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The Avian Brain Nomenclature Forum:
Terminology for a New Century in
Comparative Neuroanatomy
ANTON REINER,
1
*DAVID J. PERKEL,
2
LAURA L. BRUCE,
3
ANN B. BUTLER,
4
ANDRA
´S CSILLAG,
5
WAYNE KUENZEL,
6
LORETA MEDINA,
7
GEORGE PAXINOS,
8
TORU SHIMIZU,
9
GEORG STRIEDTER,
10
MARTIN WILD,
11
GREGORY F. BALL,
12
SARAH DURAND,
13
ONUR GU
¨TU
¨RKU
¨N,
14
DIANE W. LEE,
15
CLAUDIO V. MELLO,
16
ALICE POWERS,
17
STEPHANIE A. WHITE,
18
GERALD HOUGH,
19
LUBICA KUBIKOVA,
20
TOM V. SMULDERS,
21
KAZUHIRO WADA,
20
JENNIFER DUGAS-FORD,
22
SCOTT HUSBAND,
9
KEIKO YAMAMOTO,
1
JING YU,
20
CONNIE SIANG,
20
AND ERICH D. JARVIS
20
*
1
Department of Anatomy and Neurobiology, University of Tennessee Health Science
Center, Memphis, Tennessee 38163
2
Departments of Biology and Otolaryngology, University of Washington,
Seattle Washington 98195-6515
3
Department of Biomedical Sciences, Creighton University School of Medicine,
Omaha, Nebraska 68178
4
Krasnow Institute and Department of Psychology, George Mason University,
Fairfax, Virginia 22030-4444
5
Department of Anatomy, Semmelweis University, Faculty of Medicine,
H-1094 Budapest, Hungary
6
Department of Poultry Science, Poultry Science Center, University of Arkansas,
Fayetteville, Arkansas 72701
7
Department of Human Anatomy, Faculty of Medicine, University of Murcia,
Murcia E-30100, Spain
8
Prince of Wales Medical Research Institute, Sydney, New South Wales 2031, Australia
9
Department of Psychology, University of South Florida, Tampa, Florida 33620-8200
10
Department of Neurobiology and Behavior, University of California at Irvine,
Irvine, California 92697-4550
11
Division of Anatomy, Faculty of Medical and Health Sciences, University of Auckland,
Auckland 92019, New Zealand
12
Department of Psychological and Brain Sciences, Johns Hopkins University,
Baltimore, Maryland 21218
13
Department of Biology, Queens College–City University of New York,
Flushing, New York 11367-1597
14
Department of Biopsychology, Institute of Cognitive Neuroscience, Faculty of Psychology,
Ruhr-Universita¨t Bochum, 44780 Bochum, Germany
15
Department of Psychology, California State University, Long Beach,
Long Beach, California 90840-0901
16
Neurological Sciences Institute, Oregon Health and Science University, West Campus,
Beaverton, Oregon 97006-3499
17
Department of Psychology, St John’s University, Jamaica, New York 11439
18
Department of Physiological Science, University of California, Los Angeles,
Los Angeles, California 90095-1606
19
Department of Psychology, Bowling Green State University, Bowling Green, Ohio 43403
20
Department of Neurobiology, Duke University Medical Center,
Durham, North Carolina 27710
21
School of Biology, University of Newcastle, Newcastle upon Tyne
NE2 4HH, United Kingdom
22
Department of Organismal Biology and Anatomy, University of Chicago,
Chicago, Illinois 60637
THE JOURNAL OF COMPARATIVE NEUROLOGY 473:E1–E6 (2004)
©2004 WILEY-LISS, INC.
