4.2 The Evolution of the Nervous System
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The information regarding the evolution of the nervous system derives
from a variety of sources, including the comparison of anatomical features
of fossils with living species thought closely to resemble such fossils. In
this way, biologists and other researchers can study the nervous systems
of living species in order to gain a picture of how the nervous systems of
ancient creatures most likely had functioned. Accordingly, after I speak
about the evolution of the cell, I trace the evolution of the nervous system
in the following progression: euglena →hydra →worm →fi sh →reptile
→mammal →primate. Thankfully, we have fossilized remains of creatures
resembling each of these types of species—including what we take to be
cells and protocells (Forterre, 2005; Pennisi, 2004; Margulis & Dolan, 2002;
Berra, 1990)—making our inferences regarding the evolution of nervous
systems reliable and probable.
Some 3.5 billion years ago, approximately 1 billion years after the earth
was formed and 12 billion or so years after the Big Bang, rudimentary onecelled
life forms—bacteria—appeared here on earth. Since the famous
Stanley Miller experiment in 1953 where four amino acids, urea, and
several fatty acids (all organic molecules) were synthesized from methane,
ammonia, hydrogen, and water, numerous other laboratory experiments, thought experiments, and hypotheses have shown that it surely is possible
to generate organic molecules from inorganic molecules (Forterre, 2005;
Pennisi, 2004; Margulis & Dolan, 2002; Schopf, 2002; Line, 2002; Balter,
2000; Margulis & Sagan, 1986; Dawkins, 1976, 1986, 1996, 2005; DeDuve,
1995, 1996; Ball, 1999; Fry, 2000; Eigen, 1992; Fenchel & Finlay, 1994).
Theorists postulate that amino acid chains, proteins, and then the organelles
of the cell would have been the fi rst things to evolve, in either a primordial
soup or some crystalline formations (Pennisi, 2004; Margulis &
Dolan, 2002; Margulis & Sagan, 1986; Mayr, 2001; Eldredge, 2001; Churchland,
1984). The cell is the fi rst kind of thing that exhibits the four properties
I have spoken of in the previous chapters, namely, internal–hierarchical
data exchange, data selectivity, informational integration, and environmental–
organismic information exchange.
However, added to these properties would be mechanisms of self-replication,
protection, and Humphrey’s (1983, 1992, 1998) distinction between
what is happening out there (hereafter, OT) versus what is happening in here
(hereafter, IH) mechanisms (also see Dawkins, 1996; Churchland, 1984).
The DNA and RNA of cells act as the mechanism of self-replication, the
cell wall acts as the mechanism of protection and encapsulation, and the
processes of the organelles themselves engage in the exchange and control
of information while interacting with environments. In light of the ideas
already presented regarding genetic variation and natural selection, once
RNA and DNA emerge, the mechanisms of evolution can do their work in
cellular processes of organisms. Nervous systems came into being by the
interplay of genetic mutation and environmental fi tness, as generation
after generation of RNA and DNA have replicated in cells over and over
again.
The evolution of the nervous system really begins with microscopic
organisms that developed a light/dark sensitivity area, most likely in primordial
waters, some 3 billion years ago. In the fi rst chapter, I spoke about
the euglena, an algae that has an eyespot and uses its fl agellum to move
toward light in order to get food. This is not yet a nervous system;
however, we witness the information exchange among the processes
within the euglena, as well as the hierarchical organization of such processes
that will be found in nervous systems. We also see the beginnings
of Humphrey’s distinction between OT and IH, since the euglena uses
basic stimulus/response mechanisms to maneuver toward food and away
from danger.
It is important to note that a kind of primitive visual system would
appear to act as the catalyst for the evolutionary development of the
The Evolution of the Visual System and Scenario Visualization 97
nervous system. In fact, light/dark sensitivity emerged among a whole
multitude of organisms in the protist and monera kingdoms (Febvre-
Chevalier, Bilbaut, Febvre, & Bone, 1989). Further refi nements of the light/
dark sensitivity area can be found in animals, either in terms of stimulus/
response mechanisms or various kinds of rudimentary and advanced visual
systems (Cronly-Dillon, 1991; Blake & Truscianko, 1990; Horridge, 1987).
