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Your Environment. Your Health.

Person Details: Dr. Pamela L. Mellon

Superfund Research Program

Dr. Pamela L. Mellon

University of California-San Diego
Leichtag Biomedical Research Building, Room 3A14
Mail Code 0674
9500 Gilman Drive
La Jolla, California 92093-0674
Phone: 858-534-1312
Fax: 858-534-1438




  • Cassin J, Stamou MI, Keefe KW, Sung K, Bojo C, Tonsfeldt KJ, Rojas RA, Lopes VF, Plummer L, Salnikov KB, Keefe DL, Ozata M, Genel M, Georgopoulos NA, Hall JE, Crowley WF, Seminara SB, Mellon PL, Balasubramanian R. 2023. Heterozygous mutations in SOX2 may cause idiopathic hypogonadotropic hypogonadism via dominant-negative mechanisms. JCI Insight e164324. doi:10.1172/jci.insight.164324 PMID:36602867


  • Lavalle SN, Chou T, Hernandez J, Naing NP, He MY, Tonsfeldt KJ, Mellon PL. 2022. Deletion of the homeodomain gene Six3 from kisspeptin neurons causes subfertility in female mice. Mol Cell Endocrinol 546:111577. doi:10.1016/j.mce.2022.111577 PMID:35121076 PMCID:PMC8934285
  • Meadows JD, Breuer JA, Lavalle SN, Hirschenberger MR, Patel MM, Nguyen D, Kim A, Cassin J, Gorman MR, Welsh DK, Mellon PL, Hoffmann HM. 2022. Deletion of Six3 in post-proliferative neurons produces weakened SCN circadian output, improved metabolic function, and dwarfism in male mice. Mol Metab 57:doi:10.1016/j.molmet.2021.101431 PMID:34974160 PMCID:PMC8810556
  • Tonsfeldt KJ, Cui L, Lee J, Walbeek TJ, Brusman LE, Jin Y, Mieda M, Gorman MR, Mellon PL. 2022. Female fertility does not require Bmal1 in suprachiasmatic nucleus neurons expressing arginine vasopressin, vasoactive intestinal peptide, or neuromedin-S. Front. Endocrinol 13:doi:10.3389/fendo.2022.956169 PMID:35992114 PMCID:PMC9389073
  • Tonsfeldt KJ, Mellon PL, Hoffmann HM. 2022. Circadian rhythms in the neuronal network timing the luteinizing hormone surge. Endocrinology 163(2):doi:10.1210/endocr/bqab268 PMID:34967900 PMCID:PMC8782605


  • Bohaczuk SC, Cassin J, Slaiwa TI, Thackray VG, Mellon PL. 2021. Distal Enhancer Potentiates Activin- and GnRH-Induced Transcription of FSHB. Endocrinology 162:doi:10.1210/endocr/bqab069 PMID:33824966 PMCID:PMC8157479
  • Bohaczuk SC, Thackray VG, Shen J, Skowronska-Krawczyk D, Mellon PL. 2021. FSHB transcription is regulated by a novel 5' distal enhancer with a fertility-associated single nucleotide polymorphism. Endocrinology 162:doi:10.1210/endocr/bqaa181 PMID:33009549 PMCID:PMC7846141
  • Ho EV, Shi C, Cassin J, He MY, Nguyen RD, Ryan GE, Tonsfeldt KJ, Mellon PL. 2021. Reproductive deficits induced by prenatal Antimullerian hormone exposure require androgen receptor in kisspeptin cells. Endocrinology 162:doi:10.1210/endocr/bqab197 PMID:34529765 PMCID:PMC8507963
  • Hoffmann HM, Meadows JD, Breuer JA, Yaw AM, Nguyen D, Tonsfeldt KJ, Chin AY, Devries BM, Trang C, Oosterhouse HJ, Lee JS, Doser JW, Gorman MR, Welsh DK, Mellon PL. 2021. The transcription factors SIX3 and VAX1 are required for suprachiasmatic nucleus circadian output and fertility in female mice. J Neurosci Res PMID:34212416
  • Lavalle SN, Chou T, Hernandez J, Naing NP, Tonsfeldt KJ, Hoffmann HM, Mellon PL. 2021. Kiss1 is differentially regulated in male and female mice by the homeodomain transcription factor VAX1. Mol Cell Endocrinol 534:111358. doi:10.1016/j.mce.2021.111358 PMID:34098016 (ahead of print)
  • Ryan GE, Bohaczuk SC, Cassin J, Witham EA, Shojaei S, Ho EV, Thackray VG, Mellon PL. 2021. Androgen receptor positively regulates gonadotropin-releasing hormone receptor in pituitary gonadotropes. Mol Cell Endocrinol 530:doi:10.1016/j.mce.2021.111286 PMID:33872733 PMCID:PMC8177864
  • Schoeller EL, Tonsfeldt KJ, Sinkovich M, Shi R, Mellon PL. 2021. Growth hormone pulses and liver gene expression are differentially regulated by the circadian clock gene Bmal1. Endocrinology 162(4):doi:10.1210/endocr/bqab023 PMID:33539533 PMCID:PMC7901660


