Geeta Narlikar, PhD
Professor
Biochemistry and Biophysics
School of Medicine

Education & Training

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• Diversity, Equity, and Inclusion Champion Training University of California 2021

Websites

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• Profile at biochemistry.ucsf.edu (biochemistry.ucsf.edu)
• Narlikar lab website (narlikarlab.ucsf.edu)

Grants and Projects

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• Tetrad: Genetics, Cell Biology, Biochemistry and Molecular Biology Training Grant, NIH, 2021-2026
• Modulation and functional characterization of protein condensation in chromatin organization, NIH, 2020-2025
• ATP-dependent and independent mechanisms of regulating chromatin states, NIH, 2018-2023
• Cell Biology, Genetics, and Biochemistry Training Grant, NIH, 1979-2021
• Mechanistic Analysis of Chromatin Remodeling, NIH, 2005-2018
• Mechanistic dissection of HP1 mediated heterochromatin, NIH, 2014-2018
• Bio-Organic Biomedical Mass Spectrometry Resource, NIH, 1982-2015

Publications (103)

Top publication keywords:
SchizosaccharomycesChromatin Assembly and DisassemblyChromatinHeterochromatinAdenosine TriphosphatasesRNA, CatalyticNucleosomesAdenosine TriphosphateHistonesChromosomal Proteins, Non-HistoneFluorescence Resonance Energy TransferDNASchizosaccharomyces pombe ProteinsNucleic Acid ConformationSaccharomyces cerevisiae Proteins

• Functionalized graphene-oxide grids enable high-resolution cryo-EM structures of the SNF2h-nucleosome complex without crosslinking.
  Nature communications 2024 Chio US, Palovcak E, Smith AAA, Autzen H, Muñoz EN, Yu Z, Wang F, Agard DA, Armache JP, Narlikar GJ, Cheng Y
• Hexasomal particles: consequence or also consequential?
  Current opinion in genetics & development 2024 Kaur U, Muñoz EN, Narlikar GJ
• Understanding how genetically encoded tags affect phase separation by Heterochromatin Protein HP1α.
  bioRxiv : the preprint server for biology 2023 Zhou ZK, Narlikar GJ
• Nucleosome density shapes kilobase-scale regulation by a mammalian chromatin remodeler.
  Nature structural & molecular biology 2023 Abdulhay NJ, Hsieh LJ, McNally CP, Ostrowski MS, Moore CM, Ketavarapu M, Kasinathan S, Nanda AS, Wu K, Chio US, Zhou Z, Goodarzi H, Narlikar GJ, Ramani V
• Catalytic and non-catalytic mechanisms of histone H4 lysine 20 methyltransferase SUV420H1.
  Molecular cell 2023 Abini-Agbomson S, Gretarsson K, Shih RM, Hsieh L, Lou T, De Ioannes P, Vasilyev N, Lee R, Wang M, Simon MD, Armache JP, Nudler E, Narlikar G, Liu S, Lu C, Armache KJ

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• Reorientation of INO80 on hexasomes reveals basis for mechanistic versatility.
  Science (New York, N.Y.) 2023 Wu H, Muñoz EN, Hsieh LJ, Chio US, Gourdet MA, Narlikar GJ, Cheng Y
• Functionalized graphene-oxide grids enable high-resolution cryo-EM structures of the SNF2h-nucleosome complex without crosslinking.
  bioRxiv : the preprint server for biology 2023 Chio US, Palovcak E, Autzen AAA, Autzen HE, Muñoz EN, Yu Z, Wang F, Agard DA, Armache JP, Narlikar GJ, Cheng Y
• In diverse conditions, intrinsic chromatin condensates have liquid-like material properties.
  Proceedings of the National Academy of Sciences of the United States of America 2023 Gibson BA, Blaukopf C, Lou T, Chen L, Doolittle LK, Finkelstein I, Narlikar GJ, Gerlich DW, Rosen MK
• Catalytic and non-catalytic mechanisms of histone H4 lysine 20 methyltransferase SUV420H1.
  bioRxiv : the preprint server for biology 2023 Abini-Agbomson S, Gretarsson K, Shih RM, Hsieh L, Lou T, De Ioannes P, Vasilyev N, Lee R, Wang M, Simon M, Armache JP, Nudler E, Narlikar G, Liu S, Lu C, Armache KJ
• Satellite repeat transcripts modulate heterochromatin condensates and safeguard chromosome stability in mouse embryonic stem cells.
