Marilyn Ramenofsky

image of Marilyn Ramenofsky

Adjunct Professor


Neurobiology, Physiology and Behavior

Offices and Labs

104 Animal Communications Lab
530 400 8945


1982 PhD Zoology University of Washington, Seattle
1972 MSc Zoology University of Texas, Austin
1969 BA Botany-Biology Pomona College

Research Contribution

I am fascinated by the interplay of environmental signals and the physiological and behavioral expressions of organisms and primarily those of migratory birds. As migration is the movement to and away from areas where species breed, each trip is composed of multiple steps or substages that involve regulation of the neuroendocrine, endocrine, metabolic, and behavioral systems. During these trips, organisms are exposed to a wide range environmental conditions that include photoperiod, atmospheric and climatic, food availability, predation and unpredictable pertubations. Thus, in order for migrants to be successful they rely on environmental information to cue progression of each substage. Migrants therefore can serve as an effective model for studying how organisms rely on environmental conditions and respond to environmental changes and in particular global climate fluctuations. Former studies have focused on the effects of photoperiod, lunar condition, food availability and stressors on migratory specific characteristics that include migratory behavior (migratory restlessness, feeding, locomotor activity) and measures of the endocrine and metabolic systems. My other studies include vocal communication of nocturnal migrants, role of experience and geomagnetic field on orientation during migration, endogenous rhythms and migratory behavior, effects of prolonged migratory restlessness on the endocrine and metabolic systems. Most recently my laboratory has used the migrant/resident model present in the Pacific White-crowned Sparrows subspecies (Zonotrichia leucophrys) to compare the physiological and behavioral traits attributed to the migratory life history and assess whether these features are present in residents and may be considered more in seasonal terms rather then strictly migratory.


Howard Hughes Award for development of Ingraham High School Urban Ecology Project, 2002
NSF Travel Award to attend 19th Int’l Ethology Conference, Tolouse, France, 1985
Office of the Dean, University of Washington, 1998.
Office of the Dean, University of Washington, 2001.
University of Washington Travel Award, 24th Int'l Ethological Congress, Honolulu, Hawaii, 1995
National Science Foundation Grant from the Division of Psychobiology, "Environmental stress, aggression, corticosterone, and metabolism in birds", 1984- 1986
Obesity Core Center Feasibility Grant, St. Luke's-Roosevelt Hospital, "Hormonal regulation of migratory obesity", 1985-1986.
National Science Foundation Grant from the Division of IOS, “Physiological mechanisms of migratory traits”, 2009-2011
National Science Foundation Grant from the Division of ARC, “Physiological mechanisms of migratory traits”, 2012 - 2017..
"Thinkers in Residence Fellowship" Deakin University, Waurn Ponds, Vic, Australia, July - Dec, 2015

Department and Center Affiliations

Neurobiology, Physiology and Behavior


Board of Directors of Cooper Ornithological Society.
Society for Integrative and Comparative Biology (SICB) - Chair of Division of Animal Behavior; Comparative Endocrinology, Committee of the International Ornithologists Union,
Animal Behavior Society
Ornithological Societies of North America

CBS Grad Group Affiliations

Animal Behavior

Specialties / Focus

Animal Behavior
  • Physiology and Behavior

Graduate Groups not Housed in CBS

Animal Biology Graduate Group


Ramenofsky Lab
  • Zoltan Nemeth (Former Postdoctoral Fellow 2010 - 2014), Jonathan Perez (Graduate Student)
  • Simeon Lisovski (Postdoctoral Fellow 2016- 2017)

Field Sites

UC Davis Field House
West Putah Creek
Center for land-based Learning
Bodega Marine Laboratories Reserve
University of Alaska Fairbanks
Toolik Lake Field Station, Institue of Arctic Biology, UAF website

Teaching Interests

Animal migrations, Comparative Vertebrate Reproduction


Physiolog NPB 139 Frontiers in Physiology (Winter 2011)
Seminar PHE 198 History of the Olympics (Spring Quarter)


8/11/2017 2:05:46 PM
  • Ramenofsky, M., Faulkner D. J., and Ireland, C. (1974) Effect of juvenile hormone on cirriped metamorphosis, Biochem. Biophys. Comm. 110,172-178.
  • Ramenofsky, M. (1984) Agonistic behaviour and endogenous plasma hormones in adult Japanese quail. Anim. Behav. 32,698-708.
  • Ramenofsky, M. (1985) Acute changes in plasma steroids and agonistic behavior in male Japanese quail. Gen. Comp. Endocrinol. 60,116-128.

