# Consciousness Studies/The Neuroscience Of Consciousness#Topological mapping and cortical columns

*\"All parts of the brain may well be involved in normal conscious
processes but the indispensable substratum of consciousness lies outside
the cerebral cortex, probably in the diencephalon\" Penfield 1937.*

*\"The brain stem-thalamocortical axis supports the state, but not the
detailed contents of consciousness, which are produced by cortex\" Baars
et al 1998.*

## Introduction

*It is recommended that readers review ../The Philosophical
Problem/ before reading the
sections on the neuroscience of consciousness.*

One of the most exciting discoveries of neuroscience is that nearly all
of the brain performs functions that are not part of conscious
experience. In everyday life we are usually unaware of breathing or
heartbeats yet there are parts of the brain dedicated to these
functions. When we pick up a pencil we have no experience of the fine
control of individual muscles yet large areas of cortex and cerebellum
implement this. Things do not appear as greyscale and then have the
colour poured into them although this strange colour addition is done in
the visual cortex. Most of the brain is non-conscious but how is the
\"ghost in the machine\", the mind, created by and linked into the
mostly non-conscious brain?

Although most of the processes in the brain are non-conscious there can
be little doubt that the output of sensory processes contribute to
experience. For example, although we do not experience the process of
adding colour to visual data in cortical area V4 we do experience
coloured forms and although we have little inkling of the hugely complex
creation of words in the temporal/frontal lobes we do experience verbal
thoughts. Our experience is an integrated output of most of the brain
processes that deal with sensation as well as dreams, thoughts and
emotions. But how and where does this experience occur?

The signals that compose phenomenal consciousness have not been
elucidated. Perhaps the least likely signals for this role are
electrical impulses in nerve fibres because they are distributed
unevenly in time and space and can even be absent for relatively long
periods. Furthermore, electrical impulses across the membranes of
neurons have an all or nothing character; they cannot be easily
superimposed on one another. There are many other possibilities however,
such as: the electrical fields on the dendrites of neurons, the fields
of chemicals spreading out from synapses, the radio-frequency emissions
of action potentials, events in the microtubules in cells, the
depolarisations of glia, the varying fields measured by EEG devices, the
quantum superposition of brain states etc\...

Phenomenal consciousness could exist in the dendritic field of ten
neurons receiving 100,000 synapses or as an oscillation of fields over
the whole brain. The substrate of phenomenal consciousness could be
staring us in the face as a state of the whole brain or be like a needle
in a haystack, lurking in a tiny region of brain, unsuspected and
undiscovered.

Given that there is no widely accepted theory of phenomenal
consciousness Crick (1994) and Crick and Koch (1998) approached the
problem of the location of the substrate of consciousness by proposing
that scientists search for the **Neural Correlates of Consciousness**.
These neural correlates consist of events in the brain that accompany
events in conscious experience.

References:

Crick, F. (1994). The Astonishing Hypothesis. New York: Scribners.

Crick, F. & Koch, C. (1998).Consciousness and Neuroscience. Cerebral
Cortex, 8:97-107, 1998
<http://www.klab.caltech.edu/~koch/crick-koch-cc-97.html>

## Neuroanatomy

### General layout of the CNS

The Central Nervous System (CNS) consists of the spinal cord, the brain
and the retina.

The CNS consists of two major groups of active cells, the **neurons**
and the **glia**. The neurons conduct short impulses of electricity
along their membranes called **\'action potentials** and encode data as
frequency modulated signals (i.e.: different intensities of stimulation
are converted into different rates of firing). The glia modify the
connections between neurons and can respond to neuron activity by a
change of voltage across their membranes. Glia also have many other
roles such as sustaining neurons and providing electrical insulation.

Neurons have three principal parts: the **cell body**, the **dendrites**
and the **axon**. Impulses flow from the cell body to the axon. The axon
can be over a metre long and bundles of axons form **nerve fibres**.
Where an axon makes contact with the dendrites or cell body of another
neuron there is a special sort of junction called a **synapse**.
Transmission of data across synapses is usually mediated by chemical
signals.

Areas of the brain where there are many cell bodies have a beige/grey
tinge and are called **grey matter**. Areas that contain mainly nerve
fibres are called **white matter**. Masses of grey matter outside of the
surface of the cerebral cortex or the cerebellum are called **nuclei**.

The brain is of central interest in consciousness studies because
consciousness persists even when the spinal cord is sectioned at the
neck.

The brain can be divided into five distinct divisions or *\'vesicles* on
the basis of embryological development. These are the myelencephalon,
metencephalon, mesencephalon, diencephalon and telencephalon (See the
illustration below).

![](Constuddivis.png "Constuddivis.png")

:   Myelencephalon: Medulla oblongata.
:   Metencephalon: pons and cerebellum.
:   Mesencephalon: midbrain (tectum containing the superior colliculus
    and inferior colliculus, red nucleus, substantia nigra, cerebellar
    peduncles.
:   Diencephalon: thalamus, epithalamus, hypothalamus, subthalamus.
:   Telencephalon: corpus striatum, cerebral hemispheres.

These divisions tend to obscure the physical anatomy of the brain which
looks like a rod of spinal cord with a swelling at the top due to the
thalamus and corpus striatum. Around the top of the rod is a globe of
deeply indented cerebral cortex and at the back there is the puckered
mass of cerebellum. The physical anatomy is shown in greater detail in
the illustration below where the thalamus and corpus striatum have been
splayed out to show more detail.

![](Constudoverbrain_-_2.png "Constudoverbrain_-_2.png")

The thalamus is a complex organ with numerous nuclei. These are listed
below:

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The location of these nuclei is shown in the illustration below:

![](constudthal.gif "constudthal.gif")

The cerebral hemispheres consist of a thin layer of nerve cell bodies on
the surface (the cerebral cortex) with a mass of white, interconnecting
fibres below (the cerebral medulla). Each hemisphere is divided into
four principle **lobes** as shown in the illustration below:

![](Main_brain_lobes.gif "Main_brain_lobes.gif")

The cortex is a set of interconnected processors. The general layout of
the cortex with the location of the processors is shown in the
illustration below:

![](Constudproc.png "Constudproc.png")

The pathways in the brain tend to preserve the **topography** of the
sense organs so that particular groups of cells on the retina, cochlear
or body have corresponding groups of cells in the thalamus or cortex.
The retina is said to have a topological mapping onto the thalamus so
that the projection of the optic nerve is said to be **retinotopic**.

Nerve fibres that go to a part of the brain are called **afferents** and
fibres that come from a part of the brain are called **efferents**.

The cortex and thalamus/striatum are intimately linked by millions of
connecting fibres and there is also a direct connection from the motor
cortex to the spinal cord.

### Sensory pathways

Information from the sense organs travels along the appropriate sensory
nerve (optic, auditory, spinal etc.) and once in the brain is divided
into three principal paths that connect either with the thalamus, the
cerebellum or the reticular formation.

There are thalamic nuclei for each broad type of sensation and these
have reciprocal connections with specific areas of cortex that deal with
the appropriate mode of sensation. The large mass of nerve fibres that
mediate the connection between the thalamus and cortex are known as the
thalamo-cortical and cortico-thalamic tracts. There tend to be more
sensory nerve fibres returning from the cortex to the thalamus than
connect from the thalamus to the cortex so it is difficult to determine
whether the cortex is the destination of sensory data or a region that
supplies extra processing power to thalamic nuclei.

The cerebellum mediates reflex control of complex movements and receives
input from most of the sense organs.

The reticular formation is a group of loosely distributed neurons in the
medulla, pons and mesencephalon. It receives a large amount of autonomic
input and also input from all the sense organs. The intralaminar nuclei
of the thalamus are the principal destination of reticular output to
higher centres. In the most primitive vertebrates the reticular
formation performs most of the higher control functions of the animal.
The reticular formation is implicated in the maintenance of sleep-wake
cycles and activates the higher centres. This activity has attracted the
label **ascending reticular activating system** (ARAS) to describe how
the activity of higher centres is controlled by reticular input. This
title is unfortunate from the point of view of consciousness studies
because it implies that conscious experience is a result of activating
the cortex when it could be due to turning on or off particular systems
all the way from the reticular formation to the cortex. Destruction of
the reticular formation leads to coma.

