

Szostak JW. The eightfold path to non-enzymatic RNA replication. Journal of Systems Chemistry 2012, 3:2. PDF.

Zhu TF, Adamala K, Zhang N, Szostak JW. Photochemically driven redox chemistry induces protocell membrane pearling and division. Proc Natl Acad Sci U S A. 2012 Jun 4. PDF.

Hofmann FT, Szostak JW, Seebeck FP. In vitro selection of functional lantipeptides. J Am Chem Soc. 2012 May 16;134(19):8038-41. PDF.

Guillen Schlippe YV, Hartman MC, Josephson K, Szostak JW. In Vitro Selection of Highly Modified Cyclic Peptides That Act as Tight Binding Inhibitors. J Am Chem Soc. 2012 Mar 29. PDF.

Zhang N, Zhang S, Szostak JW. Activated ribonucleotides undergo a sugar pucker switch upon binding to a single-stranded RNA template. J Am Chem Soc. 2012 Feb 29;134(8):3691-4. PDF.

Szostak JW. Attempts to define life do not help to understand the origin of life. J Biomol Struct Dyn. 2012 Feb;29(4):599-600. PDF.


Szostak JW. An optimal degree of physical and chemical heterogeneity for the origin of life?. Philos Trans R Soc Lond B Biol Sci. 2011 Oct 27;366(1580):2894-901. Review. PDF.

Seebeck FP, Ricardo A, Szostak JW. Artificial lantipeptides from in vitro translations. Chem Commun (Camb). 2011 Jun 7;47(21):6141-3. PDF.

Zhu TF, Szostak JW. Exploding vesicles. Journal of Systems Chemistry 2011, 2:4. PDF.

Trevino SG, Zhang N, Elenko MP, Luptak A, Szostak JW. Evolution of functional nucleic acids in the presence of nonheritable backbone heterogeneity. Proc Natl Acad Sci USA. 2011 Aug 11. [Epud ahead of print]. PDF. Supporting Information.

Budin I, Szostak JW. Physical effects underlying the transition from primitive to modern cell membranes. Proc Natl Acad Sci USA. 2011 Mar 14;108:5249-5254. PDF. Supporting Information.

Subtelny AO, Hartman MCT, Szostak JW. Optimal codon choice can improve the efficiency and fidelity of N-methyl amino acid incorporation into peptides by in vitro translation. Angew Chem Int Ed. 2011 Mar 4;50:3164-3167. PDF.


Powner MW, Sutherland JD, Szostak JW. Chemoselective multicomponent one-pot assembly of purine precursors in water. J Am Chem Soc. 2010 Nov 2;132:16677-16688. PDF.

Elenko MP, Szostak JW, van Oijen AM. Single-molecule binding experiments on long time scales. Rev Sci Instrum. 2010 Aug 27;81:083705. PDF.

Schrum JS, Zhu TF, Szostak JW. The origins of cellular life. Cold Spring Harb Perspect Biol. 2010 May 19. [Epud ahead of print]. PDF.

Rajamani S, Ichida JK, Antal T, Treco DA, Leu K, Nowak MA, Szostak JW, Chen IA. Effect of Stalling after Mismatches on the Error Catastrophe in Nonenzymatic Nucleic Acid Replication. J Am Chem Soc. 2010 Apr 28;132(16):5880-5. PDF. Supporting Information.

Budin I, Szostak JW. Expanding Roles for Diverse Physical Phenomena During the Origin of Life. Annu Rev Biophys. 2010 Jun 9;39:245-63. PDF.


Bruckner RJ, Mansy SS, Ricardo A, Mahadevan L, Szostak JW. Flip-flop-induced relaxation of bending energy: implications for membrane remodeling. Biophys J. 2009 Dec 16;97(12):3113-22. PDF. Supporting Information.

Schrum JP, Ricardo A, Krishnamurthy M, Blain JC, Szostak JW. Efficient and Rapid Template-Directed Nucleic Acid Copying Using 2'-Amino-2',3'-dideoxyribonucleoside-5'-Phosphorimidazolide Monomers. J Am Chem Soc. 2009 Oct 14;131(40):14560-70. PDF. Supporting Information.

