Photoreceptors

• In animals light is detected by?

– The eyes
• The structure of the eye varies for
different organisms.
Visual receptors on diverse animals
depend on light-absorbing pigments
• Animals use a diverse set of organs for vision, but the
underlying mechanism for capturing light is the same,
suggesting a common evolutionary origin

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 Evolution of Visual Perception
• Light detectors in the animal kingdom range from
simple clusters of cells that detect direction and
intensity of light to complex organs that form images
• Light detectors all contain photoreceptors, cells that
contain light-absorbing pigment molecules

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 Light-Detecting Organs
• Most invertebrates have a light-detecting organ
• One of the simplest light-detecting organs is that of
planarians
• A pair of ocelli called eyespots are located near the head
• These allow planarians to move away from light and
seek shaded locations

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 TO REMEMBER

-Although animals use many types of light detectors

(phtoreceptors), all of them use similar pigments (opsins).

-Planaria have very simple ocelli (eye spots).

-Invertebrates have ocelli, compound eyes (artropods,

annelids), and single-lens eyes (octopi, squid).

-Compound eyes are made of omatidia (cornea, crystalline,

rhabdom, photoreceptor)..

-Single-lens eyes of vertebrates have sclera, cornea, lens,

iris, retina, optic nerve.
 Structure of Eyes
• Snails
– Have simple eyes which detect light in
the same way as we detect heat.
• Insects and Crustaceans
– Have compound eyes made up of many
separate ommatidia. These give a mosaic
vision, very good for detecting
movement.
Figure 50.15

LIGHT

DARK
(a)

Light

Photoreceptor
Ocellus Nerve to
Visual brain
pigment
Screening
Ocellus pigment
(b)
Vision Light
Photoreceptors
• Planarians: Ocelli or eye
spots in the head region
• Light stimulates
photoreceptors Light shining from
the front is detected
• Brain compares rate of action
potential coming form the
two ocelli Photoreceptor
• Brain directs the body to turn
until sensation form both
ocelli are equal and minimal Nerve to
• Animal can move to shade, brain
Visual pigment
under a rock away from
Screening
predators pigment
Ocellus

Light shining from

behind is blocked
by the screening pigment
 Vision

photoreceptors contain similar

pigment molecules that absorb light.
Most of these pigments (opsins) are
homologous (evolved from a
common ancestral molecule).

Planarians have two ocelli (light detecting

The cross-eyes in planaria
serve a purpose!!
organs, sometimes called eye spots or eye
cups) that allow the animal to sense light
(and often turn away from light). Orientation
comes about by “comparing” light entering
through each side.
 Compound Eyes
• Insects and crustaceans have compound eyes, which
consist of up to several thousand light detectors called
ommatidia
• Compound eyes are very effective at detecting
movement

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Compound eyes:

• Very good at detecting movement

• Very good at detecting flickering light (6 times
faster than human eye)
• Some bees can see in the ultraviolet range of
light
• Several thousand omatidia (facets) in every eye
• Cornea and crystalline cone form the lens which focuses
light on the rhabdom
• Light stimulates the photoreceptors to generate receptor
potential which generates action potential
Cornea

Crystalline Lens
cone

Rhabdom

Photoreceptor
Axons
2 mm

Ommatidium
Compound eyes are made of lots of
ommatidia. The cornea and
crystalline act as a lens that focuses
light. Each ommatidium detects light
on a narrow portion of the visual
field and then the brain integrates
this mosaic. Some insects can
detect color in the ultraviolet range.
We cannot extrapolate our
sensory world to other species.

We Insect (maybe)
Compound Eyes
 Single-Lens Eyes
• Single-lens eyes are found in some jellies, polychaetes,
spiders, and many molluscs
• They work on a camera-like principle: the iris changes
the diameter of the pupil to control how much light
enters
• The eyes of all vertebrates have a single lens

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Two types of image-forming eyes have evolved among invertebrates.
Compound eyes in arthropods (also some polychates) and single-lens eyes
which have evolved in some polychaetes (annelids), in spiders, and in octopi
and squid.

