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THE CEREBRUM 1
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Humans do not have a well developed sense
of smell. Not only is the sense
of smell not as developed in humans as it is in most other vertebrates,
our lineage has actually been regressing from the ancestral state. In the primitive chordates (such as lancelets),
the telencephalon became the site for the processing
of olfactory stimuli (Ariens). In jawless fish, the olfactory bulbs comprise
the majority of the cerebrum. Bipolar neurons connect the olfactory bulbs to
the olfactory epithelia in jawless fish and the olfactory pathway is structured
similarly to that of higher vertebrates (Kardong). In gnathostomes, the
olfactory lobe of the cerebrum includes a bulb which is attached to the
rest of the brain by a stalk (Weichert, 1970,
p.627). The pallial
region of the cerebrum is primarily responsible for dealing with smell
in both actinopterygian and sarcopterygian
fish. (Weichert, 1970, p.627). |
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In higher primates in general, and in humans
in particular, the olfactory bulb has become greatly reduced in size.
Humans also lack the accessory olfactory bulb and (apparently) a
functional vomeronasal system.
Although the cerebrum was primarily an olfactory processing center
at its outset, our lineage has greatly enhanced the non-olfactory processing
and reduced the importance of olfaction. The anatomy, neurochemistry, development, and gene expression in the olfactory bulb is conserved in tetrapods ( Moreno, 2008). |
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Even though the cerebrum of jawless fish
is primarily a site for the processing of olfaction, it includes the two
major regions of the cerebrum present in higher vertebrates: the pallium
which forms the cerebral cortex in mammals and the subpallium
which forms the basal nuclei in mammals.
The pallium of jawless fish includes both
the medial/hippocampal region and the dorsal and
lateral/cortex region. The subpallium of jawless fish includes both the striatum and
septum (Kardong, p. 646). Regions of the hagfish telencephalon
correspond to the hippocampus, corpus striatum, part of the pallidum,
preoptic nucleus, and cerebral hemispheres of
vertebrates. In hagfish, olfactory
pathways project to the hippocampus, as in vertebrates (Hardisty, p. 316;
Ariens). Lampreys
and all higher vertebrates share a number of features such as common striatal regions, a medial olfactory nucleus, a lobus subhippocampus (Ariens, p.1247), and medial, dorsal, and lateral subdivisions
of the pallium (Butler, 1996, p. 263). |
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HAGFISH |
LAMPREY |
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Vertebrates appear to share the expression
of specific genes in the development of regions of the telencephalon. All vertebrates share the expression of Distal-less
in the subpallium ( amygdala
and basal ganglia of tetrapods), Pax6 and Tbr genes
in the pallium, Emx
in the ventral pallium (LIM homeodomain
genes are shared in the ventral pallium of tetrapods) (Medina, 2005). In gnathostomes,
there is an increase in the non-olfactory regions of the cerebrum (Romer p. 588) and the cerebral cortex is divided by a sulcus into two hemispheres. (Weichert, 1970, p.627). Olfactory fibers no longer project to all pallial areas ( In bony fish, the corpus striatum is enlarged
so that it bulges into the lateral ventricles (Weichert,
1970). The cerebrum of bony fish
and tetrapods is involved in reproductive behavior, aggression,
sex drive, parental behavior, (Prosser, 1973, p. 691), color discrimination,
and learning (Hoar,VOl. IV, 1970, p. 19-20). The lateral subpallium
of lungfish may be homologous to all the regions of the mammalian striatum
(Butler, 1996, p. 351). The basal ganglia of lungfish have 2 major groups
of projection neurons which release substance P and enkalphins
( |
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PERCH |
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LUNGFISH |
Homologous
regions between the pallial regions of lungfish
and amphibians are depicted below. |
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In amphibians, neuron migrations form an
archpallium dorsal and medial to the ancestral palaeopallium which is still dedicated to processing olfaction.
