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Avian Brains Secondary article

Anton Reiner, University of Tennessee, Memphis, Tennessee, USA Article Contents


. Introduction
The brain in birds is large, complex and unique in a number of ways, and it underlies the . Common Features of Bird and Reptile Brains
sophisticated cognitive, social and motor behaviours that typify birds. . Bird Brains: Neither Small Nor Simple
. The Visual System of Birds and the Optic Tectum

Introduction . Evolution of the Avian Telencephalon


. Neurotransmitters and the Avian Brain

As birds are capable of sophisticated cognitive, social and . Neural Circuits for Song in Bird Brains

motor behaviours, it is not surprising that they possess . Summary

large and complex brains. Notably, the cerebrum (or


telencephalon) and cerebellum in birds are as large and well
developed as in many mammalian species (Figure 1). While tomical and functional studies during the latter half of the
the structure of the brain in birds does possess many twentieth century clearly show the DVR is akin to parts of
commonalities with that of mammals, the brain in birds the cerebral cortex of mammals in its connections with
more closely reflects the evolutionary origin of birds from thalamus and in its role in information processing and
reptiles. As a consequence, the elaborate brain structures learning (Karten, 1969). Unlike the cerebral cortex, which
underlying the sophisticated behaviour of birds, such as consists of various functionally distinct areas that each
vocal learning in songbirds, appear to have evolved in possess a laminar architecture, the various regions of the
parallel with the brain structures subserving comparable
functions in mammals. (see Aves (birds).)
Dorsal
cortex
Hippocampus
Olfactory
Dorsal
cortex
Common Features of Bird and Reptile ventricular ridge

Brains
S Striatum
Birds evolved from archosaurian reptiles about 150 million
years ago. Many investigators favour the view that this Pallidum
origin was directly from small bipedal carnivorous
dinosaurs (Chiappe, 1995). The reptilian origin of birds is Hy
evidenced by the number of brain traits shared by birds and
reptiles that neither shares with mammals. The most (a) Turtle
conspicuous of these is the presence of a so-called dorsal
ventricular ridge (DVR) in the telencephalon of both birds
and reptiles. (see Dinosaurs and the origin of birds.) Wulst
The DVR is a mass of neural tissue that protrudes from
the dorsolateral wall of the telencephalon into the lateral Hippocampus
ventricle (Figure 2). While its relatively uniform distribution Dorsal
ventricular ridge
of neurons makes the DVR superficially resemble the
striatal part of the basal ganglia in mammals, neuroana-

Striatum Olfactory
cortex
Cerebellum Cerebrum S
Pallidum

Optic lobe
Hindbrain Hy
Olfactory bulb
Spinal cord (b) Pigeon
Optic chiasm
Figure 2 Frontal views through the cerebrum of a turtle (a) and a pigeon
Figure 1 Side view of a pigeon brain showing the major brain (b), with the major regions identified; areas that are homologous for
subdivisions. The diencephalon lies between and is hidden by the reptiles and birds are shown by the same colours. Hy, hypothalamus; S,
cerebrum and the optic lobe. septum.

