AVIAN BRAIN NOMENCLATURE FORUM 18-20 JULY, 2002 - DURHAM (NC, USA) PROPOSAL FOR NEW NOMENCLATURE - 2 July 2002 Loreta Medina This proposal is a revised version of a previous one done in collaboration with Luis Puelles, to which I have incorporated some suggestions done by some avietaers (to whom I am very grateful). I thank long and productive discussions on the development and evolution of the brain, and on a possible new avian brain nomenclature maintained during several years with Luis Puelles. I also thank discussions and input on a possible new avian brain nomenclature from George Paxinos and Salvador Martínez, as well as comments and criticisms to previous versions of this proposal provided by Georg Striedter, David Perkel, Christoph Redies, Harvey Karten, Erich Jarvis, Tony Reiner, Laura Bruce, Wayne Kuenzel, Gregory F. Ball, Andras Csillag and all participants of the avi-eater list. Finally, I would like to add that -in a similar spirit to that clarified by Georg Striedter, David Perkel, Tony Reiner, Andras Csillag and all other participants at the Forum- I will go to this meeting with an open mind, prepared to debate the different issues, and happy to support any other proposal or part of it that seems appropiate. Nomenclature in colored text represents homologous names with mammals. KARTEN-HODOS-1967 and OTHER1 TERM ABBREV. ____________________________________________ MEDINA2-2002 TERM ABBREV. _______________________________________ PALLIUM - P * Hippocampal complex (medial part of the pallium) - H Hippocampus Area parahippocampalis Area corticoidea dorsolateralis Hp APH CDL NO CHANGE NO CHANGE NO CHANGE or change to caudolateral APH - APHcl * Wulst (dorsomedial part of the pallium) - W (Alternatives: dorsal pallium or dorsomedial pallium - D or DM) Hyperstriatum accesorium Hyperstriatum dorsale Hyperstriatum intercalatus superior Interstitial nucleus of HA (Karten et al., 1973) Hyperstriatum laterale HA HD HIS IHA area 1 of Wulst area 3 of Wulst area 2 of Wulst Interstitial nucleus of Wulst W1 W3 W2 WN HL superficial lateral Wulst WLs 1 Also includes areas or nuclei newly described in the pigeon/chicken brain after the Karten and Hodos atlas, as well as specific areas or nuclei described in vocal learning birds, such as oscine songbirds (zebra finch, canary, etc.) or budgerigars (parrot). Some references are given for these additional areas or nuclei. 2 For clarity, homology-based proposed terms are introduced in pink colour. In some cases, alternative names are given between parenthesis. 1 (Shimizu and Karten, 1990) KARTEN-HODOS-1967 and OTHER* TERM ABBREV. ________________________________________________ MEDINA-2002 TERM ABBREV. ____________________________________________ * Intermediate pallium (intermediate part of the pallium) - I Hyperstriatum ventrale HV Intermediate pallium Hyperstriatum ventrale dorso-ventrale HVdv Dorsointermediate pallium Hyperstriatum ventrale ventro-ventrale HVvv Ventrointermediate pallium Nucleus of the HV HVn Nucleus of the intermediate pallium (Redies et al., 2001) Specific auditory/vocal-related areas/nuclei in the intermediate pallium of vocal learning birds Oval nucleus of the anterior HV HVo Oval nucleus of the anterior I (budgerigar; Striedter, 1994; Durand et al., 1997; Jarvis and Mello, 2000). I ID IV In HVo complex (vocal anteromedial field of HV; Jarvis and Mello, 2000) HVoc Ino complex (vocal anteromed. field of I) Inoc Lateral nucleus of the anterior HV (budgerigar; Jarvis and Mello, 2000). lAHV Lateral nucleus of the anterior I Inl Caudomedial region of HV (budgerigar/songbird; Mello et al., 1992; Jarvis and Mello, 2000) CMHV Caudomedial region of interm. pall. CMI Ino * Epistriatal pallium (ventral part of the pallium) - E (Alternative: ventral pallium or V) Neostriatum Neostriatum caudale Neostriatum caudolaterale N NC NCL Epistriatal pallium Epistriatal caudal pallium Caudolateral epiestriatal pallium E EC ECL 2 (Kröner and Güntürkün, 1999) KARTEN-HODOS-1967 and