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Willard Morgan Freeman

InstitutionUniversity of Oklahoma Health Sciences Center
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    Wake Forest University, Winston-Salem, NCBAEnglish Literature & Chemistry
    Wake Forest University School of Medicine, Winston-Salem, NCPhDPharmacology
    Vollum Institute, Portland, ORPost-DocMolecular Neuroscience
    Yerkes National Primate Research Center, Atlanta, GAPost-DocMolecular Neuroscience

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    Collapse overview
    Dr. Willard Freeman's research program is centered on systems neurobiology related to cognitive decline with aging, diabetic retinopathy and substance abuse. Dr. Freeman received both his BA and his PhD from Wake Forest University. He completed post-doctoral training at the Vollum Institute at the University of Oregon Health Sciences Center and Yerkes National Primate Research Center at Emory University. Prior to coming to the University of Oklahoma in 2013 he was an Assistant and Associate Professor in the Department of Pharmacology at Penn State University. Dr. Freeman is in the Department of Physiology and is a member of the Reynolds Oklahoma Center on Aging, Harold Hamm Diabetes Center, and Stephensen Cancer Center.

    Dr. Freeman's research in aging are focused on identifying the molecular mechanisms within the brain that result in cognitive decline with age. In a series of proteomic studies, Dr. Freeman identified the myelin-associated inhibitor signaling cascade (Nogo, MAG, OMGP through NgR1) as a potential therapeutic target for preventing or reversing cognitive decline with age. Additionally his group is investigating neuroimmune alterations, specifically in the MHCI pathway, with aging and their potential contribution to synaptic dysfunction. As part of the Reynolds Oklahoma Center on Aging his group is bringing advanced epigenomic, transcriptomic, and proteomic technologies to a variety of collaborative projects with other Center investigators.

    Much of Dr. Freeman's group is currently examining epigenetic mechanisms in aging, diabetic retinopathy, cancer, and substance abuse. These share a common fundamental question: is the epigenome persistently altered by environmental factors (diet, stress, infection, drugs) that can cause disease and dysfunction years later. In their aging studies, the laboratory has identified that the changes to the epigenome with aging are sexually divergent. In other words, males and females respond differently. In the diabetes studies, Dr. Freeman is testing the hypothesis that persistent retinal cellular and molecular changes during a period of poor glycemic control persist long after re-establishment of normal glycemic control and contribute to the pathogenesis of diabetic retinopathy. This phenomenon, called metabolic memory, may be perpetuated by epigenetic mechanisms, both histone modifications and DNA methylation. His group develops next generation sequencing approaches to quantify epigenetic modifications for these studies.

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    Publications listed below are automatically derived from MEDLINE/PubMed and other sources, which might result in incorrect or missing publications. Faculty can login to make corrections and additions.
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    1. Valcarcel-Ares MN, Tucsek Z, Kiss T, Giles CB, Tarantini S, Yabluchanskiy A, Balasubramanian P, Gautam T, Galvan V, Ballabh P, Richardson A, Freeman WM, Wren JD, Deak F, Ungvari Z, Csiszar A. Obesity in Aging Exacerbates Neuroinflammation, Dysregulating Synaptic Function-Related Genes and Altering Eicosanoid Synthesis in the Mouse Hippocampus: Potential Role in Impaired Synaptic Plasticity and Cognitive Decline. J Gerontol A Biol Sci Med Sci. 2019 Feb 15; 74(3):290-298. PMID: 29893815.
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    2. Ahn B, Ranjit R, Premkumar P, Pharaoh G, Piekarz KM, Matsuzaki S, Claflin DR, Riddle K, Judge J, Bhaskaran S, Satara Natarajan K, Barboza E, Wronowski B, Kinter M, Humphries KM, Griffin TM, Freeman WM, Richardson A, Brooks SV, Van Remmen H. Mitochondrial oxidative stress impairs contractile function but paradoxically increases muscle mass via fibre branching. J Cachexia Sarcopenia Muscle. 2019 Feb 01. PMID: 30706998.
      View in: PubMed
    3. Unnikrishnan A, Freeman WM, Jackson J, Wren JD, Porter H, Richardson A. The role of DNA methylation in epigenetics of aging. Pharmacol Ther. 2018 Nov 09. PMID: 30419258.
      View in: PubMed
    4. Li X, Oh S, Song H, Shin S, Zhang B, Freeman WM, Janknecht R. A potential common role of the Jumonji C domain-containing 1A histone demethylase and chromatin remodeler ATRX in promoting colon cancer. Oncol Lett. 2018 Nov; 16(5):6652-6662. PMID: 30405805.