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Avian Brains

DVR are organized into groupings of uniformly distrib- means by which the basal ganglia in birds and reptiles may
uted neurons. The most prominent of the specialized exert an influence on movement via the visuomotor
regions of the DVR in birds and reptiles are a visual region circuitry of the tectum. In light of the similar visual outputs
and an auditory region (Figure 3). These regions are the of the tectum to the thalamus in birds and reptiles, and the
major visual and auditory processing regions of the similar motor projections from the pretectum to the
cerebrum in both birds and reptiles. The visual region, tectum, it should be clear that the tectum and pretectum
called the ectostriatum in birds, receives its input from a share structural and functional similarities in birds and
distinct round thalamic nucleus called the nucleus rotun- reptiles. Both structures are more elaborately developed
dus, which itself receives visual input from the central grey than in mammals, and individual tectal layers and
layer of the visual part of the midbrain roof (or tectum). individual pretectal cell groups can readily be homologized
The auditory region of the DVR receives its input from a between birds and reptiles.
midline thalamic nucleus, which receives its auditory input Major differences are, however, present between birds
from the auditory midbrain roof (the inferior colliculus). and reptiles in the olfactory system and the cerebellum.
The auditory region of the DVR in birds is called field L, Birds typically do not have a keen sense of smell, and so
and the auditory thalamic nucleus in birds is ovoid in shape their olfactory bulbs and the region of the cerebrum to
and consequently called the nucleus ovoidalis. (see Basal which the olfactory bulbs convey information (the
ganglia and the regulation of movement.) olfactory cortex) are much smaller in birds than in reptiles.
Noteworthy similarities between birds and reptiles are On the other hand, birds are capable of much more
also evident for the other major constituent of the cerebral complex motor behaviour than reptiles, and one neural
hemispheres, the basal ganglia (Figure 3). As in mammals, correlate of this is the highly developed cerebellum of birds.
the basal ganglia in birds and reptiles are divided into an (see Olfaction.) (see Cerebellum: movement regulation and cognitive
input zone called the striatum and an output zone called functions.)
the pallidum, with the striatum sending its major output to
the striatum (Reiner et al., 1998). Unique to birds and
reptiles, however, the pallidal part of the basal ganglia has
but one subdivision rather than the two found in mammals, Bird Brains: Neither Small Nor Simple
and among the pallidal projection targets is a major
pretectal cell group, termed the lateral spiriform nucleus in Birds are capable of sophisticated foraging strategies,
birds. As this pretectal nucleus projects prominently to the elaborate parental and social behaviour, impressive hom-
deep output layers of the tectum, this circuit provides a ing and migratory behaviour, complex vocal learning, song
production, and remarkable motor behaviours such as
self-powered flight. In light of this, it is not surprising that
Hp
Wulst birds possess a large and complex brain. Brain size–
HV Field L
bodyweight correlations, in fact, show that brain size for
Neostriatum
N Ecto N birds rivals that for many mammals (Jerison, 1985).
Archistriatum Despite any similarities to mammals in brain size or
BG
Rt behavioural output, however, the structure of the brain in
birds closely reflects the evolutionary origin of birds from
reptiles, as emphasized by the commonalities noted
Ov previously. Contributing to the similarity in brain size
Tectum between birds and mammals are the cerebrum and
Rt Rt
Hy
cerebellum (see Figure 1), which in birds are as large and
well developed as in many mammalian species, and clearly
better developed than in any extant reptilian species. The
IC
DVR, for example, is much larger and possesses more
EW subdivisions in birds, with some of these being higher-
SN
order visual and auditory areas. Additionally, birds have
Pontine
nucleus elaborated a dorsal part of their cerebrum that in reptiles is
(a) Visual pathway (b) Auditory pathway
represented only by a thin sheet of cortical tissue overlying
a lateral extension of the lateral ventricle of the cerebrum
Figure 3 Schematic diagrams of frontal views through the cerebrum, (Figure 2). In birds this region is so thickened that it creates a
diencephalon and midbrain of the pigeon showing the cell groups that bulge at the dorsal surface of the cerebrum – hence its
make up the major visual circuit (a) and the major auditory circuit (b) in name, the Wulst, which means swelling in German.
birds. BG, basal ganglia; Ecto, ectostriatum; EW, nucleus of Edinger-
Westphal; Hp, hippocampus; HV, hyperstriatum ventrale; Hy, Irrespective of its degree of development, this cortical
hypothalamus; IC, inferior colliculus; N, neostriatum; Ov, nucleus region in birds and reptiles receives somatosensory and
ovoidalis; Rt, nucleus rotundus; SN, substantia nigra. visual input from the thalamus and thereby participates in