### Motor and output pathways

Motor control of the body below the skull is accomplished by three
principle routes.

The motor cortex of the frontal lobes and related cortex in the parietal
lobes can control movement directly via nerves known as the
cortico-spinal tract (also called the pyramidal tract). The activity of
the motor cortex is modified and controlled by a loop that passes
through the corpus striatum, the substantia nigra and the subthalamic
nucleus and returns to the cortex. These controlling nuclei are, along
with the amygdala, known as the **basal ganglia**.

The cerebellum and the corpus striatum provide complex reflex control of
the body through nerves that travel through the red nucleus and form the
rubro-spinal tract.

The vestibular nucleus, which processes signals related to balance and
posture, has direct connections with the periphery via the
vestibulo-spinal tract.

Apart from the routes for controlling motor activity there are also
other outputs from the brain, for instance the autonomic nervous system
is intimately linked with the reticular formation which has areas that
control blood pressure, respiratory rhythm etc.

## Topological mapping and cortical columns

!Sensorimotor
homunculus{width="400"}The
cerebral cortex has a highly convoluted surface that provides a large
area of tissue. The parts of the cortex that are used for motor and
sensory functions are organised so that different areas correspond to
different zones of the body. This **topological** organisation is shown
classically by a drawing of the *sensorimotor homunculus* such as that
shown on the right.

Within a given area of the cortex there are further subdivisions. For
example, the occipital cortex corresponds to the eyes of the
sensorimotor homunculus and it is further organised so that areas of the
retina have corresponding areas on the cortex. This mapping of the
layout of the retina onto the cortex is known as **topological
mapping**. It results in a corresponding mapping of the receptive field
of the eye onto the cortex. The mapping is like an image on the surface
of the brain tissue and the visual scene that is presented to a subject
can be recovered by using fMRI along with computer analysis (Miyawaki
*et al.* 2008).

The human cortex is fairly deep, containing 100-200 neurons from the
surface to the white matter. It is divided into six histological and
functional layers. These layers can be further subdivided. In 1957
Mountcastle used microelectrode measurements to show that activity of
small zones of cortex about 0.1 to 1 mm in diameter corresponded to
particular points in the receptive field. These functional columns of
cortical tissue are called **cortical columns**.

![](Corticalcol.png "Corticalcol.png")

The diagram above shows the organisation of **ocular dominance
columns**. Each column represents a particular part of the receptive
field of a single eye. The columns for left and right eyes are linked
together in lines. The lines of ocular dominance form a pattern like a
fingerprint on the surface of the cortex.

The same part of cortex can have overlapping columns for different
functions. For instance there are columns that react to particular
orientations of edges at particular places in the visual field. These
columns tend to be located together on the cortex forming a **pinwheel**
of columns that cover all orientations at a particular receptive field
position.

![](Corticalorient.GIF "Corticalorient.GIF")

There are also topologically arranged columns for colour, spatial
frequency etc.

Yoichi Miyawaki, Hajime Uchida, Okito Yamashita, Masa-aki Sato, Yusuke
Morito, Hiroki C. Tanabe, Norihiro Sadato and Yukiyasu Kamitani. (2008).
Visual Image Reconstruction from Human Brain Activity using a
Combination of Multiscale Local Image Decoders. Neuron, Volume 60, Issue
5, 10 December 2008, Pages 915-929

**Modules**

-   The neurophysiology of sensation and
    perception
-   The cortex and
    thalamus
-   Rivalries and
    synchronisation


# Consciousness Studies/The Neurophysiology Of Sensation And Perception

## Vision

### The human eye

The eye is a remarkable optical instrument that is often poorly
understood by students of consciousness. The most popular misconception
is that there is a \'focus\' within the eye through which all the light
rays pass! The purpose of this article is to describe our knowledge of
the optics of the eye so that such misconceptions can be avoided.

![](Constudtheeye.png "Constudtheeye.png")

The eye consists of several surfaces at which refraction occurs:
air-cornea, cornea-aqueous humour, aqueous humour-lens, lens-vitreous
humour. The crude image forming capability of the eye can be represented
quite accurately by the *reduced eye* model which involves a single
optical surface (air-cornea). Optometrists use more accurate models such
as the Gullstrand Schematic Eye, the Le Grand Theoretical and the
LeGrand Simplified Eye.

The lens system at the front of the eye forms an inverted image on the
retina.

The eye is about 23 mm deep from the front of the cornea to the back of
the retina. The refractive index of the components of the lens system
varies from about 1.33 to 1.39.

Light from every point of a field of view falls all over the surface of
the eye. There is no \'point eye\' and there is no ordered image between
objects in the view and the retina except on the retina. The image on
the eye has the form of an inverted mapping of 3D objects to a 2D
surface. This is also the form of conscious experience so the images on
the retinas are the closest physical analogues of phenomenal, visual,
conscious experience (see Perspective below).

### Perspective

Perspective describes how light from three dimensional objects is mapped
onto a two dimensional surface as a result of the action of lenses of
the type found in the eye.

![](Constudperspective.png "Constudperspective.png")

Perspective is used by artists to create the impression of viewing a 3D
scene. To do this they create a 2D image that is similar to the image on
the retina that would be created by the 3D scene.

![](Constudperspective2.png "Constudperspective2.png")

Naive Realists and many Direct Realists believe that the 2D perspective
view is the way things are actually arranged in the world. Of course,
things in the world differ from images because they are arranged in
three dimensions.

### Colour

The colour of an object can be represented by its **spectral power
distribution** which is a plot of the power available at each
wavelength. The unit of light power is the watt but the unit that is
used to measure subjective illumination is the **candela**. One candela
is the illumination due to light of a wavelength of 555 nanometres and a
radiant intensity of 1/683 watts per steradian in the direction being
measured. A steradian is a solid angle at the centre of sphere of one
metre radius that is subtended by one square metre of the surface. The
curious number 1/683 occurs because the unit was originally based on
light emitted from a square centimetre of molten platinum. The
wavelength of 555 nm is chosen because this is the wavelength of peak
sensitivity for light adapted (photopic) vision over a large group of
subjects. Light adapted vision is largely due to photosensitive cells in
the retina called **cones**. The candela is fixed as a standard SI Unit
for light at a wavelength of 555 nanometres.

The **lumen** is a subjective measure of the flux of light energy
passing through a solid angle (a steradian). 683 lumens of light at 555
nm are equivalent to a watt passing through the solid angle. At a
wavelength of about 520 nm only 500 lumens of luminous flux occur per
watt because the visual system is less sensitive at this wavelength. The
curve of sensitivity of the visual system to light is known as the
**V-lambda Curve**. At a wavelength of about 510 nm the same radiant
intensity is seen as being half as bright as at a wavelength of 555 nm.

![](Constudluminance.png "Constudluminance.png")

Dark adapted (scotopic) vision has a peak sensitivity at a wavelength of
507 nm and is largely due to photosensitive cells called **rods** in the
retina. Spectral Luminous Efficacy Curves are also used to express how
the sensitivity to light varies with wavelength.

Phenomenal colours are due to mixtures of **spectral colours** of
varying intensities. A **spectral colour** corresponds to a wavelength
of light found on the electromagnetic spectrum of visible light. Colours
have three attributes: **brightness**, **saturation** and **hue**. The
brightness of a colour depends on the illuminance and the reflectivity
of the surface. The saturation depends on the amount of white present,
for instance white and red make pink. The hue is similar to spectral
colour but can consist of some combinations - for instance magenta is a
hue but combines two spectral colours: red and blue. It should be noted
that experiences that contain colour are dependent on the properties of
the visual system as much as on the wavelengths of light being
reflected.

Any set of three colours that can be added together to give white are
known as **primary colours**. There are a large number of colours that
can be combined to make white, or almost any other colour. This means
that a set of surfaces that all appear white could reflect a wide range
of different wavelengths of light.