  • Core N, Erni A, Hoffmann HM, Mellon PL, Saurin AJ, Beclin C, Cremer H. 2020. Stem cell regionalization during olfactory bulb neurogenesis depends on regulatory interactions between Vax1 and Pax6. Elife 9:doi:10.7554/eLife.58215 PMID:32762844 PMCID:PMC7440913
  • Gou L, Lim D, Ma W, Aubol BE, Hao Y, Wang X, Zhao J, Liang Z, Shao C, Zhang X, Meng F, Li H, Zhang X, Xu R, Li D, Rosenfeld MG, Mellon PL, Adams JA, Liu M, Fu X. 2020. Initiation of parental genome reprogramming in fertilized oocyte by splicing kinase SRPK1 catalyzed protamine phosphorylation. Cell 180:doi:10.1016/j.cell.2020.02.020 PMID:32169215
  • Nicholas DA, Knight VS, Tonsfeldt KJ, Terasaka T, Molinar-Inglis O, Stephens SB, Trejo J, Kauffman AS, Mellon PL, Lawson MA. 2020. GLUT1-mediated glycolysis supports GnRH-induced secretion of luteinizing hormone from female gonadotropes. Sci Rep 10:13063. doi:10.1038/s41598-020-69913-z PMID:32747664 PMCID:PMC7400764


  • Hoffman HM, Larder R, Lee JS, Hu RJ, Trang C, Devries BM, Clark DD, Mellon PL. 2019. Differential CRE expression in Lhrh-cre and GnRH-cre alleles and the impact on fertility in Otx2-Flox mice. Neuroendocrinology 108:328-342. doi:10.1159/000497791 PMID:30739114 PMCID:PMC6753941
  • Hoffmann HM, Pandolfi EC, Larder R, Mellon PL. 2019. Haploinsufficiency of homeodomain proteins Six3, Vax1, and Otx2 causes subfertility in mice via distinct mechanisms. Neuroendocrinology 109:200-207. doi:10.1159/000494086 PMID:30261489 PMCID:PMC6437011
  • Pandolfi EC, Breuer JA, Huu VN, Talluri T, Nguyen D, Lee JS, Hu R, Bharti K, Skowronska-Krawczyk D, Gorman MR, Mellon PL, Hoffmann HM. 2019. The homeodomain transcription factors vax1 and six6 are required for scn development and function. Mol Neurobiol 1-16. doi:10.1007/s12035-019-01781-9 PMID:31705443 PMCID:PMC7035166
  • Pandolfi EC, Tonsfeldt KJ, Hoffmann HM, Mellon PL. 2019. Deletion of the homeodomain protein Six6 from GnRH neurons decreases GnRH gene expression, resulting in infertility. Endocrinology 160:2151-2164. doi:10.1210/en.2019-00113 PMID:31211355 PMCID:PMC6821215
  • Ruf-Zamojski F, Ge Y, Pincas H, Shan J, Song Y, Hines N, Kelley K, Montagna C, Nair P, Toufaily C, Bernard DJ, Mellon PL, Nair V, Turgeon JL, Sealfon SC. 2019. Cytogenetic, Genomic, and Functional Characterization of Pituitary Gonadotrope Cell Lines. J Endocr Soc 3. doi:10.1210/js.2019-00064 PMID:31020055 PMCID:PMC6469952 (accepted for publication)
  • Tonsfeldt KJ, Schoeller EL, Brusman LE, Cui L, Lee J, Mellon PL. 2019. The Contribution of the Circadian Gene Bmal1 to Female Fertility and the Generation of the Preovulatory Luteinizing Hormone Surge. J Endocr Soc 3(4):716-733. doi:10.1210/js.2018-00228 PMID:30906911 PMCID:PMC6425515


  • Hoffman HM, Gong P, Tamrazian A, Mellon PL. 2018. Transcriptional interaction between cFOS and the homeodomain-binding transcription factor VAX1 on the GnRH promoter controls Gnrh1 expression levels in a GnRH neuron maturation specific manner. Mol Cell Endocrinol 461:143-154. doi:10.1016/j.mce.2017.09.004 PMID:28890143 PMCID:PMC5756504
  • Pandolfi EC, Hoffmann HM, Schoeller EL, Gorman MR, Mellon PL. 2018. Haploinsufficiency of SIX3 Abolishes Male Reproductive Behavior Through Disrupted Olfactory Development, and Impairs Female Fertility Through Disrupted GnRH Neuron Migration. Mol Neurobiol 55(11):8709-8727. doi:10.1007/s12035-018-1013-0 PMID:29589282 PMCID:PMC6156938
  • Que X, Hung M, Yeang C, Gonen A, Prohaska TA, Sun X, Diehl C, Maatta A, Gaddis DE, Bowden K, Pattison J, MacDonald JG, Yla-Herttuala S, Mellon PL, Hedrick CC, Ley K, Miller YI, Glass CK, Peterson KL, Binder CJ, Tsimikas S, Witztum JL. 2018. Oxidized phospholipids are proinflammatory and proatherogenic in hypercholesterolaemic mice. Nature 558:301. doi:10.1038/s41586-018-0198-8 PMID:29875409 PMCID:PMC6033669
  • Ryan GE, Malik S, Mellon PL. 2018. Antiandrogen Treatment Ameliorates Reproductive and Metabolic Phenotypes in the Letrozole-Induced Mouse Model of PCOS. Endocrinology 159(4):1734-1747. doi:10.1210/en.2017-03218 PMID:29471436 PMCID:PMC6097580