  • Wingfield, J.C., Ball, G., Dufty, A.M., Hegner, R., and Ramenofsky, M. (1986) Testosterone and aggression in birds. Am. Sci. 75, 602-608
  • Wingfield, J.C., and Ramenofsky, M. (1984) Testosterone and aggressive behaviour during the reproductive cycle of male birds, in Proceedings in Life Sciences, Neurobiology/Current Comparative Approaches. (Edited by R. Gilles and J. Balthazart), pp. 92-104, Springer Verlag, Heidelberg.
  • Schwabl, H., Ramenofsky, M., Schwabl-Benzinger, I., Farner, D.S., and Wingfield, J.C. (1988) Social status, circulating levels of hormones, and competition for food in winter flocks of the White-throated sparrow. Behaviour 107,107-121.
  • Ramenofsky, M. (1990) Fat storage and fat metabolism in relation to bird migration, in Physiology and Ecophysiological Aspects of Bird Migration, (ed. E. Gwinner), Springer Verlag, Heidelberg pp 214-231.
  • Gray, J.M., Yarian, D., Ramenofsky, M. (1990) Corticosterone, foraging behavior, and metabolism in Dark-eyed juncos, Junco hyemalis. General and Comp. Endocrinol. 79:375-384.
  • Savard, R., Ramenofsky, M.,Greenwood, M.R.C. (1991) A north-temperate migratory bird: a model for the fate of lipids during exercise of long duration. Can. J. Physiol. Pharmacol. 69:1443-1447.
  • Ramenofsky, M., Gray, J., and Johnson, R. (1992) Behavioral and physiological adjustments of birds living in winter flocks. Ornis Scand. 23:371-380.
  • Hahn, T.P., Swingle, J., Wingfield, J.C., and Ramenofsky, M. (1992) Adjustments of the prebasic molt schedules in birds. Ornis Scand. 23:314-321.
  • Rodgers, C.M., Ramenofsky, M., Ketterson, E.D., Nolan, V., and Wingfield, J.C. (1993) Relationship of corticosterone and adrenal mass with winter weather conditions in natural populations of the Dark-eyed Junco (Junco hyemalis). AUK 110:279-285.
  • Boswell, T., Richardson, R.D., Seeley, R.J., Ramenofsky, M., Wingfield, J.C., Friedman, M.I., and Woods, S.C. (1995) Regulation of food intake by metabolic fuels in White-crowned Sparrows. Am. J. Physiol. 269 (Regulatory Integrative Comp. Physiol 38):R1462-1468.
  • Boxwell, J., Ayson, P., and Ramenofsky, M. (1995) Growth and metabolic parameters of undernourished lactating rats. Physiol Behav. 57(30): 469-475.
  • Ramenofsky, M., Piersma, T., and Jukema, J. (1995) Plasma corticosterone in Bar-tailed Godwits at a major stop-over site during spring migration. Condor 97:580-584.
  • Piersma, T., Jukema, J., and Ramenofsky, M. (1997) Onvoorspelbaarheid, lange afstandstrek, en de stress-hormonen van Rosse Grutto’s, (Unpredictability, long-distance migration and the stree-hormones of Bar-tailed Godwits), Limosa 70:34-35.
  • Romero, L.M., Ramenofsky, M., and Wingfield, J.C. (1997) Season and migration alters the corticosterone response to capture and handling in an arctic migrant, the White-crowned Sparrow ( Zonotrichia leucophrys gambelii). Comp. Biochem and Physiol. 116C:171-177.
  • Boswell, T., Lehman, T.L., and Ramenofsky, M. (1997) Effects of plasma glucose manipulations on food intake in white-crowned sparrows. Comp. Biochem. Physiol. 118A:721-726
  • Wingfield, J.C., and Ramenofsky, M. (1997) Corticosterone and facultative dispersal in unpredictable events. Ardea. 85:155-166.