Mansy SS, Szostak JW. Reconstructing the Emergence of Cellular Life through the Synthesis of Model Protocells. Cold Spring Harb Symp Quant Biol. 2009 Sep 4;74:47-54. PDF

Ricardo A, Szostak JW. Origin of life on earth. Sci Am. 2009 Sep;301(3):54-61.

Lin S, Horning DP, Szostak JW, Chaput JC. Conformational analysis of DNA repair intermediates by time-resolved fluorescence spectroscopy. J Phys Chem A. 2009 Sep 3;113(35):9585-7. PDF. Supporting Information.

Budin I, Bruckner RJ, Szostak JW. Formation of protocell-like vesicles in a thermal diffusion column. J Am Chem Soc. 2009 Jul 22;131(28):9628-9. PDF. Supporting Information. Supporting Information (AVI VIDEO).

Elenko MP, Szostak JW, van Oijen AM. Single-molecule imaging of an in vitro-evolved RNA aptamer reveals homogeneous ligand binding kinetics. J Am Chem Soc. 2009 Jul 29;131(29):9866-7. PDF. Supporting Information. Supporting Information (AVI VIDEO).

Szostak JW. Origins of life: Systems chemistry on early Earth. Nature. 2009 May 14;459(7244):171-2. PDF

Zhu TF, Szostak JW. Preparation of large monodisperse vesicles. PLoS ONE. 2009;4(4):e5009. Epub 2009 Apr 6. PDF

Zhu TF, Szostak JW. Coupled Growth and Division of Model Protocell Membranes. J Am Chem Soc. 2009 Mar 26. [Epub ahead of print]. PDF. Supporting Information. Supporting Information (Videos).

Chen JJ, Tsai CH, Cai X, Horhota AT, McLaughlin LW, Szostak JW. Enzymatic primer-extension with glycerol-nucleoside triphosphates on DNA templates. PLoS ONE. 2009;4(3):e4949. Epub 2009 Mar 23. PDF

Chen JJ, Cai X, Szostak JW. N2'-->p3' phosphoramidate glycerol nucleic acid as a potential alternative genetic system. J Am Chem Soc. 2009 Feb 18;131(6):2119-21. PDF. Supporting Information.


Mansy SS, Szostak JW. Thermostability of model protocell membranes. Proc Natl Acad Sci U S A. 2008 Sep 9;105(36):13351-5. Epub 2008 Sep 3. PDF

Mansy SS, Schrum JP, Krishnamurthy M, Tobé S, Treco DA, Szostak JW. Template-directed synthesis of a genetic polymer in a model protocell. Nature. Published online 4 June 2008. PDF. Supporting Information.

Subtelny AO, Hartman MC, Szostak JW. Ribosomal synthesis of N-methyl peptides. J Am Chem Soc. 2008 May 14;130(19):6131-6. Epub 2008 Apr 11. PDF

Monnard PA, Szostak JW. Metal-ion catalyzed polymerization in the eutectic phase in water-ice: A possible approach to template-directed RNA polymerization. J Inorg Biochem. 2008 May;102(5-6):1104-11. Epub 2008 Feb 1.


Hartman MC, Josephson K, Lin CW, Szostak JW. An expanded set of amino Acid analogs for the ribosomal translation of unnatural peptides. PLoS ONE. 2007 Oct 3;2(10):e972. PDF

Tsai CH, Chen J, Szostak JW. Enzymatic synthesis of DNA on glycerol nucleic acid templates without stable duplex formation between product and template. Proc Natl Acad Sci U S A. 2007 Sep 11; 104(37): 14598-603. PDF Supporting information

Seelig B, Szostak JW. Selection and evolution of enzymes from a partially randomized non-catalytic scaffold. Nature. 2007 Aug 16;448(7155):828-31. PDF Supporting information

Mansy SS, Zhang J, Kummerle R, Nilsson M, Chou JJ, Szostak JW, Chaput JC. Structure and Evolutionary Analysis of a Non-biological ATP-binding Protein. J Mol Biol. 2007 Aug 10;371(2):501-13. PDF

Smith MD, Rosenow MA, Wang M, Allen JP, Szostak JW, Chaput JC. Structural insights into the evolution of a non-biological protein: importance of surface residues in protein fold optimization. PLoS ONE. 2007 May 23;2:e467. PDF