For example, the photosensitive pigment cells in a worm’s skin are responsive
to light, as is the rudimentary pinhole eye of the nautilus. This speaks
to the fact that an organism’s ability to “see” is crucial for its survival,
which makes sense given that the world is bathed in sunlight. Sunlight
interacts in the world in such a way that organisms have taken advantage
of this form of energy so as to optimally engage in forms of feeding, foraging,
fl eeing, fi ghting, and so forth.
The euglena is a one-celled organism. With the arrival of multicellular
organisms having simple nervous systems came the possibility that a
variety of cells could have a variety of functions. The hydra is a cnidarian
that has a simple nervous system called a nerve net. This nerve net forms
an undifferentiated network of sensory neurons, motor neurons, and interneurons
(DeDuve, 1996; Bonner, 2000; Mackie, 1989). This was a crucial
development in the evolution of the nervous system for at least three
reasons. First, the electrochemical processes of the neuron, as well as the
multiple axonal/dendritic neuronal connections, could aid in speeding up
reactions to external stimuli, enabling the animal to be more effi cient at
hunting and/or avoiding being eaten.
Second, more complex and more effi cient forms of communication
within the animal could be achieved with the introduction of sensory
neurons, motor neurons, and interneurons that are specialized in their
functions. Sensory neurons deal with the out there of an animal, since they
are located near the periphery of an animal’s body and are concerned with
relaying external environmental information to it. Motor neurons deal
with the in here of an animal, since they aid in controlling the animal’s
bodily movements, as well as internal responses to external environmental
stimuli. Interneurons connect sensory neurons to motor neurons and act
as a kind of fi lter or buffer for the exchange of information between the
two. Response times could be decreased by specialized neurons’ performing
specifi c tasks in parallel rather than having general-purpose neurons
perform a variety of tasks. A neuron that has emerged to handle only one
kind of problem likely will be able to handle that problem swiftly and
effi ciently because it has to handle only that particular kind of problem and
no other one (also see Arp, 2008b). Further, many specialized neurons working on some problem together minimizes errors and allows systems
to perform more optimally (cf. Culler & Singh, 1999; Bechtel & Abrahamsen,
2002; Feldman & Ballard, 1982). The parallel form of processing
exhibited in the tripartite neuronal network made it through an evolutionary
sieve because of its effi ciency, and such a natural selection comports
with the general evolutionary principle of economy, namely, whatever trait
gives an organism a competitive advantage most likely will be naturally
selected as fi t for that organism and will be passed on to that organism’s
progeny. I will say more about specialized versus general-purpose processing
in the next chapter.
Third, with the introduction of this tripartite neuronal system, Humphrey’s
distinction between IH and OT clearly comes into play. This is
important for more evolved organisms that require part of the nervous
system to be devoted to maintaining homeostasis within the organism (IH)
while another part of the nervous system is attending to external stimuli
(OT). Again, a nervous system having neurons of differing types devoted
to specialized tasks could perform more effi ciently than one having generalpurpose
neurons.
The next step in evolution was for sensory cells to cluster in the heads
of simple animals like the worm. The body began to take on segmentation,
and the movement of each segment came to be regulated by nerve cell
bodies, or ganglia. A central cord of nerve fi bers connected ganglia to each
other and to the head. Also, a larger clustering of nerve cell bodies (a cerebral
ganglion) appeared in the head, where a kind of brain-like command
center for integrating sensory input and directing the body began to take
shape. It makes sense that the clustering of nerve cells together in the head
was selected for, since electrochemical information would not have as far
to travel, thus allowing for conservation of energy and faster communication
between neurons.
