George R. Bousfield, PhD

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George R. Bousfield, PhD
Dr. Lawrence M. Jones Distinguished Professor
Protein Core Laboratory Director
Office:  431 Hubbard Hall
Phone: (316) 978-6088                                                         P01 Retreat Info 
E-mail: george.bousfield@wichita.edu 

Courses Taught
Research Interests
Professional Experience
Professional Organizations and Honors


Courses Taught

Number Course Name
BIOL 210 General Biology I
BIOL 310 Human Reproduction: Issues and Perspectives
BIOL 626 Reproductive Biology
BIOL 710 Glycobiology
BIOL 767 Mechanisms of Hormone Action



Institution and Location Degree Year Field of Study
Saginaw Valley State College, University Center, MI B.S. 1974 Chemistry & Biology
Indiana University, Bloomington, IN M.A. 1976 Zoology
Indiana  University, Bloomington, IN Ph.D. 1981 Zoology


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Research Interests

My laboratory is interested in the structure-function relations of the gonadotropins. These are members of the glycoprotein hormone family, which consists of the three gonadotropins, luteinizing hormone (LH), follicle-stimulating hormone (FSH), and chorionic gonadotropin (CG). The fourth member of this protein family is thyroid stimulating hormone (TSH). Together, the glycoprotein hormones form a unique subset of the cystine knot growth factor superfamily. While most other superfamily members are located at the C-terminal end of a larger precursor from which they are cleaved, and the N-terminal portion then acts as a regulatory factor, the glycoprotein subunits consist of only the cystine knot domains. The subunits are dissimilar proteins, a common alpha subunit that is differentially glycosylated in each hormone, as well as a hormone-specific beta subunit. Another unusual facet of glycoprotein hormone structure-function is these hormones are decorated with N-linked and sometimes O-linked glycans, which are necessary for function. The glycans can determine metabolic clearance rates of the hormones and at least some of them are necessary for biological activity, although the mechanism is unknown. Glycosylation varies in two ways: microheterogeneity and macroheterogeneity. The former results from a population of 2-105 glycans found at each N- or O-glycosylation site, while the latter results from partial glycosylation of the hormone-specific beta subunit. In humans an interesting pattern of glycosylation is found in FSH. The alpha subunit is always N-glycosylated, while the FSHΒ subunit appears to be glycosylated in an all-or-none manner. This produces a FSH glycoform that possesses four N-glycans that we call tetra-glycosylated hFSH and another form that possesses only the alpha subunit N-glycans that we call di-glycosylated hFSH. In young women, di-glycosylated hFSH is the more abundant glycoform present in the pituitary gland, however, there appears to be a progressive loss of this glycoform with increasing age. We are interested in determining if this can be used as a marker for reproductive aging.

