Chapter 2: Cognitive Neuroscience

Notes
Cognitive Neuroscience
the study of the physiological basis of cognition

Levels of Analysis
refers to the idea that a topic can be studied in a number of different ways, with each approach
contributing its own dimension to our understanding
we can study the physiology of cognition at levels ranging from the whole brain, to structures
within the brain, to chemicals that create electrical signals within these structures
NEURONS: BASIC PRINCIPLES
neurons
create and transmit information about what we experience and know

Early Conceptions of Neurons

 nerve net
anatomists applied special stains to brain tissue, which increased the contrast between
different types of tissue within the brain. When they viewed this stained tissue under a
microscope, they saw a network they called a nerve net
This network was believed to be continuous, like a highway system in which one street
connects directly to another, but without stop signs or traffic lights
it was called continuous because the staining techniques and microscopes used during
that period could not resolve small details, and without these details the nerve net
appeared to be continuous
the nerve net provided a complex pathway for conducting signals uninterrupted through
the network

Camilo Golgi (1843-1926)

Italian anatomist
developed a staining technique in 1870s in which a thin slice of brain tissue was immersed in a
solution of silver nitrate. This technique created pictures like the one in Figure 2.2b, in which
fewer than 1 percent of the cells were stained, so they stood out from the rest of the tissue.
Ramon y Cajal (1852–1934)
his discoveries earned a Nobel Prize in 1906
recognized as "the person who made this cellular study of mental life possible”
use two techniques to investigate the nature of the nerve net
used the Golgi stain, which stained only some of the cells in a slice of brain tissue
studied tissue from the brains of newborn animals, because the density of cells in the
newborn brain is small compared with the density in the adult brain
The property of the newborn brain combined with the Golgi stain made it possible for Cajal to
clearly see that the nerve net was not continuous but was instead made up of individual units
(neurons) connected together
centerpiece of the neuron doctrine
 the idea that individual cells transmit signals in the nervous system, and that these cells
are not continuous with other cells as proposed by nerve net theory
basic parts of a neuron

cell body - metabolic center of the neuron; it contains mechanisms to keep the cell alive
dendrites - branch out from the cell body to receive signals from other neurons
axons (nerve fibers) - are usually long processes that transmit signals to other neurons

Cajal also came to some other conclusions about neurons:

There is a small gap between the end of a neuron’s axon and the dendrites or cell body of
another neuron. This gap is called a synapse
Neurons are not connected indiscriminately to other neurons but form connections only to
specific neurons. This forms groups of interconnected neurons, which together form neural
circuits
there are also neurons that are specialized to pick up information from the environment,
such as the neurons in the eye, ear, and skin. These neurons, called receptors
The Signals That Travel in Neurons
determining the exact nature of transmitted signals in neurons had to await the development of
electronic amplifiers that were powerful enough to make the extremely small electrical signals
generated by the neuron visible
Edgar Adrian
was able to record electrical signals from single sensory neuron
awarded the Nobel Prize in 1932
recorded electrical signals from single neurons using microelectrodes
small shafts of hollow glass filled with a conductive salt solution that can pick up electrical
signals at the electrode tip and conduct these signals back to a recording device
a typical setup used for recording from a single neuron. There are two electrodes: a
recording electrode, shown with its recording tip inside the neuron, and a reference
electrode, located some distance away so it is not affected by the electrical signals. The
difference in charge between the recording and reference electrodes is fed into a computer
and displayed on the computer’s screen.
resting potential
no signals in the neurons
nerve impulse
neuron's receptor is stimulated and so this nerve impulse is transmitted down the axon
As the impulse continues past the electrode, the charge inside the fiber reverses
course and starts becoming negative again, until it returns to the resting potential.
This impulse is called the action potential
action potential
each action potential travels all the way down the axon without changing its height or
shape
ideal for sending signals over a distance
 neurotransmitter
studied by other researchers as Adrian was recording from single neurons
a chemical released when the signals reach the synapse at the end of the axon
studied the relation between nerve firing and sensory experience
measured how the firing of a neuron from a receptor in the skin changed as he applied
more pressure to the skin
the shape and height of the action potential remained the same as pressured
increased, but the rate of nerve firing—that is, the number of action potentials that
traveled down the axon per second—increased
Adrian drew a connection between nerve firing and experience. He describes this
connection in his book The Basis of Sensation (1928)
if nerve impulses “are crowded closely together the sensation is intense, if they are
separated by long intervals the sensation is correspondingly feeble”
electrical signals are representing the intensity of the stimulus
the rate of neural firing is related to the intensity of stimulation, which, in turn, is related to
the magnitude of an experience, such as feeling pressure on the skin or experiencing the
brightness of a light
how is the quality of experience represented in neural firing?
quality across the senses refers to the different experience associated with each of the
senses—perceiving light for vision, sound for hearing, smells for olfaction, and so on.
quality within a particular sense, such as for vision: color, movement, an object’s shape,
or the identity of a person’s face
If all nerve impulses are basically the same whether they are caused by seeing a red fire
engine or remembering what you did last week, how can these impulses stand for different
qualities?
different qualities of stimuli, and also different aspects of experience, activate different
neurons and areas in the brain
this pioneering research on how nerve impulses represent the intensity of a stimulus, in
which he related high nerve firing to feeling greater pressure, marks the beginning of
research on neural representation

