Lectures 4 & 5: Vision Flashcards
Light enters the Eye and reaches the Retina
What is light?
- Light can be thought of as discrete particles of energy (photons) or as waves of energy
- Light is defined as waves of electromagnetic energy that are between 380 and 760 nanometres
- Light wavelength has role in perception of colour, intensity has role in perception of brightness
Light enters the Eye and reaches the Retina:
The pupil and lens
- Amount of light reaching retinas regulated by contractile tissue – the irises (give eyes their colour)
- Light enters eye though pupil (hole in iris)
- Adjustment of pupil size in response to changes in illumination represents compromise between sensitivity (ability to detect presence of dimly lit objects) and acuity (ability to see the details of objects)
- When illumination is high, visual system constricts pupils, image falling on each retina is sharper creating greater depth of focus
- When illumination is too low to adequately activate the receptors, pupils dilate to let in more light thereby sacrificing acuity and depth of focus
- Lens focuses incoming light on retina
Light enters the Eye and reaches the Retina:
Accommodation:
- Process of adjusting configuration of lenses to bring images into focus on retina
- Ciliary muscles adjust tension of ligaments holding lens in place, when gaze is directed lens assumes natural cylindrical shape, increases ability of lens to refract (bend) light and bring close objects into sharp focus
- Lens is flattened when focusing on distant object
Light enters the Eye and reaches the Retina:
Eye Position and Binocular Disparity:
- Eyes in vertebrates come in pairs (left and right), see in almost every direction without moving heads except humans have eyes on the front of their heads
- Human arrangement is important, what is in front can be viewed simultaneously allowing visual system to create 3D perceptions to see depth from 2D retinal images
- Movements of eyes are co-ordinated so each point in visual world is projected to corresponding points on both retinas, eyes must converge (turn slightly inward) to achieve this
- Convergence is greatest when you are inspecting things that are close
- Positions of images on retinas can never correspond exactly as eyes do not view world from exactly same position
- -> Binocular disparity is the difference in the position of the same image on the two retinas
- Is greater for close objects than distant therefore visual system can use degree of binocular disparity to construct one 3D perception from 2D retinal images
The Retina and Translation of Light into Neural Signals:
Light –> Pupil –> Lens –> Retina
- Retina converts light into neural signals, conducts them towards CNS, participates in processing of signal
- Retina composed of 5 layers of different types of neurons: Receptors, Horizontal Cells, Bipolar Cells, Amacrine Cells and Retinal Ganglion Cells
- Retinal neurons communicate both chemically via synapses and electrically via gap junctions
- Amacrine and horizontal cells are specialised for lateral communication (communication across the major channels of sensory input)
- Light reaches receptor layer only after passing through other 4 layers
- Once receptors activated, neural message transmitted back out through retinal layers to retinal ganglion cells
- Retinal ganglion cell axons project across inside of retina before gathering together in a bundle exiting eyeball
Inside out arrangement creates 2 visual problems:
• Incoming light is distorted by retinal tissue through which it must pass before reaching receptors, minimised by fovea – indentation 0/33cm at centre of retina specialised for high acuity vision (seeing fine details), thinning of retinal ganglion cell layer at fovea reduces distortion of incoming light
• For bundle to leave eye there must be a gap in receptor layer –> gap is Blind Spot, problem solved by completion (or filling in)
Light enters the Eye and reaches the Retina:
Completion (Filling in):
- Visual system uses information provided by receptors around blind spot to fill in gaps in retinal images
- When visual system detects straight bar going into one side of blind spot and another leaving the other side, visual system fills in missing bit so what is seen is a continuous bar
- Not only form of completion, when looking at object visual system extracts key info about its edges and location, conducts information to cortex, perception of entire object created from that partial information
- Surface Interpolation: The process by which we perceive surfaces; the visual system extracts information about edges and from it infers the appearance of large surfaces
Light enters the Eye and reaches the Retina:
Cone and Rod Vision – Receptors
Cones: Cone shaped receptors
Rods: Rod shaped receptors
- Species active only in day tend to have cone-only retinas and species only active at night ted to have rod only retinas
• Duplexity Theory:
Theory that cones and rods mediate different kinds of vision
- Phototopic vision (cone-mediated vision) predominates in goof lighting and provides high acuity (finely detailed) coloured perceptions of the world, in dim lighting there is not enough light to reliably excite the cones, more sensitive Scotopic vision (rod mediated vision) predominates, Scotopic vision however lacks both detail and colour of Phototopic vision
–> Differences in types of vision result from a difference in the way the two systems are wired, in the Scotopic system output of several hundred rods converge on a single retinal ganglion cell whereas in the Photopic system only a few cones converge on each retinal ganglion cell to receive input from only a few cones, the effects of dim light simultaneously stimulating many rods can summate (add) to influence the firing of the retinal ganglion cell onto which the output of the stimulated rods converge whereas the effects of the same dim light applied to a sheet of cones cannot summate to the same degree and retinal ganglion cells may not respond at all to the light
- Convergent scotopic system pays for high degree of sensitivity with low level acuity, no way of knowing which portion of rods contributed to change
- More intense light is required to change firing of a retinal ganglion cell that receives signals from cones, when such a cell does react there is less ambiguity about the location of the stimulus that triggered the reaction
- There are no rods at all in the fovea, only cones, at boundaries of foveal indentation the proportion of cones decline and the density of rods reaches a maximum (there are more rods in the nasal hemiretina, the half of each retina next to the nose than in the temporal hemiretina, the half next to the temples)
Light enters the Eye and reaches the Retina:
Spectral Sensitivity:
- More intense lights appear brighter however wavelength has substantial effect on perception of brightness
