Irene Chen

Professor, Chemical And Biomolecular Engineering, University of California Los Angeles

Professor, Chemistry and Biochemistry, University of California Los Angeles

Irene A. Chen is a Professor in the Department of Chemical and Biomolecular Engineering at the University of California, Los Angeles. Her laboratory studies life-like biochemical systems to understand their fundamental properties and address emerging challenges in biotechnology and infectious disease.

Education and Training

Harvard UniversityFellow2013Systems Biology
Harvard Medical SchoolMD2007Medicine, Health Sciences and Technology
Harvard UniversityPhD2007Biophysics
Harvard UniversityAB1999Chemistry

Awards and Honors

  • Camille Dreyfus Teacher-Scholar Award, 2018-2023.
  • NIH New Innovator Award, 2016-2021.
  • Searle Scholar Award, 2014-2017.
  • Simons Investigator, Collaboration on the Origins of Life, 2013-2023.

Publications

  1. Batra J, Rutkowska M, Zhou Y, Ye C, Adavikolanu R, Young JM, Anand D, Verma S, Parthasarathy H, Gordon M, Malpotra S, Cupic A, Kehrer T, Dos Santos M, Benjamin R, Moen JM, Winters DM, Caval V, Rojc A, Mena I, Aslam S, Martinez-Romero C, Viñas IC, Khalil Z, Farrugia K, Villalón-Letelier F, Banerjee A, Tussia-Cohen D, Diallo A, Maji S, Muralidharan M, Foussard H, Chen IP, Fuchs R, San Felipe CJ, Zuliani-Alvarez L, Choudhury P, Obernier K, Gracias S, Suryawanshi RK, Bonaventure B, Ibáñez C, Johnson JR, Juste J, Pache L, Stroud RM, Verba KA, Fraser JS, van Bakel H, Taha TY, Ott M, Hagai T, Jouvenet N, Demeret C, Polacco BJ, Swaney DL, Echeverria I, Bouhaddou M, Eckhardt M, Malik HS, Martinez-Sobrido L, Miorin L, García-Sastre A, Krogan NJ. Coronavirus protein interaction mapping in bat and human cells reveals network rewiring governing immune evasion and zoonotic potential.. Cell host & microbe, 2026.
  2. Khalid MM, Chen IP, Soveg FS, Taha TY, Tabata T, Suryawanshi RK, Syed AM, Ciling A, McCavitt-Malvido M, Schulze-Gahmen U, Hayashi J, Kim IJ, Fong SW, Batra J, Kumar GR, Renia L, Ng LF, Krogan NJ, Doudna JA, Verdin E, Ott M. Regulation of virion production by the ORF8 signal peptide across SARS-CoV-2 variants.. bioRxiv : the preprint server for biology, 2026.
  3. Bagchi D, Adhikari A, McCarthy K, Kang D, Chen Y, Chen IA. Silver Nanoparticles Templated by the M13 Phage Exhibit High Antibacterial Activity against Gram-Negative Pathogens and a Reduced Rate of Bacterial Resistance In Vitro.. Langmuir : the ACS journal of surfaces and colloids, 2025.
  4. Yang Y, Kang D, Mihalache B, Vexler S, Jain S, Peng H, Annabi N, Yang S, Chen IA. Metagenome-inspired libraries to engineer phage M13 for targeted killing of Gram-negative bacterial species.. Nucleic acids research, 2025.
  5. Yang Y, Vexler S, Jordan MC, Abbondante S, Kang D, Peng H, Marshall M, Naini BV, Jain S, Lai YC, Annabi N, Roos KP, Pearlman E, Chen IA. A Synthetic Phage-Peptide Conjugate as a Potent Antibacterial Agent for Pseudomonas aeruginosa Infections.. ACS central science, 2025.
  6. Salas-Ambrosio P, Vexler S, P Sivasankaran R, Vlahakis N, Lai RS, Johnson C, Baas-Maynard SI, Min DS, Lower H, Doyle AG, Tang Y, Rodriguez JA, Chen IA, Read de Alaniz J, Maynard HD. Biosourced Functional Hydroxybenzoate-co-Lactide Polymers with Antimicrobial Activity.. Journal of the American Chemical Society, 2025.
