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All Publications

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Axxx: Original Research Articles Bxxx: Invited/Other Publications

Anthropogeny Publications
Red Meat Risk Publications


A291. Khan, Naazneen et al. Sialoglycan binding patterns of bacterial AB5 toxin B subunits correlate with host range and toxicity, indicating evolution independent of A subunits. J Biol Chem. 10:1016.101900. 2022. Epub ahead of print.


A290. Vaill M, Chen DY, Diaz S, Varki A. Improved methods to characterize the length and quantity of highly unstable PolySialic acids subject category: (Carbohydrates, chromatographic techniques). Anal Biochem. 20:114426. 2021.

A289. Kooner AS, Diaz S, Yu H, Santra A, Varki A, Chen X. Chemoenzymatic Synthesis of Sialosides Containing 7-N- or 7,9-Di-N-acetyl Sialic Acid as Stable O-Acetyl Analogues for Probing Sialic Acid-Binding Proteins. J Org Chem. 86(21):14381-14397. 2021.

A288. Kawanishi K, Coker JK, Grunddal KV, Dhar C, Hsiao J, Zengler K, Varki N, Varki A, Gordts PLSM. Dietary Neu5Ac Intervention Protects Against Atherosclerosis Associated with Human-Like Neu5Gc Loss. Arterioscler Thromb Vasc Biol. 11:2730-2739. 2021.

A287. Nason, R., Büll, C., Konstantinidi, A., Sun, L., Ye, Z., Halim, A., Du, W., Sørensen, D.M., Durbesson, F., Furukawa, S., Mandel, U., Joshi, H.J., Dworkin, L.A., Hansen, L., David, L., Iverson, T.M., Bensing, B.A., Sullam, P.M., Varki, A., Vries, E., de Haan, C.A.M., Vincentelli, R., Henrissat, B., Vakhrushev, S.Y., Clausen, H., and Narimatsu, Y. Display of the human mucinome with defined O-glycans by gene engineered cells. Nat Commun. 12:4070. 2021.

B204. Dhar, C., Sasmal, A., Diaz, S., Verhagen, A., Yu, H., Li, W., Chen, X., and Varki, A. Are sialic acids involved in COVID-19 pathogenesis? Glycobiology. 31(9):1068-1071. 2021.

A286. Martin, P.T., Kawanishi, K., Ashbrook, A., Golden, B., Samraj, A., Crowe, K.E., Zygmunt, D.A., Okerblom, J., Yu, H., Maki, A., Diaz, S., Chen, X., Janssen, P.M.L., and Varki, A. Serum Antibodies to N-Glycolylneuraminic Acid Are Elevated in Duchenne Muscular Dystrophy and Correlate with Increased Disease Pathology in Cmah−/−mdx Mice. Am J Pathol. 191:1474-1486. 2021.

A285.  Ji, Y., Sasmal, A., Li, W., Oh, L., Srivastava, S., Hargett, A.A., Wasik, B.R., Yu, H., Diaz, S., Choudhury, B., Parrish, C.R., Freedberg, D.I., Wang, L.P., Varki, A., and Chen, X. Reversible O-Acetyl Migration within the Sialic Acid Side Chain and Its Influence on Protein Recognition. ACS Chem Biol. 2021.

B203. Siddiqui, S.S., Vaill, M., and Varki, A. Ongoing selection for a uniquely human null allele of SIGLEC12 in world-wide populations may protect against the risk of advanced carcinomas. FASEB Bioadv. 3:278-279. 2021.

A284. Siddiqui, S.S., Dhar, C., Sundaramurthy, V., Sasmal, A., Yu, H., Bandala-Sanchez, E., Li, M., Zhang, X., Chen, X., Harrison, L., Xu, D., Varki, A. Sialoglycan recognition is a common connection linking acidosis, zinc, and HMGB1 in sepsisProc Natl Acad Sci USA. 118(10):e2018090118. 2021.

A283.  Kawanishi, K., Saha, S., Diaz, S., Vaill, M., Sasmal, A., Siddiqui, S.S., Choudhury, B.P., Sharma, K., Chen, X., S7choenhofen, I.C., Sato, C., Kitajima, K., Freeze, H.H., Münster-Kühnel, A., and Varki, A. Evolutionary conservation of human ketodeoxynonulosonic acid production is independent of sialoglycan biosynthesis. J Clin Invest. 131(5):e137681 2021.

A282. Saha, S., Coady, A., Sasmal, A., Kawanishi, K., Choudhury, B., Yu, H., Sorensen, R., Inostroza, J., Schoenhofen, I., Chen, X., Münster-Kühnel, A., Sato, C., Kitajima, K., Ram, S., Nizet, V., and Varki, A. Exploring the impact of ketodeoxynonulosonic acid in host-pathogen interactions using uptake and surface display by non-typeable Haemophilus influenzae. mBio. 12(1):e03226-20. 2021. Supplemental Materials 


A281. Siddiqui, S.S., Vaill, M., Do, R., Khan, N., Verhagen, A.L., Zhang, W., Lenz, H., Johnson‐Pais, T.L., Leach, R.J., Fraser, G., Wang, C., Feng, G., Varki, N., and Varki, A. Human‐specific polymorphic pseudogenization of SIGLEC12 protects against advanced cancer progression. FASEB BioAdvances. 3:69-82. 2020.    UCSD Press Release

A280. Gulati S, Schoenhofen IC, Lindhout-Djukic T, Lewis LA, Moustafa IY, Saha S, Zheng B, Nowak N, Rice PA, Varki A, Ram S. Efficacy of Antigonococcal CMP-Nonulosonate Therapeutics Require Cathelicidins. J Infect Dis. 222(10):1641-1650. 2020.

A279. Hane M, Chen DY, Varki A. Human-specific Microglial Siglec-11 Transcript Variant has the Potential to affect Polysialic Acid-Mediated Brain Functions at a Distance. Glycobiology. 31:231-242. 2020

A278. Awofiranye, A. E., Baytas, S. N., Xia, K., Badri, A., He, W., Varki, A., Koffas, M. and Linhardt, R. J. N-glycolyl chondroitin synthesis using metabolically engineered E. coli. AMB Express 10(1):144. 2020

A277.  Khan, N., de Manuel, M., Peyregne, S., Do, R., Prufer, K., Marques-Bonet, T., Varki, N., Gagneux, P., and Varki, A. Multiple Genomic Events Altering Hominin SIGLEC Biology and Innate Immunity Predated the Common Ancestor of Humans and Archaic Hominins. Genome Biol Evol. 12(7):1040-1050. 2020. Supplemental Materials      Featured in SCIENCE

A276.  Gulati, S., Schoenhofen, I.C., Lindhout-Djukic, T., Schur, M.J., Landig, C.S., Saha, S., Deng, L., Lewis, L.A., Zheng, B., Varki, A., and Ram, S. Therapeutic CMP-Nonulosonates against Multidrug-Resistant Neisseria gonorrhoeae. J Immunol. 204(12):3283-3295. 2020.

A275. Barnard, K.N., Alford-Lawrence, B.K., Buchholz, D.W., Wasik, B.R., LaClair, J.R., Yu, H., Honce, R., Ruhl, S., Pajic, P., Daugherity, E.K., Chen, X., Schultz-Cherry, S.L., Aguilar, H.C., Varki, A., and Parrish, C.R. Modified sialic acids on mucus and erythrocytes inhibit influenza A Virus Hemagglutinin and Neuraminidase functions. J Virol. 94(9):e01567-19. 2020.

A273. Kiser, Z.M., Lizcano, A., Nguyen, J., Becker, G.L., Belcher, J.D., Varki, A.P., and Vercellotti, G.M. Decreased erythrocyte binding of Siglec-9 increases neutrophil activation in sickle cell disease. Blood Cells Mol Dis. 81:102399. 2020.

B202. Varki, A. Roland Schauer (1936-2019): A tribute to “Mr. Sialic Acid”. Glycobiology. 30(3):132-133. 2020.

B201Kawanishi, K., Dhar, C., Varki, A., and Gordts, P.L.S.M. Reply to Soulillou et al.: Difficulties in extrapolating from animal models exemplify unusual human atherosclerosis susceptibility and mechanisms via CMAH lossProc Natl Acad Sci USA. 117(4):1847-48. 2020.


A274. Khan, N., Kim, S.K., Gagneux, P., Dugan, L.L., and Varki, A. Maximum reproductive lifespan correlates with CD33rSIGLEC gene number: Implications for NADPH oxidase-derived reactive oxygen species in aging. FASEB J. 34(2):1928-38. 2019. Supplemental Materials

B199. Laubli, H., and Varki, A. Sialic acid–binding immunoglobulin-like lectins (Siglecs) detect self-associated molecular patterns to regulate immune responses. Cell Mol Life Sci. 77(4):593-605. 2019.

A272. Tsai C.M., Riestra A.M., Ali S.R., Fong J.J., Liu J.Z., Hughes G., Varki A., and Nizet V. Siglec-14 Enhances NLRP3-Inflammasome Activation in MacrophagesJ Innate Immun. 12(4):333-343. 2019.

A271. Zaramela, L.S., Martino, C., Alisson-Silva, F., Rees, S.D., Diaz, S.L., Chuzel, L., Ganatra, M.B., Taron, C.H., Secrest, P., Zuñiga, C., Huang, J., Siegel, D., Chang, G., Varki, A., and Zengler, K. Gut bacteria responding to dietary change encode sialidases that exhibit preference for red meat-associated carbohydrates. Nat Microbiol. 4(12):2082-89. 2019. UCSD Press Release

B200. Varki, A. Did Human Reality Denial Breach the Evolutionary Psychological Barrier of Mortality Salience? A Theory that Can Explain Unusual Features of the Origin and Fate of Our Species. Evolutionary Perspectives on Death. Shackelford, T.K., and V. Zeigler-Hill, editors. Springer International Publishing, Cham. 109-135. 2019.

B198. Saha, S., Siddiqui, S., Khan, N., Verhagen, A., Jiang, W., Springer, S., Ghosh, P., and Varki, A. Controversies about the subcellular localization and mechanisms of action of the Alzheimer's disease–protective CD33 splice variant. Acta Neuropathol. 138(4):671-2. 2019.

B197. Neelamegham, S., Aoki-Kinoshita, K., Bolton, E., Frank, M., Lisacek, F., Lütteke, T., O'Boyle, N., Packer, N., Stanley, P., Toukach, P., Varki, A., Woods, R.J., and SNFG, D.G. Updates to the Symbol Nomenclature For Glycans (SNFG) Guidelines. Glycobiology. 29(9):620-4. 2019. 

A270. Kawanishi, K., Dhar, C., Do, R., Varki, N., Gordts, P.L.S.M., and Varki, A. Human species-specific loss of CMP-N-acetylneuraminic acid hydroxylase enhances atherosclerosis via intrinsic and extrinsic mechanisms. Proc Natl Acad Sci USA. 116(32):16036-45. 2019. UCSD Press Release   Medical News Today   The Scientist   Science Daily

A269. Narimatsu, Y., Joshi, H.J., Nason, R., Van Coillie, J., Karlsson, R., Sun, L., Ye, Z., Chen, Y.H., Schjoldager, K.T., Steentoft, C., Furukawa, S., Bensing, B.A., Sullam, P.M., Thompson, A.J., Paulson, J.C., Büll, C., Adema, G.J., Mandel, U., Hansen, L., Bennett, E.P., Varki, A., Vakhrushev, S.Y., Yang, Z., and Clausen, H. An Atlas of Human Glycosylation Pathways Enables Display of the Human Glycome by Gene Engineered Cells. Mol Cell. 75(2):394-407. 2019. Supplemental Materials

A268. Ng, P.S.K., Day, C.J., Atack, J.M., Hartley-Tassell, L.E., Winter, L.E., Marshanski, T., Padler-Karavani, V., Varki, A., Barenkamp, S.J., Apicella, M.A., and Jennings, M.P. Nontypeable Haemophilus influenzae Has Evolved Preferential Use of N-Acetylneuraminic Acid as a Host Adaptation. mBio. 10(3):e00422-19. 2019.

B196. Dhar, C., Sasmal, A., and Varki, A. From “Serum Sickness” to “Xenosialitis”: Past, Present, and Future Significance of the Non-human Sialic Acid Neu5Gc. Front Immunol. 10:807. 2019.

A267. Uchiyama, S., Sun, J., Fukahori, K., Ando, N., Wu, M., Schwarz, F., Siddiqui, S.S., Varki, A., Marth, J.D., and Nizet, V. Dual actions of group B Streptococcus capsular sialic acid provide resistance to platelet-mediated antimicrobial killing. Proc Natl Acad Sci U S A. 116(15):7465-70. 2019.

A266. Lu, N., Ye, J., Cheng, J., Sasmal, A., Liu, C.C., Yao, W., Yan, J., Khan, N., Yi, W., Varki, A., and Cao, H. Redox-Controlled Site-Specific α2-6-Sialylation. J Am Chem Soc. 141(11):4547-52.  2019.

A265. Landig, C.S., Hazel, A., Kellman, B.P., Fong, J.J., Schwarz, F., Agarwal, S., Varki, N., Massari, P., Lewis, N.E., Ram, S., and Varki, A. Evolution of the exclusively human pathogen Neisseria gonorrhoeae: Human-specific engagement of immunoregulatory Siglecs. Evol Appl. 12(2): 337-49. 2019.


A263. Stanczak, M.A., Siddiqui, S.S., Trefny, M.P., Thommen, D.S., Boligan, K.F., von Gunten, S., Tzankov, A., Tietze, L., Lardinois, D., Heinzelmann-Schwarz, V., von Bergwelt-Baildon, M., Zhang, W., Lenz, H.J., Han, Y., Amos, C.I., Syedbasha, M., Egli, A., Stenner, F., Speiser, D.E., Varki, A., Zippelius, A., and Läubli, H. Self-associated molecular patterns mediate cancer immune evasion by engaging Siglecs on T cells. J Clin Invest. 128(11):4912-23. 2018.

A264. Fong, J.J., Tsai, C.M., Saha, S., Nizet, V., Varki, A., and Bui, J.D. Siglec-7 engagement by GBS β-protein suppresses pyroptotic cell death of natural killer cells. Proc Natl Acad Sci U S A. 115(41):10410-15. 2018.

A262. Okerblom, J., Fletes, W., Patel, H.H., Schenk, S., Varki, A., and Breen, E.C. Human-like Cmah inactivation in mice increases running endurance and decreases muscle fatigability: implications for human evolution. Proc Biol Sci. 285(1886). 2018. UCSD Press Release  Featured in >100 news reports in many languages, including; Science Magazine   Psychology Today    Smithsonian Magazine   Popular Science  Scientific American and Discover Magazine

A259. Ram, S., Gulati, S., Lewis, L.A., Chakraborti, S., Zheng, B., DeOliveira, R.B., Reed, G.W., Cox, A.D., Li, J., St Michael, F., Stupak, J., Su, X.H., Saha, S., Landig, C.S., Varki, A., and Rice, P.A. A Novel Sialylation Site on Neisseria gonorrhoeae Lipooligosaccharide Links Heptose II Lactose Expression with Pathogenicity. Infect Immun. 86(8):e00285-18. 2018.

A258. Meng, C., Sasmal, A., Zhang, Y., Gao, T., Liu, C.C., Khan, N., Varki, A., Wang, F., and Cao, H. Chemoenzymatic Assembly of Mammalian O-Mannose Glycans. Angew Chem Int Ed Engl. 57(29):9003-7. 2018.

A261. Samraj, A.N., Bertrand, K.A., Luben, R., Khedri, Z., Yu, H., Nguyen, D., Gregg, C.J., Diaz, S.L., Sawyer, S., Chen, X., Eliassen, H., Padler-Karavani, V., Wu, K., Khaw, K.T., Willett, W., and Varki, A. Polyclonal human antibodies against glycans bearing red meat-derived non-human sialic acid N-glycolylneuraminic acid are stable, reproducible, complex and vary between individuals: Total antibody levels are associated with colorectal cancer risk. PLoS One. 13(6):e0197464. 2018. Supplemental Material

A260. Alisson-Silva, F., Liu, J.Z., Diaz, S.L., Deng, L., Gareau, M.G., Marchelletta, R., Chen, X., Nizet, V., Varki, N., Barrett, K.E., and Varki, A. Human evolutionary loss of epithelial Neu5Gc expression and species-specific susceptibility to cholera. PLoS Pathog. 14(6):e1007133. 2018.  Supplemental Material     UCSD News Room


A257. Owen, C.D., Tailford, L.E., Monaco, S., Šuligoj, T., Vaux, L., Lallement, R., Khedri, Z., Yu, H., Lecointe, K., Walshaw, J., Tribolo, S., Horrex, M., Bell, A., Chen, X., Taylor, G.L., Varki, A., Angulo, J., and Juge, N. Unravelling the specificity and mechanism of sialic acid recognition by the gut symbiont Ruminococcus gnavus. Nat Commun. 8:2196. 2017.

A256. Hayakawa, T., Khedri, Z., Schwarz, F., Landig, C., Liang, S.Y., Yu, H., Chen, X., Fujito, N.T., Satta, Y., Varki, A., and Angata, T. Coevolution of Siglec-11 and Siglec-16 via gene conversion in primates. BMC Evol Biol. 17:228. 2017. 

B195. Varki, A. New and updated glycoscience-related resources at NCBI. Glycobiology. 27:993. 2017.

A255. Patras, K.A., Coady, A., Olson, J., Ali, S.R., RamachandraRao, S.P., Kumar, S., Varki, A., and Nizet, V. Tamm-Horsfall glycoprotein engages human Siglec-9 to modulate neutrophil activation in the urinary tract. Immunol Cell Biol. 95:960-965. 2017.

A254. Gao, X., Deng, L., Stack, G., Yu, H., Chen, X., Naito-Matsui, Y., Varki, A., and Galán, J.E. Evolution of host adaptation in the Salmonella typhoid toxin. Nat Microbiol. 2:1592-1599. 2017.