Arthropod compound eye Single-lens eye of squid

Vertebrate eye
• Single lens system (very different from
invertebrate single eyes)
 The Vertebrate Visual System
• In vertebrates the eye detects color and light, but the
brain assembles the information and perceives the
image

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 Structure of Eyes
• Vertebrates
– Have lenses that help to form clear
images.
– If two eyes face forwards they have
overlapping fields (Binocular vision)
enabling good judgement of distance
Vertebrate Eyes
 Structure of Eyes
• Night Vision
– This involves a structure called the
tapetum, this is a layer of silvery crystals
that acts as a reflector allowing animals
like cats to pick up 50% more light at
night than human’s can detect.
– This is why cat’s eyes glow in the dark.
Night Vision
 Vision
• Animals also use their vision to detect
movement, speed of movement, shapes etc.
 Colour Vision
• We, as humans see violet, blue, green,
yellow, orange and red parts of the
light spectrum.
• Bees see ultraviolet light but are less
sensitive to red. This is why most red
flowers have some blue or a strong
scent.
How bees see
 Colour Vision cont
• Goldfish can see far red light.

• Sea birds are particularly sensitive to

red light.
The retina contains
photoreceptors (rods and cones),
cells that integrate information
across the retina (horizontal cells
and amacrine cells), and cells that
receive information from several
rods and cones (bipolar cells) and
relay it to ganglion cells which
transmit action potentials to the
brain by the optic nerve.
The photoreceptors in the retina of
vertebrate eyes have two
morphologies: they can be rod-like
(“rods”)
or
cone-like (“cones”).

Rods

Cones
Rods are very good at detecting light
at low intensities and at the low end of
the electromagnetic spectrum, but are
not good at distinguishing colors.
Cones can detect variation in colors
but are not as sensitive to light as
rods. The reason for this difference is
in the molecular structure of the
pigments (opsins) contained in these
two types of receptors. All vertebrate
classes (fishes, amphibians, reptiles,
and birds) have color vision. Most
mammals are nocturnal and hence
have mostly rods. Old World primates
(including humans) are exceptional in rod
that we have lots of cones (we will
discuss the evolution of color vision a
bit later).
In addition to the opsin found in rods (called rhodopsin),
humans (and most Old World primates) have 3 different
pigments which absorb (and hence are stimulated) at different
peaks of the spectrum. For this reason, our visual system is
called “trichromatic”
 TO REMEMBER
- The retina contains photoreceptors (rods and cones), cells
that integrate information across the retina (horizontal cells and
amacrine cells), and cells that receive information from
several rods and cones (bipolar cells) and relay it to
ganglion cells which transmit action potentials to the brain
by the optic nerve.
-Each optic nerve has axons that connects with interneurons in
a structure called the geniculate nuclei. These interneurons relay
sensations to the primary visual cortex, which is one of the
brain centers responsible in constructing visual perceptions.
-Rods are good at detecting light at low intensities but are not
good at distinguishing colors. Cones can detect variation in
colors but are not as sensitive to light as rods.
-Our visual system is called “trichromatic” because in addition to
rhodopsin, we have 3 different pigments which absorb at different
peaks of the spectrum.
Rods contain the visual pigment rhodopsin,
which is embedded in a stack of membrane
disks. Rhodopsin consists of the protein
(pigment) opsin and the light absorbing
molecule retinal (there are many opsins!).
Retinal exists in two forms. Light
converts the cis form to a trans
form and enzymes return it to its
original form.
Sea Birds Vision
Structure and Function
of the Human Eye
 palpabre

Lacrimal cornea sclera

caruncle
Medial lateral
commisure commisure

tear iris
drainage pupil
canal palpabre
bright light normal light dim light
FLOW OF TEARS
Lacrimal gland