  7. Taha TY, Ezzatpour S, Hayashi JM, Ye C, Zapatero-Belinchón FJ, Rosecrans JA, Kimmerly GR, Chen IP, Walcott K, Kurianowicz A, Jorgens DM, Chaplin NR, Choi A, Buchholz DW, Sahler J, Hilt ZT, Imbiakha B, Vagi-Szmola C, Montano M, Stevenson E, Gordon M, Swaney DL, Krogan NJ, Whittaker GR, Martinez-Sobrido L, Aguilar HC, Ott M. Enhanced RNA replication and pathogenesis in recent SARS-CoV-2 variants harboring the L260F mutation in NSP6.. PLoS pathogens, 2025.
  8. Adhikari A, Chen IA. Antibody-Nanoparticle Conjugates in Therapy: Combining the Best of Two Worlds.. Small (Weinheim an der Bergstrasse, Germany), 2025.
  9. Seelig B, Chen IA. Intellectual frameworks to understand complex biochemical systems at the origin of life.. Nature chemistry, 2025.
  10. Syed AM, Ciling A, Chen IP, Carlson CR, Adly AN, Martin HS, Taha TY, Khalid MM, Price N, Bouhaddou M, Ummadi MR, Moen JM, Krogan NJ, Morgan DO, Ott M, Doudna JA. SARS-CoV-2 evolution balances conflicting roles of N protein phosphorylation.. PLoS pathogens, 2024.
  11. Kenchel J, Vázquez-Salazar A, Wells R, Brunton K, Janzen E, Schultz KM, Liu Z, Li W, Parker ET, Dworkin JP, Chen IA. Prebiotic chiral transfer from self-aminoacylating ribozymes may favor either handedness.. Nature communications, 2024.
  12. Weinberg ZY, Soliman SS, Kim MS, Shah DH, Chen IP, Ott M, Lim WA, El-Samad H. De novo-designed minibinders expand the synthetic biology sensing repertoire.. bioRxiv : the preprint server for biology, 2024.
  13. Saha R, Choi JA, Chen IA. Protocell Effects on RNA Folding, Function, and Evolution.. Accounts of chemical research, 2024.
  14. Simoneau CR, Chen PY, Xing GK, Hayashi JM, Chen IP, Khalid MM, Meyers NL, Taha TY, Leon KE, Suryawanshi RK, McCavitt-Malvido M, Ashuach T, Fontaine KA, Rodriguez L, Joehnk B, Walcott K, Vasudevan S, Fang X, Maishan M, Schultz S, Roose JP, Matthay MA, Sil A, Arjomandi M, Yosef N, Ott M. NF-κB inhibitor alpha controls SARS-CoV-2 infection in ACE2-overexpressing human airway organoids.. Scientific reports, 2024.
  15. Saha R, Vázquez-Salazar A, Nandy A, Chen IA. Fitness Landscapes and Evolution of Catalytic RNA.. Annual review of biophysics, 2024.
  16. Chen IA. RNA life on the edge of catastrophe.. Proceedings of the National Academy of Sciences of the United States of America, 2024.
  17. Yang Y, Chen IA. Visualization of Engineered M13 Phages Bound to Bacterial Targets by Transmission Electron Microscopy.. Methods in molecular biology (Clifton, N.J.), 2024.
  18. Kang D, Bagchi D, Chen IA. Pharmacokinetics and Biodistribution of Phages and their Current Applications in Antimicrobial Therapy.. Advanced therapeutics, 2023.
  19. Saha R, Kao WL, Malady B, Heng X, Chen IA. Effect of montmorillonite K10 clay on RNA structure and function.. Biophysical journal, 2023.
  20. Suryawanshi RK, Taha TY, McCavitt-Malvido M, Silva I, Khalid MM, Syed AM, Chen IP, Saldhi P, Sreekumar B, Montano M, Foresythe K, Tabata T, Kumar GR, Sotomayor-Gonzalez A, Servellita V, Gliwa A, Nguyen J, Kojima N, Arellanor T, Bussanich A, Hess V, Shacreaw M, Lopez L, Brobeck M, Turner F, Wang Y, Ghazarian S, Davis G, Rodriguez D, Doudna J, Spraggon L, Chiu CY, Ott M. Previous exposure to Spike-providing parental strains confers neutralizing immunity to XBB lineage and other SARS-CoV-2 recombinants in the context of vaccination.. Emerging microbes & infections, 2023.
