The standard nomenclature that has been used for many telencephalic and related brainstem structures in birds is based on flawed assumptions of homology to mammals. In particular, the outdated terminology implies that most of the avian telencephalon is a hypertrophied basal ganglia, when it is now clear that most of the avian telencephalon is neurochemically, hodologically, and functionally comparable to the mammalian neocortex, claustrum, and pallial amygdala (all of which derive from the pallial sector of the developing telencephalon). Recognizing that this promotes misunderstanding of the functional organization of avian brains and their evolutionary relationship to mammalian brains, avian brain specialists began discussions to rectify this problem, culminating in the Avian Brain Nomenclature Forum held at Duke University in July 2002, which approved a new terminology for avian telencephalon and some allied brainstem cell groups. Details of this new terminology are presented here, as is a rationale for each name change and evidence for any homologies implied by the new names.Revisions for the brainstem focused on vocal control, catecholaminergic, cholinergic, and basal ganglia-related nuclei. For example, the Forum recognized that the hypoglossal nucleus had been incorrectly identified as the nucleus intermedius in the Karten and Hodos (1967) pigeon brain atlas, and what was identified as the hypoglossal nucleus in that atlas should instead be called the supraspinal nucleus. The locus ceruleus of this and other avian atlases was noted to consist of a caudal noradrenergic part homologous to the mammalian locus coeruleus and a rostral region corresponding to the mammalian A8 dopaminergic cell group. The midbrain dopaminergic cell group in birds known as the nucleus tegmenti pedunculopontinus pars compacta was recognized as homologous to the mammalian substantia nigra pars compacta and was renamed accordingly; a group of ␥-aminobutyric acid (GABA)ergic neurons at the lateral edge of this region was identified as homologous to the mammalian substantia nigra pars reticulata and was also renamed accordingly. A field of cholinergic neurons in the rostral avian hindbrain was named the nucleus pedunculopontinus tegmenti, whereas the anterior nucleus of the ansa lenticularis in the avian diencephalon was renamed the subthalamic nucleus, both for their evident mammalian homologues.For the basal (i.e., subpallial) telencephalon, the actual parts of the basal ganglia were given names reflecting their now evident homologues. For example, the lobus parolfactorius and paleostriatum augmentatum were acknowledged to make up the dorsal subdivision of the striatal part of the basal ganglia and were renamed as the medial and lateral striatum. The paleostriatum primitivum was recognized as homologous to the mammalian globus pallidus and renamed as such. Additionally, the rostroventral part of what was called the lobus parolfactorius was acknowledged as comparable to the mammalian nucleus accumbens, which, together with the...

We believe that names have a powerful influence on the experiments we do and the way in which we think. For this reason, and in the light of new evidence about the function and evolution of the vertebrate brain, an international consortium of neuroscientists has reconsidered the traditional, 100-year-old terminology that is used to describe the avian cerebrum. Our current understanding of the avian brain -in particular the neocortex-like cognitive functions of the avian pallium -requires a new terminology that better reflects these functions and the homologies between avian and mammalian brains.One hundred years ago, Edinger, the father of comparative neuroanatomy, formulated a unified theory of brain evolution that formed the basis of a nomenclature that has been used to define the cerebral subdivisions of all vertebrates 1 . This resulted in terms and associated concepts such as palaeostriatum, archistriatum, neostriatum and neocortex that are still in common use. According to this theory, the avian cerebrum is almost entirely composed of basal ganglia, the basal ganglia is involved in only instinctive behaviour, and the malleable behaviour that is thought to typify mammals exclusively requires the so-called neocortex. However, towards the end of the twentieth century, there accumulated a wealth of evidence that these viewpoints were incorrect. The avian cerebrum has a large pallial territory that performs functions similar to those of the mammalian cortex. Although the avian pallium is nuclear, and the mammalian cortex is laminar in organization, the avian pallium supports cognitive abilities similar to, and for some species more advanced than, those of many mammals. To eliminate these misconceptions, an international forum of neuroscientists (BOX 1) has, for the first time in 100 years, developed new terminology that more accurately reflects our current understanding of the avian cerebrum and its homologies with mammals. This change in terminology is part of a new understanding of vertebrate brain evolution.In this article, we summarize the traditional view of telencephalic evolution before reviewing more recent findings and insights. We then present the new nomenclature that has been Correspondence to Erich Jarvis at the

Birds have excellent visual abilities that are comparable or superior to those of primates, but how the bird brain solves complex visual problems is poorly understood. More specifically, we lack knowledge about how such superb abilities are used in nature and how the brain, especially the telencephalon, is organized to process visual information. Here we review the results of several studies that examine the organization of the avian telencephalon and the relevance of visual abilities to avian social and reproductive behavior. Video playback and photographic stimuli show that birds can detect and evaluate subtle differences in local facial features of potential mates in a fashion similar to that of primates. These techniques have also revealed that birds do not attend well to global configural changes in the face, suggesting a fundamental difference between birds and primates in face perception. The telencephalon plays a major role in the visual and visuo-cognitive abilities of birds and primates, and anatomical data suggest that these animals may share similar organizational characteristics in the visual telencephalon. As is true in the primate cerebral cortex, different visual features are processed separately in the avian telencephalon where separate channels are organized in the anterior-posterior axis roughly parallel to the major laminae. Furthermore, the efferent projections from the primary visual telencephalon form an extensive column-like continuum involving the dorsolateral pallium and the lateral basal ganglia. Such a column-like organization may exist not only for vision, but for other sensory modalities and even for a continuum that links sensory and limbic areas of the avian brain. Behavioral and neural studies must be integrated in order to understand how birds have developed their amazing visual systems through 150 million years of evolution.

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Until recently, the exact location of the avian nucleus accumbens within the basal forebrain had not been well established (Reiner et al. [2004] J Comp Neurol 473:377-414). While a number of previous studies have shown afferents and efferents of the presumptive "nucleus accumbens," detailed and accurate connection patterns of this newly recognized area are still lacking. We set out to clarify these connections using small, localized injections of cholera toxin subunit B and biotinylated dextran amine directly into the nucleus. In order to increase the accuracy of tracer injections into target sites, we first conducted a systematic comparison of three calcium-binding proteins, namely, parvalbumin, calretinin, and calbindin, to characterize the nucleus accumbens and ascertain its boundaries. The results showed that the avian and mammalian nucleus accumbens had remarkable hodological similarities, including the connections with the hippocampus, amygdala, ventral pallidum, lateral hypothalamus, and ventral tegmental area. However, the most significant aspect of the present study is that the avian nucleus accumbens had extensive reciprocal connections with medial pallial structures, the mammalian counterparts of which are unclear. Three implications of this finding are discussed. First, the avian medial pallium may correspond to part of the mammalian prefrontal cortex based on the connections with the nucleus accumbens. Second, the avian brain has a "limbic loop" involving the medial pallium, which also receives input from the avian equivalent of the mediodorsal thalamus. Third, the extensive connections between the accumbens and medial pallium just dorsal to it suggest a column-like organization of limbic-associated areas in the avian telencephalon.