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    5. Reagan AM, Gu X, Paudel S, Ashpole NM, Zalles M, Sonntag WE, Ungvari Z, Csiszar A, Otalora L, Freeman WM, Stout MB, Elliott MH. Age-related focal loss of contractile vascular smooth muscle cells in retinal arterioles is accelerated by caveolin-1 deficiency. Neurobiol Aging. 2018 Nov; 71:1-12. PMID: 30059797.
      View in: PubMed
    6. Imperio CG, McFalls AJ, Hadad N, Blanco-Berdugo L, Masser DR, Colechio EM, Coffey AA, Bixler GV, Stanford DR, Vrana KE, Grigson PS, Freeman WM. Exposure to environmental enrichment attenuates addiction-like behavior and alters molecular effects of heroin self-administration in rats. Neuropharmacology. 2018 09 01; 139:26-40. PMID: 29964093.
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    7. Mangold CA, Yao PJ, Du M, Freeman WM, Benkovic SJ, Szpara ML. Expression of the purine biosynthetic enzyme phosphoribosyl formylglycinamidine synthase in neurons. J Neurochem. 2018 Mar; 144(6):723-735. PMID: 29337348.
      View in: PubMed
    8. Hadad N, Unnikrishnan A, Jackson JA, Masser DR, Otalora L, Stanford DR, Richardson A, Freeman WM. Caloric restriction mitigates age-associated hippocampal differential CG and non-CG methylation. Neurobiol Aging. 2018 07; 67:53-66. PMID: 29631215.
      View in: PubMed
    9. Logan S, Pharaoh GA, Marlin MC, Masser DR, Matsuzaki S, Wronowski B, Yeganeh A, Parks EE, Premkumar P, Farley JA, Owen DB, Humphries KM, Kinter M, Freeman WM, Szweda LI, Van Remmen H, Sonntag WE. Insulin-like growth factor receptor signaling regulates working memory, mitochondrial metabolism, and amyloid-ß uptake in astrocytes. Mol Metab. 2018 03; 9:141-155. PMID: 29398615.
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    10. Unnikrishnan A, Hadad N, Masser DR, Jackson J, Freeman WM, Richardson A. Revisiting the genomic hypomethylation hypothesis of aging. Ann N Y Acad Sci. 2018 Apr; 1418(1):69-79. PMID: 29363785.
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    11. Masser DR, Hadad N, Porter H, Stout MB, Unnikrishnan A, Stanford DR, Freeman WM. Analysis of DNA modifications in aging research. Geroscience. 2018 02; 40(1):11-29. PMID: 29327208.
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    12. Steyn FJ, Ngo ST, Chen VP, Bailey-Downs LC, Xie TY, Ghadami M, Brimijoin S, Freeman WM, Rubinstein M, Low MJ, Stout MB. 17a-estradiol acts through hypothalamic pro-opiomelanocortin expressing neurons to reduce feeding behavior. Aging Cell. 2018 Feb; 17(1). PMID: 29168299.
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    13. Masser DR, Otalora L, Clark NW, Kinter MT, Elliott MH, Freeman WM. Functional changes in the neural retina occur in the absence of mitochondrial dysfunction in a rodent model of diabetic retinopathy. J Neurochem. 2017 12; 143(5):595-608. PMID: 28902411.
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    14. Du M, Mangold CA, Bixler GV, Brucklacher RM, Masser DR, Stout MB, Elliott MH, Freeman WM. Retinal gene expression responses to aging are sexually divergent. Mol Vis. 2017; 23:707-717. PMID: 29062222.
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    15. Masser DR, Hadad N, Porter HL, Mangold CA, Unnikrishnan A, Ford MM, Giles CB, Georgescu C, Dozmorov MG, Wren JD, Richardson A, Stanford DR, Freeman WM. Sexually divergent DNA methylation patterns with hippocampal aging. Aging Cell. 2017 12; 16(6):1342-1352. PMID: 28948711.
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    16. Mangold CA, Wronowski B, Du M, Masser DR, Hadad N, Bixler GV, Brucklacher RM, Ford MM, Sonntag WE, Freeman WM. Sexually divergent induction of microglial-associated neuroinflammation with hippocampal aging. J Neuroinflammation. 2017 07 21; 14(1):141. PMID: 28732515.
