WEEK 2 - Visual pigment

o Light-sensitive chemicals
CHAPTER 3 o React to light & trigger electrical signals
The Eye and Retina
Retinitis pigmentosa Optic nerve
- Genetic disorder of the eye resulting in total blindness - Where signals emerge from the receptors
- Conduct signals toward the brain
Bionic eye
- Array of electrodes implanted in the back of eye that
(through cam in glasses) sends signals to visual
system of what is “out there”

First 4 steps of visual process

Light, the Eye, and the Visual Receptors

The ability to see anything depends on the light being reflected.

Light: The Stimulus for Vision

Visible light
- A band of energy within the electromagnetic spectrum
- Energy that humans can perceive
- Has wavelengths ranging from 400-700nm (1nm = 10-
9 meters)
- For humans & other animals, its wavelengths are
associated w different colors of spectrum (short = B,
middle = G, long = YOR)
From the rod & cone distributions, we can conclude that
1. The fovea contains only cones. When we look directly
at an object, its image falls on this.
2. The peripheral retina (includes all retina outside of
fovea) contains both rods and cones
3. The peripheral retina contains more rods than cones
bc there are 120M rods and 6M cones in retina

Electromagnetic spectrum Macular degeneration

- Continuum of electromagnetic energy produced by
electric charges and radiated as waves
- Energy can be described by its wavelength
o Distance between peaks of electromagnetic
waves
o Short gamma rays (10-12 meters) to long
radio waves (104 meters)
- Most common in elders
- Destroys the cone-rich fovea and a small area that
surrounds it
- Macula is usually associated with medical practice
that includes fovea + small area surrounding
- Ex. when looking directly at something, u lose sight of
it

The Eye
Eyes
- Contain receptors for vision
- First eyes
o Appeared in Cambrian period (500-700
million years ago)
o Eyespots on primitive animals (ex.
flatworms) that can distinguish light from
Retinitis pigmentosa
dark
- Degeneration of retina passed from 1 generation to
the next (but NOT always affecting everyone in fam)
Pupil
- First attacks peripheral rod receptors, resulting in poor
- Light reflected from objects in environment enteres
vision in peripheral field
the eye through this
Blind spot
Cornea lens
- Place of absence of photoreceptors
- Focuses on the light to form sharp images of the
- Where the nerve fibers that make up the optic nerve
objects
leave the eye
Retina
- Network of neurons that covers the back of the eye
and contains photoreceptors

Photoreceptors
- Receptors for vision
- 2 types
o Rods
o Cones Why are we not usually aware of the blind spot?
- Outer segments 1. It is located off the side of our visual field
o Rod- and cone-shaped 2. A mechanism in the brain “fills in” the place where
o Part of receptor that contains visual pigment image disappears

Focusing Light Onto the Retina
2-element optical system of light being reflected and focused
onto retina
1. Cornea (transparent covering, 80% of eye’s focusing
power, fixed place so cannot adjust focus)
2. Lens (remaining 20%, can change shape to adjust
focus)
Ciliary muscles
- Increase the focusing power of lens (its ability to bend
light) by increasing its curvature
o Occurs in photoreceptors (rods & cones) and
transforms light into electricity
o ^ create electricity bc of millions of molecules
of light-sensitive visual pigment contained in
outer segments of photoreceptors
2 parts of visual pigments
1. Opsin (long protein)
2. Retinal (smaller light-sensitive component; CRUCIAL
part bc when combined with opsin, the molecule
absorbs visible light)

More than 20 feet away

- Parallel light rays

Accommodation
Accommodation
- Change in the lens’s shape that occurs when the
ciliary muscles at the front tighten & increase the
curvature of the lens so it gets thicker
- Increased curvature = increased bending of light rays
(focus is pulled from point B back to A)
Steps in transduction
1. When light hits retina, the visual pigment molecule
absorbs it
2. Retinal changes its shape from bent to straight
(isomerization)
3. Isomerization creates chemical reaction, activating
thousands of charged molecules to create electrical
signals in receptors
4. The initial isomerization of 1 visual pigment molecule
can lead to activation of entire photoreceptor
5. Electrical signal is created

2 aspects of visual perception

1. How we adjust to darkness
2. How well we see light in diff parts of spectrum

Adapting to the Dark

Dark adaptation
- Process of increasing sensitivity in the dark

Dark adaptation curve

Refractive Errors
- Measures dark adaptation
Refractive errors
- Errors that can affect the ability of cornea and/or lens
Measuring the Dark Adaptation Curve
to focus the visual input onto the retina
1. Looking at small fixation point, while paying attention
to flashing test light at the side
Presbyopia
2. Main image falls on the fovea, and test light image
- “old eye”
falls to the side: peripheral retina
- Age-related loss of ability to accommodate
- Can be dealt with wearing glasses

