Stephen McCormickStephen McCormick

Adjunct Professor

Senior Scientist and Research Physiologist


Phone: 413 863-3804

Office: USGS, Conte Anadromous Fish Research Center, Turners Falls, MA

Lab Website:

Primary Interests

Environmental Physiology of Migratory Fishes

Physiological demands are especially important to anadromous fish because of their need to move between freshwater and seawater and to make long, often difficult, migrations. These demands can become critical when there are barriers to migration or other changes in river ecosystems.  The McCormick Laboratory is located 20 miles north of campus at the USGS Conte Anadromous Fish Research Center. We conduct research on physiological and endocrine changes that may limit fish populations, particularly those in need of conservation or restoration. We are particularly interested in how environmental change, both of ‘natural’ and ‘human’ origins, affects normal growth and development in anadromous fish. Physiological systems such as ion regulation, stress, energetics, growth and reproduction are critical for survival of migratory fish. Environmental factors such as climate change, salinity, temperature, stress, hatchery practices, dams and pollution may affect normal development of these complex systems. Because of threats to their survival in many rivers in the northeastern US, Atlantic salmon, American shad, blueback herring, alewife, brook trout, sturgeon and lamprey are the anadromous species of primary interest.

Current Projects

  • Identification, localization and endocrine regulation of gill ion transport proteins that secrete salts as anadromous fish move between freshwater and seawater.
  • Thermal physiology of salmonids, including brook trout, Atlantic salmon and Pacific salmon.
  • Hormonal control of osmoregulation in basal vertebrates, including sea lamprey and Atlantic sturgeon.
  • Mechanisms of photoperiodic signaling in fish.

Courses Taught

Migratory Fishes


Ph.D.:   Massachusetts Institute of Technology/Woods Hole

Oceanographic Institution Joint Program in Biological Oceanography

September 1983

Dissertation: “Effects of size, age and photoperiod on hypoosmoregulation in brook trout, Salvelinus fontinalis.”


Undergraduate: Bates College, Lewiston, Maine

B.S., June 1977

Majors:  Biology and Economics

Selected Publications

Recent Publications (2015-present)

McCormick, S.D., Taylor, M.L. and Regish, A.M.  2020.  Cortisol is an osmoregulatory and glucose-regulating hormone in Atlantic sturgeon, a basal ray-finned fish.  Journal of Experimental Biology 223, jeb220251. doi:10.1242/jeb.220251.

Kjærner-Semb, E., R. B. Edvardsen, F. Ayllon, P. Vogelsang, T. Furmanek, C. J. Rubin, A. E. Veselov, T. O. Nilsen, S. D. McCormick, C. R. Primmer, and A. Wargelius.  2020.  Comparison of anadromous and landlocked Atlantic salmon genomes reveals signatures of parallel and relaxed selection across the northern hemisphere.  Evolutionary Applications Online early September 13. doi:10.1111/eva.13129.

McCormick, S. D., E. Schultz, and C. J. Brauner.  2020. Osmoregulation and acid-base balance. In S. Midway, C. Hasler, and P. Chakrabarty, editors. Methods for Fish Biology, 2nd edition. American Fisheries Society, Bethesda, Maryland. (in press).

von Biela, V.R., L. Bowen, S.D. McCormick, M.P. Carey, D.S. Donnelly, S. Waters, A. Regish, S.M. Laske, R.J. Brown, S. Larson, S. Zuray, C.E. Zimmerman.  2020.  Evidence of prevalent heat stress in Yukon River Chinook salmon. Canadian Journal of Fisheries and Aquatic Sciences (in press).

Bowen, L, V.R. von Biela, S.D. McCormick, A.M. Regish, S. Waters, B. Durbin-Johnson, M. Britton, M. Settles, D.S. Donnelly, S. Laske, M.P. Carey, R.J. Brown, C.E. Zimmerman.  2020. Transcriptomic response to elevated water temperatures in adult migrating Yukon River Chinook salmon (Oncorhynchus tshawytscha). Conservation Physiology 8(1): coaa084. doi:10.1093/conphys/coaa084.

