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Distributed Representation N the Song System of Oscines

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Distributed Representation N the Song System of Oscines Brain,Behavior and Evolution Editor-in-Chief: R. Glenn Northcutt. La Jolla. Calif. Reprint Publisher: S.Karger AG, Basel Printed in Switzerland Brain Behav Evol 1994:44:247264 Daniel Margoliash" Eric S. Fortune' DistributedRepresentation in the MitchellL. Sutterh AlbertC. Yu' SongSystem of Oscines: B. DavidWren-Hardin'' A. Dave' Evolutionarylmplications and ' Departmentof OrganismalBiology and Anatomy, The University of Chicago, FunctionalConsequences Chicago,Ill.; b Center for Neuroscience, University of Califomia, Davis, Calif.; " Committee on Neurobiology, The University of Chicago,Chicago, Ill., USA Key Words Abstract Hierarchical This paper reviews the organizationalprinciples and implications that have Parallel emergedfrom the analysisof HVc, a forebrainnucleus that is a major site of sen- Distributed sory, motor, and sensorimotorintegration in the song control systemof oscine Redundant passerinebirds (songbirds). Anatomical, physiological, and behavioral data sup- Sensorimotorintegration port the conclusion that HVc exists within a hierarchically organizedsystem Forebrainevolution with parallelpathways that convergeonto HVc. The organizationof HVc is dis- Bird songsystem tributed and redundant,and its outputsexhibit broad divergence.A similar pat- tern of connectivityexists for neostriatumadjacent to HVc. This and other data supportthe hypothesisthat the song systemarose from an elaborationor dupli- cation of pathways generally present in all birds. Spontaneousand auditory responseactivity is stronglycorrelated throughout HVc, with auditoryresponses exhibiting strongtemporal modulation in a synchronizedfashion throughout the nucleus.This suggeststhat the auditoryrepresentation of songis encodedin the synchronizedtemporal pattems of activation,and that the predominantselectiv- ity for the individual's own songthat is observedfor HVc neuronsresults from interactionsof auditory input with centralpattem generatorsfor song. Most, or all HVc neuronsare recruitedduring singing.The auditory responseand motor recruitmentpropefiies of individual HVc neuronshave no simple relationship, and the spontaneousactivity in HVc may build up in the secondspreceding a song.To the extent HVc participatesin perceptualphenomena associated with song, production and perception are not tightly linked in adults but may be linked by shareddevelopmental processes during periodsof sensorimotorleam- itrg. Dr. Daniel Margoliash O 1994S. KugerAG, Basel Department of Organismal Biology md Anatomy The University of Chicago 1025E. 57rh St. Chicago, IL 60627 (USA) ture with neuronalfunction. In particular,the nucleusHVcl Abbreviations is a major site of integrationin the song systemwhere sen- A Archistriatum sorimotor transformationsrelated to syllable and song Am Nucleus ambiguus structureprobably occur. This paperwill review recentdata 'Autogenous'song AS demonstratinga distributed,time-domain representation for cDLP Nucleus dorsolateralisposterior thalami pars caudalis song in HVc, including anatomicalobservations, physio- CPGs Central pattern generators recordingsin preparations, physio- DIP Nucleusdorsointermediusposteriorthalami logical anesthetized and DLM Nucleus dorsolateralisanterior thalami, pars medialis logical recordingsand lesion studiesin singing birds. The DLP Nucleus dorsolateralisposterior thalami analysis of HVc's anatomy and physiology has yielded DM Dorsomedial subdivision of nucleus intercollicularis insights into the organizationof episodic behaviors,audi- DMP Nucleusdorsomedialis posterior thalami tory temporalpattem processing,sensorimotor integration, HV Hyperstriatum ventrale the relationshipbetween production and perceptionof IMAN Lateral magnocellular nucleus of the anterior neostriatum and mMAN Medial magnocellular nucleus of the anterior neostriatum song.Additionally, we reinterpretthe forebraincircuitry in N Neostriatum and around HVc from an evolutionary perspective,con- NC Neostriatum caudale cluding that the caudal forebrain componentsof the song NCM Caudomedial neostriatum systemare an elaborationof a systemwhich was probably Nd Dorsal caudal neostriatum present NI Neostriatum intermedium in ancestralforms. NIDL Neostriatumintermediumparsdorsolateralis NIf Nucleus interfacialis HVc within the Contextof the SongControl System NIVL Neostriatumintermediumparsventrolateralis The motor system for song production comprisesdis- nXIIts Nucleusnervihypoglossi,tracheosyringealcomponent crete forebrain, midbrain, and brainstemnuclei that sub- OM Tractus occipitomesencephalicus servea limited setof discreteepisodic behaviors (songs and Ov Nucleusovoidalis RA Nucleus robustus archistriatalis calls) (fig. 1). The HVc is a major site of