Myopia
- Nearsightedness
- Inability to see distant objects clearly
- Affects more than 70M Americans
- Optical system brings parallel rays of light into focus
in front of retina, so the image that reaches retina is
blurred
- Caused by 2 factors
o Refractive myopia – cornea/lens bends light
too much
o Axial myopia – eyeball is too long
- Corrective lenses
Sensitivity
Hyperopia - Sensitivity = 1/threshold
- Farsightedness - High threshold = low sensitivity
- Have trouble seeing nearby objects
- Focus point for parallel is located behind the retina (bc Light-adapted sensitivity
eyeball is too short) - Sensitivity measured in the light (while the eyes are
- Young people = accommodating, older people = adapted to the light)
corrective lenses
Photoreceptor Processes
Transforming Light Energy Into Electrical Energy
Transduction
- Transformation of 1 form of energy into another form
- Visual transduction

2 phases that sensitivity increases
1. It increases rapidly for first 3-4 minutes after light is
extinguished, then levels off
2. It increases again at 7-10 minutes until he has been in
the dark for 20-30 minutes
Dark-adapted sensitivity
- Sensitivity at the end of dark adaptation
- 10,000 greater than the light-adapted sensitivity
measured before dark adaptation
The first part of dark adaptation is caused by cones, the
second is rods.
Cones
- More sensitive to light at beginning of dark adaptation,
so they control our vision during early stages
Rod monochromats
- People who have no cones bc of rare genetic defect
Dark adaptation process
1. When light is extinguished, the sensitivity of both
cones and rods increase
2. But bc the cones are more sensitive, we see with
cones right after the lights are turned out
3. After 3-5 minutes, cones have reached maximum
sensitivity
4. By 7 minutes, the rods’ sensitivity finally catches up,
becoming more sensitive than cones
Rod-cone break
- Place where rods begin to determine dark adaptation
curve instead of cones
Visual Pigment Regeneration
Visual pigment bleaching
- Change in shape & separation from opsin
- Causes molecule to become lighter in color
- Thus, no longer useful for vision
Visual pigment regeneration
- Process of reforming the visual pigment molecule
- Occurs more rapidly in cones than rods
- Cone pigment = 6 minutes, rod pigment = more than
30 minutes
2 important connections between perception & physiology
1. Out sensitivity to light depends on concentration of
chemical – visual pigment
2. The speed at which out sensitivity increases in the
dark depends on chemical reaction – regeneration
Pigment epithelium
- Layer containing enzymes necessary for pigment
regeneration
Detached retina
- Occurs due to traumatic injuries of eye or head
- Bleached pigment’s separated retinal & opsin can no
longer be recombined
Spectral Sensitivity
Spectral sensitivity
- Eye’s sensitivity to light as a function of the light’s
wavelength
- Measured by spectral sensitivity curve
o Relationship between wavelength and
sensitivity
Spectral Sensitivity Curves
Monochromatic light
- Light of single wavelength
- Created by special filters or spectrometer
Cone spectral sensitivity
- Measured by making participant look directly at test
light, so it stimulates only cones in fovea
Rod spectral sensitivity
- Measured by presenting test flashes in peripheral
retina
Rods vs Cones
- R more sensitive to short-wavelength light 500nm; C
more sensitive to light of 560nm
- R more sensitive in dark; C more sensitive in light
- Cone pigment regenerates faster than Rod pigment
As vision shifts from cones to rods, our vision shifts to become
more sensitive to short-wavelength light—near blue & green.
Purkinje shift
- Johann Purkinje
- Enhanced perception of short wavelengths during
dark adaptation
- Ex. blue flower being brighter than red in your dark-
adapted eye
Rod- and Cone-Pigment Absorption Spectra
Absorption spectrum
- Plot of the amount of light absorbed vs wavelength of
light
3 absorption cone pigments
1. Short-wavelength pigment (S) – absorbs light at 419
nm
2. Medium-wavelength pigment (M) – 531 nm
3. Long-wavelength pigment (L) – 558 nm
The increase in sensitivity occurring in the dark (dark
adaptation) and the sensitivity to diff wavelengths across the
spectrum (spectral sensitivity) are determined by the properties
of rod & cone visual pigments.
What Happens as Signals Travel Through the Retina
Rod and Cone Convergence
Neural circuits
- Interconnected groups of neurons

Bipolar cells (B)

- Where signals generated in receptors (R) travel to

Ganglion cells (G)

- Where signals travel to after ^
- Have long axons
o They transmit signals out of retina in the
optic nerve