Fish evolved from worms, and, at fi rst, there was just a brain-like bulge
on top of the spine. Then, the nerves started to sort themselves into specialized
modules—some became sensitive to certain molecules and formed
smell modules, while others became sensitive to light and formed visual
modules. The major evolutionary advance concerning fi sh has to do with
the clear differentiation of the nervous system into the CNS and the
peripheral nervous system (PNS). The CNS consists of the brain and spinal
cord and is in constant two-way communication with the PNS, which
consists of the somatosensory, afferent somatic, visceral, motor efferent,
automatic (made up of the involuntary sympathetic and parasympathetic
systems), and somatic (voluntary) systems. This is the general layout that
The Evolution of the Visual System and Scenario Visualization 99
is found in reptiles, amphibians, and mammals as well (Audesirk et al.,
2002; Kandel et al., 2000).
The distinction between IH and OT is crucial in order to understand
more complex vertebrate nervous system functioning and behavior. Parts
of the CNS and PNS—like the hypothalamus and parasympathetic systems
in mammals—are devoted to the internal homeostasis of the organism;
these are the IH processes. Other parts of the CNS and PNS—like the LGN
in the brain and the somatic system in mammals—are devoted to reaction
to external stimuli around the organism; these are the OT processes. Ultimately,
as we will see in the evolution of primates, some of the IH processes
formed a kind of feedback loop, generating emotions and conscious
thought. Instead of merely being directed toward the maintenance of OT
and IH processes, the brain evolved abilities to feed back information
within itself that is distinct from, but analogously related to, the processes
of the CNS and PNS. It is arguable that this feedback of information
allowed for the possibility of mental states such as emotion, refl ection, and
consciousness to emerge (Humphrey, 1992, 1998).
After fi sh came into being, the processes and systems of the brains of
subsequent animals evolved at a rate that is disproportionate in relation
to the rest of the body. The evolutionary story from here on out is more
about the increasing size and complexity of the brain rather than that of
the nervous system generally. Presumably, this is the case because the
primary sensory mechanisms converge in the head of most animals, and
the brain has become a kind of command center for both the CNS and
PNS. There is an adaptive payoff to having a locus for information transfer
and exchange. As this locus, the brain can go about its business of processing
information quickly and effi ciently, utilizing neurons that are crossconnected,
working in parallel, and in close proximity to one another.
Given the brain’s ability to process information effi ciently, it has been able
to pass through one environment after another in the evolutionary sieve.
At the same time, more parts and processes have been added to the brain
because, to put it crudely, more parts mean more possibilities for effi cient
information exchange. However, it is not merely the addition of new parts
to the brain that accounts for its effi cient processing, and, later in this
section, I will make some qualifi cations about how to understand the
evolutionary addition of parts to the brain.
The evolutionary development of the brain from fi sh, through reptiles,
to mammals and primates has been explained effectively in the famous
works by MacLean (1967, 1991; cf. Sternberg, 1988; Kaas, 1987, 1993, 1995,
1996). According to MacLean’s model, the primate brain is really a three-part brain having evolved the neocortex but retaining the limbic system
found in mammals and the brainstem core found only in reptiles. The base
of the primate brain is shared with reptiles and consists of the brainstem,
reticular formation, and striate cortex. These areas are where the necessary
command centers for living are located, namely, the control of sleep and
waking, respiration, body temperature, basic automatic movements, and
the primary way stations for sensory input.
Eventually, what MacLean calls the paleomammalian cortex evolved on
top of the reptilian brainstem, allowing for more modules to develop: the
thalamus, allowing sight, smell, and hearing to be used together; the
amygdala and hippocampus, apparatuses for memory and emotions; and
the hypothalamus, making it possible for the organism to react to more
stimuli by refi ning, amending, and coordinating movements. The functioning
of the paleomammalian and reptilian cortices are somewhat analogous
to the functioning of a heart, pancreas, or kidney, since they are
organized less for thought and more for automatic action and response.
This makes sense from an evolutionary perspective. Reptiles, amphibians,
and mammals out in the wild share the common problems of having to
respond quickly to environmental stimuli so as to know whether to fi ght,
fl ee, forage, or procreate in order to survive.