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Avian Brains

information processing. Note that in some avian lineages, thalamus, which projects to the visual area of the DVR
such as owls, the Wulst is even further hypertrophied, called the ectostriatum (Figure 3). This tecto-rotundo-
whereas in others such as parrots it is the DVR that is ectostriatal circuit plays a major role in visual information
enlarged. The hippocampus is also well developed in birds processing in birds (Hodos, 1976). (see Visual system.) (see Eye
(Figure 2), although not as complexly laminated as in anatomy.)
mammals. Nonetheless, as in mammals, the hippocampus Other central retinal targets in birds, however, also play
plays a role in the spatial learning that underpins homing important roles. For example, the retina also projects
behaviour, foraging and food storage. (see Hippocampus.) contralaterally to a prominent visual area of the thalamus
Consistent with the Wulst and DVR enlargement, the best referred to by the name of its mammalian homologue,
thalamic nuclei providing input to them are also enlarged the dorsal lateral geniculate nucleus. This thalamic cell
compared with their state in reptiles. Also indicative of the group has a bilateral projection to a visual region within
complexity of the avian brain is the tectum, which is more the Wulst and it plays a role in stereopsis, especially in birds
highly developed (i.e. larger and more complexly lami- such as owls with frontally placed eyes (Pettigrew, 1979).
nated) than in any other vertebrate group, as are the Note that, in mammals possessing stereopsis, it is a
associated nuclei of the pretectum. Finally, the cerebellum bilateral projection of retina to the dorsal lateral geniculate
of birds resembles that of mammals in receiving a mossy nucleus that allows the overlap of input from the two eyes
fibre input from the pontine nuclei and a climbing fibre within the brain which serves as the neural substrate for
input from the inferior olivary nucleus in the hindbrain, stereopsis. Thus, stereopsis has seemingly evolved inde-
and in being organized into lobes, which greatly increase its pendently in birds and mammals. Two final visual areas of
surface area and processing power. The expansion and note in birds are the area pretectalis of the medial
lobulation of the avian cerebellum, however, occurred pretectum, which mediates the pupillary light reflex by
independently of the superficially similar expansion and means of a projection to the nucleus of Edinger–Westphal
lobulation of the mammalian cerebellum, because the in the midbrain, and the nucleus of the basal optic root at
cerebellum is small and unlobulated in living reptiles. the anterior base of the midbrain, which mediates
(see Thalamus.) (see Cerebellum: anatomy and organization.) compensatory eye movements in response to slow move-
ments of the visual field or of the head through visual space.
(see Prey detection by bats and owls.) (see Oculomotor system.)
The Visual System of Birds and the Optic
Tectum
Birds typically possess high visual acuity. One major factor Evolution of the Avian Telencephalon
that contributes to this is that birds possess large eyes and
many large visual areas in the brain. Notable among these The avian telencephalon clearly evolved from the tele-
brain areas is the visual midbrain roof, the tectum (Figure 3). ncephalon of reptiles. Study of the telencephalon of living
Much of the retinal input to the avian brain ends in the reptiles provides clues as to the nature of the evolutionary
upper layers of the tectum, with the projection being changes that occurred as birds diverged from their now
largely crossed and organized in a retinotopic fashion. The extinct reptilian ancestors. For the basal ganglia, these
tectum makes up a considerably greater fraction of the changes were minimal and consisted largely of an
total brain volume in birds than in reptiles or mammals, expansion in the volume of the structure relative to body
apparently because of the large surface area required for size (Reiner et al., 1998). This change would have allowed
the retinotopic input from the large eyes. To provide room birds more refined motor control and motor learning
for this enlargement, the tectal lobes in birds are rotated capabilities than possessed by reptiles. (see Relationships of
outward and protrude laterally below the overlying birds - molecules v morphology.) (see Reptilia (reptiles).)
posterior pole of the cerebrum (Figures 1 and 3). By Greater changes occurred in the two telencephalic
contrast, in reptiles and mammals the tectum (which is regions comparable to the cerebral cortex of mammals
called the superior colliculus in mammals) is situated (Reiner, 2000). In reptiles these regions are called the dorsal
dorsally, and its surface area is relatively more modest. The cortex and DVR (Figure 2). In living primitive reptiles such
avian optic tectum also shows a high degree of laminar as the tuatara Sphenodon punctatum, the cortex and DVR
differentiation, with 15 cytologically distinct layers typi- appear to form one continuous structure, and the DVR is
cally being identified (Hunt and Brecha, 1984). The first not broken into separate cell groups. Even in Sphenodon,
seven layers are within the retinorecipient zone of the however, both dorsal cortex and DVR receive thalamic
tectum, and the remainder give rise to the ascending sensory input. The changes in the DVR from Sphenodon to
sensory and descending motor projections of the tectum. the grade observed in later reptiles are likely to have
As noted above, among the ascending projections of the involved differentiation of the DVR into distinct cell
tectum is an output to the nucleus rotundus of the groups, and the transition to the avian grade appears to

