Brian McRae Barnes
Professor of Zoophysiology,
Director, Institute of Arctic Biology

Institute of Arctic Biology
Department of Biology and Wildlife
University of Alaska Fairbanks
Fairbanks, AK 99775

office (907) 474-6067 or 7649
laboratory (907) 474-6067
message (907) 474-7640
fax (907) 474-6967


• Research Interests
• Teaching
• Education and Professional Experience
• Awards and Research Experience
• Graduate Students
• Memberships
• Selected Publications


Last Modified on:

Faculty Listing

Institute of
Arctic Biology

Biology & Wildlife

University of Alaska

Research Interests

Physiological ecology and endocrinology of hibernating mammals; biological rhythms and sleep; overwintering biology of animals including insects. We work in field and laboratory settings investigating behavioral and physiological mechanisms by which animals cope with high latitude winter and summer environments.


  • Reproductive Biology of Vertebrates
  • Biological Rhythms
  • Physiological Ecology of Over-Wintering
  • Graduate Seminar in Physiology and Neurobiology

Education and Professional Experience

  • University of California, Riverside. B.S. in Biology, 1977
  • Stanford University, Hopkins Marine Station. Workshop on Biological Rhythms, 1977
  • University of Washington. Ph.D. in Zoology, 1983. G.J. Kenagy, advisor
  • University of California, Berkeley. Post-doctoral fellow, Departments of Zoology and Psychology, 1983-1986. P. Licht and I. Zucker, advisors
  • University of Alaska Fairbanks, Institute of Arctic Biology and Department of Biology and Wildlife. Assistant Professor of Zoophysiology, 1986-1991, Associate Professor, 1991-1999, Professor 2000. Interim Director, IAB 2001.
  • Secretary, Section in Comparative Endocrinology, American Society of Zoologists. 1994-1995
  • N.S.F. Panel for Dissertation Improvement Grants, Ecology and Evolutionary Biology, 1995
  • University of Tromsų, Norway. Department of Arctic Biology. Visiting Scholar, 1995
  • University of Groningen, The Netherlands, Chronobiology Research Group. Visiting Scholar, 1996
  • N.S.F. Workshop developing a mission statement for the Toolik Field Station
  • Chair, Toolik Field Station Steering Committee
  • Chair, Laurence Irving and Per Scholander Memorial Lecture Committee

Awards and Research Experience

  • University of Washington, Department of Zoology.
    • N.I.H. Cell and Molecular Biology Trainee, 1982-1983.
  • University of California, Berkeley.
    • N.I.H. Individual National Research Service Award Post-doctoral Fellowship, 1985-1986.
  • University of Alaska Fairbanks
  • .
    • UAF Faculty Research Grants Program: "Periodic arousals and reproductive development in hibernating mammals," 1987; "DNA fingerprinting in arctic ground squirrels," 1989; "Sleep regulation in heterothermic mammals," 1993.
    • NIH RO1 HD23383 (PI) "Endocrine responsiveness in heterothermic mammals," 1987-1996.
    • NIH K04 HD000973 (PI) “Research Career Development Award”, 1992-1997.
    • Durfee Foundation and Earthwatch (PI) “Student Field Expedition Awards”, 1990-1992.
    • NIH and NSF conference grants for symposium "Endocrinology of arctic birds and mammals," (co PI with P. Deviche) 1994.
    • NSF shared equipment grant (co-PI with D. Roby and others) 1994.
    • American Heart Institutional Award for Graduate Research (co-PI with B. Boyer) 1995‑1999.
    • NSF (PI) “An arctic residence facility for the Toolik Field Station, Alaska,” 1997-1999.
    • American Heart Association Standard Grant Award 98-AK-303 (co-PI with B. Boyer) “Multi-tissue regulation of energy expenditure by uncoupling proteins,” 1998-2001.
    • MJ Murdock Charitable Trust (PI) “Renovation of environmental chambers for support of arctic research,” 2000-2002.
    • NSF 9819540 (PI) “Energetics, homeostasis, and life history in an arctic hibernator,” 1999-2002.
    • NSF (PI) “Physiological limits to hibernation in large mammals,” 1999-2000.
    • NSF (PI) “A cooperative proposal for the Toolik Field Station, Alaska,” 2000-2005; renewed 2005-2010.
    • NSF EPSCoR (Research Focus Leader) “Integrative Approaches to Environmental Physiology,” 2000-2003.
    • NSF (PI, collaborative research with J. Duman) “Studies on antifreeze proteins in arctic and subarctic insects,” 2001-2006, 2006-2008.
    • US Army Medical Research and Materiel Command (PI) “Hibernation genomics” 2005-2010.
    • NSF (co-PI) “International Polar Year -- Arctic Observatory Network” 2007-2010.
    • NSF (co-PI with D. O’Brien) “IPY: Extremes of hibernation physiology: patterns of expression, regulation, and limits” 2007-2010.

