Cited by 86Animal Vocal Communication and BehaviorNeuroscience and Music PerceptionNeurogenesis and neuroplasticity mechanisms

Core and shell song systems unique to the parrot brain

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Chakraborty, Mukta, Walløe, Solveig, Nedergaard, Signe, Fridel, Emma E., Dabelsteen, Torben, Pakkenberg, Bente, Bertelsen, Mads F., Dorrestein, Gerry M. +3 more·2015·PLOS ONE

Abstract

The ability to imitate complex sounds is rare, and among birds has been found only in parrots, songbirds, and hummingbirds. Parrots exhibit the most advanced vocal mimicry among non-human animals. A few studies have noted differences in connectivity, brain position and shape in the vocal learning systems of parrots relative to songbirds and hummingbirds. However, only one parrot species, the budgerigar, has been examined and no differences in the presence of song system structures were found with other avian vocal learners. Motivated by questions of whether there are important differences in the vocal systems of parrots relative to other vocal learners, we used specialized constitutive gene expression, singing-driven gene expression, and neural connectivity tracing experiments to further characterize the song system of budgerigars and/or other parrots. We found that the parrot brain uniquely contains a song system within a song system. The parrot "core" song system is similar to the song systems of songbirds and hummingbirds, whereas the "shell" song system is unique to parrots. The core with only rudimentary shell regions were found in the New Zealand kea, representing one of the only living species at a basal divergence with all other parrots, implying that parrots evolved vocal learning systems at least 29 million years ago. Relative size differences in the core and shell regions occur among species, which we suggest could be related to species differences in vocal and cognitive abilities.

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  • Parrot pallial song nuclei underwent a further duplication event to evolve a unique additional circuit, the shell song system, which is particularly well developed in large-brained parrots.

    “Intriguingly, parrot pallial song nuclei underwent a further duplication event to evolve a unique additional circuit, the so-called shell song system, which seems to be particularly well developed in large-brained parrots (45).”

    From Birds have primate-like numbers of neurons in the forebrain (2016) · cites this paper as [45]

    The Introduction and Discussion describe a parrot-specific shell system that evolved after a core-like system, and the Results report larger shell regions in larger-brained species. However, the paper does not establish that the pallial song nuclei underwent a duplication event; it presents the system’s evolutionary sequence as a hypothesis. The Discussion also notes a larger-brained exception, the kea, whose shell specialization is not robust. The indexed passages provide section labels but no page numbers. (LLM, conf=0.86)

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  • The relatively enlarged subpallium in large parrots is likely associated with their greater learning skills, including vocal learning, and enhanced foot and beak dexterity.

    “Because subpallial structures play an important role in sensory and motor learning and execution of motor behavior (15, 44), we suggest that the relatively enlarged subpallium in large parrots is likely associated with their greater learning skills, including vocal learning, and enhanced foot and beak dexterity (5, 6, 13, 45).”

    From Birds have primate-like numbers of neurons in the forebrain (2016) · cites this paper as [45]

    The Results section on core and shell area ratios reports larger song-nucleus shell regions in larger-brained parrot species typically considered to have more advanced vocal and cognitive abilities. The paper does not establish that the subpallium as a whole is relatively enlarged in large parrots or link that enlargement to foot and beak dexterity. (LLM, conf=0.83)