Hazen RM, Griffin PL, Carothers JM, Szostak JW. Functional information and the emergence of biocomplexity. Proc Natl Acad Sci U S A. 2007 May 15;104 Suppl 1:8574-81. Epub 2007 May 9. PDF

Meena, Sam M, Pierce K, Szostak JW, McLaughlin LW.  2',3'-Dideoxy-3'-thionucleoside Triphosphates: Syntheses and Polymerase Substrate Activities.  Org Lett. 2007 Mar 15;9(6):1161-3. PDF   Supporting Information     Supporting Information  


Horhota AT, Szostak JW, McLaughlin LW.  Glycerol nucleoside triphosphates: synthesis and polymerase substrate activities.  Org Lett. 2006 Nov 9;8(23):5345-7. PDF   Supporting Information  

Blackburn EH, Greider CW, Szostak JW.  Telomeres and telomerase: the path from maize, Tetrahymena and yeast to human cancer and aging.  Nat Med. 2006 Oct;12(10):1133-8. PDF  

Salehi-Ashtiani K, Luptak A, Litovchick A, Szostak JW.  A genomewide search for ribozymes reveals an HDV-like sequence in the human CPEB3 gene.  Science. 2006 Sep 22;313(5794):1788-92. Article Online   Supporting Information  

Hanczyc MM, Mansy SS, Szostak JW.  Mineral Surface Directed Membrane Assembly.  Orig Life Evol Biosph. 2007 Feb;37(1):67-82. PDF  

Carothers JM, Oestreich SC, Szostak JW.  Aptamers selected for higher-affinity binding are not more specific for the target ligand.  J Am Chem Soc. 2006 Jun 21;128(24):7929-37. PDF   Supporting Information  

Seebeck FP, Szostak JW.  Ribosomal synthesis of dehydroalanine-containing peptides.  J Am Chem Soc. 2006 Jun 7;128(22):7150-1. PDF   Supporting Information  

Hartman MC, Josephson K, Szostak JW.  Enzymatic aminoacylation of tRNA with unnatural amino acids.  Proc Natl Acad Sci U S A. 2006 Mar 21;103(12):4356-61. PDF   Supporting Information  

Carothers JM, Davis JH, Chou JJ, Szostak JW.  Solution structure of an informationally complex high-affinity RNA aptamer to GTP.  RNA. 2006 Apr;12(4):567-79. PDF  

Cho GS, Szostak JW.  Directed evolution of ATP binding proteins from a zinc finger domain by using mRNA display.  Chem Biol. 2006 Feb;13(2):139-47. PDF   Supporting Information  


Fujikawa SM, Chen IA, Szostak JW.  Shrink-wrap vesicles.  Langmuir. 2005 Dec 20;21(26):12124-9. PDF   Supporting Information  

Plummer KA, Carothers JM, Yoshimura M, Szostak JW, Verdine GL.  In vitro selection of RNA aptamers against a composite small molecule-protein surface.  Nucleic Acids Res. 2005 Sep 30;33(17):5602-10. PDF   Supporting Information  

Chen IA, Salehi-Ashtiani K, Szostak JW.  RNA catalysis in model protocell vesicles.  J Am Chem Soc. 2005 Sep 28;127(38):13213-9. PDF   Supporting Information  

Ichida JK, Horhota A, Zou K, McLaughlin LW, Szostak JW.  High fidelity TNA synthesis by Therminator polymerase.  Nucleic Acids Res. 2005 Sep 12;33(16):5219-25. PDF  

Josephson K, Hartman MC, Szostak JW.  Ribosomal synthesis of unnatural peptides.  J Am Chem Soc. 2005 Aug 24;127(33):11727-35. PDF   Supporting Information  

Horhota A, Zou K, Ichida JK, Yu B, McLaughlin LW, Szostak JW, Chaput JC.  Kinetic analysis of an efficient DNA-dependent TNA polymerase.  J Am Chem Soc. 2005 May 25;127(20):7427-34. PDF   Supporting Information  

Shen X, Valencia CA, Szostak JW, Dong B, Liu R.  Scanning the human proteome for calmodulin-binding proteins.  Proc Natl Acad Sci U S A. 2005 Apr 26;102(17):5969-74. PDF   Supporting Information  

