Describe some of the diverse types of selectivity shown by

visual receptive fields in the retina and in the visual cortex

and offer some explanation for the existence of this diversity.

The complex network of nervous pathways, which corroborate to give us

our idea of vision, relies upon several separate factors, each one being responsible
for a different parameter of sight. Vision takes into account the form, movement,
and colour of objects and the brain addresses these individual factors in different
locations. The parvocellular-interblob system concerns the form, the
magnocellular the motion, and the parvocellular- blob the colour. However
ultimately, analysis from these independent locations will combine to establish
our interpretation of a given visual field, this being defined as the view seen by
two eyes without movement of the head.
The retina and visual field are both divided into four quadrants. If one is to
imagine a vertical line through the middle of the visual field, one can establish
that there is a hemifield nearer the nose, the nasal hemiretina, and the other
hemifield, the temporal hemiretina. These two quadrants can then further be
separated into the ventral and dorsal quadrants. In terms of the visual field, there
is the imaginary left half, from which a point would project into the neurons of the
nasal hemiretina of the left eye and the temporal hemiretina of the right eye. The
same applies for a point in the right hemifield. Thus one can see that there is a
region in the middle of the visual field, where a given point would project into
both eyes and this area is known as the binocular zone. Similarly there regions
neighbouring this zone peripherally are termed monocular zones or the temporal
crescents. Here the angle of a point relative to the eyes will give rise to light
entering only one of the eyes as the position of the nose prevents projection of
light into reaching the eye on the opposite side. The optic disc, the location from
where the retinal ganglion output cells project, is insensitive to light due to the
absence of the photoreceptors. However this is located nearer the midline than the
fovea and hence light from a specified point will never fall simultaneously on
both optic discs thus allowing us to be unaware of our so-called blind spot. This
can easily be exposed by covering one eye and focusing the other on a point.
Another point in the temporal crescent will disappear when at the right distance.
Measurement of visual field function can be done through a visual field
examination. This procedure comprises several separate tests, each of which
concerns a different aspect of vision. A confrontational visual field exam is a
simple test whereby a clinician will cover one eye of the patient and then wave
their hand in front of the person to attain a sense of their visual field boundaries.
The Goldmann field exam allows mapping of a patient's visual field. The
individual is placed three feet from a screen with a target near its centre. With one
eye covered and the other eye focussed on the target, the examiner will slowly
move an object towards the patient's visual field, measuring when
acknowledgement is obtained. An automated perimetry exam involves a machine
which will shine light into a patient's retina while they focus on a single point.
Thus the computer can map a patient's visual field.
Visual field examinations are extremely relevant in patients who have
suffered conditions such as optic nerve disorders, scotomas or more generally any
optical difficulty. Inflammation of the optic nerve, optic neuritis, can lead to a
partial or complete loss of vision. The reduced capacity of the optic nerve in
carrying nerve impulses from the retina to the visual cortex results in a lack of
sight and the extent of this can be measured using these visual field exams. Severe
loss of vision can also occur from retinal arterial emboli. The sudden depletion of
blood supply causes significant levels of retinal cell death and thus reduced or
complete loss of the visual field.
Despite the abundance of photoreceptors throughout the majority of the
retinal outer nuclear layer, the output from the retina occurs at a region known as
the optic disc which is completely devoid of all cells bar the output ganglion
neurons hence the existence of the blind spot. These ganglion cells with their
circular centre-surround antagonistic receptive fields can come in two forms –
either on-centre, off surround where light projecting onto the centre excites the
neuron but light falling on the receptive field surround inhibits the cell or off-
centre, on surround which adopts the opposite format. As these ganglion cells
travel further towards the visual cortex, the two optic discs converge to form the
optic tract after which the nasal hemifields cross over at the optic chiasm and thus
the opposite hemifield is concerned with a given half of the brain i.e. The right
nasal hemifield will be analysed by the left half of the visual cortex. Therefore one
can see why compression of the optic chiasm, by a pathology such as a pituitary
tumour, which would usually be located superior to the chiasm in the pituitary
fossa, would lead to loss of peripheral vision or bitemporial hemianopsia. The
crossing nasal hemiretina no longer are able to continue on to the higher centres of
the brain.
From the chiasm, the optic tract will project forward towards the lateral