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Avian Brains

have involved further differentiation and enlargement of thought on the evolutionary relationship of avian DVR
both the dorsal cortex (into the Wulst) and the DVR. and Wulst to mammalian cerebral cortex (Reiner, 2000).
In living birds, the Wulst and DVR consist of a series of Both accept that stem amniotes possessed a structure
stacked slab-like zones, each of which possesses subregions resembling the dorsal cortex of extant reptiles, and that this
(Figure 4). For the Wulst, the slab-like zones are termed the region was the forerunner of the superior part of
hyperstriatum accessorium, intercalatus superior and mammalian cerebral cortex, the dorsal cortex in living
dorsale, whereas for the DVR they are termed the reptiles and the Wulst in birds, all of which possess a visual
hyperstriatum ventrale and the neostriatum. It is the area receiving input from the dorsal lateral geniculate
neostriatum that contains the thalamorecipient sensory nucleus of the thalamus and a primary somatosensory
zones of the DVR, while the hyperstriatum intercalatus area. (see Cerebral cortex.) (see Brain evolution and comparative
superior and dorsale receive sensory thalamic input in the neuroanatomy.)
Wulst. The hyperstriatum accessorium and a posterior The two hypotheses diverge as to the relationship of
basal region of the DVR termed the archistriatum (Figure 3) DVR to mammalian cerebral cortex. One viewpoint
are the source of the major descending motor projections proposes that the DVR, which seems to resemble the
of the Wulst and DVR. Note that the suffix ‘-striatum’ temporal sector of cerebral cortex in possessing thalamor-
appearing in the names of the parts of the Wulst and DVR ecipient auditory and visual regions, is derived from a
does not reflect a derivation or relationship of these zones subcortical pallial region in stem amniotes that in the
to the striatal part of the basal ganglia. These structures mammalian lineage came to give rise to several small
have ‘-striatum’ in their name because, at one time, most of amygdalar and periamygdalar cell groups. The evidence
the avian telencephalon was thought to be an expansion of used to argue for this viewpoint involves the similar
the striatal part of the basal ganglia. This notion is now positions of the DVR and these periamygdalar nuclei in the
known to be incorrect, but the names for these structures cerebrum and the embryology of these regions, especially
became entrenched and have not yet been displaced by in terms of similar expression patterns for certain genes
more accurate names. that regulate development called homeobox genes. The
In addition to the evolutionary relationship of the avian alternative hypothesis proposes that a region in stem
and reptilian telencephala, another key question of interest amniotes situated at the ventrolateral edge of dorsal cortex
is the evolutionary relationship of the avian telencephalon gave rise to the temporal part of cerebral cortex in the
to the telencephalon in mammals. For the basal ganglia, mammalian lineage and to DVR in reptiles and birds. The
there is little disagreement. Basal ganglia closely resem- evidence favouring this view is the high similarity in the
bling those in birds and mammals must have been present identity, connections and locations of the visual and
in the common amniote ancestor, given the many auditory areas of the DVR and temporal cerebral cortex.
similarities in cell types, projections, functions and Regardless of which viewpoint is correct, however, the
transmitters that are present (but see above regarding evidence is clear that the DVR and Wulst have both
differences between birds and mammals in pallidal evolved a high degree of sophistication in birds that allows
organization and projections to the pretectum) (Reiner birds to perform complex behaviours similar to those
et al., 1998). There are, however, two major schools of controlled by cerebral cortex in mammals. (see Cerebral
cortex development.) (see Mammalian embryo: Hox genes.)

Hyperstriatum
intercalatus
Hyperstriatum
superior Neurotransmitters and the Avian Brain
accessorium Hyperstriatum
dorsale The organization of neurotransmitter-specific systems of
Hyperstriatum the avian brain reflects both the evolutionary derivation of
ventrale
birds from reptiles, as well as the common evolutionary
Neostriatum derivation of birds, reptiles and mammals from stem
amniotes (Reiner et al., 1998). In the cerebrum, the lateral
Basal ganglia
wall of the telencephalon in birds, reptiles and mammals is
Olfactory divided into the same two fundamental ontogenetic
cortex
subdivisions: the pallium and the subpallium. The vast
majority of the neurons of the pallium, which consists of
the cerebral cortex in mammals and the Wulst/DVR in
Figure 4 Frontal view through the cerebrum of a pigeon, with the major birds, use the excitatory neurotransmitter glutamate, as do
regions identified and the slab-like zones making up the Wulst
(hyperstriatum accessorium, hyperstriatum intercalatus superior, and the thalamic neurons that convey information to the
hyperstriatum dorsale) and dorsal ventricular ridge (hyperstriatum ventrale pallium. As a consequence of this, the Wulst and DVR are
and the neostriatum) indicated by different colours. rich in glutamate receptors. By contrast, the vast majority