Lacrimal ducts

Sup. or inf. lacrimal canal

Lacrimal sac

Nasolacrimal duct

Nasal cavity
 Superior oblique
Superior rectus

optic nerve

Medial
rectus

Inferior oblique Lateral rectus

Inferior rectus
Fibrous tunic- sclera and cornea (outer
most layer)
Cornea
• 100s of sheets of collagen fibers between
sheets of epithelium and endothelium
• Clear because regular alignment
• Role in light bending
• Avascular but does have pain receptors
• Regenerates
Vascular tunic- uvea: choroid, cilliary body, iris, pupil (middle layer)
Choroid- rich vascular nutritive layer; contains a dark pigment
that prevents light scattering within the eye
Cilliary body- lens is attached; contains muscles that change the
lenses shape
Iris- pigmented ring of muscular tissue composed of circular
and radial muscles
• reflex contraction of circular muscle in bright light (small dia of
pupil)
• reflex contraction of radial muscle in dim light (large dia of pupil)
Pupil- central hole in iris 
Sensory tunic- retina (inner most layer)
Photoreceptors:
• rods (dim light, contains pigment rhodopsin)
120 million rods
• Cones (color vision, not evenly distributed,
concentrated in fovea) 6 million cones
Optic disc- blind spot because its where optic nerve
leaves the eyeball (no rods or cones)
Macula lutea- yellow spot, area of high cone
Fovea centralis- in center of macula lutea, contains only
cones, area of greatest visual acuity
Figure 50.17aa
Choroid
Sclera
Retina
Suspensory Fovea
ligament
Cornea
Iris
Optic
nerve
Pupil

Aqueous
humor
Lens Central
artery and
vein of
Vitreous humor Optic disk the retina
Vitreous humor- behind lens, gel-like substance
with fine collagenic fibrils imbedded in as
viscous ground substance- binds with water

• transmits light
• supports the posterior surface of the lens and
holds the neural retina firmly against
pigmented layer
• contributes to intraoccular pressure, helping to
counter act the pulling force of the extrinsic
eye muscles
Aqueous humor- in front of lens, anterior
segment, watery fluid

• Supplies cornea and lens with nutrients

• Helps to maintain the shape of the eye
• Produced and renewed every 4 hrs by the
cilliary body      
Lens- transparent biconvex structure, flexible
• Attached by suspensory ligaments to ciliary
body
• focuses image onto retina
• changes lens thickness to allow light to be
properly focused onto retina
Coarse Fixed Focusing
• Cornea Shape
Accommodation- adjust configuration of
• Lens Shape
• Pupil Size
 Front view of lens
and ciliary muscle
Humans and other mammals
Choroid Lens (rounder)

• spherical ( ciliary muscles Retina

contract, sensory
ligaments relax – near
Ciliary
objects) muscle

Suspensory
• flatter (ciliary muscles Near vision (accommodation) ligaments
relax, edge of choroid
moves away from lens,
suspensory ligaments Lens (flatter)
contract and pull the lens
– distant objects)

Distance vision
• Fishes, squids and octopuses focus by
moving lens forward and backward
 The Retina

blind
macula
spot
Each optic nerve has ≈
106 axons that connects
with interneurons in a
structure called the
geniculate nuclei. These
interneurons relay
sensations to the primary
visual cortex, which is
one of the brain centers
responsible in
constructing visual
perceptions.
 Figure 50.17a

Choroid Retina
Sclera
Retina Photoreceptors
Suspensory Neurons
Fovea Rod Cone
ligament
Cornea
Iris
Optic
nerve
Pupil

Aqueous
humor
Lens Central
artery and
vein of
Vitreous humor Optic disk the retina
Optic Amacrine Horizontal cell
nerve cell Pigmented
fibers Ganglion Bipolar epithelium
cell cell
photoreceptors Rod cell membrane
Figure 50.17ab
Retina
Photoreceptors
Neurons
Rod Cone

