Eye neurobiology Flashcards

(60 cards)

1
Q

Describe the three layers of the eye

A

The fibrous outer layer comprises the
cornea (anterior) and sclera (posteriorly).
It is continuous with the dura mater.
❚ The middle choroidal layer is vascular
posteriorly and forms the iris and ciliary
body anteriorly. It is continuous with the
arachnoid and pia.
❚ The inner neural layer is the retina. It is
continuous with CNS tissue

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2
Q

Describe the retina

A

The innermost layer is called the retina.
The optic nerve exits the retina at a pale
circular region termed the optic disc.
❚ There are no photoreceptors in the optic
disc, hence it is called the blind spot
❚ The macula is a circular region adjacent
the optic disc and is responsible for
central (as opposed to peripheral
vision). The centre of the macula is a
depression termed the fovea

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3
Q

How does the eye get its nourishment

A

The lens and cornea are avascular. The cornea
and aqueous humour are oxygenated directly
from the atmosphere.

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4
Q

What is the refractive index in relation to the eye

A

The Refractive Index (Ri) is a measure of how much the speed of
light is reduced traveling through a given medium relative to a
vacuum
❚ The Ri of the lens of the eye is 1.45. Hence the speed of light
traveling through the lens = 1/1.45 = 0.7x the speed of light in
a vacuum

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5
Q

Where does all the resolving power of the eye lie

A

The principal axis of the eye falls on the fovea which is a 1 mm
diameter area where all the resolving power of the retina lies.

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6
Q

How does lens accomodation work

A

In order to focus near objects on the retina the
refractive power of the eye must be increased by
increasing the curvature of the lens. This is called
accommodation

When an object is more than ~6 meters away the
rays coming from every part of it will be parallel
and an image will form at the focal plane (retina).

❚ The nearer an object is to the eye the more rays
of light diverge from different parts of it and will
be brought to a focus behind the retina.

accommodation takes place in the ciliary body which consists of the ciliary muscle and ciliary process
When relaxed, the ciliary body exerts tension
on the lens capsule tending to flatten the lens.
❚ When the ciliary muscle contracts it relaxes the
zonular fibres. This makes the lens expand

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7
Q

How does regulation of light take place

A

❚ Regulation of light is controlled mechanically by
the iris although the CNS also automatically
compensates for changes in light in intensity
❚ The iris aperture (pupil) is controlled by two
muscles:
❚ The pupillary sphincter is the stronger of the
two and constricts the pupil (miosis).
❚ The pupillary dilator muscle fibres are radially
orientated and widens the pupil (mydriasis).
❚ Innervation of the sphincter is via
parasympathetic NS, innervation of the dilator is
via the sympathetic NS

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8
Q

Describe the visual field

A

The visual field is the view
seen by the two eyes without
movement of the head
❚ The visual field is divided into
both binocular and monocular
zones

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9
Q

What is the binocular zone

A

Light from the binocular zone strikes the
retina in both eyes.
❚ Therefore central part of both visual
hemifields is viewed by both retinas

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10
Q

What are the monocular zones

A

❚ In each half of the visual field there
is also a monocular zone where
light from the temporal portion of
each visual hemifield projects only
onto the nasal hemiretina of its
corresponding eye (the ipsilateral
nasal hemiretina).

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11
Q

What are the hemiretinas

A

❚ The surface of the retina is divided
with respect to the midline (the
principal axis intersecting the fovea).
❚ The nasal hemiretina lies medial to the
fovea and contains the blind spot. The
temporal hemiretina lies lateral to the
fovea.

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12
Q

What is the significance of the optic chiasm

A

❚ The optic nerve of each eye contains the
axons of retinal ganglion cells from both
hemiretinas.
❚ A partial DECUSSATION takes place in
the optic chiasm.
❚ At the optic chiasm, fibres from the nasal
half of each retina cross.
❚ This means that each optic tract exiting
from the chiasm carries half of the visual
field to its contralateral hemisphere.

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13
Q

What is the difference between the optic tract and the optic nerve

A

Because of the optic chiasm, each optic nerve carries all of
the visual information from its corresponding eye, whereas
each optic tract carries a complete representation of one
half of the visual field.
❚ Fibres from the nasal half of each retina cross to the
opposite side of the chiasm, whereas fibres from the
temporal half do not cross
❚ Fibres in the the optic tracts project to subcortical
regions of the brain, principally to the lateral geniculate
nucleus of the thalamus

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14
Q

How light passed through the retina

A

❚ Light entering the cornea is projected onto the back of the eye where it is
converted into electrical signals by photoreceptors in the retina
The laminar organization of the retina is perhaps surprising - seemingly
inside-out
❚ Light must pass through the vitreous humour, through the ganglion cells
and bipolar cells before it reaches the photoreceptors.