ABSTRACT
Many of the assumptions of homology on which the standard nomenclature for the cell
groups and fiber tracts of avian brains have been based are in error, and as a result that
terminology promotes misunderstanding of the functional organization of avian brains and
their evolutionary relationship to mammalian brains. Recognizing this problem, a number of
avian brain researchers began an effort to revise the terminology, which culminated in the
Avian Brain Nomenclature Forum, held at Duke University from July 18 to 20, 2002. In the
new terminology approved at this Forum, the flawed conception that the telencephalon of
birds consists nearly entirely of a hypertrophied basal ganglia has been purged from the
telencephalic terminology, and the actual parts of the basal ganglia and its brainstem
afferent cell groups have been given names reflecting their now evident homologies. The
telencephalic regions that were erroneously named to reflect presumed homology to mam-
malian basal ganglia were renamed as parts of the pallium, using prefixes that retained most
established abbreviations (to maintain continuity with the replaced nomenclature). Details of
this meeting and its major conclusions are presented in this paper, and the details of the new
terminology and its basis are presented in a longer companion paper. We urge all to use this
new terminology, because we believe it will promote better communication among neurosci-
entists. More information is available at the Avian Brain Nomenclature Exchange website
http://avianbrain.org. J. Comp. Neurol. 473:E1–E6, 2004. ©2004 Wiley-Liss, Inc.
Indexing terms: pallium; basal ganglia; telencephalon; brainstem; evolution; terminology; birds;
mammals
The view of telencephalic evolution that became wide-
spread during the first 60 years of the 20th century was
that both birds and mammals shared several basal gan-
glia structures, namely, an older structure inherited from
fish called the paleostriatum (now called the globus palli-
dus in mammals) and a newer basal ganglia structure
that evolved in amphibians but expanded in reptiles and
more so in birds, called the neostriatum (then considered
equivalent to the supposedly newer parts of the caudate
and putamen; Edinger et al., 1903; Edinger, 1908; Arie¨ns-
Kapper 1922, 1928; Johnston, 1923; Arie¨ns-Kappers et al.,
1936; Herrick, 1948, 1956). Reptiles were thought to have
also elaborated the two parts of the paleostriatum of fish,
the primitivum and the augmentatum (the latter consid-
ered equivalent to older parts of mammalian caudatopu-
tamen) into distinct regions and passed on this trait to
birds, whereas the neostriatum in birds was thought to
have given rise to a novel overlying structure called the
hyperstriatum. Birds and mammals were also thought to
share a caudobasal subcortical structure termed the arch-
istriatum, now called the amygdala in mammals. Because
an equivalent of laminated mammalian neocortex was not
evident in birds, their telencephalon was considered to
consist primarily of a hypertrophied basal ganglia. Al-
though some investigators such as Kuhlenbeck, Rose,
and Ka¨lle´n dissented from these views of avian brain
organization and evolution (Rose, 1914; Kuhlenbeck,
1938; Ka¨lle´n, 1953), the accretionary theory of verte-
brate brain evolution, as espoused in major books by
Arie¨ns-Kappers et al. (1936) and Herrick (1948, 1956),
became the prevailing view and led to the predominant
use of the terms neostriatum, archistriatum, and hyper-
striatum to refer to the major sectors of the telenceph-
alon above the so-called paleostriatum in birds and to
the term neocortex for the major telencephalic sector in
mammals.
In 1967, Karten and Hodos published their stereotaxic
atlas of the pigeon brain, which provided the first compre-
hensive effort to identify and name all parts of the brain in
birds. For the diverse subtelencephalic structures, prior
studies offered simple and uncontroversial terms that
Karten and Hodos adopted (Huber and Crosby, 1929;
Craigie, 1931; Kuhlenbeck, 1937, 1939; Meessen and Ol-
szewski, 1949; Olszewski and Baxter, 1954). Karten and
Hodos, however, recognized that the choice of a terminol-
ogy for avian telencephalon was more problematic, be-
cause they were aware that the structures termed the
archistriatum, neostriatum, ectostriatum, and hyperstria-
tum by Arie¨ns-Kappers et al. (1936) were unlikely to be
parts of the basal ganglia. Despite their misgivings,
Karten and Hodos chose to use these terms because they
were already entrenched. Subsequent atlases for other
avian species (Kuenzel and Masson, 1988) largely used
the same terminology as Karten and Hodos. Although
much of this terminology has stood the test of time, many
Grant sponsor: National Science Foundation; Grant number: IBN-
0110894; Grant sponsor: National Institutes of Health; Grant number:
1R13-MH-64400-01.