Donnelly, D.S., V.R. von Biela, S.D. McCormick, S. Laske, M.P. Carey, S. Waters, L. Bowen, R.J. Brown, S. Larson, and C.E. Zimmerman.  2020.  A manipulative experimental thermal challenge protocol for adult salmonids in remote field settings. Conservation Physiology 8(1): coaa074. doi:10.1093/conphys/coaa074.

van Rijn, C.A., Jones, P.L., Schultz, A.G., Evans, B.S., McCormick, S.D., and Afonso, L.O.B.  2020.  Atlantic salmon (Salmo salar) exposed to different preparatory photoperiods during smoltification show varying responses in gill Na+/K+-ATPase, salinity-specific mRNA transcription and ionocyte differentiation. Aquaculture 529: 735744.   doi:10.1016/j.aquaculture.2020.735744

Shaughnessy, C.A., Barany-Ruiz, A. and McCormick, S.D.  2020.  11‑Deoxycortisol controls hydromineral balance in the most basal osmoregulating vertebrate, sea lamprey (Petromyzon marinus).  Scientific Reports 10: 12148.  DOI:10.1038/s41598-020-69061-4

Shaughnessy, C.A. and McCormick, S.D.  2020.  Functional characterization and osmoregulatory role of the Na+-K+-2Cl cotransporter in the gill of sea lamprey (Petromyzon marinus), a basal vertebrate.  American Journal of Physiology Regulatory Integrative and Comparative Physiology 318: R17–R29.  DOI:10.1152/ajpregu.00125.2019.

Barany-Ruiz, A., Shaughnessy, C.A., Fuentes, J., Mancera, J.M. and McCormick, S.D.  2020. Osmoregulatory role of the intestine in the sea lamprey (Petromyzon marinus).  American Journal of Physiology Regulatory Integrative and Comparative Physiology 318: R410–R417. DOI:10.1152/ajpregu.00033.2019.

Gong, N., Ferreira-Martins, D., McCormick S.D. and Sheridan, M.A.  2020.  Divergent genes encoding the putative receptors for growth hormone and prolactin in sea lamprey display distinct patterns of expression. Scientific Reports 10: 1674. DOI: 10.1038/s41598-020-58344-5

Archer, L.C., Hutton, S.A., Harman, L., McCormick, S.D., O’Grady, M.N., Kerry, J.P., Poole, W.R., Gargan, P., McGinnity, P., and Reed, T.E.  2020.  Food and temperature stressors have opposing effects in determining flexible migration decisions in brown trout (Salmo trutta). Global Change Biology 26(5): 2878-2896.  DOI: 10.1111/gcb.14990

Bayse, S., Shaughnessy, C., Regish, A. and McCormick, S.D.  2020.  Upper thermal tolerance and heat shock protein response of juvenile American shad (Alosa sapidissima).  Estuaries and Coasts 43:182–188.  DOI: 10.1007/s12237-019-00642-x

van Rijn, C.A., Jones, P.L., Evans, B.S., Huynh, C., McCormick, S.D. and Afonso, L.O.B.  2020.  Characterization of smoltification in the Tasmanian strain of Atlantic salmon (Salmo salar) in recirculation and flowthrough systems.  Aquaculture 516: 734603. DOI:10.1016/j.aquaculture.2019.734424

Roy, M.A., Sant, K.E., Venezia, O.L., Shipman, A.B., McCormick, S.D., Saktrakulkla, P., Hornbuckle, K.C. and Timme-Laragy, A.R. 2019. The emerging contaminant 3,30-dichlorobiphenyl (PCB-11) impedes Ahr activation and Cyp1a activity to modify embryotoxicity of Ahr ligands in the zebrafish embryo model (Danio rerio). Environmental Pollution 254: 113027.  DOI: 1016/j.envpol.2019.113027

Bayse, S.M., McCormick, S.D. and Castro-Santos, T.  2019.  How lipid content and temperature affect American shad (Alosa sapidissima) attempt rate and sprint swimming: implications for overcoming migration barriers.   Canadian Journal of Fisheries and Aquatic Sciences 76:2235-2244.  DOI: 10.1139/cjfas-2018-0406

Lema, S.C., Washburn, E.H., Crowley, M.E., Carvalho, P.G., Egelston, J.N. and McCormick, S.D.  2019.  Evidence for a role of arginine vasotocin (AVT) receptors in the gill during salinity acclimation by a euryhaline teleost fish.  American Journal of Physiology; Integrative, Regulatory and Comparative Physiology 316: R735–R750. DOI: 10.1152/ajpregu.00328.2018

Carey, M.P., Kevin D. Keith, Merlyn Schelske, Charlie Lean, Stephen D. McCormick, Amy Regish, and Christian E. Zimmerman.  2019.  Energy depletion and stress levels of Sockeye Salmon migrating at the northern edge of their distribution.  Transactions of the American Fisheries Society 148:785–797.  DOI: 10.1002/tafs.10172

Fleming, M.S., Maugars, G., Lafont, A.-G., Rancon, J., Fontaine, R., Nourizadeh-Lillabadi, R., Weltzien, F.A., Yebra-Pimentel, E.S., Dirks, R., McCormick, S.D., Rousseau, K, Martin, P. &  Dufour, S.  2019.  Functional divergence of thyrotropin beta-subunit paralogs gives new insights into salmon smoltification metamorphosis.  Scientific Reports 9:4561.  DOI: 10.1038/s41598-019-40019-5