Optic Amacrine Horizontal cell

nerve cell Pigmented
fibers Ganglion Bipolar epithelium
cell cell
 Figure 50.17b

Rod CYTOSOL

Synaptic Cell Outer Disks Retinal: cis isomer

terminal body segment
Light Enzymes
Cone

Rod
Retinal: trans isomer
Cone
Retinal
Rhodopsin
Opsin
INSIDE OF DISK
Figure 50.17ba

Rod

Synaptic Cell Outer Disks

terminal body segment

Cone

Rod

Cone
Figure 50.17bb

CYTOSOL

Retinal
Rhodopsin
INSIDE OF DISK Opsin
Figure 50.17bc

Retinal: cis isomer

Light Enzymes

Retinal: trans isomer

Figure 50.17bd

Rod

Cone
refraction
Fig. 6-13, p. 194
Vision or Sight
• Visible light: 400-700 nm.

Copyright 2009, John Wiley & Sons,

Inc.
 Vision
• Photoreceptors of the retina.
– rod cells - light sensitive but do not distinguish
colors.

– cone cells - not as light sensitive as rods but

provide color vision

• Most highly concentrated on the fovea – an area of the

retina that lacks rods.

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 Vision
• Rhodopsin (retinal + opsin) is the visual pigment
of rods.
• The absorption of light by rhodopsin initiates a
signal-transduction pathway
• Receptor potential is hyperpolization .

Fig. 49.13
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 Photon

Retinal changes shape

Retinal and
opsin are
reassembled
to form
rhodopsin

Bleaching
Regeneration enzyme (separation)
Retinal
restored ADP ATP

Opsin Opsin

Opsin
inactivated

Figure 9-20
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 Retinal and
opsin are
reassembled
to form
rhodopsin

Figure 9-20
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 Photon

Retinal and
opsin are
reassembled
to form
rhodopsin

Figure 9-20
Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings 3 of 7
 Photon

Retinal changes shape

Retinal and
opsin are
reassembled
to form
rhodopsin

Figure 9-20
Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings 4 of 7
 Photon

Retinal changes shape

Retinal and
opsin are
reassembled
to form
rhodopsin

Bleaching
enzyme (separation)
Retinal
restored ADP ATP

Figure 9-20
Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings 5 of 7
 Photon

Retinal changes shape

Retinal and
opsin are
reassembled
to form
rhodopsin

Bleaching
enzyme (separation)
Retinal
restored ADP ATP

Opsin Opsin

Opsin
inactivated

Figure 9-20
Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings 6 of 7
 Photon

Retinal changes shape

Retinal and
opsin are
reassembled
to form
rhodopsin

Bleaching
Regeneration enzyme (separation)
Retinal
restored ADP ATP

Opsin Opsin

Opsin
inactivated

Figure 9-20
Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings 7 of 7
 Sensory Transduction in the Eye
• Transduction of visual information to the nervous
system begins when light induces the conversion of cis-
retinal to trans-retinal
• Trans-retinal activates rhodopsin, which activates a G
protein, eventually leading to hydrolysis of cyclic GMP

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 • When cyclic GMP breaks down, Na channels close
• This hyperpolarizes the cell
• The signal transduction pathway usually shuts off again
as enzymes convert retinal back to the cis form

© 2011 Pearson Education, Inc.

 Figure 50.18

INSIDE OF DISK EXTRA-

Light Disk CELLULAR
Active membrane FLUID
rhodopsin Phosphodiesterase
Plasma
membrane
CYTOSOL

cGMP
Inactive Transducin
rhodopsin GMP
Na

Dark Light
0
potential (mV)
Membrane

Hyper-
40 polarization
Na
70
Time
 The Special Senses—Vision
• The Visual Pathway
– Ganglion cells axon converge at optic disc
– Axons leave as optic nerve (CN II)
– Some axons cross at optic chiasm
– Synapse in thalamus bilaterally
– Thalamic neurons project to visual cortex
• Located in occipital lobes
• Contains map of visual field

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Processing of Visual Information in the
Retina
• Processing of visual information begins in the retina
• In the dark, rods and cones release the neurotransmitter
glutamate into synapses with neurons called bipolar
cells
• Bipolar cells are either hyperpolarized or depolarized in
response to glutamate

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 • In the light, rods and cones hyperpolarize, shutting off
release of glutamate
• The bipolar cells are then either depolarized or
hyperpolarized

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 Bleaching and Regeneration of
Photopigment
1. Isomerization: In darkness, retinal has a
bent shape called cis-retinal. Absorption of
photon causes straightening of the retinal
(trans-retinal).
2. Bleaching: trans-retinal separates from
opsin.
3. Regeneration: trans-retinal→ cis-retinal.