The size of the cerebral hemispheres is larger than that of fish,
although the cerebrum still seems to function primarily in olfaction (Weichert,
1970, p.627) The amount of the cerebrum which performs non-olfactory
tasks is increased over that observed in fish (Romer, Ariens). A number of characteristics are typical of
tetrapod cerebrums including the accessory olfactory bulb,
amygdaloid complex, caudate nucleus, nucleus accumbens, vomeronasal organ (VNO),
a vomeronasal nerve, set of connections involving
the stria medullaris,
amygdalo-habenularis tract, median forebrain bundle,
fornix, bed nucleus of anterior commissure,
striohypothalamic tract, primordium
pallii dorsalis (Ariens, p.1293-1311), dorsal and ventral striatopallidal systems (Greenberg, 2002), and the entopeduncular nucleus (Butler, 1996, p. 272). While non-mammalian vertebrates evolved the
ability for more complex processing, higher order functions were not necessarily
centered in the cerebrum the way they are in mammals. In fish and amphibians, the tectum of the midbrain is the primary association center of
the brain. In reptiles and birds,
both the cerebrum and tectum are centers for higher
processing. In mammals the tectum was reduced as the cerebrum became the main association
center (Romer p. 585).
Comparisons of the expression patterns of regulatory genes expressed in the forebrain indicate that differences exist between tetrapods and jawless fish but that comparable subdivisions exist among tetrapods. In tetrapods, the subpallium is defined by the expression of genes of the Dlx family, Sonic Hedgehog and Gsh 1/2 while the pallium is defined by the expression of Pax6, Emx 1/2, Neurogenin2, T-brain family members and LIM homeodomain proteins. Bony fish also express Dlx genes in the subpallium and Pax6 and Emx in the pallium. Bony fish lack Sonic Hedgehog expression in the subpallium which defines the anterior entopeduncular region and jawless fish may lack a pallium since Nkx expression in not known ( Medina, 2005). |
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FROG |
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SALAMANDER |
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Although at first glance the mammalian
cerebrum appears to have a completely different organization than that
of other amniotes (birds and reptiles), in reality early reptiles evolved
many of the regions of the amniote cerebrum
which were subsequently modified in mammals.
In amniotes, the cerebral cortex is much larger than that of amphibians
and at least partially covers the diencephalon. In reptiles, the cerebral surface is still smooth.
Some reptiles form the first neopallium at the anterior dorsal end of the cerebrum which
performs higher association functions (Weichert,
1970, p.627). Amniote
cerebrums share a number of derived characteristics. These include a dorsal pallium
composed of two divisions (medial lemnopallial
division and a lateral collopallial division;
Butler, 1996, p. 377), the division
of amygdale in to several regions (Butler, 1996, p. 455), an increase
in the size and complexity of the limbic system (Butler, 1996, p. 456),
an increase in size of the telencephalon, greater
separation of the olfactory bulbs, the lentiform
nucleus, putamen, claustrum, nucleus epibasalis, nucleus
centralis, nucleus basalis
accumbens, bed nucleus of the stria
terminalis, globus
pallidus, piriform lobe cortex (Ariens, p.1310-36), medial and lateral septal
nuclei (Ariens, p.1408), homologs
of the mammalian ventral nuclear group and posterior nuclear group, lemnothalamic and collothalamic
visual pathways (Butler, 1996, p. 391), and a more medial position of
the corpus striatum (Weichert, 1970). Amniotes evolved a medial division of dorsal
pallium and a lateral division of dorsal pallium, ventricular and subventricular
divisions of the lateral division of dorsal pallium,
and an end to the overlap between lemnothalamic
and collothalamic projections to the dorsal pallium. Lemnothalamic fibers
project through the lateral forebrain bundle to the dorsal pallium and striatum, at least some lemnothalamic
tracts are bilateral, and the lateral part of the medial division of the
dorsal pallium alone receives visual projections
from the lemnothalamus. The
medial part of the medial division of the dorsal pallium
alone receives somatosensory and limbic input
from lemnothalamus, the lemnothalamus
projects to both divisions of the dorsal pallium. Amniotes also share dorsal pallium projections to dorsal thalamus, dopaminergic