- Visual systems are not equally sensitive to all wavelengths, lights of the same intensity but of different wavelengths can differ markedly in brightness
- Graph of relative brightness = spectral sensitivity curve
- Cone = Photopic spectral sensitivity curve, can be determined by having subjects judge the relative brightness of different wavelengths of light shone on the fovea
- Rod = Scotopic spectral sensitivity curve, determined by asking subjects to judge relative brightness of different wavelengths of light shone on the periphery of the retina at an intensity too low to activate the few peripheral cones that are located there
- Purkinje Effect: Tendency for the peak luminance sensitivity of the human eye to shift toward the blue end of the colour spectrum at low illumination levels as part of dark adaptation
Light enters the Eye and reaches the Retina:
Eye Movement:
- Eyes continually scan visual field, visual perception at any instant is a summation of recent visual information, it is because of this temporal integration that the world does not vanish momentarily each time we blink
- Even when we fix our gaze on an object our eyes constantly move
- Involuntary fixational eye movements are of 3 kinds: Tremor, drifts and saccades (small jerky movements or flicks)
- Normally unaware of these, have critical visual function, must fix our gaze to perceive minute details, if we were to fixate perfectly world would fade/disappear, visual neurons respond to change, if retinal images are artificially stabilised (kept from moving on the retina) the images start to disappear and reappear
- -> Fixational eye movements enable us to see during fixation by keeping the images moving on retina
Light enters the Eye and reaches the Retina:
Visual Transduction: The conversion of Light to Neural Signals:
- Transduction is the conversion of one form of energy to another
- Visual transduction is the conversion of light to neural signals by the visual receptors
- Red pigment extracted from rod (rhodopsin), exposed to continuous intense light, became bleached, lost ability to absorb light, when it was returned to dark it regained its redness and light absorbing ability, now clear that rhodopsin’s absorption of light is first step in rod-mediated vision
- Degree to which rhodopsin absorbs lights of different wavelengths is related to the ability of humans and other animal’s rods to detect the presence of different wavelengths of light under scotopic conditions
- Rhodopsin is a G-protein coupled receptor that responds to light rather than to neurotransmitter modules, initiate a cascade of intracellular chemical events when they are activated
- When rods are in darkness their sodium channels are partially open, this keeps the rods slightly depolarised allowing a steady flow of excitatory glutamate neurotransmitter molecules to emerge from them. However, when the rhodopsin receptors are bleached by light the resulting cascade of intracellular chemical events closes the sodium channels, hyperpolarises the rods and reduces the release of glutamate
- The transduction of light by rods exemplifies that signals are often transmitted through neural systems by inhibition
From Retina to Primary Visual Cortex
- Largest studied pathways are the retina-geniculate-striate pathways which conduct signals from each retina to the primary visual cortex or striate cortex, via the lateral geniculate nuclei of the thalamus
- 90% of axons of retinal ganglion cells become part of the retina-geniculate-striate pathways
- All signals from left visual field reach the right primary visual cortex, either ipsilaterally from the temporal hemiretina of the right eye or contralaterally via the optic chiasm from the nasal hemiretina of the left eye and that the opposite is true of all signals from the right visual field
- Each lateral geniculate nucleus has 6 layers and each layer of each nucleus receives input from all parts of the contralateral visual field of one eye
- -> Each lateral geniculate nucleus receives visual input from only the contralateral visual field, 3 layers receive input from one eye and 3 from the other
- ->Most of the lateral geniculate neurons that project to the primary visual cortex terminate in the lower part of the cortical layer IV producing a characteristic stripe or striation when viewed in cross section (hence striate cortex name)
From Retina to Primary Visual Cortex:
Retinotopic Organisation:
- The retina-geniculate-striate system is Retinopic: each level of the system is organised like a map of the retina
- 2 stimuli presented to adjacent areas of the retina excite adjacent neurons at all levels of the system
- Retinopic layout of the primary visual cortex has a disproportionate representation of the fovea although the fovea is only small part of the retina, large proportion (25%) is dedicated to analysis of input
From Retina to Primary Visual Cortex:
The M and P Channels:
- 2 parallel channels of communication flow through each lateral geniculate nucleus
- One channel runs through the top 4 layers (Parvocellular layers) /P layers), are composed of neurons with small cell bodies, are particularly responsive to colour, fine pattern details and stationary or slowly moving objects, cones provide majority of input to P layers
- Other channel runs through bottom 2 layers (Magnocellular layers/M layers), are composed of neurons with large cell bodies, are particularly responsive to movement, rods provide majority of input into M layers
- -> Both neurons project to different sites in the lower part of the layer IV of the striate cortex, in turn the M and P portions of lower layer IV project to different parts of visual cortex
SUMMARY TASK:
- Neural signals are carried from the retina to the lateral geniculate nuclei by the axons of retinal ganglion cells
- The axons of retinal ganglion cells leave the eyeball at the blind spot
- The area of the retina that mediates high-acuity vision is the fovea
- Cones are the receptors of the Photopic system, which functions only in good lighting
- The retinal ganglion cells from the nasal hemiretina decussate (cross over to the other side of the brain) via the optic chiasm
- The photo pigment of rods is rhodopsin
- The most important organisational principle of the retina-geniculate-striate system is that it is laid out retonotopically
- Rhodopsin was implicated in scotopic vision by the fit between the absorption spectrum of rhodopsin and the scotopic spectral sensitivity curve
- The high degree of convergence characteristic of scotopic system increases its sensitivity but decreases its acuity
Seeing Edges
A visual edge is merely the place where 2 different areas of a visual image meet, the perception of an edge is really the perception of a contrast between 2 adjacent areas of the visual field