Lecture 1 - Descent with Modification

The concept of descent with modification displays the unity of life, and how all organisms are
related in that they have all descended from a common ancestral species. The history of life can
be depicted as a sort of family tree, where the tips are contemporary organisms, branch points
represent ancestors or evolutionary lines (branches are usually extinct) and closely related
organisms share recent branch points.
Why must descent with modification happen? What drives the modification to occur? Descent
with modification is a result of natural selection which occurs when resources are limited and the
given environment necessitates unique adaptations. Adaptations are modifications resulting
over time as descendants inhabit differing environments. Those who can compete for resources
and are well suited for their environments have the reproductive advantage, and therefore the less
adapted genes leave the gene pool because those individuals whom are less adapted do not
reproduce. A gene pool with only genes suited for the given environment is a result of the
cumulative effect of natural selection over extended periods of time.
Natural selection does not occur spontaneously in any given population, however. There are
conditions necessary to bring about natural selection. Overpopulation, which causes limited
resources and therefore competition, and diversity within the gene pool so that there are traits
suitable for the environment that can survive and be passed on, are two conditions necessary for
natural selection. Natural selection happens as a gradual accumulation of small changes, and
only traits carried by gametes are passed on, not any adaptations the individual makes during its
lifetime (learning).
We know that natural selection occurs in populations of species, but what is a population in the
first place? What makes it different from a species? A population is a group of interbreeding
members of a species that share a common geographical area, while a species is a group of
populations whose individuals have the capacity to interbreed and produce fertile of offspring.
A gene pool is an aggregate of all the alleles for all the genes in a population at a given time.
The alleles are diploid and each gene has 2 alleles, and therefore can be homozygous or
heterozygous. Every gene generally has 2 or more types, and each allele type as a relative
frequency in a gene pool. The Hardy-Weinburg Theory provides a theoretical description of
allelic frequency in non-evolving populations through a mathematically proportionate
relationship between alleles present. (p = dominant probability, q=recessive probability; p2 + 2pq
+ q2 = 1)
Microevolution is the generation to generation change in a populations allelic frequencies,
which are obviously small scale changes, but overtime represent evolutionary departures.
Because the population is evolving, the Hardy-Weinburg Theory/Equilibrium cannot provide
very accurate calculations. In order for the H.W. equilibrium to be maintained, there must be
large non-isolated populations with random mating, no net mutations, and no natural selection.
Microevolution can happen, however, as a result of genetic drifting, gene flow, mutations,
nonrandom mating, and natural selection.

Genetic drifting is a change in the gene pool of small populations due to chance. A genetic drift
can happen due to the Bottleneck Effect or the Founder Effect. The Bottleneck Effect occurs
with the non-selective death of members of a small population, reduction in population size,
random survival, and therefore a remaining population that differs genetically from the original
population. The Founder Effect results from colonization, where the genetic makeup of the
founding colony differs from the original population.
Gene flow is the migration of fertile individuals between populations, resulting in the transfer of
gametes. This reduces interpopulation differences and can offset the effects of genetic drift or
natural selection. Mutations in gametes affect microevolution, though their effects are about
1/1,000,000 in a large population, but are the original source of genetic variation. Nonrandom
mating increases the homogenous loci but does not alter allelic frequency. Nonrandom mating
can happen either as a result of inbreeding or assortative mating. Inbreeding occurs because
individuals are more likely to mate with close neighbors and it changes genotypic frequencies, as
it reduces the number of heterozygotes, but not allelic. Assortative mating occurs when
individuals mate with partners possessing similar phenotypes. Natural selection describes
particular individuals being favored when there is variation and therefore those individuals being
the most likely to produce offspring, passing on alleles in disproportionate numbers to those of
the current generation. It is an adaptive process by which favorable genotypes accumulate.
Natural selection requires variation, that is, an actual assortment of genes to select from. Where
do these variations come from? Subtle gene differences occurring within or between
populations, as well as mutations and sexual recombination, bring about variation. Variation
within populations affects character traits which can be either quantitative or discrete.
Polygenetic characters vary quantitatively as the trait is controlled by multiple loci; it contributes
most to inheritable variation, such as height. Discrete characters are controlled by a single locus,
and different alleles produce distinct phenotypes. Two or more different allele forms is called
polymorphism, morph meaning form, and morphs are present is significant frequencies in a
population. Morphs are contrasting character states that are not limited to physical traits;
cellular and biochemical substrates also exist in multiple character states.
There is also variation between populations due to differing environmental factors, and genetic
drift may cause variation among different populations. Some variables may be graded along a
geographic area, such as incremental temperature differences, altitude, and food source
differences. When different populations of the same species live along an ecocline and exhibit
genetic variation that parallels the environmental differences, it is referred to as a cline.
Mutations, most intuitively, contribute to genetic variation, though they are random changes in
the genetic composition of a population. Not all mutations producing new alleles will be
heritable; most mutations occur in somatic cells but only mutations occurring in gametes
generate heritable genetic variation. Mutations have the longest impact on variation in
organisms with short generational lines, as it does not take long for the mutation to spread and
therefore the allelic frequency of the mutation locus can change rapidly; and also in asexually
reproducing microorganisms. However, nearly all genetic variation results from sexual
recombination of existing alleles. The recombination occurs in the random segregation and
independent assortment during meiosis, synapsis of prophase I, and random fertilizationall
resulting in zygotes that possess new combinations of existing alleles.

