Revised July, 2021

CURRICULUM VITAE

GARY J. ROSE Professor 317 South Biology 801-581-3974 [email protected] Citizenship: U.S.

Degrees:

B.A. University of California, San Diego, 1977 - Biology M.S. Cornell University, 1982 - Neurobiology and Behavior Ph.D. Cornell University, 1983 - Neurobiology and Behavior

Professional Experience:

1977-1983 Graduate teaching and research, Cornell University. Ph.D. Advisor: Professor Robert Capranica 1983-1985 Postdoctoral Fellow, Scripps Institution of Oceanography. Advisor: Professor Walter Heiligenberg 1985-1988 Postgraduate researcher, Scripps Institution of Oceanography 1988-1993 Assistant Professor, Department of Biology, University of Utah 1993-1999 Associate Professor, Department of Biology, University of Utah 1999- Professor, Department of Biology, University of Utah

Honors:

1977 Honors Thesis (University California, San Diego) 1984 Honorable Mention, Lindsley Prize 1986 International Society for Neuroethology Prize 1989 Research Fellowship (Sloan Foundation) ($25,000)

Research Interests:

Neural basis of electrosensory behaviors in weakly electric fish Acoustic communication in anurans Neural mechanisms of audition Behavioral and physiological determinants of sex and coloration in marine wrasses Song learning in songbirds

Grants, Scholarships and Awards:

Student Organized Science Project (from NSF) at University of California, San Diego Traineeship on NIH Training Grant, and NIMH Training Grant Postdoctoral Fellowship (NIH), 9/83 - 8/85 New Investigator Award (NIH), 9/85 - 8/88 – $136,735 Research Award (NSF), 8/89 - 2/93 – $270,313 Research Award (NSF) 4/92 - 3/95 – $108,300 Research Award (NIMH) 11/92 - 10/94 – $140,499 Research Award (NSF) 2/95 - 1/98 $289,573 Rose

Research Award (NSF) 6/98 - 2/03 – $307,817 Research Award - National Org. for Hearing Res. 1998-99 ($5,000) Shannon Award (NIH) 1999-00 ($50,000) Research Award (NIH) 1/00-2/04 $464,468 Seed Grant (U of U) 4/03 - 3/04 $30,500 Research Award (NIH) 6/1/04 - 5/31/10 $1,360,450 Research Award (NIH) 7/1/09 - 6/30/10 $375,000 Research Award (NIH) 7/1/10 - 6/30/16 $1,875,000 U of U Seed Grant 6/15 - 5/16 $35,000 Research Award (NIH, R56) 7/1/18-6/30/19 $300,000 Research Award (NIH, R01) 12/1/18-11/30/23 $1,620,300 Research Award (NSF, co-PI, Dr. Bee): 5/15/21- 5/14/24 $650,000. Research Award (NIH, R03) 5/1/21-4/31/23 $150,000. Research Award (NSF, Mentor for Dr. Emily Lemmon) 8/1/21-7/31/24, $74,500 (Utah)

Pending Funding:

Research application (NIH, R01) submitted 6/5/2021

Membership in Professional Societies: Society for Neuroscience Acoustical Society of America International Society for Neuroethology

Courses Taught:

Cornell University: Introductory Neurobiology and Behavior, Cellular and Molecular Neurobiology, Sensory Physiology.

University of Utah: (103) Princ. Biology; (317) Intro. Neuro., w/Yoshikami and Olivera; (317) Intro. Neuro.; (524) Neural mechanisms of behavior. Currently, Behavioral Neurobiology, (3330) 100%, Critical Analysis (7406) (100%).

Woods Hole: Summers of 1996 & 1997 Neural Systems and Behavior Course.

Committees:

Search, Insect Physiology Search, Animal physiology Graduate Admissions Scholarship (Flowers) Scholarship (Hughes) Executive Committee New Building Committee Search, Neurobiology Search, Animal Physiology (Chair) Internal Review (Bioengineering) Internal Review (Neurobiology and Anatomy)

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Undergraduate Scholarship Committee, Biology (Current) Academic Senate (2004-2005; 2011-2015) Admissions Committee, Neuroscience Program (2006- 2010, 2011- 2013) College Committee on Admission Standards and Degree Programs (Chair,18), (2014-2018)

Invited Presentations: (past 15 years)

