title: Investigation of musicality in birdsong.
abstract: Songbirds spend much of their time learning, producing, and listening to complex vocal sequences we call songs. Songs are learned via cultural transmission, and singing, usually by males, has a strong impact on the behavioral state of the listeners, often promoting affiliation, pair bonding, or aggression. What is it in the acoustic structure of birdsong that makes it such a potent stimulus? We suggest that birdsong potency might be driven by principles similar to those that make music so effective in inducing emotional responses in humans: a combination of rhythms and pitches-and the transitions between acoustic states-affecting emotions through creating expectations, anticipations, tension, tension release, or surprise. Here we propose a framework for investigating how birdsong, like human music, employs the above "musical" features to affect the emotions of avian listeners. First we analyze songs of thrush nightingales (Luscinia luscinia) by examining their trajectories in terms of transitions in rhythm and pitch. These transitions show gradual escalations and graceful modifications, which are comparable to some aspects of human musicality. We then explore the feasibility of stripping such putative musical features from the songs and testing how this might affect patterns of auditory responses, focusing on fMRI data in songbirds that demonstrate the feasibility of such approaches. Finally, we explore ideas for investigating whether musical features of birdsong activate avian brains and affect avian behavior in manners comparable to music's effects on humans. In conclusion, we suggest that birdsong research would benefit from current advances in music theory by attempting to identify structures that are designed to elicit listeners' emotions and then testing for such effects experimentally. Birdsong research that takes into account the striking complexity of song structure in light of its more immediate function - to affect behavioral state in listeners - could provide a useful animal model for studying basic principles of music neuroscience in a system that is very accessible for investigation, and where developmental auditory and social experience can be tightly controlled.
authors.affiliation: Department of Humanities, New Jersey Institute of Technology, University Heights, Newark, NJ 07102, USA.
authors.forename: David
authors.initials: D
authors.lastname: Rothenberg
authors.affiliation: Department of Psychology, Hunter College, City University of New York, New York, NY 10065, USA.
authors.forename: Tina C
authors.initials: TC
authors.lastname: Roeske
authors.affiliation: Department of Radiology, Citigroup Biomedical Imaging Center, Weill Cornell Medical College, New York, NY 10021, USA.
authors.forename: Henning U
authors.initials: HU
authors.lastname: Voss
authors.affiliation: Behavioural Ecology Group, Animal Science Department, Wageningen University, Wageningen, The Netherlands.
authors.forename: Marc
authors.initials: M
authors.lastname: Naguib
authors.affiliation: Department of Psychology, Hunter College, City University of New York, New York, NY 10065, USA. Electronic address: tchernichovski@gmail.com.
authors.forename: Ofer
authors.initials: O
authors.lastname: Tchernichovski
mesh_terms: D000818:Animals; D001307:Auditory Perception / Q000502:physiology; D001921:Brain / Q000502:physiology; D004644:Emotions; D005260:Female; D006309:Hearing; D006801:Humans; D008279:Magnetic Resonance Imaging; D008297:Male; D009146:Music; D010898:Pitch Perception; D020308:Songbirds / Q000502:physiology*; D014828:Vocalization, Animal
references.citation: Science. 2001 Jun 29;292(5526):2432-3
references.pmid: 11441879
references.citation: Proc Natl Acad Sci U S A. 2011 Sep 13;108(37):15510-5
references.pmid: 21876156
