Untersuchungen zu neuroanatomischen Veränderungen beim  Gesangslernen des Zebrafinken Taeniopygia guttata Viola von Bohlen und Halbach1.1231.1.4484#2371.2.5585#3821.3.1.4.6553#4011.4.1.1.4.1.1.71.4.1.2.81.4.1.3.1.4.1.4.91.4.1.5.101.4.1.6.111.4.1.7.1.4.2.121.4.2.1.131.4.2.2.14572#4151.5.151.5.1.16518#463171.5.2.1819201.6.212.2.1.2.1.1.2122316#1612.1.2.232.2.2.2.1.24400#55625360#4622.3.26272.3.1.282.3.2.2.4.2.4.1.292.4.2.2.4.2.1.30250#4032.4.3.31634#445322.4.3.1.33480#3602.4.3.2.3435363738645#2122.4.4.393.39403.1.3.1.1.41645#215423.1.2.3.1.2.1.4344405#28045635#4283.1.2.2.46636#48347635#4353.1.3.483.2.493.2.1.3.2.1.1.50624#42351624#42752520#61353553#3243.2.1.2.543.2.2.3.2.2.1.55644#21956247#2673.2.3.3.2.3.1.57645#21158293#3203.2.4.593.2.4.1.60624#427605#416613.2.4.2.532#62862553#2973.2.5.633.2.5.1.64614#424605#43165546#64566554#3203.2.5.2.3.2.6.673.2.6.1.68809#5673.2.6.2.6970930#3944.70714.1.72734.2.4.2.1.74644#22975764.2.2.7778622#2097980818283844.3.858687884.4.89466#75990ABKÜRZUNGSVERZEICHNISLITERATURVERZEICHNISANHANGVeröffentlichungsnachweisErklärungdeINHALTSVERZEICHNISHilfe LITERATURVERZEICHNIS Abercrombie M (1946) Estimation of nuclear population from microtome sections. Anat. Rec. 94: 239-247 Airey DC, Castillo-Juarez H, Casella G, Pollak EJ, DeVoogd TJ (2000) Variation in the volume of zebra finch song control nuclei is heritable: developmental and evolutionary implications. Proc R Soc Lond B Biol Sci 267: 2099-104 Airey DC, DeVoogd TJ (2000) Greater song complexity is associated with augmented song system anatomy in zebra finches. NeuroReport 11: 2339-2344 Akutagawa E, Konishi M (1994) Two separate areas of the brain differentially guide the development of a song control nucleus in the zebra finch. Proc.Natl.Acad.Sci.U.S.A. 91: 12413-12417 Alvarez-Buylla A, Kirn JR (1997) Birth, migration, incorporation, and death of vocal control neurons in adult songbirds. J. Neurobiol. 33: 585-601 Alvarez-Buylla A, Kirn JR, Nottebohm F (1990a) Birth of projection neurons in adult avian brain may be related to perceptual or motor learning. Science 249: 1444-6 Alvarez-Buylla A, Ling CY, Kirn JR (1990b) Cresyl violet: a red fluorescent Nissl stain. J.Neurosci.Methods 33: 129-133 Alvarez-Buylla A, Ling C-Y, Nottebohm F (1992) High vocal center growth and its relation to neurogenesis, neuronal replacement and song acquisition in juvenile canaries. J.Neurobiol. 23: 396-406 Arnold AP (1975) The effects of castration on song development in zebra finches (Poephila guttata). J. Exp. Zool. 191: 261-278 Arnold AP (1980a) Effects of androgens on volumes of sexually dimorphic brain regions in the zebra finch. Brain Res. 185: 441-444 Arnold AP (1980b) Quantitative analysis of sex differences in hormone accumulation in the zebra finch brain: methodological and theoretical issues. J.Comp.Neurol. 189: 421-436 Arnold AP (1980c) Sexual differences in the brain. Ann. Scientist 68: 165-173 Arnold AP, Bottjer SW, Brenowitz EA, Nordeen EJ, Nordeen KW (1986) Sexual dimorphisms in the neural vocal control system on song birds: ontogeny and phylogeny. Brain Behav.Evol. 28: 22-31 Arnold AP, Nottebohm F, Pfaff DW (1976) Hormone concentrating cells in vocal control and other areas of the brain of the zebra finch (Poephila guttata). J.Comp.Neurol. 165: 487-511 Ball GF, Riters LV, Balthazart J (2002) Neuroendocrinology of song behavior and avian brain plasticity: multiple sites of actio of sex steroid hormones. Front Neuroendocrinol 23: 137-78 Basham ME, Nordeen EJ, Nordeen KW (1996) Blockade of NMDA receptors in the anterior forebrain impairs sensory acquisition in the zebra finch (Poephila guttata). Neurobiol Learn Mem 66: 295-304. Bischof HJ, Böhner J, Sossinka R (1981) Influence of external stimuli on the quality of the song of the zebra finch (Taeniopygia guttata castanotis Gould). Z.Tierpsychol. 57: 261-267 Bischof HJ, Herrmann K (1988) Isolation-dependent enhancement of 2-[(14 )C ] deoxyglucose uptake in the forebrain of zebra finch males. Behav.Neural.Biol. 49: 386-397 Bock J, Braun K (1999) Filial imprinting in domestic chicks is associated with spine pruning in the associative area, dorsocaudal neostriatum. E J Neuroscience 11: 2566-2570. Boettiger CA, Doupe AJ (1998) Intrinsic and thalamic excitatory inputs onto songbird LMAN neurons differ in their pharmacological and temporal properties. J Neurophysiol 79: 2615-2628 Boettiger CA, Doupe AJ (2001) Developmentally restricted synaptic plasticity in a songbird nucleus required for song learning. Neuron 31: 809- 818 Böhner J (1981) Untersuchungen zur Selektivität des Gesangserwerbs beim australischen Zebrafinken (Taeniopygia guttata castanotis). Böhner J (1990) Early acquisition of song in the zebra finch, Taeniopygia guttata. Anim.Behav. 