Sacerdote MG, Szostak JW.  Semipermeable lipid bilayers exhibit diastereoselectivity favoring ribose.  Proc Natl Acad Sci U S A. 2005 Apr 26;102(17):6004-8. PDF   Supporting Information  

Zou K, Horhota A, Yu B, Szostak JW, McLaughlin LW.  Synthesis of alpha-L-threofuranosyl nucleoside triphosphates (tNTPs).  Org Lett. 2005 Apr 14;7(8):1485-7. PDF   Supporting Information     Supporting Information  

Ichida JK, Zou K, Horhota A, Yu B, McLaughlin LW, Szostak JW.  An in vitro selection system for TNA.  J Am Chem Soc. 2005 Mar 9;127(9):2802-3. PDF   Supporting Information


Hanczyc MM, Szostak JW.  Replicating vesicles as models of primitive cell growth and division.  Curr Opin Chem Biol. 2004 Dec;8(6):660-4. PDF  

Chen, IA, Roberts, RW, Szostak, JW. The emergence of competition between model protocells.  Science 2004 Sep 3;305:1474-76. PDF   Supporting Information  

Chen IA, Szostak JW. A kinetic study of the growth of fatty acid vesicles. Biophys J. 2004 Aug;87(2):988-98. PDF

Sazani PL, Larralde R, Szostak JW. A small aptamer with strong and specific recognition of the triphosphate of ATP. J Am Chem Soc. 2004 Jul 14;126(27):8370-1. PDF Supporting Information

Chaput JC, Szostak JW. Evolutionary optimization of a nonbiological ATP binding protein for improved folding stability. Chem Biol. 2004 Jun;11(6):865-74. PDF

Chen IA, Szostak JW. Membrane growth can generate a transmembrane pH gradient in fatty acid vesicles. Proc Natl Acad Sci U S A. 2004 May 25;101(21):7965-70. PDF

Carothers JM, Oestreich SC, Davis JH, Szostak JW. Informational complexity and functional activity of RNA structures.  J Am Chem Soc. 2004 Apr 28;126(16):5130-7.PDF   Supporting Information   Extra Materials


Hanczyc, MM, Fujikawa, SM, Szostak, JW. Experimental models of primitive cellular compartments: encapsulation, growth, and division. Science. 2003 Oct 24; 302: 618-622. PDF

Chaput, JC, Szostak, JW. TNA synthesis by DNA polymerases. J Am Chem Soc. 2003 Aug 6;125(31):9274-5. PDF

Vaish, NK, Larralde, R. Fraley, AW, Szostak, JW, McLaughlin, LW. A novel, modification-dependent ATP-binding aptamer selected from an RNA library incorporating a cationic functionality. Biochemistry. 2003 Jul 29;42(29):8842-51. PDF

Szostak, JW. Functional information: molecular messages. Nature. 2003 June 12; 423: 689. PDF

Huang, Z, and Szostak, JW. Selective labeling and detection of specific RNAs in an RNA mixture. Anal Biochem. 2003 Apr 1;315(1):129-33. PDF

Chaput, JC, Ichida, JK, Szostak, JW. DNA polymerase-mediated DNA synthesis on a TNA template. J Am Chem Soc. 2003 Jan 29;125(4):856-7. PDF


Davis, JH, Szostak, JW. Isolation of high-affinity GTP aptamers from partially structured RNA libraries. Proc Natl Acad Sci U S A. 2002 Sep 3; 99(18):11616-11621. PDF


Salehi-Ashtiani K, Szostak JW. In vitro evolution suggests multiple origins for the hammerhead ribozyme. Nature. 2001 Nov 1;414(6859):82-4. PD

Szostak JW, Bartel, DP and Luisi, PL. Synthesizing Life. Nature. 2001; 409: 387-390. PDF

Keefe AD, Szostak JW. Functional proteins from a random-sequence library. Nature. 2001 Apr 5;410(6829):715-8. PDF

Keefe AD, Wilson, DS, Seelig, B, and Szostak JW. One-step purification of recombinant proteins using a nanomolar-affinity streptavidin-binding peptide, the SBP-Tag. Protein Expression and Purification. 2001; 28 : 440-446. PDF

Wilson DS, Keefe AD, Szostak JW.The use of mRNA display to select high-affinity protein-binding peptides. Proc Natl Acad Sci U S A. 2001 Mar 27;98(7):3750-5. PDF


Cho, G, Keefe, AD, Wilson, DS, Liu, R and Szostak, JW. Constructing high complexity synthetic libraries of long ORFs using in vitro selection. J.Mol.Biol. 2000; 297:309-319 PDF supplementary information.