McCormick, S.D., Regish, A.M., Ardren, W.R., Björnsson, B.Th. and Bernier, N.J.  2019.  The evolutionary consequences for seawater performance and its hormonal control when anadromous Atlantic salmon become landlocked.  Scientific Reports (2019) 9:968.  DOI: 10.1038/s41598-018-37608-1

McCormick, S.D. and Regish, A.M.  2018.  Effects of ocean acidification on salinity tolerance and seawater growth of Atlantic salmon Salmo salar smolts.  Journal of Fish Biology 93:560-566.  DOI: 10.1111/jfb.13656

Vargas-Chacoff, L., Regish, A.M., Weinstock, A. and McCormick, S.D.  2018.  Effects of elevated temperature on osmoregulation and stress responses in Atlantic salmon Salmo salar smolts in fresh water and seawater.  Journal of Fish Biology 93:550-559.  DOI: 10.1111/jfb.13683

Breves, J.P., Tara A. Duffy, Ingibjörg E. Einarsdottir, Björn Thrandur Björnsson, Stephen D. McCormick.  2018.  In vivo effects of 17α-ethinylestradiol, 17β-estradiol and 4-nonylphenol on insulin-like growth-factor binding proteins (igfbps) in Atlantic salmon.  Aquatic Toxicology 203:28-39.  DOI: 10.1016/j.aquatox.2018.07.018

Lema, S.C., Carvalho, P.G., Egelston, J.N., Kelly, J.T., McCormick, S.D.  2018.  Dynamics of gene expression responses for ion transport proteins and aquaporins in the gill of a euryhaline pupfish during freshwater and high salinity acclimation. Physiological and Biochemical Zoology 91(6), 1148-1171. DOI: 10.1086/700432

Regish, A.M., Kelly, J.T., O’Dea, M.F. and McCormick, S.D.  2018.  Effect of acid and aluminum exposure on osmoregulation, gill Na+/K+-ATPase isoforms and ionocytes in Atlantic salmon smolts.  Canadian Journal of Fisheries and Aquatic Sciences 75: 1319–1328.

Shaughnessy, C.A. and McCormick, S.D.  2018.  Reduced thermal tolerance during salinity acclimation in brook trout (Salvelinus fontinalis) can be rescued by prior treatment with cortisol.

Journal of Experimental Biology 221, jeb169557. doi:10.1242/jeb.169557

Christensen, A.K., Regish, A.M. and McCormick, S. D.  2018.  Shifts in the relationship between mRNA and protein abundance of gill iontransporters during smolt development and seawater acclimation in Atlantic salmon (Salmo salar).  Comparative Biochemistry and Physiology, Part A 221: 63-73.  doi:10.1016/j.cbpa.2018.03.020

Bayse, S.M., Regish, A.M. and McCormick, S. D.  2018.  Proximate composition, lipid utilization and validation of a non-lethal method to determine lipid content in migrating

American shad Alosa sapidissimaJournal of Fish Biology 92, 1832–1848. doi:10.1111/jfb.13624

Velotta, J.P., McCormick, S.D., Jones, A.W. and Schultz, E.T.  2018.  Reduced Swimming Performance Repeatedly Evolves on Loss of Migration in Landlocked Populations of Alewife.  Physiological and Biochemical Zoology 91: 814-825.  doi: 10.1086/696877

Galt, N.J., McCormick, S.D., Froehlich, J.M. and Biga, P.R.  2018.  A comparative evaluation of crowding stress on muscle HSP90 and myostatin expression in salmonids.  Aquaculture 483: 141-148.  doi: 10.1016/j.aquaculture.2017.10.019

Chadwick, J.G. and McCormick, S.D.  2017.  Upper thermal limits of growth in brook trout and their relationship to stress physiology.  Journal of Experimental Biology 220, 3976-3987. doi:10.1242/jeb.161224

Morshedian, A., Toomey, M.B., Pollock, G.E., Frederiksen, R., Enright, J.M., McCormick, S.D., Cornwall, M.C., Fain, G.L., Corbo, J.C.  2017.  Cambrian origin of the CYP27C1-mediated

vitamin A1-to-A2 switch, a key mechanism of vertebrate sensory plasticity.  Royal Society of Open Science 4: 170362.  doi:10.1098/rsos.170362

Nyqvist, D., S. D. McCormick, L. Greenberg, W. R. Ardren, E. Bergman, O. Calles & T. Castro-Santos.  2017.  Downstream Migration and Multiple Dam Passage by Atlantic Salmon Smolts.  North American Journal of Fisheries Management 37: 816-828. doi:10.1080/02755947.2017.1327900