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Avian Brains

of the neurons of the subpallium, which consists of the disease.) (see Amine neurotransmitters.) (see Nitric oxide as a
basal ganglia in mammals and birds, use the inhibitory neuronal messenger.)
neurotransmitter g-aminobutyric acid (GABA). As com- Finally, a diversity of neuropeptides is found in various
monly true in the nervous system, many neurons of the neurons throughout the brain in birds, and these appear to
cerebrum also use one or more neuropeptides as neuro- serve as modulators of the primary neurotransmitters co-
modulators of their primary neurotransmitters. This is contained in these neurons. Details of the distribution of
especially evident for the basal ganglia, in which one these neuropeptides is beyond the scope of the present
population of GABA-containing striatal neurons contains overview, but in addition to those named above they
the neuropeptides substance P and dynorphin, while include vasoactive intestinal polypeptide (VIP), neuroten-
another contains the enkephalin neuropeptides. sin, somatostatin, neuropeptide Y, cholecystokinin, calci-
(see Glutamate as a neurotransmitter.) (see GABA as a neurotrans- tonin gene-related peptide and corticotrophin-releasing
mitter.) (see Peptide neurotransmitters and hormones.) factor. In many cases, the distribution of neurons contain-
These two types of striatal neuron each make up nearly ing these neuropeptides helps to identify homologous
half of all striatal neurons and they play distinct roles in neuronal groups in birds and mammals, while in others
movement control, with the former promoting planned they show the unique brain circuits that birds have evolved.
movements and the latter inhibiting movements that Additionally, hormones such as vasopressin and oxytocin,
potentially conflict with planned movements. The striatal and such hypophysiotropic factors as adrenocorticotropic
part of the basal ganglia receives prominent glutamatergic hormone (ACTH), somatostatin and gonadotropin-re-
inputs from cortex and thalamus (which provide the leasing hormone are found in hypothalamic neurons in
information required for the striatum to play a role in birds, and again help to define functional circuits, many of
movement control) and a prominent dopaminergic input which represent common features of amniote brain
from the midbrain substantia nigra (which plays an organization. They also express steroid receptors, such as
important role in modulating the striatum according to oestrogen receptor (ER) and glucocorticoid receptor
motivational state). Owing to these two inputs, the (GR). (see Cellular neuromodulation.) (see Somatostatin.)
striatum is rich in glutamate and dopamine receptors. As (see Peptides: biological activities of small peptides.)
the neurons of the striatum and pallidum are GABAergic, (see Neurotransmitters.)
the neurons in their projection targets possess GABA
receptors. Among the targets of striatal neurons are
included other neurons of the striatum itself and the
neurons of the pallidum. Pallidal targets include thalamic Neural Circuits for Song in Bird Brains
sites projecting to motor regions of the Wulst and the
lateral spiriform nucleus of the pretectum. The lateral Passerine songbirds, such as zebra finches (which are the
spiriform nucleus is itself noteworthy because it uses both best studied of songbirds), are born in a highly altricial
GABA and enkephalin, as does its homologue in reptiles. state and are dependent on their parents, for the first 30–
(see Dopamine.) 40 days after hatching in the case of zebra finches (Bottjer,
An additional major neurotransmitter system of the 1997). Zebra finches fledge from the nest at around 20 days
telencephalon in birds is the basal forebrain cluster of of age, which is the time at which active vocal learning in
cholinergic neurons. In mammals, this group of neurons males begins. Juvenile male zebra finches must hear
has extensive projections to the cerebral cortex and plays a conspecific song from approximately 20 to 40 days of age
role in learning and memory. It is likely that they play a in order eventually to produce a song of their own, which is
similar role in birds. Degeneration of these neurons in modelled after the tutor song. Hearing conspecific song is
humans has been implicated as a contributor to the thought to result in the formation of a stable memory of the
learning and memory defects of Alzheimer disease. In tutor song which then guides the sensorimotor learning
addition, a set of cholinergic neurons with ascending needed to produce a copy of that pattern. Male zebra
projections to midbrain and diencephalon is located in the finches begin to produce their first song-related vocaliza-
rostral hindbrain, and a series of catecholaminergic tions at around 25 days, and gradually refine their
neurons with diverse projections is distributed from utterances until they achieve a close match to the tutor
forebrain to hindbrain levels in birds, as is a series of song. By 80–90 days, male zebra finches are sexually
serotonergic neurons (Reiner et al., 1994). Similar choli- mature and produce a stereotyped song pattern that, under
nergic, catecholaminergic and serotonergic neurons are normal circumstances, will be maintained without change
present in mammals and reptiles, and they appear thus to throughout adult life. In contrast, female zebra finches do
be fundamental constituents of amniote brain organiza- not demonstrate singing ability. In addition to the need to
tion. Various neurons of the avian brain, such as some of hear an external song model during early development,
the cholinergic neurons, also make the gaseous neuro- males must also hear their own vocalizations during the
transmitter nitric oxide, as is also true in mammals. period of song learning so that they can learn to adjust their
(see Acetylcholine.) (see Learning and memory.) (see Alzheimer vocal output to match the tutor song. Once a stereotyped