  21. Ma T, Suryawanshi RK, Miller SR, Ly KK, Thomas R, Elphick N, Yin K, Luo X, Kaliss N, Chen IP, Montano M, Sreekumar B, Standker L, Münch J, Heath Damron F, Palop JJ, Ott M, Roan NR. Post-acute immunological and behavioral sequelae in mice after Omicron infection.. bioRxiv : the preprint server for biology, 2023.
  22. Taha TY, Suryawanshi RK, Chen IP, Correy GJ, McCavitt-Malvido M, O'Leary PC, Jogalekar MP, Diolaiti ME, Kimmerly GR, Tsou CL, Gascon R, Montano M, Martinez-Sobrido L, Krogan NJ, Ashworth A, Fraser JS, Ott M. A single inactivating amino acid change in the SARS-CoV-2 NSP3 Mac1 domain attenuates viral replication in vivo.. PLoS pathogens, 2023.
  23. Charest N, Shen Y, Lai YC, Chen IA, Shea JE. Discovering pathways through ribozyme fitness landscapes using information theoretic quantification of epistasis.. RNA (New York, N.Y.), 2023.
  24. Taha TY, Chen IP, Hayashi JM, Tabata T, Walcott K, Kimmerly GR, Syed AM, Ciling A, Suryawanshi RK, Martin HS, Bach BH, Tsou CL, Montano M, Khalid MM, Sreekumar BK, Renuka Kumar G, Wyman S, Doudna JA, Ott M. Rapid assembly of SARS-CoV-2 genomes reveals attenuation of the Omicron BA.1 variant through NSP6.. Nature communications, 2023.
  25. Salas-Ambrosio P, Vexler S, P S R, Chen IA, Maynard HD. Caffeine and Cationic Copolymers with Antimicrobial Properties.. ACS bio & med chem Au, 2023.
  26. Zhou B, Mobberley J, Shi K, Chen IA. Effects of Preservation and Propagation Methodology on Microcosms Derived from the Oral Microbiome.. Microorganisms, 2022.
  27. Walter M, Chen IP, Vallejo-Gracia A, Kim IJ, Bielska O, Lam VL, Hayashi JM, Cruz A, Shah S, Soveg FW, Gross JD, Krogan NJ, Jerome KR, Schilling B, Ott M, Verdin E. SIRT5 is a proviral factor that interacts with SARS-CoV-2 Nsp14 protein.. PLoS pathogens, 2022.
  28. Tran CH, Saha R, Blanco C, Bagchi D, Chen IA. Modulation of α-Synuclein Aggregation In Vitro by a DNA Aptamer.. Biochemistry, 2022.
  29. Syed AM, Ciling A, Taha TY, Chen IP, Khalid MM, Sreekumar B, Chen PY, Kumar GR, Suryawanshi R, Silva I, Milbes B, Kojima N, Hess V, Shacreaw M, Lopez L, Brobeck M, Turner F, Spraggon L, Tabata T, Ott M, Doudna JA. Omicron mutations enhance infectivity and reduce antibody neutralization of SARS-CoV-2 virus-like particles.. Proceedings of the National Academy of Sciences of the United States of America, 2022.
  30. Chen IP, Longbotham JE, McMahon S, Suryawanshi RK, Khalid MM, Taha TY, Tabata T, Hayashi JM, Soveg FW, Carlson-Stevermer J, Gupta M, Zhang MY, Lam VL, Li Y, Yu Z, Titus EW, Diallo A, Oki J, Holden K, Krogan N, Fujimori DG, Ott M. Viral E protein neutralizes BET protein-mediated post-entry antagonism of SARS-CoV-2.. Cell reports, 2022.
  31. Janzen E, Shen Y, Vázquez-Salazar A, Liu Z, Blanco C, Kenchel J, Chen IA. Emergent properties as by-products of prebiotic evolution of aminoacylation ribozymes.. Nature communications, 2022.
