Evolution of Sight and Mimicry

Contributed by: Chen Raymundo

Evolution of SIGHT
Eyes are the organs of vision. They detect light and convert it into electro-
chemical impulses in neurons. The simplest photoreceptor cells in conscious
vision connect light to movement.
Higher organisms: eye is a complex optical system which collects light from the
surrounding environment, regulates its intensity through a diaphragm, focuses it
through an adjustable assembly of lenses to form an image, converts this image
into a set of electrical signals, and transmits these signals to the brain through
complex neural pathways that connect the eye via the optic nerve to the visual
cortex and other areas of the brain.
Eyes with resolving power have come in ten fundamentally different forms, and
96% of animal species possess a complex optical system. Image-resolving eyes
are present in molluscs, chordates and arthropods.
The simplest eyes, i.e. microorganisms, do nothing but detect whether the
surroundings are light or dark, which is sufficient for the entrainment of circadian
rhythms. From more complex eyes, retinal photosensitive ganglion cells send
signals along the retinohypothalamic tract to the suprachiasmatic nuclei to effect
circadian adjustment.
Complex eyes can distinguish shapes and colours. The visual fields of many
organisms, especially predators, involve large areas of binocular vision to
improve depth perception. In other organisms, eyes are located so as to
maximise the field of view, such as in rabbits and horses, which have monocular
vision.
The first proto-eyes evolved among animals 600 million years ago about the time
of the Cambrian explosion. The last common ancestor of animals possessed the
biochemical toolkit necessary for vision, and more advanced eyes have evolved
in 96% of animal species in six of the ~35 main phyla.
In most vertebrates and some molluscs, the eye works by allowing light to enter
and project onto a light-sensitive panel of cells, known as the retina, at the rear of
the eye. The cone cells (for colour) and the rod cells (for low-light contrasts) in
the retina detect and convert light into neural signals for vision. The visual signals
are then transmitted to the brain via the optic nerve. Such eyes are typically
roughly spherical, filled with a transparent gel-like substance called the vitreous
humour, with a focusing lens and often an iris; the relaxing or tightening of the
muscles around the iris change the size of the pupil, thereby regulating the
 amount of light that enters the eye, and reducing aberrations when there is
enough light.
The eyes of most cephalopods, fish, amphibians and snakes have fixed lens
shapes, and focusing vision is achieved by telescoping the lenssimilar to how a
camera focuses.
Compound eyes are found among the arthropods and are composed of many
simple facets which, depending on the details of anatomy, may give either a
single pixelated image or multiple images, per eye. Each sensor has its own lens
and photosensitive cell(s). Some eyes have up to 28,000 such sensors, which
are arranged hexagonally, and which can give a full 360 field of vision.
Compound eyes are very sensitive to motion. Some arthropods, including many
Strepsiptera, have compound eyes of only a few facets, each with a retina
capable of creating an image, creating vision. With each eye viewing a different
thing, a fused image from all the eyes is produced in the brain, providing very
different, high-resolution images.
Possessing detailed hyperspectral colour vision, the Mantis shrimp has been
reported to have the worlds most complex colour vision system. Trilobites, which
are now extinct, had unique compound eyes. They used clear calcite crystals to
form the lenses of their eyes. In this, they differ from most other arthropods,
which have soft eyes. The number of lenses in such an eye varied, however:
some trilobites had only one, and some had thousands of lenses in one eye.
In contrast to compound eyes, simple eyes are those that have a single lens. For
example, jumping spiders have a large pair of simple eyes with a narrow field of
view, supported by an array of other, smaller eyes for peripheral vision. Some
insect larvae, like caterpillars, have a different type of simple eye (stemmata)
which gives a rough image. Some of the simplest eyes, called ocelli, can be
found in animals like some of the snails, which cannot actually see in the
normal sense. They do have photosensitive cells, but no lens and no other
means of projecting an image onto these cells. They can distinguish between
light and dark, but no more. This enables snails to keep out of direct sunlight. In
organisms dwelling near deep-sea vents, compound eyes have been secondarily
simplified and adapted to spot the infra-red light produced by the hot ventsin
this way the bearers can spot hot springs and avoid being boiled alive.

EVOLUTION OF MIMICRY

The effectiveness of warning systems

There is considerable experimental evidence to illustrate how effectively

predators learn to avoid certain adverse stimuli. Chickens conditioned by
electric shock to avoid drinking dark green water drank progressively more
 from paler solutions in proportion to the intensity of the colour. This
experiment suggests that even an incomplete warning system provides
a modicum of protection. The degree of protection provided is also affected by
the strength of the punishment; after strong shocks the chickens drank only
from very light coloured solutions. In the presence of severe punishment, an
improved warning system made little additional effect once a threshold level
was reached.
In other experiments, starlings (Sturnus vulgaris) were fed normal mealworms,
two segments of which had been painted orange. To provide aposematic
models, the experimenter made other mealworms distasteful and painted the
same segments green. Mimics were marked with green but not rendered
unpalatable. There is no known instance in nature in which animals employ
green for warning; there was therefore no possibility that the birds had already
learned to avoid the experimental colour pattern. Before long the green-marked
worms were completely avoided, regardless of palatability, even when the ratio
of edible to distasteful was 60:40. This indicates that the number of mimics can
exceed that of the model, when the resemblance is close, without loss of
protection. When the ratio was increased to 90:10, 17 percent of the mimics
were avoided, probably sufficient to a selective advantage in nature. Although a
test bird would occasionally peck at a model, then reject it, the same action was
sometimes shown to a mimic that it had picked up, suggesting that a premature
response had been subsequently corrected.