39: 369-374 Böhner J (1991) Ontogenetische Entwicklung von Gesangsaktivität und -struktur beim Zebrafinken (Taeniopygia guttata). Verh.Dtsch.Zool.Ges. 84: 468-469 Böhner J, Harding CF, Marler P (1992) Androstenedione therapy reinstates normal, not supernormal, song structure in castrated adult male zebra finches. Hormones and Behavior 26:136-142 Bolhuis JJ, Van Mil DP, Houx BB (1999) Song learning with audiovisual compound stimuli in zebra finches. Anim Behav 58:1285-1292 Bottjer SW (2004) Developmental regulation of basal ganglia circuitry during the sensitve period for vocal learning in songbirds. Ann. N. Y. Acad. Sci. 1016:395-415 Bottjer SW, Arnold AP (1984) The role of feedback from the vocal organ. I. Maintenance of stereotypical vocalizations by adult zebra finches. J. Neurosci. 4: 2387-2396 Bottjer SW, Brady JD, Cribbs B (2000) Connections of a motor cortical region in zebra finches: relation to pathways for vocal learning. J Comp Neurol 420:244-260 Bottjer SW, Glaessner SL, Arnold AP (1985) Ontogeny of brain nuclei controlling song learning and behavior in zebra finches. J.Neurosci. 5: 1556-1562 Bottjer SW, Halsema KA, Brown SA, Miesner EA (1989) Axonal connections of a forebrain nucleus involved wiht vocal learning in zebra finches. J.Comp.Neurol. 279:312-326 Bottjer SW, Johnson F (1997) Circuits, Hormones, and Learning: vocal Behavior in Songbirds. J. Neurobiol. 33:602-618 Bottjer SW, Miesner EA, Arnold AP (1984) Forebrain lesions disrupt development but not maintenance of song in passerine birds. Science 224:901-903 Bottjer SW, Miesner EA, Arnold AP (1986) Changes in neuronal number, density and size account for increases in volume of song-control nuclei during song development in zebra finches. Neuroscience Letters 67:263-268 Bottjer SW, Sengelaub DR (1989) Cell death during development of a forebrain nucleus involved with vocal learning in zebra finches. J.Neurobiol. 20:609-618 Braaten RF, Reynolds K (1999) Auditory preference for conspecific song in isolation-reared zebra finches. Animal Behaviour 58:105-111 Brainard MS (2004) Contributions of the anterior forebrain pathway to vocal plasticity. Ann. N. Y. Acad. Sci. 1016:377-394 Brainard MS, Doupe AJ (1997) Anterior forebrain in lesions eliminate deafening induced song deterioration in adult zebra finches. Soc. Nuerosci. Abstr. 23:796 Brainard MS, Doupe AJ (2000a) Auditory feedback in learning and maintenance of vocal behaviour. Nature Neurosci. Reviews 1:31-40 Brainard MS, Doupe AJ (2000b) Interruption of a basal ganglia-forebrain circuit prevents plasticity of learned vocalizations. Nature 404:762-766 Brown ES, Woolston J, Frol A, Bobadilla L, Khan DA, Hanczyc M, Rush AJ, Fleckenstein J, Babcock E, Cullum CM (2004) Hippocampal volume, spectroscopy, cognition, and mood in patients receiving, corticosteroid therapy. Biol Psychiatry 55:538-545 Burek MJ, Nordeen KW, Nordeen EJ (1991) Neuron loss and addition in developing zebra finch song nuclei are independent of auditory experience during song learning. J.Neurobiol. 22:215-223 Burek MJ, Nordeen KW, Nordeen EJ (1993) Ontogeny of sex differences among HVC projection neurons. Soc. Neurosci. Abstr. 19: 1312 Cammermeyer J (1978) Is the solitary dark neuron a manifestation of postmortem trauma to the brain inadequately fixed by perfusion? J. Histochem. 56:97-115 Canady RA, Burd GD, DeVoogd TJ, Nottebohm F (1988) Effect of testosterone on input received by an identified neuron type of the canary song system: a Golgi / electron microscopy / degeneration study. J. Neurosci. 8: 3770-3784 Changeux JP, Danchin A (1976) Selective stabilization of developing synapses as a mechanism for the specification of neuronal networks. Nature 264:705-771 Clayton DF (1997) Role of gene regulation in song circuit development and song learning. J. Neurobiol. 33:549-571 Clayton NS (1989 ) The effects of cross-fostering on selective song learning in estrildid finches. Behaviour 109:163-175 Clayton NS (1998) Memory and the hippocampus in food-storing birds: a comparative approach. Neuropharmacology 37:441-452 Coggeshall RE (1992) A consideration of neural counting methods. Trends Neurosci 15:9-13 Coggeshall RE, Lekan HA (1996) Methods for determining numbers of cells and synapses: a case for more uniform standarts of review. The Journal of Comparative Neurology 364:6-15 Cole AJ, Saffen DW, Barban JM, Worley PF (1989) Rapid increase of an immediate early gene messenger RNA in hippocampal neurons by synaptic NMDA receptor activation. Nature 340:474-476 Cynx J (1990) Experimental determination of a unit of song production in the zebra finch (Taeniopygia guttata). J.Comp.Psychol. 104:3-10 Cynx J, Nottebohm F (1992) Role of gender, season and familiarity in discrimination of conspecific song by zebra finches (Taeniopygia guttata). Proc.Natl.Acad.Sci.USA 89:1368-1371 DeVoogd TJ, Cardin JA, Szekeley T, Büki J, Newman SW (1996) Relative volume of LMAN in female wrablers species varies with the number of songs produced by conspecific males. Soc. Neurosci. Abstr. 