Francis
Crick; Astonishing Hypothesis |
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"You" are the behavior of a vast assembly
of nerve cells and their associated molecules. |
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Not all neuroscientists believe that the idea of the soul is a myth -- Sir
John Eccles is the most notable exception -- but certainly the majority do. |
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Many educated
people, especially in the Western World, share the belief
that the soul is a metaphor and that there is no personal life either before conception or after death. They may call themselves atheists, agnostics, humanists, or
just lapsed believers, but they all deny
the major claims of the
traditional religions. |
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Evolution
is not a clean design. As Francis Jacob the French molecular biologist has
written, "Evolution is a tinkerer." It builds, mainly in a series of smallish steps, on what was there before. |
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The mature brain is the product of both Nature and Nurture. |
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The ability to handle complex language fluently appears
to be unique to human beings. |
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Much of the behavior
of the brain is emergent -- i.e. behavior does not exist in its separate parts, such as the individual
neurons. |
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Scientific meaning of emergent - The whole may not be the
simple sum of the separate
parts.
The whole can
be understood from (1) behavior of the parts plus (2) knowledge of how all the parts interact. |
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In the behaviorist
movement,
all behavior
had to be explained in terms of the stimulus and the response. |
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Behaviorism
was especially strong
in the United States, where was started by John B. Watson and others before
World War I. |
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Behaviorism
flourished
in the 1930s
and 1940s when B. F.
Skinner
was its most celebrated exponent. |
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Neurobiologists David Hubel and Torsten Wiesel in the late 1950s found that nerve cells in the visual cortex of the brain of an anesthetized cat showed a whole series of interesting responses when light
was shown on the cat's
opened eyes,
even though it's brain waves showed it to be more asleep them
awake.
For this and subsequent work they were awarded a Nobel Prize in 1981. |
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It is better to avoid a precise definition of consciousness. Until the problem is understood much better, a formal definition is likely to be misleading or overly restrictive. |
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A language system of the type
found in humans is not
essential for consciousness. |
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Not profitable to argue about
whether lower animals (octopus) are conscious. It is probable that
consciousness
correlates to some extent with complexity of any nervous system. |
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Attention
involves some form of short term memory. Global unity of shape, color, movement, location,
etc, may be expressed by the correlated firing of the neurons involved. Neurons that respond to the properties of a particular object fire in synchrony. Other active neurons do not fire in synchrony. |
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Necker cube. |
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Gestalt psychology -- started in Germany at around 1912. When the Nazis came to power all three left Germany for
India and the United States. |
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Gestalt laws
of grouping include proximity, similarity, continuation, and closure. |
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Gestalt law
of proximity
states that we tend to group things together that are close
to one another,
and more distant from other (similar)
objects. |
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Gestalt law
of similarity means that we group
things together if they have some obvious visual property in common, such as color or direction of movement. |
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Gestalt laws
of perception
should not be regarded as rigid laws but as useful heuristics. |
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An important
operation in vision, as the Gestaltists recognized, is to separate
figure from ground. |
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"Everyone likes to show
that philosophers are
wrong." |
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Psychology
distinguishes between "arousal" (or alertness) and "attention." |
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All forms of attention are likely to have both reflex and willed components. |
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Attention filters
out unattended events. |
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The cortex consists of two separate sheets of nerve
cells, one on each side of the
head. The sheet varies somewhat in
thickness but is typically 2 to 5 millimeters thick. |
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About 100,000
neurons per square millimeter in the cortex. |
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The hippocampus stores,
probably for a few weeks or so, new
long-term episodic memories before the
information is established more permanently in the
cortex. |
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The thalamus is divided into about two dozen
regions,
each of which is concerned with some
particular subdivision of the neocortex. Each thalamic area also receives massive connections from
the cortical areas to which it sends information. These
neocortical areas can also connect directly to other parts of the brain. |
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The hypothalamus has many small subregions whose functions are to regulate hunger, thirst,
temperature, sexual
behavior, and similar body operations. |
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The inactivity of the locus ceruleus during REM sleep may help explain why we are unable
to recall the majority of our dreams. |
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When nothing
much is happening, a neuron usually sends spikes down its axon
at a background rate between 1 and 5 Hz. |
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When a neuron becomes excited, because it receives many excitatory signals, its firing
rate increases to 50-100
Hz or more. |
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For short
intervals,
a neuron's firing rate may reach 500 Hz. |
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Neuron firing rates - background rate;
average rate; as fast as it
can -
(illustration) |
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Neurons of
the brain stem that project to the cortex use transmitters such as serotonin, norepinephrine, and dopamine. Other neurons in the brain use acetylcholine. |
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In REM
sleep, brain waves are similar to awake brain, hence the name paradoxical sleep, since the person
is asleep
but
the brain appears to be awake. |
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Two types
of brain scan: (1) those that respond to some aspect of
the static structure of the brain and (2) those that detect activity. CAT scan uses X-rays. MRI records the density of protons, especially
sensitive to water, static, do not register activity. |
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Six layers of LGN of a macque monkey. - (diagram). Each layer gets input from
only one eye. |
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Inhibitory neurons in the reticular nucleus of the thalamus. |
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Reticular nucleus of the thalamus is a thin sheet of cells
surrounding much of the thalamus. It's neurons are all inhibitory. |
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Neurons of the reticular nucleus of the thalamus
receive excitation from most of the axons passing
to and from the neocortex. Their output is mapped onto the underlying
part of the thalamus immediately beneath them. |
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If the thalamus can be described as the gateway to
the cortex,
the reticular nucleus looks like the guardian of the
gateway. |
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LGN neurons
also get input coming
back from the visual
cortex.