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    17. Unnikrishnan A, Jackson J, Matyi SA, Hadad N, Wronowski B, Georgescu C, Garrett KP, Wren JD, Freeman WM, Richardson A. Role of DNA methylation in the dietary restriction mediated cellular memory. Geroscience. 2017 06; 39(3):331-345. PMID: 28477138.
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    18. Hopiavuori BR, Masser DR, Wilkerson JL, Brush RS, Mandal NA, Anderson RE, Freeman WM. Isolation of Neuronal Synaptic Membranes by Sucrose Gradient Centrifugation. Methods Mol Biol. 2017; 1609:33-41. PMID: 28660571.
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    19. Masser DR, Clark NW, Van Remmen H, Freeman WM. Loss of the antioxidant enzyme CuZnSOD (Sod1) mimics an age-related increase in absolute mitochondrial DNA copy number in the skeletal muscle. Age (Dordr). 2016 Aug; 38(4):323-333. PMID: 27444179.
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    20. Hadad N, Masser DR, Logan S, Wronowski B, Mangold CA, Clark N, Otalora L, Unnikrishnan A, Ford MM, Giles CB, Wren JD, Richardson A, Sonntag WE, Stanford DR, Freeman W. Absence of genomic hypomethylation or regulation of cytosine-modifying enzymes with aging in male and female mice. Epigenetics Chromatin. 2016; 9:30. PMID: 27413395; PMCID: PMC4942942.
    21. Masser DR, Stanford DR, Hadad N, Giles CB, Wren JD, Sonntag WE, Richardson A, Freeman WM. Bisulfite oligonucleotide-capture sequencing for targeted base- and strand-specific absolute 5-methylcytosine quantitation. Age (Dordr). 2016 Jun; 38(3):49. PMID: 27091453; PMCID: PMC5005917 [Available on 06/01/17].
    22. Mangold CA, Masser DR, Stanford DR, Bixler GV, Pisupati A, Giles CB, Wren JD, Ford MM, Sonntag WE, Freeman WM. CNS-wide Sexually Dimorphic Induction of the Major Histocompatibility Complex 1 Pathway With Aging. J Gerontol A Biol Sci Med Sci. 2017 01; 72(1):16-29. PMID: 26786204.
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    23. Imperio CG, McFalls AJ, Colechio EM, Masser DR, Vrana KE, Grigson PS, Freeman WM. Assessment of individual differences in the rat nucleus accumbens transcriptome following taste-heroin extended access. Brain Res Bull. 2016 05; 123:71-80. PMID: 26733446.
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    24. McFalls AJ, Imperio CG, Bixler G, Freeman WM, Grigson PS, Vrana KE. Reward devaluation and heroin escalation is associated with differential expression of CRF signaling genes. Brain Res Bull. 2016 05; 123:81-93. PMID: 26655889.
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    25. Deak F, Freeman WM, Ungvari Z, Csiszar A, Sonntag WE. Recent Developments in Understanding Brain Aging: Implications for Alzheimer's Disease and Vascular Cognitive Impairment. J Gerontol A Biol Sci Med Sci. 2016 Jan; 71(1):13-20. PMID: 26590911; PMCID: PMC4851715 [Available on 01/01/17].
    26. Dozmorov MG, Adrianto I, Giles CB, Glass E, Glenn SB, Montgomery C, Sivils KL, Olson LE, Iwayama T, Freeman WM, Lessard CJ, Wren JD. Detrimental effects of duplicate reads and low complexity regions on RNA- and ChIP-seq data. BMC Bioinformatics. 2015; 16 Suppl 13:S10. PMID: 26423047.
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    27. Masser DR, Stanford DR, Freeman WM. Targeted DNA methylation analysis by next-generation sequencing. J Vis Exp. 2015 Feb 24; (96). PMID: 25741966; PMCID: PMC4354667.
    28. Masser DR, Bixler GV, Brucklacher RM, Yan H, Giles CB, Wren JD, Sonntag WE, Freeman WM. Hippocampal subregions exhibit both distinct and shared transcriptomic responses to aging and nonneurodegenerative cognitive decline. J Gerontol A Biol Sci Med Sci. 2014 Nov; 69(11):1311-24. PMID: 24994846; PMCID: PMC4271093.
    29. Masser DR, VanGuilder Starkey HD, Bixler GV, Dunton W, Bronson SK, Freeman WM. Insulin treatment normalizes retinal neuroinflammation but not markers of synapse loss in diabetic rats. Exp Eye Res. 2014 Aug; 125:95-106. PMID: 24931083; PMCID: PMC4122586.