*Correspondence to: Anton Reiner, Department of Anatomy & Neurobi-
ology, University of Tennessee Health Science Center, 855 Monroe Avenue,
Memphis, TN 38163. E-mail: areiner@utmem.edu and/or Erich D. Jarvis,
Department of Neurobiology, Box 3209, Duke University Medical Center,
Durham, NC 27710. E-mail:jarvis@neuro.duke.edu
Received 18 April 2003; Revised 11 December 2003; Accepted 21 January
2004
DOI 10.1002/cne.20119
Published online in Wiley InterScience (www.interscience.wiley.com).
E2 A. REINER ET AL.
of the interpretations of telencephalic homology implied
by the terminology of Arie¨ns-Kappers et al. (1936) have
been overwhelmingly shown to be erroneous. Additionally,
the mammalian homologues of some brainstem cell groups
connected with the telencephalon, which were not known
at the time the Karten and Hodos atlas was completed,
have also become clear. As deeper insight has been gained
into the evolution, development, and function of the brains
of birds and mammals, the flawed homologies implied by
the terms of Arie¨ns-Kappers et al. for avian telencephalon
and some now evident errors in brainstem terminology
have greatly hindered communication among avian and
mammalian brain research specialists and perpetuated an
outdated view of avian brain evolution.
This issue came to be of increasing concern to avian
neurobiologists over the past 10 years, and formal efforts
to revise avian brain nomenclature were begun 5 years
ago by a small group of avian brain specialists. To develop
widely acceptable new terms, this group sought to involve
a greater number of researchers than had participated in
two previous attempts to standardize avian neuroana-
tomical nomenclature (Baumel, 1979, 1993). Accordingly,
the group discussing such a revision eventually grew to an
international collection of multidisciplinary neuroscien-
tists, and 2 years ago the group decided to hold an open
Nomenclature Forum, at which a new terminology would
be adopted. This Forum was held July 18 –20, 2002 at
Duke University in Durham, North Carolina, and it was
preceded by in-depth discussions by E-mail and telephone
of the need for a terminology change and specific recom-
mendations as to the nature of the new terms. This report
describes the pre-Forum preparatory period, the Forum
logistics, and the decision-making process. The new ter-
minology itself and the rationale for individual changes
are presented in detail in a companion paper (Reiner et
al., 2004).
AVIAN BRAIN NOMENCLATURE FORUM
Rationale and overview of meeting
Armed with 2 years of formal preparation, an interna-
tional team of experts in the fields of avian, mammalian,
reptilian, and fish brain research assembled at Duke Uni-
versity and took on the task of devising a new avian
telencephalic nomenclature. This group critically evalu-
ated the evidence, as detailed in various published and
soon-to-be-published studies, for specific, possible new
terms. We concluded that an overwhelming body of data
supports the interpretation that most of the dorsal three-
fourths of the cerebrum in birds (including what has been
termed the neostriatum, hyperstriatum, and archistria-
tum) is pallial in nature and therefore homologous as a
field to the brain sector that in mammals includes the
neocortex, claustrum, piriform cortex, and pallial amyg-
dala (Karten, 1969, 1991; Gu¨ntu¨rku¨n, 1991; Butler, 1994;
Reiner et al., 1998; Smith-Fernandez et al., 1998; Medina
and Reiner, 2000; Puelles et al., 2000). Accordingly, we
have now designated the major subdivisions of the dorsal
three-fourths of the telencephalon in birds with terms that
contain the root word “-pallium”rather than the Arie¨ns-
Kappers et al. term “-striatum.”We have also revised
prefixes with questionable evolutionary implications. We
further concluded that the approximately ventral one-
fourth of the cerebrum in birds contains the homologues of
such subpallial structures in mammals as the basal gan-
glia proper (including dorsal striatal and pallidal subdivi-
sions), the more ventrally located limbic striato-pallidal
complex (sometimes called the ventral basal ganglia), the
medial and lateral bed nuclei of the stria terminalis, the
basal nucleus of Meynert, and part of the subpallial amyg-
dala. The new names chosen for these subpallial struc-
tures reflect these homologies.
