The Evolution of the Eye!!!

By Dan Zembrosky and Dr. Pepper, M.D.

Types of Eyes
The Simple Eye (3 Main Kinds):

1. The Positive Lens or Camera Eye

Found in vertebrates and some
invertebrates. This is the type of
eye humans have. You can even
poke it with your finger right now.
Go ahead, gently. Thats right.

Types of Eyes
The Simple Eye
2. The Concave Mirror Eye
Found in the clam Pecten and a few
ostracod crustaceans. This produces
bright but reasonably hazy picture.
3. The Pinhole Eye
Pit or Cup eyes are found mainly in
mollusks and can only resolve
location of objects.

The Compound Eye

Made up of ommatidia receptors, each of which
functions as a separate visual receptor.
The Apposition Eye
Ommatidia function
independently.
The Superposition Eye
Ommatidia cooperate to produce a brighter,
superimposed image on the retina.

The Two Kinds of Photoreceptors

All photoreceptors use a light sensitive pigment
derived from vitamin A which is bound to an opsin.
After being exposing the photopigment to light the
opsin binds to a G-protein (common
neurotransmitter). These similarities suggest a
shared ancestry.
Rhabdomeric Photoreceptors (in protosomes)
Found mainly in the compund eyes of arthropods.
Ciliary Photoreceptors (in deuterosomes)
Common in vertebrates.

Rhabdomeric Photoreceptors
Increase their surface area by
throwing up their apical
surfaces into numerous folds
(think Bart Simpsons hair).
Many morphologies for this
receptor exist.

Ciliary Photoreceptor
Increases membrane surface area by
modifying the cilium. The ciliary
membrane is expanded and thrown into
deep folds, so that the actual receptor
region of the cell looks like a stack of
discs.

Evidence of Common Receptor Ancestry

Rods and Cones seem to have evolved from common ciliary photoreceptor
precursors, while retinal ganglion, amacrine and horizontal cells seem to have
evolved from a rhabdomeric precursor.

How Could Eyes Evolve?

Creationists often use the eye as a debate point
to show that evolution is flawed, citing that the
eye is too complex and perfect to have evolved.
Darwin himself noted that To suppose that the
eye could have been formed by natural
selection, seems, I freely confess, absurd in the
highest possible degree, (The Origin of
Species). He follows, however, with the
assertion that eyes could likely evolve from light
sensitive neurons. This seems to be the case.

The Evolution of the Eye

Research by Dan-E Nilsson and Susanne Pelger
indicates that it is in fact easier to estimate the
number of generations necessary to evolve an
eye than complex organs. This is because these
changes can be viewed as quantitative local
modifications to a pre-existing tissue.
In order to determine the number of generations
needed to evolve an eye, Nilsson simply made
calculations outlining the plausible sequence of
alterations leading from a light sensitive spot to
a fully developed lens eye.

The Evolution of the Eye

Nilsson assumed an organism with a light sensitive
patch of cells resting on a dark pigmented background
and placed in a selection for spatial recognition. The
first method to create a spatial recognition is either for a
depression to form in the center of the patch, or for the
edges of the patch to constrict and raise. This cupping
would allow for the vague correlation of light to position
where the exposure of an area on the patch is
dependent on its angle to the light source.

The Evolution of the Eye

This cupping evolution should
first favor the formation of a
depression in the patch, than
the constriction of an aperture
via the raising and constriction
of the surrounding pigment
epithelium. This results in a
sunken eye cup that resembles
that of some mollusks.

The Evolution of the Eye

This pinhole-like eye is not very
good at resolving detail and
creates a very dim image.
Because of this, any change that
improves clarity and illumination
will be favored. The two routes of
change for this would be the
development of a lens, or the
increase in the size of the eye.
Increasing the size of the eye,
however, presents physical
problems and less acute vision
than a lens would.

The Evolution of the Eye

Nillson gauged number of 1%
changes in structures in this
diagram. The number of 1%
steps comes out to 1829
necessary steps to progress
from a light sensitive disk to a
camera-eye.
But how many generations
will that take? Prepare for
math on the next slide.
I am so, so sorry.