22:1401 DeVoogd TJ, Krebs JR, Healy SD, Purvis A (1993) Relations between song repertoire size and the volume of brain nuclei related to song: comparative evolutionary analyses amongst oscine birds. Proc R Soc Lond B Biol Sci 254:75-82 DeVoogd TJ, Nottebohm F (1981a) Gonadal hormones induce dendritic growth in the adult avian brain. Science 214:202-4 DeVoogd TJ, Nottebohm F (1981b) Sex differences in dendritic morphology of a song control nucleus in the canary: a quantitative Golgi study. J comp Neurol 196:309-316 Doupe AJ, Brainard MS, Hessler NA (1999) The song system: neural circuits essential throughout life for vocal behavior and plasticity. In: Gazzaniga MS ( ed ) The new cognitive neuronsciences, second edition edn , pp 451-467 Doupe AJ (1997) Song selective and order selective neurons in the songbird anterior forebrain emerge during vocal development. Biomed Res 18:17-37 Doupe AJ, Konishi M (1991) Song-selective auditory circuits in the vocal control system of the zebra finch. Proc.Natl.Acad.Sci.U.S.A. 88:11339-11343 Doupe AJ, Kuhl PK (1999) Birdsong and human speech common themes and mechanisms. Annu. Rev. Neurosci. 22:567-631 Doupe AJ, Solis MM (1997) Song- and order-selective neurons develop in the songbird anterior forebrain during vocal learning. J. Neurobiol. 33:694-709 Doupe AJ, Solis MM, Kimpo R, Boettiger CA (2004) Cellular, circuit, and synaptic mechanisms in song learning. Ann. N. Y. Acad. Sci. 1016:495-523 Doya K, Sejnowski T (1995) A novel reinforcement model of birdsong vocalization learning. In: Tesauro G, Touretzky DS, Leen TK (eds) Advances in Neural Information Processing Systems, vol 7. MIT Press,

Camebridge

, pp 101-108 Driscoll I, Hamilton DA, Petropoulos H, Yeo RA, Brooks WM, Baumgartner RN, Sutherland RJ (2003) The aging hippocampus: cognitive, biochemical and structural findings. Cereb Cortex. 13:1344-51 Eales LA (1985) Song learning in zebra finches: some effects of song modes availability on what is learnt and when. Anim.Behav. 33:1295-1300 Eales LA (1987) Song learning in female-raised zebra finches: another look at the sensitive phase. Anim.Behav. 35:1356-1365 Eales LA (1989) The influences of visual and vocal interaction on song learning in zebra finches. Anim. Behav. 37:507-508 Ehret G (1996) Auditorische Systeme. In: Dudel J, Menzel R, Schmidt RF ( eds ) Neurowissenschaft. Springer, pp 353-379 Farries MA (2004) The avian song system in comparative perspective. Ann. N. Y. Acad. Sci. 1016:61-76 Farries MA, Perkel DJ (2000) Electrophysiological properties of avian basal ganglia neurons recorded in vitro. J Neurophysiol 84:2502-13. Fortune ES, Margoliash D (1992) Cytoarchitectonic organization and morphology of cells of the field L complex in male zebra finches (Taenopygia guttata). J.Comp.Neurol. 325:388-404 Fortune ES, Margoliash D (1995) Parallel pathways and convergence onto HVC and adjacent neostriatum of adult zebra finches (Taeniopygia guttata). J. Comp. Neurol. 360:413-441 Foster EF, Bottjer SW (1992) Axonal connections of a forebrain nucleus in male zebra finches. Soc. Neurosci. Abstr.:18: 528 Foster EF, Bottjer SW (1993) Lesion of mMAN produce slight disruption in vocal behavior of juvenile male zebra finches. Soc. Neurosci. Abstr. 19: 1016 Foster EF, Mehta RP, Bottjer SW (1997) Axonal connections of the medial magnocellular nucleus of the anterior neostriatum in zebra finches. J. Comp. Neurol. 382:364-381 Garamszegi LZ, Eens M (2004) Brain space for a learned task: strong intraspecific evidence for neural correlates of singing behavior in songbirds. Brain Res Brain Res Rev. 44:187-93 Gehr DD, Hofer SB, Marquardt D, Leppelsack H (2000) Functional changes in field L complex during song development of juvenile male zebra finches. Brain Res Dev Brain Res 125:153-65 Grzimek B (1980) Grzimeks Tierleben, Enzyklopädie des Tierreichs, vol Band 9. Deutscher Taschenbuch Verlag GmbH Gundersen HJ, Jensen EB, Kieu K, Nielsen J (1999) The efficiency of systematic sampling in stereology-reconsidered. J. Microsc. 193:199-211 Gurney ME ( 1981 ) Hormonal control of cell form and number in the zebra finch song system. J.Neurosci. 1: 658-673 Gurney ME (1982) Behavioral correlates of sexual differentiation in the zebra finch song system. Brain Res. 231:153-172 Gurney ME, Konishi M (1980) Hormone induced sexual differntiation of brain and behavior in zebra ficnhes. Science 208:1380-1382 Halsema KA, Bottjer SW (1992) Chemical lesions of a thalamic nucleus disrupt song development in male zebra finches. Soc Neurosci Abstr 18: 529 Hamilton KS, King AP, Sengelaub DR, West MJ (1997) A brain of her own: a neural correlate of song assessment in female songbird. Neurobiology of learning and memory 68:325-332 Hamilton KS, King AP, Sengelaub DR, West MJ (1998) Visual and song nuclei correlate with courtship skills in brown-headed cowbirds. Animal Behaviour 56:973-982 Hedreen JC (1998a) Lost caps in histological counting methods. Anat. Rec. 250:366-372 Hedreen JC (1998b) What was wrong with the Abercrombie and empirical cell counting mehtods? Anat. Rec. 250:373-380 Herrmann K, Arnold AP (1991a) The development of afferent projections to the robust archistriatal nucleus in male zebra finches: a quantitative electron microscopic study. J.Neurosci. 11:2063-2074 Herrmann K, Arnold AP (1991b) Lesions of HVc block the developmental masculinizing effects of estradiol in the female zebra finch song system. J.Neurobiol. 22:29-39 Herrmann K, Bischof HJ (1986) Delayed development of song control nuclei in the zebra finch is related to behavioral development. J.Comp.Neurol. 245:167-175 Hessler NA, Doupe AJ (1999) Singing-related neural activity in a dorsal forebrain-basal ganlia circuit of adult zebra finches. J. Neurosci. 19:10461-10481 Hessler NH, Doupe AJ (1997) Singing-related activity in anterior forebrain nuclei of adult zebra finch. Soc. Neurosci. Abstr. 23:245 Houx BB, Ten Cate C (1998) Do contingencies with tutor behaviour influence song learning in zebra finches? Behaviour 135:599-614 Immelmann K (1969) Song development in the zebrea finch and other estrildid finches. In: Hinde RA ( ed ) Bird vocalizations: Their relation to current problems in biology and psychology. Cambridge Univ. Press,