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Inc.
 Light and Dark Adaptation
• Light adaptation: Dark → light. Faster.
• Dark adaptation: Light →dark. Slow.
• Cones regenerate rapidly whereas
rhodopsin regenerates more slowly.

Copyright 2009, John Wiley & Sons,

Inc.
Figure 50.19
Dark Responses Light Responses

Rhodopsin inactive Rhodopsin active

Na channels open Na channels closed

Rod depolarized Rod hyperpolarized

Glutamate released No glutamate

released

Bipolar cell either Bipolar cell either

depolarized or hyperpolarized or
hyperpolarized, depolarized,
depending on depending on
glutamate receptors glutamate receptors
 • Three other types of neurons contribute to information
processing in the retina
– Ganglion cells transmit signals from bipolar cells to the
brain
– Horizontal and amacrine cells help integrate visual
information before it is sent to the brain
• Interaction among different cells results in lateral
inhibition, enhanced contrast in the image

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Binocular vision
Processing of Visual Information in the
Brain
• The optic nerves meet at the optic chiasm near the
cerebral cortex
• Sensations from the left visual field of both eyes are
transmitted to the right side of the brain
• Sensations from the right visual field are transmitted to
the left side of the brain

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 • Most ganglion cell axons lead to the lateral geniculate
nuclei
• The lateral geniculate nuclei relay information to the
primary visual cortex in the cerebrum
• At least 30% of the cerebral cortex, in dozens of
integrating centers, is active in creating visual
perceptions

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 Figure 50.20

Right Optic chiasm

visual
field

Right
eye

Left
eye
Primary
Left Optic nerve visual
visual Lateral
cortex
field geniculate
nucleus
 Color Vision
• Among vertebrates, most fish, amphibians, and reptiles,
including birds, have very good color vision
• Humans and other primates are among the minority of
mammals with the ability to see color well
• Mammals that are nocturnal usually have a high
proportion of rods in the retina

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 • In humans, perception of color is based on three types
of cones, each with a different visual pigment: red,
green, or blue
• These pigments are called photopsins and are formed
when retinal binds to three distinct opsin proteins

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 • Abnormal color vision results from alterations in the
genes for one or more photopsin proteins
• In 2009, researchers studying color blindness in squirrel
monkeys made a breakthrough in gene therapy

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Figure 50.21
Focusing on a Near Object
Focusing on a Far Object
• Emmetropia- normal 20:20
• Hyperopia- farsighted
• Myopia-near sighted
• Presbyopia- mature eyes
• Astigmatism
20 ft:20 ft
You see Normal vision
 What condition
does this
person have?

What condition
20/10 does this
person have?
                                                                           

                                  
Clouding of lens (hardening or thickening
causes: diabetes mellitus, smoking, UV
damage
Glaucoma
Effect: Subjective or illusory contours
 Geometrical illusions

The apparent tilt of the mortar lines is caused by

orientation-sensitive simple cells in the striate cortex.
 Geometrical illusions

Cushion Illusion: This drawing consists solely of

rectangles and squares, set straight and true; the
curvature is all in your mind.
Luminance & Contrast
Luminance & Contrast
The scintillating grid illusion
Effect: Illusory visual motion
Effect: Illusory visual motion
 It’s a spiral right?

No, these are a bunch of independent circles

 Successive contrast : afterimages ...

fixate the black dot in the center for 60 seconds ...

… and then look at a the black dot in the right panel !

                                            what do you see?