genticulate nucleus (LGN). Here the ganglion cells synapse allowing the initial
nervous impulse to be processed by the brain. Anatomically the LGN is organised
into six numbered layers, the first ones comprising the ventral regions and the last
the dorsal regions. Ipsilateral layers from the retina are found in layers 2, 3 and 5
while contralateral layers will be found in layers 1, 4 and 6. Also, layers 1 and 2
are magnocellular layers while the remaining are parvocellular layers. Those cells
found in the first two layers, M cells are monochromatic with a low degree of
spatial resolution but have good temporal resolution and larger receptive fields
than the P cells. Cells found in the magnocellular regions are believed to be
responsible for processing of motion by the visual cortices. P cells concern colour
and have limited temporal resolution and smaller receptive fields but have better
spatial resolution.
Neurons from the LGN then collect to form the optic radiation, a group of
axons which will continue towards several higher centres of the brain, the
moajority of which will travel to the primary visual cortex or V1. Some fibres will
continue towards the pretectum, which concerns pupillary reflexes through the
oculomotor cranial nerve and others to the superior colliculus, which is associated
with the saccadic eye movements via the oculomotor, trochlear and abduces
cranial nerves. Some intial fibres will project forwards to cover the front of the
temporal horn resulting in a loop structure, known as Meyer's loop. Lesioning of
this loop has been seen to result in loss of vision in the superior quadrant or
quadrantopia.
Having reached V1, or the striate cortex, which is located on the medial
surface of both cerebral hemispheres in the occipital pole, the fibres arrange so
that the visual field is analysed by the brain contralaterally in retinotopic order.
The central visual field is represented at the most posterior end with the upper half
of the visual field associated with the inferior banks of the calacrine culcus and
the lower half with the superior banks of the sulcus. V1 is once again divided into
six layers with fibres projecting from the LGN only going into layer 4. Layer 4
can be further separated into layers 4A, 4B, 4Cα and 4Cß. Neurons from the
magnocellular layer of the LGN travel towards 4Cα and those from the
parvocellular layer travel to 4Cß.
Whereas the receptive fields in the retina were based arround the centre-
surround antagonistic form, those in the primary visual cortex comprise a different
structure. Here, the “on” and “off” bands run side by side thus making them more
selective for the specific orientation of visual features. Throughout all six layers,
V1 is arranged into achromatic cell columns, all having similar receptive fields.
The orientation format slightly differs between each layer in such a way that over
the whole course of the visual cortex, orientation selectivity gradually varies in a
smooth fashion. Columns found between the orientation columns respond to
particular wavelengths and these are known as blobs due to their so-called
appearance when viewed after staining with cytochrom oxidase enzyme. Depth of
a viewed object is also processed initially in V1. Retinal disparity, which the
cortex has a sensitivity towards, involves the mismatching of a given visual
feature's relative position of space.
The primary visual cortex does not complete the vision system. Higher
centres of the brain will specifically address the issues of colour, depth, movement
and orientation. Fibres from V1 will pass to the first extrastriate area, V2, and
eventually other visual areas of the brain. Most of these pathways will be divided
into dorsal and ventral streams, referring back the original arrangement of the
magnocellular and parvocellular neurons. The parvocellular pathway, or ventral
streams, involves areas V4 and the inferotemporal lobe and concerns primarily
colour and depth contours giving it its characteristic name, the “what” pathway.
The dorsal stream includes V5 and the medial superior temporal area (MST). Here
the cells have strong selectivities for motion direction and motion in-depth and
thus has been labelled the “where” pathway.
Studies on separate parallel pathways being associated with different aspects
of vision has provided the evidence for their existence. For example, localised
damage to the temporal cortex, which is considered to be the equivalent of V4,
results in a patient suffering from achromatopsia or loss of colour vision.
However, surprisingly they still enjoy a relatively accurate vision for form. The
design of the visual system in humans has adopted a form such that despite
damage being inflicted to a certain area of the pathway, the resulting consequence
will most likely still consist of vision to some extent. If the visual system relied
upon one pathway alone, then damage to this would cause total blindness every
time. The individual segments also have evolved so as best to benefit our
circumstances, whether the compromise between colour and acuity has increased
our accuracy in sight or the superior colliculus has coordinated some movement to
sudden stimuli in our periphery, perhaps originating from rapid predators. For this
reason, one can easily see that the optical centre is convoluted as are the array of
conditions, which can alter it.