  • Hilgarth, N., Ramenofsky, M., and Wingfield, J.C. (1997) Testosterone and sexual selection. Behavioral Ecology 8(1):108-109.
  • Meddle, S.L., King, V.M., Follett, B.K., Wingfield, J.C., Ramenofsky, M., Foidart, A., and Balthazart, J. (1997) Copulation activities of Fos-like immunoreactivity in the male quail forebrain. Behav. Brain Res. 85: 143-159
  • Wingfield, J.C., Maney, D.L., Bruener, C.W., Jacobs, J.D., Lynn, S., Ramenofsky, M., and Richardson, R.D. (1998) Ecological bases of hormone-behavior interactions; the “emergency life history stage”. Amer. Zool. 38:191-206.
  • Piersma, T. and Ramenofsky, M. (1998) Long-term decreases of corticosterone in captive migrant shorebirds that maintain seasonal mass and moult cycles. J Avian Biology 29:97-104.
  • Ramenofsky, M., Savard, R., and Greenwood, M.R.C. (1999) Seasonal and diel transitions in physiology and behavior in the migratory dark-eyed junco. Comp Biochem Physiol A 122:385-397.
  • Meddle SL, Foidart A, Wingfield JC, Ramenofsky,and M, Balthazart (1999) Effects of sexual interactions with a male on fos-like immunoreactivity in the female quail brain. J Neuroendocrinol 11(10):771-84
  • Wingfield, J.C., Jacobs, J.D., Soma, K., Maney, D.L., Hunt, K., Wisti-Peterson., D., Meddle, S., Ramenofsky, M., and Sullivan, K. (1999) Testosterone, aggression, and communication: ecological bases of endocrine phenomena. In “The biology of communication” (M. Konishi and M. Hauser, eds.) M.I.T. Press, Cambridge
  • Wingfield, J.C., and Ramenofsky, M. (1999) Hormones and behavioral ecology of stress. In “Stress Phyisology” (P. Balm, ed.), Sheffield University Press, Sheffield, U.K.pp 1-51.
  • Hiebert, S.M., Ramenofsky, M., Salvante, K, Wingfield, J.C., and Gass, C.L. (2000) Noninvasive methods for measuring and manipulating corticosterone in hummingbirds, Gen Comp Endocrinol, 120:235-247
  • Piersma, T., Reneerkens, J., and Ramenofsky, M. (2000) Baseline corticosterone peaks in shorebirds with maximal energy stores for migration: a general preparatory mechanism for rapid behavioral and metabolic transitions? Gen Comp Endocrinol 120:118-126
  • Hiebert, S.M.,Salvante, K., Ramenofsky, M., and Wingfield, J.C. (2000) Corticosterone and nocturnal torpor in the rufous hummingbirds (Selasphorus rufus), Gen Comp Endocrinol, 120:220-234.
  • Reneerkens, J., Morrison, R.I., Ramenofsky, M., Piersma, T., Wingfield, J.C. (2002) Baseline and stress-induced levels of corticosterone after a long migratory flight, preparation for breeding and during incubation of a high arctic breeding shorebird (Red Knot, Calidris canutus). Physiol. Biochem. Zool. 75(2):200-208.
  • Landys-Ciannelli, M.M., Ramenofsky, M. Piersma, T., Jukema, J., Wingfield, J.C. (2002) Baseline and stress-induced plasma corticosterone during long-distance migration in the bar-tailed godwit, Limosa lapponica. Physiol. Biochem. Zool. 75:101-110.
  • Reneerkens, J., Ramenofsky, M., Piersma, T. (2002) An experimental test of the relationship between temporal variability of feeding opportunities and baseline levels of corticosterone in a shorebird. J. Exp. Zool. 293:81-88.
  • Landys, M.M., Ramenofsky, M., Piersma, T., Jukema, J., Castricum Ringing Group, Wingfield, J.C. (2002) Baseline and stress-induced plasma corticosterone during long-distance migration in the bar-tailed godwit, Limosa lapponica. Physiol. Biochem. Zool. 75(1):101-110,.