University of Illinois, Chicago, Seminar Speaker International Brain Research Organization meeting, Prague: Co-chair and speaker, Symposium on Temporal Processing in Sensory Systems. University of Chicago, Neuroethology Conference: Speaker University of Virginia, Seminar speaker Wesleyan University, Seminar speaker UCLA Conference on Neural Control of Behavior: Speaker University of Texas, Austin: Seminar speaker Cornell University: Seminar speaker University of Utah, Neuroscience Symposium: Speaker University of Arizona: Seminar speaker University of Texas, San Antonio: Seminar speaker Hayama Institute, Japan: Symposium speaker Neuroethology meeting, Spain: Symposium speaker Workshop on Auditory Processing, Cody WY: Speaker University of Minnesota: Seminar speaker Royal Society conference, London: Speaker

Training:

Post Docs: (last 10 years)

Rishi Alluri 2018-present Jessica Hanson 2014- 2016 Chris Edwards 2001-2009 Chris Leary 2005-2009 Elizabeth Wilkinson 2009-2010

Grad Students: (last 5 years)

Jeremy Wilkerson Stephen Odom Kyphuong Luong Rishi Alluri Anwesha Mukhopadhyay

Undergraduate Students: (last 5 years) Sarah Campbell Rickey Bell Dina Krikova Caleb Herrick

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Current: Maddie Bernardo Anthony Khong Michael Lindstrom Jaime McDowell Alexandra Strich (ACCESS)

Publications:

Geyer, M.A., L.R. Petersen, G.J. Rose, D.D. Horwitt, R.K. Light, L.M. Adams, J.A. Zook, R.L. Hawkins, and A.J. Mandell (1978) The effects of Lysergic acid diethylamide and mescaline-derived hallucinogens on sensory-integrative function: Tactile startle. J. of Pharm. and Exp. Therapeutics 207:837-847.

Geyer, M.A., R.K. Light, G.J. Rose, L.R. Petersen, D.D. Horwitt, L.M. Adams, and R.L. Hawkins (1979) A characteristic effect of hallucinogens on investigatory responding of rats. Psychopharmacol. 65:35-40.

Geyer, M.A., G.J. Rose and L.R. Petersen (1979) Mescaline increases startle responding equally in normal and raphe-lesioned rats. Pharm. Biochem. and Behav. 10:293-298.

Geyer, M.A., L.R. Petersen and G.J. Rose (1980) Effects of serotonergic lesions on investigatory responding by rats in a hole board. Behavioral and Neural Biology 30:160- 177.

Rose, G.J. and R.R. Capranica (1983) Temporal processing in the central auditory system of the leopard frog (Rana pipiens). Science 219:1087-1089.

Rose, G.J. and R.R. Capranica (1984) Processing amplitude-modulated sounds by the auditory midbrain of two species of toads: Matched temporal filters. J. Comp. Physiol. 154:211- 219.

Rose, G.J., and W. Wilczynski (1984) The anuran superficial reticular nucleus: Evidence for homology with the nucleus of the lateral lemniscus. Brain Res. 304:170-172.

Rose, G.J. and R.R. Capranica (1985) Sensitivity to amplitude modulated sounds in the anuran auditory system. J. Neurophysiol. 53:446-465.

Heiligenberg, W. and G.J. Rose (1985) Phase and amplitude computations in an electric fish's midbrain. J. Neuroscience 5:515-531.

Rose, G.J. and W. Heiligenberg (1985) Processing phase and amplitude information by the midbrain of an electric fish: Relationship between structure and function. J. Neuroscience 5:2269-2280.

Rose, G.J. and W. Heiligenberg (1985) Temporal hyperacuity in the electric sense of fish. Nature 319:178-180.

Brenowitz, E.A., G.J. Rose, and R.R. Capranica (1985) Neural correlates of temperature coupling in the vocal communication system of the Gray treefrog, Hyla versicolor. Brain Research 359:354-367.

Rose, G.J. (1986) A temporal processing mechanism for all species? Brain Behav. and Evol. 28:134-144.

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Rose, G.J. and W. Heiligenberg (1986a) Neural coding of difference frequencies in the midbrain of the electric fish Eigenmannia: Reading the sense of rotation in an amplitude- phase plane. J. Comp. Physiol. A158:613-624.

Rose, G. and W. Heiligenberg (1986b) Limits of phase and amplitude sensitivity in the torus semicircularis of Eigenmannia. J. Comp. Physiol. A159:813-822.

