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Anthropogeny Publications

*PDF links are provided are as a courtesy to Varki lab members and collaborators.


Axxx: Original Research Articles
Bxxx: Invited/Other Publications

Anthropogeny Publications
Red Meat Risk Publications


A296.Thiesler H., Gretenkort L., Hoffmeister L., Albers I., Ohlmeier L., Röckle I., Verhagen A., Banan R., Köpcke N., Krönke N., Feuerhake F., Behling F., Barrantes-Freer A., Mielke D., Rohde V., Hong B., Varki A., Schwabe K., Krauss J.K., Stadelmann C., Hartmann C., Hildebrandt H. Proinflammatory macrophage activation by the polysialic acid-Siglec-16 axis is linked to increased survival of glioblastoma patients. Clin Cancer Res. 29(12):2266-2279. Jun 2023.

B218. Angata T., Varki A. Discovery, classification, evolution and diversity of Siglecs. Mol Aspects Med.  2023 Apr. Epub 2022 Aug.

A292 (PDF). Saha, S., Khan, N., Comi, T., Verhagen, A., Sasmal, A., Diaz, S., Yu, H., Chen, X., Akey, J.M., Frank, M., Gagneux, P., and Varki, A. Evolution of Human-specific Alleles Protecting Cognitive Function of Grandmothers. Mol Biol Evol. 2022 Aug.

A288 (PDF). 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.

A286 (PDF). 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.

B203 (PDF). 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.

A281 (PDF). 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

A277 (PDF). 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 Material

A274 (PDF). 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. 2020.

B201 (PDF). Kawanishi, 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 loss. Proc Natl Acad Sci USA. 117(4):1847-48. 2020.

B200 (PDF). 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.

A272 (PDF). 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 Macrophages. J Innate Immun. 12(4):333-343. 2019.

A271 (PDF). 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

A270 (PDF). 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 U S A. 116:16036-16045. 2019. UCSD Press Release Medical News Today The Scientist Science Daily

B196 (PDF). 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.

A265 (PDF). 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. 12:337-349. 2019.

A262 (PDF). 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. 28510.1098/rspb. 2018. Featured in >100 news reports in many languages, including; UCSD Press Release, Science Magazine, Psychology Today, Smithsonian Magazine, Popular Science, Scientific American, Discover Magazine

A260 (PDF). 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:e1007133. 2018. UC San Diego News Room

A256 (PDF). 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.

A254 (PDF). 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.

A251 (PDF). 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 fossils. Proc Natl Acad Sci U S A. 114:E8155-E8164. 2017. UC San Diego Press Release

A250 (PDF). 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.

A247 (PDF). 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 (PDF). 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.

B193 (PDF). Varki, A. Are humans prone to autoimmunity? Implications from evolutionary changes in hominin sialic acid biology. (Invited Review). J Autoimmun. 83: 134-142, 2017.

A248 (PDF). 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 (PDF). 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.

B178 (PDF). Varki, A, 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.

B175 (PDF). 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 (PDF). Varki, A. Why are there no persisting hybrids of humans with Denisovans, Neanderthals, or anyone else. Proc Natl Acad Sci USA. 10.1073/pnas.1602270113. 2016.

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

A236 (PDF). 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 Natl Acad Sci USA, doi: 10.1073/pnas.1517951112. 2015. UCSD Press Release

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

B169 (PDF). 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.

A231 (PDF). 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

A229 (PDF). 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

A227 (PDF). 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.

B167 (PDF). 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, 2015.

B166 (PDF). 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, 2014.

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

A224 (PDF). 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.

A220 (PDF). 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.

A218 (PDF). 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 Primates. FASEB J. 28:1280-93, 2014.

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

A214 (PDF). 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 (PDF). 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.

A211 (PDF). 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 (PDF). 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.

A208 (PDF). 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 (PDF). O'Bleness, M., Serales, V.B., Varki, A., Gagneux, P., Sikela, J.M. Evolution of Genetic and Genomic Features Unique to the Human Lineage (Invited Review). Nat. Rev. Genet., 13:853-866, 2012.

A204 (PDF). Bergfeld, A.K. Pearce, O.M., Diaz, S.L., Pham, T., 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., 287:28868-28881, 2012.

A203 (PDF). Banda, K., Gregg, C.J., Chow, R., Varki, N.M., 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.

A201 (PDF). 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., Varki, A. Specific Inactivation of Two Immunomodulatory SIGLEC Genes During Human Evolution. Proc. Natl. Acad. Sci. U.S.A., 109:9935-9940, 2012.