Cambridge

, pp 61-74 Iyengar S, Bottjer SW (2002a) Development of individual axon arbors in a thalamocortical circuit necessary for song learning in zebra finches. The Journal of Neuroscience 22 :901-911 Iyengar S, Bottjer SW (2002b) The role of auditory experience in the formation of neuronal circuits underlying vocal learning in zebra finches. The Journal of Neuroscience 22:946-958 Iyengar S, Viswanathan SS, Bottjer SW (1999) Development of topography within song control circuitry of zebra finches during the sensitive period for song learning. J. Neurosci. 19:6037-6057 Jacobs EC, Grisham W, Arnold AP (1995) Lack of a synergistic effect between estradiol and dihydrotestosteron in the masculization of the zebra finch song system. J. Neurol. 27:513-519 Janata P, Margoliash D (1999) Gradual emergence of song selectivity in sensorimotor structures of the male zebra finch song system. J Neurosci 19:5108-18 Jarvis ED (2004) Learned birdsong and the neurobiology of human language. Ann. N. Y. Acad. Sci. 1016:749-777 Jarvis ED, Nottebohm F (1997) Motor-driven gene expression. Proc. Natl. Acad. Sci. USA 94:4097-4102 Jarvis ED, Scharff C, Grossman MR, Ramos JA, Nottebohm F (1998) For whom the bird sings: context-dependent gene expression. Neuron 21:775-788 Johnson F (1998) Disruption of song learning following lMAN lesions in fledgling male zebra finches. Congress of Neuroethology:301 Johnson F, Bottjer SW ( 1992 ) Growth and regression of thalamic efferents in the song-control system of male zebra finches. J.Comp.Neurol. 326:442-450 Johnson F, Bottjer SW (1994) Afferent influences on cell death and birth during development of a cortical nucleus necessary for learned vocal behavior in zebra finches. Development 120:13-24 Johnson F, Jones C, Taylor P, Thompson J, Turner S (2003) Does vocal practice contribute to individual differnences in brain space for singing in zebra finches? Soc. Neurosci. Abstr. 33: 294.8 Johnson F, Sablan MM, Bottjer SW (1995) Topographic organization of a forebrain pathway involved with vocal learning in zebra finches. J.Comp.Neurol. 358:260-278 Johnson F, Sellix M (2000) Reorganisation of an telencephalic motor region during sexual differentiation and vocal learning in zebra finches. Brain Res Dev Brain Res 121:253-63 Jones AE, Ten Cate C, Slater PJB (1996) Early experience and plasticity of song in adult male zebra finches (Taeniopygia guttata). Journal of Comparative Psychology 110:354-369 Kafitz KW, Herth G, Bartsch U, Güttinger HR, Schachner M (1992) Application of testosterone accelerates oligodendrocyte maturation in brains of zebra finches. NeuroReport 3: 315-318 Kandel ER, Schwartz JH, Jessell TM (2000) Principles of Neuronal Science, 4 edn. Appleton & Lange Karnovsky MH (1965) A formaldehyde - glutaraldehyde fixative of high osmolarity for use in electron microscopy. J. Cell Biol. 27: 137 A Karten HJ (1991) Homology and evolutionary origins of the "neocortex". Brain. Behav. Evol. 38:264-272 Kelley DB, Nottebohm F (1979) Projections of a telencephalic auditory nucleus -field L- in the canary. J. Comp. Neurol. 183:445-470 Kimpo RR, Doupe AJ (1997) FOS is induced by singing in distinct neuronal populations in a motor network. Neuron 18:315-25 Kirn JR, DeVoogd TJ (1989) Genesis and death of vocal control neurons during sexual differentiation in the zebra finch. J.Neurosci. 9: 3176-3187 Kreck G, Nixdorf-Bergweiler BE (1999) Evidence for the existence of an anterior forebrain pathway in female zebra finches identified by in vivo and in vitro tracing techniques. Proc. of the 27th Göttingen Neurobiology Conference:280 Kreck G, Nixdorf-Bergweiler BE (2005) Evidence for a cortical-basal ganglia projection pathway in female zebra finches. Neuroreport 16: 21-24 Konishi M (1965a) Effects of deafening on song development in American robins and black-headed grosbeaks. Z. Tiepsychol. 22:584-599 Konishi M (1965b) The role of auditory feedback in the control of vocalization in the white crowned sparrow. Z. Tierpsychol. 22:770-783 Konishi M (1985) Birdsong: from behavior to neuron. Ann. Rev. Neurosci.:125-170 Konishi M (2004) The role of auditory feedback in birdsong. Ann. N. Y. Acad. Sci. 1016:463-475 Konishi M, Akutagawa E (1985) Neuronal growth, atrophy and death in a sexually dimorphic song nucleus in the zebra finch brain. Nature 315:145-147 Konishi M, Akutagawa E (1987) Hormonal control of cell death in a sexually dimorphic song nucleus in the zebra finch. Ciba.Found.Symp. 126:173-185 Korsia S, Bottjer SW (1991) Chronic testosterone treatment impairs vocal learning in male zebra finches during a restricted period of development. J.Neurosci. 