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15
Q

What is the role of the pigment epithelium

A

One advantage of this structure is that the pigment epithelium lies below the
photoreceptors. These cells absorb light passing through the retina and prevent
reflection of light within the eye

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16
Q

What is the macula

A

Pigmented area ~5.5 mm
diameter. Yellow
pigment helps by
absorbing blue and
ultraviolet light

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17
Q

What is the fovea

A

Pit shaped area ~1.5 mm
diameter located near the
center of the macula. Cone
photoreceptors only (no
rods)

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18
Q

What are the different retinal cell types

A

The rod and cone photoreceptors are involved in phototransduction
3 types of interneurons connect
photoreceptors to the retinal ganglion cells
– bipolar (B) cells, horizontal (H) cells and
amacrine (A) cells.
These are involved in pre-processing which enable the receptive fields of ganglion cells
and bipolar cells to respond with precision
to different spatial and temporal patterns of
light

Transmission of takes place through M type (magnocellular LGN) and P type (Parvocellular LGN) retinal ganglion cells. M type cells have larger receptive fields than P type cells

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19
Q

Describe the layered structure of the retina

A

PIGMENT CELL LAYER (RPE)
❚ OUTER SEGMENTS (OS) OF
PHOTORECEPTORS - contains light-
sensitive elements of retina
❚ OUTER NUCLEAR (ON) LAYER - cell
bodies of inner segments of
photoreceptors
❚ OUTER PLEXIFORM (OP)- synapses
of bipolar cells with photoreceptors
❚ INNER NUCLEAR (IN) - cell bodies of
bipolar cells
❚ INNER PLEXIFORM (IP) LAYER -
synapses of bipolar cells with
ganglion cells
❚ GANGLION CELL (GC) LAYER
-
Output cells
❚ NERVE FIBERS (NF)

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20
Q

Describe differences between rods and cones

A

Rods mediate night vision while cones mediate day vision

Rods are achromatic while cones are chromatic

Cones perform better than rods at
all tasks except the detection of
dim stimuli: they have a higher
visual acuity, better temporal
resolution (they respond better to
rapid changes in the visual image)
and they mediate colour vision

Rods have a lower threshold to stimulation compared to cones

All cones can respond to light of any
wavelength
❚ But each type of cone requires fewer
photons of their characteristic
wavelength to respond

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21
Q

Describe the organization of the outer layer in photoreceptors

A

The outer segments of both
types of photoreceptor are filled
with light absorbing pigment in
an elaborate array of stacked
membranous discs, this is where rhodopsin is contained.
❚ This organization dramatically
increases the surface area of
the membrane in this cellular
compartment.
Thus, providing space for large
numbers of photopigments
anchored in these membranes

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22
Q

Describe rhodopsin and hence the first stage of phototransduction

A
  • The visual pigment in Rod cells is called Rhodopsin.
  • Rhodopsin is large complex comprised of two covalently bound compounds
  • a 348 amino acid containing transmembrane protein called Opsin (MW = 40 kDa).
  • and a small light absorbing hydrocarbon - retinal.
  • Opsin is anchored in the membrane in the outer segment stacks via 7 transmembrane domains
  • In its non-activated form Retinal is attached to the side chain of lysine 296, which is found in the
    seventh membrane spanning region of Opsin.

Absorption of light by 11-cis retinal causes a
rotation around the 11-cis double bond, forming
the 11-trans isomer. This reaction is the only light
dependent step in the visual pathway.
❚ The 11-trans conformation can no longer fit into
the binding site in opsin, and this promotes a
conformational change in opsin into the semi-
stable metarhodopsin-II conformation

Metarhodopsin-II is unstable and splits within minutes into opsin
and all-trans retinal.
❚ The all-trans retinal is transported to pigment epithelial cells
where it is reduced to all-trans retinol (a.k.a. Vitamin A). This is
the precursor of 11-cis retinal, which is then transported back to
the rods

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23
Q

How is retinal recycled

A

Metarhodopsin-II is unstable and splits within minutes into opsin
and all-trans retinal.
❚ The all-trans retinal is transported to pigment epithelial cells
where it is reduced to all-trans retinol (a.k.a. Vitamin A). This is
the precursor of 11-cis retinal, which is then transported back to
the rods

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24
Q

What is the second step of phototransduction

A

Activation of pigment molecules by light leads to stimulation of 100s of G proteins (called transducin in rods). This in turn activates a cGMP phosphodiesterase,
which catalyzes the breakdown of 1000s of cGMP into 5’GMP.