The reconstruction of evolutionary pathways

Analysis and understanding of a given mimicry system require a
rather comprehensive knowledge of morphology, behaviour, ecology, and
mutual relationships of animals usually in different classesfor example,
wasps (Hymenoptera), flies (Diptera), insect-eating amphibians, reptiles, birds,
and small mammals. Tracing the evolution of such a complicated system
requires a detailed acquaintance with a large group of forms related to each of
the animals involved.

Reconstructing the evolution of a case of mimicry within the same species is

valuable, because mimicry is an indispensable tool in the study of the evolution
of animal communication, and usually starts from conspicuously elaborated
signals, which postulate a signal receiver interested in them. The receiver
practically always has undergone a special molding toward optimal receiving of
the signal. The mutual adaptations of the sender and the receiver must be
examined separately.
 This examination is easily made, so far as the evolution of a reaction or of a
receiving mechanism is concerned, in all predators trying to find their prey and
in all prey animals attempting to escape an approaching predator. The
suppression of signals may be studied in predators trying to sneak up on a prey
unnoticed. The elaboration of a signal, which must, of course, be important to
the receiver, can only be studied after consideration of compensatory
adaptations in the receiver and in situations where the sender has a one-sided
interest in the signal. The deceiving signal can be derived only from one of two
types: a signal developed by the receiver and another signal sender in their
common interest or a signal emitted by another signal sender and made use of
by the receiver only in its own interest. Both cases, by the definition given
above, are called mimicry. An additional advantage is that the model is known
to be the final stage toward which the mimic will evolve (so far as the signal
characters are concerned), thus indicating a trend in evolution that is still
operating and that probably over time will further elaborate the mimetic
signals.

If the female Haplochromis fish were to discriminate between real eggs and the
egg dummies of the male and were to stop reacting toward the latter, her eggs
would remain unfertilized. In such cases of deceptive signals developed within
the same species, natural selection operates against better
signal discrimination on the part of the signal receiver.

The importance of the signal receiver

Fundamental characteristics of mimicry are determined mainly by behavioral
properties of the signal receiver. A precise knowledge of the identity of the
receiver and a thorough study of its behaviour are therefore indispensable for
the understanding of mimicry.

Mimicry gradually merges into other senderreceiver systems. Palatability is a

matter of degree; whole ranges of distastefulness therefore exist, even in the
mimics, model and mimic in the case of Mllerian mimicry being equally
unpalatable and sharing the same warning coloration. Mllerian mimicry could
be considered not to be true mimicry, after all, because no one is deceived, and
it is impossible to designate one as model and the other as mimic.

Individuals of a given wasp species look alike and are all equally protected, this
phenomenon is not usually called Mllerian mimicry, simply because the
 signals were not independently evolved, a property known as convergence.
Because, however, the male wasps have no protective properties but retain their
group-specific warning coloration, this is Batesian mimicry, although model
and mimic are of the same species and their signals homologous (evolved from
the same source)

Convergence (or independent evolution) of the signal characters, therefore, is

essential only for the so-called Mllerian mimicry, and thus Mllerian mimicry
is distinguished from other cases of signal standardization. The typical
(Batesian) mimicry merges into Mllerian mimicry if the difference between
the consequences for the receiver of reacting similarly to model and mimic
diminishes; and by homology of the signal characters it further merges into
general signal standardization.

An insect may be protectively coloured to resemble, for example, a wasp or a

twig. In the first case the coloration is called mimicry, in the second, mimesis,
or protective coloration. The difference lies within the signal receiver. If the
mimetic signal does not release any reaction in the receiver, the mimic is said to
exhibit mimesis. This distinction is illustrated by the experiments of the Dutch
biologist L. de Ruiter with stick caterpillars, which, by virtue of their close
resemblance to twigs, are protected against insect-eating birds. As soon as the
number of twigs becomes too large, however, the bird develops an interest in
them, attacks some real twigs, and also finds some caterpillars. If one positive
experience with the caterpillar has the same weight as a negative one with the
twig (the signal remaining unchanged), the relative abundance of caterpillars
and twigs determines whether all twigs are mistakenly exterminated or whether
the feeding reaction toward twiglike objects disappears, thus protecting the
caterpillars.

The birds ability to decide correctly which is the model and further shows how
easily an object (the twig) may quite involuntarily become a mimic.