    30. Govey PM, Jacobs JM, Tilton SC, Loiselle AE, Zhang Y, Freeman WM, Waters KM, Karin NJ, Donahue HJ. Integrative transcriptomic and proteomic analysis of osteocytic cells exposed to fluid flow reveals novel mechano-sensitive signaling pathways. J Biomech. 2014 Jun 03; 47(8):1838-45. PMID: 24720889; PMCID: PMC4037855.
    31. Bezinover D, Ramamoorthy S, Postula M, Weller G, Mahmoud S, Mani H, Kadry Z, Uemura T, Mets B, Spiess B, Brucklacher R, Freeman W, Janicki PK. Effect of cold perfusion and perfluorocarbons on liver graft ischemia in a donation after cardiac death model. J Surg Res. 2014 May 15; 188(2):517-26. PMID: 24582069.
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    32. Masser DR, Berg AS, Freeman WM. Focused, high accuracy 5-methylcytosine quantitation with base resolution by benchtop next-generation sequencing. Epigenetics Chromatin. 2013 Oct 11; 6(1):33. PMID: 24279302; PMCID: PMC3907040.
    33. Sonntag WE, Deak F, Ashpole N, Toth P, Csiszar A, Freeman W, Ungvari Z. Insulin-like growth factor-1 in CNS and cerebrovascular aging. Front Aging Neurosci. 2013; 5:27. PMID: 23847531; PMCID: PMC3698444.
    34. VanGuilder Starkey HD, Bixler GV, Sonntag WE, Freeman WM. Expression of NgR1-antagonizing proteins decreases with aging and cognitive decline in rat hippocampus. Cell Mol Neurobiol. 2013 May; 33(4):483-8. PMID: 23525710; PMCID: PMC3651369.
    35. VanGuilder Starkey HD, Sonntag WE, Freeman WM. Increased hippocampal NgR1 signaling machinery in aged rats with deficits of spatial cognition. Eur J Neurosci. 2013 May; 37(10):1643-58. PMID: 23438185; PMCID: PMC3660548.
    36. Torrente MP, Freeman WM, Vrana KE. Protein biomarkers of alcohol abuse. Expert Rev Proteomics. 2012 Aug; 9(4):425-36. PMID: 22967079; PMCID: PMC3535006.
    37. Huber-Keener KJ, Liu X, Wang Z, Wang Y, Freeman W, Wu S, Planas-Silva MD, Ren X, Cheng Y, Zhang Y, Vrana K, Liu CG, Yang JM, Wu R. Differential gene expression in tamoxifen-resistant breast cancer cells revealed by a new analytical model of RNA-Seq data. PLoS One. 2012; 7(7):e41333. PMID: 22844461; PMCID: PMC3402532.
    38. Jones NR, Sun D, Freeman WM, Lazarus P. Quantification of Hepatic UDP glucuronosyltransferase 1A splice variant expression and correlation of UDP glucuronosyltransferase 1A1 variant expression with glucuronidation activity. J Pharmacol Exp Ther. 2012 Sep; 342(3):720-9. PMID: 22661630; PMCID: PMC3422519.
    39. VanGuilder Starkey HD, Van Kirk CA, Bixler GV, Imperio CG, Kale VP, Serfass JM, Farley JA, Yan H, Warrington JP, Han S, Mitschelen M, Sonntag WE, Freeman WM. Neuroglial expression of the MHCI pathway and PirB receptor is upregulated in the hippocampus with advanced aging. J Mol Neurosci. 2012 Sep; 48(1):111-26. PMID: 22562814; PMCID: PMC3414657.
    40. Kaiser JM, Imai H, Haakenson JK, Brucklacher RM, Fox TE, Shanmugavelandy SS, Unrath KA, Pedersen MM, Dai P, Freeman WM, Bronson SK, Gardner TW, Kester M. Nanoliposomal minocycline for ocular drug delivery. Nanomedicine. 2013 Jan; 9(1):130-40. PMID: 22465498; PMCID: PMC3762261.
    41. Vanguilder HD, Bixler GV, Sonntag WE, Freeman WM. Hippocampal expression of myelin-associated inhibitors is induced with age-related cognitive decline and correlates with deficits of spatial learning and memory. J Neurochem. 2012 Apr; 121(1):77-98. PMID: 22269040; PMCID: PMC3341628.
    42. Umstead TM, Lu CJ, Freeman WM, Myers JL, Clark JB, Thomas NJ, Icitovic N, Chinchilli VM, Undar A, Phelps DS. The kinetics of cardiopulmonary bypass: a dual-platform proteomics study of plasma biomarkers in pediatric patients undergoing cardiopulmonary bypass. Artif Organs. 2012 Jan; 36(1):E1-20. PMID: 22250822.
