The Evolutionary Theory of
Sex:
Mechanisms of Regulation of the Population Parameters

Since there are many different
environmental factors (temperature, pressure, humidity,
amount of food, numbers of enemies and pests etc.) it is
natural to think that evolution could not combine each
environmental factor directly with the population parameters
by means of independent mechanism. Evolution must have
produced some “generalizing lever” by means of which any
environmental factor could affect the population parameters.
A definite range of values exists for
each factor which corresponds to comfort conditions.
Discomfort and elimination zones adjoin the comfort one on
both sides. Similar zones can be distinguished in the
population habitat where in a stable environment the comfort
conditions are more often in the habitat center, the
discomfort ones—in the periphery; and the elimination zones
correspond to the territories outside of the habitat.
The control mechanisms of population
parameters are switched on as a result of ecological
information received by the organisms in the discomfort
zones. They get the information because of their contact
with the front of the environment factor. Particular nature
of the environmental factor which causes the discomfort of
the organism seems to have no significance for starting up
these mechanisms. The cause of the discomfort (frost, dry
periods, famine or enemies) makes no difference. The
“generalized” ecological information has one dimension only
(“good” or “bad”) and its cause is unimportant.
Stress as a
transmitter of ecological information between the animals
Such a “generalizing lever”, a
non-specific factor which transmits ecological information
from the environment to population in animals is the
mechanism realized through the stress. Animals
actively “move and communicate” with each other (fight, look
after etc.) and thus receive the ecological information. For
example, if a population of animals luck females, males more
often need to fight for the female, or experience a sex
starvation longer.
All the unfavorable environmental
factors regardless of their specific nature bring about
stress. Stress, initiated as a result of discomfort
conditions, transforms ecological information into the
physiological one, which is coded by concentrations of
various hormones in the organism. Further control in the
organism is performed by the hormones.
Consequently, frequent stresses in
animals and man must enlarge phenotypic (and, probably
genetic) variation of the offspring, increase the quota
of male offspring and produce more pronounced sex dimorphism
in the progeny. The data of demographic statistics show that
during long wars and other social or climatic shifts
(famine, people migrations etc.) the secondary sex ratio
grows statistically significant.
Chronic stress can create steady
changes of hormonal balance which may lead to sharp raise of
the form modification process. This can be due to modifying
influence of hormones on regulatory processes in
Ontogeny, and it is also possible, by virtue of a mutagen
role of hormones.
Sex Hormones
It is well-known that male and female
sex hormones androgenes (An) and estrogenes
(Es), which are chemical antagonists, are
developed and are present in both sexes, but in different
proportions. Androgen concentration in man is approximately
hundred times higher, and estrogen—hundred times higher at
women.
Sex hormones form phenotypical
realization of sex, therefore they should determine the
relation of an organism and environment in Ontogeny.
Androgene/estrogene ratio in the body regulates its
information contact with the environment. This leads to a
new interpretation of androgens as an environmental
hormones “bringing the system closer” to the environment,
while their antagonists, estrogens, insulate or protect the
system from the environment.
By analogy to a sex ratio (M/F),
ratio An/Es is a regulator of a “distance” from the
environment, and evolutionary plasticity as well. The
evolutionary plasticity should decrease in optimum
environment, and increase in extreme one.
Previously described phenomenon of
the increased male mortality represents that total price
which the male sex has to pay to the environment for the new
information. This price develops of many components: at
animals brighter, appreciable coloring and plumage, more
risky behavior, in humans—a choice of dangerous
professions, the higher susceptibility to “new” illnesses
(the illnesses of a “century” or “civilization”) and to
social defects (smoking, alcoholism, drug addiction, crime
and gambling).
Pollen as a
carrier of ecological information in plants
What is the plant's source of
ecological information? How the plants “know” about
ecological situation? Of course, plants can get information
about temperature or humidity directly. But how, for example
they can get information about sex ratio changes?
The application of the ideas which
have been developed to the plant populations, the
distinctive trait of which is an immobile form of life, an
attachment to a place, makes it possible to disclose certain
new regularities. When analyzing the habitat of a plant
species it is clear that in the average more optimal comfort
conditions are in the center of the habitat (in its depth),
while more the extreme, the discomfort ones exist in its
periphery (on its borders). The existence of the habitat
boundary itself indicates that in its natural expansion, the
species came upon a “wall” of the ecological niche at this
place because of one environmental factor or another. Such
factors may be either low or high temperature, or various
concentrations of moisture, nutritive substances, enemies,
parasites, etc. It is evident that in the center of the area
the density of the population is maximal, and at the
periphery, minimal.
In realizing the informational
connection between plants, the principal role may belong to
the amount of pollen. In dioecious or
cross-pollinated species in otherwise equal circumstances,
the falling of a large amount of pollen on a female plant
means that there are many male plants in the area.
Conversely, a small amount of pollen means that there are
few male plants in the vicinity. In the center the amount of
pollen getting on female flower is always on the average
greater than that on the periphery due to different
population density. Male plants are first who die from the
extreme environmental conditions. This also decreases pollen
count.
The pollen amount gives information
to the female flower about population density, tertiary sex
ratio around it, about its location in the habitat center or
periphery, and about optimal or extreme environmental
conditions. Reception of the great amount of pollen always
carries on the information about favorable environmental
conditions and requires increased production of females with
small phenotypic variation. On the contrary, the
reception of small amount of pollen carries on the
information about unfavorable conditions. This can occur
either at the periphery where the density of the population
drops sharply, or at the center, but when extreme conditions
occur at the center, they eliminate the male individuals
first. Both require higher production of male offspring with
higher phenotypic variation to speed up the search of
evolutionary pathways (leading selection).
So, in any optimal environmental
conditions female flower receives more pollen than in
extreme conditions. In other words, small amount of pollen
always mean “bad”, and big amount of pollen always mean
“good”.
Consequently, the statement of
classical genetics should be reconsidered, that pollen
carries only genetic information and the pollen amount which
gets on the female flower is of no importance, since one
pollen grain is sufficient for fertilization. The pollen
transmits not only genetic information, but also the
ecological one, which controls the ratio between stabilizing
and leading selection. The information is coded by the
pollen amount.
Characteristics of the
Habitat
Center |
← |
Location |
→ |
Periphery |
|
|
↓ |
|
|
Optimal |
← |
Environmental
conditions |
→ |
Extreme |
|
|
↓ |
|
|
Max |
← |
Density of
population |
→ |
Min |
|
|
↓ |
|
|
Max |
← |
Pollen amount |
→ |
Min |
|
|
↓ |
|
|
Min |
← |
Pollen of other
species |
→ |
Max |
|
|
↓ |
|
|
Min |
← |
Dispersion of offspring
Sex Ratio
Sexual Dimorphism |
→ |
Max |
|
|
↓ |
|
|
Min |
← |
Evolutionary
plasticity |
→ |
Max |
Back
to
Receiving the Ecological Information from the Environment
More about
Pollen Amount:
The Amount of Pollen as a Regulator of Evolutionary
Plasticity of Cross-Pollinating Plants.
Geodakyan V. A. Doklady Biological Sciences, 1977, v.
234, N 1-6, p. 193–196. Translated from Doklady Akademii
Nauk, Vol. 234, No. 6, pp. 1460-1463, May, 1977.
|