A253. Graustein, A.D., Horne, D.J., Fong, J.J., Schwarz, F., Mefford, H.C., Peterson, G.J., Wells, R.D., Musvosvi, M., Shey, M., Hanekom, W.A., Hatherill, M., Scriba, T.J., Thuong, N.T.T., Mai, N.T.H., Caws, M., Bang, N.D., Dunstan, S.J., Thwaites, G.E., Varki, A., Angata, T., and Hawn, T.R. The SIGLEC14 null allele is associated with Mycobacterium tuberculosis- and BCG-induced clinical and immunologic outcomes. Tuberculosis (Edinb). 104:38-45. 2017.

A252. Wasik B, Barnard K, Ossiboff R, Khedri Z, Feng K, Yu, H., Chen, X., Perez, D., Varki, A., Parrish, C. The distribution of O-acetylated sialic acids among target host tissues for influenza virus. mSphere. Sep 6;2(5). pii: e00379-16. 2017.

A251. Bergfeld, A.K., Lawrence, R., Diaz, S.L., Pearce, O.M.T., Ghaderi, D., Gagneux, P., Leakey, M.G., and Varki, A. N-glycolyl groups of nonhuman chondroitin sulfates survive in ancient fossilsProc Natl Acad Sci U S A. 114:E8155-E8164. 2017. UC San Diego Press Release

A250. Siddiqui, S.S., Springer, S.A., Verhagen, A., Sundaramurthy, V., Alisson-Silva, F., Jiang, W., Ghosh, P., and Varki, A. The Alzheimer’s disease-protective CD33 splice variant mediates adaptive loss of function via diversion to an intracellular pool. J Biol Chem. 292:15312-15320. 2017.

B194. Varki, A. Rename the Impact Factor. (Letter) Nature. 548:393, 2017.
B193. Varki, A. Are humans prone to autoimmunity? Implications from evolutionary changes in hominin sialic acid biology. J Autoimmun. 83: 134-142, 2017. (Invited Review). 

B192. Nizet, V., Varki, A., and Aebi, M. Chapter 37: Microbial Lectins: Hemagglutinins, Adhesins, and Toxins. (Book Chapter) IN Essentials of Glycobiology 3rd edition. Edited by Varki, A., et al. Cold Spring Harbor Laboratory Press, N.Y. 2017.

B191. Taylor, M., Drickamer, K., Schnaar, R., Etzler, M., and Varki, A. Chapter 28: Discovery and Classification of Glycan-Binding Proteins. (Book Chapter) IN Essentials of Glycobiology 3rd edition. Edited by Varki, A., et al. Cold Spring Harbor Laboratory Press, N.Y. 2017.

B190. Hart, G.W., and Varki, A. Chapter 60: Future Directions in Glycosciences. (Book Chapter) IN Essentials of Glycobiology 3rd edition. Edited by Varki, A., et al. Cold Spring Harbor Laboratory Press, N.Y. 2017.

B189. Varki, A., Kannagi, R., Toole, B., Stanley, P. Chapter 47: Glycosylation Changes in Cancer. (Book Chapter) IN Essentials of Glycobiology 3rd edition. Edited by Varki, A., et al. Cold Spring Harbor Laboratory Press, N.Y. 2017.

B188. Freeze, H., Kinoshita, T., Varki, A. Chapter 46: Glycans in Acquired Human Diseases. (Book Chapter) IN Essentials of Glycobiology 3rd edition. Edited by Varki, A., et al. Cold Spring Harbor Laboratory Press, N.Y. 2017.

B187. Varki, A., Freeze, H., Baum, L. Chapter 41: Glycans in Systemic Physiology. (Book Chapter) IN Essentials of Glycobiology 3rd edition. Edited by Varki, A., et al. Cold Spring Harbor Laboratory Press, N.Y. 2017.

B186. Varki, A., Schnaar, R., Crocker, P. Chapter 35: I-type Lectins. (Book Chapter) IN Essentials of Glycobiology 3rd edition. Edited by Varki, A., et al. Cold Spring Harbor Laboratory Press, N.Y. 2017.

B185. Varki, A., Kornfeld, S. Chapter 33: P-type Lectins. (Book Chapter) IN Essentials of Glycobiology 3rd edition. Edited by Varki, A., et al. Cold Spring Harbor Laboratory Press, N.Y. 2017.

B184. Varki, A. Aebi, M., Gagneux, P. Chapter 20: Evolution of Glycan Diversity. (Book Chapter) IN Essentials of Glycobiology 3rd edition. Edited by Varki, A., et al. Cold Spring Harbor Laboratory Press, N.Y. 2017.

B183. Varki, A., Schnaar, R., Schauer, R. Chapter 15: Sialic Acids and other Nonulosonic Acids (Book Chapter) IN Essentials of Glycobiology. 3rd edition. Edited by Varki, A., et al. Cold Spring Harbor Laboratory Press, N.Y. 2017.

B182. Varki, A., Gagneux, P. Chapter 7: Biological Functions of Glycans. (Book Chapter) IN Essentials of Glycobiology. 3rd edition. Edited by Varki, A., et al. Cold Spring Harbor Laboratory Press, N.Y. 2017.

B181. Colley, K., Varki, A., Kinoshita, T. Chapter 4: Cellular Organization of Glycosylation. (Book Chapter) IN Essentials of Glycobiology. 3rd edition. Edited by Varki, A., et al. Cold Spring Harbor Laboratory Press, N.Y. 2017.

B180. Varki, A., Kornfeld, S. Chapter 1: Historical Background and Overview. (Book Chapter) IN Essentials of Glycobiology. 3rd edition. Edited by Varki, A., et al. Cold Spring Harbor Laboratory Press, N.Y. 2017.

A249. Lizcano, A., Secundino, I., Döhrmann, S., Corriden, R., Rohena, C., Diaz, S., Ghosh, P., Deng, L., Nizet, V., and Varki, A. Erythrocyte sialoglycoproteins engage Siglec-9 on neutrophils to suppress activation. Blood. 129:3100-3110. 2017.

A248. Dharmadhikari, G., Stolz, K., Hauke, M., Morgan, N.G., Varki, A., de Koning, E., Kelm, S., and Maedler, K. Siglec-7 restores β-cell function and survival and reduces inflammation in pancreatic islets from patients with diabetes. Sci Rep. 7:45319. 2017.

B179. Okerblom, JJ, and Varki, A. Biochemical, Cellular, Physiological and Pathological Consequences of Human loss of N-glycolylneuraminic Acid. (Invited Review) ChemBioChem. 18: 1155-1171. 2017.

A247. Schwarz, F., Landig, C.S., Siddiqui, S., Secundino, I., Olson, J., Varki, N., Nizet, V., and Varki, A. Paired Siglec receptors generate opposite inflammatory responses to a human-specific pathogen. EMBO J. 36:751-760. 2017.

A246. Okerblom, J.J., Schwarz, F., Olson, J., Fletes, W., Ali, S.R., Martin, P.T., Glass, C.K., Nizet, V., and Varki, A. Loss of CMAH during Human Evolution Primed the Monocyte-Macrophage Lineage toward a More Inflammatory and Phagocytic State. J Immunol. 198:2366-2373. 2017.

A245. Naito-Matsui, Y., Davies, L.R., Takematsu, H., Chou, H.H., Tangvoranuntakul, P., Carlin, A.F., Verhagen, A., Heyser, C.J., Yoo, S.W., Choudhury, B., Paton, J.C., Paton, A.W., Varki, N.M., Schnaar, R.L., and Varki, A. Physiological Exploration of the Long Term Evolutionary Selection against Expression of N-Glycolylneuraminic Acid in the Brain. J Biol Chem. 292:2557-2570. 2017. UCSD Press Release

B177. Varki, A. Biological Roles of Glycans. Glycobiology 27: 3-49, 2017.

A244. Siddiqui, S., F. Schwarz, S. Springer, Z. Khedri, H. Yu, L. Deng, A. Verhagen, Y. Naito-Matsui, W. Jiang, D. Kim, J. Zhou, B. Ding, X. Chen, N. Varki, and Varki, A. Studies on the detection, expression, glycosylation, dimerization and ligand binding properties of mouse Siglec-E. J Biol Chem. 1029-1037. 2017.

A243. Khedri, Z., A. Xiao, H. Yu, C.S. Landig, W. Li, S. Diaz, B.R. Wasik, C.R. Parrish, L.P. Wang, Varki, A, and X. Chen. A Chemical Biology Solution to Problems with Studying Biologically Important but Unstable 9-O-Acetyl Sialic Acids. ACS Chem Biol. 214-224. 2017.


B178. Varki, A, and Gagneux, P. Chapter 9: How Different Are Humans and "Great Apes"? A Matrix of Comparative Anthropogeny. In: Tibayrenc, M, Ayala, FJ, eds. On human nature: evolution, diversity, psychology, ethics, politics and religion. Boston, MA: Elsevier. 151–160. 2016.

A242. Hentrich, K., J. Löfling, A. Pathak, V. Nizet, A. Varki, and B. Henriques-Normark. Streptococcus pneumoniae Senses a Human-like Sialic Acid Profile via the Response Regulator CiaR. Cell Host Microbe. 20:307-317. 2016. 

A241. Mahajan, V.S., E. Demissie, H. Mattoo, V. Viswanadham, A. Varki, R. Morris, and S. Pillai. Striking Immune Phenotypes in Gene-Targeted Mice Are Driven by a Copy-Number Variant Originating from a Commercially Available C57BL/6 Strain. Cell Rep. 15:1901-1909. 2016.

B176. Läubli, H., A. Varki, and L. Borsig. Antimetastatic Properties of Low Molecular Weight Heparin. J Clin Oncol. 34:2560-2561. 2016.

B175. Alisson-Silva, F., Kawanishi, K., and Varki, A. Human risk of diseases associated with red meat intake: analysis of current theories and proposed role for metabolic incorporation of a non-human sialic acid. Mol Aspects Med. 51:16-30, 2016. 

B174. Varki, A. Why are there no persisting hybrids of humans with Denisovans, Neanderthals, or anyone else? Proc Natl Acad Sci U.S.A. 113: E2354, 2016.

B173. Springer, S.A., Schwarz, F., Altheide, T.K., Varki, N.M., Varki, A., Gagneux, P. Reply to Liu and Jiang: Maintenance of post reproductive cognitive capacity by inclusive fitness. Proc Natl Acad Sci USA. 113, 12, E1591-2. DOI=10.1073/pnas.1601419113. 2016.

A240. Bensing, B. A., Khedri, Z., Deng, L., Yu, H., Prakobphol, A., Fisher, S. J., Chen, X., Iverson, T. M., Varki, A., and Sullam, P. M. Novel aspects of sialoglycan recognition by the Siglec-like domains of streptococcal SRR glycoproteins. Glycobiology. 10.1093/glycob/cww042. 2016.

A234. Secundino, I., Lizcano, A., Roupé, K.M., Wang, X., Cole, J.N., Olson, J., Ali, S.R., Dahesh, S., Amayreh, L.K., Henningham, A., Varki, A. and Nizet, V. Host and pathogen hyaluronan signal through human siglec-9 to suppress neutrophil activation. J. Mol. Med. 94(2):219-33. doi: 10.1007/s00109-015-1341-8. 2016.

B172. Agre P, Bertozzi C, Bissell M, Campbell KP, Cummings RD, Desai UR, Estes M, Flotte T, Fogleman G, Gage F, Ginsburg D, Gordon JI, Hart G, Hascall V, Kiessling L, Kornfeld S, Lowe J, Magnani J, Mahal LK, Medzhitov R, Roberts RJ, Sackstein R, Sarkar R, Schnaar R, Schwartz N, Varki A, Walt D, Weissman I. Training the next generation of biomedical investigators in glycosciences. J Clin Invest. 126(2):405-8. doi:10.1172/JCI85905. 2016.

A238. Song, H, Qi, J., Khedri, Z., Diaz, S., Yu, H., Chen, X., Varki, A., Shi, Y., Gao, G.F. An Open Receptor-Binding Cavity of Hemagglutinin-Esterase-Fusion Glycoprotein from Newly-Identified Influenza D Virus: Basis for Its Broad Cell Tropism. PLoS Pathog. 12(1):e1005411. doi:10.1371/journal.ppat.1005411. 2016.

A239. Tang, X.X., Ostedgaard, L.S., Hoegger, M.J., Moninger, T.O., Karp, P.H., McMenimen, J.D., Choudhury, B., Varki, A., Stoltz, D.A., Welsh, M.J. Acidic pH increases airway surface liquid viscosity in cystic fibrosis. J Clin Invest. 126(3):879-91. 2016.


B171. Varki, A., Cummings, R.D., Aebi, M., Packer, N.H., Seeberger, P.H., Esko, J.D., Stanley, P., Hart, G., Darvill, A., Kinoshita, T., Prestegard, J.J., Schnaar, R.L., Freeze, H.H., Martin, J.D., Bertozzi, C.R., Etzler, M.S., Frank, M., Vliegenthart, J.F., Lütteke, T., Perez, S., Bolton, E., Rudd, P., Paulson, J., Kanehisa, M., Toukach, P., Aoki-Kinoshita, K.F., Dell, A., Narimatsu, H., York, W., Taniguchi, N., and Kornfeld, S. Symbol Nomenclature for Graphical Representations of Glycans. Glycobiology. 25(12):1323-4. Doi: 10.1093/glycob/cwv091. 2015.

A237. Gulati, S., Schoenhofen, I.C., Whitfield, D., Cox, A.D., Li, J., St. Michael, F., Vinogradov, E., Stupak, J., Zheng, B., Ohnishi, M., Unemo, M., Lewis, L.A., Taylor, R.E., Landig, C.S., Diaz, S., Reed, G., Varki, A., Rice, P.A., and Ram, S. Utilizing CMP-sialic acid analogs to unravel Neisseria gonorrhoeae lipooligosaccharide-mediated complement resistance and design novel therapeutics. PloS Pathog. DOI: 10.1371/journal.ppat.1005290. 2015.

A236. Schwarz, F., Springer, S., Altheide, T., Varki, N., Gagneux, P., and Varki, A. Human-specific Derived Alleles of CD33 and Other Genes Protect Against Post-Reproductive Cognitive Decline. Proc. Nat'l. Acad. Sci. USA doi: 10.1073/pnas.1517951112. 2015.  UCSD Press Release

A235. Fong, J.J., Sreedhara, K., Deng, L., Varki, N., Angata, T., Liu, Q., Nizet, V. and Varki, A. Immunomodulatory Activity of Extracellular Hsp70 Mediated Via Paired Receptors Siglec-5 and Siglec-14. EMBO J. 34:2775-2788. 2015.

A233. Langereis, M.S., Bakkers, M.J.G., Deng, L., Padler-Karavani, V., Vervoort, S.J., Hulswit, R.J.G., van Vliet, A.L.W., Gerwig, G.J., de Poot, S.A.H., Boot, W., van Ederen, A.M., Heesters, B.A., van der Loos, C.M., van Kuppeveld, F.J.M., Yu, H., Hulzinga, E.G., Chen, A., Varki, A., Kamerling, J.P., and de Groot, R.J. Complexity and Diversity of the Mammalian Sialome Revealed by Nidovirus Virolectins. Cell Reports. 2015.

A232. Feng, KH, Gonzalez, G, Deng, L, Yu, H, Tse, VL, Huang, L, Huang, K, Wasik, BR, Zhou, B, Wentworth, DE, Holmes, EC, Chen, X, Varki, A, Murcia, PR, Parrish, CR. Equine and canine influenza H3N8 viruses show minimal biological differences despite phylogenetic divergence. J Virol. 89:6860-6873. 2015.

A231. Schwarz, F., Pearce, O.M.T, Wang, X., Samraj, A.N., Läubli, H., Garcia, J.O., Lin, H., Fu, X., Garcia-Bingman, A., Secrest, P., Romanoski, C.E., Heyser, C., Glass, C.K., Hazen, S.L., Varki, N., Varki, A., Gagneux, P. Siglec receptors impact mammalian lifespan by modulating oxidative stress. eLife. 4:e06184. DOI: 10.7554/eLife.06184. 2015. UCSD Press Release

B170. Varki, N.M. and Varki, A. On the Apparent Rarity of Epithelial Cancers in Captive Chimpanzees. Philos Trans R Soc Lond B Biol Sci. 370:20140225. 2015.

A230. Sumida, M., Hane, M., Yabe, U., Shimoda, Y., Pearce, O.M.T., Kiso, M., Miyagi, T., Sawada, M., Varki, A., Kitajima, K. and Sato, C. Rapid trimming of cell surface polysialic acid (PolySia) by exovesicular sialidase triggers release of preexisting surface neurotrophin. J Biol Chem. 290:13202-13214. 2015.

B169. Pearce, O.M.T., Samraj, A.N., Läubli, H., A., Varki, N., Varki, A. Reply to Mackenzie: A comparison of Neu5Gc and alpha-gal xenoantigens. (Invited Letter) Proc Natl Acad Sci U S A. 112:E1405. 2015. 

A229. Samraj, A.N., Pearce, O.M.T., Läubli, H., A., Crittenden, A.N., Bergfeld, A.K., Banda, K., Gregg, C.J., Bingman A.E., Secrest, P., Diaz, S.L., Varki, N., Varki, A. A Red Meat-Derived Glycan Promotes Inflammation and Cancer Progression. Proc Natl Acad Sci U S A. 112:542-7. 2015. UCSD Press Release


A228. Deng, L., Bensing, B.A., Thamadilok, S., Yu, H., Lau, K., Chen, X., Ruhl, S., Sullam, P.A., Varki, A. Oral Streptococci Utilize a Siglec-like Domain of Serine-Rich Repeat Adhesins to Preferentially Target Platelet Sialoglycans in Human Blood. PloS Pathog. 10(12):e1004540. 2014. 