REPRESENTATION BY NEURAL FIRING

principle of neural representation
everything a person experiences is based on representations in the person’s nervous system

The Story of Neural Representation and Cognition: A Preview

 as research progressed, researchers began recording from neurons in areas outside the primary
visual area and discovered two key facts:
1. many neurons at higher levels of the visual system fire to complex stimuli like geometrical
patterns and faces
2. a specific stimulus causes neural firing that is distributed across many areas of the cortex
Feature Detectors
the idea of neurons that respond to specific qualities was brought to the forefront by a series of
papers by David Hubel and Thorsten Wiesel, which would win them the Nobel Prize in 1981
they presented a visual stimuli to cats
They found that each neuron in the visual area of the cortex responded to a specific type of
stimulation presented to a small area of the retina
they called the neurons feature detectors
neurons that respond to specific stimulus features such as orientation, movement, and length
the idea that feature detectors are linked to perception was supported by many different
experiments
experience-dependent plasticity
the structure of the brain is changed by experience
e visual cortex is an early stage of visual processing, and that vision depends on signals that
are sent from the visual cortex to other areas of the brain
Neurons That Respond to Complex Stimuli

Charles Gross
How are complex stimuli represented by the firing of neurons in the brain?
One answer to this question began to emerge in his laboratory
he recorded from single neurons in the monkey’s temporal lobe
neurons in the temporal lobe respond to complex stimuli
 neurons in another area of the temporal lobe respond to faces
hierarchal processing
progression from lower to higher areas of the brain
neurons in the visual cortex that respond to relatively simple stimuli send their axons to
higher levels of the visual system, where signals from many neurons combine and interact;
neurons at this higher level, which respond to more complex stimuli such as geometrical
objects, then send signals to even higher areas, combining and interacting further and
creating neurons that respond to even more complex stimuli such as faces

Sensory Coding
sensory code
how neurons represent various characteristics of the environment
specificity coding
the idea that an object could be represented by the firing of a specialized neuron that
responds only to that object
unlikely to be correct
an alternative to the idea of specificity coding is that a number of neurons are involved in
representing an object
population coding
representation of a particular object by the pattern of firing of a large number of neurons
advantage of population coding is that a large number of stimuli can be represented,
because large groups of neurons can create a huge number of different patterns
sparse coding
occurs when a particular object is represented by a pattern of firing of only a small group of
neurons, with the majority of neurons remaining silent
 There is also other evidence that the code for representing objects in the visual system, tones in
the auditory system, and odors in the olfactory system may involve the pattern of activity across
a relatively small number of neurons, as sparse coding suggests