Graduate Students

  • Sylvia Feder, M.S. 1990
  • Mark Reed, M.S., 1993
  • Olav Ormseth, M.S. 1997 (co-advised with B. Boyer)
  • C. Loren Buck, Ph.D. 1998
  • Kelly Hochstetler, Ph.D. (withdrew)
  • William Toxvard, M.S. 2001
  • Shawna Karpovitch M.S. 2002
  • Jamie Barger, Ph.D. 2002 (co-advised with B. Boyer)
  • Øivind Tøien, Ph.D., post-doc then Research Associate, 1997-current
  • Petter Kvadsheim, Ph.D., post-doc, 2001
  • Todd Sfromo, Ph.D., 2009
  • Melanie Griffith, Ph.D., current
  • Benjamin Gaglioti MS., current.


  • Sigma Xi
  • AAAS
  • American Society of Mammalogists
  • American Society of Physiology
  • Society for the Study of Reproduction
  • Society for Research on Biological Rhythms
  • Society for Neuroscience
  • Society for Integrative and Comparative Biology

Selected Publications

  • Tøien Ø, J Blake, DM Edgar, DA Grahn, HC Heller, and BM Barnes (2011) Hibernation in black bears: Independence of metabolic suppression from body temperature. Science 331:906-909.
  • Williams CT, AV Goropashnaya, CL Buck, VB Fedorov, F Kohl, TN Lee, and BM Barnes (2011) Hibernating above the permafrost: Effects of ambient temperature and season on expression of metabolic genes in liver and brown adipose tissue of arctic ground squirrels. J Exp Biol
  • Walters Jr KR, A S Serianni, Y Voituron, T Sformo, BM Barnes, JG Duman (2011) A thermal hysteresis-producing xylomannan glycolipid antifreeze associated with cold tolerance is found in diverse taxa. Journal of Comparative Physiology – B
  • Sheriff MJ, Kenagy GJ, Richter M, Lee T, Toien O, Kohl F, Buck CL, Barnes BM (2011) Phenological variation in annual timing of hibernation and breeding in nearby populations of arctic ground squirrels. Proceedings of the Royal Society B, doi:10.1098/rspb.2010.2482
  • Coltrane J, Farley S, Barboza P, Kohl F, Sinnott R, Barnes BM (2011) Seasonal regulation of body composition, water turnover, and field metabolic rates in captive and free-ranging North American porcupines (Erethizon dorsatum) in Alaska. J Mammal
  • Stieler JT, Bullmann T, Kohl F, Tøien Ø, Bruckner MK, Hartig W, Barnes BM, Arendt T. (2011) The physiological link between metabolic rate depression and tau phosphorylation in mammalian hibernation. PLoS ONE
  • Liu Y, Hu W, Wang H, Lu M, Shao C, Menzel C, Yan Z, Ying L, Zhao S, Khaitovich P, Liu M, Chen W, Barnes BM, and Yan J (2010) Genomic analysis of miRNAs in an extreme mammalian hibernator, the Arctic Ground Squirrel. Physiological Genomics 42A:39-51.
  • Zhao S, Shao C, Goropashnaya AV, Stewart NC, Xu Y, Tøien Ø, Barnes BM, Fedorov VB and Yan J (2010) Genomic analysis of expressed sequence tags in American black bear Ursus americanus. BMC Genomics 9:313-326.
  • Sformo T, Walters K, Jeannet K, Wowk B, Fahy G, Barnes BM, Duman JG (2010) Deep supercooling, vitrification, and limited survival to -100 C in the Alaskan beetle larvae Cucujus clavipes puniceus. (Coleoptera: Cucujidae) J Exp Biol 213:502-509.
  • Shao C, Liu Y, Ruan H, Li Y, Wang H, Kohl, Goropashnaya AV, Fedorov VB, Zeng R, Barnes BM, and Yan J (2010) Shotgun Proteomic Analysis of Hibernating Arctic Ground Squirrels. Mol Cell Proteomics 9:313-326.
  • Lee TN, Barnes BM, Buck CL (2009) Body temperature patterns during hibernation in a free-living Alaska marmot (Marmota broweri). Ethology Ecology & Evolution 21: 403-413.
  • Walters KR, Serianni A, Sformo T, Barnes BM, Duman JG (2009) A nonprotein thermal hysteresis-producing zylomannan antifreeze in the freeze tolerant Alaskan beetle Upis ceramboides. PNAS 106(48): 20210-20215.
  • Stieler JT, Bullmann T, Kohl F, Barnes BM, Arendt T (2009) PHF-like tau phosphorylation in mammalian hibernation is not associated with p25-formation. J Neural Transm 116:345-350.
  • Karpovich SA, Tøien Ø, Buck CL, Barnes BM (2009) Energetics of arousal episodes in hibernating arctic ground squirrels. J Comp Physiol B 179:691-700.