11:2362-2371 Kubota M, Saito N (1991) NMDA receptors participate differentially in two different synaptic inputs in neurons of the zebra finch robust nucleus of the archistriatum in vitro. Neurosci.Lett. 125:107-109 Lee DW, Smith GT, Tramontin AD, Soma KK, Brenowitz EA, Clayton NS (2001) Hippocampal volume does not change seasonally in a non food-storing songbird. Neuroreport 12:1925-8 Leonardo A, Konishi M (1999) Decrystallization of adult birdsong by perturbation of auditory feedback. Nature 399:466-470 Li R, Zuo MX, Sakaguchi H (1999) Auditory-vocal cholinergic pathway in zebra finch brain. Neuroreport 10:165-9 Lombardino AJ, Nottebohm F (2000) Age at deafening affects the stability of learned song in adult male zebra finches. J Neurosci 20:5054-5064 Luo M, Ding L, Perkel DJ (1999) Topographic mapping throughout the anterior forebrain pathway of the zebra finch song system. Soc. Neurosci. Abstr. 25: 1367 Luo M, Ding L, Perkel DJ (2001) An avian basal ganglia pathway essential for vocal learning forms a closed topographic loop. The Journal of Neuroscience 21:6836-6845 Luo M, Perkel DJ (1999) Long-range GABAergic projection in a circuit essential for vocal learning. Journal of Comparative Neurology 403:68-84 MacDougall-Shackleton SA, Hulse SH, Ball GF (1998a) Neural correlates of singing behavior in male zebra finches (Taeniopygia guttata). J Neurobiol 36:412-430 MacDougall-Shackleton SA, Hulse SH, Ball GF (1998b) Neural bases of song preferences in female zebra finches (Taeniopygia guttata). Neuroreport 9: 3047-52 Maekawa M (1997) Song note selective responses of the neurons in a song control nucleus of the zebra finch. Biomed Res 18:39-43 Margoliash D (1997) Functional organization of forebrain pathways for song production and perception. J. Neurobiol. 33:671-693 Marler P (1970) A comparative approach to vocal learning: song development in white-crowned sparrows. J Comp Physiol Psychol 71:1-25 Marler P (1991a) The instinct to learn. In: Gelman SCR ( ed ) The Epigenesis of Mind: Essays on Biology and Cognition. L. Erlbaum, Hillsdale,

New Jersey,

pp 37-66 Marler P (1991b) Song-learning behavior: the interface with neuroethology. TINS 14:199-206 Marler P (1992) Communication: behavior and neurobiology. seminars in The Neurosciences 4: 373-375 Marler P, Peters S (1987) A sensitive period for song acquisition in the song sparrow: a case of agelimited learning. Ethology 76:89-100 McCasland JP, Konishi M (1981) Interaction between auditory and motor activities in an avian songcontrol nuceus. Proc Natl Acad. Sci. US 78:7815-7819 Mello CV, Clayton DF (1994) Song-induced ZENK gene expression in auditory pathways of song bird brain and its relation to the song control system. J. Neurosci. 14:6652-6666 Mello CV, Vates GE, Okuhata S, Nottebohm F (1998) Descending auditory pathways in the adult male zebra finch (Taeniopygia guttata). J Comp Neurol 395:137-60 Miller DB (1979) Long-term recognition of father`s song by female zebra finches. Nature 280:389-391 Mooney R (1992) Synaptic basis for developmental plasticity in a birdsong nucleus. J.Neurosci. 12:2464-2477 Mooney R, Konishi M (1991) Two distinct inputs to an avian song nucleus activate different glutamate receptor subtypes on individual neurons. Proc.Natl.Acad.Sci.USA 88:4075-4079 Mooney R, Rao M (1994) Waiting periods versus early innervation: the development of axonal connections in the zebra finch song system. J.Neurosci. 14:6532-6543 Morrison RG, Nottebohm F (1993) Role of a telencephalic nucleus in the delayed song learning of socially isolated zebra finches. J.Neurobiol. 24:1045-1064 Nixdorf BE, Davis SS, DeVoogd TJ (1989) Mophology of Golgi-impregnated neurons in hyperstriatum ventralis, pars caudalis in adult male and female canaries. J Comp Neurol 284:337-349. Nixdorf-Bergweiler BE, Lips MB, Heinemann U (1995a) Electrophysiological and morphological evidence for a new projection of lMAN-neurons towards area X. NeuroReport 6: 1729-1732 Nixdorf-Bergweiler BE, Wallhäusser-Franke E, DeVoogd TJ (1995b) Regressive development in neuronal structure during song learning in birds. J Neurobiol 27:204-215. Nixdorf-Bergweiler BE (1996) Divergent and parallel development in volume sizes of telencephalic song control regions in male and female zebra finches. J.Neurobiol. 375:445-456 Nixdorf-Bergweiler BE (1997) Nucleolar size and shape is sexually dimorphic in neurons of lateral magnocellular nucleus of the anterior neostriatum (LMAN) in birds. Cytogenetics and Cell Genetics 76:77-86. Nixdorf-Bergweiler BE (1998) Enlargement of neuronal somata in the LMAN coincides with the onset of sensorimotor learning for song. Neurobiology of learning and memory 69:258-273 Nixdorf-Bergweiler BE, Bindrich A, Freyer C, Hintz V (1999a) Neuronal and behavioral effects of deprivation of memory formating for song are multifaceted. Soc. Neurosci. Abstr. 