  32. Vázquez-Salazar A, Chen IA. In vitro evolution: From monsters to mobs.. Current biology : CB, 2022.
  33. Suryawanshi RK, Chen IP, Ma T, Syed AM, Brazer N, Saldhi P, Simoneau CR, Ciling A, Khalid MM, Sreekumar B, Chen PY, Kumar GR, Montano M, Gascon R, Tsou CL, Garcia-Knight MA, Sotomayor-Gonzalez A, Servellita V, Gliwa A, Nguyen J, Silva I, Milbes B, Kojima N, Hess V, Shacreaw M, Lopez L, Brobeck M, Turner F, Soveg FW, George AF, Fang X, Maishan M, Matthay M, Morris MK, Wadford D, Hanson C, Greene WC, Andino R, Spraggon L, Roan NR, Chiu CY, Doudna JA, Ott M. Limited cross-variant immunity from SARS-CoV-2 Omicron without vaccination.. Nature, 2022.
  34. Verbanic S, Deacon JM, Chen IA. The Chronic Wound Phageome: Phage Diversity and Associations with Wounds and Healing Outcomes.. Microbiology spectrum, 2022.
  35. Peng H, Rossetto D, Mansy SS, Jordan MC, Roos KP, Chen IA. Treatment of Wound Infections in a Mouse Model Using Zn2+-Releasing Phage Bound to Gold Nanorods.. ACS nano, 2022.
  36. Syed AM, Ciling A, Khalid MM, Sreekumar B, Chen PY, Kumar GR, Silva I, Milbes B, Kojima N, Hess V, Shacreaw M, Lopez L, Brobeck M, Turner F, Spraggon L, Taha TY, Tabata T, Chen IP, Ott M, Doudna JA. Omicron mutations enhance infectivity and reduce antibody neutralization of SARS-CoV-2 virus-like particles.. medRxiv : the preprint server for health sciences, 2022.
  37. Peng H, Lelievre A, Landenfeld K, Müller S, Chen IA. Vesicle encapsulation stabilizes intermolecular association and structure formation of functional RNA and DNA.. Current biology : CB, 2021.
  38. Syed AM, Taha TY, Tabata T, Chen IP, Ciling A, Khalid MM, Sreekumar B, Chen PY, Hayashi JM, Soczek KM, Ott M, Doudna JA. Rapid assessment of SARS-CoV-2-evolved variants using virus-like particles.. Science (New York, N.Y.), 2021.
  39. Kanwar N, Blanco C, Chen IA, Seelig B. PacBio sequencing output increased through uniform and directional fivefold concatenation.. Scientific reports, 2021.
  40. Shen Y, Pressman A, Janzen E, Chen IA. Kinetic sequencing (k-Seq) as a massively parallel assay for ribozyme kinetics: utility and critical parameters.. Nucleic acids research, 2021.
  41. Peng H, Latifi B, Müller S, Lupták A, Chen IA. Self-cleaving ribozymes: substrate specificity and synthetic biology applications.. RSC chemical biology, 2021.
  42. Lai YC, Liu Z, Chen IA. Encapsulation of ribozymes inside model protocells leads to faster evolutionary adaptation.. Proceedings of the National Academy of Sciences of the United States of America, 2021.
  43. Peng H, Chen IA. Phage engineering and the evolutionary arms race.. Current opinion in biotechnology, 2020.
  44. Blanco C, Verbanic S, Seelig B, Chen IA. EasyDIVER: A Pipeline for Assembling and Counting High-Throughput Sequencing Data from In Vitro Evolution of Nucleic Acids or Peptides.. Journal of molecular evolution, 2020.
  45. Lai YC, Chen IA. Protocells.. Current biology : CB, 2020.
  46. Peng H, Borg RE, Nguyen ABN, Chen IA. Chimeric Phage Nanoparticles for Rapid Characterization of Bacterial Pathogens: Detection in Complex Biological Samples and Determination of Antibiotic Sensitivity.. ACS sensors, 2020.
  47. Verbanic S, Shen Y, Lee J, Deacon JM, Chen IA. Microbial predictors of healing and short-term effect of debridement on the microbiome of chronic wounds.. NPJ biofilms and microbiomes, 2020.
  48. Janzen E, Blanco C, Peng H, Kenchel J, Chen IA. Promiscuous Ribozymes and Their Proposed Role in Prebiotic Evolution.. Chemical reviews, 2020.
  49. Blanco C, Verbanic S, Seelig B, Chen IA. High throughput sequencing of in vitro selections of mRNA-displayed peptides: data analysis and applications.. Physical chemistry chemical physics : PCCP, 2020.