OTHER* MEDINA-2002 TERM ABBREV. TERM ABBREV. ________________________________________________ ____________________________________________ Neostriatum dorsale (or dorsocaudal) Nd/Ndc Dorsocaudal epistriatal pallium Ed or Edc (Wild, 1994; Metzger et al., 1998; Kröner and Güntürkün, 1999) Neostriatum frontale NF Epistriatal frontal pallium EF Neostriatum intermedium NI Epistriatal intermediate pallium EI Neostriatal island field NIF Epistriatal island field EIF (Redies et al., 2001) Lateral Neostriatum NL Lateral epistriatal pallium EL (Striedter, 1984; Redies et al., 2001) Ectostriatum Ec Ectostriatal pallium Ec Field L L Field L L Nucleus basalis Bas Nucleus basalis trigemini BT Specific auditory/vocal-related areas/nuclei in the epistriatal/caudolateral pallium of vocal learning birds Magnocellular nucleus of MAN Magnocellular nucleus of the MAE anterior neostriatum ( songbirds; anterior epistriatal pallium Nottebohm et al.,1982; Bottjer et al., 1989) Oval nucleus of the anterior neostriatum NAo (budgerigar, Brauth et al., 1994; Durand et al., 1997; Jarvis and Mello, 2000; proposed comparable to MAN of songbirds) Oval nucleus of the anterior E EAo NAo complex (budgerigar; Jarvis and Mello, 2000) EAo complex EAoc Lateral nucleus of the frontal or anterior E EFl or EAl NAoc Lateral nucleus of the frontal or anterior NFl or lAN neostriatum (budgerigar, Striedter, 1994; Durand et al., 1997; Jarvis and Mello 2000) 3 KARTEN-HODOS-1967 and OTHER* TERM ABBREV. ________________________________________________ MEDINA-2002 TERM ABBREV. ____________________________________________ Central nucleus of the lateral neostriatum NLc (budgerigar; Striedter, 1994, Durand et al., 1997; Jarvis and Mello, 2000) Lateral nucleus of the lateral E ELc Supracentral nucleus of NL (budgerigar; Striedter, 1994) NLs Supracentral nucleus of EL ELs High vocal center (in NCL) (songbirds; Nottebohm et al., 1976; Fortune and Margoliah, 1995) HVc/HVC High vocal center (in CL) HVC HVC shelf shelf (songbirds; Wild, 1994; Vates et al., 1996; Mello et al., 1998; proposed comparable to Nd of pigeon/chicken) NO CHANGE Para-HVC (songbirds; Gahr et al., 1987; Foster and Bottjer, 1998) NO CHANGE pHVC Neostriatum intermedium p. dorsolateralis NIDL (budgerigar; Brauth and McHale, 1988; Jarvis and Mello, 2000; proposed comparable to Nd of pigeon/chicken and to HVC shelf of songbirds) Dorsolateral epistriatal pallium (located in ECL) KARTEN-HODOS-1967 and OTHER* MEDINA-2002 EDL 4 TERM ABBREV. ________________________________________________ TERM ABBREV. ____________________________________________ SUBPALLIUM/ENTOPEDUNCULAR AREA - SP/AEP Lobus parolfactorius LPO Medial Striatum STm Rostromedial part of LPO LPO Nucleus Accumbens Acc Nucleus accumbens Acc Bed nucleus of the stria terminalis, lateral part (Aste et al., 1998) BSTl Paleostriatum augmentatum Paleostriatum primitivum Nucleus intrapeduncularis Septal Nuclei Tuberculum olfactorium PA PP INP SL, SM, etc. TO Lateral Striatum Dorsal Pallidum NO CHANGE NO CHANGE NO CHANGE STl PAd GABAergic neurons inside the fasciculus prosencephalicus medialis inside FPM Ventral Pallidum (Veenman and Reiner, 1994; Medina and Reiner, 1997) PAv Large cholinergic neurons of the basal none forebrain partially invading the medial and lateral prosencephalic bundles (FPM, FPL) and pallidum (Medina and Reiner, 1994) Basal magnocellular cell population BM KARTEN-HODOS-1967 and OTHER* TERM ABBREV. MEDINA-2002 TERM ABBREV. 