29: 752.7 Nixdorf-Bergweiler BE, Bindrich A, Frommolt KH (1999b) Are introductory notes in zebra finches song genetically determined? IBAC Nixdorf-Bergweiler BE (2001) Lateral magnocellular nucleus of the anterior neostriatum (LMAN) in the zebra finch: neuronal connectivity and the emergence of sex differences in cell morphology. Microsc Res Tech 54:335-53 Nixdorf-Bergweiler BE, von Bohlen und Halbach V (2004) Major sex differences in the development of myelination are prominent in song system nucleus HVC but not lMAN. Animal Biology 54:27-43 Nixdorf-Bergweiler BE, von Bohlen und Halbach V (2005) Neuronal number in song nucleus LMAN remains high in female zebra finches throughout development and in adulthood. Acta Zool Sincica (in press) Nordeen EJ, Grace A , Burek MJ, Nordeen KW (1992) Sex-dependent loss of projection neurons involved in avian song learning. J.Neurobiol. 23:671-679 Nordeen EJ, Nordeen KW (1988a) Sex and regional differences in the incorporation of neurons born during song learning in zebra finches. J. Neurosci. 8:2869-2874 Nordeen EJ and Nordeen KW (1989) Estrogen stimulates the incorporation of new neurons into avian song nuclei during adolescence. Dev. Brain Res.49:27-32 Nordeen EJ, Nordeen KW (1996) Sex difference among nonneuronal cells precedes sexually dimorphic neuron growth and survival in an avian song control nucleus. Journal of Neurobiology 30:531-542 Nordeen EJ, Nordeen KW, Arnold AP (1987a) Sexual differentiation of androgen accumulation within the zebra finch brain through selective cell loss and addition. J.Comp.Neurol. 259:393-399 Nordeen KW, Nordeen EJ, Arnold AP (1986) Estrogen establishes sex differences in androgen accumulation in zebra finch brain. J. Neuroscience 6: 734-738 Nordeen KW, Marler P, Nordeen EJ (1989) Addition of song-related neurons in swamp sparrows coincides with memorization, not production, of learned songs. J.Neurobiol. 20:651-661 Nordeen KW, Nordeen EJ (1988b) Projection neurons within a vocal motor pathway are born during song learning in zebra finches. Nature 334:149-151 Nordeen KW, Nordeen EJ (1992) Auditory feedback is necessary for the maintenance of stereotyped song in adult zebra finches. Behav.Neural.Biol. 57:58-66 Nordeen KW, Nordeen EJ (1993) Long-term maintance of song in adult zebra finches is not affected by lesions of a forebrain region involved in song learning. Behavioral and neural Biology 57:79-82 Nordeen KW, Nordeen EJ (1997) Anatomical and synaptic substrates for avian song learning. J. Neurobiol. 33:532-548 Nordeen KW, Nordeen EJ, Arnold AP (1986) Estrogen establishes sex differences in androgen accumulation in zebra finch brain. J.Neurosci. 6: 734-738 Nordeen KW, Nordeen EJ, Arnold AP (1987b) Estrogen accumulation in zebra finch song control nuclei: implications for sexual differentiation and adult activation of song behavior. J.Neurobiol. 18:569-582 Nottebohm F (1981) A brain for all seasons: cyclical anatomical changes in song control areas of the songbird brain. Science 214:1368-1370 Nottebohm F (1985) Neuronal replacement in adulthood. In: Nottebohm F ( ed ) Hope for a New Neuorlogy.

New York

Academy Science, New York, pp 143-161 Nottebohm F (1987) Birdsong. In: Adelman G (ed) Incyclopedia of Neuroscience, vol I. Birkhauser Boston,

Boston

, pp 133-136 Nottebohm F (1989) Vom Vogelgesang zur Bildung neuer Nervenzellen. Spektrum der Wissenschaft 4: 112-117 Nottebohm F (1993) The search for neural mechanisms that define the sensitive period for song learning in birds. Netherlands Journal of Zoology 43:193-234 Nottebohm F, Arnold AP (1976) Sexual dimorphism in vocal control areas of the songbird brain. Science 194:211-213 Nottebohm F, Kasparian S, Pandazis C (1981) Brain space for a learned task. Brain Res. 213:99-109 Nottebohm F, Kelley DB, Paton JA (1982) Connections of vocal control nuclei in the canary telencephalon. J. Comp. Neurol. 207:344-357 Nottebohm F, Nottebohm ME, Crane L (1986) Developmental and seasonal changes in canary song and their relation to changes in the anatomy of song-control nuclei. Behav.Neural Biol. 46:445-471 Nottebohm F, Stokes TM, Leonard CM (1976) Central control of song in the canary Serinus canarius. J. Comp. Neurol. 165:457-486 Okuhata S, Saito N (1987) Synaptic connections of thalamo-cerebral vocal nuclei of the canary. Brain Res. Bull. 18:35-44 Paton JA, Manogue KR, Nottebohm F (1981) Bilateral organization of the vocal control pathway in the budgerigar. J. Neuroscience 11:1279-1288 Perkel DJ, Farries MA (2000) Comlementary "bottom-up" and "top-down" approaches to basal ganglia function. Current Opinion Neurobiology 10:725-731 Pohl-Apel G, Sossinka R (1984) Hormonal determination of song capacity in females of the zebra finch: critical phase of treatment. Z.Tierpsychol. 64:330-336 Price PH (1979) Developmental determinations of structure in zebra finch song. J. Comp. Physiol. Psychol. 93:260-277 Rehkämper G, Zilles K (1991) Parallel evolution in mammalian and avian brains: Comparative cytoarchitectonic and cytochemical analysis. Cell Tissue Res. 263:3-28 Reiner A, Laverghetta AV, Meade CA, Cuthbertson SL, Bottjer SW (2004) An immunohistochemical and pathway tracing study of the striatopallidal organization of area X i the male zebra finch. J. Comparative Neurology 469:239-261 Reiner A, Perkel DJ, Bruce LL, Butler AB, Csillag A, Kuenzel W, Medina L, Paxinos G, Shimizu T, Striedter G, Wild M (2004) Revised nomenclatur for avian telencephalon and some related brainstem nuclei. J. Comp. Neurol. 