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  1. [1]Nottebohm F. The origins of vocal learning. Am Nat. 1972; 106: 116– 140.
  2. [2]Janik VM, Slater PJB. Vocal learning in mammals. Adv Study Behav. 1997; 26: 59– 99.
  3. [3]Arriaga G, Zhou EP, Jarvis ED. Of mice, birds, and men: The mouse ultrasonic song system has some features similar to humans and song learning birds. PLoS ONE. 2012; 7: e46610 10.1371/journal.pone.0046610 23071596 PMC3468587 · DOI
  4. [4]Fitch WT, Jarvis ED. Birdsong and other animals models for human speech, song, and vocal learning In: Arbib MA, editor. Language, Music, and the Brain. Cambridge, MA: MIT Press 2013 pp. 499– 539.
  5. [5]Jarvis ED. Learned birdsong and the neurobiology of human language. Ann N Y Acad Sci. 2004; 1016: 749– 777. 15313804 PMC2485240
  6. [6]Fitch WT. The evolution of speech: a comparative review. Trends Cogn Sci. 2000; 4: 258– 267. 10859570
  7. [7]Suh A, Paus M, Kiefmann M, Churakov G, Franke FA, Brosius J, et al Mesozoic retroposons reveal parrots as the closest living relatives of passerine birds. Nature Commun. 2011; 2: 443. 21863010 PMC3265382
  8. [8]Jarvis ED, Mirarab S, Aberer AJ, Li B, Houde P, Li C, et al Whole-genome analyses resolve early branches in the tree of life of modern birds. Science. 2014; 12 12; 346(6215): 1320– 1321. 10.1126/science.1253451 25504713 PMC4405904 · DOI · Paper page
  9. [9]Cruickshank AJ, Gautier J-P, Chappuis C. Vocal mimicry in wild African Grey parrots Psittacus erithacus. Ibis. 1993; 135: 293– 299.
  10. [10]Gramza AF. Vocal mimicry in captive budgerigars (Melopsittacus undulatus). Z Tierpsychol. 1970; 27: 971– 983.
  11. [11]Kelley LA, Coe RL, Madden JR, Healy SD. Vocal mimicry in songbirds. Anim Behav. 2008; 76: 521– 528.
  12. [12]Kelley LA, Healy SD. Vocal mimicry in male bowerbirds: who learns from whom? Biol Lett. 2010; 6: 626– 629. 10.1098/rsbl.2010.0093 20236967 PMC2936137 · DOI
  13. [13]Kelley LA, Healy SD. Vocal mimicry. Curr Biol. 2011; 21(1): R9– R10. 10.1016/j.cub.2010.11.026 21215939 · DOI
  14. [14]Klatt DH, Stefanski RA. How does a mynah bird imitate human speech? The J Acoust Soc Am. 1974; 55: 822– 832. 4833078
  15. [15]Pepperberg I. The Alex Studies: Cognitive and Communicative Abilities of Grey Parrots. Cambridge, MA: Harvard University Press; 1999
  16. [16]Schachner A. Auditory-motor entrainment in vocal mimicking species: Additional ontogenetic and phylogenetic factors. Commun Integr Biol. 2010; 3: 290– 293. 20714417 PMC2918780
  17. [17]Pepperberg IM. Vocal learning in Grey parrots: A brief review of perception, production, and cross-species comparisons. Brain Lang. 2010; 115: 81– 91. 10.1016/j.bandl.2009.11.002 20199805 · DOI
  18. [18]Colbert-White EN, Covington MA, Fragaszy DM. Social context influences the vocalizations of a home-raised African Grey parrot (Psittacus erithacus erithacus). J Comp Psychol. 2011; 125: 175– 184. 10.1037/a0022097 21381810 · DOI
  19. [19]Poole JH, Tyack PL, Stoeger-Horwath AS, Watwood S. Anim Behav. Elephants are capable of vocal learning. Nature. 2005; 434: 455– 456. 15791244
  20. [20]Richards DG, Wolz JP, Herman LM. Vocal mimicry of computer-generated sounds and vocal labeling of objects by a bottlenosed dolphin, Tursiops truncatus. J Comp Psychol. 1984; 98: 10– 28. 6705501
  21. [22]Ridgway S, Carder D, Jeffries M, Todd M. Spontaneous human speech mimicry by a cetacean. Curr Biol. 2012; 22: R860– 861. 10.1016/j.cub.2012.08.044 23098588 · DOI
  22. [23]Stoeger AS, Mietchen D, Oh S, de Silva S, Herbst CT, Kwon S, et al An Asian elephant imitates human speech. Curr Biol. 2012; 22: 2144– 2148. 10.1016/j.cub.2012.09.022 23122846 PMC3548412 · DOI