 Awarded Grants

12/01/1989 - 11/30/1990 Carbohydrate Chemistry of Equine Gonadotoropins. M.D. Anderson Cancer Center Institutional Research Grant.
10/01- 08/31/1996 Role of Carbohydrate in FSH Specificity Determination.  National Institutes of Health, HD 29047.
01/01/1994 - 06/30/1994 Chimeric Gonadotropin Alpha Subunits. Wichita State University, University Research Grant.
08/01/1997 - 07/31/2000 Functional Consequences of Carbohyrate Differences Between eLH and eCG. National Institutes of Health, DK 52383.
02/01/2000 - 01/31/2001 Acquisition of a Protein Sequencer, NSF EPSCoR.
07/01/2001 - 06/30/2002 Adaptation of Adsorptive Sequencer to Solid-Phase Edman Degradation, Wichita State University, University Research Grant
07/01/2002 - 06/30/2003 Carbohydrate Modulation of Gonadotropin Action. National Institutes of Health/KUMC-KBRIN RR-16475, Joan Hunt, PI.
09/30/2002 to 08/31/2005 Structural Characterization of Glycoproteins using Mass Spectrometry. (Sub-contract, Heather Desaire, KU, PI) National Institutes of Health/KU-COBRE RR-017708 Bob Hanzlik PI
12/15/2002 - 08/31/2003 Replacement of Dionex Carbohydrate Analyzer Detector. National Institutes of Health/KUMC-KBRIN RR-16475, Joan Hunt PI.
09/01/03 - 06/30/2004 Replacement of Dionex Carbohydrate Analyzer Detector. National Institutes of Health/KUMC-KBRIN RR-16475, Joan Hunt PI.
10/01/2003 - 09/30/2004 Follicle-Stimulating Hormone Antagonist Inhibition of Prostate Tumor Cell Division, Flossie E. West Memorial Foundation.
12/19/2003 - 06/30/2004 Replacement Of Dionex High pH Liquid Chromatograph For Carbohydrate Analysis. National Institutes of Health/KUMC-KBRIN RR-16475, Joan Hunt PI.
03/01/2004 - 02/28/2005 Acquisition of Fluorescence and Phospho Imaging Instruments, NSF-EPSCoR, EPS-0236913, Thomas N. Taylor, Project Director.
07/01/2005 - 06/30/2006 Procise sequencer upgrade. National Institutes of Health/KUMC-KBRIN RR-16475, Joan Hunt PI.
04/01/2005 to 03/31/2006 Glycan Inhibition of LH and FSH Signaling. National Institutes of Health/KUMC-KBRIN RR-16475, Joan Hunt PI.
09/01/2005 to 12/31/2006 Gonadotropin Characterization, Serono, Inc., Rockland, MA,  Research Contract.
08/01/08 to 12/31/09 Monoclonal Antibody Production for Follicle Stimulating Hormone Glycoforms, Kansas Technology Enterprise Corporation
03/01/2009 to 02/28/2014 Aging Pituitary/Gonad Axis, National Institutes of Health/NIA, P01AG029531
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Professional Experience