Rose, G.J., E.A. Brenowitz, and R.R. Capranica (1986) Species specificity and temperature dependency of temporal processing by the auditory midbrain of two species of treefrogs. J. Comp. Physiol. 157:763-769.

Heiligenberg, W. and G.J. Rose (1986) Gating of sensory information: Joint computations of phase and amplitude data in the midbrain of the electric fish Eigenmannia. J. Comp. Physiol. 159:311-324.

Carr, C.E., W. Heiligenberg, and G.J. Rose (1986) A time-comparison circuit in the electric fish midbrain. I. Behavior and Physiology. J. Neuroscience 6:107-119.

Heiligenberg, W. and G.J. Rose (1987) The optic tectum of the gymnotiform fish, Eigenmannia: Labeling of physiologically identified cells. Neuroscience 22:331-340.

Rose, G., C. Keller and W. Heiligenberg (1987) 'Ancestral' neural mechanisms of electrolocation suggest a substrate for the evolution of the jamming avoidance response. J. Comp. Physiol. 160:491-500.

Rose, G.J., R. Zelick and A.S. Rand (1988) Auditory processing of temporal information in a neotropical frog is independent of signal intensity. Ethology 77:330-336.

Rose, G.J., M. Kawasaki and W. Heiligenberg (1988) 'Recognition units' at the top of a neuronal hierarchy? Prepacemaker neurons in Eigenmannia code the sign of frequency differences unambiguously. J. Comp. Physiol. 162:759-772.

Kawasaki, M., L. Maler, G. Rose, and W. Heiligenberg (1988) Anatomical and functional organization of the prepacemaker nucleus in Gymnotiform electric fish: The accommodation of two behaviors in one nucleus. J. Comp. Neurol. 276:113-131.

Kawasaki, M., G. Rose and W. Heiligenberg (1988) Temporal hyperacuity in single neurons of electric fish. Nature 336:173-176.

Seger, J., W. Stubblefield, H.S. Horn, and G.J. Rose (1990) How to bug a bug. Sphecos 19:24- 26.

Zelick, R., G.J. Rose and A. S. Rand (1991) Differential response to frequency modulation rate and direction by the neotropical frog, Physalaemus pustulosus. Animal Behavior, 42: 413- 421.

Rose, G.J. and J.G. Canfield (1991) Discrimination of the sign of frequency differences by Sternopygus, an electric fish without a Jamming Avoidance Response. J. Comp. Physiol. 168: 461-467.

Rose, G.J. and E.A. Brenowitz (1991). Aggressive thresholds of male Pacific treefrogs for advertisement calls vary with amplitude of neighbor's calls. Ethology. 89: 244-252.

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Rose, G.J., and S.J. Call (1992). Differential distribution of ampullary and tuberous processing in the torus semicircularis of Eigenmannia. J. Comp. Physiol. 170: 253-261.

Rose, G. J., and S. J. Call. (1992) Evidence for the role of dendritic spines in the temporal filtering properties of neurons: The decoding problem and beyond. Proc. Natl. Acad. Sci. 89: 9662-9665.

Canfield J. G., and G. J. Rose. (1993) Activation of mauthner neurons during prey capture. J. Comp. Physiol. 172: 611-618.

Rose, G. J., and S. J. Call. (1993) Temporal filtering properties of neurons in the midbrain of an electric fish: Implications for the function of dendritic spines. J. Neurosci. 13(3): 1178-1189.

Rose, G. J., and J. G. Canfield. (1993) Longitudinal tracking responses of the weakly electric fish, Sternopygus. J. Comp. Physiol. 171: 791-798.

Canfield J. G., and G. J. Rose. (1993) Electrosensory modulation of escape responses. J. Comp. Physiol. 173: 463-474.

Rose, G. J., and J. G. Canfield. (1993) Longitudinal tracking responses of Eigenmannia and Sternopygus. J. Comp. Physiol. 173: 698-700.

Brenowitz E. A., and G. J. Rose. (1994) Behavioural plasticity mediates intermale aggression in choruses of the Pacific treefrog. Animal Behav. 47: 633-641.

Rose, G. J., N. Etter and T. B. Alder. (1994) Responses of electrosensory neurons in the torus semicircularis of Eigenmannia to complex beat stimuli: testing hypotheses of temporal filtering. J. Comp. Physiol. 175: 467-474.