473:377-414 Riebel K (2003) Developmental influences on auditory perception in female zebra finches - is there a sensitive phase for song preference learning? Animal. Biol. 53:73-87 Riebel K, Smallegange IM, Terpstra NJ, Bolhuis JJ (2002) Sexual equality in zebra finch song preference: evidence for a dissociation between song recognition and production learning. Proc R Soc Lond B Biol Sci 269:729-33. Rollenhagen A, Bischof HJ (1991) Rearing conditions affect neuron morphology in a telencephalic area of the zebra finch. Neuroreport 2: 711-4 Rosen MJ, Mooney R (2000) Intrinsic and extrinsic contributions to auditory selectivity in a song nucleus critical for vocal plasticity. J Neurosci 20:5437-5448 Ryan SM, Arnold AP (1981) Evidence for cholinergic participation in the control of bird song; acetylcholinesterase distribution and muscarinic receptor autoradiography in the zebra finch brain. J.Comp.Neurol. 202:211-219 Saini KD, Leppelsack HJ (1981) Cell types of the auditory caudomedial neostriatum of the starling (Sturnus vulgaris). J. Comp. Neurol. 198:209-229 Saito N, Maekawa M (1993) Birdsong: The interface with human language. Brain Dev. 15:31-40 Sakaguchi H (1997) Molecular Mechanisms of Plasticity in the Avian Brain. Biomed Res 18:71-78 Sakaguchi H, Yamaguchi A (1997) Early song-deprivation affects the expression of protein kinase C in the song control nuclei of the zebra finch during a sensitive period of song learning. Neuroreport 8: 2645-50 Scharff C, Nottebohm F (1991) A comparative study of the behavioral deficits following lesions of various parts of the zebra finch song system: implications for vocal learning. J.Neurosci. 11:2896-2913 Scharff C, Nottebohm F, Cynx J (1998) Conspecific and heterospesific song discrimination in male zebra finches with lesions in the anterior forebrain pathway. J Neurobiol 36:81-90 Scharff C, Kirn JR, Grossman M, Macklis JD, Nottebohm F (2000) Targeted neuronal death affects neuronal replacement and vocal behavior in adult songbirds. Neuron 25:481-492 Schmidt MF (1996) Neural responses to auditory feedback during singing in Field L of adult zebra finches. Soc. Neurosci. Abstr. 22 Sellix MT, Johnson F ( 1999 ) Reorganization of a motor cortical region during song learning in the zebra finch. Society for Neuroscience 25:1367 Shepherd GM (1994) Neurobiology. Oxford University Press,

New York - Oxford

Simpson HB, Vicario DS (1990) Brain pathways for learned and unlearned vocalizations differ in zebra finches. J.Neurosci. 10:1541-1556 Simpson HB, Vicario DS (1991a) Early estrogen treatment alone causes female zebra finches to produce learned, male-like vocalizations. J.Neurobiol. 22:755-776 Simpson HB, Vicario DS (1991b) Early estrogen treatment of female zebra finches masculinizes the brain pathway for learned vocalizations. J.Neurobiol. 22:777-793 Slater PJB (1974) Bouts and gaps in the behaviour of zebra finches, with special referece to preening. Rev. Comp. Animal. 8: 47-61 Slater PJB, Eales LA, Clayton NS (1988) Song learning in zebra finches (Taeniopygia guttata): progress and prospects. Department of Biology and preclinical medicine University of St.Andrews 0:0 Slater PJB , Jones A, Ten Cate C (1993) Can lack of experience delay the end of the sensitive phase for song learning ? Netherlands Journal of Zoology 43: 80-90 Sohrabji F, Nordeen EJ, Nordeen KW (1990) Selective impairment of song learning following lesions of a forebrain nucleus in the juvenile zebra finch. Behav.Neural.Biol. 53:51-63 Sohrabji F, Nordeen KW, Nordeen EJ (1989) Projections of androgen-accumulating neurons in a nucleus controlling avian song. Brain Res. 448:253-259 Solis MM, Doupe AJ (2000 ) Compromised neural selectivity for song in birds with impaired sensorimotor learning. Neuron 25:109-121 Sossinka R, Böhner J (1980) Song types in the zebra finch Poephila guttata castanotis. Z. Tierpsychol. 53:123-132 Spiro JE, Dalva MB, Mooney R (1999) Long-range inhibition within the zebra finch song nucleus RA can coordinate the firing of multiple projection neurons. J Neurophysiol 81:3007-20 Starkman MN, Giordani B, Gebarski SS, Schteingart DE (2003) Improvement in learning associated with increase in hippocampal formation volume. Biol Psychiatry 53:233-238 Sterio DC (1984) The unbiased estimation of number and sizes of arbitrary particles using the disector. J Microscopy 134:127 -136 Stocker S, Güttinger HR, Herth G (1994) Exogenous testosterone differentially affects myelination and neurone soma sizes in the brain of canaries. NeuroReport 5: 1449-1452 Thorpe WH (1958) The learning of song in birds, with special reference to the song of the chaffinch Fringilla coelebs. Zool. Jb. Syst. 100:535-570 Thorpe WH (1961) Bird Song. Cambridge University Press,

Cambridge