  Nature communications 2022 Novo CL, Wong EV, Hockings C, Poudel C, Sheekey E, Wiese M, Okkenhaug H, Boulton SJ, Basu S, Walker S, Kaminski Schierle GS, Narlikar GJ, Rugg-Gunn PJ
• ATP Hydrolysis Coordinates the Activities of Two Motors in a Dimeric Chromatin Remodeling Enzyme.
  Journal of molecular biology 2022 Johnson SL, Narlikar GJ
• A hexasome is the preferred substrate for the INO80 chromatin remodeling complex, allowing versatility of function.
  Molecular cell 2022 Hsieh LJ, Gourdet MA, Moore CM, Muñoz EN, Gamarra N, Ramani V, Narlikar GJ
• A hexasome is the preferred substrate of the INO80 chromatin remodeling complex.
  Biophysical Journal 2022 Geeta Narlikar, Laura Hsieh, Muryam Gourdet, Vijay Ramani
• Histone dynamics play a critical role in SNF2h-mediated nucleosome sliding.
  Nature structural & molecular biology 2021 Gamarra N, Narlikar GJ
• Generation and Biochemical Characterization of Phase-Separated Droplets Formed by Nucleic Acid Binding Proteins: Using HP1 as a Model System.
  Current protocols 2021 Sanulli S, Narlikar GJ
• Is transcriptional regulation just going through a phase?
  Molecular cell 2021 Narlikar GJ, Myong S, Larson D, Maeshima K, Francis N, Rippe K, Sabari B, Strader L, Tjian R
• HP1 proteins compact DNA into mechanically and positionally stable phase separated domains.
  eLife 2021 Keenen MM, Brown D, Brennan LD, Renger R, Khoo H, Carlson CR, Huang B, Grill SW, Narlikar GJ, Redding S
• Collaboration through chromatin: motors of transcription and chromatin structure.
  Journal of molecular biology 2021 Gamarra N, Narlikar GJ
• Massively multiplex single-molecule oligonucleosome footprinting.
  eLife 2020 Abdulhay NJ, McNally CP, Hsieh LJ, Kasinathan S, Keith A, Estes LS, Karimzadeh M, Underwood JG, Goodarzi H, Narlikar GJ, Ramani V
• Enzymatic Reactions inside Biological Condensates.
  Journal of molecular biology 2020 Zhang Y, Narlikar GJ, Kutateladze TG
• Biophysical Properties of HP1-Mediated Heterochromatin.
  Cold Spring Harbor symposia on quantitative biology 2020 Sanulli S, Gross JD, Narlikar GJ
• ATP Hydrolysis by the SNF2 Domain of Dnmt5 Is Coupled to Both Specific Recognition and Modification of Hemimethylated DNA.
  Molecular cell 2020 Dumesic PA, Stoddard CI, Catania S, Narlikar GJ, Madhani HD
• Zscan4 binds nucleosomal microsatellite DNA and protects mouse two-cell embryos from DNA damage.
  Science advances 2020 Srinivasan R, Nady N, Arora N, Hsieh LJ, Swigut T, Narlikar GJ, Wossidlo M, Wysocka J
• Evolutionary Persistence of DNA Methylation for Millions of Years after Ancient Loss of a De Novo Methyltransferase.
  Cell 2020 Catania S, Dumesic PA, Pimentel H, Nasif A, Stoddard CI, Burke JE, Diedrich JK, Cooke S, Shea T, Gienger E, Lintner R, Yates JR, Hajkova P, Narlikar GJ, Cuomo CA, Pritchard JK, Madhani HD
• Evolutionary Persistence of DNA Methylation for Millions of Years after Ancient Loss of a De Novo Methyltransferase.
  Cell 2020 Catania S, Dumesic PA, Pimentel H, Nasif A, Stoddard CI, Burke JE, Diedrich JK, Cook S, Shea T, Geinger E, Lintner R, Yates JR, Hajkova P, Narlikar GJ, Cuomo CA, Pritchard JK, Madhani HD
• Phase-separation in chromatin organization.
  Journal of biosciences 2020 Narlikar GJ
• Topical collection on Chromatin Biology and Epigenetics.