1974 - 1977 Associate Instructor, Indiana University
1978 - 1979 Predoctoral Fellowship, Robert A. Welch Foundation, The University of Texas M.D. Anderson Hospital and Tumor Institute, Houston,TX
1979 - 1981 Research Assistant, The University of Texas M.D. Anderson Hospital and Tumor Institute, Houston, TX
1981 - 1983 Postdoctoral Fellowship, Robert A. Welch Foundation, The University of Texas M.D. Anderson Hospital and Tumor Institute, Houston, TX
1986 Visiting Scientist, National Institute of Immunology, New Dehli, India
1983 - 1987 Research Associate, The University of Texas M.D. Anderson Hospital and Tumor Institute, Houston, TX
1988 - 1990 Assistant Biochemist and Assistant Professor of Biochemistry and Molecular Biology (non-tenured research appointment), M.D. Anderson Cancer Center, Houston, TX
1990 Visiting Scientist, Institute of Physical and Chemical Research, RIKEN, Wako, Japan
1991 Visiting Assistant Professor, Department of Biological Sciences, Wichita State University, Wichita, KS
1991 - Present Director, Protein Chemistry Core Laboratory, Department of Biological Sciences, Wichita State University, Wichita, KS
1992 - 1997 1992-1997 Assistant Professor, Department of Biological Sciences, Wichita State University, Wichita, KS
1994 & 1994 Ad hoc member, NIH DRG, Biochemical Endocrinology Study Section
1994 & 1995 NIH, NICHD site visit
1995 - 2001 Scientist, Women’s Research Institute, University of Kansas Medical School at Wichita, Wichita, KS
1997 - 2005 Associate Professor, Department of Biological Sciences, Wichita State University, Wichita, KS
2005 - 2010 Professor, Department of Biological Sciences, Wichita State University, Wichita, KS
2010 - present Lawrence M. Jones Distinguished Professor, Department of Biological Sciences, Wichita State University, Wichita, KS
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  1. Teng, C.T., Teng, C-S., Bousfield, G.R., Liu, W-K., and Ward, D.N.:  "Differential Response  of Growing and Regressing Chicken Ovaries to Gonadotropic Hormones."  Gen. Comp. Endocrinol.  48:  325-332 (1982).
  2. Jirgensens, B., Bousfield, G.R., and Moore, W.T., Jr.,:  "Circular Dichroism and Conformational Properties of Modecin, A Toxic Protein."  J. Protein Chem.  2:  105-112   (1983).
  3. Bousfield, G.R., and Ward, D.N.: "Purification of Lutropin and Follitropin in High Yield from Horse Pituitary Glands."  J. Biol. Chem.  259: 1911-1921 (1984).
  4. Bousfield, G.R., Liu, W-K., and Ward, D.N.:  "Hybrids from equine LH: alpha enhances, beta diminishes activity."  Molec. Cell. Endocrinol.  40:  60-77 (1985).
  5. Bousfield, G.R., Sugino, H., and Ward, D.N.:  "Demonstration of a COOH-terminal Extension on Equine Lutropin by Means of a Common Acid-labile Bond in Equine Lutropin and Chorionic Gonadotropin."  J. Biol. Chem.  260:  9531-9533 (1985).
  6. Liu, W-K., Bousfield, G.R., Moore, W.T., Jr., and Ward, D.N.: Priming Procedure and  Hormone Preparations Influence Rat Granulosa Cell Response."  Endocrinology  116:  1454-1459 (1985).
  7. Bousfield, G.R. and  Ward, D.N.: "Direct Demonstration of Intrinsic Follicle-Stimulating Hormone Receptor-Binding Activity in Acid-treated Equine Luteinizing Hormone."  Biochim. Biophys. Acta  885:  327-334  (1986).
  8. Liehr, J.G., Stancel, G.M., Chorich, L.P., Bousfield, G.R., and Ulubelen, A.A.:  "Hormonal Carcinognesis:  Separation of Estrogenicity from Carcinogenicity."  Chem.-Biol.  Interactions  59:  173-184 (1986).
  9. Sugino, H., Bousfield, G.R., Moore, W.T., Jr., and Ward, D.N.: "Structural Studies on Equine Glycoprotein Hormones:  Amino acid Sequence of Equine Chorionic Gonadotropin Beta-Subunit."  J. Biol. Chem.  262: 8603-8609 (1987).
  10. Bousfield, G.R., Liu, W-K., Sugino, H., and Ward, D.N.:  "Structural Studies on Equine Glycoprotein Hormones:  Amino acid Sequence of Equine Lutropin Beta Subunit."  J. Biol. Chem. 262: 8610-8620 (1987).