B155 (PDF). Varki A. Nothing in Medicine Makes Sense, Except in the Light of Evolution (Invited Commentary). J. Mol. Med., 5:481-494, 2012.

B154 (PDF). 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 (PDF). Hayakawa, T. and Varki, A. Chapter 8: Human-specific Changes 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.

B151 (PDF). 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 (PDF). 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.

A197 (PDF). 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 alleles. Proc. Natl. Acad. Sci. U.S.A., 108:17743-17748, 2011.

A193 (PDF). Mitra, N., Banda, K., Altheide, T., Schaffer, L., Johnson-Pais, T.L., Beuten, J., Leach, R.J., Angata, T., Varki, N., 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 (PDF). 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 (PDF). 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.

B148 (PDF). Padler-Karavani, V., 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 (PDF). 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. Mech. Dis. 6:365-393, 2011. 

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

A188 (PDF). 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 (PDF). Ghaderi, D., R.E. Taylor, Padler-Karavani, V., Diaz, S., and Varki, A. Implications of the presence of N-glycolylneuraminic acid in recombinant therapeutic glycoproteins. Nat. Biotechnol., 28:863-7, 2010.

A186 (PDF). Taylor, R.E., Gregg, C.J., Padler-Karavani, V., Ghaderi, D., Yu, H., Huang, S., Sorenson, 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.

B143 (PDF). Varki A. Uniquely Human Evolution of Sialic Acid Genetics and Biology. Proc. Natl. Acad. Sci. U.S.A., 107 (S2): 8939-8946, 2010.

A182 (PDF). Soto, P., 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.

A180 (PDF). Pham, T.*, Gregg, C.*, Karp, F., Chow, R., Padler-Karavani, 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 (PDF). 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 (PDF). 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.

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

B133 (PDF). Loefling, 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.

A173 (PDF). 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.

A172 (PDF). Diaz, S.L., Padler-Karavani, V., Ghaderi, D., Hurtado-Ziola, N., Yu, H., 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.

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

A170 (PDF). Hedlund, M., Padler-Karavani, V., Varki, N., 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.

A169 (PDF). Winn, V.D., Gormley, M., Paquet, A.C., Kjar-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 Siglec-6 and Pappalysin-2. Endocrinology, 150:452-462, 2008.

A168 (PDF). Byres, E., Paton, A.W., Paton, J.C., Loefling, J.C., Smith, D.F., Wilce, M.C.J., Talbot, U.M., Chong, D.C., Yu, H., Huang, S., Chen, X., Varki, N.M., 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 (PDF). Varki, A., Geschwind, D., and Eichler, E.E. Explaining Human Uniqueness: Genome Interactions with Environment, Behavior and Culture. Nat. Rev. Genet., 9:749-763, 2008.

A166 (PDF). 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.

A161 (PDF). 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.

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

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

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

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

A155 (PDF). Angata, T., Hayakawa, T, Yamanaka, M., Varki, A., 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 (PDF). Nguyen, D.H., Hurtado-Ziola, N., Gagneux, P., Varki, A. Loss of Siglec Expression on T lymphocytes during Human Evolution. Proc. Natl. Acad. Sci. U.S.A., 103:7765-7770, 2006. Link to UCSD press release

A148 (PDF). Hayakawa T., Aki I., Varki A., Satta Y., 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.

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

B102 (PDF). Martin, M.J., Muotri, A., Gage, F.H., 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.

B099 (PDF). 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.

A147 (PDF). 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. Link to UCSD press release

A146 (PDF). 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. Link to UCSD press release

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

A143 (PDF). 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.

A141 (PDF). 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 (PDF). 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. Link to UCSD press release

B096 (PDF). 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.

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

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

B091 (PDF). 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 (PDF). 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.

A135 (PDF). 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.

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

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

B084 (PDF). 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 (PDF). 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 (PDF). 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. Supplementary material:, Link to UCSD press release

B081 (PDF). 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 (PDF). 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. Link to UCSD press release

A121 (PDF). 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. Link to UCSD press release

B078 (PDF). 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.

B076 (PDF). 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.

A117 (PDF). 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 (PDF). 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.

A113 (PDF). 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.

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

A106 (PDF). 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.

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

B067 (PDF). Gagneux, P., and Varki, A. Genetic Differences Between Humans and Great Apes. (Invited Review) Molecular Phylogenetics and Evolution 18:2-13, 2001.

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

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

B064 (PDF). 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.

A097 (PDF). 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. Link to UCSD press release

A096 (PDF). 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.

B048 (PDF). 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.