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    43. Helms CM, Messaoudi I, Jeng S, Freeman WM, Vrana KE, Grant KA. A longitudinal analysis of circulating stress-related proteins and chronic ethanol self-administration in cynomolgus macaques. Alcohol Clin Exp Res. 2012 Jun; 36(6):995-1003. PMID: 22141444; PMCID: PMC3324628.
    44. VanGuilder HD, Bixler GV, Brucklacher RM, Farley JA, Yan H, Warrington JP, Sonntag WE, Freeman WM. Concurrent hippocampal induction of MHC II pathway components and glial activation with advanced aging is not correlated with cognitive impairment. J Neuroinflammation. 2011 Oct 11; 8:138. PMID: 21989322; PMCID: PMC3216278.
    45. Yan H, Mitschelen M, Bixler GV, Brucklacher RM, Farley JA, Han S, Freeman WM, Sonntag WE. Circulating IGF1 regulates hippocampal IGF1 levels and brain gene expression during adolescence. J Endocrinol. 2011 Oct; 211(1):27-37. PMID: 21750148; PMCID: PMC3395434.
    46. Vanguilder HD, Freeman WM. The hippocampal neuroproteome with aging and cognitive decline: past progress and future directions. Front Aging Neurosci. 2011; 3:8. PMID: 21647399; PMCID: PMC3102218.
    47. Bixler GV, Vanguilder HD, Brucklacher RM, Kimball SR, Bronson SK, Freeman WM. Chronic insulin treatment of diabetes does not fully normalize alterations in the retinal transcriptome. BMC Med Genomics. 2011 May 15; 4:40. PMID: 21575160; PMCID: PMC3113923.
    48. Van Kirk CA, VanGuilder HD, Young M, Farley JA, Sonntag WE, Freeman WM. Age-related alterations in retinal neurovascular and inflammatory transcripts. Mol Vis. 2011; 17:1261-74. PMID: 21633715; PMCID: PMC3103744.
    49. VanGuilder HD, Farley JA, Yan H, Van Kirk CA, Mitschelen M, Sonntag WE, Freeman WM. Hippocampal dysregulation of synaptic plasticity-associated proteins with age-related cognitive decline. Neurobiol Dis. 2011 Jul; 43(1):201-12. PMID: 21440628; PMCID: PMC3096728.
    50. Freeman WM, Vanguilder HD, Guidone E, Krystal JH, Grant KA, Vrana KE. Plasma proteomic alterations in non-human primates and humans after chronic alcohol self-administration. Int J Neuropsychopharmacol. 2011 Aug; 14(7):899-911. PMID: 21303580; PMCID: PMC3107900.
    51. Carkaci-Salli N, Salli U, Kuntz-Melcavage KL, Pennock MM, Ozgen H, Tekin I, Freeman WM, Vrana KE. TPH2 in the ventral tegmental area of the male rat brain. Brain Res Bull. 2011 Apr 05; 84(6):376-80. PMID: 21272616; PMCID: PMC3070366.
    52. VanGuilder HD, Bixler GV, Kutzler L, Brucklacher RM, Bronson SK, Kimball SR, Freeman WM. Multi-modal proteomic analysis of retinal protein expression alterations in a rat model of diabetic retinopathy. PLoS One. 2011 Jan 13; 6(1):e16271. PMID: 21249158; PMCID: PMC3020973.
    53. Lebold KM, Grant KA, Freeman WM, Wiren KM, Miller GW, Kiley C, Leonard SW, Traber MG. Individual differences in hyperlipidemia and vitamin E status in response to chronic alcohol self-administration in cynomolgus monkeys. Alcohol Clin Exp Res. 2011 Mar; 35(3):474-83. PMID: 21118275; PMCID: PMC3116096.
    54. Rountree CB, Van Kirk CA, You H, Ding W, Dang H, VanGuilder HD, Freeman WM. Clinical application for the preservation of phospho-proteins through in-situ tissue stabilization. Proteome Sci. 2010 Nov 22; 8:61. PMID: 21092202; PMCID: PMC3000382.
    55. Ali M, Umstead TM, Haque R, Mikerov AN, Freeman WM, Floros J, Phelps DS. Differences in the BAL proteome after Klebsiella pneumoniae infection in wild type and SP-A-/- mice. Proteome Sci. 2010 Jun 17; 8:34. PMID: 20565803; PMCID: PMC2911411.