  • Duman JG, Walters KR, Sformo T, Carrasco MA., Nickell P. and Barnes BM (2009) Antifreeze and ice nucleator proteins. In Insects at Low Temperature (eds. D. L. Denlinger and R. E. Lee) 2nd edition Cambridge Press pp 59-90.
  • Sformo T, Kohl F, McIntyre, Kerr  P, Duman JG, and Barnes BM (2009) Simultaneous freeze tolerance and avoidance in individual fungus gnats, Exechia nugatoria. J Comp Physiol B. 179:897-902.
  • Fedorov VB, Goropashnaya AV, Tøien Ø, Stewart NC, Gracey AY, Chang C, Qin S, Pertea G, Quackenbush J, Showe LC, Showe MK, Boyer BB and Barnes BM (2009) Elevated expression of protein biosynthesis genes in liver and muscle of hibernating black bears (Ursus americanus). Physiological Genomics 37:108-118.
  • Walters KR, Sformo T, Barnes BM, and Duman JG (2009) Freeze tolerance in an Arctic Alaska stonefly. Journal of Experimental Biology 212: 305-312
  • Frank, CL, Karpovich SA, Barnes BM (2008) Dietary fatty acid composition and the hibernation patterns in free-ranging arctic ground squirrels. Physiological and Biochemical Zoology 81(4):486-495.
  • Stieler JT, AS Boerrema, T Bullmann, F Kohl, AM Strijkstra, BM Barnes, T Arendt (2008) Activity-state profile of tau kinases in hibernating animals. In Lovegrove BG and McKechnie AE (eds.) Hypometabolism in animals: torpor, hibernation and cryobiology. University of KwaZulu-Natal, Pietermaritzburg  133-142.
  • Buck CL, Breton A, Tøien Ø, Barnes BM (2008) Overwinter body temperature patterns of free-living arctic ground squirrels (Spermophilus parryii). In Lovegrove BG and McKechnie AE (eds.) Hypometabolism in animals: torpor, hibernation and cryobiology. University of KwaZulu-Natal, Pietermaritzburg  317-326.
  • Yan J., BM Barnes, F. Kohl, and TG Marr (2008) Modulation of gene expression in hibernating arctic ground squirrels. Physiol Genomics 32:170-181.
  • Long RA, RA Hut, BM Barnes (2007) Simultaneous collection of body temperature and activity data in burrowing mammals: a new technique. J Wildlife Management 71(4):1375-1379.
  • Drew KL, CL Buck, BM Barnes, SL Christian, BT Rasley, and MB Harris (2007) Central nervous system regulation of mammalian hibernation: implications for metabolic suppression and ischemia tolerance. J of Neurochemistry 10.1111/j.1471-4159.2007.04675.x
  • Van der Heyden MAG, B Kok, En Kouwenhoven, Ø Tøien, BM Barnes, VG Federov, IR Efimov, and T Opthof (2007) Cloning, sequence analysis and phylogeny of connexin43 isolated from American black bear. DNA Sequence 1-5.
  • Barger JL, BM Barnes, BB Boyer (2006) Regulation of UCP1 and UCP3 in arctic ground squirrels and relation with mitochondrial leak. J Appl Physiol 101:339-347. (Highlighted topic, with invited editorial).
  • Yan J, A Burman , C Nichols, L Alila, LC Showe, MK Showe, BB Boyer, BM Barnes and TG Marr (2006) Detection of differential gene expression in brown adipose tissue of hibernating arctic ground squirrels with mouse microarrays. Physiol Genomics 25: 346-353, 2006.
  • Bennett VA, T Sformo, K Walters, O Toien, K Jeannet, R Hochstrasser, Q Pan, AS Serianni, BM Barnes, JG Duman (2005) Comparative overwintering physiology of Alaska and Indiana populations of the beetle Cucujus clavipes (Fabricius): roles of antifreeze proteins, polyols, dehydration and diapause. J Exp Biol 208:4467-4477.
  • Long RA, TJ Martin, and BM Barnes (2005) Body temperature and activity patterns in free-living arctic ground squirrels. J Mammal 86(2):312-322.
  • Duman JG, VA Bennett, N Li, L Wang, L Huang, T Sformo, and BM Barnes (2004) Antifreeze proteins in terrestrial arthropods. in Barnes, BM, CV Carey, editors. (2004) Life in the Cold: Evolution, mechanisms, adaptation, and applications. Biological Publications of the University of Alaska, Institute of Arctic Biology no 27., pp 527-542.