Meeting planning and preparation
In organizing and planning the Nomenclature Forum,
several goals were paramount. First, it was necessary to
devise means by which the researchers interested in the
issue of nomenclature change could communicate and de-
velop their thoughts about suitable name changes. This
was achieved through two E-mail list servers, one for all
avian brain researchers and one for songbird specialists.
Typically messages were posted to both lists. By means of
these two E-mail list servers, the discussion of nomencla-
ture change was open to the broad community of avian
brain researchers, and all had the opportunity to contrib-
ute. Additionally, those interested in nomenclature
change met as a group one evening at the annual Society
for Neuroscience Meeting for each of the several years
preceding the Forum and discussed issues related to avian
brain nomenclature revision.
Second, a planning group of 13 neurobiologists was es-
tablished for the Nomenclature Forum. This group also
communicated openly by E-mail, supplemented by indi-
vidual face-to-face or telephone conversations. In forming
this group, which constituted the core of those attending
the Forum, we sought to include major experts in avian
neurobiology, as well as in fish, amphibian, reptilian, and
mammalian neurobiology. This group included Laura L.
Bruce, Ann B. Butler, Andra´s Csillag, Erich D. Jarvis,
Harvey J. Karten, Wayne Kuenzel, Loreta Medina,
George Paxinos, David J. Perkel, Anton Reiner, Toru
Shimizu, Georg Striedter, and Martin Wild. Many mem-
bers of this group possess expertise in more than one
vertebrate group, some have considerable experience in
brain atlas construction, and some are conversant with
classical languages.
Third, to provide concrete ideas for nomenclature revi-
sion, proposals for specific nomenclature change were so-
licited by the core group and then extensively discussed
(principally by E-mail) as to their strengths and weak-
nesses prior to the meeting. The open nature of the E-mail
communication within the avian brain research commu-
nity made it possible to gauge the reactions of diverse
members of the community to specific proposals.
Fourth, to foster planning for the meeting and dissem-
ination of ideas and information related to the nomencla-
ture revision effort, a website called the Avian Brain No-
menclature Exchange (http://jarvis.neuro.duke.edu/nomen/
index.html, renamed recently to http://avianbrain.org/
nomen/index.html) was established. The expectation was
that Forum attendees would be well versed in the need for
the terminology change, in specific suggestions as to which
structures needed a name change, specific proposals as to
the new names, and the rationale or data supporting any
given proposed name change.
Meeting format
Schedule. The 3-day Forum was organized into three
major goal-oriented blocks. On the first day, the rationales
E3AVIAN BRAIN NOMENCLATURE FORUM
for specific suggested name changes for subpallial and
some related brainstem cell groups were reviewed and
evaluated, and the Forum voted on the name changes. For
the subpallium and related brainstem cell groups, the
parties recommending name changes differed little among
each other in the cell groups recommended for name
change or in the specifics of the proposed new names. The
discussion mainly focused on the evidence for the homol-
ogies underpinning the specific recommended name
changes. On the second day, the rationales and merits for
various specific suggested name changes for the neostria-
tum and hyperstriatum were presented and discussed.
Voting was completed on new names for these structures
on the morning of the third day, and the remainder of the
third day was devoted to the rationales for specific pro-
posed name changes for the archistriatal complex and
voting for new names for this region. The discussion about
the pallial terminology focused on cytoarchitectonic
boundaries, the limits of what the data could conclusively
prove about homology to mammalian structures, and the
esthetics and practicalities (impact on the accessibility of
the avian brain literature) of the specific proposed name
changes for the pallium. The overall discussion and eval-
uation process on each day involved use of computers,
projectors, video-interfaced microscopes, and internet con-
nections to display the data and images required to assess
published and unpublished data favoring or opposing par-
ticular proposals. Discussion was open, and focus was
maintained by a moderator for each session. Discussion/
Data sessions typically were 2–3 hours in length, sepa-
rated by 15–30-minute breaks that were largely charac-
terized by spontaneous small group discussions on the
topic of the preceding formal session.