    56. Abcouwer SF, Lin CM, Wolpert EB, Shanmugam S, Schaefer EW, Freeman WM, Barber AJ, Antonetti DA. Effects of ischemic preconditioning and bevacizumab on apoptosis and vascular permeability following retinal ischemia-reperfusion injury. Invest Ophthalmol Vis Sci. 2010 Nov; 51(11):5920-33. PMID: 20554620.
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    57. Umstead TM, Lu CJ, Freeman WM, Myers JL, Clark JB, Thomas NJ, Chinchilli VM, Vrana KE, Undar A, Phelps DS. Dual-platform proteomics study of plasma biomarkers in pediatric patients undergoing cardiopulmonary bypass. Pediatr Res. 2010 Jun; 67(6):641-9. PMID: 20308938.
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    58. Freeman WM, Vrana KE. Future prospects for biomarkers of alcohol consumption and alcohol-induced disorders. Alcohol Clin Exp Res. 2010 Jun; 34(6):946-54. PMID: 20374220; PMCID: PMC2900387.
    59. VanGuilder HD, Yan H, Farley JA, Sonntag WE, Freeman WM. Aging alters the expression of neurotransmission-regulating proteins in the hippocampal synaptoproteome. J Neurochem. 2010 Jun; 113(6):1577-88. PMID: 20374424; PMCID: PMC3010414.
    60. Freeman WM, Salzberg AC, Gonzales SW, Grant KA, Vrana KE. Classification of alcohol abuse by plasma protein biomarkers. Biol Psychiatry. 2010 Aug 01; 68(3):219-22. PMID: 20299005; PMCID: PMC2890298.
    61. Freeman WM, Lull ME, Patel KM, Brucklacher RM, Morgan D, Roberts DC, Vrana KE. Gene expression changes in the medial prefrontal cortex and nucleus accumbens following abstinence from cocaine self-administration. BMC Neurosci. 2010 Feb 26; 11:29. PMID: 20187946; PMCID: PMC2837051.
    62. Blake DC, Mikse OR, Freeman WM, Herzog CR. FOXO3a elicits a pro-apoptotic transcription program and cellular response to human lung carcinogen nicotine-derived nitrosaminoketone (NNK). Lung Cancer. 2010 Jan; 67(1):37-47. PMID: 19380174.
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    63. Achur RN, Freeman WM, Vrana KE. Circulating cytokines as biomarkers of alcohol abuse and alcoholism. J Neuroimmune Pharmacol. 2010 Mar; 5(1):83-91. PMID: 20020329; PMCID: PMC2862254.
    64. Freeman WM, Bixler GV, Brucklacher RM, Walsh E, Kimball SR, Jefferson LS, Bronson SK. Transcriptomic comparison of the retina in two mouse models of diabetes. J Ocul Biol Dis Infor. 2009 Dec 12; 2(4):202-213. PMID: 20157355.
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    65. Lull ME, Freeman WM, VanGuilder HD, Vrana KE. The use of neuroproteomics in drug abuse research. Drug Alcohol Depend. 2010 Feb 01; 107(1):11-22. PMID: 19926406; PMCID: PMC3947580.
    66. Undar A, Pauliks L, Clark JB, Zahn J, Rosenberg G, Kunselman AR, Sun Q, Pekkan K, Saliba K, Carney E, Thomas N, Freeman W, Vrana K, El-Banayosy A, Ural SH, Wilson R, Umstead TM, Floros J, Phelps DS, Weiss W, Snyder A, Yang S, Kimatian S, Cyran SE, Chinchilli VM, Guan Y, Rider A, Haines N, Rogerson A, Alkan-Bozkaya T, Akcevin A, Sun K, Wang S, Cun L, Myers JL. Penn State Hershey--center for pediatric cardiovascular research. Artif Organs. 2009 Nov; 33(11):883-7. PMID: 20021467.
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    67. Kuntz-Melcavage KL, Brucklacher RM, Grigson PS, Freeman WM, Vrana KE. Gene expression changes following extinction testing in a heroin behavioral incubation model. BMC Neurosci. 2009 Aug 07; 10:95. PMID: 19664213; PMCID: PMC2733140.
    68. Jaishankar A, Barthelery M, Freeman WM, Salli U, Ritty TM, Vrana KE. Human embryonic and mesenchymal stem cells express different nuclear proteomes. Stem Cells Dev. 2009 Jun; 18(5):793-802. PMID: 18821827.