  • Barnes, BM, CV Carey, editors (2004) Life in the Cold: Evolution, mechanisms, adaptation, and applications. Biological Publications of the University of Alaska, Institute of Arctic Biology no 27. 602pp.
  • Kenagy, GJ, RA Vasquez, BM Barnes, F Bozinovic (2004) Microstructure of summer activity bouts of degus in a thermally heterogeneous habitat. J Mammal 85(2):2600-267.
  • Duman JG, Bennett V, Sformo T, Hochstrasser R, Barnes BM (2004). Antifreeze proteins in Alaskan insects and spiders. J Insect Physiol 50:259-66. (Abstract online)
  • Barger JL, Brand MD, Barnes BM, and Boyer BB (2003). Tissue-specific depression of mitochondrial proton leak and substrate oxidation in hibernating arctic ground squirrels. Am J Physiol Regul Integr Comp Physiol 284:1306-1313. (Abstract online)
  • Buck CL and Barnes BM (2003). Androgen in free-living arctic ground squirrels: seasonal changes and influence of staged male-male aggressive encounters. Horm Behav 43:318-26 (Abstract online)
  • Hut RA, Barnes BM, and Daan S (2002) Body temperature patterns during semi-natural hibernation in the European ground squirrel. J Comp Physiology and Biochem B 172(1):47-58. (Abstract online)
  • Barnes BM and Buck CL (2000) Hibernation in the extreme: burrow and body temperatures, metabolism, and limits to torpor bout length in arctic ground squirrels. in Living in the Cold. G Heldmaier, M Klingenspor, eds. Springer-Verlag, Berlin. pp. 65-72.
  • Buck, CL and BM Barnes. (2000). Effects of ambient temperature on metabolic rate, respiratory quotient, and torpor in an arctic hibernator. Amer. J. Physiol R255-R262. (online)
  • Knight JE, EN Narus, SL Martin, A Jacobson, BM Barnes, BB Boyer. (2000). mRNA stability and translational inhibition in hibernating arctic ground squirrels (Spermophilus parryii) Molecular and Cellular Biology 20:6374-6379.
  • Linnell JDC, Swenson JE, Andersen R, Barnes BM (2000) How vulnerable are denning bears to disturbance? Wildlife Society Bull 28(2): 400-413.
  • Boyer BB and BM Barnes (1999) Molecular and metabolic aspects of hibernation. Bioscience. 49:713-724.
  • Buck CL and BM Barnes (1999) Temperatures of hibernacula and changes in body composition of arctic ground squirrel over winter. Journal of Mammalogy 80:1264-1276.
  • Buck CL and BM Barnes (1999) Annual cycle of body condition and hibernation in free-living arctic ground squirrels. Journal of Mammalogy. 80(2):430-442.
  • Osborne, PG, Hu Y, Covey DN, Barnes BM, Katz Z, and Drew KL. (1999) Determination of striatal extracellular gama-aminobutyric acid in non-hibernating and hibernating arctic ground squirrels using quantitative microdialysis. Brain Research 839:1-6.
  • Boyer BB, BM Barnes, BB Lowell, and D Grujic (1998) Differential regulation of uncoupling protein gene homologs in multiple tissues of hibernating ground squirrels. American J Physiology: Regulatory, Integrative & Comparative. 275: R1232-R1238.
  • Boyer BB, OA Ormseth, L Buck, M Nicolson, MA Pelleymounter, and BM Barnes (1998). Leptin prevents post-hibernation weight gain but does not reduce energy expenditure in arctic ground squirrels. Comp Biochem Physiol:118C:405-412. (Abstract online)
  • Barnes BM. (1996). Relationships between hibernation and reproduction in male ground squirrels. In: Adaptations to the Cold: Tenth International Hibernation Symposium. Geiser F et al, eds. University of New England Press, Armidale, Australia. 71-80.
  • Barnes BM. (1996). Torpor in arctic ground squirrels. The Sciences September/October:12-14.
  • Barnes BM, Barger JL, Seares J, Tacquard PC, and Zuercher GL. (1996). Overwintering in yellowjacket queens (Vespula vulgaris) and green stinkbugs (Elasmostethus interstinctus) in subarctic Alaska. Physiol Zool 69: (1469-1480)
  • Cossins AR and BM Barnes. (1996). Hibernation: Southern discomfort. Nature 382:582-583