  11. Bidart, J-M., Troalen, F., Salesse, Bousfield, G.R., Bohouon, C.J., and Bellet, D.H.:  "Topographic Antigenic Determinants Recognized by Monoclonal Antibodies on Human Choriogonadotropin Beta-Subunit."  J. Biol. Chem.  262: 8551-8556 (1987).
  12. Ward, D.N., Wen, T., and Bousfield, G.R.:  "Sulfate and Phosphate Analysis in Glycoproteins and Other Biologic Compounds Using Ion Chromatography:  Application to Glycoprotein Hormones and Sugar Esters."  J. Chromatog.  398:  255-264 (1987).
  13. Bidart, J-M., Troalen, F., Bousfield, G.R., Birken, S., and Bellet, D.: "Antigenic Determinants on Human Choriogonadotropin Beta-Subunit.  I.  Characterization of Topographic Sites Recognized by Monoclonal Antibodies."  J. Biol. Chem.  263:  10364-10369  (1988).
  14. Bousfield, G.R. and Ward, D.N.:  "Selective Proteolysis of Ovine Lutropin or its Beta Subunit by Endoproteinase Arg-C:  Properties of the Arg (Β-43) cleaved Hormone."  J. Biol. Chem.   263:  12602-12607 (1988).
  15. Ward, D.N., Bousfield,G.R., Gordon, W.L., and Sugino, H.:  "Chemistry of the Peptide Components of Glycoprotein Hormones."  In: Keel, B.A. and Grotjan, H.E., Jr., eds.  Microheterogeneity of Glycoprotein Hormones,  CRC Press, Boca Raton,  1988 , 1-21. 
  16. Ward, D.N., Hines, K.K., Gordon, W.L. and  Bousfield, G.R.: "The Purification of Native Inhibin and Chemical Characterization."  In: Hodgen, G.D., Rosenwaks, Z., and  Spieler, J.M., eds., Nonsteroidal Gonadal Factors: Physiological Roles and Possibilities in Contraceptive Development, The Jones Institute Press, Norfolk, 1988,  1-16.
  17. Bousfield, G. R., Liu, W-K. , and  Ward, D. N.: "Effects of Removal of Carboxyterminal Extension From Equine LH Beta-Subunit."   Endocrinology 124: 379-387 1989.
  18. Bidart, J., Troalen, F., Bousfield, G. R., Bohuon, C., & Bellet, D.  "Monoclonal Antibodies Directed to Human and Equine Chorionic Gonadotropins as Probes for the Topographic Analysis of Epitopes on the Human Beta-Subunit. " Endocrinology, 124:  923-929,  1989.
  19. Gordon, W. L., G. R. Bousfield and D. N. Ward   "Comparative binding of FSH to chicken and rat testis. " J. Endocrinol. Invest. 12: 383-392.  1989.
  20. Ward, D.N., Bousfield, G.R., and Mar, A.O. : "Chemical Reduction-Reoxidation of the Glycoprotein Hormone Disulfide Bonds."  In: Bellet, D. and  Bidart, J.-M., eds., Structure-Function Relationship of Gonadotropins, Raven Press, New York, 1989, 1-19.
  21. Moore, K.H., Dunbar, B.S., Bousfield, G.R., and Ward, D.N.:  "The Heterogeneity of  Porcine 32,000 Mr Inhibin alpha-subunit:  A Gel Electrophoresis and Immunoblot Study."  Endocrinology 127:  1477-1486, 1990.
  22. Ward, D.N., Bousfield, G.R., and Moore, K.H.:  "Gonadotropins."  In:  Cupps, P.T., Reproduction in Domestic Animals, 4th ed., Academic  Press, Orlando, Fl, 1990, pp 25-80.
  23. Matsui, T., Sugino, H., Miura, M., Bousfield, G.R., Ward, D.N., Titani, K., and Mizouchi, T.: "Beta-subunits of equine chorionic gonadotropin and luteinizing hormone with an identical amino acid sequence have different asparagine-linked oligosaccharide chains."  Biochem. Biophys. Res. Commun.  174: 940-945, 1991.
  24. Mizuno M., Farach–Carson M.C., Pinero G.J., Fujisawa R., Brunn J.C., Seyer J.M., Bousfield G.R., Mark M.P., Butler W.T. Identification of the rat bone 60K acidic glycoprotein as alpha2HS–glycoprotein. Bone Miner 13: 1-21,1992.
  25. Bousfield, G.R. and Ward, D.N.: “Reduction and reoxidation of equine gonadotropin alpha–subunits. Endocrinology  131: 2986-2998, 1992.
  26. Smith P.L., Bousfield G.R., Kumar S., Fiete D., Baenziger J.U.  “Equine lutropin and chorionic gonadotropin bear oligosaccharides terminating with SO4–4–GalNAc and siaα2,3Gal respectively”. J Biol Chem 268: 795-802, 1993
  27. Couture L., Lemonnier J.P., Troalen F., Roser J.F., Bousfield G.R., Bellet D., Bidart J.M. “Immunochemical studies of equine chorionic gonadotropin (eCG), eCG–α and eCG–Β.”. Endocrinology 132: 205-211, 1993