A227. Deng, L., Song, J., Gao, Xiang., Wang, J., Yu, H., Chen, X., Varki, N., Naito-Matsui, Y., Galán, J.E., Varki, A. Host adaptation of a bacterial toxin from the human pathogen Salmonella Typhi. Cell. 159:1290-1299. 2014. UCSD Press Release

A226. Läubli, H., da Silva, F.A., Stanczak, M.A., Siddiqui, S.S., Deng, L., Verhagen, A., Varki, N., , Varki, A. Lectin galactoside-binding soluble 3 binding protein (LGALS3BP) Is a tumor-associated immunomodulatory ligand for CD33-related Siglecs. J Biol Chem., 289:33481-33491. 2014.

A225. Läubli, H., Pearce, O.M.T., Schwarz, F., Siddiqui, S.S., Deng, L., Stanczak, M., Deng, L., Verhagen, A., Secrest, P., Lusk, C., Schwartz, A.G., Varki, N., Bui, J.D., Varki, A. Engagement of Myelomonocytic Siglecs by Tumor-associated Ligands Modulates the Innate Immune Response to Cancer. Proc Natl Acad Sci U.S.A., 111:14211-14216. 2014. UCSD Press Release

B168. Pearce, O.M., Läubli, H., Bui, J., Varki, A. Hormesis in cancer immunotherapy: Does the quantity of an immune reactant matter? OncoImmunology, 3:e29312. 2014.

B167. Schwarz, F., Fong, J.J., and Varki, A. Human-Specific Evolutionary Changes in the Biology of Siglecs. (Invited Review) Adv. Exp. Med. Biol., 842:1-16. 2014.

A224. Ali, S.R., Fong, J.J., Carlin, A.F., Busch, T.D., Linden, R., Angata, T., Areschoug, T., Parast, M., Varki, N., Murray, J., Nizet, V., and Varki, A. Siglec-5 and Siglec-14 are polymorphic paired receptors that modulate neutrophil and amnion signaling responses to group B Streptococcus. J. Exp. Med. 211:1231-1242. 2014.

A223. Chang, Y.-C., Olson, J., Louie, A., Crocker, P.R., Varki, A., Nizet, V. Role of macrophage sialoadhesin in host defense against the sialylated pathogen group B Streptococcus. J. Mol. Med. 92:951-959. 2014.

B165. Angata, T. and Varki, A. Siglec Interactions with Pathogens. (Invited Essay). IN Glycoscience: Biology and Medicine, edited by Endo, T., Seeberger, P.H., Hart, G.W., Wong, C.-H., and Taniguchi, N. Springer, Tokyo, pp 633-642. 2014.

B166. Bergfeld A., Samraj, A. and Varki, A. Metabolism of N-glycolylneuraminic Acid in Human and Non-Human Cells, and Potential Relationships to Human Disease. (Invited Essay) IN Glycoscience: Biology and Medicine, edited by Endo, T., Seeberger, P.H., Hart, G.W., Wong, C.-H., and Taniguchi, N. Springer, Tokyo, pp1-8. 2014.

A221. Muto, J., Morioka, Y., Yamasaki, K., Kim, M., Garcia, A., Carlin, A.F., Varki, A., Gallo, R.L. Hyaluronan digestion controls DC migration from the skin. J Clin Invest. 124:1309-19. 2014.

A222. Pearce, O.M., Laubli, H., Verhagen, A., Secrest, P., Zhang, J., Varki, N.M., Crocker, P., Bui, J., and Varki, A. Inverse Hormesis of Cancer Growth Mediated by Narrow Ranges of Tumor-Directed Antibodies. Proc Natl Acad Sci U.S.A. 111:5998-6003. 2014.

A220. Martin, P.T., Golden, B., Okerblom, J., Camboni, M., Chandrasekharan, K., Xu, R., Varki, A., Flanigan, K.M., Kornegay, J.N. A Comparative Study of N-glycolylneuraminic Acid (Neu5Gc) and Cytotoxic T Cell (CT) Carbohydrate Expression in Normal and Dystrophin-Deficient Dog and Human Skeletal Muscle. PLoS One. 9(2):e88226. 2014.

B163. Samraj, A., Läubli, H., Varki, N., Varki, A. Involvement of a non-human sialic acid in human cancer. Front. Oncol. 4:33. 2014.

B162. Cohen, M., Varki, A. Modulation of glycan recognition by clustered saccharide patches. (Invited Review) Cell Mol Biol. 308:75-125. 2014.

A219. Chang, Y.-C., Olson, J.; Beasley, F.C., Tung, C., Zhang, J., Crocker, P.R., Varki, A., Nizet, V. Group B Streptococcus Engages an Inhibitory Siglec through Sialic Acid Mimicry to Blunt Innate Immune and Inflammatory Responses In Vivo. PLoS Pathog. 10:e1003846. 2014.

A218. Padler-Karavani, V., Hurtado-Ziola, N., Chang, Y.-C., Sonnenburg, J., Ronaghy, A., Yu, H., Verhagen, A., Nizet, V., Chen, X., Varki, N., Varki, A., Angata, T. Rapid Evolution of Binding Specificities and Expression Patterns of Inhibitory CD33-related Siglecs in PrimatesFASEB J. 28:1280-93. 2014. UCSD Press Release

B164. Bergfeld, A.K., Varki, A. Cytidine Monophospho-N-acetylneuraminic Acid Hydroxylase (CMAH). IN: Handbook of Glycosyltransferases and Related Genes, Taniguchi, N., Honke, K., Fukuda, M., Narimatsu, H., Yamaguchi, Y., Angata, T. (eds).  Springer, New York. 2014.


A217. Ogata, S., Shimizu, C., Franco, A., Touma, R., Kanegaye, J.T., Choudhury, B.P., Naidu, N.N., Kanda, Y., Hoang, L.T., Hibberd, M.L., Tremoulet, A.H., Varki, A., Burns, J.C. Treatment response in kawasaki disease is associated with sialylation levels of endogenous but not therapeutic intravenous immunoglobulin G. PLoS One. 8(12):e81448. 2013. 

B161. Varki, A. Thought Experiment: Dating the Origin of Us.  The Scientist 27(11):28-29. 2013.

A216. Scobie, L., Padler-Karavani, V., Le Bas-Bernardet, S., Crossan, C., Blaha, J., Matouskova, M., Hector, R.D., Cozzi, E., Vanhove, B., Charreau, B., Blancho, G., Bourdais, L., Tallacchini, M., Ribes, J.M., Yu, H., Chen, X., Kracikova, J., Broz, L., Hejnar, J., Vesely, P., Takeuchi, Y., Varki, A., Soulillou, J.P. Long-Term IgG Response to Porcine Neu5Gc Antigens without Transmission of PERV in Burn Patients Treated with Porcine Skin Xenografts. J Immunol., 191:2907-2915. 2013.

B160. Deng, L., Chen, X., Varki, A. Exploration of Sialic Acid Diversity and Biology Using Sialoglycan Microarrays. Biopolymers. 99:650-65. 2013. 

A215. Suzukawa, M., Miller, M., Rosenthal, P., Cho, J.Y., Doherty, T.A., Varki, A., Broide, D. Sialyltransferase ST3Gal-III Regulates Siglec-F Ligand Formation and Eosinophilic Lung Inflammation in Mice. J Immunol. 190:5939-5948. 2013. 

A214. Buchlis, G., Odorizzi, P., Soto, P.C., Pearce, O.M.T., Hui, D.J., Jordan, M.S., Varki, A. Wherry, E.J., High, K.A. Enhanced T Cell Function in a Mouse Model of Human Glycosylation. J Immunol. 191:228-237. 2013. 

A213. Martin, P.T., Camboni, M., Xu, R., Golden, B., Chandrasekharan, K., Wang, C.-M., Varki, A., Janssen, P.M.L. N-glycolylneuraminic acid deficiency worsens cardiac and skeletal muscle pathophysiology in α-sarcoglycan-deficient mice. Glycobiology. 23:833-843. 2013.

B158. Varki, A. Account for the 'dark matter' of biology. Nature. 497:565, 2013.

B159. Davies, L.R.L., Varki, A. Why is N-Glycolylneuraminic Acid Rare in the Vertebrate Brain? Top. Curr. Chem. doi: 10.1007/128_2013_419, 2013.

A212. Kikkeri, R., Padler-Karavani, V., Diaz, S., Verhagen, A., Yu, H., Cao, H., Langereis, M.J., De Groot, R.J., Chen, X., Varki, A. Quantum Dot-Nanometal Surface Energy Transfer Based Biosensing of Sialic Compositions and Linkages in biological Samples. Anal Chem. 85:3864-70. 2013.

A211. Angata, T., Ishii, T., Motegi, T., Oka, R., Taylor, R.E., Campos Soto, P., Chang, Y-C.,Secundino, I., Gao, C-X., Ohtsubo, K., Kitazume, S., Nizet, V., Varki, A., Gemma, A., Kida, K., Taniguchi, N. Loss of Siglec-14 reduces the risk of chronic obstructive pulmonary disease exacerbation. Cell. Mol. Life Sci., 70:3199-210. 2013.

A210. Löfling, J., Lyi, S.M., Parrish, C.R., Varki, A. Canine and Feline Parvoviruses Preferentially Recognize the Non-Human Cell Surface Sialic Acid N-glycolylneuraminic acid. Virology, 440:89-96. 2013.

A209. Padler-Karavani, V., Tremoulet, A.H., Yu, H., Chen, X., Burns, J.C., Varki, A. A Simple Method for Assessment of Human Anti-Neu5Gc Antibodies Applied to Kawasaki Disease. PLoS ONE, 8:e58443. 2013.

A208. Soto, P.C., Karris, M.Y., Spina, C.A., Richman, D.D., Varki, A. Cell-intrinsic mechanism involving Siglec-5 associated with divergent outcomes of HIV-1 infection in human and chimpanzee CD4 T cells. J Mol Med. 91:261-70. 2013.


B157. O'Bleness, M.O., Searles, V.B., Varki, A., Gagneux, P., Sikela, J.M. Evolution of genetic and genomic features unique to the human lineage. Nat. Rev. Genet., 13:853-66. 2012.

A207. Davies, L.R., Pearce, O.M., Tessier, M.B., Assar, S., Smutova, V., Pajunen, M., Sumida, M., Sato, C., Kitajima, K., Finne, J., Gagneux, P., Pshezhetsky, A., Woods, R., and Varki, A. Metabolism of Vertebrate Amino Sugars with N-glycolyl Groups: Resistance of α2-8-linked N-glycolylneuraminic Acid to Enzymatic Cleavage. J. Biol. Chem., 287:28917-28931. 2012.

A206. Bergfeld, A.K., Pearce, O.M., Diaz, S.L., Lawrence, R., Vocadlo, D.J., Choudhury, B., Esko, J.D., and Varki, A. Metabolism of Vertebrate Amino Sugars with N-glycolyl Groups: Incorporation of N-glycolylhexosamines into mammalian glycans by feeding N-glycolylgalactosamine. J. Biol. Chem., 287:28898-28916, 2012.

A205. Macauley, M.S., Chan, J., Zandberg, W.F., He, Y., Whitworth, G.E., Stubbs, K.A., Yuzwa, S.A., Bennet, A.J., Varki, A., Davies, G.J., and Vocadlo, D.J. Metabolism of vertebrate amino sugars with N-glycolyl groups: intracellular β-O-linked N-glycolylglucosamine (GlcNGc), UDP-GlcNGc, and the biochemical and structural rationale for the substrate tolerance of β-O-linked β-N-acetylglucosaminidase. J. Biol. Chem., 287:28882-28897, 2012.

A204. Bergfeld, A.K. Pearce, O.M., Diaz, S.L., Pham, T., and Varki, A. Metabolism of Vertebrate Amino Sugars with N-glycolyl Groups: Elucidating the intracellular fate of the non-human sialic acid N-glycolylneuraminic acid. J. Biol. Chem., 287:28865-28881, 2012.

A203. Banda, K., Gregg, C.J., Chow, R., Varki, N.M., and Varki, A. Metabolism of Vertebrate Amino Sugars with N-glycolyl Groups: Mechanisms underlying gastrointestinal Incorporation of the non-human sialic acid Xeno-autoantigen, N-glycolylneuraminic acid.  J. Biol. Chem., 287:28852-28864, 2012.

A202. Padler-Karavani, V., Song, X., Yu, H., Hurtado-Ziola, N., Huang, S., Muthana, S., Chokhawala, H.A., Cheng, J., Verhagen, A., Langereis, M.A., Kleene, R., Schachner, M., de Groot, R.J., Lasanajak, Y., Matsuda, H., Schwab, R., Chen, X., Smith, D.F., Cummings, R.D., and Varki, A. Cross-comparison of protein recognition of sialic acid diversity on two novel sialoglycan microarrays. J. Biol. Chem., 287:22593-22608, 2012.

A201. Wang, X., Mitra, N., Secundino, I., Banda, K., Cruz, P., Padler-Karavani, V., Verhagen, A., Reid, C., Lari, M., Rizzi, E., Balsamo, C., Corti, G., De Bellis, G., Longo, L., NISC Comparative Sequencing Program, Beggs, W.,
Caramelli, D., Tishkoff, S. A., Hayakawa, T., Green, E. D., Mullikin, J. C., Nizet, V., Bui, J., and Varki, A. Specific Inactivation of Two Immunomodulatory SIGLEC Genes During Human Evolution. Proc. Natl. Acad. Sci. U.S.A., 109:9935-9940, 2012. UCSD Press Release.

B156. Varki, A. Siglecs. In: Lennarz, W.J. and Lane, M.D. (eds.) Encyclopedia of Biological Chemistry, Vol. 4. Academic Press,167-171, 2013.

B155. Varki A. Nothing in Medicine Makes Sense, Except in the Light of Evolution (Invited Commentary). J. Mol. Med., 90:481-494, 2012.

B154. Varki, A. Invited Preface IN: Post-genome Biology in Primates. Edited by Hirai, H., Imani, H., and Go, Y. (Series Editors: Matsuzawa, T., and Yamagiwa, J.) Springer, Japan, 2012.

B153. Hayakawa, T. and Varki, A. Chapter 8: Human-specific Chances in Sialic Acid Biology, IN: Post-genome Biology of Primates. Edited by Hirai, H., Imani, H., and Go, Y. (Series Editors: Matsuzawa, T., and Yamagiwa, J.) Springer, Japan, 2012.

B152. Varki, A., and Gagneux, P. Multifarious Roles of Sialic Acids in Immunity (Invited Review). Ann. N.Y. Acad. Sci., 1253:16-36, 2012.

B151. Ghaderi, D., Zhang, M., Hurtado-Ziola, N., and Varki, A. Production platforms for biotherapeutic glycoproteins. Occurrence, impact, and challenges of non-human sialylation.  Biotechnol. Genet. Eng. Rev., 28:147-175, 2012.

A200. Wang, X., Mitra, N., Cruz, P., Deng, L., NISC Comparative Sequencing Program, Varki, N., Angata, T., Green, E.D., Mullikin, J., Hayakawa, T., and Varki, A. Evolution of Siglec-11 and Siglec-16 Genes in Hominins. Mol. Biol. Evol., 29:2073-2086, 2012.

A199. Lu, Q., Padler-Karavani, V., Yu, H., Chen, X., Wu, S.L., Varki, A., and Hancock, W.S. LC-MS Analysis of Polyclonal Human Anti-Neu5Gc Xeno-Autoantibodies Immunoglobulin G Subclass and Partial Sequence Using Multistep Intravenous Immunoglobulin Affinity Purification and Multienzymatic Digestion. Anal. Chem., 84:2761-2768, 2012.

A198. Chang, Y.-C., Uchiyama, S., Varki, A. and Nizet, V. Leukocyte Inflammatory Responses Provoked by Pneumococcal Sialidase.  mBio, 3:e00220-11, 2012.


A197. Ghaderi, D., Springer, S. A., Ma, F., Cohen, M., Secrest, P., Taylor, R.E., Varki, A., and Gagneux, P. Sexual selection by female immunity against paternal antigens can fix loss of function allelesProc. Natl. Acad. Sci. U.S.A., 108:17743-17748, 2011. UCSD Press Release.

A196. Rubinstein, E., Cho J.Y., Rosenthal, P., Chao, J., Miller, M., Pham, A., Aceves, S.S., Varki, A., and Broide, D.H. Siglec-F inhibition reduces esophageal eosinophilia and angiogenesis in a mouse model of eosinophilic esophagitis. J. Pediatr. Gastroenterol. Nutr., 53:409-416, 2011.

A195. Shao, B., Wahrenbrock, M. G., Yao, L., David, T., Coughlin, S.R., Xia, L., Varki, A., and McEver, R. P. Carcinoma mucins trigger reciprocal activation of platelets and neutrophils in a murine model of Trousseau syndrome. Blood, 118:4015-4023, 2011.

A194. Nordström, T., Movert, E., Olin, A. I., Ali, S.R., Nizet, V., Varki, A.,and Areschoug, T. Human Siglec-5 inhibitory receptor and immunoglobulin A (IgA) have separate binding sites in streptococcal beta protein. J. Biol. Chem., 286:33981-33991, 2011.

A193. Mitra, N., Banda, K., Altheide, T., Schaffer, L., Johnson-Pais, T.L., Beuten, J., Leach, R.J., Angata, T., Varki, N., and Varki, A. SIGLEC12, a human-specific segregating (pseudo)gene, encodes a signaling molecule expressed in prostate carcinomas. J. Biol. Chem., 286:23003-23011, 2011.

A192. Wang, X., Chow, R., Deng, L., Anderson, D., Weidner, N., Godwin, A.K., Bewtra, C., Zlotnik, A., Bui, J., Varki, A., and Varki, N. Expression of Siglec-11 by Human and Chimpanzee Ovarian Stromal Cells, with Uniquely Human Ligands: Implications for Human Ovarian Physiology and Pathology. Glycobiology, 21:1038-1048, 2011.