  50. Peng H, Borg RE, Dow LP, Pruitt BL, Chen IA. Controlled phage therapy by photothermal ablation of specific bacterial species using gold nanorods targeted by chimeric phages.. Proceedings of the National Academy of Sciences of the United States of America, 2020.
  51. Verbanic S, Kim CY, Deacon JM, Chen IA. Improved single-swab sample preparation for recovering bacterial and phage DNA from human skin and wound microbiomes.. BMC microbiology, 2019.
  52. Pascal R, Chen IA. From soup to peptides.. Nature chemistry, 2019.
  53. Saha R, Chen IA. Effect of UV Radiation on Fluorescent RNA Aptamers' Functional and Templating Ability.. Chembiochem : a European journal of chemical biology, 2019.
  54. Guan J, Zhou W, Hafner M, Blake RA, Chalouni C, Chen IP, De Bruyn T, Giltnane JM, Hartman SJ, Heidersbach A, Houtman R, Ingalla E, Kategaya L, Kleinheinz T, Li J, Martin SE, Modrusan Z, Nannini M, Oeh J, Ubhayakar S, Wang X, Wertz IE, Young A, Yu M, Sampath D, Hager JH, Friedman LS, Daemen A, Metcalfe C. Therapeutic Ligands Antagonize Estrogen Receptor Function by Impairing Its Mobility.. Cell, 2019.
  55. Blanco C, Chen IA. Phage therapy administered noninvasively could be effective in thin tubes subject to episodic flow despite washout: a simulation study.. Physical biology, 2019.
  56. Pressman AD, Liu Z, Janzen E, Blanco C, Müller UF, Joyce GF, Pascal R, Chen IA. Mapping a Systematic Ribozyme Fitness Landscape Reveals a Frustrated Evolutionary Network for Self-Aminoacylating RNA.. Journal of the American Chemical Society, 2019.
  57. Blanco C, Janzen E, Pressman A, Saha R, Chen IA. Molecular Fitness Landscapes from High-Coverage Sequence Profiling.. Annual review of biophysics, 2019.
  58. Peng H, Chen IA. Rapid Colorimetric Detection of Bacterial Species through the Capture of Gold Nanoparticles by Chimeric Phages.. ACS nano, 2019.
  59. Saha R, Verbanic S, Chen IA. Lipid vesicles chaperone an encapsulated RNA aptamer.. Nature communications, 2018.
  60. Blanco C, Bayas M, Yan F, Chen IA. Analysis of Evolutionarily Independent Protein-RNA Complexes Yields a Criterion to Evaluate the Relevance of Prebiotic Scenarios.. Current biology : CB, 2018.
  61. Pressman A, Moretti JE, Campbell GW, Müller UF, Chen IA. Analysis of in vitro evolution reveals the underlying distribution of catalytic activity among random sequences.. Nucleic acids research, 2017.
  62. Pressman A, Moretti JE, Campbell GW, Müller UF, Chen IA. Analysis of in vitro evolution reveals the underlying distribution of catalytic activity among random sequences.. Nucleic acids research, 2017.
  63. Chen IA, de Vries MS. From underwear to non-equilibrium thermodynamics: physical chemistry informs the origin of life.. Physical chemistry chemical physics : PCCP, 2016.
  64. Xulvi-Brunet R, Campbell GW, Rajamani S, Jiménez JI, Chen IA. Computational analysis of fitness landscapes and evolutionary networks from in vitro evolution experiments.. Methods (San Diego, Calif.), 2016.
  65. Saha R, Chen IA. Origin of Life: Protocells Red in Tooth and Claw.. Current biology : CB, 2015.
  66. Pressman A, Blanco C, Chen IA. The RNA World as a Model System to Study the Origin of Life.. Current biology : CB, 2015.
  67. Chen IA. Prebiotic chemistry: Replicating towards complexity.. Nature chemistry, 2015.
  68. Saha R, Pohorille A, Chen IA. Molecular crowding and early evolution.. Origins of life and evolution of the biosphere : the journal of the International Society for the Study of the Origin of Life, 2015.
  69. Athavale SS, Spicer B, Chen IA. Experimental fitness landscapes to understand the molecular evolution of RNA-based life.. Current opinion in chemical biology, 2014.