  28. Bousfield, G.R., Perry W.M., and Ward, D.N.  “Gonadotropins: chemistry and biosynthesis”. In: Knobil E., Neill J.D. (ed) The Physiology of Reproduction. Raven Press, New York,  pp 1749-1792, 1994
  29. Moore KH, Dunbar BS, Bousfield GR, Ward DN: “Initial Characterization of Equine Inhibin.” Biol Reprod 51: 63-71, 1994
  30. Bousfield, G.R. and Ward, D.N.  “Evidence for Two Folding Domains in Glycoprotein Hormone Alpha-Subunits” Endocrionology  135:  624-635, 1994
  31. Matsui, T., Mizuochi, T., Titani, K., Oinaga, T., Hoshi, M., Bousfield, G.R., Sugino, H., and Ward, D.N.  “Structural Analysis of N–Linked Oligosaccharides of Equine Chorionic Gonadotropin and Lutropin Beta–Subunits”  Biochemistry 33: 14039-14048, 1994
  32. Robert, P., Troalen, F., Bellet, D., Bousfield, G. R. and Bidart, J.-M. Immunochemical mapping of human lutropin: I. delineation of a conformational antigenic determinant. Mol. Cell. Endocrinol. 101: 11-20, 1994
  33. Butnev V.Y., Gotschall R.R., Baker V.L., Moore W.T., Bousfield G.R.: "Negative Influence of O–linked Oligosaccharides of High Molecular Weight Equine Chorionic Gonadotropin (eCG) on its LH and FSH Receptor Binding Activities." Endocrinology  137: 2530-2542, 1996
  34. Gotschall R.R. and Bousfield G.R.:  "Oligosaccharide mapping reveals hormone–specific glycosylation patterns on eqiune gonadotropin alpha-subunit Asn56." Endocrinology  137: 2543-2557, 1996
  35. Butnev V.Y., Gotschall R.R., Baker V.L., Moore, W.T., Gout, P.W. and Bousfield .GR.:  Glycosylated equine prolactin and its carbohydrate moiety. J Protein Chem 15: 413-426, 1996
  36. Peng, K.-C., Bousfield, G.R., Puett, D., and Ward, D.N.:  “Circular dichroic spectroscopy of Arg46-nicked ovine lutropin α and derived fragments.” J Protein Chem 15: 547-552, 1996
  37. Bousfield GR, Butnev VY, Gotschall RR, Baker VL, Moore WT:  Structural features of mammalian gonadotropins. Mol. Cell. Endocrinol. 125: 3–19, 1996
  38. Butnev VY, Gotschall RR, Butnev, VY, Baker VL, Moore WT, Bousfield GR:  Inhibitory influence of alpha-subunit Asn56 oligosaccharide on in vitro subunit association and FSH receptor binding of equine gonadotropins. Biol. Reprod. 58: 458–469, 1998
  39. Bousfield, G.R. LH (Luteinizing Hormone), in Encyclopedia of Reproduction, Knobil, E and Neill, JD, eds. Academic Press, San Diego,Vol. 2, 1034-1054, 1999
  40. Bousfield, G.R., Baker, V.L., Gotschall, RR., Butnev, V.Y., and Butnev, V.Y.: Carbohydrate Analysis of Glycoprotein Hormones. Methods. 21: 15-39, 2000
  41. Bousfield, G.R., Butnev, V.Y. and Butnev, V.Y.: Identifcation of 12 O-glycosylation sites in eLH and eCG alpha subunits by solid-phase Edman degradation. Biol. Reprod. 64: 136-147, 2001
  42. Walton, W.J., Nguyen, V.T., Butnev, V.Y., Singh, V., Moore, W.T., and Bousfield, G.R.:  Characterization of Human Follicle-Stimulating Hormone Isoforms Reveals a Non-Glycosylated Beta-subunit In Addition to the Conventional Glycosylated Alpha-subunit. J. Clin. Endocrinol. Metab. 86:3675-3585, 2001.
  43. Butnev, V.Y., Singh, V., Nguyen, V.T., and Bousfield, G.R.:  Truncated eLHΒ, Asparagine56-deglycosylated-eLH alpha Combine to Produce a Potent Follicle-Stimulating Hormone Antagonist.  J. Endocrinol. 172: 545-555, 2002
  44. Bousfield, G.R. and Butnev, V.Y.: Equine gonadotropins as models for glycosylation studies. Indian J. Exptl. Biology  40: 486-506, 2002.
  45. Nguyen VT, Singh V, Butnev VY, Gray CM, Westfall S, Davis JS, Dias JA, Bousfield GR. Inositol Phosphate Stimulation by LH Requires the Entire Alpha Asn56 Oligosaccharide. Molec. Cell. Endocrinol. 199: 73-86, 2003.
  46. Jiang H, Butnev VY, Bousfield GR, Desaire H. Glycoprotein profiling by electrospray mass spectrometry. J. Am. Soc. Mass Spectrom. 15: 750-758, 2004.
  47. Bousfield, G.R., Butnev, V.Y., Butnev, V.Y., Nguyen, V.T., Gray, C.M., Dias, J.A., MacColl, R., Eisele, L., and Harvey, D.J., “Differential Effects of Alpha Asparagine56 Oligosaccharide Structure on Equine Lutropin and Follitropin Hybrid Conformation and Receptor-Binding Activity.”  Biochemistry 43:  10817-10813, 2004.