Another example illustrating the importance of a correct model is found in the

common farming relationship between ants and aphids. The protuberances,
called the siphones and cauda, on the abdomens of aphids resemble
respectively the bases of the antennae and labium of the ants head. The aphids
abdomen is thus mistaken by the ant for the head of a fellow ant, thereby
eliciting the food-begging response, which is identical with milking. Saturated
ants in turn even try to feed the abdomens of the aphids. Aphid species with
 reduced abdominal siphones use their hind legs as antennae dummies, the
movements elicited being originally defensive movements. This situation is
exactly the way in which mimicry arises. Mimetic characters need not have
evolved under the selection pressure of mimicking; in fact, their earliest
evolutionary stages could not even have been brought about in this way. All
cases studied thus far can be traced back to an incipient stage of deceptive
resemblance, initiated as a preadaptive, nondirected by-product of pre-existing
species-specific features, thus providing a point of attack for new selective
pressure.

The effects of selective pressure

The selective consequences for the signal receiver of responding to the model
are always positive (the reaction would disappear if, on balance, it were
unfavourable to the receiver). The mimic always has a selective advantage in
releasing the reaction from the receiver. An unfavourable signal by the mimic
would also disappear by natural selection.

The selective consequence for the model eliciting and obtaining the reaction
from the receiver may be of several types. Consequences may be absent, if the
model is an inanimate object on which natural selection does not act. They may
be negative, if the model is non-aposematic (non-warning), such as the tiny
crustacean, usually eaten by the signal receiver and mimicked by the
male Corynopoma characin in order to attract the female. Or they may instead
be positive, as in the wasp, which remains alive if it is avoided by the predator;
in the cleaner, which feeds on parasites harmful to other fish; or in those
hymenopteran females whose male-attracting signals are mimicked by certain
orchids. Mutual interest is present between model and receiver in cases of
aggressive mimicry where both parties belong to the same particular species
and also in typical Batesian mimicry.

Constant learning by the signal receiver results in a strong selective pressure on

the mimic against detectable differences from the model, but at the same time it
also exerts a complementary strong pressure on the model to develop just such
new differences from the mimic. Typically, it is the group of songbirds
parasitized by cuckoos that has developed the most divergent egg-colour
patterns; the group of estrildine finches parasitized by whydahs that has
developed particular gape patterns; and among the cleaner wrasses the
species Labroides dimidiatus mimicked by the blenny Aspidontus that develops
into many different local races.
 There is a boomerang effect, characteristic of the parasitehost relationship,
that the more successfully a bird rears young cuckoos, the more certain it is that
it will lose its own young, because they are killed off by the young cuckoo.
Parasites that are too successful, therefore, harm themselves, for each female
cuckoo needs several nests of the same host species for her eggs. In an area that
contains particularly successful cuckoos, the number of reed warbler nests has
been found to decrease from year to year, while the percentage of nests
parasitized by cuckoos increases from year to year. This ratio means that a
cuckoo that is too well adapted reduces the availability of its own hosts, while
one insufficiently adapted kills off its own offspring. Presumably, selection in
both directions produces a continual oscillation in the densities of hosts and
cuckoos.

A similar dilemma is inflicted on human beings, who act as predators

against weeds in crop fields and by winnowing select the wanted seeds from
the usually smaller weed seeds. The flax dodder (Cuscuta epilinum), for
example, which grows as a creeper around flax and linseed plants and damages
them, originally had small seeds that could be easily separated from the larger
flax seeds. By a mutation that produced twin seeds, the dodder has evolved the
capability of being separated out and planted with the desirable flax seeds. This
mutant of the flax dodder is now cultivated and spread by growers, despite
being against their interests. In this case, the parasite mimics the protected
plant, receiving the same protection.

The effects of geographic distribution and population density

It has been postulated that the model and the mimic should always occur in the
same areai.e., be sympatric. They need not always be sympatric, however,
but must always have a signal receiver in common: a model might be in Africa,
for instance, and its mimic might be in Europe (or vice versa), functionally
connected by a migratory bird.

Another postulate, that mimics must naturally be less numerous than their
models, means, correctly stated, that the receiver has to meet the mimic less
often than the model; this postulate is based on the assumption that one
experience with the model has the same aftereffect, the same weight, as has one
with the mimic. This assumption, however, has been proved not always to be
so; in fact, the negative experience seems usually to be the stronger one. This
negative experience may result from an encounter with the model (such as a
 wasp) or with the mimic (for example, the sabre-toothed blenny). There might
be more wasp mimics than wasps, but in cases such as that of the cleaner
wrasse mimic, the mimic probably has to be less numerous than the model. The
protective power of the model, of course, is reduced with an increasing number
of mimics, because the predator may eat larger numbers of them before his first
encounter with the model.

The importance of mimicry to evolutionary theory

The mimicry hypothesis emerged in the middle of the Darwinian controversy

and provided an ideal test case for the views of Charles Darwin and his
contemporary Alfred Russel Wallace on the operation of natural selection in the
evolutionary change of living organisms. It is now quite evident that the basic
theory of natural selection is correct and that the theory is strengthened by
many detailed studies of the process by which a mimetic resemblance is
brought about and selected for. In addition, investigating suitable cases of
mimicry provides important insight into the evolution of signals and the
semantization process by which signals get their meaning.

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