  • Ramenofsky, M., R. Agatsuma, M. Barga, R. Cameron, J. Harm, M. Landys, T. Ramfar (2003) Migratory behavior: new insights from captive studies. In Berhold P, Gwinner, E, Sonnenschein, E (eds), Avian Migration, Springer, Berlin, pp 97-112
  • Landys, M.M., J.C. Wingfield, and M. Ramenofsky (2004) Plasma corticosterone increases during migratory restlessness in the captive white-crowned sparrow Zonotrichia leucophrys gambelii, Hormones and Behavior 46:574-581.
  • Landys, M.M., M. Ramenofsky, Guglielmo, C.G.,and J.C. Wingfield. (2004) The low-affinity glucocorticoid receptor regulates feeding and lipid breakdown in the migratory white-crowned sparrow Zonotrichia leucophrys gambelii. Journal of Experimental Biology, 207:143-154.
  • Landys, M.M., T. Piersma, M. Ramenofsky, and J.C. Wingfield (2004) The low-affinity glucocorticoid receptor regulates behavior and energy metabolism in the migratory red knot Calidris canutus islandica. Physiological and Biochemical Zoology. 77(4):658-668
  • Landys, M.M., Piersma, T., Guglielmo, C.G., Jukema, J., M. Ramenofsky, & J.C. Wingfield, (2005) Metabolic profile of long-distance migratory flight and stopover in a shorebird. Proc. Royal Society, 272:295-302.
  • Agatsuma R and Ramenofsky M (2006) Migratory behaviour of captive white-crowned sparrows, Zonotrichia leucophrys gambelii differs during autumn and fall migration, Behaviour, 143(10):1219-1240.
  • Ramenofsky, M. and Wingfield, J.C. (2006). Behavioral and physiological conflicts in migrants: the transition between migration and breeding, J of Ornithology 147 (2):135-145.
  • Landys, M., Ramenofsky M., Wingfield JC (2006) Actions of glucocorticoids at a seasonal baseline as compared to stress-related levels in the regulation of periodic life processes Gen Comp Endocrinol.148:132-149.
  • Bonier, F, Martin, PR, Jensen, JP, Butler, LK, Ramenofsky, M, and Wingfield, JC. (2007) Pre-migratory life history stages of juvenile arctic birds: costs, constraints and tradeoffs. Ecology 88(11):2729-2735.
  • Ramenofsky, M and Wingfield, J.C. (2007) Regulation of migration. Bioscience 57(2): 135-143.
  • Lynn, Sharon, E., Egar, Joseph M., Walker, Brian G., Sperry, Todd S., and Ramenofsky, Marilyn. (2007) Fish on Prozac: a simple, noninvasive physiology laboratory investigating the mechanisms of aggressive behavior in Betta splendens. Advances in Physiology Education 31:358-363.
  • Thorup, Kasper, Bisson, Isabelle-A., Bowlin, Melissa S., Holland, Richard A., Wingfield, John C., Ramenofsky, Marilyn, and Wikelski, Martin. (2007) Evidence for a navigational map stretching across the continental U.S. in a migratory songbird. PNAS 104:18115-18119.
  • Coverdill, A.J, Bentley, G.E. and Ramenofsky, M. (2008) Circadian and masking control of migratory restlessness in Gambel’s White-crowend sparrow (Zonotrichia leucophrys gambelii), Journal of Biological Rhythms 23(1):59-68.
  • Ramenofsky, M., Agatsuma, R., and Ramfar, T. (2008) Environmental conditions affect behavior of captive, migratory White-crowned Sparrows, Condor 110:658-671