fibers to dorsal pallium, GABA-containing neurons
in both divisions of dorsal pallium, and projections
of dorsal pallium to striatum ( All amniotes seem to have similar cerebral
circuitry. The organization of
the mammalian cerebral cortex is similar to that of the dorsal ventricular
ridge in reptiles and birds (Karten, 1997). The anterior dorsal ventricular ridge is a singular
structure which seems contain the cells that,
after migrations which occur during the development of the mammalian brain,
are homologous to the basolateral amygdala
complex, the lateral neocortex, and the claustrum-endopiriform nucleus. In monotremes the
first two structures, but not the claustrum-endopiriform
nucleus, are present. In mammals,
the formation of these regions requires the Pax-6
gene and these regions in Pax-6
mutant mice are very similar to those observed in reptiles and birds ( Evidence supports the conclusion that large brains in birds increases the likelihood of adaptation to a novel type of environment (Sol, 2005). Despite the fact that the 27g echidna brain is three times the size of the 9 g platypus brain and the two are adapted for very distinct lifestyles, the regional subdivision of their brains are comparable and neither possesses a subdivision which is absent in the other. Monotreme brain organization possesses subdivisions which are lacking in therian brains and lack certain subdivisions found in therian brains. The electroreception of platypuses has not resulted in novel brain regions nor has the loss of this ancestral system resulted in a brain reorganization in echidnas (Manger, 2005). A number of animal lineages, such as crows, parrots, owls, anthropoid primates, and whales have evolved higher intelligence (Roth, 2005). Among whales, brain size can vary from 2.6 kg to 9 kg (Roth, 2005).Gibbon brains vary from 88 to 105 g and their encephalization quotient varies from 1.9 to 2.7. Capuchin monkey brains vary from 26 to 80 grams and their encephalization quotients vary from 2.4 to 4.8. Chimp brains range from 330 to 430 g with an EQ of 2.2 to 2.5. Normal human brains vary from 1250-1450 g with an EQ of 7.4-7.8, although microcephalic humans may possess brain sizes within chimp range (Roth, 2005).Humans posses about 11.5 billion cortical neurons, African elephants possess 11 billion, false killer whales possess 10.5 billion, bottlenose dolphins possess 5.8, chimps possess 6.2, gorillas 4.3, and dogs possess 0.6 billion cortical neurons (Roth, 2005). The brain has evolved as a mosaic in which the expansion of one region has led to the expansion of those specific regions connected to it. Correlated expansion of connected areas of the neocortex, cerebellar cortex, and vestibular complex is evident in recent human evolution (Whiting, 2003). Increased brain size does not necessarily result in an increased complexity in the subdivision of the brain. For example, mice possess a brain weighing 3 g which includes one more region of the locus coeruleus complex than that of the dolphin whose brain weighs 1500 g. The subdivision of brain regions is often conserved within a mammalian order. Cats and dogs possess the same nuclear subdivisions of the midbrain despite a 3 fold difference in brain size. The human brain possesses the same subdivisions of the midbrain as macaques despite a 15 fold increase in brain size. The nuclear subdivision of the thalamus seems to be conserved across primates despite brain sizes which vary from 2 to 1300 grams (Manger, 2005). Cats and ferrets possess the same subdivisions of the visual cortex despite a six fold difference in brain size (Manger, 2005).A number of mammals have evolved reduced brain sizes. An extinct island species of bovid genus Myotragus reduced its brain size by half compared to other living and fossil bovids. Reduction of brain size has occurred in multiple species in at least 8 families of bats (Niven, 2005). In general, mammals of the same order typically possess equivalent complexity in the subdivision of brain regions (Manger, 2005). The brain of human infants is only a quarter of its adult size which is much lower than the fraction of adult brain size in infant chimps (40%) and macaques (70%) (McCollum, 2006). |
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TURTLE |
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ALLIGATOR |
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CHICKEN |
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