2 types of neurons that connect neurons across retina

1. Horizontal cells
2. Amacrine cells
Perception Is Shaped by Neural Convergence
Neural convergence
- Occurs when a number of neurons synapse onto a
single neuron
- Occurs in retina (bc each eye has 126 million
photoreceptors but only 1 million ganglion cells, thus
each ganglion cell receives signals from 126
photoreceptors)
The signals from the rods converge more than the signals from
cones. 120 rods send signals to 1 ganglion cell, but only 6
cones send signals.
2 differences in perception
1. The rods result in better sensitivity than cones
2. The cones result in better detail vision than the rods
Convergence Causes the Rods to Be More Sensitive Than the
Cones
Greater sensitivity of rods
- Bc it takes less light to generate a response from an
individual rod receptor than cone receptor
- Bc rods have greater convergence than cones
The cones’ lack of convergence causes cone vision to have
higher acuity than rod vision. (bc 2 separate ganglion cells fire)
High convergence = high sensitivity but poor acuity (rods)
Low convergence = low sensitivity but high acuity (cones)
Ganglion Cell Receptive Fields
Hartline’s Discovery of Receptive Fields
H. Keffer Hartline
- Isolated 1 ganglion cell axon in the opened eyecup of
frog by teasing apart the optic nerve
Receptive field
- Region of retina that must receive illumination to
obtain a response in any given fiber
- Area that caused the neuron to fire the ganglion cell
- Ganglion cell’s receptive field covers greater area
than single photoreceptor
- Ex. football field (retina) analogy with spectators
(ganglion cell) having binoculars and viewing small
areas (receptive fields) each
Kuffler’s Discovery of Center-Surround Receptive Fields
Stephen Kuffler
- Measured ganglion cell receptive fields in cat
- Cat and other mammals’ (monkeys & humans)
ganglion cells have center-surround receptive fields

Center-surround receptive fields

- Arranged like concentric circles in a center-surround
organization
- The area in the “center” of receptive field responds
differently to light than area in the “surround”

Unit of excitation
- When a photoreceptor is stimulated by 1 spot of light
- It takes 10 units for ganglion cell to fire
- 10-unit threshold
- Ex. spots of light with 1 intensity = rod ganglion cell
receives 5 units of excitation (1 from each rod
receptor); cone ganglion cell receives only 1 (1 from
each cone receptor) BUT not enough to fire
Excitatory-center, inhibitory-surround receptive field
It takes less light to stimulate a ganglion cell receiving inputs - Excitatory area
from rods. (bc they converge) o Presenting spot of light to center increases
firing
Less Convergence Causes the Cones to Have Better Acuity - Inhibitory area
Than the Rods o Stimulation of surround causes decreases
Visual acuity firing
- Acuity refers to the ability to see details
- Cones have better/high visual acuity bc of less Inhibitory-center, excitatory-surround receptive field
convergence - Inhibition when center is stimulated, excitation when
- Ex. scanning for an item with cone-rich fovea (used surround is stimulated
when u need to look directly at something), not clear
in periphery Center-surround antagonism
- Ex. bookshelf turning dark at the bottom (color vision - Effect where response begins to decrease when the
and detail depends on the cones) size of the spot is increased so it stimulates part of the
inhibitory surround (c) and (d)
Foveal cones vs Peripheral cones
- FC tightly packed; PC widely spaced
 - Stimulation of inhibitory surround counteracts the - Although the intensity is same, the perception of light
center’s excitatory response, causing a DECREASE is NOT
in neuron’s firing rate
Chevreul illusion
- Michel-Eugene Chevreul
o Became interested in how placing colors side
by side can alter their appearance
- Perceived light and dark bands at the borders which
are NOT present in the actual physical stimuli

Mach bands
- Ernst Mach
- Borders where light and dark bands are created at
Lateral inhibition
- Inhibition transmitted across the retina (literally)

Lateral Inhibition Underlies Center-Surround Antagonism

Limulus
- Horseshow crab
- The structure of its eye makes it possible to stimulate
individual receptors
- Its eye is made up of hundreds of tiny structures
called ommatidia
o Each ommatidium has small lens on eye’s
surface located directly over single receptor
o Each lens & receptor is diameter of a pencil
point (larger compared to human receptors)

Perception is the outcome of neural processing – neural

processing that occurs in the center-surround receptive fields
in retina.

Lateral plexus
- Transmits signals laterally in the Limulus

Center-Surround Receptive Fields and Edge Enhancement

Edge enhancement
- Increase in perceived contrast at borders between
regions of the visual field
- Helps make edges look more distinct so we can see
them more easily