LOCALIZED REPRESENTATION
specific functions are served by specific areas of the brain
Many cognitive functions are served by the cerebral cortex
Other functions are served by subcortical areas that are located below the cortex
Early evidence for localization of function came from neuropsychology
the study of the behavior of people with brain damage

Localization Determined by Neuropsychology

cortical equipotentiality
the idea that the brain operated as an indivisible whole as opposed to specialized areas

Paul Broca
in 1861, Paul Broca published work based on his study of patients who had suffered brain
damage due to strokes that caused disruption of the blood supply to the brain
These strokes caused damage to an area in the frontal lobe that came to be called Broca’s
area
 Patients with this problem—slow, labored, ungrammatical speech caused by damage to
Broca’s area—are diagnosed as having Broca’s aphasia
"Alright. . . . Uh … stroke and un. . . . I . . . huh tawanna guy. . . . H . . . h . . . hot tub and. . . . And the. . . . Two days
when uh. . . . Hos . . . uh. . . . Huh hospital and uh . . . amet . . . am . . . ambulance."
Carl Wernicke
described a number of patients who had damage to an area in their temporal lobe that came to
be called Wernicke’s area
patients produced speech that was fluent and grammatically correct but tended to be
incoherent which became called as Wernicke's aphasia
"It just suddenly had a feffort and all the feffort had gone with it. It even stepped my horn. They took them from earth
you know. They make my favorite nine to severed and now I’m a been habed by the uh stam of fortment of my
annulment which is now forever"

Patients such as this not only produce meaningless speech but are unable to
understand other people’s speech
inability to match words with their meanings, with the defining characteristic of
Wernicke’s aphasia being the absence of normal grammar

damage to the occipital lobe of the brain where the visual cortex is located resulted in
blindness, and that there was a connection between the area of the occipital lobe that was
damaged and the place in visual space where the person was blind
damage to the left part of the occipital lobe caused an area of blindness in the upper-right
part of visual space
The auditory cortex, which receives signals from the ears, is in the upper temporal lobe and is
responsible for hearing
 The somatosensory cortex, which receives signals from the skin, is in the parietal lobe and is
responsible for perceptions of touch, pressure, and pain
The frontal lobe receives signals from all of the senses and is responsible for coordination of
the senses, as well as higher cognitive functions like thinking and problem solving
prosopagnosia
another effect of brain damage on visual functioning
damage to the temporal lobe on the lower-right side of the brain
an inability to recognize faces

Localization Determined by Recording from Neurons

Another tool for demonstrating localization of function is recording from single neurons
The idea that our perception of faces is associated with a specific area of the brain is also
supported by research using the technique of brain imaging
Localization Demonstrated by Brain Imaging
brain scanning, and especially fMRI, has played an important role in understanding the
physiological basis of cognition
 Many of the brain-imaging experiments that have provided evidence for localization of function
have involved determining which brain areas were activated when people observed pictures of
different objects

Looking at Pictures
fusiform face area (FFA)
A face area has also been identified by having people in a brain scanner look at
pictures of faces
in the fusiform gyrus on the underside of the temporal lobe
 the same part of the brain that is damaged in cases of prosopagnosia
parahippocampal place area (PPA)
is activated when there's perceiving pictures representing indoor and outdoor scenes
information about spatial layout
increased activation occurs when viewing pictures both of empty rooms and of rooms
that are completely furnished
extrastriate body area (EBA)
is activated by pictures of bodies and parts of bodies (but not by faces)

Looking at Movies
Alex Huth and coworkers (2012) conducted an fMRI experiment using stimuli similar to what we see in
the environment, by having participants view film clips. Huth’s participants viewed 2 hours of film clips
while in a brain scanner. To analyze how the voxels in these participants’ brains responded to different
objects and actions in the films, Huth created a list of 1,705 different objects and action categories and
determined which categories were present in each film scene
DISTRIBUTED REPRESENTATION
different areas respond to different features of humans—illustrates a central principle of
cognition: most of our experience is multidimensional