  23. [24]Jarvis ED, Mello CV. Molecular mapping of brain areas involved in parrot vocal communication. J Comp Neurol. 2000; 419: 1– 31. 10717637 PMC2538445
  24. [25]Jarvis ED. Brains and birdsong In: Marler P, Slabberkoom H, editors. Nature's music: The Science of Birdsong. New York: Elsevier- Academic Press; 2004.
  25. [26]Gahr M. Neural song control system of hummingbirds: comparison to swifts, vocal learning (songbirds) and nonlearning (suboscines) passerines, and vocal learning (budgerigars) and nonlearning (dove, owl, gull, quail, chicken) nonpasserines. J Comp Neurol. 2000; 426: 182– 196. 10982462
  26. [27]Jarvis ED, Ribeiro S, da Silva ML, Ventura D, Vielliard J, Mello, CV. Behaviourally driven gene expression reveals song nuclei in hummingbird brain. Nature. 2000; 406: 628– 632. 10949303 PMC2531203
  27. [28]Katz LC, Gurney ME. Auditory responses in the zebra finch's motor system for song. Brain Res. 1981; 221: 192– 197. 6168335
  28. [29]McCasland JS, Konishi M. Interaction between auditory and motor activities in an avian song control nucleus. Proc Natl Acad Sci USA. 1981; 78: 7815– 7819. 6950421 PMC349362
  29. [30]Jarvis ED, Scharff C, Grossman MR, Ramos JA, Nottebohm F. For whom the bird sings: context-dependent gene expression. Neuron. 1988; 21: 775– 788. 9808464
  30. [31]Brauth SE, Liang W, Roberts TF. Projections of the oval nucleus of the hyperstriatum ventrale in the budgerigar: relationships with the auditory system. J Comp Neurol. 2001; 432: 481– 511. 11268010
  31. [32]Striedter GF. The vocal control pathways in budgerigars differ from those in songbirds. J Comp Neurol. 1994; 343: 35– 56. 8027436
  32. [33]Paton JA, Manogue KR, Nottebohm F. Bilateral organization of the vocal control pathway in the budgerigar, Melopsittacus undulatus. J Neurosci. 1981; 1: 1279– 1288. 6171631 PMC6564219
  33. [34]Durand SE, Heaton JT, Amateau SK, Brauth SE. Vocal control pathways through the anterior forebrain of a parrot (Melopsittacus undulatus). J Comp Neurol. 1997; 377: 179– 206. 8986880
  34. [35]Feenders G, Liedvogel M, Rivas M, Zapka M, Horita H, Hara E, et al Molecular mapping of movement-associated areas in the avian brain: a motor theory for vocal learning origin. PLoS ONE. 2008; 3: e1768 10.1371/journal.pone.0001768 18335043 PMC2258151 · DOI
  35. [36]Farabaugh SM, Wild JM. Reciprocal connections between primary and secondary auditory pathways in the telencephalon of the budgerigar (Melopsittacus undulatus). Brain Res. 1997; 747(1): 18– 25. 9042523
  36. [38]Hara E, Rivas MV, Ward JM, Okanoya K, Jarvis ED. Convergent differential regulation of parvalbumin in the brains of vocal learners. PLoS One 7 2012; e29457 10.1371/journal.pone.0029457 22238614 PMC3253077 · DOI
  37. [40]Celio MR. Parvalbumin in most gamma-aminobutyric acid-containing neurons of the rat cerebral cortex. Science. 1986; 231: 995– 997. 3945815
  38. [41]Braun K, Scheich H, Zuschratter W, Heizmann CW, Matute C, Streit P. Postnatal development of parvalbumin-, calbindin- and adult GABA-immunoreactivity in two visual nuclei of zebra finches. Brain Res. 1988; 475: 205– 217. 3214731
  39. [42]Chen CC, Winkler CM, Pfenning AR, Jarvis ED. Molecular profiling of the developing avian telencephalon: regional timing and brain subdivision continuities. J Comp Neurol. 2013; 521: 3666– 3701. 10.1002/cne.23406 23818174 PMC3863995 · DOI
  40. [43]Jarvis ED, Yu J, Rivas MV, Horita H, Feenders G, Whitney O, et al Global view of the functional molecular organization of the avian cerebrum: mirror images and functional columns. J Comp Neurol. 2013; 521: 3614– 3665. 10.1002/cne.23404 23818122 PMC4145244 · DOI
  41. [44]Jarvis ED. Neural systems for vocal learning in birds and humans: a synopsis. J Ornithol. 2007; 143: S35– 44. PMC2726745 19684872