  Journal of biosciences 2020 Narlikar G, Galande S
• HP1 reshapes nucleosome core to promote phase separation of heterochromatin.
  Nature 2019 Sanulli S, Trnka MJ, Dharmarajan V, Tibble RW, Pascal BD, Burlingame AL, Griffin PR, Gross JD, Narlikar GJ
• Chromatin topology, condensates and gene regulation: shifting paradigms or just a phase?
  Development (Cambridge, England) 2019 Mir M, Bickmore W, Furlong EEM, Narlikar G
• Cryo-EM structures of remodeler-nucleosome intermediates suggest allosteric control through the nucleosome.
  eLife 2019 Armache JP, Gamarra N, Johnson SL, Leonard JD, Wu S, Narlikar GJ, Cheng Y
• Ion counting demonstrates a high electrostatic field generated by the nucleosome.
  eLife 2019 Gebala M, Johnson SL, Narlikar GJ, Herschlag D
• The Role of Phase-Separation in Heterochromatin.
  Biophysical Journal 2019 Geeta Narlikar, Serena Sanulli, John D. Gross, Patrick Griffin, Mike Trnka
• A Nucleosome Bridging Mechanism for Activation of a Maintenance DNA Methyltransferase.
  Molecular cell 2018 Stoddard CI, Feng S, Campbell MG, Liu W, Wang H, Zhong X, Bernatavichute Y, Cheng Y, Jacobsen SE, Narlikar GJ
• Visualization and Quantitation of Phase-Separated Droplet Formation by Human HP1α.
  Methods in enzymology 2018 Keenen MM, Larson AG, Narlikar GJ
• The Role of Phase Separation in Heterochromatin Formation, Function, and Regulation.
  Biochemistry 2018 Larson AG, Narlikar GJ
• The nucleosomal acidic patch relieves auto-inhibition by the ISWI remodeler SNF2h.
  eLife 2018 Gamarra N, Johnson SL, Trnka MJ, Burlingame AL, Narlikar GJ
• The Yeast INO80 Complex Operates as a Tunable DNA Length-Sensitive Switch to Regulate Nucleosome Sliding.
  Molecular cell 2018 Zhou CY, Johnson SL, Lee LJ, Longhurst AD, Beckwith SL, Johnson MJ, Morrison AJ, Narlikar GJ
• Dissecting the Mechanism of HP1 Mediated Chromatin Compaction via Single Molecule DNA Curtains.
  Biophysical Journal 2018 Madeline M. Keenen, Adam G. Larson, Geeta J. Narlikar, Sy Redding
• Biochemical Basis for Distinct Roles of the Heterochromatin Proteins Swi6 and Chp2.
  Journal of molecular biology 2017 Isaac RS, Sanulli S, Tibble R, Hornsby M, Ravalin M, Craik CS, Gross JD, Narlikar GJ
• Liquid droplet formation by HP1α suggests a role for phase separation in heterochromatin.
  Nature 2017 Larson AG, Elnatan D, Keenen MM, Trnka MJ, Johnston JB, Burlingame AL, Agard DA, Redding S, Narlikar GJ
• Regulation of Rvb1/Rvb2 by a Domain within the INO80 Chromatin Remodeling Complex Implicates the Yeast Rvbs as Protein Assembly Chaperones.
  Cell reports 2017 Zhou CY, Stoddard CI, Johnston JB, Trnka MJ, Echeverria I, Palovcak E, Sali A, Burlingame AL, Cheng Y, Narlikar GJ
• Distortion of histone octamer core promotes nucleosome mobilization by a chromatin remodeler.
  Science (New York, N.Y.) 2017 Sinha KK, Gross JD, Narlikar GJ
• Mechanisms of ATP-Dependent Chromatin Remodeling Motors.
  Annual review of biophysics 2016 Zhou CY, Johnson SL, Gamarra NI, Narlikar GJ
• Nucleosome breathing and remodeling constrain CRISPR-Cas9 function.
  eLife 2016 Isaac RS, Jiang F, Doudna JA, Lim WA, Narlikar GJ, Almeida R
• Analysis of Nucleosome Sliding by ATP-Dependent Chromatin Remodeling Enzymes.
  Methods in enzymology 2016 Zhou CY, Narlikar GJ
• Single-Molecule Mechanistic Dissection of a Chromatin Remodeling Motor.