Many more axons coming back from the cortex than going to it. |
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Inputs from the brainstem that modulate the behavior of the thalamus and especially its reticular nucleus. |
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Major pathways within cortical
layer V1 - (illustration) |
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Parts of cortical
layer V1 show heavy
striations. (hence the name 'striate cortex') - (photo) |
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Albert Einstein, "We should make things as
simple as possible, but not
simpler." |
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Area V1 develops
so that broad features, such as which
region corresponds to the fovea, are probably laid
down by genes. Finer details are likely to be modifications made during input from the eyes, perhaps by whether the firing of
neurons are correlated. |
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Almost as many neurons project backward from V2
as project forward
from V1. The forward projection goes heavily into layer 4 of V2, whereas the backward projection to V1 avoids layer 4 altogether. |
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At least twenty distinct visual areas have been identified,
plus about seven more that are partly visual. |
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Many connections between visual areas; RGC, retinal
ganglion cells; LGN of thalamus; V1, V2, . . . ER, entorhinal cortex; HC,
hippocampus; each line in the diagram symbolizes millions of axons running in
both directions - (diagram) |
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There are many cross-connections between cortical areas at the same
level. |
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Connections
between areas in the cortex are almost always reciprocal. |
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Adjacent areas in the cortex almost always
connect to each other. |
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The general pattern from LGN to hippocampus is that each area receives several inputs from lower layers. |
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Back pathways
-- urgently need more study; may help to synchronize neuronal oscillations. |
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Whole system does not look like
a one-shot static response mechanism. Likely to operate by many transient, dynamic interactions,
conducted at a fairly fast rate. |
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Although there are many different visual regions, each of which analyzes
visual input in different and complex ways, we can locate no
single region in which the neural activity corresponds exactly to the visual picture of the world we see in front of our eyes. |
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We can see how the brain takes the picture apart, but
do not yet understand how it puts it together. How does it
construct the well-organized and detailed visual
awareness of all the objects, and the behavior of
these objects, in our visual fields? |
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A remarkable degree of functional specialization in the cortex. |
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Living cortex has the texture of a rather soft jelly. Bits of it can easily
be removed by sucking on a pipette. |
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"Blindsight" -- even with a badly damaged V1, the brain can detect some fairly simple visual stimuli and act on them, although the patient will firmly deny his awareness of them. |
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Hippocampus
and its closely associated cortical areas are not
necessary for visual
awareness. |
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Typical firing
rate of active
neurons is in the range of 100 spikes/sec. |
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Functionality
of the individual neurons are subject to much variability. Neurons are
subjected to signals that can modulate their
behavior,
and some neuron properties can change while the neurons are
active. |
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Typical neuron can have anywhere from a few hundred to many tens of thousands of inputs, and its axon projects as multitudeinously. |
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A brain does not
look even a little bit
like a general-purpose computer. |
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Different parts of the brain, even different parts of the neocortex, specialize, at least to some extent, in handling
different sorts of information. |
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Most memory appears to be stored in the very same locations that carry out current operations. |
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In 1949, Canadian psychologist Donald Hebb published a book called
Organization of Behavior. In the process of learning, one of the key factors is a modification of the strength of the neuronal connections -- the synapses. |
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Donald Hebb's rule -- "When an axon
of cell A is near enough to excite a cell B, and repeatedly
and persistently takes part in firing it, some growth process or metabolic change
takes place in one or both cells such that A's efficiency, as one of the
cells firing B, is increased." Such a mechanism, or one somewhat
resembling it, is called "Hebbian." |
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Memory is "content addressable." Any
appreciable part of the input pattern will act as an address. |
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Memory is embedded in the pattern of weights -- the strength of the connections between all the various
neurons. |
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Any one memory is distributed over many
connections. |
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Memory is superimposed, because any
one connection
can be involved in several memories. |
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Memory is robust, since altering a few connections will usually not alter its behavior very much. |
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Connections of a region of the hippocampus called CA3 do in fact look like a content-addressable network. |