references.citation: Proc Biol Sci. 2004 Mar 7;271(1538):483-91
references.pmid: 15129958
references.citation: Proc Natl Acad Sci U S A. 1995 Apr 11;92(8):3406-10
references.pmid: 7724575
references.citation: Neurosci Biobehav Rev. 2011 Oct;35(9):1837-45
references.pmid: 21251924
references.citation: Nature. 2006 Apr 27;440(7088):1204-7
references.pmid: 16641998
references.citation: J Neurosci. 2006 Aug 30;26(35):9010-4
references.pmid: 16943558
references.citation: Proc Biol Sci. 2004 May 7;271 Suppl 4:S167-9
references.pmid: 15252973
references.citation: J Neurophysiol. 2008 Apr;99(4):1700-11
references.pmid: 18216221
references.citation: NMR Biomed. 2010 Nov;23(9):1027-32
references.pmid: 20806227
references.citation: Cognition. 2006 May;100(1):173-215
references.pmid: 16412411
references.citation: Hum Brain Mapp. 2007 Nov;28(11):1150-62
references.pmid: 17290372
references.citation: J Ornithol. 2007 Dec 1;148(1):35-44
references.pmid: 19684872
references.citation: J Neurobiol. 2003 Sep 15;56(4):303-14
references.pmid: 12918015
references.citation: Nat Neurosci. 2011 Jul 26;14(8):947-8
references.pmid: 21792190
references.citation: Perspect Biol Med. 2005 Summer;48(3):426-43
references.pmid: 16085998
references.citation: Neuron. 1998 Oct;21(4):775-88
references.pmid: 9808464
references.citation: Science. 2001 Jan 5;291(5501):52-4
references.pmid: 11192008
references.citation: Eur J Neurosci. 2007 Jun;25(11):3406-16
references.pmid: 17553009
references.citation: Nat Neurosci. 2005 Oct;8(10):1371-9
references.pmid: 16136039
references.citation: Nature. 2013 Jun 6;498(7452):104-8
references.pmid: 23719373
references.citation: Proc Biol Sci. 2009 Jun 7;276(1664):2045-50
references.pmid: 19324798
references.citation: Anim Behav. 2000 Feb;59(2):443-458
references.pmid: 10675267
references.citation: Proc Natl Acad Sci U S A. 1992 Feb 15;89(4):1372-5
references.pmid: 1741391
references.citation: Nature. 2008 Jan 17;451(7176):305-10
references.pmid: 18202651
references.citation: J Neurosci. 2008 Dec 3;28(49):13232-47
references.pmid: 19052215
references.citation: J Neurophysiol. 2010 Nov;104(5):2474-86
references.pmid: 20884763
references.citation: Anim Behav. 2000 Sep;60(3):297-306
references.pmid: 11007638
references.citation: J Comp Physiol A Neuroethol Sens Neural Behav Physiol. 2002 Dec;188(11-12):879-95
references.pmid: 12471488
references.citation: Nature. 2009 May 28;459(7246):564-8
references.pmid: 19412161
references.citation: Ann N Y Acad Sci. 2009 Jul;1169:209-13
references.pmid: 19673783
references.citation: Trends Cogn Sci. 2009 Dec;13(12):505-10
references.pmid: 19892586
references.citation: Anim Behav. 2000 Jun;59(6):1167-1176
references.pmid: 10877896
references.citation: Dev Neurobiol. 2010 Jan;70(1):28-40
references.pmid: 19937773
references.citation: Trends Neurosci. 2005 Jul;28(7):353-63
references.pmid: 15935486
references.citation: Nat Rev Neurosci. 2005 Feb;6(2):151-9
references.pmid: 15685220
references.citation: J Comp Neurol. 2005 Jul 18;488(1):48-60
references.pmid: 15912499
references.citation: PLoS One. 2010 Dec 23;5(12):e14415
references.pmid: 21203446
references.citation: Nat Rev Neurosci. 2010 Nov;11(11):747-59
references.pmid: 20959859
references.citation: Trends Neurosci. 2009 May;32(5):257-66
references.pmid: 19307029
references.citation: Behav Brain Res. 2011 Dec 1;225(2):511-4
references.pmid: 21856337
references.citation: Trends Neurosci. 2005 Mar;28(3):127-32
references.pmid: 15749165
references.citation: Curr Biol. 1997 May 1;7(5):R289-91
references.pmid: 9115379
references.citation: Percept Psychophys. 1980 Nov;28(5):381-9
references.pmid: 7208247
references.citation: J Neurosci. 2009 Feb 18;29(7):2252-8
references.pmid: 19228978
references.citation: J Neurobiol. 1992 Oct;23(8):1006-20
references.pmid: 1460461
references.citation: Brain Lang. 2012 Sep;122(3):142-50
references.pmid: 22284300