integrationwithin RAm Nucleus retroambigualis the song system and participatesin the two major song SMS Syllable of maximum synchronization system forebrain pathways. One forebrain pathway in- TCS Temporalcombinationsensitive cludesnucleus interface (NIf), HVc, and the robustnucleus TMS Time of maximal synchronization (RA) in the and Uva Nucleus uvaeformts of the archistriatum caudal telencephalon WTS White+hroated sparrows the caudal subdivision of the posterior thalamic nucleus X Area X (cDLP) in the thalamus- previously called nucleusuvae- formis (Uva); seebelow andWild tl993al - andultimately projectsto the brainstemhypoglossal nucleus, which inner- lntroduetion vates the syrinx, the vocal organ in birds. Lesion and chronic recording experimentsdemonstrate that these nu- The richnessof behavioralphenomena associated with clei are involved in moment-to-momentcontrol of song songleaming, production, and perceptionin oscinepasser- production [Nottebohm et al., 1976, 1982; McCasland, ine birds [seeKroodsma and Miller, 1982] is supportedby 1987;Williams andVicario, 19931.A second,more rostral a richnessof neurobiologicalcomplexity of the song con- pathwayinvolves projections between HVc, areaX (X), the trol system.Although the analysisof songsystem circuitry lateral magnocellularnucleus of the anterior neostriatum at the cellular level is incomplete,for somenuclei there is (MAN), and RA in the telencephalonand the medial divi- now sufficientinformation to besin to relateneuronal struc- sion of the dorsolateralthalamic nucleus (DLM) in the tha- lamus.Lesion studiesdemonstrate X. IMAN. and DLM are not necessaryfor songmaintenance in adult birds, but they 1 TheHVc is anucleus originally identified as ventral caudal hyper- implicate thesenuclei in juvenile song development striatum [Nottebohm et al.,1976] but is now thought to be in the dor- [Bott- sal caudal neostriatum [Nottebohm, 1987]. The suggested revision jer et al., 1984;Scharff and Nottebohm,1991; Sohrabji et 'High (or Higher) Vocal Center' (HVC) retains the acronym for con- a1.,19901. Lesions of the rostralpathway may also result in sistencywith the substantialquantity ofprevious literature but is inad- song degradationin open-endedleamers, such as canaries equatebecause it has funcitonal implications, and becauseit does not (Serinuscanarius), during seasonalperiods of adult song describe the position, shape,or size of the nucleus or its cellular con- stituents.We propose adopting the acronym as the proper name. This development[Nottebohm et al., 1990], or in other species is not inconsistent with old terminology (e.g. field L) nor with some during experimentallyinduced periods of adult song plas- new termhology (e.9. areaX). ticity [Monison and Nottebohm,1993]. 248 Margoliash/Fotune/Sutter/YuAVren-Hardin/ Organization of Song System: Dave Functions and Origins I I Telencephalon Fig. 1. Schematic of the song system. I I Arows indicate the direction of major pro- I jections. Subdivisions of nuclei (esp. Ov, Dorsal thalanurs Field L, and RA) are not represented. The Midbrain projectionof RA onto DLM lwild, 1993b]is Brainstem difiicult to confirm with retrograde tracers. RAm Am nXIIts Note HVc is the major site of convergence, /tt and RA is the major output structure [seealso respfatory laryngea-l synnx wild.1994t. The unidirectional,and mostly unilateralconnectivity of own song than to conspeci{icsongs [seeMargoliash, 1987]. song system nuclei supports a hierarchical organization Such responseselectivity implies hierarchical organization scheme,within which HVc has a specialposition. HVc is and a high degreeofconvergence, organizationalprinciples the major site of convergencein the telencephalon,and it generally observedin auditory systems [e.g. Konishi, l99l; gives rise to two efferentroutes to RA, the major telence- Carr,19921. Furthermore, HVc is a nexus for auditory input phalic output structure(fig 1). One efferentpathway is the to the song system: lesions of HVc abolish auditory song direct projection of HVc caudally and ventrally onto RA selectivity otherwise seenin both of HVc's efferent targets, [Nottebohmet a1.,I976];the secondpathway is the indirect area X and RA fDoupe and Konishi, l99l:' Vicario and projection to RA via the rostral pathway, HVc-+area Yohay, 19931.The concept of a hierarchical organizationin X+DLM -+ IMAN-+RA [Nottebohmet al., 1916, 1982; the song system is also supported by motor properties of Okuhataand Saito, 1987; Bottjer et al., 19891.The indirect HVc. In zebra finches (Taeniopygiaguttata), HVc stimula- 'recursive' pathwayhas alsobeen called lWilliams, 1989; tion during singing causesa re-settingof the pattem of song Nottebohm,19911, however multiple but separateroutes syllables,whereas stimulation in RA affects only the acous- from the samesource (HVc) to the sametarget (RA) does tic structure of the syllable
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