Tramontin AD, Smith GT, Breuner CW, Brenowitz EA ( 1998) Seasonal plasticity and sexual dimorphism in the avian song control system: Stereological measurement of neuron density and number. J Comp Neurol 396:186-192 Tramontin AD, Hartman VN, Brenowitz EA (1999) Rapid and sequential growth of adult avian song nuclei in response to seasonal cues. Soc. Neurosci. Abstr. 25: 864 van der Knaap MS, Valk J, Bakker CJ, Schooneveld M, Faber JA, Willemse J, Gooskens RH (1991) Myelination as an expression of the functional maturity of the brain. Dev-Med-Child-Neurol. 33:849-57 Vates GE, Broome BM, Mello CV, Nottebohm F (1996) Auditory pathways of caudal telencephalon and their relation to the song system of adult male zebra finches (Taeniopygia guttata). J.Comp.Neurol. 366:613-642 Vates GE, Nottebohm F (1995) Feedback circuitry within a song-learning pathway. Proc.Natl.Acad.Sci.USA 92:5139-5143 Vates GE, Vicario DS, Nottebohm F (1997) Reafferent thalamo- "cortical" loops in the song system of oscine songbirds. The Journal of Comparative Neurology 380:275-290 Vicario DS (1991) Organization of the zebra finch song control system: II. Functional organization of outputs from nucleus Robustus archistriatalis. J.Comp.Neurol. 309:486-494 Vicario DS (1993) A new brain stem pathway for vocal control in the zebra finch song system. Neuroreport. 4: 983-986 Vicario DS (1994) Motor mechanisms relevant to auditory-vocal interactions in songbirds. Brain Behav Evol 44:265-278 Vicario DS, Nottebohm F (1988) Organization of the zebra finch song control system: I. Representation of syringeal muscles in the hypoglossal nucleus. J.Comp.Neurol. 271:346-354 Vicario DS, Raksin JN (2000) Possible roles for GABAergic inhibition in the vocal control system of the zebra finch. Neuroreport 11:3631-5. Volman SF, Khanna H (1995) Convergence of untutored song in group-reared zebra finches (Taeniopygia guttata). J.Comp.Psychol. 109:211-221 von Bartheld CS (2002) Counting particles in tissue sections: choices of methods and importance of calibration to minimize biases. Histol. Histopathol. 17:639-648 Vu ET, Mazurek MW, Kuo YC (1994) Identification of a forebrain motor programming network for the learned song of zebra finches. J.Neurosci. 14:6924-6934 Wallhäusser E, Scheich H (1987) Auditory imprinting leads to differential 2-deoxyglucose uptake and dendritic spine loss in the chick rostral forebrain. Brain Res 428:29-44. Wallhäusser-Franke E, Nixdorf Bergweiler BE, DeVoogd TJ (1995) Song isolation is associated with maintaining high spine frequencies on zebra finch LMAN neurons. Neurobiol.Learn.Mem. 64:25-35 Wang J, Sakaguchi H, Sokabe M (1999) Sex differences in the vocal motor pathway of the zebra finch revealed by real-time optical imaging technique. NeuroReport 10:2487-2491 Ward BC, Nordeen EJ, Nordeen KW (1998) Individual variation in neuron number predicts differences in the propensity for avian vocal imitation. Proc Nat Acad Sci USA 95:1277-1282 West MJ (1993) New stereological methods for counting neurons. Neurobiol. Aging 14:275-285 West MJ (1999) Stereological methods for estimating the total number of neurons and synapses: issues of precision and bias. TINS 22:51-61 West MJ, Slomanka L (2001) 2-D versus 3-D cell counting - a debate. What is an optical disector? TINS 24:374-380 Whitney O, Johnson F (2003) Singing - driven zenk expression: regulation by auditory feedback during zebra finch vocal learning. Soc. Neurosci. Abstr. 33:522.6 Whitney O, Soderstrom K, Johnson F (2000) Post-transcriptional regulation of zenk expression associated with zebra finch vocal development. Brain Res Mol Brain Res 80:279-90. Wild JM (1993a) Descending projections of the songbird nucleus robustus archistriatalis. The Journal of Comparative Neurology 338:225-241 Wild JM (1993b) The avian nucleus retroambigualis: a nucleus for breathing, singing and calling. Brain Research 606:319-324 Wild JM (1994) The auditory-vocal-respiratory axis in birds. Brain Behav. Evol. 44:192-209 Wild JM (1997) Neural pathways for the control of birdsong production. J. Neurobiol. 33:653-670 Williams H (1985) Sexual dimorphism of auditory activity in the zebra finch song system. Behav.Neural.Biol. 44:470-484 Williams H (1989) Multiple representations and auditory-motor interactions in the avian song system. Ann NY Acad Sci 563: 148-164 Williams H, Cynx J, Nottebohm F (1989) Timbre control in zebra finch (Taeniopygia guttata) song syllables. J.Comp.Psychol. 103:366-380 Williams H, Metha N (1999) Changes in adult zebra finch song require a forebrain nucleus that is not necessary for song production. J Neurobiol 39:14-28 Williams H, Staples K (1992) Syllable chunking in zebra finch (Taeniopygia guttata) song. J.Comp.Psychol. 106:278-286 Wisden W, Errington ML, Williams S, Dunnett SB, Waters C, Hitchcock D, Evan G, Bliss TV, Hunt SP (1990) Differential expression of immediate early genes in the hippocampus and spinal cord. Neuron 4:603-614 Zann R (1984) Structural variation in the zebra finch distance call. Z. Tierpsychol. 66:328-345 Zann R (1985) Ontogeny of the zebra finch distance call: I) Effects of cross-fostering to bengalese finches. Z. Tierpsychol 68:1 ,23 Zann R (1996) The Zebra Finch. Oxford University Press