  • Ramenofsky, M., Moffat and Bentley, G. (2008) Corticosterone and migratory behavior of captive White-crowned Sparrows, Proceedings 4th CPB Meeting, Maasai Mara, Kenya “Molecules to migration: The pressures of life” (Ed S. Morris & A. Vosloo). Medimond Publishing Co, Bologna, Italy. Pages 575-582.
  • Ramenofsky, M., Agatsuma, R., Ramfar, T. (2008) Environmental conditions affect behavior of captive migratory White-crowned Sparrows, Condor 110(4):658-671.
  • Ramenofsky, M., Moffat, J., Bentley, G.E. (2008) Corticosterone and migratory behaviour of captive White-crowned Sparrows, Proceedings of the 4th International ICA-CPB, "Pressures of Life: Molecules to Migration". pp. 575-582.
  • Ramenofsky M. (2010) Behavioral Endocrinology of Migration. In: Breed M.D.and Moore J., (eds.) Encyclopedia of Animal Behavior, volume 1, pp. 191-199 Oxford: Academic Press.
  • Bowlin, M.S., Bisson, I-A., Shamoun-Baranes, J., Reichard, J.D., Sapir, Nir, Marra, P.P., Kunz, T.H., Wilcove, D.S., Hedenström, A., Guglielmo, C.G., Åkesson, S., Ramenofsky, M., Wikelski, M. (2010) Grand Challenges in migration biology, Int. Comp. Biol.50(3): 261-279.
  • Blumstein, D.T., Ebensperger, L.A., Hayes, L.D., Vasquez, R.A., Ahern, T.H, Burger, J. R., Dolezal, A.G., Dosmann, A., Gonzales-Mariscal, G., Harris, B., Herrara, E.A., Lacey, E.A., Mateo, J., McGraw, L.A., Olazabal, D., Ramenofsky, M., Rubenstein, D.R., Sakhai, S.A., Saltzman, W., Sainz-Borgo, C., Soto-Gamboa, M., Stewart, M.L., Wey, T.W., Wingfield, J.C., Young, L.J. (2010) Towards an integrative understanding of social behavior: new models and new opportunities. Front. Behav. Neurosci. 4:34. doi
  • Kumar, V., Wingfield, J.C., Dawson, A., Ramenofsky, M., Rani, S., Bartell, P. (2010). Biological clocks and regulation of seasonal reproduction and migration in birds. Physiol. Biochem. Zool. 83(5):827-35.
  • Ramenofsky, M. (2010). Hormones in migration and reproductive cycles of birds. In Hormones and Reproduction in Vertebrates, D. Norris and K. H. Lopez. (Eds.), Academic Press. Vol 8. pp. 205 – 236.
  • Sih, A, Stamps, J., Yang, L.H., McElreath, R., Ramenofsky, M. (2010) Grand challenges: behavior as a key component of integrative biology in a human-altered world, Integrative and Comparative Biology 50 (6): 934-944.
  • Coverdill, A.J., Clark, A.D, Wingfield, J.C. & Ramenofsky, M. (2011) Examination of nocturnal activity and behaviour in resident white-crowned sparrows (Zonotrichia leucophrys nuttalli). Behaviour 148:859-876. DOI:10.1163/000579511X579038
  • Wingfield, J.C. & Ramenofsky, M. 2011. Hormone-behavioral interrelationships of birds in response to weather. In H. J. Brockmann, T. J. Roper, M. Naguib, J. C. Mitani, Leigh W. Simmons, (eds), Advances in the study of behavior 43: 93-188. DOI: 10.1016/B978-0-12-380896-7.00003-4
  • Ramenofsky, M. (2011) Reconsidering the role of photoperiod in relation to effects of precipitation and food availability on spring departure of a migratory bird. Proc R Soc B 279: 15-16. doi: 10.1098/rspb.1591
  • Wingfeld, J.C., Ramenofsky, M. 2012. Hormone-behavior interrelationship of birds in response to weather. Advances in the Study of behavior, 43:93–188.
  • Wingfield, J.C., Ramenofsky, M. 2012. Hormone-behavior interrelationship of birds in response to weather. Advances in the Study of behavior, 43:93–188.