  • Kauffman AS, Thackray VG, Ryan GE, Tolson KP, Glidewell-Kenney CA, Semaan SJ, Poling MC, Iwata N, Breen KM, Duleba AJ, Stener-Victorin E, Shimasaki S, Webster NJ, Mellon PL. 2015. A novel letrozole model recapitulates both the reproductive and metabolic phenotypes of polycystic ovary syndrome in female mice. Biol Reprod 93(3):69. doi:10.1095/biolreprod.115.131631 PMID:26203175 PMCID:PMC4710190
  • Skowronska-Krawczyk D, Zhao L, Zhu J, Weinreb RN, Cao G, Luo J, Flagg K, Patel S, Wen C, Krupa M, Luo H, Ouyang H, Lin D, Wang W, Li G, Xu Y, Li O, Chung C, Yeh E, Jafari M, Ai M, Zhong Z, Shi W, Zin C, Mellon PL, Gao W, Abagyan R, Zhang L, Sun X, Zhong Z, Zhuo Y, Rosenfeld MG, Liu Y, Zhang K. 2015. P16INK4a upregulation mediated by SIX6 defines retinal ganglion cell pathogenesis in glaucoma. Mol Cell 59(6):931-940. doi:10.1016/j.molcel.2015.07.027 PMID:26365380 PMCID:PMC4648709
  • Stephens SB, Tolson KP, Rouse ML, Poling MC, Hashimoto-Partyka MK, Mellon PL, Kauffman AS. 2015. Absent progesterone signaling in kisspeptin neurons disrupts the LH surge and impairs fertility in female mice. Endocrinology 156(9):3091-3097. doi:10.1210/en.2015-1300 PMID:26076042 PMCID:PMC4541622
  • Xie H, Hoffman HM, Meadows JD, Mayo S, Trang C, Leming SS, Maruggi C, Davis SW, Larder R, Mellon PL. 2015. Homeodomain proteins SIX3 and SIX6 regulate gonadotrope-specific genes during pituitary development. Mol Endocrinol 29(6):842-855. doi:10.1210/me.2014-1279 PMID:25915183 PMCID:PMC4447639


  • Ahow M, Min L, Pampillo M, Nash C, Wen J, Soltis K, Carroll RS, Glidewell-Kenney CA, Mellon PL, Bhattacharya M, Tobet SA, Kaiser UB, Babwah AV. 2014. KISS1R signals independently of Gαq/11 and triggers LH secretion via the β-arrestin pathway in the male mouse. Endocrinology 155(11):4433-4446. doi:10.1210/en.2014-1304 PMID:25147978 PMCID:PMC4197989
  • Breen KM, Mellon PL. 2014. Influence of stress-induced intermediates on gonadotropin gene expression in gonadotrope cells. Mol Cell Endocrinol 385(1-2):71-77. doi:10.1016/j.mce.2013.08.014 PMID:24012628 PMCID:PMC3942370
  • Glidewell-Kenney CA, Trang C, Shao PP, Gutierrez-Reed N, Uzo-Okereke AM, Coss D, Mellon PL. 2014. Neurokinin B induces c-fos transcription via protein kinase C and activation of serum response factor and Elk-1 in immortalized GnRH neurons. Endocrinology 155(10):3909-3919. doi:10.1210/en.2014-1263 PMID:25057795 PMCID:PMC4164922
  • Hoffmann HM, Tamrazian A, Xie H, Perez-Millan MI, Kauffman AS, Mellon PL. 2014. Heterozygous deletion of ventral anterior homeobox (vax1) causes subfertility in mice. Endocrinology 155(10):4043-4053. doi:10.1210/en.2014-1277 PMID:25060364 PMCID:PMC4164932
  • Roybal LL, Hambarchyan A, Meadows JD, Barakat AI, Pepa PA, Breen KM, Mellon PL, Coss D. 2014. Roles of binding elements, FOXL2 domains, and interactions with cJUN and SMADs in regulation of FSHβ. Mol Endocrinol 28(10):1640-1655. doi:10.1210/me.2014-1008 PMID:25105693 PMCID:PMC4179627




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