references.citation: Neuroimage. 2005 May 1;25(4):1242-55
references.pmid: 15850742
references.citation: Front Evol Neurosci. 2012 Nov 28;4:14
references.pmid: 23226128
references.citation: J Neurobiol. 1993 Apr;24(4):488-505
references.pmid: 8515252
references.citation: Proc Biol Sci. 2010 Jan 7;277(1678):153-60
references.pmid: 19812084
references.citation: Horm Behav. 1997 Feb;31(1):25-34
references.pmid: 9109596
references.citation: J Exp Zool. 1971 Jun;177(2):229-61
references.pmid: 5571594
references.citation: Proc Natl Acad Sci U S A. 2007 Jun 19;104(25):10667-72
references.pmid: 17563380
references.citation: Science. 2013 Apr 12;340(6129):216-9
references.pmid: 23580531
references.citation: Proc Natl Acad Sci U S A. 2010 Feb 2;107(5):2301-6
references.pmid: 20133876
references.citation: Proc Biol Sci. 2000 Dec 22;267(1461):2553-8
references.pmid: 11197134
references.citation: PLoS One. 2011;6(5):e20131
references.pmid: 21625389
references.citation: Brain Lang. 2010 Oct;115(1):21-8
references.pmid: 19913899
references.citation: J Neurosci. 1997 Feb 1;17(3):1147-67
references.pmid: 8994068
references.citation: Neuroimage. 2006 Jul 1;31(3):1327-42
references.pmid: 16546406
references.citation: Proc Natl Acad Sci U S A. 1985 Sep;82(17):5997-6000
references.pmid: 16593601
references.citation: PLoS One. 2008 Sep 10;3(9):e3184
references.pmid: 18781203
references.citation: Neuropsychologia. 2008 Jan 15;46(1):362-8
references.pmid: 17709121
references.citation: J Neurophysiol. 2003 Nov;90(5):2884-99
references.pmid: 12878713
references.citation: Hum Brain Mapp. 2006 Mar;27(3):239-50
references.pmid: 16078183
references.citation: Proc Natl Acad Sci U S A. 2010 Apr 20;107(16):E65; author reply E66-7
references.pmid: 20388905
references.citation: J Neurosci. 2006 Mar 1;26(9):2499-512
references.pmid: 16510728
references.citation: Proc Natl Acad Sci U S A. 2009 Dec 1;106(48):20538-43
references.pmid: 19918074
references.citation: Proc Natl Acad Sci U S A. 2001 Sep 25;98(20):11818-23
references.pmid: 11573015
references.citation: Neuroscience. 2003;122(2):521-9
references.pmid: 14614916
references.citation: Nat Neurosci. 2011 Jun 26;14(8):1067-74
references.pmid: 21706017
references.citation: PLoS One. 2010 Dec 16;5(12):e13812
references.pmid: 21179549
references.citation: J Chem Neuroanat. 2010 Mar;39(2):112-23
references.pmid: 19900537
references.citation: Front Evol Neurosci. 2012 Apr 13;4:5
references.pmid: 22518103
references.citation: Magn Reson Imaging. 2001 Jul;19(6):821-6
references.pmid: 11551722
references.citation: Eur J Neurosci. 2007 Nov;26(9):2613-26
references.pmid: 17970728
references.citation: Proc Biol Sci. 2002 Jul 22;269(1499):1479-85
references.pmid: 12137578
references.citation: Nat Neurosci. 2011 Feb;14(2):257-62
references.pmid: 21217764
references.citation: J Cogn Neurosci. 2001 Aug 15;13(6):786-92
references.pmid: 11564322
references.citation: Trends Cogn Sci. 2011 Mar;15(3):113-21
references.pmid: 21296608
references.citation: Annu Rev Neurosci. 1999;22:567-631
references.pmid: 10202549
references.citation: J Neurosci. 1998 May 15;18(10):3786-802
references.pmid: 9570809
references.citation: J Cogn Neurosci. 2003 Jul 1;15(5):673-82
references.pmid: 12965041
references.citation: J Neurobiol. 1990 Dec;21(8):1236-48
references.pmid: 2273402
references.citation: J Neurosci. 1997 Aug 15;17(16):6447-62
references.pmid: 9236252
references.citation: Proc Natl Acad Sci U S A. 2012 Jul 31;109(31):12782-7
references.pmid: 22802637
references.citation: J Neurophysiol. 2008 Feb;99(2):931-8
references.pmid: 17881485
references.citation: Front Evol Neurosci. 2011 Dec 23;3:9
references.pmid: 22207847
references.citation: Proc Natl Acad Sci U S A. 2010 Feb 23;107(8):3852-7
references.pmid: 20133597
references.citation: PLoS One. 2011;6(11):e27241
references.pmid: 22110619