A191. Padler-Karavani, V., Hurtado-Ziola, N., Pu, M., Yu, H., Huang, S., Muthana, S., Chokhawala, H.A., Cao, H., Secrest, P., Friedmann-Morvinski, D., Singer, O., Ghaderi, D., Verma, I.M., Liu, Y.-T., Messer, K., Chen, X., Varki, A., and Schwab, R. Human Xeno-autoantibodies Against a Non-human Sialic Acid Serve As Novel Serum Biomarkers and Immunotherapeutics in Cancer. Cancer Res., 71:3352-3363, 2011.

B150. Varki, A. Since there are PAMPs and DAMPs, there must be SAMPs? Glycan "self-associated molecular patterns" dampen innate immunity, but pathogens can mimic them. Glycobiology, 21:1121-1124, 2011.

A190. Kavaler, S., Morinaga, H., Jih, A., Fan, W., Hedlund, M., Varki, A., and Kim, J.J. Pancreatic beta cell failure in obese mice with human-like CMP-Neu5Ac hydroxylase deficiency. FASEB J., 25:1887-1893, 2011.

B149. Varki, A. Evolutionary Forces Shaping The Golgi Glycosylation Machinery: Why Cell Surface Glycans are Universal to Living Cells. CSH Perspectives in Biology (Invited Review), doi: 10.1101/cshperspect.a005462, 2011.

B148. Padler-Karavani, V., and Varki, A. Potential Impact of the Non-human Sialic Acid N-glycolylneuraminic Acid on Transplant Rejection Risk (Invited Commentary). Xenotransplantation, 18:1-5, 2011.

B147. Varki, N.M., Strobert, E., Dick Jr., E.J., Benirschke, K. and Varki, A. Biomedical Differences Between Humans and Nonhuman Hominids: Potential Roles for Uniquely Human Aspects of Sialic Acid Biology (Invited Review). Annu. Rev. Pathol., 6:365-393, 2011.


A189. Cho, J.Y., Song, D.J., Pham, A., Rosenthal, P., Miller, M., Dayan, S., Doherty, T.A., Varki, A., and Broide, D.H. Chronic OVA allergen challenged Siglec-F deficient mice have increased mucus, remodeling, and epithelial Siglec-F ligands which are up-regulated by IL-4 and IL-13. Respir. Res., 11:154, 2010.

B146. Varki, A. Fate of ‘Retired’ Research Chimps (Correspondence). Nature, 467:1047, 2010.

A188. Chandrasekharan, K., Yoon, J.H., Xu, Y., deVries, S., Camboni, M., Janssen, P.M., Varki., A., and Martin, P.T. A Human-Specific Deletion in Mouse Cmah Increases Disease Severity in the mdx Model of Duchenne Muscular Dystrophy. Sci. Transl. Med., 2:42ra54, 2010. 

A187. Ghaderi, D., Taylor, R.E., Padler-Karavani, V., Diaz, S., and Varki, A. Implications of the presence of N-glycolylneuraminic acid in recombinant therapeutic glycoproteins.  Nat. Biotechnol., 28:863-867, 2010. 

B145. Varki, A. Introduction of Stuart Kornfeld. J. Clin. Invest., 120:2635-2638, 2010.

A186. Taylor, R.E., Gregg, C.J., Padler-Karavani, V., Ghaderi, D., Yu, H., Huang, S., Sorensen, R.U., Chen, X., Inostroza, J., Nizet, V., and Varki, A. Novel mechanism for the generation of human xeno-autoantibodies against the nonhuman sialic acid N-glycolylneuraminic acid. J. Exp. Med., 207:1637-1646, 2010.

A185. Surolia, I., Pirnie, S.P., Chellappa, V., Taylor, K.N., Cariappa, A., Moya, J., Liu, H., Bell, D.W., Driscoll, D.R., Diederichs, S., Haider, K., Netravali, I., Le, S., Elia, R., Dow, E., Lee, A., Freudenberg, J., De Jager, P.L., Chretien, Y., Varki, A., MacDonald, M.E., Gillis, T., Behrens, T.W., Bloch, D., Collier, D., Korzenik, J., Podolsky, D.K., Hafler, D., Murali, M., Sands, B., Stone, J.H., Gregersen, P.K., and Pillai, S. Functionally defective germline variants of sialic acid acetylesterase in autoimmunity. Nature, 466:243-247, 2010.

A184. Pearce, O.M.T. and Varki, A. Chemo-enzymatic synthesis of the carbohydrate antigen N- glycolylneuraminic acid from glucose. Carbohydr. Res., 345:1225–1229, 2010.

B144. Cohen, M. and Varki, A. The Sialome – Far More Than the Sum of its Parts (Invited Review). OMICS, 14:455-464, 2010. 

A183. Weiman, S., Uchiyama, S., Lin, F.Y., Chaffin, D., Varki, A., Nizet, V., and Lewis, A.L. O-Acetylation of Sialic Acid on Group B Streptococcus Inhibits Neutrophil Suppression and Virulence. Biochem. J., 428:163–168, 2010.

B143. Varki A. Colloquium Paper: Uniquely Human Evolution of Sialic Acid Genetics and Biology. Proc. Natl. Acad. Sci. U.S.A., 107 (Suppl 2): 8939-8946, 2010.

A182. Soto, P.C., Stein, L.L., Hurtado-Ziola, N., Hedrick, S.M., and Varki A. Relative Over-reactivity of Human versus Chimpanzee Lymphocytes: Implications for the Human Diseases Associated with Immune Activation. J. Immunol., 184:4185-4195, 2010.

B142. Chen, X., and Varki, A. Advances in the Biology and Chemistry of Sialic Acids (Invited Review). ACS Chem. Biol., 5:163-176, 2010.


B141. Varki, A., Cummings, R.D., Esko, J.D., Freeze, H.H., Stanley, P., Marth, J.D., Bertozzi, C.R., Hart, G.W., and Etzler, M.E. Symbol Nomenclature for Glycan Representation (Viewpoint Article). Proteomics, 9: 5398–5399, 2009.

A181. Song, D.J., Cho, J.Y. Lee, S.Y., Miller, M., Rosenthal, P., Soroosh, P., Croft, M., Zhang, M., Varki, A., and Broide, D.H. Anti-Siglec-F Antibody Reduces Allergen-Induced Eosinophilic Inflammation and Airway Remodeling.  J. Immunol., 183:5333-5341, 2009.

A180. Pham, T.*, Gregg, C.J.*, Karp, F., Chow, R., Padler-Karavani, V., Cao, H., Chen, X., V., Witztum, J.L., Varki, N.M. and Varki, A. Evidence for a Novel Human-specific Xeno-Auto-Antibody Response Against Vascular Endothelium.  Blood, 114:5225-5235, 2009. * = The contributions of the first two authors are equal.

B140. Varki, A and Gagneux, P.G. Human-Specific Evolution of Sialic Acid Targets: Explaining the Malignant Malaria Mystery?  Proc. Natl. Acad. Sci. U.S.A., 106:14739-14740, 2009.

B137. Varki, A. Evolutionary Perspectives on the Origin of Diseases IN, Current Trends in Sciences (Invited Book Chapter), Ed. N. Mukunda. Indian Academy of Sciences, Bangalore, India, pp. 395-402, 2009. 

A179. Cohen, M., Hurtado-Ziola, N., and Varki, A. ABO Blood Group Glycans Modulate Sialic Acid Recognition on Erythrocytes.  Blood, 114:3668-3676, 2009.

A178. Weiman, S., Dahesh, S., Carlin, A.F., Varki, A., Nizet, V., and Lewis, A.L. Genetic and Biochemical Modulation of Sialic Acid O-Acetylation on Group B Streptococcus: Phenotypic and Functional Impact.  Glycobiology, 19:1204-1213, 2009.

B139. Varki, A. Human Uniqueness and the Denial of Death (Invited Correspondence).  Nature, 460:684, 2009.

B138. Diaz, S., and Varki, A. Metabolic Radiolabeling of Animal Cell Glycoconjugates. IN Curr. Protoc. Mol. Biol., 87:17.4.1-17.4.15, 2009.

A177. Lewis, A.L., Desa, N., Hansen, E.E., Knirel, Y., Gordon, J.I., Gagneux, P., Nizet, V., and Varki, A. Innovations in Host and Microbial Sialic Acid Biosynthesis Revealed by Phylogenomic Prediction of Nonulosonic Acid Structure.  Proc. Natl. Acad. Sci. U.S.A., 106:13552-13557, 2009.

A176. Carlin, A.F., Chang, Y.C., Areschoug, T., Lindahl, G., Hurtado-Ziola, N., King, C.C., Varki, A. and Nizet, V. Group B Streptococcus Suppression of Phagocyte Functions by Protein-Mediated Engagement of Human Siglec-5.  J. Exp. Med., 206:1691-1699, 2009.

B136. Varki, A. Natural Ligands for CD33-related Siglecs?  Glycobiology, 19:810-812, 2009.

B135. Varki, A., Freeze, H.H., and Manzi, A.E. Overview of Glycoconjugate Analysis. IN Curr. Protoc. in Protein Sci., 57:12.1.1-12.1.10, 2009.

B134. Varki, A., Freeze, H.H., and Manzi, A.E. Preparation and Analysis of Glycoconjugates. IN Curr. Protoc. Mol. Biol., 88:17.0.1-17.0.12, 2009.

B133. Löfling, J.C., Paton, A.W., Varki, N.M., Paton, J.C., and Varki, A. A Dietary Non-Human Sialic Acid May Facilitate Hemolytic-Uremic Syndrome.  Kidney Int., 76:140-144, 2009.

B132. Varki, A, Varki, N.M., and Borsig, L. Molecular Basis of Metastasis (Correspondence).  N. Engl. J. Med., 360:1678-1680, 2009.

A175. Carlin, A.F., Uchiyama, S., Chang, Y.C., Lewis, A.L., Nizet, V., and Varki, A Molecular Mimicry of Host Sialylated Glycans Allows A Bacterial Pathogen to Engage Neutrophil Siglec-9 and Dampen the Innate Immune Response.  Blood, 113:3333-3336, 2009. 

A174. Song, D.J., Cho, J.Y., Miller, M., Strangman, W., Zhang, M., Varki, A. and Broide, D.H. Anti-Siglec-F Antibody Inhibits Oral Egg Allergen Induced Intestinal Eosinophilic Inflammation in a Mouse Model. Clin. Immunol., 131:157-169, 2009.

A173. Varki, N., Anderson, D., Herndon, J.G., Pham, T., Gregg, C.J., Cheriyan, M., Murphy, J., Strobert, E., Fritz, J., Else, J.G., and Varki, A. Heart Disease is Common in Humans and Chimpanzees, but is Caused by Different Pathological Processes.  Evolutionary Applications, 2:101-112, 2009.

B130. Banda, K., Kang, G., and Varki, A. "Sialidase-Sensitivity" of Rotavirus Revisited (Invited Editorial).  Nat. Chem. Biol., 5:71-72, 2009.

A172. Diaz, S.L., Padler-Karavani, V., Ghaderi, D., Hurtado-Ziola, N., Yu, H., Chen, X.,Brinkman-Van der Linden, E.C.M., Varki, A., and Varki, N.M. Sensitive and Specific Detection of the Non-Human Sialic Acid N-Glycolylneuraminic Acid In Human Tissues and Biotherapeutic Products.  PLoS ONE, 4:e4241, 2009.

A171. Cariappa, A., Takematsu, H., Liu, H., Diaz, S., Haider, K., Boboila, C., Kalloo, G., Connole, M., Shi, H.N., Varki, N., Varki, A., and Pillai, S. B cell antigen receptor signal strength and peripheral B cell development are regulated by a 9-O-acetyl sialic acid esterase.  J. Exp. Med., 206:125-138, 2009.

B131. Varki, A. Multiple Changes in Sialic Acid Biology During Human Evolution.  Glycoconj. J., 26:231-245, 2009.

B129. Bertozzi, C.R., Freeze, H.H., Varki, A. and Esko, J.D. Chapter 51: Glycans in Biotechnology and the Pharmaceutical Industry. (Book Chapter) IN Essentials of Glycobiology 2nd edition. Edited by Varki, A., Cummings, R.D., Esko, J.D., Freeze, H.H., Stanley, P., Bertozzi, C.R., Hart, G.W., and Etzler, M.E. Cold Spring Harbor Laboratory Press, N.Y., 2009.

B128. Varki, A., Kannagi, R., and Toole, B.P. Chapter 44: Glycosylation Changes in Cancer. (Book Chapter) IN Essentials of Glycobiology 2nd edition. Edited by Varki, A., Cummings, R.D., Esko, J.D., Freeze, H.H., Stanley, P., Bertozzi, C.R., Hart, G.W., and Etzler, M.E. Cold Spring Harbor Laboratory Press, N.Y., 2009.

B127. Varki, A. and Freeze, H.H. Chapter 43: Glycans in Acquired Human Diseases. (Book Chapter) IN Essentials of Glycobiology 2nd edition. Edited by Varki, A., Cummings, R.D., Esko, J.D., Freeze, H.H., Stanley, P., Bertozzi, C.R., Hart, G.W., and Etzler, M.E. Cold Spring Harbor Laboratory Press, N.Y., 2009.

B126. Varki, A., Freeze, H.H., and Vacquier, V.D. Chapter 38: Glycans in Development and Systemic Physiology. (Book Chapter) IN Essentials of Glycobiology 2nd edition. Edited by Varki, A., Cummings, R.D., Esko, J.D., Freeze, H.H., Stanley, P., Bertozzi, C.R., Hart, G.W., and Etzler, M.E. Cold Spring Harbor Laboratory Press, N.Y., 2009.

B125. Varki, A. and Crocker, P.R. Chapter 32: I-type Lectins. (Book Chapter) IN Essentials of Glycobiology 2nd edition. Edited by Varki, A., Cummings, R.D., Esko, J.D., Freeze, H.H., Stanley, P., Bertozzi, C.R., Hart, G.W., and Etzler, M.E. Cold Spring Harbor Laboratory Press, N.Y., 2009.

B124. Varki, A. and Kornfeld, S. Chapter 30: P-type Lectins. (Book Chapter) IN Essentials of Glycobiology 2nd edition. Edited by Varki, A., Cummings, R.D., Esko, J.D., Freeze, H.H., Stanley, P., Bertozzi, C.R., Hart, G.W., and Etzler, M.E. Cold Spring Harbor Laboratory Press, N.Y., 2009.

B123. Varki, A., Etzler, M.E., Cummings, R.D., and Esko, J.D. Chapter 26: Discovery and Classification of Glycan-Binding Proteins. (Book Chapter) IN Essentials of Glycobiology 2nd edition. Edited by Varki, A., Cummings, R.D., Esko, J.D., Freeze, H.H., Stanley, P., Bertozzi, C.R., Hart, G.W., and Etzler, M.E. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, N.Y., 2009.

B122. Varki, A., Freeze, H.H., and Gagneux, P. Chapter 19: Evolution of Glycan Diversity. (Book Chapter) IN Essentials of Glycobiology 2nd edition. Edited by Varki, A., Cummings, R.D., Esko, J.D., Freeze, H.H., Stanley, P., Bertozzi, C.R., Hart, G.W., and Etzler, M.E. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, N.Y., 2009.

B121. Varki, A. and Schauer, R. Chapter 14: Sialic Acids. (Book Chapter) IN Essentials of Glycobiology 2nd edition. Edited by Varki, A., Cummings, R.D., Esko, J.D., Freeze, H.H., Stanley, P., Bertozzi, C.R., Hart, G.W., and Etzler, M.E. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, N.Y., 2009.

B120. Varki, A. and Lowe, J.B. Chapter 6: Biological Roles of Glycans. (Book Chapter) IN Essentials of Glycobiology. 2nd. Edition. Edited by Varki, A., Cummings, R.D., Esko, J.D., Freeze, H.H., Stanley, P., Bertozzi, C.R., Hart, G.W., and Etzler, M.E. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, N.Y., 2009.

B119. Rini, J.M., Esko, J.D., and Varki, A. Chapter 5: Glycosyltransferases and Glycan-processing Enzymes. (Book Chapter) IN Essentials of Glycobiology 2nd edition. Edited by Varki, A., Cummings, R.D., Esko, J.D., Freeze, H.H., Stanley, P., Bertozzi, C.R., Hart, G.W., and Etzler, M.E. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, N.Y., 2009.

B118. Varki, A., Esko, J.D., and Colley, K.J. Chapter 3: Cellular Organization of Glycosylation. (Book Chapter) IN Essentials of Glycobiology 2nd edition. Edited by Varki, A., Cummings, R.D., Esko, J.D., Freeze, H.H., Stanley, P., Bertozzi, C.R., Hart, G.W., and Etzler, M.E. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, N.Y., 2009.

B117. Varki, A., and Sharon, N. Chapter 1: Historical Background and Overview. (Book Chapter) IN Essentials of Glycobiology 2nd edition. Edited by Varki, A., Cummings, R.D., Esko, J.D., Freeze, H.H., Stanley, P., Bertozzi, C.R., Hart, G.W., and Etzler, M.E. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, N.Y., 2009.

A169. Winn, V.D., Gormley, M., Paquet, A.C., Kjaer-Sorensen, K., Kramer, A., Rumer, K.K., Haimov-Kochman, R., Yeh, R.-F., Overgaard, M.T., Varki, A., Oxvig, C., and Fisher, S.J. Severe Preeclampsia-Related Changes in Gene Expression at the Maternal-Fetal Interface Include Sialic Acid-Binding Immunoglobulin-Like Lectin-6 and Pappalysin-2.  Endocrinology, 150:452-462, 2009.