  Biophysical Journal 2016 Stephanie L. Johnson, Nathan I. Gamarra, Matthew J. Johnson, Geeta J. Narlikar
• A multilaboratory comparison of calibration accuracy and the performance of external references in analytical ultracentrifugation.
  PloS one 2015 Zhao H, Ghirlando R, Alfonso C, Arisaka F, Attali I, Bain DL, Bakhtina MM, Becker DF, Bedwell GJ, Bekdemir A, Besong TM, Birck C, Brautigam CA, Brennerman W, Byron O, Bzowska A, Chaires JB, Chaton CT,…
• A nucleotide-driven switch regulates flanking DNA length sensing by a dimeric chromatin remodeler.
  Molecular cell 2015 Leonard JD, Narlikar GJ
• Mechanisms of functional promiscuity by HP1 proteins.
  Trends in cell biology 2014 Canzio D, Larson A, Narlikar GJ
• The histone H4 tail regulates the conformation of the ATP-binding pocket in the SNF2h chromatin remodeling enzyme.
  Journal of molecular biology 2014 Racki LR, Naber N, Pate E, Leonard JD, Cooke R, Narlikar GJ
• Mechanism of Nucleosome Remodeling by INO80 from S. Cerevisiae.
  Biophysical Journal 2014 Coral Y. Zhou, Geeta J. Narlikar
• The Mechanism of Nucleosome Spacing by a Dimeric Chromatin Remodeling Enzyme.
  Biophysical Journal 2014 John D. Leonard, Jean-Paul Armache, Nariman Naber, Shenping Wu, Edward Pate, Roger Cooke, Yifan Cheng, Geeta J. Narlikar
• Mechanisms and functions of ATP-dependent chromatin-remodeling enzymes.
  Cell 2013 Narlikar GJ, Sundaramoorthy R, Owen-Hughes T
• Division of labor between the chromodomains of HP1 and Suv39 methylase enables coordination of heterochromatin spread.
  Molecular cell 2013 Al-Sady B, Madhani HD, Narlikar GJ
• Abstract IA23: Functional and structural studies of HP1 heterochromatin.
  DNA Methylation 2013 Daniele Canzio, Moafu Liang, Nariman Naber, Adam Larson, Diana Marina, Jennifer Garcia, Hiten Madhani, Roger Cooke, Peter Schuck, Yifan Cheng, Geeta J. Narlikar
• Reconstitution of nucleosome demethylation and catalytic properties of a Jumonji histone demethylase.
  Chemistry & biology 2013 Shiau C, Trnka MJ, Bozicevic A, Ortiz Torres I, Al-Sady B, Burlingame AL, Narlikar GJ, Fujimori DG
• Mechanistic Analysis of HP1 heterochromatin assembly.
  The FASEB Journal 2013 Geeta Narlikar, Daniele Canzio, Maofu Liao, Nariman Naber, Ed Pate, Adam Larson, Shenping Wu, Roger Cooke, Peter Schuck, Yifan Cheng
• A conformational switch in HP1 releases auto-inhibition to drive heterochromatin assembly.
  Nature 2013 Canzio D, Liao M, Naber N, Pate E, Larson A, Wu S, Marina DB, Garcia JF, Madhani HD, Cooke R, Schuck P, Cheng Y, Narlikar GJ
• Ers1 links HP1 to RNAi.
  Proceedings of the National Academy of Sciences of the United States of America 2012 Rougemaille M, Braun S, Coyle S, Dumesic PA, Garcia JF, Isaac RS, Libri D, Narlikar GJ, Madhani HD
• Analysis of changes in nucleosome conformation using fluorescence resonance energy transfer.
  Methods in molecular biology (Clifton, N.J.) 2012 Shahian T, Narlikar GJ
• Mechanisms of Chromatin Remodeling Motors.
  Biophysical Journal 2012 Geeta Narlikar, John Leonard, Lisa Racki, Nariman Naber, Roger Cooke, Yifan Cheng
• Structural basis of silencing: Sir3 BAH domain in complex with a nucleosome at 3.0 Å resolution.
  Science (New York, N.Y.) 2011 Armache KJ, Garlick JD, Canzio D, Narlikar GJ, Kingston RE
• Site-specific acetylation mark on an essential chromatin-remodeling complex promotes resistance to replication stress.