Finally, in the evolutionary history of primates what MacLean calls the
neomammalian cortex (or cerebral cortex) evolved on top of the paleomammalian
and reptilian brains. This area is responsible for the fi ne-tuning of
lower functions, complex multimodular sensory associations, voluntary
motor control, abstract thinking, planning abilities, and responsiveness to
novel challenges. MacLean’s model is powerful because it not only comports
well with the fossil evidence but is consistent with experiments and
studies performed on humans, other primates, mammals, reptiles, and fi sh
(Harvey & Pagel, 1991; Kaas, 1987, 1993, 1995, 1996; Reiner, 1993; Karten
& Shimazu, 1989; Jerison, 1973, 1991, 1997; Wise, 1996; Frith, 1996;
Fuster, 1997; Sternberg, 1988; Northcutt & Kaas, 1995).
We must make at least one qualifi cation regarding MacLean’s model. It
would be an oversimplifi cation to claim simply that structures were added
onto existing structures in the evolution of the brain. It is true in the
evolution of primates that the brain expanded, thereby ultimately allowing
for humans to do such things as invent tools, develop language, and
contemplate Goldbach’s conjecture. However, it is more accurate to say
that with the increase of brain size came the addition of new brain structures
to provide new functions, as well as a reorganization of the connections
of existing brain structures to allow them to serve novel functions,
The Evolution of the Visual System and Scenario Visualization 101
and the expansion of certain structures to augment particular abilities
(Karten, 1998; Karten & Shimazu, 1989; Keverne, Martel, & Nevison,
1996).
For example, Deacon (1990) has put together a convincing case that the
six-layered mammalian neocortex is not homologous to a single structure
in a reptile but instead derives from the merger of the dorsal ventricular
plate (or pallium) and the dorsal ventricular ridge of the reptile. Such a
development would require the addition of new cortical material and the
reorganization of existing cortical material (Rakic & Kornack, 2001; Kaas,
1987, 1993; Reiner, 1993). Also, Gannon, Holloway, Broadfi eld, & Braun
(1998) and Gannon & Kheck (1999) have made a similar case regarding
the evolution of Wernicke’s area in humans as being homologous to the
planum temporale and planum parietale in macaque monkeys and chimpanzees
(cf. the recent research concerning Broca’s area in Petrides et al.,
2005).
The information regarding the evolution of the nervous system derives
from a variety of sources, including the comparison of anatomical features
of fossils with living species thought closely to resemble such fossils. In
this way, biologists and other researchers can study the nervous systems
of living species in order to gain a picture of how the nervous systems of
ancient creatures most likely had functioned. Accordingly, after I speak
about the evolution of the cell, I trace the evolution of the nervous system
in the following progression: euglena →hydra →worm →fi sh →reptile
→mammal →primate. Thankfully, we have fossilized remains of creatures
resembling each of these types of species—including what we take to be
cells and protocells (Forterre, 2005; Pennisi, 2004; Margulis & Dolan, 2002;
Berra, 1990)—making our inferences regarding the evolution of nervous
systems reliable and probable.
Some 3.5 billion years ago, approximately 1 billion years after the earth
was formed and 12 billion or so years after the Big Bang, rudimentary onecelled
life forms—bacteria—appeared here on earth. Since the famous
Stanley Miller experiment in 1953 where four amino acids, urea, and
several fatty acids (all organic molecules) were synthesized from methane,
ammonia, hydrogen, and water, numerous other laboratory experiments, thought experiments, and hypotheses have shown that it surely is possible
to generate organic molecules from inorganic molecules (Forterre, 2005;
Pennisi, 2004; Margulis & Dolan, 2002; Schopf, 2002; Line, 2002; Balter,
2000; Margulis & Sagan, 1986; Dawkins, 1976, 1986, 1996, 2005; DeDuve,
1995, 1996; Ball, 1999; Fry, 2000; Eigen, 1992; Fenchel & Finlay, 1994).
Theorists postulate that amino acid chains, proteins, and then the organelles
of the cell would have been the fi rst things to evolve, in either a primordial
soup or some crystalline formations (Pennisi, 2004; Margulis &
Dolan, 2002; Margulis & Sagan, 1986; Mayr, 2001; Eldredge, 2001; Churchland,
1984). The cell is the fi rst kind of thing that exhibits the four properties
I have spoken of in the previous chapters, namely, internal–hierarchical
data exchange, data selectivity, informational integration, and environmental–
organismic information exchange.
