WEST VIRGINIA UNIVERSITY AT
PARKERSBURG
BIOLOGY/GEOLOGY 397
INTRODUCTION TO ORGANIC EVOLUTION
CO-EVOLUTION AND THE
EVOLUTIONARY ARMS RACE
INTRODUCTION
Co-evolution
is concerned with the joint evolution of two or more species resulting from
their ecological interaction (Kardong, 2005).
Organisms
must continually change their genetic make-up (to adapt to a changing
environment) if they are going to survive and
reproduce. The competition for resources is fierce. Although
organisms certainly must constantly adapt to a changing physical environment,
uncommonly is the physical environment the reason for death or the cause of
non-reproduction. Most often it is other organisms (parasites, predators,
and competitors) that prevent survival and/or reproductive success (Ridley,
1993). Van Valen (1973) referred to the concept (he considered it to be a
new evolutionary law) of the need for continual change of the genome of an
organism , as the "Red Queen Hypothesis".
The Red Queen is a reference to the Red Queen chess piece in Alice Through
the Looking Glass (by Lewis Carroll, 1872), who told Alice that here (in
this land) ".....it takes all the running you can do, to keep in the
same place." (Prothero, 2004; Ridley, 1993). So, if species are
going to avoid extinction, they must constantly improve their survival and
reproductive fitness (Prothero, 2004). Thus, an evolutionary
arms race can co-evolve between a predator and its prey, parasites
and their host, or competitors for the same resources. For example, a newt
develops a toxin within its flesh so that predators will avoid it.
However, natural selection will favor the variants of the predator that are more
resistant to the toxin and that can eat the newt and withstand the
toxin. So, the newt becomes more toxic to the predator. The predator
evolves more resistance to the toxin, and so on. There are many examples
in nature of this evolutionary arms race.
Also,
according to the Red Queen Hypothesis,
sexual reproduction persists because it enables many species to rapidly evolve
new genetic defenses against parasites that attempt to live off them. For
example, if a certain bacterium species infects humans, some people are more
resistant to the bacteria than others. The bacterium species may kill
large numbers of people, but those exposed that were not killed have a higher
degree of immunity to the effects of the bacteria, however, they still may get
sick. But their defense mechanisms (like white blood cells) were better
able to kill the bacteria that invaded the body. However, the variants of
the bacterium species that were not killed by the body defenses of the more
resistant humans, multiplied and became even more deadly. The humans that
were not killed by the more deadly form of the bacterium species, produced
offspring that inherited their higher degree of resistance to these
bacteria. Any mutations in the bacteria to make them better able to cope
with the defenses of the humans would be selected for. The same is true
for the humans, any mutations or different variants of defense mechanisms that
arose through sexual reproduction (and reshuffling of the genes) that favored
resistance to the bacteria would be favored and passed on to offspring.
Thus, the arms race would go, but in the end neither the bacterium species nor
the human hosts would have gained much ground (a zero sum game).
The following
statements fromWikipedia, the free encyclopedia on the internet at http://en.wikipedia.org/wiki/The_Red_Queen
will help explain the Red Queen Hypothesis:
"The recognition of a simple example of
a biological arms race (from Richard
Dawkins) can be achieved by considering the contrast between two adaptations
of the polar
bear. This animal has a coat of hair which is thick to help the bear survive
the cold of the arctic and white in order that the bear can stalk seals
for food. For the first case the selection pressure is likely to be constant or
subject to random change, in the second case the selection pressure is likely to
increase steadily as selection for cautiousness in seals makes the average seal
harder and harder for the bear to stalk successfully. As a result both the bear
and the seal find themselves running a red queen's race over evolutionary time,
each becoming better and better adapted (to stealth and caution respectively)
but neither becoming any more successful (as they are engaged in a zero
sum game).
Science writer Matt
Ridley wrote a book The Red Queen in which he discussed the debate
in theoretical biology over the adaptive benefit of sexual reproduction to
those species in which it appears. The connection of the Red Queen to this
debate arises from the fact that the traditionally accepted theory (The
Vicar of Bray) only showed adaptive benefit at the level of the
species or group, not at the level of the gene.
By contrast, a Red-Queen-type theory that organisms are running cyclic
arms races with their parasites
can explain the utility of sexual reproduction at the level of the gene by
positing that the role of sex is to preserve genes which are currently
disadvantageous, but which will become advantageous against the background of
a likely future population of parasites.
Sex is an evolutionary puzzle. In most sexual species, males make up half
the population, yet they bear no offspring directly and generally contribute
little to the survival of offspring. In addition, males and females must find
each other to mate, and sexual
selection often favors traits that reduce the survival of organisms. Thus,
sex is highly inefficient.
One possible explanation for the fact that nearly all vertebrates
are sexual is that sex increases the rate at which adaptation
can occur. This is for two reasons. First, if an advantageous mutation
occurs in an asexual line, it is impossible for that mutation to spread
without wiping out all other lines, which may have different advantageous
mutations of their own. Second, it mixes up genes. Some genes might be
advantageous only when paired with other genes, and sex increases the
likelihood that such pairings will occur.
For sex to be advantageous for these reasons requires constant selection
for changing conditions. One factor that might cause this is the constant arms
race between parasites and their hosts. Parasites
generally evolve quickly, due to their short lifespans. As they evolve, they
attack their hosts in a variety of ways. Two consecutive generations might be
faced with very different selective pressures. If this change is rapid enough,
it might explain the persistence of sex.".
SYMBIOSIS
Symbiosis (living together) is a
co-evolutionary relationship between two organisms. The relationship may
be good for both organisms, good for one and neutral for the other, or good for
one at the detriment of the other.
- Mutualism is a symbiotic relationship that is to the benefit of
both partners. Honeybees and angiosperms are illustrative of a
mutualistic symbiotic relationship. The bees visit flowers to eat
nectar and collect pollen to take back to the hive, thus as they move from
one flower to another they carry pollen from one flower to another and
contribute the fertilization process.
- Commensalism is a symbiotic relationship in which one of the
species involved receives a benefit, whereas the other species does not
benefit (or benefits little) but is not harmed by the relationship.
Insect eating birds that follow cattle or other large herbivores is an
example, the birds catch the insects that are stirred-up by the cattle as
they forage, whereas the cattle are not harmed but do not gain anything
either.
- Parasitism, Predation, and Herbivory are symbiotic
relationships in which one species loses and the other gains. Bacteria
that use the human body for room and board and give off harmful toxins is
certainly a good example of parasitism. The polar bear that preys on
arctic seals is but one of thousands of examples of predation, and bison
that graze on grasses is an example of herbivory.
PLANT-ANIMAL HERBIVORY CO-EVOLUTION
- Spines, thorns, bristles and other adaptations of plants to make
themselves less palatable to animals that eat them.
- Secondary chemical compounds.
MUTUALISM
- Ants and aphids. Aphids are the cattle of the
ants. The aphids excrete a sugary liquid ("honeydew") that
the ants ingest. The ants protect the aphids from their predators,
carnivorous insects.
- Ants and fungus. Attini ants (also called leaf-cutter ants)
of Central and South American Tropics culture an eat the fungus Leucocoprini.
The fungus is grown on pulverized plant material and excrement from the
ants. The ants gain a food source. The fungus is cared and
nutured. The ants even carry on their bodies a bacterium that produces
antibiotics that suppresses the growth of a parasitic fungus that can attack
the Leucoprini fungus that the ants grow and harvest.