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Avian Brains

song is produced, adult males rehearse their song and rely birds correlates precisely with the specific song syllables
on the auditory feedback during the rehearsals to maintain heard, whereas the activity of RA neurons is more related
a stable song pattern. Song learning in other songbirds to the motor aspects of song production. Thus, the HVC–
shows similar characteristics to those in zebra finches. RA pathway appears to be part of the circuitry for
(see Bird song: steroid hormones and plasticity.) producing learned song in adult birds, with RA directly
Two functionally distinct telencephalic circuits underlie linking the telencephalic song control circuitry with
vocal learning and song production in passerine songbirds descending motor circuitry that activates the vocal
(Figure 5): (1) a direct pathway from the cortical higher musculature (Figure 5). (see Auditory processing.)
vocal centre (HVC) to a cortical motor region termed the The multisynaptic pathway connecting HVC to RA
robust nucleus of the archistriatum (RA), and (2) a (HVC–X–DLM–LMAN–RA) is necessary for song
multisynaptic pathway connecting the HVC to the RA, learning. Neurons in HVC that are distinct from those
whose components are: HVC–area X of the basal ganglia– projecting to RA project to area X in the basal ganglia,
the dorsolateral medial nucleus of the thalamus (DLM)– which relays through the thalamic nucleus DLM to the
the lateral magnocellular anterior neostriatum (LMAN)– cortical region LMAN, which in turn projects on to the
RA (Bottjer, 1997). These pathways differ in their roles, same RA neurons that receive inputs from HVC. Lesions
with the former important for adult song production and of the X–DLM–LMAN pathway have no major effect on
the latter for song learning in juveniles and maintenance in song production in adult birds, but lesions in this pathway
adults. In keeping with the fact that female songbirds do during song acquisition profoundly disrupt vocal produc-
not sing, the size of song control nuclei is several times tion in juveniles and prevent song acquisition. Thus, the
greater in males than in females, and some (such as area X) anterior forebrain song control circuit involving area X
are not even evident in females. plays a critical role in shaping song-related activity in RA
The HVC–RA circuit appears to be required for during song learning so that song output comes to match
ongoing production of stable song in adult male birds the tutor song. The area X circuit has also been shown to be
(Bottjer, 1997). HVC receives input from the auditory active during adult rehearsal of song and for the long-term
cortex (field L) in birds and is the source from which the maintenance of song (Jarvis et al., 1998). It is of interest
other song control nuclei derive their song-related auditory that similar vocal control circuitry to that in songbirds has
information. Song-related auditory responses are weak in evolved independently in the other two vocalizing orders of
avian auditory cortex but are prominent in HVC in adult birds, parrots and hummingbirds.
songbirds. The activity of HVC neurons in awake, singing

Summary
HVC
During the early part of the twentieth century, birds were
Aud thought to possess an inflexible behavioural repertoire that
CTX consisted largely of innate behaviours. Concomitant with
LMAN this view, the forebrain anatomy of birds was thought to be
RA
dominated by a hypertrophied basal ganglia and related
Cb circuitry, which at that time were thought to mediate
Area X reflexive behaviour. It is now clear that birds possess a
DLM
complex and malleable behavioural repertoire mediated by
a brain that rivals the brain of many mammals in size and
sophistication. The cerebral hemispheres of birds are
nXII
especially enlarged, and the part of them devoted to
higher-order processes such as learning and cognition,
namely the Wulst and DVR, are comparable to the
Vocal organ
cerebral cortex of mammals, which subserves similar
functions. (see Neural networks and behaviour.) (see Synaptic
Figure 5 Side view of a songbird brain showing the song control cell plasticity as a mechanism of learning.)
groups of the forebrain and their interconnections. The green arrow
indicates the serially connected structures forming the forebrain motor
circuit for song control, while the red arrows show the connections of the References
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