Canfield, J. G., and G. J. Rose. (1996) Hierarchical sensory guidance of mauthner-mediated escape responses in goldfish (Carassius auratus) and cichlids (Haplochromis burtoni). Brain, Behav. and Evol. 48: 137-156.

Rose, G. J., and E. S. Fortune. (1996) New techniques for making whole-cell recordings from CNS neurons in vivo. Neurosci. Res. 26: 89-94.

Rose, G. J., and E. A. Brenowitz. (1997) Plasticity of aggressive thresholds in male Pacific treefrogs: Discrete accommodation to the encounter call. Animal Behav. 53: 353-361.

Fortune, E. S., and G. J. Rose. (1997) Temporal filtering properties of ampullary electrosensory neurons in the torus semicircularis of Eigenmannia: evolutionary and computational implications. Brain, Behav. and Evol. 49: 312-323.

Fortune, E. S. and G. J. Rose. (1997) Passive and active membrane properties contribute to the temporal filtering properties of midbrain neurons in vivo. J. Neurosci. 17(10): 3815- 3825.

Alder TB, Rose GJ (1998) Long-term integration in the anuran auditory system. Nature Neuroscience 1(6): 519-523.

Takizawa Y, Rose GJ, Kawasaki M. (1999) Resolving competing theories for control of the jamming avoidance response: The role of amplitude modulations in electric organ discharge decelerations. J. Exp. Biol. 202; 1377-1386.

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Brenowitz EA, Rose GJ (1999) Female choice and plasticity of male calling behavior in the Pacific treefrog. Animal Behavior 57: 1337-1342

Fortune ES, Rose GJ (1999) Frequency-dependent PSP depression contributes to low-pass temporal filtering in Eigenmannia. J. Neuroscience 19(17): 7629-7639.

Fortune E.S., Rose G.J. (1999) Mechanisms for generating temporal filters in the electrosensory system. J. Exp. Biol. 202: 1281-1289.

Fortune E.S., Rose G.J. (2000) Short-Term Synaptic Plasticity Contributes to the Temporal Filtering of Electrosensory Information. J. Neurosci. 20(18): 7122-7130.

Alder TB, Rose GJ (2000) Integration and recovery processes contribute to the temporal selectivity of neurons in the northern leopard frog, Rana pipiens. J. Comp. Physiol. 186: 923-937.

Fortune E.S., Rose G.J. (2001) Short-term synaptic plasticity as a temporal filter. TRENDS Neurosci. 24:381-385.

Rose GJ, Brenowitz EA, (2002), Pacific Tree Frogs Use Temporal Integration to Differentiate Advertisement from Encounter Calls. Animal Behavior 63:1183-1190.

Edwards CJ, Alder TB, Rose GJ (2002) Auditory midbrain neurons that count. Nature Neuroscience 5(10):934-936.

Fortune ES and Rose GJ (2003) Voltage-Gated Na+ Channels Enhance the Temporal Filtering Properties of Electrosensory Neurons in the Torus. J. Neurophysiol. 90:924-929.

Edwards CJ and Rose GJ (2003) Interval-integration underlies AM band-suppression selectivity in the anuran midbrain. J. Comp. Physiol. A. 189:907-914.

Rose, GJ, Goller, F, Gritton, HJ, Plamondon, SL, Baugh AT, Cooper BG (2004) Species- typical songs in white-crowned sparrows tutored with only phrase pairs. Nature 432:753- 758.

Edwards CJ, Alder TB, Rose GJ (2005) Pulse rise time but not duty cycle affects the temporal selectivity of neurons in the anuran midbrain that prefer slow AM rates. J. Neurophysiol. 93: 1336-1341.

Edwards CJ, Leary CJ, Rose GJ (2007) Counting on inhibition and rate-dependent excitation in the auditory system. J. Neurosci. 27:13384-13392.

Leary CJ, Edwards CJ, Rose GJ (2008) Midbrain auditory neurons integrate excitation and inhibition to generate duration selectivity: an, in vivo, whole-cell patch study in anurans. J. Neurosci. 28:5481-5493.

Plamondon SL, Goller F, Rose GJ (2008) Tutor model syntax influences the syntactical and phonological structure of crystallized songs of white-crowned sparrows. Anim. Behav. 76(6):1815-1827.