The Evolution of the Eye

R=h2iVm

m=mean increase/decrease in a feature.

h2=heritability.
i=intensity of selection.
V=coefficient of variation=ratio between standard deviation and mean in a
population
n=number of generations.

h2=0.5 (common heritability), i and V both = .01 (low values for conservative
estimation), therefore

R=.00005m, so small variation and weak selection produce

a .005% change per generation. So
1.00005n=80129540 so n=363992 generations.

What the Heck was that?

So basically, it takes 363992 generations, roughly
364,000 years to evolve camera-type eyes given that
reproduction occurs yearly and the brain of the animal
can handle such visual processing.
Things to note:
Nilssons simulation does not take into account more
specialized structures such as sclera and capillaries
because they are not necessary for all types of camera
eyes (gastropod mollusks lack these). This simulation
also does not take into account the evolution of
photoreceptors.

Are We Done Yet? NO!

Sit down and shut up!
Nilssons simulation demonstrated basic
structural evolution, but what about
genetic evidence? Did eyes evolve
independently or is there one common
ancestor for all eyes?
Get ready for some crazy, messed up,
stuff.

PAX-6 and Aniridia

Prior to 1993 all evidence pointed to independent
evolution of the eye. Then, while looking for transcription
factors in fruit flies Walter Gehring and Rebecca Quiring
discovered a gene nearly identical to the PAX-6 gene in
mice and Aniridia in humans. All of which control the
expression of eyes in a major way.
Mutations in these analogues can truncate the
development of eyes in mice and cause serious defects
in the human eye.
Could this be evidence against independent evolution of
the eye? Asked Gehring. In order to find out he
created

Flytato: All the Eyes of a Potato, All

the Deliciousness of a Fly
By turning on this gene,
dubbed eyeless in
developing cells that do
not normally express it, it
caused the fly to develop

EXTRA EYES IN
ODD PLACES. AS
DID THE
ADDITION OF THE
PAX-6 AND
ANIDIRIA GENES.

WTF!?!?!

Flytato: Continued
Extra eyes ARE light sensitive, ARE NOT wired into the
brain like normal eyes.
Is this evidence for a single origin of the eye? MAYBE.
Ernst Mayr contests many eyeless organisms have
similar genes.
Mayr believes that this gene was originally part of a
group of genes that shape the nervous system. As
different organisms evolved, its role shifted.
PAX-6 also regulates expression of the nose in mice and
the production of tentacles in naughty children squid.

Summary
There are two basic types of eyes, the Simple and Compound eyes.
Simple eyes include the Pinhole Eye, the Concave Mirror Eye and
the Positive Lens Eye.
Compound eyes are composed of multiple Ommatidia and have
Apposition types and Superposition types.
Two types of photoreceptors are believed to have evolved from a
proto-receptor, Rhabdomeric and Ciliary.
Nillson demonstrated how the structure of the eye could evolve from
a light sensitive region to a Camera Eye structure in less than half a
million years.
PAX-6, Aniridia and eyeless are relatively analogues genes that
control the expression of eyes.
Flytato: WTF?

Bibliography

Barnett, Adrian. Evolution for Creationists. 14 Mar. 1999.

<http://www.abarnett.demon.co.uk/atheism/evolution.html#EYES>.

Evolution Library. 2001. <http://www.pbs.org/wgbh/evolution/library/01/1/l_011_01.html>.

Fernald, Russel D. Karger Gazette. Stanford University.

<http://www.karger.com/gazette/64/fernald/art_1_0.htm>.

Land, M F., and Dan E. Nilsson. Animal Eyes. 2nd ed. N.p.: Oxford UP, 2004.

Meyers, Paul Z. Pharyngula. 09 Nov. 2004. University of Minnesota, Morris.

<http://pharyngula.org/index/weblog/comments/rhabdomeric_and_ciliary_eyes/>.

Nillson, Dan E. "A Pessemistic Estimate of the Time Required for an Eye to Evolve." Biological
Sciences 256 (1994): 53-58.

Semo, Ma'ayan. Ma'ayan's Vanity Website. 09 Feb. 1998.

<http://www.maayan.uk.com/evoeyes1.html>.

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<http://www.sciencenews.org/pages/sn_arc97/5_10_97/bob1.htm>.