Attendees. The meeting was attended by 19 faculty-
level neuroscientists (Gregory F. Ball, Laura L. Bruce,
Ann B. Butler, Andra´s Csillag, Sarah Durand, Onur Gu¨n-
tu¨rku¨n, Erich D. Jarvis, Wayne Kuenzel, Diane Lee,
Loreta Medina, Claudio V. Mello, George Paxinos, David
J. Perkel, Alice Powers, Anton Reiner, Toru Shimizu,
Georg Striedter, Stephanie White, and Martin Wild), four
postdoctoral fellows (Gerald Hough, Lubica Kubikova,
Tom V. Smulders, and Kazuhiro Wada), five graduate
students (Jennifer Dugas-Ford, Haruhito Horita, Scott
Husband, Keiko Yamamoto, and Jing Yu), and one under-
graduate student (Connie Siang). All Forum attendees are
co-authors of this and the companion paper (Reiner et al.,
2004). Attendance at the meeting was open to all who
wished to attend, and travel and lodging costs, as well as
the costs of the meeting itself, were supported by awards
from NSF and NIH for the Forum. An additional 12 indi-
viduals assisted in technical aspects of the Forum, includ-
ing computer network specialists, audio-visual specialists,
a web specialist, microscopy specialists, administrators,
and graduate and undergraduate student assistants.
These persons are listed in the acknowledgments. Erich
Jarvis and Anton Reiner served as co-organizers of the
meeting.
Voting. The planning committee had decided prior to
the meeting that any nomenclature change needed a high
degree of concurrence among Forum attendees if it was to
be widely accepted by the field as a replacement for any
existing term. We therefore decided changes on a
structure-by-structure basis, with 80% approval required
for acceptance of each new term. Each faculty member
attending was accorded a full vote and each postdoctoral
fellow a half vote. Graduate and undergraduate students
did not vote, but their input was considered. The limits on
student voting were put in place because of the perception
that students were not yet adequately conversant with the
issues of relevance to the terminology revision, whereas
postdoctoral fellows were considered at least partly famil-
iar. For pallial structures, it proved necessary to eliminate
some of the proposed options by simple majority votes,
before a final vote of approval for a given name change
could be conducted. The final set of approved new names
for pallial structures was largely an amalgam of the most
highly favored choices from the different sets of proposals.
Guiding principles. In adopting a new terminology,
several guiding principles were embraced. The overall
goals were to remove inaccurate implications of homology
where they existed (notably for the pallium) and to recog-
nize homology where it was amply demonstrated (notably
for brainstem and subpallial structures). Because our in-
tent was to improve communication among avian and
mammalian brain specialists, in the case of instances in
which one-to-one homology (also termed discrete homol-
ogy by Smith, 1967) had been clearly demonstrated, we
believed it highly advantageous to adopt for birds the
same name as used for that structure in mammals (e.g.,
globus pallidus instead of paleostriatum primitivum). The
gain in communication and the already established famil-
iarity of the new avian term (because of its use in mam-
mals) were thought to far outweigh any disadvantages
inherent in abandoning the old name and old abbrevia-
tion. For the pallium, we confronted the issue of whether
sufficient data were available to conclude safely and un-
equivocally that given structures possessed one-to-one ho-
mology with specific structures in mammals. In the end,
we concluded that sufficient evidence did not exist for such
one-to-one equivalences at the pallial level, other than for
the piriform cortex (Karten, 1969, 1991; Bruce and Neary,
1995; Striedter, 1997; Smith-Fernandez et al., 1998; Me-
dina and Reiner, 2000; Puelles et al., 2000; Reiner, 2000;
Butler and Molna´r, 2002; Butler et al., 2002; Wada et al.,
2001).
The goal then became to remove any incorrect connota-
tion of homology to the basal ganglia in the case of those
pallial structures with the term “striatum”in the name.