There are numerous systems for predicting how colours will combine to
make other colours; the CIE Chromaticity Diagram, the Munsell Colour
System and the Ostwald Colour System have all been used. The 1931 CIE
Chromaticity Diagram is shown below:

![](CIExy1931.png "CIExy1931.png"){width="400"}

See Chromaticity diagram for more
information.

### The retina

The retina contains photoreceptive cells called rods and cones and
several types of neurons. The rods are generally sensitive to light and
there are three varieties of cones sensitive to long, medium and short
wavelengths of light (L, M and S type cones). Some of the ganglion cells
in the retina (about 2%) are also slightly light sensitive and provide
input for the control of circadian rhythms. A schematic diagram of the
retina is shown below.

![](Constudretina.png "Constudretina.png")

The photoreceptors hyperpolarise (their membrane potential becomes more
negative) in response to illumination. Bipolar cells make direct contact
with the photoreceptors and come in two types, *on* and *off*. The
on-bipolar cells are also known as *invaginating* bipolars and the
off-bipolars as *flat bipolars*. On-bipolars depolarise when light falls
on the photoreceptors and off-bipolars hyperpolarise. Action potentials
do not occur in the bipolar or photoreceptor cells.

The retinal neurons perform considerable preprocessing before submitting
information to the brain. The network of horizontal and ganglion neurons
act to produce an output of action potentials that is sensitive to
boundaries between areas of differing illumination (edge detection) and
to motion.

Kuffler in 1953 discovered that many retinal ganglion cells are
responsive to differences in illumination on the retina. This
**centre-surround** processing is shown in the illustration below.

![](Constudcentresurr.png "Constudcentresurr.png")

The centre-surround effect is due to **lateral inhibition** by
horizontally arranged cells in the retina.

The structure of the response fields of ganglion cells is important in
everyday processing and increases the definition of boundaries in the
visual field. Sometimes it gives rise to effects that are not directly
related to the physical content of the visual field. The most famous of
these effects is the Hermann Illusion. The Hermann Grid Illusion is a
set of black squares separated by white lines. Where the white lines
cross it appears as if there are grey dots.

![](Hermann_grid.jpg "Hermann_grid.jpg")

The grey dots are due to the relative suppression of on-centre ganglion
cells where the white lines cross. This is explained in the illustration
below.

![](Constudillfld.png "Constudillfld.png")

Notice how the grey dots disappear when the crossed white lines are at
the centre of the visual field. This is due to way that ganglion cell
fields are much smaller in the fovea.

There are many other retinal illusions. White\'s illusion is
particularly strong and was believed to be due to centre-surround
activity but is now thought to have a complex origin.

![](constudwhites.gif "constudwhites.gif")

The grey lines really are the same shade of grey in the illustration.
Mach\'s Illusion is another example of a centre-surround effect.
Centre-surround effects can also occur with colour fields, red/green and
yellow/blue contrasts having a similar effect to light/dark contrasts.

Lateral inhibition and the resultant centre-surround effect increases
the number of cells that respond to boundaries and edges in the visual
field. If it did not occur then small boundaries might be missed
entirely if these fell on areas of the retina outside of the fovea. The
result of this effect is everywhere in our normal visual phenomenal
experience so not only is visual experience a mapping of 3D on to a 2D
surface, it also contains shading and brightening at edges that will not
be found by photometers that measure objective light intensities.

Photoreceptors become less responsive after continuous exposure to
bright light. This gives rise to **afterimages**. Afterimages are
usually of the opponent colour (white light gives a dark afterimage,
yellow light gives a blue afterimage, red gives a green afterimage
etc.). Afterimages when the eyes are open are generally due to a lack of
response to a particular frequency of light within the white light that
bathes the retina.

It is clear that visual phenomenal experience is related more directly
to the layout and type of activity in the retinal cells than to things
in the visual field beyond the eye.

### Visual pathways

![](Constudeyepath.png "Constudeyepath.png")

### The lateral geniculate nucleus

Retinal ganglion cells project to the Lateral Geniculate Nuclei which
are small bumps on the back of the thalamus. (Only 10-15% of the input
to the LGN comes from the retina, most (c.80%) comes from the visual
cortex). The neurons in the LGN are arranged retinotopically so preserve
the layout of events on the surface of the retina.

The LGN are arranged in 6 layers. The top two are known as Magnocellular
layers (about 100,000 neurons with large cell bodies) and the bottom
four are called Parvocellular layers (about 1,000,000 neurons with small
cell bodies). Between the main layers are the Koniocellular layers that
consist of large numbers of tiny neurons.

The left Lateral Geniculate Nucleus receives input from the right visual
field and the right LGN receives input from the left visual field. Each
nucleus receives input from both eyes but this input is segregated so
that input from the eye on the same side goes to layers 1, 3, 5 and from
the other side to layers 2,4, 6.

The magnocellular layers contain neurons that have a large receptive
field, are sensitive to contrast, a transient response and are not
colour sensitive. The parvocellular layers contains neurons that have
small receptive fields, are colour sensitive, have a prolonged response
and are less sensitive to contrast.

The LGN pathway from the retina is largely connected to the striate part
of the visual cortex (cortical area V1) via a set of fibres called the
optic radiation. There are reciprocal connections between the Thalamic
Reticular Nucleus and the LGN. The LGN are also interconnected with the
Superior Colliculus and brainstem.

The LGN may be involved in controlling which areas of the visual field
are subjected to attention (O\'Connor *et al.* 2002).

### The visual cortex

The input from the LGN goes mainly to area V1 of the cortex. The cortex
is arranged in six layers and divided up into **columns**. Each column
in the visual cortex corresponds to a particular area of the retina in
one eye. The columns are arranged in rows called **hypercolumns**. Each
column within a hypercolumn responds to a different orientation of an
optical stimulus at a given location (so responds to edges/boundaries
that are oriented in the visual field). Hypercolumns from each eye are
arranged alternately and form a small block of cortex called a
**pinwheel**. At the centre of each pinwheel are colour sensitive cells
that are usually not orientation sensitive. These coincide with the
\"blobs\" that are seen when visual cortex is viewed using cytochrome
oxidase dependent stains. It is important to note that the
\"hypercolumns\" merge into one another and respond to line stimuli that
cover an area of retina so they may be physiological rather than
anatomical entities.

The blind spot in each eye is represented by an area of visual cortex
that only receives monocular input from the other eye (Tong & Engel
2001). The effect of the blind spot is illustrated below:

![](constudblindspot.gif "constudblindspot.gif")

Normally it seems that the blindspot is \'filled in\' with background
when one eye is used. However, Lou & Chen (2003) demonstrated that
subjects could respond to quite complex figures in the blind spot,
although how far they were investigating \'blindsight\' rather than
visual experience in the blind spot is difficult to determine.

Different layers in the visual cortex have outputs that go to different
locations. Layer 6 sends nerve fibres to the Lateral Geniculate Nuclei
and thalamus, layer 5 to superior colliculus and pons, layer 2 & 3 to
other cortical areas.

There are two important outputs to other cortical areas, the **ventral
stream** and the **dorsal stream**. The ventral stream processes colour,
form and objects. It proceeds to the inferior (lower) temporal cortex.
The dorsal stream processes motion, position and spatial relationships.
It proceeds towards the parietal cortex. Lesions in the ventral stream
can result in patients knowing where an object is located but being
unable to enumerate its properties, on the other hand, lesions to the
dorsal stream can result in patients being able to label an object but
unable to tell exactly where it is located.

There is also a large output from the visual cortex back to the
thalamus, this output contains more fibres than the thalamo-cortical
input.

### Depth perception

The world is three dimensional but the image on the back of the retinas
is two dimensional. How does the brain give the subject a perception of
depth?

Depth perception relies on **cues** which are data about the
displacement of things relative to the body. These cues consist of:

-   the convergence of the eyes
-   the accommodation of the lens
-   binocular disparity -the difference between the images on the
    retinas- this was first suggested by Wheatstone.
-   motion parallax - distant objects move slower when the observer
    moves - first suggested by Helmholtz.
-   optical flow - the rate of expansion/contraction of a scene with
    movement towards or away from it (Lee & Aronson 1974).
-   binocular occlusion - parts of a scene are invisible to each eye.
-   body motion provides cues about near objects.
-   vanishing points - the convergence of parallel lines.
-   numerous other cues such as size constancy, texture etc.