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Edwards CJ, Leary CJ, Rose GJ (2008) Mechanisms of long-interval selectivity in midbrain auditory neurons: roles of excitation, inhibition and plasticity. J. Neurophysiol. 100:3407- 3416. Plamondon SL, Rose GJ, Goller F (2010) Roles of syntax information in directing song development in white-crowned sparrows (Zonotrichia leucophrys) J. Comp. Psychol. 124(2):117-132.

Rose GJ, Leary CJ, Edwards CJ (2011) Interval-counting neurons in the anuran auditory midbrain: factors underlying diversity of interval tuning. J. Comp. Physiol. 197:97-108.

Rose GJ, Alluri RK, Vasquez-Opazo GA, Odom SE, Graham JA, Leary CJ (2013) Combining pharmacology and whole-cell patch recording from CNS neurons, in vivo. J. Neurosci. Methods 213:99-104.

Rose GJ (2014) Time computations in anuran auditory systems. Frontiers in Physiology 5(206):1-7.

Naud R, Houtman D, Rose GJ, Longtin A (2014) Modeling sound pulse counting in inferior colliculus. BMC Neuroscience. 15:p113.

Rose GJ, Hanson JL, Leary CJ, Graham JA, Alluri RK, Vasquez-Opazo GA (2015) Species- specificity of temporal processing in the auditory midbrain of gray treefrogs: Interval- counting neurons. J. Comp. Physiol. 201:485-503.

Naud R, Houtman D, Rose GJ, Longtin A (2015) Counting on dis-inhibition: a circuit motif for interval counting and selectivity in the anuran auditory system. J. Neurophysiol. 114(4): 2804-2815.

Hanson JL, Rose GJ, Leary CJ, Graham JA, Alluri RK, Vasquez-Opazo GA (2016) Species- specificity of temporal processing in the auditory midbrain of gray treefrogs: Long-interval neurons. J. Comp. Physiol. 202: 67-79.

Alluri RK, Rose GJ, Hanson JL, Leary CJ, Graham JA, Vasquez-Opazo GA, Wilkerson J (2016) Delayed, phasic excitation and sustained inhibition underlie neural selectivity for short-duration sounds. PNAS 113(13): 1927-1935.

Price SM, Kyphuong Luong, Bell R, Rose GJ (2018) Latency of facultative expression of male-typical courtship behaviour in female bluehead wrasses: The ‘priming and gating’ hypothesis J. Exp. Biol. 221:1-8.

Alluri RK, Rose GJ, Leary CJ, Palaparthi A, Hanson JL, Graham JA, Vasquez-Opazo GA (2021) How auditory selectivity for sound timing arises: multiple mechanisms for decoding long-intervals. Prog. In Neurobiology 199: 101962.

Rose GJ, Alluri RK, Hanson JL, Leary CJ, Graham JA, Vasquez-Opazo GA (in prep.) Mechanisms of short-interval counting and selectivity in the auditory midbrain of anurans: role of dynamic inhibition.

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Review Articles & Chapters:

Capranica, R.R. and G.J. Rose (1983) Frequency and temporal processing in the auditory system of anurans. In: Neuroethology and Behavioral Physiology, F. Huber and H. Markl (Eds.), Springer-Verlag, Berlin, Heidelberg: 136-152.

Capranica, R.R., G.J. Rose and E.A. Brenowitz (1985) Time resolution in the auditory systems of anurans. In: Time Resolution in Auditory Systems, A. Michelsen (Ed.), Springer- Verlag, Berlin, Heidelberg: 58-72.

Heiligenberg, W. and G.J. Rose (1985) Neural correlates of the jamming avoidance response (JAR) in the weakly electric fish Eigenmannia. Trends in Neuroscience 5:442-449.

Rose, G. J. (1996). Representation of Temporal Patterns of Signal Amplitude in the Anuran Auditory System and Electrosensory System. In: Neural Representation of Temporal Patterns (ed. E. Covey) pp 1-24.

Rose GJ, Fortune ES (1999) Mechanisms for generating temporal filters in the electrosensory system. J. Exp. Biol. 202, 1281-1289.

Brenowitz EA, Rose GJ, Alder TB (2001) The neuroethology of Acoustic Communication in Pacific treefrogs. In: Recent advances in anuran communication. MJ Ryan (ed.), Smithsonian Institution Press.