Looking at a Face
Looking at a face triggers responses to many different aspects of the face
we also respond to:
emotional aspects
where someone is looking
how parts of the face move
how attractive a face is
whether the face is familiar
distributed representation
looking at a face activates many areas of the brain
 Cognitions, be they perceptions from looking at something, or processes such as remembering
or thinking, activate numerous, sometimes widely separated, areas of the brain

Remembering
Episodic memories are memories for events in a person’s life, like remembering what you did
yesterday
Semantic memories are memories for facts, like knowing that the capital of California is
Sacramento
 episodic and semantic memories activates different areas of the brain
remembering activates areas throughout the brain
Memories can be visual (picturing someplace you often visit), auditory (remembering a
favorite song), or olfactory (smell triggering memories for a familiar place). Memories often
have emotional components, both good and bad (thinking about someone you miss). Most
memories are combinations of many of these components, each of which activates different
areas of the brain. Memories, therefore, create a symphony of neural activity throughout
the brain

Producing and Understanding Language

Wernicke suggested that language goes beyond isolated regions to include connections
between them, and to other areas as well
explaining the physiology of language involved more than just two separate, localized language
areas
Modern researchers have shown that damage to areas outside of Broca’s and Wernicke’s areas
can cause problems in producing and understanding language
NEURAL NETWORKS
neural networks
are interconnected areas of the brain that can communicate with each other
1. There are complex structural pathways called networks that form the brain’s information
highway.
2. Within these structural pathways there are functional pathways that serve different
functions.
3. These networks operate dynamically, mirroring the dynamic nature of cognition.
4. There is a resting state of brain activity, so parts of the brain are active all the time, even
when there is no cognitive activity.

Structural Connectivity
is the brain’s “wiring diagram” created by nerve axons that connect different brain areas
Early researchers determined these connections using classical neuroanatomical techniques in
which slices of brain tissue were stained to highlight axons, which enabled them to see the
neural pathways with a microscope
recently, new techniques have been developed that make more extensive mapping of the
brain’s connections possible
 track-weighted imaging (TWI)
based on detection of how water diffuses along the length of nerve fibers

Pictures of the brain’s pathways obtained by these new techniques led to the coining of the
term connectome to indicate the “structural description of the network of elements and
connections forming the human brain”, or more simply, the “wiring diagram” of neurons in
the brain
communication depends on structural connections

Functional Connectivity
functional connectivity
determined by the extent to which neural activity in two brain areas are correlated
answers the question how is it possible to determine what parts of a neural network are
involved in different functions?
If the responses of two brain areas are correlated with each other, this means that they are
functionally connected.
One method of determining whether the responding of two areas is correlated is based on
resting-state f MRI
the fMRI response measured while a person is at rest (that is, not performing a
cognitive task)
 Resting-state functional fMRI connectivity has become one of the main methods for
determining functional connectivity.

The Dynamics of Cognition

the flow of activity within and across the functional networks in the brain changes, depending
on conditions
rapid switching between networks
changes in connectivity can also occur more slowly
 functional connectivity changes in memory networks from morning to evening as memories are
accumulated during the day and are then strengthened at night
Connectivity changes have also been reported to occur in response to eating food or drinking
coffee, with some networks being strengthened and others weakened when a person who had
fasted for a day resumed eating and drinking
Functional networks are not, therefore, simply static diagrams but involve constantly changing
activity within and across networks

The Default Mode of Network

is a network of structures that respond when a person is not involved in specific tasks
There has been a great deal of speculation and research about the purpose of the DMN
One interesting observation is that when the DMN is active, people’s minds tend to wander

Technology Determines the Questions We Can Ask