5 ________________________________________________ ____________________________________________ Large cholinergic neurons in the fasciculus FDB diagonalis Brocae Nucleus of the diagonal band (Medina and Reiner, 1994) DB Cell population around the anterior commissure at the subpallium-preoptic transition Bed nucleus of the stria terminalis, medial part (Aste et al., 1998) BSTm none Specific auditory/vocal-related areas/nuclei in the subpallium of vocal learning birds Area X of LPO Area X Area X of the medial striatum (songbirds; Nottebohm et al., 1976; Bottjer et al., 1989; Farries and Perkel, 2002) Area X Magnocellular nucleus of LPO LPOm (budgerigar; Jarvis and Mello, 2000; proposed comparable to area X of songbirds) Magnocellular nucleus of the medial striatum STmc Caudal paleostriatum PC (budgerigar and songbirds; Jarvis and Mello, 2000) Caudal striatum CST Amygdaloid complex A AMYGDALOID COMPLEX - A Archistriatum A (divisions generally follow Zeier and Karten, 1971) Archistriatum anterior AA Archistriatum intermedium Ai Archistriatum interm. pars dorsalis Ad/Aid Archistriatum interm. pars ventralis Av/Aiv KARTEN-HODOS-1967 and OTHER* TERM ABBREV. _____________________________________________ Anterior amygdaloid area AA Intermediate amygdaloid area Ai Dorsointermediate amygdaloid area Aid Ventrointermediate amygdaloid area Aiv MEDINA-2002 TERM ABBREV. ____________________________________________ 6 Ventromedial nucleus of Ai (Wild et al., 1993) Archistriatum mediale Archistriatum posterior Nucleus Taeniae Aivm Ventromedial nucleus of Ai Aivm Am Ap Tn Medial amygdaloid area Posterior amygdaloid area NO CHANGE Am Ap Specific auditory/vocal-related areas/nuclei in the amygdaloid complex of vocal learning birds Nucleus robustus archistriatalis RA Nucleus robustus of the amygd. compl. (songbirds; Nottebohm et al., 1976; (or robust amygdaloid nucleus) Vicario, 1991; Wild, 1993; 1994) RA RA cup (songbird; comparable to Aivm of pigeon;Wild, 1994) cup NO CHANGE Caudomedial archistriatum (budgerigar; Jarvis and Mello, 2000; proposed comparable to Aivm of pigeons and to RA cup of songbirds) ACM Caudomedial amygdaloid area ACM Central nucleus of the anterior archistriatum (bugderigar; Striedter, 1994; Durand et al., 1997; Jarvis and Mello, 2000) AAc Central nucleus of the AA AAc KARTEN-HODOS-1967 and OTHER* TERM ABBREV. _____________________________________________ MEDINA-2002 TERM ABBREV. ____________________________________________ 7 INTRATELENCEPHALIC BOUNDARIES Lamina medullaris dorsalis Lamina hyperstriatica Lamina frontalis superior Lamina frontalis suprema Lamina archistriatalis dorsalis LMD LH LFS LFM LAD Pallial-subpallial boundary psb Ventral pallial lamina vpl Intermediate pallial lamina ipl Dorsal pallial lamina dpl Dorsal amygdaloid lamina lad or dal (we propose to use lowcase letters for boundary abbreviations) FIBER TRACTS Tractus archistriatalis dorsalis Tractus frontoarchistriaticus Tractus thalamostriaticus Tractus septomesencephalicus Tractus occipitomesencephalicus DA FA TTS TSM OM Dorsal amygdaloid tract Frontoamygdaloid tract Thalamoepistriatal tract Pallio-mesencephalic tract Ventral amygdalofugal tract DA or da FA or fa TE or te PM or pm VA or va (capitals or lowcase?) SOME BRAINSTEM AREAS RELATED TO THE TELENCEPHALON Nucleus tegmenti pedunculo-pontinus, pars compacta TPc KARTEN-HODOS-1967 and OTHER* TERM ABBREV. _____________________________________________ Substantia nigra, pars compacta (A9 dopaminergic cell group) (Bailhache and Balthazar, 1993; Reiner et al., 1994) SNc (A9) MEDINA-2002 TERM ABBREV. ____________________________________________ 8 Substantia nigra pars lateralis SNL (group of GABAergic neurons laterally adjacent to SNc and connected with the basal ganglia; Veenman and Reiner, 1994; Medina and Reiner, 1997). Substantia nigra pars reticulata SNr Area Ventralis of Tsai AVT Ventral tegmental area (A10 dopaminergic cell group; Bailhache and Balthazar, 1993; Reiner et al., 1994) VTA (A10) Locus coeruleus, rostral part (levels 2.75-2.25 of pigeon atlas) LoC (rostral part) A8 dopaminergic cell group (Bailhache and Balthazar, 1993; Reiner et al., 1994) A8 Tegmental pedunculopontine nucleus (Medina and Reiner, 1994) PPN Subthalamic nucleus (reasons explained in Jiao et al., 2000; also Medina and Reiner, 1997; Reiner et al., 1998). STN Cholinergic cell group of the rhombomere 1 tegmentum, that extends from the MPv to the nucleus subcoeruleus (SC) Anterior nucleus of the ansa lenticularis ALa * See footnote in first page. 9