Fortune ES, Rose GJ (2002) Roles for short-term synaptic plasticity in behavior. J. Physiology – Paris 96:539-545

Rose GJ (2004) Insights into neural mechanisms and evolution of behaviour from electric fish. Nature Reviews Neuroscience 5, 1-9

Bass AH, Rose GJ, Pritz MB (2005) Auditory midbrain of fish, amphibians and reptiles. In: The Inferior Colliculus, (Eds. JA Winer and CE Shreiner), Springer Verlag, New York: 467-477.

Rose GJ, Gooler DM (2006) Function of the amphibian central auditory system. In : Hearing and Sound Communication in Amphibians. (Eds. P Narins , AS Feng, RR Fay and A Popper), Springer Verlag, New York.

Rose GJ (2014) Time computations in anuran auditory systems. Frontiers in Physiology 5(206):1-7.

Rose GJ (2018) The numerical abilities of anurans and their neural correlates: Insights from neuroethological studies of acoustic communication. Phil Trans B 373:20160512.

Rose GJ, Leary CJ, Bee MA (2020) Anuran auditory systems as models for understanding sensory processing and the evolution of communication. In: The Senses. Eds: B. Fritsch, B. Grothe.

Research Interests

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We study animal behavior at both 'proximate' and 'ultimate' levels. At the proximate level, we investigate how neural circuits in fish and anuran amphibians control natural behaviors. At the ultimate level, we study the adaptive significance and evolution of these behaviors. Our research methodology, therefore, ranges from neurophysiological analysis of single neuron function to behavioral studies in the lab and field. Behavioral studies allow us to generate testable hypotheses concerning neural control. Conversely, neurophysiological experiments provide clues as to the evolution of behaviors. Specific research programs are described below.

Neural Mechanisms of Audition in Anurans Acoustic communication plays an important role in the reproductive behavior of anuran amphibians (frogs and toads). Much of the information in these vocalizations is encoded in the temporal structure (e.g. pulse repetition rate). The anuran auditory system, therefore, is well suited for investigating how the temporal structure of sound is represented at various stages in the auditory nervous system. We are particularly interested in understanding the mechanisms that underlie transformations in these representations. For example, the periodic modulations in the amplitude of sound are coded in the peripheral auditory system by the periodic fluctuations in the discharge rate of these neurons. At the midbrain, however, this 'periodicity' coding is replaced by a 'temporal filter' coding scheme wherein individual neurons selectively respond to particular rates of amplitude modulation. The mechanisms that underlie this transformation are unknown. Recent neurophysiological and behavioral studies indicate that integration of time-dependent excitation and inhibition underlies selectivity of midbrain neurons for temporal features of communication signals. Recent efforts to focally apply pharmacological agents in conjunction with whole-cell recording, in vivo, promise to provide deep insights into the mechanisms that underlie temporal computations in the auditory system. Comparative studies of gray treefrogs are also underway. One project is aimed at investigating the neural basis of the Franssen effect (illusion). The Franssen effect (FE) is an illusion in which the listener incorrectly perceives sound as coming from a speaker that broadcasts a leading, short- duration sound pulse with a fast rise in amplitude and slow decay. However, most of the sound originates from a lagging speaker that produces a long-duration sound with a slow onset, followed by a quick decay. Thus, both spatial and temporal features of sound are critical in producing the FE. Illusions such as these are distortions in perception that provide a window into the workings of the auditory system and perception. We utilize the anuran auditory system and a constellation of techniques, including in vivo whole-cell recording, focal iontophoresis of pharmacological agents, and analytical methods for estimating stimulus-related changes in excitatory and inhibitory conductances to elucidate the neural basis of the FE. The anuran auditory system has unique advantages for investigating the neural mechanisms that underlie the FE. An important advantage relates to the goal of linking neural function to perception. For precedence-type effects that have been demonstrated in animals, subjects indicate whether sound originated from left or right loudspeakers; it is unknown whether they perceive the SR-LD stimulus coming from the loudspeaker that broadcasted the transient tone. To address this question of perception, we take advantage of the natural preference of a species of gray treefrogs (Hyla versicolor) for slow-rise long-duration (SR-LD) sound pulses that are representative of those used to elicit FE illusions; in choice tests, H. versicolor females strongly prefer SR-LD sound pulses over the fast-rise short-duration (FR-SD) pulses seen in the advertisement calls of its sister species, H. chrysoscelis. Although female H. versicolor generally do not approach a loudspeaker that broadcasts FR-SD sound pulses, they will do so if SR-LD pulses are presented from a spatially separate transducer. This result tells us that the subject actually perceives the long-duration sound