However, added to these properties would be mechanisms of self-replication,
protection, and Humphrey’s (1983, 1992, 1998) distinction between
what is happening out there (hereafter, OT) versus what is happening in here
(hereafter, IH) mechanisms (also see Dawkins, 1996; Churchland, 1984).
The DNA and RNA of cells act as the mechanism of self-replication, the
cell wall acts as the mechanism of protection and encapsulation, and the
processes of the organelles themselves engage in the exchange and control
of information while interacting with environments. In light of the ideas
already presented regarding genetic variation and natural selection, once
RNA and DNA emerge, the mechanisms of evolution can do their work in
cellular processes of organisms. Nervous systems came into being by the
interplay of genetic mutation and environmental fi tness, as generation
after generation of RNA and DNA have replicated in cells over and over
again.
The evolution of the nervous system really begins with microscopic
organisms that developed a light/dark sensitivity area, most likely in primordial
waters, some 3 billion years ago. In the fi rst chapter, I spoke about
the euglena, an algae that has an eyespot and uses its fl agellum to move
toward light in order to get food. This is not yet a nervous system;
however, we witness the information exchange among the processes
within the euglena, as well as the hierarchical organization of such processes
that will be found in nervous systems. We also see the beginnings
of Humphrey’s distinction between OT and IH, since the euglena uses
basic stimulus/response mechanisms to maneuver toward food and away
from danger.
It is important to note that a kind of primitive visual system would
appear to act as the catalyst for the evolutionary development of the
The Evolution of the Visual System and Scenario Visualization 97
nervous system. In fact, light/dark sensitivity emerged among a whole
multitude of organisms in the protist and monera kingdoms (Febvre-
Chevalier, Bilbaut, Febvre, & Bone, 1989). Further refi nements of the light/
dark sensitivity area can be found in animals, either in terms of stimulus/
response mechanisms or various kinds of rudimentary and advanced visual
systems (Cronly-Dillon, 1991; Blake & Truscianko, 1990; Horridge, 1987).
For example, the photosensitive pigment cells in a worm’s skin are responsive
to light, as is the rudimentary pinhole eye of the nautilus. This speaks
to the fact that an organism’s ability to “see” is crucial for its survival,
which makes sense given that the world is bathed in sunlight. Sunlight
interacts in the world in such a way that organisms have taken advantage
of this form of energy so as to optimally engage in forms of feeding, foraging,
fl eeing, fi ghting, and so forth.
The euglena is a one-celled organism. With the arrival of multicellular
organisms having simple nervous systems came the possibility that a
variety of cells could have a variety of functions. The hydra is a cnidarian
that has a simple nervous system called a nerve net. This nerve net forms
an undifferentiated network of sensory neurons, motor neurons, and interneurons
(DeDuve, 1996; Bonner, 2000; Mackie, 1989). This was a crucial
development in the evolution of the nervous system for at least three
reasons. First, the electrochemical processes of the neuron, as well as the
multiple axonal/dendritic neuronal connections, could aid in speeding up
reactions to external stimuli, enabling the animal to be more effi cient at
hunting and/or avoiding being eaten.
Second, more complex and more effi cient forms of communication
within the animal could be achieved with the introduction of sensory
neurons, motor neurons, and interneurons that are specialized in their
functions. Sensory neurons deal with the out there of an animal, since they
are located near the periphery of an animal’s body and are concerned with
relaying external environmental information to it. Motor neurons deal
with the in here of an animal, since they aid in controlling the animal’s
bodily movements, as well as internal responses to external environmental
stimuli. Interneurons connect sensory neurons to motor neurons and act
as a kind of fi lter or buffer for the exchange of information between the
two. Response times could be decreased by specialized neurons’ performing
specifi c tasks in parallel rather than having general-purpose neurons
perform a variety of tasks. A neuron that has emerged to handle only one
kind of problem likely will be able to handle that problem swiftly and
effi ciently because it has to handle only that particular kind of problem and
no other one (also see Arp, 2008b). Further, many specialized neurons working on some problem together minimizes errors and allows systems
to perform more optimally (cf. Culler & Singh, 1999; Bechtel & Abrahamsen,
2002; Feldman & Ballard, 1982). The parallel form of processing
exhibited in the tripartite neuronal network made it through an evolutionary
sieve because of its effi ciency, and such a natural selection comports
with the general evolutionary principle of economy, namely, whatever trait
gives an organism a competitive advantage most likely will be naturally
selected as fi t for that organism and will be passed on to that organism’s
progeny. I will say more about specialized versus general-purpose processing
in the next chapter.