Binocular disparity has been most extensively studied as a source of
depth cues. When the eyes converge to focus on an object in from of them
there is very little disparity in the images of that object on the two
retinas. The angle at the object formed between the lines that project
back to the pupils is known as the **vergence** at the object. The
sphere where all objects have the same vergence is known as the
**horopter**.

![](Constudverge.png "Constudverge.png")

When the disparity between the retinas is small a single image occurs in
phenomenal experience which is accompanied by a sensation of objects
with depth. This is known as **stereopsis**. If the disparity between
the retinas is large double vision ensues, this is known as
**diplopia**. The curious feature of stereopsis is that we can see no
more of the object than is visible on the retinas and certainly cannot
see behind the object. Stereopsis is more like a stretching of 2D space
than actual 3D.

The **empirical horopter** is a zone where things are seen without
diplopia. The empirical and **Veith Muller** (geometric) horopters are
different. This difference is the result of both processing by the CNS
and optical factors.

**Physiological diplopia** refers to the stimulation of receptors in
different parts of the retinas of the two eyes by the same object.
Physiological diplopia does not always give rise to subjective diplopia,
objects close to the empirical horopter do not give rise to double
vision and the zone in which this occurs is known as **Panum\'s Fusion
Area**. It is widest for objects that are distributed away from the nose
(with \'temporal\' locations) and for objects that are slow moving and
poorly focussed.

In the review by Cutting and Vishton (1995) the contributions of each
type of cue is discussed. Cutting and Vishton also present evidence that
there are several zones of depth perception that are informed by
different sets of cues. These are **personal space**, which is the zone
of things within arms reach, **action space**, which is the zone in
which we interact and where our motions have a large impact on the
perceived layout, and **vista space** which is the zone beyond about 30m
that is informed by long range cues.

The interesting feature of 3D perceptual space is that it is not seen.
The sides of a solid object appear as intrusions or lateral extensions
in 2D space, when we close an eye that has access to the side of the
object and then open it again the side grows out into 2D space. The lack
of \'seeing\' depth is also evident when we close one eye when looking
at a vista - nothing seems to change even though stereopsis has gone.
This leaves the problem of what it is that constitutes the \'feeling\'
of depth. We have feelings that we can fall into space or move into it
or around in it. Depth seems to be defined by premotor modelling and the
potential for occupancy by our bodies and limbs. As such it involves
qualia that are different from those of vision and more akin to those
that accompany movement, as an example, if you reach out to touch
something, move the hand back, then consider the distance to the object
it is evident that a feeling of the movement is still present. Is depth
a quale of movement modelled during the extended present of perception?

-   Cutting, J.E. & Vishton, P.M. (1995) Perceiving layout and knowing
    distances: The integration, relative potency, and contextual use of
    different information about depth. In W. Epstein & S. Rogers (eds.)
    Handbook of perception and cognition, Vol 5; Perception of space and
    motion. (pp. 69-117). San Diego, CA: Academic Press.
    <http://pmvish.people.wm.edu/cutting&vishton1995.pdf>

```{=html}
<!-- -->
```
-   Gregory, R.L. (1997). Knowledge in perception and illusion. From:
    Phil. Trans. R. Soc. Lond. B (1997) 352, 1121--1128.
    <http://www.richardgregory.org/papers/knowl_illusion/knowledge-in-perception.pdf>

```{=html}
<!-- -->
```
-   Hubel, D.H. (1981). EVOLUTION OF IDEAS ON THE PRIMARY VISUAL CORTEX,
    1955-1978: A BIASED HISTORICAL ACCOUNT.
    <http://nobelprize.org/medicine/laureates/1981/hubel-lecture.pdf>

```{=html}
<!-- -->
```
-   Lou, L. & Chen, J. (2003). Attention and Blind-Spot Phenomenology.
    PSYCHE, 9(02), January 2003.
    <http://psyche.cs.monash.edu.au/v9/psyche-9-02-lou.html>

```{=html}
<!-- -->
```
-   O\'Connor, D.H., Fukui, M.M., Pinsk, M.A. & Kastner, S. (2002).
    Attention modulates responses in the human lateral geniculate
    nucleus. Nature Neuroscience 5, 1203 - 1209 (2002).
    <http://www.nature.com/neuro/journal/v5/n11/full/nn957.html#B34>

```{=html}
<!-- -->
```
-   Tong, F., & Engel, S. A. (2001). Interocular rivalry revealed in the
    human cortical blind-spot representation. Nature, 411, 195-199.
    <http://www.psy.vanderbilt.edu/tonglab/publications/Tong&Engel2001.pdf>

```{=html}
<!-- -->
```
-   Tong, F. (2003). PRIMARY VISUAL CORTEX AND VISUAL AWARENESS. Nature
    Reviews of Neuroscience. 4:219-229.
    <http://www.psy.vanderbilt.edu/tonglab/publications/Tong_NRN2003.pdf>

**Modules**

-   The cortex and
    thalamus
-   Rivalries and
    synchronisation


# Consciousness Studies/The Neurophysiology Of Sensation And Perception#Vision

## Vision

### The human eye

The eye is a remarkable optical instrument that is often poorly
understood by students of consciousness. The most popular misconception
is that there is a \'focus\' within the eye through which all the light
rays pass! The purpose of this article is to describe our knowledge of
the optics of the eye so that such misconceptions can be avoided.

![](Constudtheeye.png "Constudtheeye.png")

The eye consists of several surfaces at which refraction occurs:
air-cornea, cornea-aqueous humour, aqueous humour-lens, lens-vitreous
humour. The crude image forming capability of the eye can be represented
quite accurately by the *reduced eye* model which involves a single
optical surface (air-cornea). Optometrists use more accurate models such
as the Gullstrand Schematic Eye, the Le Grand Theoretical and the
LeGrand Simplified Eye.

The lens system at the front of the eye forms an inverted image on the
retina.

The eye is about 23 mm deep from the front of the cornea to the back of
the retina. The refractive index of the components of the lens system
varies from about 1.33 to 1.39.

Light from every point of a field of view falls all over the surface of
the eye. There is no \'point eye\' and there is no ordered image between
objects in the view and the retina except on the retina. The image on
the eye has the form of an inverted mapping of 3D objects to a 2D
surface. This is also the form of conscious experience so the images on
the retinas are the closest physical analogues of phenomenal, visual,
conscious experience (see Perspective below).

### Perspective

Perspective describes how light from three dimensional objects is mapped
onto a two dimensional surface as a result of the action of lenses of
the type found in the eye.

![](Constudperspective.png "Constudperspective.png")

Perspective is used by artists to create the impression of viewing a 3D
scene. To do this they create a 2D image that is similar to the image on
the retina that would be created by the 3D scene.

![](Constudperspective2.png "Constudperspective2.png")

Naive Realists and many Direct Realists believe that the 2D perspective
view is the way things are actually arranged in the world. Of course,
things in the world differ from images because they are arranged in
three dimensions.

### Colour

The colour of an object can be represented by its **spectral power
distribution** which is a plot of the power available at each
wavelength. The unit of light power is the watt but the unit that is
used to measure subjective illumination is the **candela**. One candela
is the illumination due to light of a wavelength of 555 nanometres and a
radiant intensity of 1/683 watts per steradian in the direction being
measured. A steradian is a solid angle at the centre of sphere of one
metre radius that is subtended by one square metre of the surface. The
curious number 1/683 occurs because the unit was originally based on
light emitted from a square centimetre of molten platinum. The
wavelength of 555 nm is chosen because this is the wavelength of peak
sensitivity for light adapted (photopic) vision over a large group of
subjects. Light adapted vision is largely due to photosensitive cells in
the retina called **cones**. The candela is fixed as a standard SI Unit
for light at a wavelength of 555 nanometres.