  70. Freese PD, Korolev KS, Jiménez JI, Chen IA. Genetic drift suppresses bacterial conjugation in spatially structured populations.. Biophysical journal, 2014.
  71. Ivica NA, Obermayer B, Campbell GW, Rajamani S, Gerland U, Chen IA. The paradox of dual roles in the RNA world: resolving the conflict between stable folding and templating ability.. Journal of molecular evolution, 2013.
  72. Jiménez JI, Xulvi-Brunet R, Campbell GW, Turk-MacLeod R, Chen IA. Comprehensive experimental fitness landscape and evolutionary network for small RNA.. Proceedings of the National Academy of Sciences of the United States of America, 2013.
  73. Bianconi G, Zhao K, Chen IA, Nowak MA. Selection for replicases in protocells.. PLoS computational biology, 2013.
  74. Leconte AM, Dickinson BC, Yang DD, Chen IA, Allen B, Liu DR. A population-based experimental model for protein evolution: effects of mutation rate and selection stringency on evolutionary outcomes.. Biochemistry, 2013.
  75. Leu K, Kervio E, Obermayer B, Turk-MacLeod RM, Yuan C, Luevano JM, Chen E, Gerland U, Richert C, Chen IA. Cascade of reduced speed and accuracy after errors in enzyme-free copying of nucleic acid sequences.. Journal of the American Chemical Society, 2012.
  76. Vaidya N, Manapat ML, Chen IA, Xulvi-Brunet R, Hayden EJ, Lehman N. Spontaneous network formation among cooperative RNA replicators.. Nature, 2012.
  77. Chen IA, Nowak MA. From prelife to life: how chemical kinetics become evolutionary dynamics.. Accounts of chemical research, 2012.
  78. Derr J, Manapat ML, Rajamani S, Leu K, Xulvi-Brunet R, Joseph I, Nowak MA, Chen IA. Prebiotically plausible mechanisms increase compositional diversity of nucleic acid sequences.. Nucleic acids research, 2012.
  79. Leu K, Obermayer B, Rajamani S, Gerland U, Chen IA. The prebiotic evolutionary advantage of transferring genetic information from RNA to DNA.. Nucleic acids research, 2011.
  80. Lin A, Jimenez J, Derr J, Vera P, Manapat ML, Esvelt KM, Villanueva L, Liu DR, Chen IA. Inhibition of bacterial conjugation by phage M13 and its protein g3p: quantitative analysis and model.. PloS one, 2011.
  81. Chen IA, Schindlinger M. Quadruplet codons: one small step for a ribosome, one giant leap for proteins: an expanded genetic code could address fundamental questions about algorithmic information, biological function, and the origins of life.. BioEssays : news and reviews in molecular, cellular and developmental biology, 2010.
  82. Chen IA, Walde P. From self-assembled vesicles to protocells.. Cold Spring Harbor perspectives in biology, 2010.
  83. 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.. Journal of the American Chemical Society, 2010.
  84. Manapat ML, Chen IA, Nowak MA. The basic reproductive ratio of life.. Journal of theoretical biology, 2009.
  85. Chen IA. Cell division: breaking up is easy to do.. Current biology : CB, 2009.
  86. Chen IA. GE Prize-winning essay. The emergence of cells during the origin of life.. Science (New York, N.Y.), 2006.
  87. Fujikawa SM, Chen IA, Szostak JW. Shrink-wrap vesicles.. Langmuir : the ACS journal of surfaces and colloids, 2005.
  88. Chen IA, Salehi-Ashtiani K, Szostak JW. RNA catalysis in model protocell vesicles.. Journal of the American Chemical Society, 2005.
  89. Chen IA, Roberts RW, Szostak JW. The emergence of competition between model protocells.. Science (New York, N.Y.), 2004.
  90. Chen IA, Szostak JW. A kinetic study of the growth of fatty acid vesicles.. Biophysical journal, 2004.
  91. Chen IA, Szostak JW. Membrane growth can generate a transmembrane pH gradient in fatty acid vesicles.. Proceedings of the National Academy of Sciences of the United States of America, 2004.
  92. Kast P, Grisostomi C, Chen IA, Li S, Krengel U, Xue Y, Hilvert D. A strategically positioned cation is crucial for efficient catalysis by chorismate mutase.. The Journal of biological chemistry, 2000.