- Photos from:
http://en.wikipedia.org/wiki/Leafcutter_ant
- Ants and Acacia Trees
- Honey Bees and Flowers
- Hummingbirds and Flowers
- Hermatypic (reef-building) Corals and Zooxanthellae Algae.
The following summary of an article entitled The Role of Symbiotic Algae
in Marine Invertebrates discusses the symbiotic relationship between
reef-building corals and zoozanthellae algae and was taken from the internet
at http://www2.hawaii.edu/~delbeek/delb17.html:
- "In summary, using light energy, zooxanthellae
convert carbon dioxide (from bicarbonate taken from seawater and the
carbon dioxide produced by cellular respiration of the coral tissue)
into carbohydrates and alanine. These products are then passed on to the
animal tissue which subsequently provides a source of nitrogen (ammonia)
and phosphate to the algae. Without the zooxanthellae, the coral host
would soon suffocate in it's own wastes. As was mentioned earlier,
zooxanthellae also contribute to the production of coral skeletons.
Corals which are deprived of their zooxanthellae, or are kept in the
dark, deposit calcium at a much slower rate than normal. It is thought
that algal photosynthesis may increase the calcium carbonate production
by removing carbon dioxide and driving the following reaction to the
right: Ca(HCO3)2 <--> CaCO3 + H2CO3
<--> H2O + CO2 (Barnes, 1974) It is this
ability to rapidly deposit calcium carbonate which has helped the corals
to become the dominant animals on the reef.".
COMMENSALISM
- "Looking-glass" Orchids and Wasps
- Skunk Cabbage and Flies, Scavenging Beetles.
PROTECTIVE COLORATION AND SHAPE
- Camouflage
- Dwarf Seahorse
- Inchworm Caterpillar
- Warning Coloration (Aposematic)
- Poison Arrow Frogs of Tropical South America - brightly colored
to advertise the toxic skin secretions they produce that would be
harmful to predators that would eat them.
- Hide and Advertise - Screech Owl - will try to hide if
approached by tucking itself up against a tree with its eyes closed, but
if threatened by too close an approach, will puff-up, spread its wings
and open its eyes to look more formidable.
- Startle Response - Eyespots on Butterfly Wings
MIMICRY
- Co-evolutionary relationships that result from natural selection of
superficial resemblances between organisms is termed mimicry.
- Batesian Mimicry - The model for this type of mimicry is one
species that is dangerous or distasteful to a predator. The species
that has evolved to look like the model is the mimic, which is not
dangerous or distasteful. Example: The monarch butterfly (the model)
contains noxious chemicals in its tissues. Birds, like the blue jay,
tend to avoid monarch butterflies because of their toxic chemicals.
The mimic, the viceroy butterfly, benefits from looking like the monarch
because birds will avoid eating it also. That is, unless the mimics
become too abundant, then the model may become threatened also. This
is because birds may learn (by some trying to eat the mimic) that the mimic
(the viceroy butterfly) is a tasteful meal. They will then seek out
the bright colored mimics and also the bright colored models (until they
again learn that the monarch is toxic and distasteful). So, it is to
the benefit of the model to evolve such that it does not resemble the
mimic. Thus, again an evolutionary arms race develops.
-

- The Monarch Butterfly. From: http://www.kidzone.ws/animals/monarch_butterfly.htm

- The Viceroy Butterfly. Photo by William T. Hark. From: http://www.nhptv.org/natureworks/viceroy.htm#2
- MULLERIAN MIMICRY
- In Mullerian mimicry, both species have evolved to superficially resemble
each other, but both (or more) species are unpalatable or dangerous to their
predators.
- The predator community is reinforced in its aversion of these species,
because both (or all) are unpalatable.
- The more species that mimic each other in this type of mimicry the better
- the more reinforcement that occurs.
- OTHER TYPES OF MIMICRY
- Agressive mimicry - Examples: Angler fish, cleaner fish and
their aggressive mimics.