  42. [45]Horita H, Kobayashi M, Liu WC, Oka K, Jarvis ED, Wada K. Specialized motor-driven dusp1 expression in the song systems of multiple lineages of vocal learning birds. PLoS One. 2012; 7: e42173 10.1371/journal.pone.0042173 22876306 PMC3410896 · DOI
  43. [46]Wada K, Howard JT, McConnell P, Whitney O, Lints T, Rivas MV, et al A molecular neuroethological approach for identifying and characterizing a cascade of behaviorally regulated genes. Proc Natl Acad Sci USA. 2006; 103: 15212– 15217. 17018643 PMC1622802
  44. [47]Jarvis ED, Nottebohm F. Motor-driven gene expression. Proc Natl Acad Sci USA. 1997; 94: 4097– 4102. 9108111 PMC20574
  45. [48]Mello CV, Clayton DF. Song-induced ZENK gene expression in auditory pathways of songbird brain and its relation to the song control system. J Neurosci. 1994; 14: 6652– 6666. 7965067 PMC6577290
  46. [49]Mello CV, Vates GE, Okuhata S, Nottebohm F. Descending auditory pathways in the adult male zebra finch (Taeniopygia guttata). J Comp Neurol. 1988; 395: 137– 160. 9603369
  47. [51]Bottjer SW. Developmental regulation of basal ganglia circuitry during the sensitive period for vocal learning in songbirds. Ann N Y Acad Sci. 2004; 1016: 395– 415. 15313787
  48. [52]Bottjer SW, Altenau B. Parallel pathways for vocal learning in basal ganglia of songbirds. Nat Neurosci. 2010; 13: 153– 155. 10.1038/nn.2472 20023650 PMC2846604 · DOI
  49. [53]Fisher SE, Scharff C. FOXP2 as a molecular window into speech and language. Trends Genet. 2009; 25(4): 166– 177. 10.1016/j.tig.2009.03.002 19304338 · DOI
  50. [54]Haesler S, Wada K, Nshdejan A, Morrisey EE, Lints T, Jarvis ED, et al FoxP2 expression in avian vocal learners and non-learners. J Neurosci. 2004; 24: 3164– 3175. 15056696 PMC6730012
  51. [55]Whitney O, Voyles T, Hara E, Chen Q, White SA, White TF. Differential FoxP2 and FoxP1 expression in a vocal learning nucleus of the developing budgerigar. Dev Neurobiol. 2014; 10.1002/dneu.22247 PMC4437895 25407828 · DOI
  52. [56]Wada K, Sakaguchi H, Jarvis ED, Hagiwara M. Differential expression of glutamate receptors in avian neural pathways for learned vocalization. J Comp Neurol. 2004; 476: 44– 64. 15236466 PMC2517240
  53. [57]Brauth SE, Liang W, Amateau SK, Roberts TF. Sexual dimorphism of vocal control nuclei in budgerigars (Melopsittacus undulatus) revealed with Nissl and NADPH-d staining. J Comp Neurol. 2005; 484: 15– 27. 15717302
  54. [58]Durand SE, Liang W, Brauth SE. Methionine enkephalin immunoreactivity in the brain of the budgerigar (Melopsittacus undulatus): similarities and differences with respect to oscine songbirds. J Comp Neurol. 1998; 393: 145– 168. 9548694
  55. [59]Durand SE, Brauth SE, Liang W. Calcitonin gene-related peptide immunoreactive cells and fibers in forebrain vocal and auditory nuclei of the budgerigar (Melopsittacus undulatus). Brain Behav Evol. 2001; 58: 61– 79. 11805374
  56. [60]Matsunaga E, Okanoya K. Vocal area-related expression of the androgen receptor in the budgerigar (Melopsittacus undulatus) brain. Brain Res. 2008; 1208: 87– 94. 10.1016/j.brainres.2008.02.076 18394591 · DOI
  57. [61]Roberts TF, Cookson KK, Heaton KJ, Hall WS, Brauth SE. Distribution of tyrosine hydroxylase-containing neurons and fibers in the brain of the budgerigar (Melopsittacus undulatus): general patterns and labeling in vocal control nuclei. J Comp Neurol. 2001; 429: 436– 454. 11116230
  58. [62]Joseph L, Toon A, Schirtzinger EE, Wright TF, Schodde R. A revised nomenclature and classification for family-group taxa of parrots (Psittaciformes). Zootaxa. 2012; 3205: 26– 40.
  59. [63]Jarvis ED, Gunturkun O, Bruce L, Csillag A, Karten H, Kuenzel W, et al Avian brains and a new understanding of vertebrate brain evolution. Nat Rev Neurosci. 2005; 6: 151– 159. 15685220 PMC2507884