A170. Hedlund, M., Padler-Karavani, V., Varki, N.M., and Varki, A. Evidence for a Human-Specific Mechanism for Diet and Antibody-Mediated Inflammation in Carcinoma Progression. Proc. Natl. Acad. Sci. U.S.A., 105:18936-18941, 2008. PNAS Online

A168. Byres, E., Paton, A.W., Paton, J.C., Löfling, J.C., Smith, D.F., Wilce, M.C., Talbot, U.M., Chong, D.C., Yu, H., Huang, S., Chen, X., Varki, N.M., and Varki, A., Rossjohn, J., and Beddoe, T. Incorporation of a non-human glycan mediates human susceptibility to a bacterial toxin.  Nature, 456:648-652, 2008. # = corresponding authors.

B115. Varki, A., Geschwind, D.H., and Eichler, E.E. Explaining Human Uniqueness: Genome Interactions with Environment, Behavior and Culture.  Nat. Rev. Genet., 9:749-763, 2008.

A167. Chokhawala, H.A., Huang, S., Lau, K., Yu, H., Cheng, J., Thoh, V., Hurtado-Ziola, N., Guerrero J.A., Varki, A. and Chen, X. Combinatorial Chemoenzymatic Synthesis and High-Throughput Screening of Sialosides. ACS Chem. Biol., 3:567-576, 2008.

A166. Padler-Karavani, V., Yu, H., Cao, H., Chokhawala, H., Karp, F., Varki, N., Chen, X., and Varki, A. Diversity in Specificity, Abundance and Composition of Anti-Neu5Gc Antibodies in Normal Humans: Potential Implications for Disease.  Glycobiology, 18:818-830, 2008.

B116. Varki, A.P., Baum, L.G., Bellis, S.L., Cummings, R.D., Esko, J.D., Hart, G.W., Linhardt, R.J., Lowe, J.B., McEver, R.P., Srivastava, A., and Sarkar, R. Working Group Report: The Roles of Glycans in Hemostasis, Inflammation and Vascular Biology.  Glycobiology, 18:747-749, 2008.

B114. Varki, A. Letter to the Glyco-Forum.  Glycobiology, 18:659, 2008.

B113. Varki, A. Sialic Acids in Human Health and Disease. (Invited Essay) Trends Mol. Med., 14:351-360, 2008.

B112. Varki, A. Addressing the Major Challenge for Women in Academia: "It's Proximate Childcare, Stupid!". (Invited Opinion Piece) ASCB Newsletter, April 2008.

A165. Hedlund, M., Ng, E., Varki, A., and Varki, N.M. Alpha2-6-linked Sialic Acids on N-Glycans Modulate Carcinoma Differentiation In Vivo. Cancer Res., 68:388-394, 2008.


A164. Stevenson, J.L., Varki, A., and Borsig, L. Heparin attenuates metastasis mainly due to inhibition of P- and L-selectin, but non-anticoagulant heparins can have additional effects. Thromb. Res., 120 Suppl2 :S107-S111, 2007.

B110. Varki, A. Obituary: Rosalind Hauk Kornfeld (1935-2007). Glycobiology, 17:1148-1149, 2007.

B098. Lewis, A. and Varki, A. Chapter 4: Evolutionary Considerations in Studying the Sialome: Sialic Acids and the Host-Pathogen Interface. (Invited Book Chapter) IN, Bioinformatics for Glycobiology and Glycomics. Ed. Von der Lieth C-W., John Wiley & Sons, pp. 69-88, 2007.

A163. Lewis, A.L., Cao, H., Patel, S.K., Diaz, S., Ryan, W., Carlin, A.F., Thon, V., Lewis, W.G., Varki, A., Chen, X., and Nizet, V. NeuA Sialic Acid O-acetylesterase Activity Modulates O-acetylation of Capsular Polysaccharides in Group B Streptococcus. J. Biol. Chem., 282:27562-27571, 2007.

B111. Borsig, L., Stevenson J.L., and Varki, A. Heparin in Cancer: Role of Selectin Interactions. IN, Cancer-Associated Thrombosis, (Invited Book Chapter) Eds. Alok A. Khorana and Charles W. Francis. Publisher, Taylor-Francis, Chapter 7:95-111, 2007.

B106. Lanctot, P.M., Gage, F.H. and Varki, A.P. The Glycans of Stem Cells. Curr. Opin. Chem Biol., 11:373-380, 2007.

A162. Yu, H., Chokhawala, H.A., Varki, A. and Chen, X. Efficient chemoenzymatic synthesis of biotinylated human serum albumin-sialoglycoside conjugates containing O-acetylated sialic acids. Org. Biomol. Chem., 5:2458-2463, 2007.

A161. Brinkman-Van der Linden, E.C.M., Hurtado-Ziola, N., Hayakawa, T., Wiggleton, L., Benirschke, K., Varki, A. and Varki, N. Human-Specific Expression of Siglec-6 in the Placenta. Glycobiology, 17:922-931, 2007.

B107. Varki, A. Trousseau's Syndrome: Multiple Definitions and Multiple Mechanisms (Invited Review). Blood, 110:1723-1729, 2007.

B108. Varki, N.M. and Varki, A. Diversity in Cell Surface Sialic Acid Presentations: Implications for Biology and Disease (Invited Review). Lab Invest., 87:851-857, 2007.

B109. Varki, A. and Nelson, D.L. Genomic Comparisons of Humans and Chimpanzees (Invited Review). Annu. Rev. Anthropol., 36:191-209, 2007.

A160. Hedlund, M., Tangvoranuntakul, P., Takematsu, H., Long, J.M., Housley, G.D., Kozutsumi, Y., Suzuki, A., Wynshaw-Boris, A., Ryan, A.F., Gallo, R.L., Varki, N. and Varki, A. N-glycolylneuraminic Acid Deficiency in Mice: Implications for Human Biology and EvolutionMol. Cell. Biol., 27:4340-4346, 2007.

A159. Sakamoto, N., Tsujik, K., Muul, L.M., Lawler, A.M., Petricoin, E.F., Candotti, F., Metcalf, J.A., Tavel, J.A., Lane, H.C., Urba, W.J., Fox, B.A., Varki, A., Lunney, J.K., and Rosenberg, A.S. Bovine apolipoprotein B-100 is a dominent immunogen in therapeutic cells populations cultured in fetal calf serum in mice and humans. Blood, 110:501-508, 2007.

A158. Zhang, M., Angata, T., Cho, J.Y., Miller, M., Broide, D. H., and Varki, A. Defining the in vivo function of Siglec-F, a CD33-related Siglec expressed on mouse eosinophils. Blood, 109:4280-4287, 2007.

B105. Varki, A. Glycan-based Interactions Involving Vertebrate Sialic Acid-Recognizing Proteins (Invited Review). Nature, 446:1023-1029, 2007.

B104. Varki, A. The uncertain future of research chimpanzees.  Science, 315:1493-1494, 2007.

B103. Crocker, P.R., Paulson, J.C., and Varki, A. Siglecs and Their Roles in the Immune System (Invited Review). Nat. Rev. Immunol., 7:255-266, 2007.

A157. Carlin, A.F., Lewis, A.L., Varki, A., and Nizet, V. Group B Streptococcal Capsular Sialic Acids Interact with Siglecs (Immunoglobulin-Like Lectins) on Human Leukocytes. J. Bacteriol., 189:1231-1237, 2007.


A156. Altheide, T.K., Hayakawa, T., Mikkelsen, T.S., Diaz, S., Varki, N., and Varki, A. System-wide Genomic and Biochemical Comparisons of Sialic Acid Biology Among Primates and Rodents: Evidence for Two Modes of Rapid Evolution.   J. Biol. Chem., 281:25689-25702, 2006. 

A155. Angata, T., Hayakawa, T, Yamanaka, M., Varki, A., and Nakamura, M.Discovery of Siglec-14, a Novel Sialic Acid Receptor Undergoing Concerted Evolution with Siglec-5 in Primates. FASEB J, 20:1964-1973, 2006.

A154. Nguyen, D.H., Hurtado-Ziola, N., Gagneux, P., and Varki, A. Loss of Siglec Expression on T lymphocytes during Human Evolution. Proc. Natl. Acad. Sci. U.S.A., 103:7765-7770, 2006. UCSD Press Release

A153. Nguyen, D.H., Ball, E.D., and Varki, A. Myeloid Precursors and Acute Myeloid Leukemia Cells Express Multiple CD33-related Siglecs. Exp. Hematol., 34:728-735, 2006. 

A152. Lewis, A.L., Hensler, M.E., Varki, A., and Nizet, V. The Group B Streptococcal Sialic Acid O-acetyltransferase is encoded by neuD, a Conserved Component of Bacterial Sialic Acid Biosynthetic Gene Clusters. J. Biol. Chem., 281:11186-11192, 2006.

A151. Chen, H.Y., Challa, A.K., and Varki, A. 9-O-acetylation of Exogenously Added Ganglioside GD3. The GD3 Molecule Induces its Own O-acetylation Machinery. J. Biol. Chem., 281:7825-7833, 2006.

A150. Wahrenbrock, M.G., and Varki, A. Multiple Hepatic Receptors Cooperate to Eliminate Secretory Mucins Aberrantly Entering the Bloodstream: Are Circulating Cancer Mucins the "Tip of the Iceberg"? Cancer Res., 66:2433-2441, 2006.

A149. Läubli, H., Stevenson, J.L., Varki, A., Varki, N.M., and Borsig, L. L-selectin Facilitation of Metastasis Involves Temporal Induction of Fut7-Dependent Ligands at Sites of Tumor Cell Arrest. Cancer Res., 66:1536-1542, 2006.

A148. Hayakawa T., Aki I., Varki A., Satta Y., and Takahata N. Fixation of the Human-specific CMP-N-acetylneuraminic Acid Hydroxylase Pseudogene and Implications of Haplotype Diversity for Human Evolution. Genetics, 172:1139-1146, 2006.

B102. Martin, M.J., Muotri, A., Gage, F.H., and Varki, A. Response to: Complement Targeting of Nonhuman Sialic Acid Does Not Mediate Cell Death of Human Embryonic Stem Cells. Nat. Med., 12:1115, 2006. 

B101. Varki A. Nothing in Glycobiology Makes Sense, Except in the Light of Evolution (Invited Essay). Cell 126:841-845, 2006. 

B100. Varki, A., Holmes, E., Yamada, T., Agre, P., and Brenner, S. Physician-Scientists are Needed Now More Than Ever. Nature (Correspondence) 440:740, 2006.

B099. Varki, A. Explaining the Human Condition: Problems and Prospects. IN Proceedings of Center for Human Evolution Workshop 5, The Evolution of the Human Brain, March 19-20, 2005. Bellevue, WA: Foundation for the Future, (Invited Symposium Presentation), 2006.

B097. Varki, A., and Angata T. Siglecs: The Major Subfamily of I-type Lectins (Invited Review) Glycobiology, 16:1R-27R, 2006.


A147. Hayakawa T., Angata, T., Lewis, A.L., Mikkelsen T.S., Varki, N.M. & Varki A. A Human-Specific Gene in Microglia. Science, 309:1963, 2005. UCSD Press Release

A146. Martin M.J., Rayner J.C., Gagneux P., Barnwell J.W., & Varki A. Evolution of Human-Chimpanzee Differences in Malaria Susceptibility: Relationship to Human Genetic Loss of N-glycolylneuraminic Acid. Proc. Natl. Acad. Sci. U.S.A., 102:12819-12824, 2005. UCSD Press Release

A145. Stevenson, J.L., Choi, S.H. & Varki, A. Differential Metastasis Inhibition by Clinically-Relevant Levels of Heparins - Correlation with Selectin Inhibition, not Antithrombotic Activity. Clin. Cancer Res. 11:7003-7011, 2005.

A144. Chimpanzee Sequencing and Analysis Consortium. Initial Sequence of the Chimpanzee Genome and Comparison with the Human Genome. Nature 437:69-87, 2005.

A143. Nguyen, D.H., Tangvoranuntakul, P., Varki, A. Effects of natural human antibodies against a non-human sialic acid that metabolically incorporates into activated and malignant immune cells. J. Immunol. 175:228-236, 2005.

A142. Jessani, N., Young, J.A., Diaz, S.L., Patricelli, M.P., Varki, A., Cravatt, B.F. Class Assignment of Sequence-Unrelated Members of Enzyme Superfamilies by Activity-Based Protein Profiling. Angew Chem Int Ed Engl, 44:2400-2403, 2005.

A141. Bardor, M., Nguyen, D., Diaz, S., Varki, A. Mechanism of Uptake and Incorporation of the Non-Human Sialic Acid N-glycolylneuraminic acid into Human Cells. J. Biol. Chem, 280:4228-4237, 2005.

A140. Martin, M.J., Moutri, A., Gage, F., Varki, A. Human Embryonic Stem Cells Express an Immunogenic Non-Human Sialic Acid. Nat. Med. 11:228-232, 2005. UCSD Press Release

B096. Varki, A. and Altheide, T.K. Comparing the human and chimpanzee genomes: Searching for needles in a haystack (Invited Perspective). Genome Research. 15:1746-1758, 2005.

B095. Stevenson, J.L., Choi, S.H., Wahrenbrock, M., Varki, A. & Varki, N.M. Heparin Effects in Metastasis and Trousseau's Syndrome: Anticoagulation is not the Primary Mechanism (Invited Article). Haematologica. 1:59-60, 2005.

B094. McConkey, E. H. and Varki A. Genomics: Thoughts on the Future of Great Ape Research (Invited Perspective). Science 309:1499-1501, 2005.

B093. Gagneux P., Moore J.J. and Varki, A. The Ethics of Research on Great Apes (Invited Commentary). Nature 437:27-29, 2005. UCSD Press Release

B092. Lewis, A., Altheide, T., Varki, A., Arden, K., Varki, N. Combining Parenting and a Science Career (Letter). Science 307:1720, 2005.

B091. Hayakawa T., Altheide, TK and Varki, A. Genetic Basis of Human Brain Evolution: Accelerating along the Primate Speedway (Invited Commentary). Dev. Cell 8:2-4, 2005.


A139. Angata T, Margulies E.H., Green, E. D., Varki, A. Large-Scale Sequencing of the CD33-Related Siglec Gene Cluster in Five Mammalian Species Reveals Rapid Evolution by Multiple Mechanisms. Proc. Natl. Acad. Sci. U.S.A., 101:13251-13256, 2004.

A138. Zhang, M. and Varki, A. Cell Surface Sialic Acids do not Affect Primary CD22 Interactions with CD45 and Surface IgM, nor the rate of Constitutive CD22 Endocytosis. Glycobiology 14:939-949, 2004.

A137. Lewis, A.L., Nizet, V., and Varki, A. Discovery and Characterization of Sialic Acid O-Acetylation in Group B Streptococcus. Proc. Nat'l. Acad. Sci. U.S.A., 101:11123-11128, 2004. UCSD Press Release

A136. Taverna, D., Moher, H., Crowley, D., Borsig, L., Varki, A., and Hynes, R.O. Increased primary tumor growth in mice null for β3- or β3/β5-integrins or selectins. Proc Natl Acad Sci USA. 101:763-768, 2004. Supporting Figure 8

A135. Sonnenburg, J.L., Altheide, T.K., and Varki, A. A Uniquely Human Consequence of Domain-specific Functional Adaptation in a Sialic Acid-binding Receptor. Glycobiology 14:339-46, 2004.

B090. Insel, P.A., Kornfeld, S., Majerus, P.W., Marks, A.R., Marks, P.A., Relman, A.S., Scharschmidt, B.F., Stossel, T.P., Varki, A.P., Weiss, S.J., and Wilson, J.D. Blasts from the past. (Invited Commentary from former Chief Editors for 80th. Anniversary issue). J. Clin. Invest. 114:1017-1033, 2004.

B089. Varki, A. How to Make an Ape Brain (Invited “News & Views”) Nature Genetics 36:1034-1036, 2004.

B088. Varki, A. Glycosylation engineering enhances stem cell homing. (Invited Commentary) Blood, 104:3005, 2004.

B087. Olson, M.V. and Varki A. The Chimpanzee Genome: a Bittersweet Celebration (Invited Perspective) Science 305:191-192, 2004.

B085. Varki, A. Siglecs (Invited Contribution) Encyclopedia of Biological Chemistry. Eds. Lennarz, W.J. and Lane, M.D. Elsevier Science, San Diego 4:38-40, 2004. 

B084. Varki, A. Genetic Differences Between Humans and Great Apes –Implications for the Evolution of Humans (Invited Article). International Astronomy Union Symposium #213, Eds. Ray Norris, Carol Oliver, and Frank Stoolman, Astronomical Union of the Pacific, 2004.


A134. Gagneux, P., Cheriyan, M., Hurtado-Ziola, N., Brinkman van der Linden, E.C.M., Anderson, D., McClure, H., Varki, A., and Varki, N.M. Human-specific Regulation of α2-6-linked Sialic Acids. J. Biol. Chem. 278:48245-4825 2003. 

A133. Tangvoranuntakul, P., Gagneux, P., Diaz, S., Bardor, M., Varki, N.M., Varki, A., and Muchmore, E. Human Uptake and Incorporation of an Immunogenic Non-human Dietary Sialic Acid. Proc. Nat'l. Acad. Sci. U.S.A., 100:12045-12050, 2003.

A132. Wahrenbrock, M., Borsig, L., Le, D., Varki, N. and Varki, A. Selectin-Mucin Interactions as a Probable Molecular Explanation for the Association of Trousseau Syndrome with Mucinous Adenocarcinomas. J. Clin. Invest. 112:853-862, 2003.

A131. Brinkman-Van der Linden, E.C.M., Angata, T., Reynolds, S.A., Powell, L.D., Hedrick, S.M., and Varki, A. CD33/Siglec-3: binding specificity, expression pattern and consequences of gene deletion in mice. Mol. Cell Biol. 23:4199-4206, 2003. 