Zak, M.A., Regish, A.M. and McCormick, S. D. and Manzon, R.G.  2017.  Exogenous thyroid hormones regulate the activity of citrate synthase and cytochrome c oxidase in warm- but not cold-acclimated lake whitefish (Coregonus clupeaformis).  General and Comparative Endocrinology 247: 215-222.  doi:10.1016/j.ygcen.2017.02.005

McCormick, S.D. and Romero, J.M.  2017.  Conservation Endocrinology.  Bioscience 67: 429-442.  doi:10.1093/biosci/bix026

Velotta, J.P., Wegrzyn, J.L., Ginzburg, S., Kang, L., Czesny, S., O’Neill, R.J., McCormick, S.D., Michalak, P. and Schultz, E.T.  2017.  Genomic imprints of adaptation to freshwater: parallel transcriptional responses to osmotic challenge are associated with physiological trade-offs in landlocked Alewives.  Molecular Ecology 26:831-848.  doi: 10.1111/mec.13983

Hollemaa, H.M., Kneebone, J. McCormick, S.D., Skomal, G.B. and Danylchuk, A.J.  2017.   Movement patterns of striped bass (Morone saxatilis) in a tidal coastal embayment in New England. Fisheries Research 187: 168-177. doi: 10.1016/j.fishres.2016.11.006

Breves, J.P., Fujimoto, C.K., Phipps-Costin, S.K., Einarsdottir, I.E., Björnsson, B.Th. and McCormick, S.D.  2017.  Variation in branchial expression among insulin-like growth-factor binding proteins (igfbps) during Atlantic salmon smoltification and seawater exposure. BMC Physiology (2017) 17:2.  doi: 10.1186/s12899-017-0028-5

Balk, L. Hägerroth, P.-A. Gustavsson, H., Sigg, L., Åkerman, G., Muñoz, Y.R., Honeyfield, D.C.  Tjärnlund, U., Oliveira, K., Ström, K., McCormick, S.D., Karlsson, S., Ström, M., Manen, M.,   Berg, A.-L., Halldórsson, H.P., Strömquist, J., Collier, T.K., Börjeson, H., Mörner, T. and Hansson, T.  2016.  Widespread episodic thiamine deficiency in Northern Hemisphere

wildlife. Scientific Reports 6, 38821; doi: 10.1038/srep38821.

Galt, N.J., McCormick, S.D., Froehlich, J.M. and Biga, P.R.  2016.  A comparative examination of cortisol effects on muscle myostatin and HSP90 gene expression in salmonids.  General and Comparative Endocrinology 237: 19-26.  doi: 10.1016/j.ygcen.2016.07.019

Ferreira-Martins, D., McCormick, S.D., Campos, A., Lopes-Marques, M, Osório, H.,

Coimbra, J, Castro, L. F. C. and Wilson, J.M.  2016.  A cytosolic carbonic anhydrase molecular switch occurs in the gills of metamorphic sea lamprey. Scientific Reports 6, 33954. doi: 10.1038/srep33954.

Divino, J.N., Monette, M.Y., McCormick, S.D., Yancey, P.H., Flannery, K.G., Bell, M.A., Rollins, J.L., von Hippel, F.A., and Schultz, E.T.  2016.  Osmoregulatory physiology and rapid evolution of salinity tolerance in recently lake-introduced Threespine Stickleback.  Evolution and Ecology Research 17: 179-201.

Breves, J.P., Phipps-Costin, S.K., Fujimoto, C.K., Einarsdottir, I.E., Regish, A.M., Björnsson, B.Th. and McCormick, S.D.  2016.  Hepatic insulin-like growth-factor binding protein (igfbp) responses to food restriction in Atlantic salmon smolts.  General and Comparative Endocrinology 233: 79-87.

Velotta, J.P., McCormick, S.D. and Schultz, E.T.  2015.  Trade-offs in osmoregulation and parallel shifts in molecular function following ecological transitions to freshwater in the Alewife.  Evolution 69: 2676-2688.

Kelly, J.T., Lerner, D. T., O’Dea, M. F., Regish, A. M., Monette, M.Y., Hawkes, J.P., Nislow, K.H. and McCormick, S.D.  2015.  Evidence for episodic acidification impacts on migrating Atlantic salmon smolts in the United States.  Journal of Fish Biology 87: 1129-1146.  doi:10.1111/jfb.12763

Chadwick, J.G., Nislow, K. and McCormick, S.D.  2015.  Thermal onset of cellular and endocrine stress responses correspond to ecological limits in brook trout, an iconic cold-water fish.  Conservation Physiology 3: doi:10.1093/conphys/cov017.

Other Publications (prior to 2015)