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25
What is the third step of phototransduction
Channels that depolarize photoreceptors are cGMP gated, in the dark state when no light is coming into the cell, this channel is constantly open leading to depolarization Since cGMP is hydrolyzed to 5'GMP when light comes in, this closes the ion channel and causes hyperpolarization of the cell and reduced glutamate release (an excitatory neurotransmitter)
26
What is the important role of Ca2+ in phototransduction
* Intracellular calcium is an important modulator in promoting the return of the photoreceptor to the dark state * Ca2+ controls at least 3 components in the transduction cascade * Low levels phosphorylation of Rhodopsin * Guanylate cyclase is accelerated to generate new cGMP * Increased affinity of cGMP to the Na2+ channels
27
How do photoreceptors respond to light
Rods and cones do not produce action potentials ❚ They respond to light with graded fluctuations of membrane potentials ❚ Rods respond slowly so that the effects of all photons absorbed over a ~100 msec period are summed together.
28
What is the difference between rods and cones in terms of flickers
This means that although rods can detect small amounts of light, they cannot resolve flickers faster than 12Hz ❚ Cones respond much faster and can resolve flickers up to at least 55Hz ❚ Generally speaking, we cannot separate flashes of light faster than about 16Hz the CRITICAL FUSION FREQUENCY. Conventional movies (24Hz), analogue TV (50-60Hz) and video monitors (>60Hz) or HDTV devices have high frame and/or refresh rates to avoid flickering
29
Which are the only cells in the retina that transmit AP's
In the retina, only the ganglion cells (G) transmit information as a series of action potentials
30
What is the function of the interneurons in the retina
Three specialized interneuron cell types connect the photoreceptors to the ganglion cells: bipolar cells (B), horizontal cells (H) and amacrine cells (A). These cells also use graded potentials. ❚ These interneurons do not simply transmit signals from the photoreceptor to the ganglion cells – they process signals from multiple photoreceptors (receptive fields) in response to different patterns of light stimulating the retina
31
What is the difference between ON and OFF bipolar neurons
ON-centre and OFF-centre bipolar cells establish parallel pathways for the signal from a single cone ❚ When the cone cell is activated by light it becomes hyperpolarized leading to a reduction of Glutamate release ❚ This causes the ON-centre bipolar cell to become excited (depolarized) and the OFF-centre bipolar cell to be inhibited (hyperpolarized). ❚ OFF centre RGC are hyperpolarized by OFF-centre bipolar neuron, whereas ON-centre RGC show a very typical neuronal response to depolarisation by firing APs.
32
Why do the different bipolar cells respond differently to Glu
The two different classes of bipolar cells respond differently to Glu because they have different post- synaptic receptors that gate different ion channels. The on bipolar cell is gated by metabotropic glutamate receptors while the off bipolar is gated by ionotropic receptors
33
What are the regional differences in the retina
In the fovea * Ganglion cells (both M and P types are found) have smaller receptive fields * Low convergence: Ratio of photoreceptors to ganglion cells is <10:1 * Cone photoreceptors only Peripheral retina * Higher ratio of rods to cones * High convergence: Higher ratio of photoreceptors to ganglion cells (>100:1) * More sensitive to light
34
What is the lateral pathway
Signals from cones in the periphery of the receptive field reach bipolars by way of horizontal cells, which are inhibitory. This creates the opposite effect to illumination at the receptive field centre.
35
Describe the indirect pathway
Signals from the surround are mediated via horizontal cells ❚ Light falling on a photoreceptor in the surround of an ON-centre bipolar cell causes a hyperpolarization. ❚ This causes the horizontal cell to hyperpolarize. This reduces the amount of inhibitory neurotransmitter (GABA)* released by the horizontal cell onto postsynaptic cones in the receptive field centre. ❚ Thus, when the surround is illuminated the horizontal cells depolarize the photoreceptors in the field centre, causing the opposite effect to light falling directly on them (hyperpolarization). ❚ This causes an ON-centre bipolar cell to hyperpolarize When the surround area of the receptive field is illuminated, the horizontal cell becomes hyperpolarized by the surrounding photoreceptors, this leads to the horizontal cell inhibiting the central 'ON' bipolar cell
36
How do ON and OFF centre bipolar neurons respond to light falling on their receptive fields
ON-centre cells depolarize in response to light falling on the centre of the field but hyperpolarize in response to light falling on the surround. ❚ OFF-centre cells hyperpolarize in response to light falling on the centre of the field but depolarize in response to light falling on the surround.
37
How is edge detection observed