The **lumen** is a subjective measure of the flux of light energy
passing through a solid angle (a steradian). 683 lumens of light at 555
nm are equivalent to a watt passing through the solid angle. At a
wavelength of about 520 nm only 500 lumens of luminous flux occur per
watt because the visual system is less sensitive at this wavelength. The
curve of sensitivity of the visual system to light is known as the
**V-lambda Curve**. At a wavelength of about 510 nm the same radiant
intensity is seen as being half as bright as at a wavelength of 555 nm.

![](Constudluminance.png "Constudluminance.png")

Dark adapted (scotopic) vision has a peak sensitivity at a wavelength of
507 nm and is largely due to photosensitive cells called **rods** in the
retina. Spectral Luminous Efficacy Curves are also used to express how
the sensitivity to light varies with wavelength.

Phenomenal colours are due to mixtures of **spectral colours** of
varying intensities. A **spectral colour** corresponds to a wavelength
of light found on the electromagnetic spectrum of visible light. Colours
have three attributes: **brightness**, **saturation** and **hue**. The
brightness of a colour depends on the illuminance and the reflectivity
of the surface. The saturation depends on the amount of white present,
for instance white and red make pink. The hue is similar to spectral
colour but can consist of some combinations - for instance magenta is a
hue but combines two spectral colours: red and blue. It should be noted
that experiences that contain colour are dependent on the properties of
the visual system as much as on the wavelengths of light being
reflected.

Any set of three colours that can be added together to give white are
known as **primary colours**. There are a large number of colours that
can be combined to make white, or almost any other colour. This means
that a set of surfaces that all appear white could reflect a wide range
of different wavelengths of light.

There are numerous systems for predicting how colours will combine to
make other colours; the CIE Chromaticity Diagram, the Munsell Colour
System and the Ostwald Colour System have all been used. The 1931 CIE
Chromaticity Diagram is shown below:

![](CIExy1931.png "CIExy1931.png"){width="400"}

See Chromaticity diagram for more
information.

### The retina

The retina contains photoreceptive cells called rods and cones and
several types of neurons. The rods are generally sensitive to light and
there are three varieties of cones sensitive to long, medium and short
wavelengths of light (L, M and S type cones). Some of the ganglion cells
in the retina (about 2%) are also slightly light sensitive and provide
input for the control of circadian rhythms. A schematic diagram of the
retina is shown below.

![](Constudretina.png "Constudretina.png")

The photoreceptors hyperpolarise (their membrane potential becomes more
negative) in response to illumination. Bipolar cells make direct contact
with the photoreceptors and come in two types, *on* and *off*. The
on-bipolar cells are also known as *invaginating* bipolars and the
off-bipolars as *flat bipolars*. On-bipolars depolarise when light falls
on the photoreceptors and off-bipolars hyperpolarise. Action potentials
do not occur in the bipolar or photoreceptor cells.

The retinal neurons perform considerable preprocessing before submitting
information to the brain. The network of horizontal and ganglion neurons
act to produce an output of action potentials that is sensitive to
boundaries between areas of differing illumination (edge detection) and
to motion.

Kuffler in 1953 discovered that many retinal ganglion cells are
responsive to differences in illumination on the retina. This
**centre-surround** processing is shown in the illustration below.

![](Constudcentresurr.png "Constudcentresurr.png")

The centre-surround effect is due to **lateral inhibition** by
horizontally arranged cells in the retina.

The structure of the response fields of ganglion cells is important in
everyday processing and increases the definition of boundaries in the
visual field. Sometimes it gives rise to effects that are not directly
related to the physical content of the visual field. The most famous of
these effects is the Hermann Illusion. The Hermann Grid Illusion is a
set of black squares separated by white lines. Where the white lines
cross it appears as if there are grey dots.

![](Hermann_grid.jpg "Hermann_grid.jpg")

The grey dots are due to the relative suppression of on-centre ganglion
cells where the white lines cross. This is explained in the illustration
below.

![](Constudillfld.png "Constudillfld.png")

Notice how the grey dots disappear when the crossed white lines are at
the centre of the visual field. This is due to way that ganglion cell
fields are much smaller in the fovea.

There are many other retinal illusions. White\'s illusion is
particularly strong and was believed to be due to centre-surround
activity but is now thought to have a complex origin.

![](constudwhites.gif "constudwhites.gif")

The grey lines really are the same shade of grey in the illustration.
Mach\'s Illusion is another example of a centre-surround effect.
Centre-surround effects can also occur with colour fields, red/green and
yellow/blue contrasts having a similar effect to light/dark contrasts.

Lateral inhibition and the resultant centre-surround effect increases
the number of cells that respond to boundaries and edges in the visual
field. If it did not occur then small boundaries might be missed
entirely if these fell on areas of the retina outside of the fovea. The
result of this effect is everywhere in our normal visual phenomenal
experience so not only is visual experience a mapping of 3D on to a 2D
surface, it also contains shading and brightening at edges that will not
be found by photometers that measure objective light intensities.

Photoreceptors become less responsive after continuous exposure to
bright light. This gives rise to **afterimages**. Afterimages are
usually of the opponent colour (white light gives a dark afterimage,
yellow light gives a blue afterimage, red gives a green afterimage
etc.). Afterimages when the eyes are open are generally due to a lack of
response to a particular frequency of light within the white light that
bathes the retina.

It is clear that visual phenomenal experience is related more directly
to the layout and type of activity in the retinal cells than to things
in the visual field beyond the eye.

### Visual pathways

![](Constudeyepath.png "Constudeyepath.png")

### The lateral geniculate nucleus

Retinal ganglion cells project to the Lateral Geniculate Nuclei which
are small bumps on the back of the thalamus. (Only 10-15% of the input
to the LGN comes from the retina, most (c.80%) comes from the visual
cortex). The neurons in the LGN are arranged retinotopically so preserve
the layout of events on the surface of the retina.

The LGN are arranged in 6 layers. The top two are known as Magnocellular
layers (about 100,000 neurons with large cell bodies) and the bottom
four are called Parvocellular layers (about 1,000,000 neurons with small
cell bodies). Between the main layers are the Koniocellular layers that
consist of large numbers of tiny neurons.

The left Lateral Geniculate Nucleus receives input from the right visual
field and the right LGN receives input from the left visual field. Each
nucleus receives input from both eyes but this input is segregated so
that input from the eye on the same side goes to layers 1, 3, 5 and from
the other side to layers 2,4, 6.

The magnocellular layers contain neurons that have a large receptive
field, are sensitive to contrast, a transient response and are not
colour sensitive. The parvocellular layers contains neurons that have
small receptive fields, are colour sensitive, have a prolonged response
and are less sensitive to contrast.

The LGN pathway from the retina is largely connected to the striate part
of the visual cortex (cortical area V1) via a set of fibres called the
optic radiation. There are reciprocal connections between the Thalamic
Reticular Nucleus and the LGN. The LGN are also interconnected with the
Superior Colliculus and brainstem.

The LGN may be involved in controlling which areas of the visual field
are subjected to attention (O\'Connor *et al.* 2002).

### The visual cortex

The input from the LGN goes mainly to area V1 of the cortex. The cortex
is arranged in six layers and divided up into **columns**. Each column
in the visual cortex corresponds to a particular area of the retina in
one eye. The columns are arranged in rows called **hypercolumns**. Each
column within a hypercolumn responds to a different orientation of an
optical stimulus at a given location (so responds to edges/boundaries
that are oriented in the visual field). Hypercolumns from each eye are
arranged alternately and form a small block of cortex called a
**pinwheel**. At the centre of each pinwheel are colour sensitive cells
that are usually not orientation sensitive. These coincide with the
\"blobs\" that are seen when visual cortex is viewed using cytochrome
oxidase dependent stains. It is important to note that the
\"hypercolumns\" merge into one another and respond to line stimuli that
cover an area of retina so they may be physiological rather than
anatomical entities.

The blind spot in each eye is represented by an area of visual cortex
that only receives monocular input from the other eye (Tong & Engel
2001). The effect of the blind spot is illustrated below:

![](constudblindspot.gif "constudblindspot.gif")

Normally it seems that the blindspot is \'filled in\' with background
when one eye is used. However, Lou & Chen (2003) demonstrated that
subjects could respond to quite complex figures in the blind spot,
although how far they were investigating \'blindsight\' rather than
visual experience in the blind spot is difficult to determine.