Vaish NK, Fraley AW, Szostak JW, McLaughlin LW. Expanding the structural and functional diversity of RNA: analog uridine triphosphates as candidates for in vitro selection of nucleic acids. Nucleic Acids Res. 2000 Sep 1;28(17):3316-22. PDF

Lee N, Bessho Y, Wei K, Szostak JW, Suga H. Ribozyme-catalyzed tRNA aminoacylation. Nat Struct Biol. 2000 Jan;7(1):28-33. PDF

Burgstaller P, Hale S, Sun L, Roberts RW, Szostak JW, Wagner R. Identification of epitope-like consensus motifs using mRNA display. J Mol Recognit. 2002 May-Jun;15(3):126-34.

Szostak JW. Constraints on the sizes of the earliest cells. In, 'Size Limits of Very Small Microorganisms', Proceedings of a Workshop of the Space Studies Board, National Research Council. 1999, National Academy Press.

Liu R, Barrick JE, Szostak JW, Roberts RW. Optimized synthesis of RNA-protein fusions for in vitro protein selection. Methods Enzymol. 2000;318:268-93.


Wilson DW, Szostak JW. In vitro Selection of Functional Nucleic Acids. Ann. Rev. Biochem., 1999; 68:611-648. PDF


Szostak JW. In Vitro Selection and Directed Evolution. 1998 Harvey Lecture Series. 1997-98;93:95-118. Review.

Holeman LA, Robinson SL, Szostak JW and Wilson C. Isolation and characterization of fluorophore-binding RNA aptamers. Folding & Design, 1998; 3:423-431.

Wilson C, Szostak JW. Isolation of a fluorophore-specific DNA aptamer with weak redox activity. Chem. Biol., 1998; 5: 609-617.

Wilson C, Nix J, Szostak JW. Functional requirements for specific ligand recognition by a biotin-binding RNA pseudoknot. Biochemistry 1998; 37:14410-9. PDF

Suga H, Cowan JA, Szostak JW. Unusual metal ion catalysis in an acyl transferase ribozyme. Biochemistry 1998; 37:10118-10125. PDF

Suga H, Lohse P, Szostak JW. Structural and Kinetic Characterization of an Acyl Transferase Ribozyme. J. Am. Chem. Soc. 1998; 120: 1151-1156. PDF


Roberts RW, Szostak JW. RNA-peptide fusions for the in Vitro selection of peptides and proteins. Proc. Natl. Acad. Sci. USA 1997; 94: 12297-12302. PDF

Hager AJ, Szostak JW. Isolation of Novel Ribozymes that Ligate AMP-Activated RNA Substrates. Chem. Biol. 1997; 4:607-617.

Dieckmann T, Butcher SE, Sassanfar M, Szostak JW and Feigon J. Mutant ATP-Binding RNA aptamers reveal the structural basis for ligand binding. J. Mol. Biol. 1997; 273: 467- 478. PDF

Szostak JW. Introduction: Combinatorial Chemistry. Chem. Rev. 1997; 97 (2): 347-348, in: Chemical Reviews: Combinatorial Chemistry, a Thematic Issue, Guest Editor Jack W. Szostak.


Hager AJ, Pollard JD, Szostak JW. Ribozymes: aiming at RNA replication and translation. Chem. Biol. 1996; 3:717-725.

Huang Z, Szostak JW. 3' End-Labeling of RNA with DNA Polymerase. Nuc. Acids Res. 1996; 24: 4360-4361. PDF

Lohse P, Szostak JW. Ribozyme-catalysed amino acid transfer reactions. Nature 1996; 381: 442-444.