  • Ramenofsky, M., Cornelius, J.M., Helm, B. (2012) Physiological and behavioural responses of migrants to environmental cues, J. Ornithology, DOI: 10.1007/s10336-012-0817-3
  • Cornelius, J.M., T. Boswell, S. Jenni-Eiermann, C.W. Breuner and M. Ramenofsky (2013) Contributions of endocrinology to the migration life history of birds. General and Comparative Endocrinology, 190 (1):47-60,

  • Ramenofsky, M. (2013) Physiological mechanisms of migration, Guest Editorial, Current Zoology 59(3): 347-348

    Wang, G., Ramenofsky, M., Wingfield, J.C. (2013) Apparent dissociation of photoperiodic time measurement between vernal migration and breeding under dim green light conditions in Gambel’s White-crowned Sparrow, Zonotrichia leucophrys gambelii, Current Zoology. 59(3): 349 -359.

    Ramenofsky, M, Németh, Z. (2014) Regulatory mechanisms for the development of the migratory phenotype: roles for photoperiod and the gonad. Hormones and Behavior 66:148–158,

    Németh, Z., Adams, R.H., Ramenofsky, M. (2015) Androgens increase persistence but do not affect neophobia in a problem-solving context in a songbird, Ethology DOI: 10.1111/eth.12353

  • Krause, J., Németh, Z., Perez, J., Chmura, H.E., Ramenofsky, M., Wingfield, J.C. 2015. Annual hematocrit profiles in two sub-species of White-crowned Sparrows: a migrant and resident comparison, Physiol Biochem Zoology, 89(1):51-60.

    Pérez, J.H., Furlow, J.D., Wingfield, J.C., Ramenofsky, M. 2016. Regulation of vernal migration in Gambel’s white-crowned sparrows: role of thyroxine and triiodothyronine.  Hormones and Behavior 84:50–56.

    Ramenofsky, M., Wingfield, J.C. 2016. Regulation of complex behavioural transitions: migration to breeding. Animal Behaviour.

    Velton, B.P.,Welch, K.C. Jr., Ramenofsky, M. 2016 Altered expression of pectoral myosin heavy chain isoforms corresponds to migration status in thewhite-crowned sparrow (Zonotrichia leucophrys gambelii). R. Soc. open sci. 3: 160775.

    Wingfield, J.C. and Ramenofsky, M. (2016) Control of the migration life history in birds, in Migration, Edited by A. Urano and H. Ando, published by Japan Society for Comparative Endocrinology, Tokyo, Chap 7, Pages 101 – 142.

    Watts, H.E., Robart, A.R., Chopra, J., Asinas, C., Hahn, T.P., Ramenofsky, M. 2016. Seasonal expression of migratory behavior in a facultative migrant, the pine siskin, Behavioral Ecology and Sociobiology, DOI 10.1007/s00265-016-2248-2

    Ramenofsky, M., Campion, A.W., Pérez, J.H., Krause, J.S., Németh, Z. 2017 Behavioral and physiological traits of migrant and resident White-crowned Sparrows: a common garden approach, Journal of Experimental Biology, 220, 1330-1340 doi:10.1242/jeb.148171

  • Franson, K., Nemeth, Z., Ramenofsky, M. WIngfield, J.C., Buchanan, K. 2017 Inter-laboratory variation in corticosterone measurement: implications for comparative ecological and evolutionary studies, Methods Ecological and Evolutionary Studies: DOI: 10.1016/j.ygcen.2017.07.027

  • Watts, H.E., Cornelius, J.M., Fudickar, A.M., Perez, J.H., Ramenofsky, M. 2017 Understanding varation in migratory movements: a mechanistic approach, General and Comparative Endocrinology