Different layers in the visual cortex have outputs that go to different
locations. Layer 6 sends nerve fibres to the Lateral Geniculate Nuclei
and thalamus, layer 5 to superior colliculus and pons, layer 2 & 3 to
other cortical areas.

There are two important outputs to other cortical areas, the **ventral
stream** and the **dorsal stream**. The ventral stream processes colour,
form and objects. It proceeds to the inferior (lower) temporal cortex.
The dorsal stream processes motion, position and spatial relationships.
It proceeds towards the parietal cortex. Lesions in the ventral stream
can result in patients knowing where an object is located but being
unable to enumerate its properties, on the other hand, lesions to the
dorsal stream can result in patients being able to label an object but
unable to tell exactly where it is located.

There is also a large output from the visual cortex back to the
thalamus, this output contains more fibres than the thalamo-cortical
input.

### Depth perception

The world is three dimensional but the image on the back of the retinas
is two dimensional. How does the brain give the subject a perception of
depth?

Depth perception relies on **cues** which are data about the
displacement of things relative to the body. These cues consist of:

-   the convergence of the eyes
-   the accommodation of the lens
-   binocular disparity -the difference between the images on the
    retinas- this was first suggested by Wheatstone.
-   motion parallax - distant objects move slower when the observer
    moves - first suggested by Helmholtz.
-   optical flow - the rate of expansion/contraction of a scene with
    movement towards or away from it (Lee & Aronson 1974).
-   binocular occlusion - parts of a scene are invisible to each eye.
-   body motion provides cues about near objects.
-   vanishing points - the convergence of parallel lines.
-   numerous other cues such as size constancy, texture etc.

Binocular disparity has been most extensively studied as a source of
depth cues. When the eyes converge to focus on an object in from of them
there is very little disparity in the images of that object on the two
retinas. The angle at the object formed between the lines that project
back to the pupils is known as the **vergence** at the object. The
sphere where all objects have the same vergence is known as the
**horopter**.

![](Constudverge.png "Constudverge.png")

When the disparity between the retinas is small a single image occurs in
phenomenal experience which is accompanied by a sensation of objects
with depth. This is known as **stereopsis**. If the disparity between
the retinas is large double vision ensues, this is known as
**diplopia**. The curious feature of stereopsis is that we can see no
more of the object than is visible on the retinas and certainly cannot
see behind the object. Stereopsis is more like a stretching of 2D space
than actual 3D.

The **empirical horopter** is a zone where things are seen without
diplopia. The empirical and **Veith Muller** (geometric) horopters are
different. This difference is the result of both processing by the CNS
and optical factors.

**Physiological diplopia** refers to the stimulation of receptors in
different parts of the retinas of the two eyes by the same object.
Physiological diplopia does not always give rise to subjective diplopia,
objects close to the empirical horopter do not give rise to double
vision and the zone in which this occurs is known as **Panum\'s Fusion
Area**. It is widest for objects that are distributed away from the nose
(with \'temporal\' locations) and for objects that are slow moving and
poorly focussed.

In the review by Cutting and Vishton (1995) the contributions of each
type of cue is discussed. Cutting and Vishton also present evidence that
there are several zones of depth perception that are informed by
different sets of cues. These are **personal space**, which is the zone
of things within arms reach, **action space**, which is the zone in
which we interact and where our motions have a large impact on the
perceived layout, and **vista space** which is the zone beyond about 30m
that is informed by long range cues.

The interesting feature of 3D perceptual space is that it is not seen.
The sides of a solid object appear as intrusions or lateral extensions
in 2D space, when we close an eye that has access to the side of the
object and then open it again the side grows out into 2D space. The lack
of \'seeing\' depth is also evident when we close one eye when looking
at a vista - nothing seems to change even though stereopsis has gone.
This leaves the problem of what it is that constitutes the \'feeling\'
of depth. We have feelings that we can fall into space or move into it
or around in it. Depth seems to be defined by premotor modelling and the
potential for occupancy by our bodies and limbs. As such it involves
qualia that are different from those of vision and more akin to those
that accompany movement, as an example, if you reach out to touch
something, move the hand back, then consider the distance to the object
it is evident that a feeling of the movement is still present. Is depth
a quale of movement modelled during the extended present of perception?

-   Cutting, J.E. & Vishton, P.M. (1995) Perceiving layout and knowing
    distances: The integration, relative potency, and contextual use of
    different information about depth. In W. Epstein & S. Rogers (eds.)
    Handbook of perception and cognition, Vol 5; Perception of space and
    motion. (pp. 69-117). San Diego, CA: Academic Press.
    <http://pmvish.people.wm.edu/cutting&vishton1995.pdf>

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```
-   Gregory, R.L. (1997). Knowledge in perception and illusion. From:
    Phil. Trans. R. Soc. Lond. B (1997) 352, 1121--1128.
    <http://www.richardgregory.org/papers/knowl_illusion/knowledge-in-perception.pdf>

```{=html}
<!-- -->
```
-   Hubel, D.H. (1981). EVOLUTION OF IDEAS ON THE PRIMARY VISUAL CORTEX,
    1955-1978: A BIASED HISTORICAL ACCOUNT.
    <http://nobelprize.org/medicine/laureates/1981/hubel-lecture.pdf>

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```
-   Lou, L. & Chen, J. (2003). Attention and Blind-Spot Phenomenology.
    PSYCHE, 9(02), January 2003.
    <http://psyche.cs.monash.edu.au/v9/psyche-9-02-lou.html>

```{=html}
<!-- -->
```
-   O\'Connor, D.H., Fukui, M.M., Pinsk, M.A. & Kastner, S. (2002).
    Attention modulates responses in the human lateral geniculate
    nucleus. Nature Neuroscience 5, 1203 - 1209 (2002).
    <http://www.nature.com/neuro/journal/v5/n11/full/nn957.html#B34>

```{=html}
<!-- -->
```
-   Tong, F., & Engel, S. A. (2001). Interocular rivalry revealed in the
    human cortical blind-spot representation. Nature, 411, 195-199.
    <http://www.psy.vanderbilt.edu/tonglab/publications/Tong&Engel2001.pdf>

```{=html}
<!-- -->
```
-   Tong, F. (2003). PRIMARY VISUAL CORTEX AND VISUAL AWARENESS. Nature
    Reviews of Neuroscience. 4:219-229.
    <http://www.psy.vanderbilt.edu/tonglab/publications/Tong_NRN2003.pdf>

**Modules**

-   The cortex and
    thalamus
-   Rivalries and
    synchronisation


# Consciousness Studies/The Neurophysiology Of Sensation And Perception#Depth perception

## Vision

### The human eye

The eye is a remarkable optical instrument that is often poorly
understood by students of consciousness. The most popular misconception
is that there is a \'focus\' within the eye through which all the light
rays pass! The purpose of this article is to describe our knowledge of
the optics of the eye so that such misconceptions can be avoided.

![](Constudtheeye.png "Constudtheeye.png")

The eye consists of several surfaces at which refraction occurs:
air-cornea, cornea-aqueous humour, aqueous humour-lens, lens-vitreous
humour. The crude image forming capability of the eye can be represented
quite accurately by the *reduced eye* model which involves a single
optical surface (air-cornea). Optometrists use more accurate models such
as the Gullstrand Schematic Eye, the Le Grand Theoretical and the
LeGrand Simplified Eye.

The lens system at the front of the eye forms an inverted image on the
retina.

The eye is about 23 mm deep from the front of the cornea to the back of
the retina. The refractive index of the components of the lens system
varies from about 1.33 to 1.39.

Light from every point of a field of view falls all over the surface of
the eye. There is no \'point eye\' and there is no ordered image between
objects in the view and the retina except on the retina. The image on
the eye has the form of an inverted mapping of 3D objects to a 2D
surface. This is also the form of conscious experience so the images on
the retinas are the closest physical analogues of phenomenal, visual,
conscious experience (see Perspective below).