Though variation is necessary for natural selection, natural selection actually reduces genetic
variability, selecting those most perfectly fit for their environment. However, supposing the
environment to which they are ideally adapted undergoes a sudden drastic change, that entire
species would quickly become extinct. Therefore there must be mechanisms at play that
preserve variation as natural selection occurs, and those are diploidy, balanced polymorphism,
and neutral variation.
Diploidy allows recessive alleles to be preserved in the heterozygote, though they are often less
favorable and sometimes harmful. Balanced polymorphism is when the heterozygote actually
has greater reproductive success than either homozygote. Some variations confer no advantage
or disadvantage, and this is neutral variation. The relative frequencies of neutral alleles are not
affected by natural selection.
Lecture 2 - Mechanisms of Adaptive Evolution
Adaptive evolution results from the combined effect of chance events that cause genetic
variation, and natural selection that favors some of those variations over others. The
evolutionary fitness of an individual refers to the relative contribution an individual makes to the
gene pool of the next generation, and this is measured by fecundity. Fecundity is the number of
progeny produced, which is determined by the reproductive lifespan of the individual and its
survival.
There are a few different types of genotypic selection that can occur, each having different
effects and serving different purposes. Stabilizing selection favors intermediates, as opposed to
extreme phenotypes, reducing phenotypic variation, and is best suited for when environmental
conditions are stable. Directional selection favors variants of one extreme, shifting the frequency
curve toward rare variants, and this occurs during periods of environmental change.
Diversifying selection actually favors opposite phenotypic extremes over intermediate.
The process by which new biological species arise is conveniently called speciation, though there
are different types of speciation. The basis for naming new species is anagenesis, so called when
there is a transformation of an unbranched lineage of organisms to a state different from the
ancestral population. The more common pattern of speciation is cladogenesis, which is the
budding of one or more new species from a parent species that continues to exist, and which also
produces more biological diversity.
The criteria for a biological species consists in that it must be a population whose members have
the potential to interbreed and the breeding results in the production of viable and fertile
offspring, but who also cannot produce viable, fertile offspring with other species, therefore they
are reproductively isolated from other species in nature. A species is the largest unit of a
population where gene flow is possible. There are also other means of characterizing species,
basing the conceptualization on criteria such as morphology, ecology, evolution, or
characteristics that maximize successful mating.

What factors would cause the reproductive isolation that brings about a discrete species? Most
species are isolated by more than a single type of barrier. There are three types of barriers:
prezygotic, postzygotic, and geographic. Prezygotic barriers are intrinsic barriers that impede
mating or fertilization.
Postzygotic barriers are also intrinsic, and they prevent hybrids from developing into a viable,
fertile adult.
Lecture 9 Blood Vessels