  • Linnell JDC, Swenson J, Barnes B, and Andersen R. (1996). How vulnerable are denning bears to disturbance? A review. A study in connection with plans for the establishment of a military training aria in Ostlandet, Norway. Paart 2. - NINA Oppdragsmelding 413:1-19.
  • Deviche P and Barnes BM. (1995). Introduction to the symposium: Endocrinology of arctic birds and mammals. Am Zool 35:189-190.
  • Barnes BM, Omtzigt C and Daan S. (1993). Hibernators periodically arouse in order to sleep. In: Life in the Cold: Ecological, Physiological, and Molecular Mechanisms. Carey, C et al, ed. Westview Press, Boulder, CO. 555-558.
  • Barnes BM and Ritter D. (1993). Patterns of body temperature change in hibernating arctic ground squirrels. In: Life in the Cold: Ecological, Physiological, and Molecular Mechanisms. Carey C et al, ed. Westview Press, Boulder, CO. 119-130.
  • Boyer BB, Barnes BM, Kopecky J, Jacobsson A, and Hermanska J. (1993). Molecular control of prehibernation brown fat growth in arctic ground squirrels. In: Life in the Cold: Ecological, Physiological, and Molecular Mechanisms. Carey C et al, ed. Westview Press, Boulder, CO. 483-492.
  • Daan S, Strijkstra A, and Barnes BM. (1992). The elusive function of sleep: New insight from hibernators? In: Circadian clocks from cell to human. Hiroshige T and Honma K, ed. Hokkaido Univ. Press, Hokkaido. 165-176.
  • Daan S, Barnes BM, and Strijkstra AM (1991). Warming up to sleep? Hibernating arctic ground squirrels sleep during arousals. Neurosci Lett 128:265-268.
  • Barnes BM and York AD (1990). Effect of winter temperatures on reproduction and circannual rhythms in hibernating ground squirrels. J Biol Rhythms: 5:119-130. (Abstract online)
  • Barnes BM (1989). Freeze avoidance in a mammal: body temperatures below 0°C in an arctic hibernator. Science 244:1593-1595.
  • Ruby NF, Ibuka N, Barnes BM, and Zucker I (1989) Suprachiasmatic nuclei influence torpor and circadian temperature rhythms in hamsters. Am J Physiol Regul Integr Comp Physiol 257:210-215 (Abstract online)
  • Barnes BM, Kretzmann M, Zucker I, and Licht P (1988). Plasma androgen and gonadotropin levels during hibernation and testicular maturation in golden-mantled ground squirrels. Biol Reproduction 38:616-622 (Abstract online)
  • Barnes BM (1986). Annual cycles of gonadotropins and androgens in the hibernating golden-mantled ground squirrel. Gen Comp Endocrinol 62:13-22. (Abstract online)
  • Barnes BM, Kretzmann M, Licht P, and Zucker I (1986). The influence of hibernation on testis growth and spermatogenisis in the golden-mantled ground squirrel, Spermophilus lateralis. Biol Reproduction 35:1289-1297. (Abstract online)
  • Forger NG, Dark J, Barnes BM and Zucker I (1986). Fat ablation and food restriction influence reproductive development and hibernation in ground squirrels. Biol Reprod 34:831-840. (Abstract online)
  • Barnes BM (1984). Influence of energy stores on activation of reproductive function in male golden-mantled ground squirrels. J Comp Physiol B 154:421-425.
  • Kenagy GJ and Barnes BM (1984). Environmental and endogenous control of reproductive function in the Great Basin pocket mouse Perognathus parvus. Biol Reprod 31:637-645. (Abstract online)
  • Pengelly ET, Aloia RC, Barnes BM, and Whitson D (1979). Differential temporal behavior between males and females in the hibernating ground squirrel, Citellus lateralis. Comp Biochem Physiol 64A:593-596.
  • Pengelly ET, Aloia RC, and Barnes BM (1978). Circannual rhythmicity in the hibernating ground squirrel Citellus lateralis under constant light and hyperthermic ambient temperature. Comp Biochem Physiol 61A:599-603.
  • Pengelly ET, Asmundson SJ, Barnes B, and Aloia RC (1976). Relationship of light intensity and photoperiod to circannual rhythmicity in the hibernating ground squirrel Citellus lateralis. Comp Biochem Physiol 53A:273-277.

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