### Perspective

Perspective describes how light from three dimensional objects is mapped
onto a two dimensional surface as a result of the action of lenses of
the type found in the eye.

![](Constudperspective.png "Constudperspective.png")

Perspective is used by artists to create the impression of viewing a 3D
scene. To do this they create a 2D image that is similar to the image on
the retina that would be created by the 3D scene.

![](Constudperspective2.png "Constudperspective2.png")

Naive Realists and many Direct Realists believe that the 2D perspective
view is the way things are actually arranged in the world. Of course,
things in the world differ from images because they are arranged in
three dimensions.

### Colour

The colour of an object can be represented by its **spectral power
distribution** which is a plot of the power available at each
wavelength. The unit of light power is the watt but the unit that is
used to measure subjective illumination is the **candela**. One candela
is the illumination due to light of a wavelength of 555 nanometres and a
radiant intensity of 1/683 watts per steradian in the direction being
measured. A steradian is a solid angle at the centre of sphere of one
metre radius that is subtended by one square metre of the surface. The
curious number 1/683 occurs because the unit was originally based on
light emitted from a square centimetre of molten platinum. The
wavelength of 555 nm is chosen because this is the wavelength of peak
sensitivity for light adapted (photopic) vision over a large group of
subjects. Light adapted vision is largely due to photosensitive cells in
the retina called **cones**. The candela is fixed as a standard SI Unit
for light at a wavelength of 555 nanometres.

The **lumen** is a subjective measure of the flux of light energy
passing through a solid angle (a steradian). 683 lumens of light at 555
nm are equivalent to a watt passing through the solid angle. At a
wavelength of about 520 nm only 500 lumens of luminous flux occur per
watt because the visual system is less sensitive at this wavelength. The
curve of sensitivity of the visual system to light is known as the
**V-lambda Curve**. At a wavelength of about 510 nm the same radiant
intensity is seen as being half as bright as at a wavelength of 555 nm.

![](Constudluminance.png "Constudluminance.png")

Dark adapted (scotopic) vision has a peak sensitivity at a wavelength of
507 nm and is largely due to photosensitive cells called **rods** in the
retina. Spectral Luminous Efficacy Curves are also used to express how
the sensitivity to light varies with wavelength.

Phenomenal colours are due to mixtures of **spectral colours** of
varying intensities. A **spectral colour** corresponds to a wavelength
of light found on the electromagnetic spectrum of visible light. Colours
have three attributes: **brightness**, **saturation** and **hue**. The
brightness of a colour depends on the illuminance and the reflectivity
of the surface. The saturation depends on the amount of white present,
for instance white and red make pink. The hue is similar to spectral
colour but can consist of some combinations - for instance magenta is a
hue but combines two spectral colours: red and blue. It should be noted
that experiences that contain colour are dependent on the properties of
the visual system as much as on the wavelengths of light being
reflected.

Any set of three colours that can be added together to give white are
known as **primary colours**. There are a large number of colours that
can be combined to make white, or almost any other colour. This means
that a set of surfaces that all appear white could reflect a wide range
of different wavelengths of light.

There are numerous systems for predicting how colours will combine to
make other colours; the CIE Chromaticity Diagram, the Munsell Colour
System and the Ostwald Colour System have all been used. The 1931 CIE
Chromaticity Diagram is shown below:

![](CIExy1931.png "CIExy1931.png"){width="400"}

See Chromaticity diagram for more
information.

### The retina

The retina contains photoreceptive cells called rods and cones and
several types of neurons. The rods are generally sensitive to light and
there are three varieties of cones sensitive to long, medium and short
wavelengths of light (L, M and S type cones). Some of the ganglion cells
in the retina (about 2%) are also slightly light sensitive and provide
input for the control of circadian rhythms. A schematic diagram of the
retina is shown below.

![](Constudretina.png "Constudretina.png")

The photoreceptors hyperpolarise (their membrane potential becomes more
negative) in response to illumination. Bipolar cells make direct contact
with the photoreceptors and come in two types, *on* and *off*. The
on-bipolar cells are also known as *invaginating* bipolars and the
off-bipolars as *flat bipolars*. On-bipolars depolarise when light falls
on the photoreceptors and off-bipolars hyperpolarise. Action potentials
do not occur in the bipolar or photoreceptor cells.

The retinal neurons perform considerable preprocessing before submitting
information to the brain. The network of horizontal and ganglion neurons
act to produce an output of action potentials that is sensitive to
boundaries between areas of differing illumination (edge detection) and
to motion.

Kuffler in 1953 discovered that many retinal ganglion cells are
responsive to differences in illumination on the retina. This
**centre-surround** processing is shown in the illustration below.

![](Constudcentresurr.png "Constudcentresurr.png")

The centre-surround effect is due to **lateral inhibition** by
horizontally arranged cells in the retina.

The structure of the response fields of ganglion cells is important in
everyday processing and increases the definition of boundaries in the
visual field. Sometimes it gives rise to effects that are not directly
related to the physical content of the visual field. The most famous of
these effects is the Hermann Illusion. The Hermann Grid Illusion is a
set of black squares separated by white lines. Where the white lines
cross it appears as if there are grey dots.

![](Hermann_grid.jpg "Hermann_grid.jpg")

The grey dots are due to the relative suppression of on-centre ganglion
cells where the white lines cross. This is explained in the illustration
below.

![](Constudillfld.png "Constudillfld.png")

Notice how the grey dots disappear when the crossed white lines are at
the centre of the visual field. This is due to way that ganglion cell
fields are much smaller in the fovea.

There are many other retinal illusions. White\'s illusion is
particularly strong and was believed to be due to centre-surround
activity but is now thought to have a complex origin.

![](constudwhites.gif "constudwhites.gif")

The grey lines really are the same shade of grey in the illustration.
Mach\'s Illusion is another example of a centre-surround effect.
Centre-surround effects can also occur with colour fields, red/green and
yellow/blue contrasts having a similar effect to light/dark contrasts.

Lateral inhibition and the resultant centre-surround effect increases
the number of cells that respond to boundaries and edges in the visual
field. If it did not occur then small boundaries might be missed
entirely if these fell on areas of the retina outside of the fovea. The
result of this effect is everywhere in our normal visual phenomenal
experience so not only is visual experience a mapping of 3D on to a 2D
surface, it also contains shading and brightening at edges that will not
be found by photometers that measure objective light intensities.

Photoreceptors become less responsive after continuous exposure to
bright light. This gives rise to **afterimages**. Afterimages are
usually of the opponent colour (white light gives a dark afterimage,
yellow light gives a blue afterimage, red gives a green afterimage
etc.). Afterimages when the eyes are open are generally due to a lack of
response to a particular frequency of light within the white light that
bathes the retina.

It is clear that visual phenomenal experience is related more directly
to the layout and type of activity in the retinal cells than to things
in the visual field beyond the eye.

### Visual pathways

![](Constudeyepath.png "Constudeyepath.png")

### The lateral geniculate nucleus

Retinal ganglion cells project to the Lateral Geniculate Nuclei which
are small bumps on the back of the thalamus. (Only 10-15% of the input
to the LGN comes from the retina, most (c.80%) comes from the visual
cortex). The neurons in the LGN are arranged retinotopically so preserve
the layout of events on the surface of the retina.

The LGN are arranged in 6 layers. The top two are known as Magnocellular
layers (about 100,000 neurons with large cell bodies) and the bottom
four are called Parvocellular layers (about 1,000,000 neurons with small
cell bodies). Between the main layers are the Koniocellular layers that
consist of large numbers of tiny neurons.

The left Lateral Geniculate Nucleus receives input from the right visual
field and the right LGN receives input from the left visual field. Each
nucleus receives input from both eyes but this input is segregated so
that input from the eye on the same side goes to layers 1, 3, 5 and from
the other side to layers 2,4, 6.

The magnocellular layers contain neurons that have a large receptive
field, are sensitive to contrast, a transient response and are not
colour sensitive. The parvocellular layers contains neurons that have
small receptive fields, are colour sensitive, have a prolonged response
and are less sensitive to contrast.