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    69. Barthéléry M, Jaishankar A, Salli U, Freeman WM, Vrana KE. 2-D DIGE identification of differentially expressed heterogeneous nuclear ribonucleoproteins and transcription factors during neural differentiation of human embryonic stem cells. Proteomics Clin Appl. 2009 Apr; 3(4):505-14. PMID: 21136975.
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    70. Haque R, Umstead TM, Freeman WM, Floros J, Phelps DS. The impact of surfactant protein-A on ozone-induced changes in the mouse bronchoalveolar lavage proteome. Proteome Sci. 2009 Mar 26; 7:12. PMID: 19323824; PMCID: PMC2666657.
    71. Lull ME, Erwin MS, Morgan D, Roberts DC, Vrana KE, Freeman WM. Persistent proteomic alterations in the medial prefrontal cortex with abstinence from cocaine self-administration. Proteomics Clin Appl. 2009 Mar 17; 3(4):462-472. PMID: 20161123.
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    72. Miller A, Lu CK, Wang S, Umstead TM, Freeman WM, Vrana K, Yang S, Myers JL, Phelps DS, Zahn JD, Undar A. Pediatric cardiopulmonary bypass circuits: a review of studies conducted at the Penn State Pediatric Cardiac Research Laboratories. J Extra Corpor Technol. 2009 Mar; 41(1):P50-8. PMID: 19361042; PMCID: PMC4680234.
    73. Fort PE, Freeman WM, Losiewicz MK, Singh RS, Gardner TW. The retinal proteome in experimental diabetic retinopathy: up-regulation of crystallins and reversal by systemic and periocular insulin. Mol Cell Proteomics. 2009 Apr; 8(4):767-79. PMID: 19049959; PMCID: PMC2667355.
    74. Prokopczyk B, Sinha I, Trushin N, Freeman WM, El-Bayoumy K. Gene expression profiles in HPV-immortalized human cervical cells treated with the nicotine-derived carcinogen 4-(methylnitrosamino)-1-(3-pyridyl)-1-butanone. Chem Biol Interact. 2009 Feb 12; 177(3):173-80. PMID: 19038236; PMCID: PMC2723790.
    75. Umstead TM, Freeman WM, Chinchilli VM, Phelps DS. Age-related changes in the expression and oxidation of bronchoalveolar lavage proteins in the rat. Am J Physiol Lung Cell Mol Physiol. 2009 Jan; 296(1):L14-29. PMID: 18931054.
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    76. Lull ME, Freeman WM, Vrana KE, Mash DC. Correlating human and animal studies of cocaine abuse and gene expression. Ann N Y Acad Sci. 2008 Oct; 1141:58-75. PMID: 18991951.
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    77. Kuntz-Melcavage KL, Freeman WM, Vrana KE. CNS genes implicated in relapse. Subst Abuse. 2008; 2:1-12. PMID: 25922574; PMCID: PMC4395042.
    78. VanGuilder HD, Brucklacher RM, Patel K, Ellis RW, Freeman WM, Barber AJ. Diabetes downregulates presynaptic proteins and reduces basal synapsin I phosphorylation in rat retina. Eur J Neurosci. 2008 Jul; 28(1):1-11. PMID: 18662330.
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    79. Brucklacher RM, Patel KM, VanGuilder HD, Bixler GV, Barber AJ, Antonetti DA, Lin CM, LaNoue KF, Gardner TW, Bronson SK, Freeman WM. Whole genome assessment of the retinal response to diabetes reveals a progressive neurovascular inflammatory response. BMC Med Genomics. 2008 Jun 13; 1:26. PMID: 18554398; PMCID: PMC2442612.
    80. Lull ME, Carkaci-Salli N, Freeman WM, Myers JL, Midgley FM, Thomas NJ, Kimatian SJ, Vrana KE, Undar A. Plasma biomarkers in pediatric patients undergoing cardiopulmonary bypass. Pediatr Res. 2008 Jun; 63(6):638-44. PMID: 18317239.
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    81. Kuntz KL, Patel KM, Grigson PS, Freeman WM, Vrana KE. Heroin self-administration: II. CNS gene expression following withdrawal and cue-induced drug-seeking behavior. Pharmacol Biochem Behav. 2008 Sep; 90(3):349-56. PMID: 18466961; PMCID: PMC4304653.
    82. VanGuilder HD, Vrana KE, Freeman WM. Twenty-five years of quantitative PCR for gene expression analysis. Biotechniques. 2008 Apr; 44(5):619-26. PMID: 18474036.