* An edge of a light stimulus moves into the receptive field surround of ON bipolar cell B. * This edge is also falling on the receptive field centre of ON bipolar cell C. * The light will cause bipolar cell C to depolarize because of the direct synapse with the photoreceptor. * The light will also cause bipolar cell B to hyperpolarize because of the indirect synapses through the horizontal cell. * The larger membrane potential difference between the cells B and C compared to without lateral inhibition. This leads to an enhancement in the perception between the dark and light side of the edge. * In parallel, this effect is enhanced through opposite modulation of OFF centre cell activity Works through lateral inhibiton of bipolar cell B Edge detection is pretty common sense, if the light falls on the edge of a receptive field, the photoreceptors nearest that edge of light will depolarize and the surrounding neurons will be inhibited
38
What do receptive fields allow for
The structure of ganglion receptive fields allows ganglion cells to respond best to changes in light intensities (contrast) between the centre and surround of the receptive field
39
How does resolution work in the retina
❚ Resolution of visual image is determined by density of sensory receptors and size of receptive fields in the retina. ❚ As the density of receptors increases and the size of the receptive field decreases spatial detail increases. ❚ Rods contribute to larger receptive fields and are relatively much more numerous than cones toward the periphery
40
What is contrast
Contrast is the difference in visual properties (brightness, colour) that makes an object distinguishable from other objects and the background.
41
What is the biological basis of contrast perception in retina
Contrast detection results from the structure of ganglion cell receptive fields, which allow ganglion cells to respond best to changes in light intensities (contrast) between the centre and surround of the receptive field.
42
Explain spatiotemporal contrast sensitivity
A moving object elicits stronger firing rate in the ganglion cell population near the edges of the objects image These are the only regions of spatial contrast and only regions where the light intensity changes over time The centre-surround antagonism is the basis for our ability to processes differences in illumination that falls onto photosensitive cells of the retina and are critical for spatiotemporal contrast sensitivity in addition to visual acuity
43
What is the difference between colour and contrast
Colour is not essential for perception of detail. The majority of our ability to perceive detail results from our ability to discriminate variations in brightness within the visual image.
44
What is the difference between reflection and adsorption of light in terms of colour vision?
Note that the light falling on the apple is broad spectrum (white light). ❚ The red pigment in the apple appears red because the apple adsorbs other light frequencies but reflects predominately red light. Similarly the banana adsorbs most light frequencies but reflects predominantly yellow light.
45
What is the trichromacy theory?
According to the Young-Helmholtz trichromacy theory, originally proposed in the 19th century, the brain assigns colours based on an additive colour model, in which the three primary colours of red (R), green (G) and blue (B) light are added together to produce a broad array of different colours. . Today, we know the human eye contains 3 types of cones, which preferentially absorb blue (S), green (M) and red (L) light. This theory explains how each type of cone photoreceptor detects different wavelengths in light
46
What are the different spectral photosensitivies of the cone photoreceptors
❚ Each class of cones contains a different photopigment that gives it a distinct spectral sensitivity. ❚ S-cones are most susceptible to blue light (~420 nM peak), M-cones to green (~530 nM peak) and L-cones to red (~560 nM peak).
47
How is colour percieved from surface reflection
Light reflected from real surfaces in the natural world rarely contains a single wavelength. In fact most surfaces reflect light that represents a continuous distribution of wavelengths. ❚ A system that uses more than two types of cones would be able to differentiate more physically different surfaces that appear to be the similar colour.
48
Describe how overlapping spectral sensitivities generate transmit colour vision
Because the reflectance functions of the flowers shown here vary slowly across the spectrum, and the spectral sensitivities of the L and M cones are so similar across a broad spectral region, these two classes of cones will generate highly correlated signals when they absorb light from natural surfaces such as these ❚ This would create a considerable redundancy of information entering the visual pathway if each cone was connected independently to the visual pathway ❚ The solution is to transmit DIFFERENCES between signals, not the signals themselves.
49
Describe the colour opponent theory