  60. [65]Fitch WT. Evolutionary Developmental Biology and Human Language Evolution: Constraints on Adaptation. Evo Biol. 2012; 39(4): 613– 637. 23226905 PMC3514691
  61. [66]Fitch WT. The evolution of language. New York, USA: Cambridge University Press; 2012.
  62. [67]Petkov CI, Jarvis ED. Birds, primates, and spoken language origins: behavioral phenotypes and neurobiological substrates. Front Evol Neurosci. 2012; 4: 12 10.3389/fnevo.2012.00012 22912615 PMC3419981 · DOI
  63. [69]Liu W, Wada K, Jarvis ED, Nottebohm F. Rudimentary substrates for vocal learning in a suboscine. Nat Commun. 2013; 4: 2082: 1– 12. 10.1038/ncomms3082 23823977 · DOI
  64. [70]Kroodsma D, Hamilton D, Sanchez JE, Byers BE, Fandino-Marino H, Stemple DW, et al Behavioral Evidence for Song Learning in the Suboscine Bellbirds (Procnias Spp.; Cotingidae). Wilson J Ornithol. 2013; 125: 1– 14. 10.1159/000360034 24662179 · DOI
  65. [71]Touchton JM, Seddon N, Tobias JA. Captive Rearing Experiments Confirm Song Development without Learning in a Tracheophone Suboscine Bird. PLoS One. 2014; 9: e95746 10.1371/journal.pone.0095746 24788343 PMC4005748 · DOI
  66. [72]Harpøth SW. Vocal modification abilities and brain structures in parrots- how do they correlate? Ph.D. Thesis, Department of Biology, Graduate School at the Faculty of Science, University of Copenhagen. 2013.
  67. [73]Bond AB, Diamond J. Geographic and ontogenetic variation in the contact calls of the kea (Nestor notabilis). Behaviour. 2005; 142: 1– 20.
  68. [74]Lavenex PB. Lesions in the budgerigar vocal control nucleus NLc affect production, but not memory, of english words and natural vocalizations. J Comp Neurol. 2000; 421: 437– 460. 10842208
  69. [76]Wong K, Park HT, Wu JY, Rao Y. Slit proteins: molecular guidance cues for cells ranging from neurons to leukocytes. Curr Opin Genet Dev. 2002; 12: 583– 591. 12200164
  70. [77]Kim M, Roesener AP, Mendonca PR, Mastick GS. Robo1 and Robo2 have distinct roles in pioneer longitudinal axon guidance. Dev Biol. 2011; 358: 181– 188. 10.1016/j.ydbio.2011.07.025 21820427 PMC3171630 · DOI
  71. [79]Hannula-Jouppi K, Kaminen-Ahola N, Taipale M, Eklund R, Nopola-Hemmi J, Kaariainen H, et al The axon guidance receptor gene ROBO1 is a candidate gene for developmental dyslexia. PLoS Genetics. 2005; 1: e50 16254601 PMC1270007
  72. [80]Cook P, Rouse A, Wilson M, Reichmuth C. A California Sea Lion (Zalophus californianus) Can Keep the Beat: Motor Entrainment to Rhythmic Auditory Stimuli in a Non Vocal Mimic. J Comp Psychol. 2013; 127: 412– 427. 10.1037/a0032345 23544769 · DOI
  73. [81]Patel AD, Iversen JR, Bregman MR, Schulz I. Experimental evidence for synchronization to a musical beat in a nonhuman animal. Curr Biol. 2009:CB 19: 827– 830. 10.1016/j.cub.2009.03.038 19409790 · DOI
  74. [82]Schachner A, Brady TF, Pepperberg IM, Hauser MD. Spontaneous motor entrainment to music in multiple vocal mimicking species. Curr Biol. 2009:CB 19: 831– 836. 10.1016/j.cub.2009.03.061 19409786 · DOI
  75. [83]Chen CC, Wada K, Jarvis ED. Radioactive in situ hybridization for detecting diverse gene expression patterns in tissue. J Vis Exp. 2012; 3764 [pii] 10.3791/3764 22566072 PMC3466668 · DOI
  76. [84]Horita H, Wada K, Rivas MV, Hara E, Jarvis ED. The dusp1 immediate early gene is regulated by natural stimuli predominantly in sensory input neurons. J Comp Neurol. 2010; 518: 2873– 2901. 10.1002/cne.22370 20506480 PMC2904818 · DOI
  77. [85]Ball GF. Neurochemical specializations associated with vocal learning and production in songbirds and budgerigars. Brain Behav Evol. 1994; 44: 234– 246. 7842283

