ISOLATING MECHANISMS

The term isolating mechanisms was introduced by T Dobzhansky in the 1930s, and has been
popularized in a number of books by E Mayr. Both authors originally proposed that isolating
mechanisms were group traits beneficial at the level of the species; today, this is generally
disbelieved. Recent authors have pointed out that the word "mechanism" is particularly
misleading as pre-mating and post-mating isolation are likely to evolve as a by-product of
natural selection or genetic drift within species, rather than as a direct result of their utility
as barriers to fertilization and gene mixing between species (a process known
as reinforcement). A leading critic of the biological species concept and of the term isolating
mechanisms is HEH Paterson, who argues that species are cohesive wholes as a result
of pre-zygotic sexual signalling within species, rather than due to isolating mechanisms
between species. Paterson therefore introduced a competing idea of species,
the recognition concept of species, in which isolating mechanisms were replaced by specific
mate recognition systems as an alternative. Unfortunately, the word "system" has as many
group-benefit connotations as "mechanism", and the recognition concept of species has not
gained universal acceptance.

There is also the terminological problem that reproductive isolation combines traits that
reduce gene flow, such as mate choice or fertilization barriers, with traits that select against
genes that have flowed, such as hybrid incompatibility. Lumping these two antagonistic
features is confusing, since they are unrelated and evolve in very different ways. For
instance, whereas it is conceivable that reinforcement might evolve to reduce an individual's
tendency to mate with another species and produce inviable offspring, it is almost
impossible to imagine that hybrid inviability itself would evolve as an adaptation.
This reproductive isolation terminology leads also to a muddled use of the term gene flow as
the opposite of reproductive isolation; in other words, gene flow comes to include not only
the flow of genes, but also the effects of any natural selection on the frequency of such
genes within each population.

Perhaps the most fundamental problem with isolating mechanisms (and specific mate
recognition systems) is that species are implied to be qualitatively different from subspecies,
races, or forms by their possession of these traits. Races cannot, in theory, differ in either
type of trait because only species are defined by their possession. Arguably, by making
species seem qualitatively different from races, these terms have spawned a number of
special models of speciation where geographic isolation, also known as allopatry, or sudden
bursts of evolution in small founder populations (founder events or punctuated equilibria)
play important roles. Only such unusual conditions were thought to be able to give rise to
new species that differ in isolating mechanisms (or specific mate recognition systems). In
reality, there is little to distinguish mate choice and disruptive natural selection commonly
observed within species from pre-mating and post-mating isolation between species; and,
indeed, it is hard to distinguish species from races in many actual organisms.

Isolation, in evolutionary term, means segregation of different populations into smaller

units by certain mechanism so as to prevent interbreeding among individuals.
1. Geographical isolation
When the populations are separated by a geographical barrier, such as river, sea,
mountain, deserts and for aquatic animals land, they are physically prevented from
interbreeding. Such populations are termed as allopatric and are forced to evolve
independently and accumulate genetic differences. Geographical isolation may be
different for different species. For example, a small stream may be an effective barrier for
land insects and small mammals while for birds even mountain and oceans may not be
barriers.

2. Reproductive isolation
It is the property of individuals that prevents interbreeding in populations that are
actually sympatric (living in the same area).

Classification of reproductive isolating mechanisms

A. Premating mechanisms
They prevent interspecific crosses in sympatric populations.
1. Seasonal isolation: Also called temporal isolation, in which potential mates do not
come in contact with each other because of differences in breeding seasons of two
species, e.g. different flowering seasons in plants. Bufo americanus breeds in early rainy
season (May), while Bufo fowleri breeds in late rainy season (July) in USA.

2. Habitat isolation: Also called Ecological isolation, in which also potential mates do not
meet each other due to differences in habitats, requirements of food, space, climate etc.
Potential mates live in different areas and therefore do not come in contact with one
another. For example spawning grounds of riverine fishes are in different tributaries,
which prevents interbreeding.

3. Ethological isolation: It is a behavioral isolation, in which potential mates meet but

cannot mate, due to differences in courtship displays or other specific signals that are
necessary rituals before mating. The signals may be of the following three types, which
stimulate the opposite sex for mating:
(a) Visual stimuli: Feather displays and dancing in male birds is necessary to
attract the female, e.g. peacocks, pheasants and birds of paradise. The colour and shape
of the feathers as well as display pattern is so unique for each species that mating
between two different species is not possible. Collection of the nest material and
construction of the nest as by the weaverbird male is also a very specific display that
cannot be imitated by the other species.
(b) Auditory stimuli: Songbirds like cuckoos, mynas, nightingales, parakeets etc.
use auditory signals to attract the opposite sex. Sometimes the singing goes on for
several days before the pair can actually come together for mating. Auditory
communication is used by a large number of animals, viz. frogs, toads, cicadas, gibbons ,
monkeys, jackals etc.
(c) Chemical stimuli: This includes odors of the animals that attract the opposite
sex for mating. For example scent of musk deer and musthing in elephants attract the
females. In insects, particularly Lepidoptera, females produce highly specific pheromones
that can be detected by the highly specialized antennae of males from a distance of
about 2 kilometers.

4. Mechanical isolation: In this case the above isolating mechanisms are not present and
therefore mating is attempted but is not successful due to mechanical problems such as
differences in the structure of genitalia. Dufour (1844) described “Lock and key
mechanism” in the genitalia of insects. In the species of Drosophila the genitalia are so
different that copulation is mechanically not possible.

B. Postmating mechanisms
These reduce the success of interspecific crosses. In case premating mechanisms fail to
prevent mating then several postmating mechanisms prevent the success of mating and
hybridization. There are 4 such mechanisms, which are given below:

1. Gamete mortality: Mating and sperm transfer takes place but egg is not fertilized.
In Drosophila vaginal wall swells killing spermatozoa should interspecific crosses take
place. If mating takes place between Bufo fowleri and Bufo valliceps, sperms cannot
penetrate the egg membrane of each other, leading to mortality of gametes.

2. Zygote mortality: Egg is fertilized but the zygote dies. Eggs of many species of fishes
may be present in the spawning grounds and some may be fertilized by the sperms of
different species forming zygote but such zygotes fail to develop due to differences in
chromosomes.

3. Zygote inviability: Zygote develops and hybrid is produced but is physically weak and
inviable due to physiological disturbances in the body. It fails to survive for long and
prematurely dies. Such cases have been recorded in different species of ducks.

4. Hybrid sterility: Hybrid is viable, physically strong and physiologically sound but is
sterile due to differences in chromosomes and different gene arrangements. Mule is a
cross between male donkey and female horse and Hinny between female donkey and
male horse and both are sterile, albeit physically strong.
Sometimes all isolating mechanisms break, leading to fertile hybrids, which are generally
not reproductively isolated from the parents and can produce fertile offsprings by
Introgression (hybrids backcrossing with parents to produce fertile offsprings). This will
be instant speciation.
Significance of isolating mechanism
 Wasteful courtship is avoided. If isolating mechanisms are distinct and specific only
individuals of the same species indulge in courtship.
 Isolating mechanism protects gene pool of a species and prevents hybridization.
 It prevents wastage of gametes and energy.
 A weak isolating mechanism leads to production of new species through
hybridization.
 Absence of isolating mechanism leads to production of new species by instant
speciation.
 Geographical isolation followed by reproductive isolation ultimately leads to
production of new species.
 Isolating mechanisms protect the identity of a species, which all species fiercely
guard.