  Proceedings of the National Academy of Sciences of the United States of America 2011 Charles GM, Chen C, Shih SC, Collins SR, Beltrao P, Zhang X, Sharma T, Tan S, Burlingame AL, Krogan NJ, Madhani HD, Narlikar GJ
• EPR Spectroscopy Shows That the H4 Tail of the Nucleosome Influences the Nucleotide-Binding Pocket of Acf.
  Biophysical Journal 2011 Lisa Racki, Nariman Naber, Roger Cooke, Geeta Narlikar
• Chromodomain-mediated oligomerization of HP1 suggests a nucleosome-bridging mechanism for heterochromatin assembly.
  Molecular cell 2011 Canzio D, Chang EY, Shankar S, Kuchenbecker KM, Simon MD, Madhani HD, Narlikar GJ, Al-Sady B
• The ATP-dependent remodeler RSC transfers histone dimers and octamers through the rapid formation of an unstable encounter intermediate.
  Biochemistry 2010 Rowe CE, Narlikar GJ
• A proposal for kinetic proof reading by ISWI family chromatin remodeling motors.
  Current opinion in chemical biology 2010 Narlikar GJ
• The Chromatin-Remodeling Complex ACF Functions as a Dimeric Motor to Space Nucleosomes.
  Biophysical Journal 2010 Lisa R. Racki, Geeta J. Narlikar, Roger Cooke, Nariman Naber, Thomas J. Purcell, Janet G. Yang, Yifan Cheng, Ashley Acevedo, Peretz Partensky
• Dynamics of nucleosome remodelling by individual ACF complexes.
  Nature 2009 Blosser TR, Yang JG, Stone MD, Narlikar GJ, Zhuang X
• The chromatin remodeller ACF acts as a dimeric motor to space nucleosomes.
  Nature 2009 Racki LR, Yang JG, Naber N, Partensky PD, Acevedo A, Purcell TJ, Cooke R, Cheng Y, Narlikar GJ
• Chromatin remodelers act globally, sequence positions nucleosomes locally.
  Journal of molecular biology 2009 Partensky PD, Narlikar GJ
• Mechanisms of ATP-dependent Chromatin Remodeling Enzymes.
  The FASEB Journal 2009 Geeta J Narlikar, Lisa Racki, Janet Yang, Yifan Cheng, Nariman Naber, Roger Cooke
• Nucleosome Sliding by ACF is Processive and Bidirectional.
  Biophysical Journal 2009 Timothy R. Blosser, Michael D. Stone, Janet Yang, Geeta Narlikar, Xiaowei Zhuang
• MacroH2A allows ATP-dependent chromatin remodeling by SWI/SNF and ACF complexes but specifically reduces recruitment of SWI/SNF.
  Biochemistry 2008 Chang EY, Ferreira H, Somers J, Nusinow DA, Owen-Hughes T, Narlikar GJ
• Epigenomics: a roadmap, but to where?
  Science (New York, N.Y.) 2008 Madhani HD, Francis NJ, Kingston RE, Kornberg RD, Moazed D, Narlikar GJ, Panning B, Struhl K
• ATP-dependent chromatin remodeling enzymes: two heads are not better, just different.
  Current opinion in genetics & development 2008 Racki LR, Narlikar GJ
• Mechanism of the ATP-dependent chromatin remodeling complex ACF.
  The FASEB Journal 2007 Geeta J Narlikar, Janet G Yang, Lisa Racki, Peretz Partensky
• The site-specific installation of methyl-lysine analogs into recombinant histones.
  Cell 2007 Simon MD, Chu F, Racki LR, de la Cruz CC, Burlingame AL, Panning B, Narlikar GJ, Shokat KM
• FRET-based methods to study ATP-dependent changes in chromatin structure.
  Methods (San Diego, Calif.) 2007 Yang JG, Narlikar GJ
• The chromatin-remodeling enzyme ACF is an ATP-dependent DNA length sensor that regulates nucleosome spacing.
  Nature structural & molecular biology 2006 Yang JG, Madrid TS, Sevastopoulos E, Narlikar GJ
• Human ACF1 alters the remodeling strategy of SNF2h.
  The Journal of biological chemistry 2006 He X, Fan HY, Narlikar GJ, Kingston RE
• Heterogeneous nuclear ribonucleoprotein C1/C2, MeCP1, and SWI/SNF form a chromatin remodeling complex at the beta-globin locus control region.