A130. Satake, H., Chen, H.Y. and Varki, A. Genes Modulated by Expression of GD3 Synthase in Chinese Hamster Ovary Cells: Evidence that the Tis21 gene is involved in the induction of GD3 9-O-acetylation. J. Biol. Chem. 278:7942-7948, 2003.

B086. Varki, A. Open access: the JCI has already shown it works (Letter). Nature 426:383, 2003. 

B083. Brinkman-Van der Linden, E.C.M. and Varki, A. Probing for masked and unmasked Siglecs on Cell surfaces. (Invited Article) IN Recognition of Carbohydrates in Biological Systems Methods in Enzymology Lee, Y.T. and Lee R.T.(eds). Academic Press, 363:113-120, 2003.

B082. Martin, L., Verhagen, A, and Varki, A. Recombinant Influenza C Hemagglutinin-Esterase as a probe for Sialic Acid 9-O-acetylation. (Invited Article) IN Recognition of Carbohydrates in Biological Systems. Methods in Enzymology Lee, Y.T. and Lee R.T.(eds). Academic Press, 363:489-498, 2003.

B081. Olson, M. and Varki A. Sequencing the Chimpanzee Genome: Insights into Human Evolution and Disease. (Invited Review) Nat. Rev. Genetics, 4:20-28, 2003.


A129. Chou H.H., Hayakawa, T., Diaz, S., Krings, M., Indriati, E., Leakey, M., Paabo, S., Satta, Y., Takahata, N., and Varki, A. Inactivation of CMP-N-acetylneuraminic Acid Hydroxylase Occurred Prior to Brain Expansion during Human Evolution. Proc. Nat'l. Acad. Sci. U.S.A., 99:11736–11741, 2002. UCSD Press Release

A128. Matsui, N.M., Varki, A., and Embury, S.H. Heparin Inhibits the Flow Adhesion of Sickle Red Blood Cells to P-selectin. Blood 100:3790-3796, 2002.

A127. Martin, L.T., Marth, J.D., Varki, A., and Varki, N.M. Genetically altered mice with different sialyltransferase deficiencies show tissue-specific alterations in sialylation and sialic acid 9-0-acetylation. J. Biol. Chem. 277:32930-32938, 2002.

A126. Wang, L., Brown, J.R., Varki, A., and Esko, J.D. Heparin’s anti-inflammatory effects require glucosamine 6-O-sulfation and are mediated by blockade of L- and P-selectins. J. Clin. Invest. 110:127-136, 2002.

A125. Ellies, L.G., Ditto, D., Levy, G.G., Wahrenbrock, M., Ginsburg, D., Varki, A., Le, D.T., and Marth, J.D. Sialyltransferase ST3Gal-IV operates as a dominant modifier of hemostasis by concealing asialoglycoprotein receptor ligands. Proc. Nat'l. Acad. Sci. U.S.A., 99:10042-10047, 2002. UCSD Press Release

A124. Angata, T., Kerr, S.C., Greaves, D.R., Varki, N.M., Crocker, P.R., and Varki, A. Cloning and characterization of human Siglec-11. A recently evolved signaling molecule that can interact with SHP-1 and SHP-2 and is expressed by tissue macrophages, including brain microglia. J. Biol. Chem. 277:24466-24474, 2002.

A123. Onami, T.M., Lin, M-Y., Page, D.M., Reynolds, S.A., Katayama, C.D., Marth, J.D., Irimura, T., Varki, A., Varki, N. and Hedrick, S.M. Generation of mice deficient for macrophage galactose- and N-acetylgalactosamine-specific lectin: Limited role in lymphoid and erythroid homeostasis and evidence for multiple lectins. Mol. Cell. Biol. 22:5173-5181, 2002. 

A122. Sonnenburg, J.L., van Halbeek, H., and Varki, A. Characterization of the acid stability of glycosidically-linked neuraminic acid: Use in detecting De-N-acetyl-gangliosides in human melanoma. J Biol Chem. 277:17502-17510, 2002.

A121. Enard, W., Khaitovich, P., Klose, J., Zöllner, S., Heissig, F., Giavalisco, P., Nieselt-Struwe, K., Muchmore, E., Varki, A., Ravid, R., Doxiadis, G.M., Bontrop, R.E., and Pääbo, S. Intra- and Interspecific Variation in Primate Gene Expression Patterns. Science, 296:340-343, 2002.  Supplemental data    UCSD Press Release

A120. Borsig, L., Wong, R., Hynes, R.O., Varki, N.M., and Varki, A. Synergistic Effects of L- and P-selectin in Facilitating Tumor Metastasis Can Involve Non-Mucin Ligands And Implicates Leukocytes as Enhancers of Metastasis. Proc. Nat'l. Acad. Sci. U.S.A., 99:2193-2198, 2002.

A119. Brinkman-Van der Linden, E. C.M., Sonnenburg, J.L. and Varki, A. Effects of Sialic Acid Substitutions on Recognition by Sambucus Nigra Agglutinin and Maackia Amurensis Hemagglutinin. Anal Biochem. 303:98-104, 2002.

B080. Chen, H. and Varki, A. O-acetylation of GD3: an enigmatic modification regulating Apoptosis? (Invited Commentary) J. Exp. Med., 196:1529-1533, 2002.

B079. Varki, A. and Rosenberg, L.E. Emerging opportunities and career paths for the young physician-scientist. Nat. Med., 8:437-439, 2002.

B078. Gagneux, P. and Varki, A. Reply to Comments by Previc: Endocrine differences between humans and great apes - did environmental factors provide genetic opportunities? Am. J. Phys. Anthro., 118:404-405, 2002.

B077. Varki, A. Six Blind Men and the Elephant - the Many Faces of Heparan Sulfates. (invited Commentary) Proc. Natl. Acad. Sci. U.S.A., 99:543-545, 2002. 

B076. Varki, A. Loss of N-Glycolylneuraminic acid in Humans: Mechanisms, Consequences and Implications for Hominid Evolution. (Invited Review) Yearbook of Physical Anthropology 44:54-69, 2002. 

B075. Angata T., and Varki, A. Chemical Diversity in the Sialic Acids and Related Alpha-keto Acids: an Evolutionary Perspective (Invited Review) Chemical Reviews, 102:439-469, 2002. Supporting info.

B074. Varki, N. and Varki, A. Heparin inhibition of Selectin-mediated mechanisms in the Hematogenous Phase of Carcinoma Metastasis: Rationale for Clinical Trials in Humans (Invited Review) Seminars in Thrombosis and Hemostasis, 28:53-66, 2002.


A118. Angata, T., Hingorani, R., Varki, N.M., and Varki, A. Cloning and characterization of a novel mouse Siglec, mSiglec-F: Differential evolution of the mouse and human (CD33) Siglec-3-related gene clusters. J. Biol. Chem. 276:45128-45136, 2001.

A117. Angata, T., Varki N.M. and Varki, A. A Second Uniquely Human Mutation Affecting Sialic Acid Biology. J. Biol. Chem. 276:40282-40287, 2001.

A116. Hayakawa T., Satta Y., Gagneux P., Varki A., and Takahata N. Alu-mediated inactivation of the human CMP-N-acetylneuraminic acid hydroxylase gene. Proc. Nat'l. Acad. Sci. U.S.A., 98:11399–11404, 2001.

A115. Matsui, N.M., Borsig, L., Rosen, S. D., Yaghmai, M., Varki, A., and Embury, S.H. P-Selectin Mediates the Adhesion of Sickle Erythrocytes to the Endothelium. Blood, 98:1955-1962, 2001.

A114. Bai, X., Brown, J.R., Varki, A., and Esko, J.D. Enhanced 3-O-Sulfation of Galactose in Asn-linked Glycans and Maackia amurenesis Lectin binding in a new Chinese Hamster Ovary Cell Line. Glycobiology 11:621-632, 2001.

A113. Gagneux, P., Amess, B., Diaz, S., Moore, S., Patel, T., Dillmann, W., Parekh, R., and Varki, A. Proteomic Comparison of Human and Great Ape Blood Plasma Reveals Conserved Glycosylation and differences in Thyroid Hormone Metabolism. Amer. J. Physical Anthropology 115:99-109, 2001. 

A112. Srikrishna, G., Panneerselvam, K., Westphal, V., Abraham, V., Varki, A., and Freeze, H.H. Two proteins modulating transendothelial migration of leukocytes recognize novel carboxylated glycans on endothelial cells. J. Immunol. 166:4678-4688, 2001. 

A111. Borsig, L., Wong, R., Feramisco, J., Nadeau, D.R., Varki, N.M., and Varki, A. Heparin and cancer revisited: Mechanistic connections involving platelets, P-Selectin, carcinoma mucins and tumor metastasis. Proc. Nat'l. Acad. Sci. U.S.A., 98:3352-3357, 2001.  Supplemental moviesUCSD Press Release.

A110. Srikrishna, G., Toomre, D.K., Manzi, A., Panneerselvam, K., Freeze, H.H., Varki, A., and Varki, N.M. A novel anionic modification of N-glycans on mammalian endothelial cells is recognized by activated neutrophils and modulates acute inflammatory responses. J. Immunol. 166:624-632, 2001.

B073. Borsig, L., Wong, R., Feramisco, J., Nadeau, D.R., Varki, N.M., and Varki, A. Pictures in Molecular Medicine: Three-dimensional visualization of intravascular tumor cells in mice. Trends in Mol. Med. 7:377, 2001.

B072. Varki, A. N-Glycolyl-Neuraminic Acid Deficiency in Humans. (Invited Review) Biochimie 83:615-622, 2001.

B071. Crocker P. and Varki, A. Siglecs in the Immune System. (Invited Review) Immunology 103:137-145, 2001.

B070. Varki, A. and Varki N. P-Selectin, Carcinoma Metastasis and Heparin: Novel Mechanistic Connections with Therapeutic Implications (Invited Review) Brazilian J. Med. Biol. Res. 34:711-717, 2001.

B069. Crocker P. and Varki, A. Siglecs, Sialic Acids and Innate Immunity. (Invited Review) Trends in Immunology 22:337, 2001.


A109. Angata, T., Varki, A. Cloning, Characterization, and Phylogenetic Analysis of Siglec-9, a New Member of the CD33-Related Group of Siglecs. J. Biol. Chem. 275:22127-22135, 2000.

A108. Norgard-Sumnicht, K., Bai, X., Esko, J.D., Varki, A., and Manzi, A.E. Exploring the Outcome of Genetic Modifications of Glycosylation in Cultured Cell Lines by Concurrent Isolation of the Major Classes of Vertebrate Glycans. Glycobiology 10:691-700, 2000.

A107. Manzi, A.E., Norgard-Sumnicht, K., Argade, S., Marth, J.D., van Halbeek, H., and Varki, A. Exploring the Glycan Repertoire of Genetically Modified Mice by Isolation and Profiling of the Major Glycan Classes and Nano-NMR Analysis of Glycan Mixtures. Glycobiology 10:669-689, 2000.

A106. Brinkman-Van der Linden, E.C.M., Sjoberg, E.R., Juneja, L.R., Crocker, P.R., Varki, N., and Varki, A. Loss Of N-Glycolyl-Neuraminic Acid In Human Evolution: Implications For Sialic Acid Recognition By Siglecs. J. Biol. Chem. 275:8633-40. 2000.

A105. Brinkman-Van der Linden, E.C.M., and Varki, A. New Aspects Of Siglec Binding Specificities, Including The Significance Of Fucosylation And Of The Sialyl-Tn Epitope. J. Biol. Chem. 275:8625-8632. 2000.

A104. Angata, T., Varki, A. Siglec-7: A Sialic Acid-Binding Lectin of the Immunoglobulin Superfamily. Glycobiology 10:431-438. 2000.

A103. Todeschini, A.R., Mendonca-Previato, L., Previato, J.O., Varki, A., and van Halbeek, H. Trans-sialidase from Trypanosoma cruzi catalyzes sialoside hydrolysis with retention of configuration. Glycobiology 10:213-21. 2000.

B068. McConkey, E. H. and Varki, A. Examining priorities for a primate genome project - Response.Science 290:1505. 2000.

B067. Gagneux, P., and Varki, A. Genetic Differences Between Humans and Great Apes. (Invited Review) Molecular Phylogenetics and Evolution 18:2-13. 2000.

B066. McConkey, E.H. and Varki A. A Primate Genome Project Deserves High Priority. (Letter to the Editor) Science 289:1295-96, 2000.

B065. Varki, A. A Chimpanzee Genome Project is a Biomedical Imperative (Invited Opinion Article) Genome Research 10:1065-1070, 2000.

B064. McConkey, E. H., Fouts, R., Goodman, M., Nelson, D., Penny, D., Ruvolo, M., Sikela, J., Stewart, C.B., Varki, A., and Wise, S. Proposal for a Human Genome Evolution Project. Molecular Phylogenetics and Evolution.15:1-4, 2000.


A102. Takematsu, H., Diaz, S., Stoddart, A., Zhang, Y., and Varki, A. Lysosomal and cytosolic sialic acid 9-O-acetylesterase activities can be encoded by one gene via differential usage of a signal peptide-encoding exon at the N terminus. J. Biol. Chem. 274:25623-25631, 1999.

A101. Patel, N., Brinkman-Van der Linden, E.C.M., Altmann, S.W., Gish, K., Balasubramanian, S., Timans, J.C., Peterson, D., Bell, M.P., Bazan, J.F., Varki, A., Kastelein, R.A OB-BP1/Siglec-6: a leptin and sialic acid-binding protein of the immunoglobulin superfamily. J. Biol. Chem. 274:22729-22738, 1999.

A100. Kim, Y.J., Borsig, L., Han, H-L., Varki, N.M., and Varki, A. Distinct selectin ligands on colon carcinoma mucins can mediate pathological interactions among platelets, leukocytes, and endothelium. Amer. J. Pathol.155:461-472, 1999.

A099. Razi, N. and Varki, A. Cryptic sialic acid binding lectins on human blood leukocytes can be unmasked by sialidase treatment or cellular activation. Glycobiology 9:1225–1234, 1999.

A098. Chammas, R., Sonnenburg, J.L., Watson, N.E., Tai, T., Farquhar, M.G., Varki, N.M., and Varki, A. De-N-acetyl-gangliosides in Humans: Unusual Subcellular Distribution of a Novel Tumor Antigen. Cancer Research 59:1337-1346, 1999.

B063. Varki, A. Diversity at the 5-Carbon Position of Sialic Acids. IN: Sialobiology and Other Forms of Glycosylation. Edited by Inoue, Y., Lee, Y.C., and Troy II, F.A. Gakushin Publishing Co., Osaka, Japan, 1999.

B062. Varki, A. Executive Summary of The Nerflex Commission Report, April 25th. 2049 (based on the 1999 ASCI Presidential Address). J. Clin. Invest (Supplement). v 104, S1-S10, 1999.

B061. Gagneux, P. and Varki, A. Evolutionary Considerations in Relating Oligosaccharide Diversity to Biological Function. Glycobiology. 9:747-755, 1999.

B060. Paulson, J.C.,Varki, A., and Esko, J. Glycobiology in Biotechnology and Medicine. IN: Essentials of Glycobiology. Edited by Varki, A., Esko, J., Cummings, R., Freeze, H.H., Hart, G.W., and Marth, J. Plainview, N.Y., Cold Spring Harbor Laboratory Press, 1999.

B059. Varki, A. Acquired Glycosylation Changes in Human Disease. IN Essentials of Glycobiology. Edited by Varki, A., Esko, J., Cummings, R., Freeze, H.H., Hart, G.W., and Marth, J. Plainview, N.Y., Cold Spring Harbor Laboratory Press, 1999.

B058. Varki, A. Glycosylation Changes in Cancer. IN Essentials of Glycobiology. Edited by Varki, A., Esko, J., Cummings, R., Freeze, H.H., Hart, G.W., and Marth, J. Plainview, N.Y., Cold Spring Harbor Laboratory Press, 1999.

B057. Varki, A. P-Type Lectins. IN Essentials of Glycobiology. Edited by Varki, A., Esko, J., Cummings, R., Freeze, H.H., Hart, G.W., and Marth, J. Plainview, N.Y., Cold Spring Harbor Laboratory Press, 1999.

B056. Varki, A. Discovery and Classification of Animal Lectins. IN Essentials of Glycobiology. Edited by Varki, A., Esko, J., Cummings, R., Freeze, H.H., Hart, G.W., and Marth, J. Plainview, N.Y., Cold Spring Harbor Laboratory Press, 1999.

B055. Lowe, J., and Varki, A. Glycosyltransferases. IN Essentials of Glycobiology. Edited by Varki, A., Esko, J., Cummings, R., Freeze, H.H., Hart, G.W., and Marth, J. Plainview, N.Y., Cold Spring Harbor Laboratory Press, 1999.

B054. Varki, A. Sialic Acids. IN Essentials of Glycobiology. Edited by Varki, A., Esko, J., Cummings, R., Freeze, H.H., Hart, G.W., and Marth, J. Plainview, N.Y., Cold Spring Harbor Laboratory Press, 1999.

B053. Varki, A. Glycosphingolipids. IN Essentials of Glycobiology. Edited by Varki, A., Esko, J., Cummings, R., Freeze, H.H., Hart, G.W., and Marth, J. Plainview, N.Y., Cold Spring Harbor Laboratory Press, 1999.

B052. Varki, A. Exploring the Biological Roles of Glycans. IN Essentials of Glycobiology. Edited by Varki, A., Esko, J., Cummings, R., Freeze, H.H., Hart, G.W., and Marth, J. Plainview, N.Y., Cold Spring Harbor Laboratory Press, 1999.

B051. Varki, A. Evolution of Oligosaccharide Diversity. IN Essentials of Glycobiology. Edited by Varki, A., Esko, J., Cummings, R., Freeze, H.H., Hart, G.W., and Marth, J. Plainview, N.Y., Cold Spring Harbor Laboratory Press, 1999.