Third, with the introduction of this tripartite neuronal system, Humphrey’s
distinction between IH and OT clearly comes into play. This is
important for more evolved organisms that require part of the nervous
system to be devoted to maintaining homeostasis within the organism (IH)
while another part of the nervous system is attending to external stimuli
(OT). Again, a nervous system having neurons of differing types devoted
to specialized tasks could perform more effi ciently than one having generalpurpose
neurons.
The next step in evolution was for sensory cells to cluster in the heads
of simple animals like the worm. The body began to take on segmentation,
and the movement of each segment came to be regulated by nerve cell
bodies, or ganglia. A central cord of nerve fi bers connected ganglia to each
other and to the head. Also, a larger clustering of nerve cell bodies (a cerebral
ganglion) appeared in the head, where a kind of brain-like command
center for integrating sensory input and directing the body began to take
shape. It makes sense that the clustering of nerve cells together in the head
was selected for, since electrochemical information would not have as far
to travel, thus allowing for conservation of energy and faster communication
between neurons.
Fish evolved from worms, and, at fi rst, there was just a brain-like bulge
on top of the spine. Then, the nerves started to sort themselves into specialized
modules—some became sensitive to certain molecules and formed
smell modules, while others became sensitive to light and formed visual
modules. The major evolutionary advance concerning fi sh has to do with
the clear differentiation of the nervous system into the CNS and the
peripheral nervous system (PNS). The CNS consists of the brain and spinal
cord and is in constant two-way communication with the PNS, which
consists of the somatosensory, afferent somatic, visceral, motor efferent,
automatic (made up of the involuntary sympathetic and parasympathetic
systems), and somatic (voluntary) systems. This is the general layout that
The Evolution of the Visual System and Scenario Visualization 99
is found in reptiles, amphibians, and mammals as well (Audesirk et al.,
2002; Kandel et al., 2000).
The distinction between IH and OT is crucial in order to understand
more complex vertebrate nervous system functioning and behavior. Parts
of the CNS and PNS—like the hypothalamus and parasympathetic systems
in mammals—are devoted to the internal homeostasis of the organism;
these are the IH processes. Other parts of the CNS and PNS—like the LGN
in the brain and the somatic system in mammals—are devoted to reaction
to external stimuli around the organism; these are the OT processes. Ultimately,
as we will see in the evolution of primates, some of the IH processes
formed a kind of feedback loop, generating emotions and conscious
thought. Instead of merely being directed toward the maintenance of OT
and IH processes, the brain evolved abilities to feed back information
within itself that is distinct from, but analogously related to, the processes
of the CNS and PNS. It is arguable that this feedback of information
allowed for the possibility of mental states such as emotion, refl ection, and
consciousness to emerge (Humphrey, 1992, 1998).
After fi sh came into being, the processes and systems of the brains of
subsequent animals evolved at a rate that is disproportionate in relation
to the rest of the body. The evolutionary story from here on out is more
about the increasing size and complexity of the brain rather than that of
the nervous system generally. Presumably, this is the case because the
primary sensory mechanisms converge in the head of most animals, and
the brain has become a kind of command center for both the CNS and
PNS. There is an adaptive payoff to having a locus for information transfer
and exchange. As this locus, the brain can go about its business of processing
information quickly and effi ciently, utilizing neurons that are crossconnected,
working in parallel, and in close proximity to one another.
Given the brain’s ability to process information effi ciently, it has been able
to pass through one environment after another in the evolutionary sieve.