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Consciousness
will correspond to a particular type of activity in a transient set of neurons that are a fraction of a much larger set. [dynamic core] |
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Awareness
is likely to involve some form of attention -- we should study the mechanism the brain uses to attend to one visual object rather than another. |
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Awareness
is likely to involve some form a very short-term
memory. |
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It seems probable that, at any moment, any particular
object In the visual field is represented by
the firing of a set of
neurons. |
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Any object
will have different characteristics (form, color, motion, etc.) that are processed in several different visual areas. |
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Seeing any one object often involves neurons in many different visual areas. |
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How the neurons in different areas temporarily become active as a unit is often described as the "binding problem." |
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An object seen is often also heard, smell, or felt. This binding must also occur across different sensory modalities. |
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Brain can sometimes be tricked
into making an incorrect binding, as when you hear the voice coming not from the ventriloquist but from his dummy. |
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Attentional mechanism makes use of correlated firing. -- what matters is not just the average rate of firing
of a neuron, but the exact moments at which each neuron fires. |
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Correlated firing -- spikes
arriving at a neuron at
the same moment will produce a
larger effect than the same number of spikes arriving at different times. |
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Theoretical requirement is that the firing of the neurons
in each set should be strongly correlated with each other, while at the same time firing of
neurons in different sets should be weakly correlated, or not at all. |
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Necker cube
-- bistable vision --
visual information coming into the eyes remains the same, but the percept
changes. |
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Cortical area MT is
that area mainly concerned with movement, but largely indifferent to color. |
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Thalamus
("gateway to the cortex") has many
fairly distinct regions, some of which are
interested in vision. |
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Superior colliculus is closely involved with the control
of eye movements, another form of
visual attention. |
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"anesthetize a patient
during an operation in such a way
that he is unaware of what is going on,
partly to spare the patient pain
and partly to prevent him from suing them." |
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"unwise to embark on
experiments on consciousness on alert people until he had obtained the
security of academic tenure." |
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In the somatosensory
system,
a weak or brief signal can influence behavior without
producing awareness, while a stronger
or longer signal of the same type can make awareness occur. |
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Awareness
may involve reentrant
pathways
as Gerald Edelman
has suggested. |
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Hippocampus
is reentrant, since
it gets most of its input from the entorhinal cortex and sends most of its output back there. |
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Theoretical arguments about reentrant
pathways to give it an
air of intellectual respectability. |
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By the cortical
system I mean the cerebral
cortex and such
regions as the thalamus
and claustrum that
are very closely associated with it. |
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Although the dendrites and axon of a neuron often extend over several layers, the layer in which its soma (the cell body) is located is probably determined
genetically during normal embryonic development. |
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Hippocampal system involved in the temporary storage, or coding, of episodic memory. |
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Long-term changes in synaptic strength. |
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Temporary maintenance of reverberating circuits. |
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Reverberation
-- associated with active short-term memory. |
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Circuit from the thalamus to the type of pyramidal neuron in cortical layer 6, which sends signals back to the same part of the thalamus. |
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Back pathway,
from layer 6 to the LGN has perhaps
five or ten times as many axons as the major forward connections from LGN to layer
4. |
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Neural pathways from the brainstem can alter the activity of the LGN during slow wave sleep. |
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It is mainly the lower cortical layers whose activity correlates with consciousness in general, and with visual
awareness in particular. |
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One example where short-term memory involves the
continued firing of neurons. Working memory for visual spatial location. |
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Claustrum - thin sheet of neurons next to lower cortical layers, project very
widely over the cortex. |
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Philosophers
have had such a poor record over the last two thousand years that they would do better to show a certain modesty rather than the lofty superiority that they usually display. |
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