Rohatgi R, Bartel D, Szostak JW. Non-enzymatic, Template-Directed Ligation of Oligoribonucleotides is Highly Regioselective for the Formation of 3'-5' Phosphodiester Bonds. J. Am. Chem. Soc. 1996; 118: 3340-3334. PDF

Rohatgi R, Bartel D, Szostak JW. Kinetic and Mechanistic Analysis of Nonenzymatic, Template-Directed Oligonucleotide Ligation. J. Am. Chem. Soc. 1996; 118: 3332-3339 PDF

Lorsch JR, Szostak JW. Chance and Necessity in the Selection of Nucleic Acid Catalysts. Accounts Chem. Res. 1996; 29: 103-110. PDF

Lorsch JR, Szostak JW. In Vitro Selection of Nucleic Acid Sequences that Bind Small Molecules. 1996. Chpt. 4 in Combinatorial Libraries: Synthesis, Screening and Application Potential. Ed. R. Cortese. Publ. Walter de Gruyter, Berlin, New York.


Lorsch JR, Bartel DP, Szostak JW. Reverse transcriptase reads through a 2'-5' linkage and a 2'-thiophosphate in a template. Nuc. Acids Res. 1995; 23: 2811-2814.

Ekland EH, Szostak JW, Bartel DP. Structurally complex and highly active RNA ligases derived from random RNA sequences. Science 1995; 269: 364-370. PDF

Cuenod B, Szostak JW. A DNA metalloenzyme with DNA ligase activity. Nature 1995; 375: 611-614.

Chapman KB, Szostak JW. Isolation of a ribozyme with 5'-5' ligase activity. Chemistry and Biology 1995; 2: 325-333.

Wilson CW, Szostak JW. In vitro evolution of a self-alkylating ribozyme. Nature 1995; 374: 777-782.

Whoriskey S, Usman N, Szostak JW. Total chemical synthesis of a ribozyme derived from a group I intron. Proc. Natl. Acad. Sci. U.S.A. 1995; 92: 2465-2469.

Lauhon CT, Szostak JW. RNA Aptamers that bind Flavin and Nicotinamide Redox Cofactors. J. Am. Chem. Soc. 1995; 117: 1246-1257. PDF

Huizenga DE, Szostak JW. DNA Aptamer that Binds Adenosine and ATP. Biochemistry 1995; 34: 656-665.


Bartel DP, Szostak JW. Study of RNA-protein recognition by in vitro selection. in RNA- Protein Interactions, eds. K. Nagai and I.W. Mattaj, Oxford University Press 1994.

Hanna M, Szostak JW. Suppression of mutations in the core of the Tetrahymena ribozyme by spermidine, ethanol and by substrate stabilization. Nuc. Acids Res. 1994; 22: 5326-5331.

Lorsch JR, Szostak JW. In vitro evolution of new ribozymes with polynucleotide kinase activity. Nature 1994; 371:31-36.

Chapman KB, Szostak JW. In Vitro Selection of Catalytic RNAs. Current Opinions in Structural Biology 1994; 4:618-622.

Green R, Szostak JW. Template-directed oligonucleotide assembly reactions catalyzed by a minimally structured ribozyme derived from a group I intron. pp. 277-283 in, Structural Biology: The State of the Art, Proceedings of the Eighth Conversation, State University of New York, Albany, NY 1993 Eds. Ramaswamy H. Sarma and Mukti H. Sarma, Adenine Press 1994

Peterson RD, Bartel DP, Szostak JW, Horvath SJ, Feigon J. H NMR Studies of the High- Affinity Rev Binding Site of the Rev Responsive Element of HIV-1 mRNA: Base Pairing in the Core Binding Element. Biochemistry 1994; 33: 5357-5366.

Lorsch J, Szostak, JW. In Vitro Selection of RNA Aptamers Specific for Cyanocobalamin. Biochemistry 1994; 33:973-982.

Lin, C-W, Szostak JW. Evidence That the Guanosine Substrate of the Tetrahymena Ribozyme is Bound in the Anti Conformation and That N7 Contributes to Binding. Biochemistry 1994; 33:2703-2707.

Green R, Szostak JW. In vitro genetic analysis of the hinge region between helical elements P5-P4-P6 and P7-P3-P8 in the sunY group I self-splicing intron. J. Mol. Biol. 1994; 235:140-155.