At the same time, more parts and processes have been added to the brain
because, to put it crudely, more parts mean more possibilities for effi cient
information exchange. However, it is not merely the addition of new parts
to the brain that accounts for its effi cient processing, and, later in this
section, I will make some qualifi cations about how to understand the
evolutionary addition of parts to the brain.
The evolutionary development of the brain from fi sh, through reptiles,
to mammals and primates has been explained effectively in the famous
works by MacLean (1967, 1991; cf. Sternberg, 1988; Kaas, 1987, 1993, 1995,
1996). According to MacLean’s model, the primate brain is really a three-part brain having evolved the neocortex but retaining the limbic system
found in mammals and the brainstem core found only in reptiles. The base
of the primate brain is shared with reptiles and consists of the brainstem,
reticular formation, and striate cortex. These areas are where the necessary
command centers for living are located, namely, the control of sleep and
waking, respiration, body temperature, basic automatic movements, and
the primary way stations for sensory input.
Eventually, what MacLean calls the paleomammalian cortex evolved on
top of the reptilian brainstem, allowing for more modules to develop: the
thalamus, allowing sight, smell, and hearing to be used together; the
amygdala and hippocampus, apparatuses for memory and emotions; and
the hypothalamus, making it possible for the organism to react to more
stimuli by refi ning, amending, and coordinating movements. The functioning
of the paleomammalian and reptilian cortices are somewhat analogous
to the functioning of a heart, pancreas, or kidney, since they are
organized less for thought and more for automatic action and response.
This makes sense from an evolutionary perspective. Reptiles, amphibians,
and mammals out in the wild share the common problems of having to
respond quickly to environmental stimuli so as to know whether to fi ght,
fl ee, forage, or procreate in order to survive.
Finally, in the evolutionary history of primates what MacLean calls the
neomammalian cortex (or cerebral cortex) evolved on top of the paleomammalian
and reptilian brains. This area is responsible for the fi ne-tuning of
lower functions, complex multimodular sensory associations, voluntary
motor control, abstract thinking, planning abilities, and responsiveness to
novel challenges. MacLean’s model is powerful because it not only comports
well with the fossil evidence but is consistent with experiments and
studies performed on humans, other primates, mammals, reptiles, and fi sh
(Harvey & Pagel, 1991; Kaas, 1987, 1993, 1995, 1996; Reiner, 1993; Karten
& Shimazu, 1989; Jerison, 1973, 1991, 1997; Wise, 1996; Frith, 1996;
Fuster, 1997; Sternberg, 1988; Northcutt & Kaas, 1995).
We must make at least one qualifi cation regarding MacLean’s model. It
would be an oversimplifi cation to claim simply that structures were added
onto existing structures in the evolution of the brain. It is true in the
evolution of primates that the brain expanded, thereby ultimately allowing
for humans to do such things as invent tools, develop language, and
contemplate Goldbach’s conjecture. However, it is more accurate to say
that with the increase of brain size came the addition of new brain structures
to provide new functions, as well as a reorganization of the connections
of existing brain structures to allow them to serve novel functions,
The Evolution of the Visual System and Scenario Visualization 101
and the expansion of certain structures to augment particular abilities
(Karten, 1998; Karten & Shimazu, 1989; Keverne, Martel, & Nevison,
1996).
For example, Deacon (1990) has put together a convincing case that the
six-layered mammalian neocortex is not homologous to a single structure
in a reptile but instead derives from the merger of the dorsal ventricular
plate (or pallium) and the dorsal ventricular ridge of the reptile. Such a
development would require the addition of new cortical material and the
reorganization of existing cortical material (Rakic & Kornack, 2001; Kaas,
1987, 1993; Reiner, 1993). Also, Gannon, Holloway, Broadfi eld, & Braun
(1998) and Gannon & Kheck (1999) have made a similar case regarding
the evolution of Wernicke’s area in humans as being homologous to the
planum temporale and planum parietale in macaque monkeys and chimpanzees
(cf. the recent research concerning Broca’s area in Petrides et al.,
2005).