B050. Varki, A. Historical Background and Overview. IN Essentials of Glycobiology. Edited by Varki, A., Esko, J., Cummings, R., Freeze, H.H., Hart, G.W., and Marth, J. Plainview, N.Y., Cold Spring Harbor Laboratory Press, 1999.

B049. Varki, A. Disappearing Physician-Scientists (Letter to the Editor) Science 283:791-792, 1999.


A097. Chou, H-H., Takematsu, H., Diaz, S., Iber, J., Nickerson, E., Wright, K.L., Muchmore, E.A., Nelson, D.L., Warren, S.T. and Varki, A. A mutation in human CMP-sialic acid hydroxylase occurred after the Homo-Pan divergence. Proc. Nat'l. Acad. Sci. U.S.A. 95:11751-11756, 1998. UCSD Press Release.

A096. Muchmore, E.A., Diaz, S. and Varki, A. A structural difference between the cell surfaces of humans and the great apes. Amer. J. Physical Anthropology 107:187-198, 1998.

A095. Kim, Y.J., Borsig, L., Varki, N.M. and Varki, A. P-Selectin deficiency attenuates tumor growth and metastasis. Proc. Nat'l. Acad. Sci. U.S.A. 95:9325-9330, 1998.

A094. Razi, N. and Varki, A. Masking and unmasking of the sialic acid-binding lectin activity of CD22 (Siglec-2) on B lymphocytes. Proc. Nat'l. Acad. Sci. U.S.A. 95:7469-7474, 1998.

A093. Jain R.K., Piskorz, C.F., Huang, B-G, Locke, R.D., Han, H-L., Koenig, A., Varki, A. and Matta, K.L. Inhibition of L- and P-selectin by a rationally synthesized novel core 2-like branched structure containing GalNAc-Lewisx and Neu5Acα2-3Galβ1-3GalNAc sequencesGlycobiology 8:707-717, 1998.

A092. Koenig, A., Norgard-Sumnicht, K., Linhardt, R. and Varki, A. Differential interactions of heparin and heparan sulfate glycosaminoglycans with the selectins. Implications for the use of unfractionated and low molecular weight heparins as therapeutic agents. J. Clin. Invest. 101:877-889, 1998.

A091. Shi, W-X, Chammas, R. and Varki, A. Induction of sialic acid 9-O-acetylation by diverse gene products: implications for the expression cloning of sialic acid O-acetyltransferases. Glycobiology 8:199-205, 1998.

B048. Varki, A., Wills, C., Perlmutter, D., Woodruff, D., Gage, F., Moore, J., Semendeferi, K.,Benirschke, K., Katzman, R., Doolittle, R., and Bullock, T. Great Ape Phenome Project? (Letter to the Editor) Science 282:239-240, 1998.

B047. Crocker, P.R., Clark, E.A., Filbin, M., Gordon, S., Jones, Y., Kehrl, J.H., Kelm, S., Le Douarin, N., Powell, L.D., Roder, J., Schnaar, R.L., Sgroi, D.C., Stamenkovic, I., Schauer, R., Schachner, M., van den Berg, T.K., Anton van der Merwe, P., Watt, S.M., Varki, A. Siglecs – a family of sialic acid-binding lectins. Glycobiology 8 v (2),1998.

B046. Varki, A. Factors Controlling the Glycosylation Potential of the Golgi Apparatus. (Invited Review) Trends in Cell Biology 8:34-40, 1998.


A090. Srikrishna, G., Varki, N.M., Newell, P.C., Varki, A. and Freeze, H.H. An IgG monoclonal antibody against Dictyostelium discoideum glycoproteins specifically recognizes Fucα1,6GlcNAcβ in the core of N-linked glycans: localized expression of core fucosylated glycoconjugates in human tissues. J Biol Chem. 272:25743-25752. 1997.

A089. Krishna, M. and Varki, A. 9-O-acetylation of sialomucins: a novel marker of murine CD4 T cells that is regulated during maturation and activation. J Exp Med. 185:1997-2013. 1997. 

A088. Koenig, A., Jain, R., Vig, R., Norgard-Sumnicht, K., Matta, K.L. and Varki, A. Selectin inhibition. Synthesis and evaluation of novel sialylated, sulfated and fucosylated oligosac-charides, including the major capping group of GlyCAM-1. Glycobiology 7:79-93. 1997.

B045. Kim, Y.J. and Varki, A. Perspective on the significance of altered glycosylation of glycoproteins in cancer. (Invited Review). Glycoconjugate J. 14:569-576. 1997.

B044. Varki, A. Sialic acids as ligands in recognition phenomena. (Invited Review) FASEB J. 11:248-255. 1997

B043. Varki, A., Kagnoff, M.F., and Insel, P.A. Passing the baton at high speed: time to hand over to a new editorial board (Editorial) J Clin Invest. 99:553-554. 1997.

B042.2. Varki, A. Selectin ligands. Will the real ones please stand up? (Invited Perspective) J Clin Invest 99:158-162. 1997.

B039. Toomre, D.K. and Varki, A. Simultaneous fluorescent labeling and biotinylation of oligo-saccharides: a versatile approach to the analysis of oligosaccharides structure and function. IN A Laboratory Guide to Glycoconjugate Analysis, Jackson, P. and Gallagher, J.T. (eds). Birkhauser, Basel, Switzerland. 1997.


B042.1.  Varki, A.P. The times they are still a’changing: keeping up with the times. J Clin Invest. 97:1. 1996.

A087. Hicke, B.J., Watson, S.R., Koenig, A., Lynott, C.K., Bargatze, R.F., Chang, Y-F., Ringquist, S., Moon-McDermott, L., Jennings, S., Fitzwater, T., Han, H-L, Varki, N., Albinana, I., Willis, M.C., Varki, A., and Parma, D. DNA aptamers block L-selectin function in vivo - Inhibition of human lymphocyte trafficking in SCID mice. J Clin Invest. 98:2688-2692. 1996.

A086. Shi, W-X., Chammas, R., and Varki, A. Regulation of sialic acid 9-O-acetylation during the growth and differentiation of murine erythroleukemia cells. J Biol Chem. 271:31517-31525. 1996.

A085. Shi, W-X., Chammas, R., Varki, N.M., Powell, L., and Varki, A. Sialic acid 9-O-acetylation on murine erythroleukemia cells affects complement activation, binding to I-type lectins, and tissue homing. J Biol Chem. 271:31526-31532. 1996.

A084. Chammas, R., McCaffery, J. M., Klein, A., Ito, Y., Saucan, L., Palade, G., Farquhar, M.G., and Varki, A. Uptake and incorporation of an epitope-tagged sialic acid donor into intact rat liver Golgi compartments. Functional localization of sialyltransferase overlaps with beta-galactosyl-transferase but not with sialic acid O-acetyltransferase. Mol Biol Cell. 7:1691-1707. 1996.

A083. Shi, W.X., Chammas, R., and Varki, A. Linkage-specific action of endogenous sialic acid O-acetyltransferase in Chinese hamster ovary cells. J Biol Chem. 271:15130-15138. 1996.

A082. Guimarães, M.J., Bazan, J.F., Castagnola, J., Diaz, S., Copeland, N.G., Gilbert, D.J., Jenkins, N.A., Varki, A., and Zlotnik, A. Molecular cloning and characterization of lysosomal sialic acid O-acetylesterase. J Biol Chem. 271:13697-13705. 1996.

A081. Crottet, P., Kim, Y., and Varki, A. Subsets of sialylated, sulfated mucins of diverse origins are recognized by L-selectin. Lack of evidence for unique oligosaccharide sequences mediating binding. Glycobiology 6:191-208. 1996.

A080. O'Connell, D., Koenig, A., Jennings, S., Hicke, B., Han, H-L., Fitzwater, T., Chang, Y-F., Varki, N., Parma, D., and Varki, A. Calcium-dependent oligonucleotide antagonists specific for L-selectin. Proc Natl Acad Sci USA. 93:5883-5887. 1996.
B041. Ginsberg, M.H., Ruggeri, Z.M., and Varki, A. Cell adhesion in vascular biology: Series introduction. J Clin Invest. 98:1505. 1996.

B040. Varki, A. "Unusual" modifications and variations of vertebrate oligosaccharides: Are we missing the flowers for the trees?Glycobiology 6:707-710. 1996. 

B038. Varki, A. Does DG42 synthesize hyaluronan or chitin?: A controversy about oligosaccharides in vertebrate development. Proc Natl Acad Sci USA. 93:4523-4525. 1996.

B028. Hayes B.K., and Varki, A. Fractionation and analysis of neutral oligosaccharides by HPLC. John Wiley & Sons, New York. 0:17-21B. 1996. 

B027. Varki, A. Analysis of oligosaccharide negative charge by anion-exchange chromatography. John Wiley & Sons, New York. 0:17-20. 1996. 

B026. Manzi, A., Powell, L.D., and Varki, A. Thiobarbituric acid assay for sialic acids with spectrophotometric of HPLC detection.John Wiley & Sons, New York. 0:17-Unit 18. 1996.


A079. Norgard-Sumnicht, K.E., L. Roux, L., Toomre, D., Manzi, A., Freeze. H.H., and Varki, A. Unusual anionic N-linked oligosaccharides from bovine lung. J Biol Chem. 270:27634-27645. 1995.

A078. Mannori, G., Crottet, P., Cecconi, O., Hanasaki, K., Aruffo, A., Nelson, R.M., Varki, A. and Bevilacqua, M.P. Differential colon cancer cell adhesion to E-, P-, and L-selectin: Role of mucin type glycoproteins. Cancer Research 55:4425-4431. 1995.

A077. Hayes, B.K., Esquivel, F., Bennink, J.R., Yewdell, J. and Varki, A. Structure of the N-linked oligosaccharides of MHC class I molecules from cells deficient in the antigenic peptide transporter - Implications for the site of peptide association. J Immunol. 7:3780-3787. 1995.

A076. Norgard-Sumnicht, K., and Varki, A. Endothelial heparan sulfate proteoglycans that bind to L-selectin have glucosamine residues with unsubstituted amino groups. J Biol Chem. 270:12012-12024. 1995.

A075. Butor, C., Griffiths, G., Aronson, N.N. Jr., and Varki, A. Co-localization of hydrolytic enzymes with widely disparate pH optima: Implications for the regulation of lysosomal pH. J Cell Sci. 108:2213-2219. 1995. 

A074. Hanasaki, K., Powell, L.D., and Varki, A. Binding of human plasma sialoglycoproteins by the B cell-specific lectin CD22. Selective recognition of immunoglobulin M and haptoglobin. J Biol Chem. 270:7543-7550. 1995.

A073. Hanasaki, K., Varki, A., and Powell, L.D CD22-mediated cell adhesion to cytokine-activated human endothelial cells. Positive and negative regulation by alpha2-6 sialylation of cellular glycoproteins. J Biol Chem. 270:7533-7542. 1995.

A072. Powell, L.D., Jain, R.K., Matta, K.L., Sabesan, S., and Varki, A. Characterization of sialyloligosaccharide binding by recombinant soluble and native cell-associated CD22. Evidence for a minimal structural recognition motif and the potential importance of multisite binding. J Biol Chem. 270:7523-7532. 1995.

A071. Sjoberg, E.R., Chammas, R., Ozawa, H., Kawashima, I., Khoo, K-H, Morris, H.R., Dell, A., Tai, T., and Varki, A. Expression of de-N-acetyl-gangliosides in human melanoma cells is induced by genistein or nocodazole. J Biol Chem. 270:2921-2930. 1995.

B037. Powell, L.D., and Varki, A. I-Type lectins. J Biol Chem. 270:14243-14246. 1995.

B036. Varki, A. Neoglycobiology. Trends in Biochemical Sci. 20:214. 1995.

B035. Varki, A., and Marth, J. Oligosaccharides in vertebrate development. Semin Dev Biol. 6:127-138. 1995.


A070. Powell, L.D., Paneerselvam, K., Vij, R., Diaz, S., Manzi, A., Buist, N., Freeze, H., and Varki, A. Carbohydrate-deficient glycoprotein syndrome: Not an N-linked oligosaccharide processing defect, but an abnormality in lipid-linked oligosaccharide biosynthesis? J Clin Invest. 94:1901-1909. 1994.

A069. Manzi, A.E., Higa, H.H., Diaz, S., and Varki, A. Intramolecular self-cleavage of polysialic acid. J Biol Chem. 269:23617-23624. 1994.

A068. Toomre, D., and Varki, A. Advances in the use of biotinylated diaminopyridine (BAP) as a versatile fluorescent tag for oligosaccharides. Glycobiology 4:653-664. 1994.

A067. Sjoberg, E.R., Powell, L.D., Klein, A., and Varki, A. Natural ligands of the B cell adhesion molecule CD22 beta can be masked by 9-O-acetylation of sialic acids. J Cell Biol. 126:549-562. 1994.

A066. Klein, A., Krishna, M., Varki, N.M. and Varki, A. 9-O-acetyl-sialic acids have widespread but selective expression: analysis using a chimeric dual-function probe derived from Influenza C hemagglutinin-esterase. Proc Natl Acad Sci USA. 91:7782-7786. 1994.

A065. Hanasaki, K., Varki, A., Stamenkovic, I., and Bevilacqua, M. Cytokine-induced beta-galactoside alpha-2,6-sialyltransferase in human endothelial cells mediates alpha 2.6-sialylation of adhesion molecules and CD22 ligands. J Biol Chem. 269:10637-10643. 1994.

A064. Powell, L.D., and Varki, A. The oligosaccharide binding specificities of CD22beta, a sialic acid-specific lectin of B cells. J Biol Chem. 269:10628-10636. 1994.

B034. Varki, A. The screening review system: Fair or Foul? J Clin Invest. 93:1871-1874. 1994. 

B033. Varki, A. Res Ipsa Loquitur: Positive and negative trends affecting the JCI. J Clin Invest. 93:1. 1994. 

B032. Varki, A. Selectin ligands. Proc. Natl Acad Sci USA. 91:7390-7397. 1994.

B031. Manzi, A.E., and Varki, A. Compositional Analysis of Glycoproteins. Oxford University Press, Oxford. 0:27-77. 1994.

B030. Varki, A. Metabolic radiolabelling of glycoconjugates. Methods Enzymol. 230:16-32. 1994. 

B029. Varki, A., and Freeze, H.H. The Major Glycosylation Pathways of Mammalian Membranes; a summary. Plenum Press, New York.  0:71-100. 1994.


A063. Rothenberg, B., Hayes, B.K., Toomre, D., Manzi, A.E., and Varki, A. Biotinylated diaminopyridine: An approach to tagging oligosaccharides and exploring their biology. Proc Natl Acad Sci USA. 90:11939-11943. 1993.

A062. Norgard, K.E., Varki, N.M., and Varki, A. Calcium-dependent heparin-like ligands for L-selectin in non-lymphoid endothelial cells. Science. 261:480-483. 1993.

A061. Hayes, B. K., and Varki, A. The biosynthesis of oligosaccharides in intact Golgi preparations from rat liver. Analysis of N-linked and O-linked glycans labeled by UDP-[6-3H]N-acetyl-galactosamine. J Biol Chem. 268:16170-16184. 1993. 

A060. Hayes, B. K., and Varki, A. Biosynthesis of oligosaccharides in intact Golgi preparations from rat liver. Analysis of N-linked glycans labeled by UDP-[6-3H] galactose, CMP-[9-3H]N-acetylneuraminic acid, and [acetyl-3H]acetyl-coenzyme A. J Biol Chem. 268:16155-16169. 1993.

A059. Hayes, B. K. , Freeze, H. H., and Varki, A. Biosynthesis of oligosaccharides in intact Golgi preparations from rat liver. Analysis of N-linked glycans labeled by UDP-[6-3H]N-acetyl-glucosamine. J Biol Chem. 268:16139-16154. 1993.

A058. Norgard, K.E., Moore, K.L., Diaz, S., Stults, N.L., Ushiyama, S., McEver, R.P., Cummings, R.D., and Varki, A. Characterization of a specific ligand for P-selectin on myeloid cells. A minor glycoprotein with sialylated O-linked oligosaccharides. J Biol Chem. 268:12764-12774. 1993.

A057. Butor, C., Higa, H.H., and Varki, A. Structural, immunological, and biosynthetic studies of a sialic acid-specific O-acetylesterase from rat liver. J Biol Chem. 268:10207-10213. 1993.

A056. Butor, C., Higa, H.H., and Varki, A. High level O-acetylation of sialic acids on N-linked oligosaccharides of rat liver membranes. Differential subcellular distribution of 7- and 9-O-acetyl groups and of enzymes involved in their regulation. J Biol Chem. 268:10197-10206. 1993.

A055. Sjoberg, E.R., and Varki, A. Kinetic and spatial interrelationships between ganglioside glycosyltransferases and O-acetyltransferase(s) in human melanoma cells. J Biol Chem. 268:10185-10196. 1993.

A054. Powell, L.D., Sgroi, D., Sjoberg, E.R., Stamenkovic, I., and Varki, A. Natural ligands of the B cell adhesion molecule CD22beta carry N-linked oligosaccharides with alpha-2,6-linked sialic acids that are required for recognition. J Biol Chem. 268:7019-7027. 1993.

A053. Sgroi, D., Varki, A., Braesch-Andersen, S., and Stamenkovic, I. CD22, a B cell-specific immunoglobulin superfamily member, is a sialic acid-binding lectin. J Biol Chem. 268:7011-7018. 1993.

A052. Norgard, K., Han, H., Powell, L., Kriegler, M., Varki, A., and Varki, N.M. Enhanced interaction of L-selectin with the high endothelial venule ligand via selectively oxidized sialic acids. Proc Natl Acad Sci USA. 90:1068-1072. 1993.

A051. Freeze, H.H., Sampath, D., and Varki, A. Alpha and beta xylosides alter glycolipid synthesis in human melanoma and Chinese hamster ovary cells. J Biol Chem. 268:1618-1627. 1993.