10 Rose pulses as coming from the loudspeaker broadcasting the FR-SD pulses. Neurophysiological studies of the anuran inferior colliculus (ICan), a homolog of mammalian IC, in H. versicolor have revealed neurons that show strong selectivity for slow-rise pulses; these cells respond strongly to SR-LD tones, but show little, if any, response to FR-SD sound pulses. These neurons, therefore, show selectivity appropriate for contributing to the FE and we plan to investigate whether their activity is modulated in accordance with presentation of FS stimuli; for example, broadcast of FR-SD pulses should augment responses to ipsilaterally presented SR-LD stimuli. Further, anurans respire cutaneously, making neurophysiological preparation highly-stable. Thus, H. versicolor are well suited for studying natural auditory perception and neural processing. Other studies are aimed at further elucidating mechanisms that underlie selectivity for temporal features of communication sounds, and how changes in temporal processing contribute to speciation; for example, we are interested in how polyploidy influences the temporal processing properties of midbrain neurons e.g., pulse shape selectivity. This work should provide insight into how changes in the functional organization of the central auditory system contribute to the evolution of new species.

Song Learning in Songbirds In collaboration with Franz Goller's lab, we have studied how songbirds learn their songs. Songbirds must hear song early in life in order to later develop a good copy of the song of their local dialect; they are not innately able to produce a correct song. During song development, birds compare what they produce to the memorized representation (template) of the song(s) that they heard during their 'sensitive period' early in life. We study song learning in the species of white- crowned sparrows that is found in our local mountains. Our work is directed at exploring the nature of the 'template', how experience shapes it, and how it is used to guide song development. Recent advances in digital signal processing now enable us to track the developmental paths that these birds take in producing complete song.

Social Control of Sex, Behavior and Coloration in Wrasses Wrasses are coral-reef fishes that exhibit highly plastic reproductive behavior and life histories. Individuals begin life in an 'initial phase', wherein males and females are similarly cryptically colored. Later in life, particular individuals may undergo a transformation, becoming more brilliantly colored and, if genetically female, switch sex. These 'supermales' maintain control over a harem of females. Remarkably, dominant females can display male-typical courtship behavior within 1 day after removal of the supermale. We have elected to study bluehead wrasses. Bluehead wrasses show remarkable gender plasticity that is governed by their social environment. Thus far, only field studies of sex transformation and behavior in these fish have been conducted. The difficulties of controlling in the field the social configuration and capturing fish that are engaged in particular behaviors have limited the utility of these fish as a model for investigating how changes in social environment can trigger ‘gender’ (behavioral sex phenotype) and sex transformation. In our project, we have constructed a novel captive environment in which these fish show natural reproductive behavior and rapidly transform from female to male in response to manipulations of their social environment; our lab was equipped with two 800-gallon custom built aquaria (4’x6’ floor, 4’ deep) that meet the minimum required space to express all natural courtship behaviors observed in the field. This advance now enabled us to do controlled behavioral tests, collect video recordings of behavior and extract brain tissue within minutes after capture. We have recently investigated the social factors that govern the decision to undergo this transformation. We are

11 Rose currently determining the expression of the c-Fos gene across brain regions in an effort to identify neural circuits that are active during male-typical courtship behaviors. We are using microiontophoresis to identify and map regions of the brain that cause changes in coloration typical of those observed during courtship behaviors. Eventually, we hope to understand the physiological processes that underlie transformations of behavioral sex phenotype.

Neural Control and Evolution of Electrosensory Behaviors In many animal behaviors, information about the environment is detected by sensory receptors and then transmitted to the central nervous system where stimulus patterns of relevance must be discriminated. Often, sensory signals are then translated into motor commands. The cellular mechanisms by which these operations are performed are poorly understood. Electric fish are particularly suitable for studying these questions. Behaviors such as the 'jamming avoidance responses' remain intact in neurophysiological preparations, permitting analysis of the entire neural circuit for generating these behaviors. We have found that short-term synaptic plasticity mechanisms are important in generating neural filters of temporal information. We have also pursued comparative neurophysiological studies of the electrosensory system of closely related species that lack jamming avoidance responses. These studies have shed light on how neural circuits change during evolution to generate new behaviors. While not an area of current research, I still remain interested in this system.

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