The LGN pathway from the retina is largely connected to the striate part
of the visual cortex (cortical area V1) via a set of fibres called the
optic radiation. There are reciprocal connections between the Thalamic
Reticular Nucleus and the LGN. The LGN are also interconnected with the
Superior Colliculus and brainstem.

The LGN may be involved in controlling which areas of the visual field
are subjected to attention (O\'Connor *et al.* 2002).

### The visual cortex

The input from the LGN goes mainly to area V1 of the cortex. The cortex
is arranged in six layers and divided up into **columns**. Each column
in the visual cortex corresponds to a particular area of the retina in
one eye. The columns are arranged in rows called **hypercolumns**. Each
column within a hypercolumn responds to a different orientation of an
optical stimulus at a given location (so responds to edges/boundaries
that are oriented in the visual field). Hypercolumns from each eye are
arranged alternately and form a small block of cortex called a
**pinwheel**. At the centre of each pinwheel are colour sensitive cells
that are usually not orientation sensitive. These coincide with the
\"blobs\" that are seen when visual cortex is viewed using cytochrome
oxidase dependent stains. It is important to note that the
\"hypercolumns\" merge into one another and respond to line stimuli that
cover an area of retina so they may be physiological rather than
anatomical entities.

The blind spot in each eye is represented by an area of visual cortex
that only receives monocular input from the other eye (Tong & Engel
2001). The effect of the blind spot is illustrated below:

![](constudblindspot.gif "constudblindspot.gif")

Normally it seems that the blindspot is \'filled in\' with background
when one eye is used. However, Lou & Chen (2003) demonstrated that
subjects could respond to quite complex figures in the blind spot,
although how far they were investigating \'blindsight\' rather than
visual experience in the blind spot is difficult to determine.

Different layers in the visual cortex have outputs that go to different
locations. Layer 6 sends nerve fibres to the Lateral Geniculate Nuclei
and thalamus, layer 5 to superior colliculus and pons, layer 2 & 3 to
other cortical areas.

There are two important outputs to other cortical areas, the **ventral
stream** and the **dorsal stream**. The ventral stream processes colour,
form and objects. It proceeds to the inferior (lower) temporal cortex.
The dorsal stream processes motion, position and spatial relationships.
It proceeds towards the parietal cortex. Lesions in the ventral stream
can result in patients knowing where an object is located but being
unable to enumerate its properties, on the other hand, lesions to the
dorsal stream can result in patients being able to label an object but
unable to tell exactly where it is located.

There is also a large output from the visual cortex back to the
thalamus, this output contains more fibres than the thalamo-cortical
input.

### Depth perception

The world is three dimensional but the image on the back of the retinas
is two dimensional. How does the brain give the subject a perception of
depth?

Depth perception relies on **cues** which are data about the
displacement of things relative to the body. These cues consist of:

-   the convergence of the eyes
-   the accommodation of the lens
-   binocular disparity -the difference between the images on the
    retinas- this was first suggested by Wheatstone.
-   motion parallax - distant objects move slower when the observer
    moves - first suggested by Helmholtz.
-   optical flow - the rate of expansion/contraction of a scene with
    movement towards or away from it (Lee & Aronson 1974).
-   binocular occlusion - parts of a scene are invisible to each eye.
-   body motion provides cues about near objects.
-   vanishing points - the convergence of parallel lines.
-   numerous other cues such as size constancy, texture etc.

Binocular disparity has been most extensively studied as a source of
depth cues. When the eyes converge to focus on an object in from of them
there is very little disparity in the images of that object on the two
retinas. The angle at the object formed between the lines that project
back to the pupils is known as the **vergence** at the object. The
sphere where all objects have the same vergence is known as the
**horopter**.

![](Constudverge.png "Constudverge.png")

When the disparity between the retinas is small a single image occurs in
phenomenal experience which is accompanied by a sensation of objects
with depth. This is known as **stereopsis**. If the disparity between
the retinas is large double vision ensues, this is known as
**diplopia**. The curious feature of stereopsis is that we can see no
more of the object than is visible on the retinas and certainly cannot
see behind the object. Stereopsis is more like a stretching of 2D space
than actual 3D.

The **empirical horopter** is a zone where things are seen without
diplopia. The empirical and **Veith Muller** (geometric) horopters are
different. This difference is the result of both processing by the CNS
and optical factors.

**Physiological diplopia** refers to the stimulation of receptors in
different parts of the retinas of the two eyes by the same object.
Physiological diplopia does not always give rise to subjective diplopia,
objects close to the empirical horopter do not give rise to double
vision and the zone in which this occurs is known as **Panum\'s Fusion
Area**. It is widest for objects that are distributed away from the nose
(with \'temporal\' locations) and for objects that are slow moving and
poorly focussed.

In the review by Cutting and Vishton (1995) the contributions of each
type of cue is discussed. Cutting and Vishton also present evidence that
there are several zones of depth perception that are informed by
different sets of cues. These are **personal space**, which is the zone
of things within arms reach, **action space**, which is the zone in
which we interact and where our motions have a large impact on the
perceived layout, and **vista space** which is the zone beyond about 30m
that is informed by long range cues.

The interesting feature of 3D perceptual space is that it is not seen.
The sides of a solid object appear as intrusions or lateral extensions
in 2D space, when we close an eye that has access to the side of the
object and then open it again the side grows out into 2D space. The lack
of \'seeing\' depth is also evident when we close one eye when looking
at a vista - nothing seems to change even though stereopsis has gone.
This leaves the problem of what it is that constitutes the \'feeling\'
of depth. We have feelings that we can fall into space or move into it
or around in it. Depth seems to be defined by premotor modelling and the
potential for occupancy by our bodies and limbs. As such it involves
qualia that are different from those of vision and more akin to those
that accompany movement, as an example, if you reach out to touch
something, move the hand back, then consider the distance to the object
it is evident that a feeling of the movement is still present. Is depth
a quale of movement modelled during the extended present of perception?

-   Cutting, J.E. & Vishton, P.M. (1995) Perceiving layout and knowing
    distances: The integration, relative potency, and contextual use of
    different information about depth. In W. Epstein & S. Rogers (eds.)
    Handbook of perception and cognition, Vol 5; Perception of space and
    motion. (pp. 69-117). San Diego, CA: Academic Press.
    <http://pmvish.people.wm.edu/cutting&vishton1995.pdf>

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-   Gregory, R.L. (1997). Knowledge in perception and illusion. From:
    Phil. Trans. R. Soc. Lond. B (1997) 352, 1121--1128.
    <http://www.richardgregory.org/papers/knowl_illusion/knowledge-in-perception.pdf>

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-   Hubel, D.H. (1981). EVOLUTION OF IDEAS ON THE PRIMARY VISUAL CORTEX,
    1955-1978: A BIASED HISTORICAL ACCOUNT.
    <http://nobelprize.org/medicine/laureates/1981/hubel-lecture.pdf>

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-   Lou, L. & Chen, J. (2003). Attention and Blind-Spot Phenomenology.
    PSYCHE, 9(02), January 2003.
    <http://psyche.cs.monash.edu.au/v9/psyche-9-02-lou.html>

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-   O\'Connor, D.H., Fukui, M.M., Pinsk, M.A. & Kastner, S. (2002).
    Attention modulates responses in the human lateral geniculate
    nucleus. Nature Neuroscience 5, 1203 - 1209 (2002).
    <http://www.nature.com/neuro/journal/v5/n11/full/nn957.html#B34>

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-   Tong, F., & Engel, S. A. (2001). Interocular rivalry revealed in the
    human cortical blind-spot representation. Nature, 411, 195-199.
    <http://www.psy.vanderbilt.edu/tonglab/publications/Tong&Engel2001.pdf>

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-   Tong, F. (2003). PRIMARY VISUAL CORTEX AND VISUAL AWARENESS. Nature
    Reviews of Neuroscience. 4:219-229.
    <http://www.psy.vanderbilt.edu/tonglab/publications/Tong_NRN2003.pdf>

**Modules**

-   The cortex and
    thalamus
-   Rivalries and
    synchronisation