Ewald Hering (1892) noticed that the hues red, yellow, green and blue have special properties. ❚ They are fundamental because all other hues (spectral colours) can be described as mixtures of them, and yet they seem to be related as mutually exclusive pairs. ❚ These led Hering to propose that vision depended on three opponent mechanisms ❚ All of the colours of the spectrum could be derived in terms of variation in two opposing colour pairs: red-green and blue-yellow. He proposed that a third mechanism would capture the light-dark variation
50
How did this theory be used to accurately model natural surface reflections
The Hering theory can be used to accurately model natural surface reflections ❚ The natural surface reflection curve can be realistically modeled by combining three separate component functions: one comprising the brightness of the image, one comprising the red- green variation and the other yellow-blue variation. ❚ These three component functions can account for >99% of the variance in surface reflective functions of natural objects.
51
Describe second stage transformation in relation to the colour opponent theory
Hering’s theory basically describes a way to transform the visual information provided by the three cone types in the Young-Helmholtz theory into a more compact form. This is called “second-stage transformation”. ❚ He proposed that the ideal transformation is one that yields, for the kinds of visual stimuli the eye normally receives, the smallest correlation among signals in the three pathways. These are: ❚ The sum of all three cone signals: ❙ L + M + S ❚ The difference between the L and M cones: ❙ L – M ❚ The difference between the S cones and some combination of L and M cones: ❙ S - LM
52
What are the different pairs in colour blindness
❚ Dichromats have only two classes of receptors, instead of three. ❚ Two kinds of dichromacy are most common: loss of either L cones or M cones (protanopia or deuteranopia), with each having about 1% prevalence. ❚ Since either will principally affect the L-M channel, they are collectively known as “red-green colour-blindness”
53
What is the 2/3 stage model of colour vision
1st stage - light is transduced by three types of cone photoreceptors, L (long), M (middle), and S (short) -wavelength sensitive. 2nd stage - cone signals are integrated through four cone-opponent mechanisms (L+M−, M+L−, S+(L+M)−, and (L+M)+S−) instantiated by retinal circuits connecting specific cone types to retinal ganglion cell types. Hypothesized 3rd stage - cone-opponent signals are predicted to combine in four specific combinations to generate colour-opponent mechanisms, as well as neurons whose activities underlie the perception of four “unique colours”
54
Describe the function of the 2 P cells for colour vision
P cells are more numerous and have small receptive fields. They respond selectively to specific wavelengths and are believed to be involved in the perception of form and colour. ❚ P cells fall into two subtypes: ❙ 1. Neurons that receive opposed signals from L and M cones. ❙ 2. Neurons that receive signals from S cones opposed to some combined signal from L and M cones. ❚ Therefore these two subtypes of P cells appear to have the necessary characteristics to provide the red-green and yellow-blue channels postulated by Hering
55
What is used to compute the brightness signal
Red-green subtype of P-cells receives inputs from only L and M cones, yellow-blue subtype receives inputs from all three classes L, S and M. ❚ S cones are much less frequent (<10%). And because it is much harder to focus short wavelength light, they are largely absent from the centre of the fovea. ❚ Therefore it seems that only the inputs from L and M cones are significantly used to compute the brightness signal.
56
How are these different types of signals (colour vs brightness) transmitted?
Both types of information must be conveyed through the discharge of action potentials by the P cells and ultimately this must be resolved by the visual cortex.
57
How are P cells suited for colour detection
P-cells respond well to brightness variations in the fine structure of the image, but also respond well to colour variation in the coarse structure of the image The spatial organization of the P-cell’s receptive field allows the cell to convey BOTH brightness and colour information. When light falling on a P cell’s receptive field covers both centre and surround, the P-cell will respond well to variations in colour between centre and surround – being excited by some hues and inhibited by others
58
What is the organization of P cells in relation to the LGN
The four ventral layers are known as parvocellular layers and receive inputs mainly from P cells ❚ Both the magnocellular and parvocellular layers contain on-centre and off-centre cells, just as in the retina ❚ P-cells in the lateral geniculate nucleus are also tightly clustered into ‘red-green” and “yellow- blue” groups
59
How can the functions of M and P cells be studied
❚ The functions of the M and P pathways can be investigated using selective lesioning techniques because the two pathways are separated into different layers of the lateral geniculate nucleus. ❚ In this case the lesions were induced using an excitotoxin, ibotenic acid.
60