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    83. Freeman WM, Patel KM, Brucklacher RM, Lull ME, Erwin M, Morgan D, Roberts DC, Vrana KE. Persistent alterations in mesolimbic gene expression with abstinence from cocaine self-administration. Neuropsychopharmacology. 2008 Jul; 33(8):1807-17. PMID: 17851536; PMCID: PMC2810407.
    84. Antonetti DA, Barber AJ, Bronson SK, Freeman WM, Gardner TW, Jefferson LS, Kester M, Kimball SR, Krady JK, LaNoue KF, Norbury CC, Quinn PG, Sandirasegarane L, Simpson IA. Diabetic retinopathy: seeing beyond glucose-induced microvascular disease. Diabetes. 2006 Sep; 55(9):2401-11. PMID: 16936187.
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    85. Lull ME, Freeman WM, Myers JL, Midgley F, Kimatian SJ, Undar A, Vrana KE. Plasma proteomics: a noninvasive window on pathology and pediatric cardiac surgery. ASAIO J. 2006 Sep-Oct; 52(5):562-6. PMID: 16966860.
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    86. Freeman WM, Lull ME, Guilford MT, Vrana KE. Depletion of abundant proteins from non-human primate serum for biomarker studies. Proteomics. 2006 May; 6(10):3109-13. PMID: 16619306.
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    87. Freeman WM, Gooch RS, Lull ME, Worst TJ, Walker SJ, Xu AS, Green H, Pierre PJ, Grant KA, Vrana KE. Apo-AII is an elevated biomarker of chronic non-human primate ethanol self-administration. Alcohol Alcohol. 2006 May-Jun; 41(3):300-5. PMID: 16581821.
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    88. Worst TJ, Tan JC, Robertson DJ, Freeman WM, Hyytia P, Kiianmaa K, Vrana KE. Transcriptome analysis of frontal cortex in alcohol-preferring and nonpreferring rats. J Neurosci Res. 2005 May 15; 80(4):529-38. PMID: 15846778.
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    89. Freeman WM, Hemby SE. Proteomics for protein expression profiling in neuroscience. Neurochem Res. 2004 Jun; 29(6):1065-81. PMID: 15176464.
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    90. Stredrick DL, Stokes AH, Worst TJ, Freeman WM, Johnson EA, Lash LH, Aschner M, Vrana KE. Manganese-induced cytotoxicity in dopamine-producing cells. Neurotoxicology. 2004 Jun; 25(4):543-53. PMID: 15183009.
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    91. Tang W, Wesley M, Freeman WM, Liang B, Hemby SE. Alterations in ionotropic glutamate receptor subunits during binge cocaine self-administration and withdrawal in rats. J Neurochem. 2004 May; 89(4):1021-33. PMID: 15140200; PMCID: PMC3843358.
    92. Walker SJ, Worst TJ, Freeman WM, Vrana KE. Functional genomic analysis in pain research using hybridization arrays. Methods Mol Med. 2004; 99:239-53. PMID: 15131342.
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    93. Vrana KE, Freeman WM, Aschner M. Use of microarray technologies in toxicology research. Neurotoxicology. 2003 Jun; 24(3):321-32. PMID: 12782098.
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    94. Worst TJ, Freeman WM, Walker SJ, Vrana KE. Systematic screening of gene expression using a cDNA macroarray. Methods Mol Med. 2003; 79:243-59. PMID: 12506701.
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    95. Stokes AH, Freeman WM, Mitchell SG, Burnette TA, Hellmann GM, Vrana KE. Induction of GADD45 and GADD153 in neuroblastoma cells by dopamine-induced toxicity. Neurotoxicology. 2002 Dec; 23(6):675-84. PMID: 12520757.
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    96. Freeman WM, Brebner K, Patel KM, Lynch WJ, Roberts DC, Vrana KE. Repeated cocaine self-administration causes multiple changes in rat frontal cortex gene expression. Neurochem Res. 2002 Oct; 27(10):1181-92. PMID: 12462416.
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    97. Freeman WM, Brebner K, Lynch WJ, Patel KM, Robertson DJ, Roberts DC, Vrana KE. Changes in rat frontal cortex gene expression following chronic cocaine. Brain Res Mol Brain Res. 2002 Jul 15; 104(1):11-20. PMID: 12117546.
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    98. Freeman WM, Dougherty KE, Vacca SE, Vrana KE. An interactive database of cocaine-responsive gene expression. ScientificWorldJournal. 2002 Mar 12; 2:701-6. PMID: 12805995.
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