  Proceedings of the National Academy of Sciences of the United States of America 2005 Mahajan MC, Narlikar GJ, Boyapaty G, Kingston RE, Weissman SM
• Noncovalent modification of chromatin: different remodeled products with different ATPase domains.
  Cold Spring Harbor symposia on quantitative biology 2004 Fan HY, Narlikar GJ, Kingston RE
• Distinct strategies to make nucleosomal DNA accessible.
  Molecular cell 2003 Fan HY, He X, Kingston RE, Narlikar GJ
• Nucleosome remodeling by the human SWI/SNF complex requires transient global disruption of histone-DNA interactions.
  Molecular and cellular biology 2002 Aoyagi S, Narlikar G, Zheng C, Sif S, Kingston RE, Hayes JJ
• Cooperation between complexes that regulate chromatin structure and transcription.
  Cell 2002 Narlikar GJ, Fan HY, Kingston RE
• Generation and interconversion of multiple distinct nucleosomal states as a mechanism for catalyzing chromatin fluidity.
  Molecular cell 2001 Narlikar GJ, Phelan ML, Kingston RE
• Functional differences between the human ATP-dependent nucleosome remodeling proteins BRG1 and SNF2H.
  The Journal of biological chemistry 2001 Aalfs JD, Narlikar GJ, Kingston RE
• Assembly of nucleosomal templates by salt dialysis.
  Current protocols in molecular biology 2001 Lee KM, Narlikar G
• Rvb1p and Rvb2p are essential components of a chromatin remodeling complex that regulates transcription of over 5% of yeast genes.
  The Journal of biological chemistry 2001 Jónsson ZO, Dhar SK, Narlikar GJ, Auty R, Wagle N, Pellman D, Pratt RE, Kingston R, Dutta A
• Stability of a human SWI-SNF remodeled nucleosomal array.
  Molecular and cellular biology 2001 Guyon JR, Narlikar GJ, Sullivan EK, Kingston RE
• Use of duplex rigidity for stability and specificity in RNA tertiary structure.
  Biochemistry 2000 Narlikar GJ, Bartley LE, Herschlag D
• Characterization of a local folding event of the Tetrahymena group I ribozyme: effects of oligonucleotide substrate length, pH, and temperature on the two substrate binding steps.
  Biochemistry 1999 Narlikar GJ, Bartley LE, Khosla M, Herschlag D
• ATP-dependent remodeling and acetylation as regulators of chromatin fluidity.
  Genes & development 1999 Kingston RE, Narlikar GJ
• Protonated 2'-aminoguanosine as a probe of the electrostatic environment of the active site of the Tetrahymena group I ribozyme.
  Biochemistry 1999 Shan SO, Narlikar GJ, Herschlag D
• Stable remodeling of tailless nucleosomes by the human SWI-SNF complex.
  Molecular and cellular biology 1999 Guyon JR, Narlikar GJ, Sif S, Kingston RE
• Reconstitution of a core chromatin remodeling complex from SWI/SNF subunits.
  Molecular cell 1999 Phelan ML, Sif S, Narlikar GJ, Kingston RE
• Direct demonstration of the catalytic role of binding interactions in an enzymatic reaction.
  Biochemistry 1998 Narlikar GJ, Herschlag D
• Quantitating tertiary binding energies of 2' OH groups on the P1 duplex of the Tetrahymena ribozyme: intrinsic binding energy in an RNA enzyme.
  Biochemistry 1997 Narlikar GJ, Khosla M, Usman N, Herschlag D
• Mechanistic aspects of enzymatic catalysis: lessons from comparison of RNA and protein enzymes.
  Annual review of biochemistry 1997 Narlikar GJ, Herschlag D
• Isolation of a local tertiary folding transition in the context of a globally folded RNA.
  Nature structural biology 1996 Narlikar GJ, Herschlag D
• Use of binding energy by an RNA enzyme for catalysis by positioning and substrate destabilization.
  Proceedings of the National Academy of Sciences of the United States of America 1995 Narlikar GJ, Gopalakrishnan V, McConnell TS, Usman N, Herschlag D
• Dissection of the role of the conserved G.U pair in group I RNA self-splicing.
  Biochemistry 1994 Knitt DS, Narlikar GJ, Herschlag D