Bartel DP, Szostak JW. Isolation of a new ribozymes from a large pool of random sequences. Science 1993; 261:1411-1418. PDF

Sassanfar M, Szostak JW. An RNA motif that binds ATP. Nature 1993; 364: 550-553.

Famulok M, Szostak JW. Selection of Functional RNA and DNA Molecules from Randomized Sequences. pp. 271-284, in Nucleic Acids and Molecular Biology, ed. F. Eckstein and D.M.J. Lilley, Springer-Verlag, 1993.

Szostak, J.W. and Ellington, A.E. In Vitro Selection of Functional RNA Sequences. pp. 511-533 in "The RNA World", ed. R. Gesteland and J. Atkins; Cold Spring Harbor Press, 1993.

Doudna JA, Szostak JW. Triplet extension: a novel strategy for RNA catalysed RNA replication. Biochemistry 1993; 32:2111-2115.


Green R and Szostak JW. Selection of a Ribozyme that Functions as a Superior Template in a Self-Copying Reaction. Science 1992; 258:1910-1915. PDF

Wilson CL, Szostak JW. Ribozyme Catalysis. Curr. Opin. in Struct. Biol. 1992; 2:749-756.

Famulok M, Szostak JW. In Vitro Selection of Specific Ligand Binding Nucleic Acids. Angew. Chem. Int. Ed. Engl. 1992; 31: 979-988.

Famulok M, Szostak JW. Stereospecific Recognition of Tryptophan Agarose by in Vitro Selected RNA. J. Am. Chem. Soc. 1992; 114: 3990-3991.

Szostak JW. In Vitro Genetics. Trends in Biochem. Sci. 1992; 17: 89-93.

Ellington AE, Szostak JW. Selection in vitro of single-stranded DNA molecules that fold into specific ligand binding structures. Nature 1992; 355:850-852.


Green R, Szostak JW, Benner SA, Rich A, Usman N. Synthesis of RNA containing inosine: analysis of the sequence requirements for the 5' splice-site of the Tetrahymena group I intron.Nuc. Acids Res., 1991; 19: 4161-4166.

Bartel DP, Zapp M, Green M, Szostak JW. HIV-1 Rev Regulation Involves Recognition of Non-Watson-Crick Base-pairs in Viral RNA . Cell, 1991; 67: 529-536.

Green R, Ellington AD, Bartel DP, Szostak JW. In Vitro Genetic Analysis: Selection and Amplification of Rare Functional Nucleic Acids. Methods: A Companion to Methods in Enzymol., 1991; 2: 75-86.

Bartel DP, Szostak JW. Template-Directed Primer Extension Catalyzed by the Tetrahymena Ribozyme. Mol. Cell. Biol. 1991; 11: 3390-3394.

Doudna JA, Couture S, Szostak JW. A multi-subunit ribozyme that is a catalyst of and a template for complementary strand RNA synthesis. Science 1991; 251: 1605-1608.


Doudna JA, Szostak JW, Rich A, Usman N. Chemical Synthesis of Oligoribonucleotides Containing 2-Aminopurine: Substrates for the Investigation of Ribozyme Function. J. Org. Chem. 1990; 55: 5547-5549.

Couture S, Ellington AD, Gerber AS, Cherry JM, Doudna JA, Green R, Hanna M, Pace U, Rajogopal J, Szostak JW. Phylogenetic and genetic evidence for base-triples in the catalytic domain of group I introns. Nature, 1990; 347: 578-580.

Ellington AE, Szostak JW. In vitro Selection of RNA Molecules that Bind Specific Ligands. Nature, 1990; 346: 818-822.


Michel F, Hanna M, Green R, Bartel DB, Szostak JW. Miniribozymes, Small Derivatives of the sunY Intron, Are Catalytically Active. Mol. Cell. Biol. 1989; 9: 5480-5483.

Doudna JA, Szostak JW. Stereochemical Course of Catalysis by the Tetrahymena Ribozyme. Science, 1989; 244: 692-694.


Doudna JA, Cormack BP, Szostak JW. Genetic Dissection of an RNA Enzyme. Cold Spring Harbor Symposia, Vol. LII. 1987.

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