  48. Harvey D.J., Bousfield G.R., Differentiation between sulphated and phosphated carbohydrates in low resolution MALDI mass spectra. Rapid Communications in Mass Spectrometry 19: 287-288, 2005
  49. Bousfield, G.R., Jia, L., and Ward, D.N. Gonadotropins: Chemistry and Biosynthesis. In: Neill J.D. (ed) Knobil and Neill: Physiology of Reproduction, 3rd edition. Elsevier Press, San Diego,  1581-1634, 2006
  50. Irungu, J.A., Dalpathado, D.S., Go, E.P., Jiang, H., Ha, H.-V., Bousfield, G.R., and Desaire, H., A Method for Characterizing Sulfated Glycoproteins in a Glycosylation Site-Specific Fashion, Using Ion-Pairing and Tandem Mass Spectrometry, Anal. Chem. 78: 11801-1190, 2006
  51. Dalpathado, D.S., Irungu, J.A., Go, E.P., Butnev, V.Y., Norton, K., Bousfield, G.R., and Desaire, H. Comparative glycomics of the glycoprotein follicle-stimulating hormone (FSH): Glycopeptide analysis of isolates from two mammalian species, Biochemistry, 45:8665-8673, 2006
  52. Bousfield, G.R., Butnev, V.Y., Walton, W.J., Nguyen, V.T., Huneidi, J., Singh, V., Kolli, K., Harvey, D.J., and Rance, N.E., All-or-None N-Glycosylation in Primate Follicle-Stimulating Hormone Beta-Subunits, Mol. Cell. Endocrinol. 260-262:  40-48, 2007
  53. Jablonka-Shariff , A, Roser, J.F., Bousfield, G.R., Wolfe, M.W., Sibley, L.E., Colgin, M., and Boime, I., Expression And Bioactivity Of A Single Chain Recombinant Equine Luteinizing Hormone (ReLH), Theriogenology  67:  311-320, 2007
  54. Bousfield, G.R., Butnev, V.Y., Bidart, J.M., Dalpathado, D., Irungu, J., and Desaire, H., Chromatofocusing fails to separate hFSH isoforms on the basis of glycan structure, Biochemistry 47:  1708-1720, 2008.
  55. Ulloa-Aguirre, A., Dias, J.A,. and Bousfield, G.R. Gonadotropins and the importance of glycosylation, In: Walsh, G. (ed) Posttranslational Modifications of Protein Biopharmaceuticals, pp 109-147, 2009.
  56. Wehbi, V., Tranchant, T., Durand, G., Musnier, A., Decourtye, J., Piketty, V., Butnev, V.Y., Bousfield, G.R., Crepieux, R., Maurel, M.C., and Reiter, E. Partially deglycosylated equine LH preferentially activates Beta-arrestin-dependent signaling at the follicle-stimulating hormone receptor, Mol. Endocrinol.  24: 561-573, 2010. 
  57. Visvanathan, M., Siddam, S.R., Lee, I.H., Lushington, G.H., Bousfield, G.R.  GlycomicsDB – a data integration platform for glycans and their structures. Open Med. Inform. J.  5:  9-16, 2011. PMC3098536
  58. Bousfield, G.R. and Dias, J.A., Synthesis and secretion of gonadotropins including structure-function correlates. Rev. Endocr. Metab. Disord. 12(4):  289-302, 2011.
  59. Ulloa-Aguirre, A., Dias, J.A., George R. Bousfield, G.R., Huhtaniemi,I.,  and Reiter, E. Trafficking of the Follitropin Receptor. Meth. Enzymol. 251:  17-45, 2013.
  60. Bousfield, G.R., Butnev, V.Y., Butnev, V.Y., Hiromasa, Y., Harvey, D.J., and May, J.V. Hypo-glycosylated human follicle-stimulating hormone (hFSH21/18) is much more active in vitro than fully-glycosylated hFSH (hFSH24), Mol. Cell. Endocrinol. 382:  989-997, 2014.

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Professional Organizations and Honors

  • Sigma Xi
  • American Association for the Advancement of Science
  • American Chemical Society
  • American Society for Biochemistry and Molecular Biology
  • The Endocrine Society
  • The Protein Society
  • Society for the Study of Reproduction
  • Society for Glycobiology
  • Association of Biomedical Resource Facilities
  • 1974, William B. Owsley Outstanding Student in Biology Award, Presented by Saginaw Valley State College
  • 1994 Wichita State University Mortar Board, Educator Appreciation Award
  • 2003 Kansas Biomedical Research Infrastructure Network Faculty Scholar Award
  • 2010 Wichita State University Excellence in Research Award
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