Although it was agreed by all that the new names for
these structures should have pallium in the name, several
issues needed to be considered in renaming the pallial
structures that possessed “-striatum”as a root word in
their outdated name. One major issue was the extent to
which developing new names that allowed retention of
existing abbreviations was desirable and could be
achieved with esthetically pleasing new terms. Alterna-
tively, consideration needed to be given to the possibility
that a new and simple descriptive terminology that did not
retain abbreviations could make avian brain structures
easy to learn and more broadly accessible to neuroscien-
tists. As is clear from the region-by-region commentary in
the companion paper (Reiner et al., 2004), in the end,
retention of abbreviations was found to be highly desirable
for the most intensely studied structures of avian pallium,
so there would be easy linkage and clear continuity be-
tween the literature using old and new terms. Further
information and avian brain images depicting this new
nomenclature are available in Reiner et al. (2004) and on
the Avian Brain Nomenclature Exchange website: http://
avianbrain.org. Further details on the terminology op-
E4 A. REINER ET AL.
tions discussed during the Forum meeting by its attendees
will be presented in a later publication, in a special edition
of Brain, Behavior and Evolution dedicated to the nomen-
clature revision.
Final comments
The Avian Brain Nomenclature Forum was the result of
growing awareness of the communication problems
caused by the faulty and outdated avian brain terminol-
ogy. The Forum sought to devise a new terminology that is
free of errors and promotes accurate understanding of
avian brain organization and evolution. We have been
scrupulous to use only names implying homology that we
are certain would not themselves later prove to be in error.
We believe that the nomenclature changes we have de-
vised can serve the field well, and we urge all investigators
to use this new terminology. In making its recommenda-
tions for terminology change for specific structures, the
Forum does not mean to imply that the names for all other
structures in the avian brain are adequate and suitable.
Nonetheless, we believe that the names changed by the
Forum are those that were in greatest need of change and
were the greatest hindrance to accurate understanding of
avian brain organization.
ACKNOWLEDGMENTS
Individuals at Duke University who helped with admin-
istrative, technical, and logistical support for the Forum
include: Deepa Bharanidharan (Jarvis Laboratory Associ-
ate in Research), Eunice Chang (Graduate Student), Mar-
garet Couvillon (Graduate Student), Haruhito Horita
(Graduate Student), Susan Havrilesky (Department of
Neurobiology Manager), Michael McElroy (Jarvis Labora-
tory Research Technician), Dawn Kernagis (Jarvis Labo-
ratory Associate in Research), Lisa Moore (Jarvis Labora-
tory Manager), Martha Musson (Department of
Neurobiology Secretary), and Netfriends computer assis-
tants, Ann Sink (Department of Neurobiology coordina-
tor), David Stokes (Web designer), and Tony Zimmermann
(Jarvis Laboratory Research Analyst). We note the valu-
able contributions of Drs. Harvey J. Karten and Luis
Puelles to the discussions on avian brain organization,
development, and evolution that preceded the nomencla-
ture meeting, and we thank Drs. Steve Brauth and Todd
Roberts for making their data on the parrot telencephalon
available to us prior to its publication. A number of other
researchers, too numerous to list here, made valuable
on-line contributions to the discussions in the years lead-
ing up to the Forum. Preparation for the Forum, the
Forum itself, and the dissemination of the conclusions of
the Forum were supported by grants from National Sci-
ence Foundation (NSF) and National Institutes of Health
(NIH). We thank Dr. Israel Lederhendler of National In-
stitute of Mental Health and Drs. Carol van Hartesveldt
and Christopher Platt of NSF for their support and en-
couragement of the Forum enterprise. We also thank the
following who did not attend the Forum for their letters of
support in applying for NIH and NSF funding for the
Forum: Verner P Bingman, Chao Deng, Timothy De-
Voogd, Alison Doupe, Barrie Frost, William Hodos, Gab-
riel Horn, Harvey Karten, Lubor Kostal, Daniel Margo-
liash, Richard Mooney, Sarah Newman, Mary Ottinger,
Giancarlo Panzica, Luis Puelles, Christoph Redies, Lesley
Rogers, and Constance Scharff.
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