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Authors
Chakraborty, M., Walløe, S., Nedergaard, S., Fridel, E. E., Dabelsteen, T., Pakkenberg, B., Bertelsen, M. F., Dorrestein, G. M., Brauth, S. E., Durand, S. E., & Jarvis, E. D.
Affiliations
  • Howard Hughes Medical Institute (US)
  • University of Copenhagen (DK)
  • Danish National Police (DK)
  • Duke Medical Center (US)
  • Bispebjerg Hospital (DK)
  • Copenhagen Zoo (DK)
  • University of Maryland, College Park (US)
  • LaGuardia Community College (US)
Corresponding author
  • Chakraborty, Mukta
  • Walløe, Solveig
  • Nedergaard, Signe
  • Fridel, Emma E.
  • Dabelsteen, Torben
  • Pakkenberg, Bente
  • Bertelsen, Mads F.
  • Dorrestein, Gerry M.
  • Brauth, Steven E.
  • Durand, Sarah E.
  • Jarvis, Erich D.
Venue
PLOS ONE
Published
2015-06-24
Volume
10
Issue
6
Pages
e0118496
ISSN
1932-6203 (ISSN-L)
OpenAlex
W1598996539
Cited by
86
References
89
Citations by year
2026: 8 · 2025: 9 · 2024: 8 · 2023: 6 · 2022: 6 · 2021: 15 · 2020: 9 · 2019: 4 · 2018: 6 · 2017: 3 · 2016: 9 · 2015: 2 · 2014: 1
FWCI
5.80
Type
journal article
Language
en
Open access
Locations
  • PLoS ONE (publishedVersion, cc-by)
  • Europe PMC (PubMed Central) (publishedVersion, cc-by)
  • PubMed Central (publishedVersion, cc-by)
  • PubMed (publishedVersion)
  • Research at the University of Copenhagen (University of Copenhagen) (publishedVersion, cc-by)
  • CUNY Academic Works (City University of New York) (submittedVersion, cc-by)
  • DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
  • DukeSpace (Duke University) (submittedVersion, cc-by-nc-nd)
  • Figshare (submittedVersion, cc-by)
Funders
  • Howard Hughes Medical Institute
  • Danmarks Frie Forskningsfond
  • National Institutes of Health
Topics
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Developmental Biology › Animal Vocal Communication and Behavior
  • Life Sciences › Neuroscience › Cognitive Neuroscience › Neuroscience and Music Perception
  • Life Sciences › Neuroscience › Developmental Neuroscience › Neurogenesis and neuroplasticity mechanisms
Keywords
vocal learning, vocal mimicry, budgerigars, avian brain evolution
MeSH terms
Animals, Brain, Humans, Learning, Music, New Zealand, Parrots, Vocalization, Animal

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