B025. Varki, A. Biological roles of oligosaccharides: all of theories are correct. Glycobiology. 3(2):97-130. 1993.

B024. Powell, L.D., and Varki, A. Sialidases. Analysis of Glycoconjugates, Current Protocols in Molecular Biology, Unit 17.13A, F. Ausubel et al. Eds., Greene Publishing/Wiley Interscience. 17.12.2–17.12.8. 1993. 

B023. Diaz, S., and Varki, A. Metabolic Radiolabelling of Animal Cell Glycoconjugates. Analysis of Glycoconjugates, Current Protocols in Molecular Biology, Unit 17.4, F. Ausubel et al. Eds., Greene Publishing/Wiley Interscience. 17.4.2–17.4.16. 1993. 

B022. Varki, A., Freeze, H.H., and Manzi, A.E. Preparation and Analysis of Glycoconjugates: General Introduction, Overview, Choice of Techniques, and Glossary. Analysis of Glycoconjugates, Current Protocols in Molecular Biology, Unit 17.01, F. Ausubel et al. Eds., Green Publishing/Wiley Interscience. 17.0.5–17.0.14. 1993. 

B021. Varki, A., and Higa, H.H. Studies of the O-acetylation and (in)stability of polysialic acid, In Polysialic Acids. J. Roth (ed.), Birkhauser Verlag AG, Zurich. 165-170 1993. 

B020. Varki, A. The Editorial Policies and Practices of the Journal of Clinical Investigation. J Clin Invest. 91:4-6. 1993.


A050. Sjoberg, E.R., Manzi, A.E., Khoo, K-H., Dell, A., and Varki, A. Structural and immunological characterization of O-acetylated GD2. Evidence that GD2 is an acceptor for ganglioside O-acetyltransferase in human melanoma cells. J Biol Chem. 267:16200-16211. 1992.

A049. Moore, K.L., Stults, N.L., Diaz, S., Smith, D.E., Cummings, R.D., Varki, A., and McEver, R.P. Identification of a specific glycoprotein ligand for P-selectin (CD62) on myeloid cells. J Cell Biol. 118:445-456. 1992.

A048. Sampath, D., Varki, A., and Freeze, H.H. The spectrum of incomplete N-linked oligosaccharides synthesized by endothelial cells in the presence of Brefeldin A. J Biol Chem. 267:4440-55. 1992.

A047. Hayes, B., and Varki, A. The synthesis of high-specific-activity UDP-[6-3H]galactose, UDP-N-[6-3H] acetylgalactosamine, and their corresponding monosaccharides. Anal Biochem. 201:140-145. 1992.

B019. Varki, A. Of pride, prejudice and discrimination: why generalizations can be unfair to the individual. Annal Int Med. 116:762-764. 1992.

B018. Varki, A. The times they are a-changin’: changing with the times. J Clin Invest. 89:721-722. 1992.

B017. Varki, A. Selectins and other mammalian sialic acid-binding lectins. Curr Opin Cell Biol. 4:257-266. 1992. 

B016. Varki, A. Diversity in the sialic acids. Glycobiology. 2:25-40. 1992.


A046. Zhou, Q., Moore, K.L., Smith, D.F., Varki, A., McEver, R.P., and Cummings, R.D. The selectin GMP-140 binds to sialylated, fucosylated lactosaminoglycans on both myeloid and nonmyeloid cells. J Cell Biol. 115:557-564. 1991.

A045. Varki, A., Hooshmand, F., Diaz, S., Varki, N.M., and Hedrick, S.M. Developmental abnormalities in transgenic mice expressing sialic acid-specific 9-O-acetylesteraseCell. 65:65-74. 1991.

A044. Moore, K.L., Varki, A., and McEver, R.P. GMP-140 binds to a glycoprotein receptor on human neutrophils: Evidence for a lectin-like interaction. J Cell Biol. 112:491-499. 1991.

B015. Bevilacqua, M., Butcher, E., Furie,B., Gallatin,M., Gimbrone, M., Harlan, J., Kishimoto, K., Lasky, L., McEver, R., Paulson, J., Rosen, S., Seed, B., Siegelman, M., Springer,T., Stoolman, L., Tedder, T., Varki, A., Wagner, D., Weissman, I., and Zimmerman, G. Selectins: A family of adhesion receptors. Cell. 67:233. 1991.

B014. Varki, A. The Selectins - a family of sialic-acid binding lectins directly involved in cell-cell adhesion. Trends in Glycoscience and Glycotechnology (TIGG). 3:122-125. 1991. 

B013. Varki, A. De-N-acetyl gangliosides: do they exist? If so, how are they made? What do they do? Trends in Glycoscience and Glycotechnology (TIGG). 3:4-9. 1991.

B012. Varki, A. Role of oligosaccharides in the intracellular and intercellular trafficking of glyco- proteins. Cell Surface Carbohydrates and Cell Development, M. Fukuda (ed). CRC Press. 2:25-69. 1991.

B011. Varki, A. Radioactive tracer techniques in the sequencing of glycoprotein oligosaccharides. FASEB J. 5:226-235. 1991.


A043. Manzi, A.E., Sjoberg, E., Diaz, S.L., and Varki, A. Biosynthesis and turnover of O-acetyl and N-acetyl groups in the gangliosides of human melanoma cells. J Biol Chem. 265:13091-13103. 1990.

A042. Manzi, A.E., Diaz, S.L., and Varki, A. High-pressure liquid chromatography of sialic acids on a pellicular resin anion-exchange column with pulsed amperometric detection: A comparison with six other systems. Anal Biochem. 188:20-32. 1990.

A041. Manzi, A., E., Dell, A., Azadi, P., and Varki, A. Studies of naturally occurring modifications of sialic acids by fast-atom bombardment-mass spectrometry. Analysis of positional isomers by periodate cleavage. J Biol Chem. 265:8094-8107. 1990.


A040. Muchmore, E.A., Milkowski, M., Varki, A., and Diaz, S. Biosynthesis of N-glycolyneuraminic acid. The primary site of hydroxylation of N-acetylneuraminic acid is the cytosolic sugar nucleotide pool. J Biol Chem. 264:20216-20223. 1989.

A039. Hayes, B., and Varki, A. O-Acetylation and de-O-acetylation of sialic acids. Sialic acid esterases of diverse evolutionary origins have serine active sites and essential arginine residues. J Biol Chem. 264:19443-19448. 1989.

A038. Higa, H., Manzi, A., and Varki, A. O-Acetylation and de-O-acetylation of sialic acids. Purification, characterization, and properties of a glycosylated rat liver esterase specific for 9-O-acetylated sialic acids. J Biol Chem. 264:19435-19442. 1989.

A037. Higa, H., Butor, C., Diaz, S., and Varki, A. O-Acetylation and de-O-acetylation of sialic acids. O-Acetylation of sialic acids in the rat liver Golgi apparatus involves an acetyl intermediate and essential histidine and lysine residues--A transmembrane reaction? J Biol Chem. 264:19427-19434. 1989.

A036. Diaz, S., Higa, H., Hayes, B.K., and Varki, A. O-acetylation and de-O-acetylation of sialic acids. 7- and 9-O-acetylation of alpha2,6-linked sialic acids on endogenous N-linked glycans in rat liver Golgi vesicles. J Biol Chem. 264:19416-19426. 1989.

B010. Diaz, S., Higa, H., and Varki, A. Glycoprotein sialate 7(9)-O-acetyltransferase from rat liver Golgi vesicles. Methods Enzymol. 179:416-421. 1989.

B009. Higa, H., Manzi, A., Diaz, S., and Varki, A. Sialate 9-O-acetylesterase from rat liver. Methods in Enzymol. 179:409-415. 1989.


A035. Higa, H., and Varki, A. Acetyl-coenzyme A: polysialic acid O-acetyltransferase from K1-positive Esherichia coli: the enzyme responsible for the O-acetyl plus phenotype and O-acetyl form variation. J Biol Chem. 263:8872-8878. 1988.

A034. Sundblad G., Sajiji, S., Quaranta, V., Freeze, H.H., and Varki, A. Sulfated N-linked oligosaccharides in mammalian cells III: Characterization of a 140 kDa pancreatic tumor cell surface glycoprotein with N- and O-sulfate esters on asparagine-linked glycans. J Biol Chem. 263:8897-8903. 1988.

A033. Sundblad, G., Holoyda, S., Roux, L., Varki, A., and Freeze, H.H. Sulfated N-linked oligosaccharides in mammalian cells II: Identification of glycosaminoglycan-like chains attached to complex-type glycans. J Biol Chem. 263:8890-8896. 1988.

A032. Roux, L., Holoyda, S., Sundblad, G., Freeze, H.H., and Varki, A. Sulfated N-linked oligosaccharides in mammalian cells I: Complex-type chains with sialic acids and O-sulfate esters. J Biol Chem. 263:8879-8889 1988.


A031. Muchmore, E., Varki, N., Fukuda, M., and Varki, A. Developmental regulation of sialic acid modifications in rat and human colon. FASEB J. 1:229-235. 1987.

A030. Higa, H., Diaz, S., and Varki, A. Biochemical and genetic evidence for distinct membrane-bound and cytosolic sialic acid O-Acetyl-Esterases: Serine-Active-Site EnzymesBiochem Biophys Res Commun. 144:1099-1108. 1987.

A029. Muchmore, E., and Varki, A. Selective inactivation of influenza C esterase: a probe for 9-O-acetylated sialic acids. Science. 236:1293-1295. 1987.


A028. Fukuda, M.N., Dell, A., Tiller, P.R., Varki, A., Klock, J.C., and Fukuda, M. Structure of a novel sialylated fucosyl lacto-N-nor-hexaosylceramide isolated from chronic myelogeneous leukemia cells. J Biol Chem. 260:2376-2383. 1986.

A027. Varki, A., Muchmore, E. and Diaz, S. A sialic acid-specific O-acetylesterase in human erythrocytes: possible identity with esterase D, the genetic marker of retinoblastomas and Wilson disease. Proc Natl Acad Sci USA. 83:882-886. 1986.

A026. Freeze, H.H., and Varki, A. Endo-Glycosidase F and peptide N-glycosidase F release the great majority of total cellular N-linked oligosaccharides: use in demonstrating that sulfated N-linked oligosaccharides are frequently found in cultured cells. Biochem Biophys Res Commun. 140:967-973. 1986.


A025. Fukuda, M.N., Fukuda, M.,Bothner, B., Ramsamooj, P., Dell, A., Tiller,P.R., Varki, A., and Klock, J.C. Structures of sialylated fucosyl polylactosaminoglycans isolated from chronic myelogenous leukemia cells. J Biol Chem. 260:12957-12967. 1985.

A024. Diaz, S., and Varki, A. Metabolic labelling of sialic acids in tissue culture cell lines: methods to identify substituted and modified radioactive neuraminic acids. Anal Biochem. 150:32-46. 1985.

A023. Varki, A., and Diaz, S. The transport and utilization of acetyl-coenzyme A by rat liver Golgi vesicles. O-acetylated sialic acids are a major product. J Biol Chem. 260:6600-6608. 1985.


A022. Cheresh, D.A., Reisfeld, R.A., and Varki, A. O-acetylation of disialoganglioside GD3 by human melanoma cells creates a unique antigenic determinant. Science. 225:844-846. 1984.

A021. Cheresh, D.A., Varki, A., Varki, N.M., Stallcup, W.B., Levine, J., and Reisfeld, R.A. A monoclonal antibody recognizes an O-acylated sialic acid in a human melanoma-associated ganglioside. J Biol Chem. 259:7453-7459. 1984. 

A020. Varki, A., and Diaz, S. The release and purification of sialic acids from glycoconjugates: methods to minimize the loss and migration of O-acetyl groups. Anal Biochem. 137:236-247. 1984.

B008. Howard, R.F., Varki, A., and Reese, R.T. Merozoite proteins synthesized in P. Falciparum schizonts. Prog Clin Biol Res. 155:45-61. 1984.


A019. Varki, A., and Diaz, S. A neuraminidase from Streptococcus sanguis that can release O-acetylated sialic acids. J Biol Chem. 258:12465-12471. 1983.

A018. Varki, A., Lottenberg, R., Griffith, R., and Reinhard, E. The syndrome of idiopathic myelofibrosis. A clinicopathologic review with emphasis on the prognostic variables predicting survival. Medicine (Baltimore). 62:353-371. 1983.

A017. Varki, A., and Kornfeld, S. The spectrum of anionic oligosaccharides released by endo-beta-N-acetylglucosaminidase H from glycoproteins. Structural studies and interactions with the phosphomannosyl receptor. J Biol Chem. 258:2808-2818. 1983.

A016. Varki, A., Sherman, W., and Kornfeld, S. Demonstration of the enzymatic mechanisms of alpha-N-acetyl-D-glucosamine-1-phosphodiester N-acetylglucosaminidase (formerly called alpha-N-acetylglucosaminylphosphodiesterase) and lysosomal alpha-N-acetylglucosaminidase. Arch Biochem Biophys. 222:145-149. 1983.

B007. Varki, A., Reitman, M.L., Tabas, I, and Kornfeld, S. Studies of the synthesis, structure and function of the phosphorylated oligosaccharides of lysosomal enzymes. Biosci. 5:101-104. 1983.

B006. Clinicopathological Conferences: Washington U. SOM. Neurologic deterioration and death in a 27-year-old man with Chondrosarcoma. Amer J Med. 74:663-671. 1983. 

B005. Clinicopathological Conferences: Washington University SOM. Anemia and thrombocytopenia in a 63-year-old man. Amer J Med. 74:502-506. 1983.

B003. Varki, N., and Varki, A. Anemia and red cell transfusion. (Chapter 15), Man Med Therap. 24th edition:15-10. 1983.


A015. Thomas, G.H., Miller, C.S., Toomey, K.E., Reynolds, L.W., Reitman,M.L., Varki, A., Vannier, A., Rosenbaum, K.N., Bias, W.B., and Schofield, B.H. Two clonal cell populations (mosaicism) in a 46, XY male with mucolipidosis II (I-cell disease)--an autosomal recessive disorder. Am J Hum Genet. 34:611-622. 1982.

A014. Varki, A., Reitman, M.L., Vannier, A., Kornfeld, S., Grubb, J.H., and Sly, W.S. Demonstration of the heterozygous state for I-cell disease and pseudo-Hurler polydystrophy by assay of N-acetylglucosaminyl-phosphotransferase in white blood cells and fibroblasts. Am J Hum Genet. 34:717-729. 1982.

B004. Clinicopathological Conferences: Washington University SOM. Life threatening infection in an older man with leukemia. Amer J Med. 72:688-694. 1982.

B002. Kornfeld, S., Reitman, M.L., Varki, A., Goldberg D., and Gabel, C.A. Steps in the phosphorylation of the high mannose oligosaccharides of lysosomal enzymes. Ciba Found Symp. 92:138-156. 1982.


A013. Varki, A., Reitman, M.L., and Kornfeld, S. Identification of a variant of mucolipidosis III (pseudo-Hurler polydystrophy): a catalytically active N-acetylglucosaminylphosphotransferase that fails to phosphorylate lysosomal enzymes. Proc Natl Acad Sci USA. 78:7773-7777. 1981.

A012. Varki, A., and Kornfeld, S. Purification and characterization of rat liver alpha-N-acetylglucosaminyl phosphodiesterase. J Biol Chem. 256:9937-9943. 1981. 

A011. Reitman, M.L., Varki, A., and Kornfeld, S. Fibroblasts from patients with I-cell sisease and pseudo-Hurler polydystrophy are deficient in uridine 5'-diphosphate-N-acetylglucosamine: glycoprotein N-acetylglucosaminylphosphotransferase activity. J Clin Invest. 67:1574-1579. 1981.


A010. Varki, A., and Kornfeld, S. Structural studies of phosphorylated high mannose-type oligosaccharides. J Biol Chem. 255:10847-10858. 1980.

A009. Varki, A., and Kornfeld, S. Identification of a rat liver alpha-N-acetylglucosaminyl phosphodies-terase capable of removing "blocking" alpha-N-acetylglucosamine residues from phosphorylated high mannose oligosaccharides of lysosomal enzymesJ Biol Chem. 255:8398-8401. 1980.

A008. Varki, A., and Kornfeld, S. An autosomal dominant gene regulates the extent of 9-O-acetylation of murine erythrocyte sialic acids. A probable explanation for the variation in capacity to activate the alternate complement pathway. J Exp Med. 152:532-544. 1980.

A007. Davis, H.D., Varki, A., Heaton, W. A., and Siegel, B.A. Detection of accessory spleens with indium-111 labelled autologous platelets. Am J Hematol. 8:81-86. 1980.


A006. Varki, A., and Puthuran, P. Value of second lumbar puncture in confirming a diagnosis of aseptic meningitis. A prospective study. Arch Neurol. 36:581-582. 1979.

A005. Varki, A., Fritz, J.L., and Davis, R.B. Effects of cervical dislocation on colony-forming cells in murine marrow culture. Exp Hematol. 7:397-400. 1979.

A004. Varki, A., Armitage, J.O., and Feagler, J.R. Typhlitis in acute leukemia: successful treatment by early surgical intervention. Cancer. 43:695-697. 1979.

B001. Clinicopathological Conferences: Washington University SOM. Fever, hepatic lesions and ascites. Amer J Med. 67:105-112. 1979.


A003. Varki, A., Roby, D.S., Watts, H., and Zatuchni, J. Serum myoglobin in acute myocardial infarction: a clinical study and review of the literature. Amer Heart J. 96:680-688. 1978.


A002. Goldkrand, J.W., Varki, A., and McClurg, J.E. Rapid prediction of pulmonary maturity by amniotic fluid lipid globule formation. Obstet Gynecol. 50:191-196. 1977.

A001. Goldkrand, J.W., Varki, A., and McClurg, J.E. Surface tension of amniotic fluid lipid extracts: prediction of pulmonary maturityAmer J Obstet Gynecol. 128:591-598. 1977.