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Modern Synthesis

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DOI: 10.1007/978-3-319-47829-6_203-1

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M
Modern Synthesis
Vertika Singh and Kiran Singh
Department of Molecular & Human Genetics,
Banaras Hindu University, Varanasi, India
Synonyms
Modern synthetic theory
Definition
The modern synthetic theory of evolution
describes the evolution in terms of genetic varia-
tions in a population that leads to the formation of
a new species.
Introduction
Darwin’s book, The Origin of Species by Means of
Natural Selection, founded a strong ground for
evolutionary theories that favored the idea that
all the organisms have descended from a common
ancestor. Darwin proposed natural selection as a
mechanism of evolution. Then came the neo-
Darwinian theory of evolution that explained the
significance of mutations and variations within a
population, as a driver factor of evolution. This
perspective was followed for many decades.
# Springer International Publishing AG 2018
J. Vonk, T.K. Shackelford (eds.), Encyclopedia of Animal Cognition and Behavior,
https://doi.org/10.1007/978-3-319-47829-6_203-1
Ronald Fisher and his colleagues set Darwin’s
concept of natural selection on a new foundation
of genetics. They left an equally major project
open for later biologists: to explain in the lan-
guage of genes what species are and how they
originate. While the Darwin’s theory largely dem-
onstrated the “facts” of evolution, the “mecha-
nism” of evolution was still unexplained. The
answer only began to emerge in the 1930s, thanks
in large part to the work of a Soviet-born geneti-
cist named Theodosius Dobzhansky, who pro-
posed the idea of modern synthesis.
However, a quiet majority of geneticists
thought of species’ origin as the gradual accumu-
lation of small mutations, and in 1940 this allowed
for propagation of “the synthetic theory of evolu-
tion” by geneticist Theodosius Dobzhansky
(1900–1975) and other zoologists.
The neo-Darwinian view was then replaced by
a new concept which considered several other
mechanisms in addition to natural selection.
These ideas on evolution are usually referred to
as the modern synthesis which is described by
Futuyma as “The major tenets of the evolutionary
synthesis, then, were that populations contain
genetic variation that arises by random (i.e. not
adaptively directed) mutation and recombination;
that populations evolve by changes in gene fre-
quency brought about by random genetic drift,
gene flow, and especially natural selection; that
most adaptive genetic variants have individually
slight phenotypic effects so that phenotypic
changes are gradual (although some alleles with
2 Modern Synthesis
discrete effects may be advantageous, as in certain
color polymorphisms); that diversification comes
about by speciation, which normally entails the
gradual evolution of reproductive isolation among
populations; and that these processes, continued
for sufficiently long, give rise to changes of such
great magnitude as to warrant the designation of
higher taxonomic levels (genera, families, and so
forth)” (Futuyma 1986).
Modern Synthetic Theory of Evolution
The modern synthetic theory of evolution
describes the evolution in terms of genetic varia-
tions in a population that leads to the formation of
a new species. It explains the contribution of
factors such as genetic variations, reproductive
and geographical isolation, and natural selection.
It also describes about the Mendelian population,
gene pool, and gene frequency. It merges the
concepts of Darwinian theory of evolution and
Mendelian genetics to form a unified theory of
evolution. Synthetic theory of evolution was
introduced by few renowned evolutionary biolo-
gists naming T. Dobzhansky, J.B.S. Haldane,
R.A. Fisher, Sewall Wright, G.L. Stebbins, and
Ernst Mayr in the years 1930–1940.
The modern synthetic theory of evolution pro-
posed a new definition to the evolution as “the
changes occurring in the allele frequencies within
the populations,” which accentuates on the genet-
ics of evolution.
Aspects of Modern Synthetic Theory of
Evolution
Some of the factors describing the modern theory
of synthetic evolution can be explained as
follows.
Recombination or Variation
Recombination occurs between the existing genes
to form a new genotype. It is an exchange of the
chromosomal pairs of alleles during the meiosis at
the time of sexual reproduction to form new gene
combinations. Cytogenetic events such as
mutations (including deletion, inversion, duplica-
tion, translocation, and polyploidy) result in the
recombination.
Mutation
A mutation is a change in DNA, the hereditary
material of life. An organism’s DNA affects its
phenotype, its behavior, and its physiology. Muta-
tions are the raw materials of evolution. Evolution
is categorically dependent on mutations because it
enables the formation of new alleles and new
regulatory regions.
Heredity
The transmission of variations from the parents to
their offsprings is a principal mechanism in the
evolution. The organisms which retain hereditary
characteristics are preferred in the struggle for the
existence.
Natural Selection
Natural selection is a phenomenon by which spe-
cies adapt to their environment. Natural selection
results in an evolutionary change when individ-
uals with certain characteristics have a greater
survival or reproductive rate than other individ-
uals in a population and pass on these inheritable
genetic characteristics to their offspring.
Reproductive Isolation
It is one of the crucial factors responsible for the
synthetic theory of evolution. Reproductive isola-
tion helps in preventing the interbreeding of
related organisms. Additionally, some other fac-
tors that contribute to the synthetic theory of evo-
lution include migration of individuals from one
form of the population to another and hybridiza-
tion between the races of species which increases
the genetic variability within the population.
It differs from Darwinism in three important
aspects:
1. In addition to natural selection, it considers
several mechanisms such as random genetic
drift to be an important element of evolution.
2. It describes that the characters are inherited as
discrete units called genes. Occurrence of
Modern Synthesis
multiple alleles of a gene within a population
gives rise to variation.
3. It postulates that speciation is (usually) due to
the gradual accumulation of small genetic
changes. This is equivalent to saying that mac-
roevolution is simply a lot of microevolution.
Modern synthesis was one of the utmost
accomplishments of evolutionary biology. By
combining the concepts of Darwin and Mendel,
it expounded that diversity within a population
ascended from random occurrence of mutations
and environment acted to select the phenotype
that is more fit. Potential reproducing animals
transmitted the genes that were advantageous.
These types of genes included those coding for
enzymes with better rates of synthesis and globins
with better oxygen-carrying capacity. It was pre-
sumed that the genetic variations, that caused
evolution within a species, also caused evolution
of a new species. If a new phenotype is to be
produced, it is essential that there should be an
accumulation of these mutations followed by
reproductive isolation. The theory of modern syn-
thesis not only did explain the evolution within
species but also explained many relevant ques-
tions such as why some alleles that seem to be
deleterious get selected in certain populations, for
example, one of the hemoglobin gene variants
results in sickle cell anemia. The population
genetic approach to evolution was summed up
by Theodosius Dobzhansky, who stated that
“Evolution is a change in the genetic composition
of populations. The study of the mechanisms of
evolution falls within the province of population
genetics” (Dobzhansky 1951).
Thus, some of the core assumptions of modern
synthesis include five major postulations which
include:
1. Phenotypic variations of evolutionary signifi-
cance arise from genetic mutations occurring at
a low rate and independent of the strength and
direction of natural selection.
2. Most of the advantageous mutations show
small phenotypic effects which progressively
lead to the change in phenotype.
3. Inheritance is genetic in nature.
3
4. Natural selection is an exclusive explanation
for adaptation.
5. Accumulation of variations that arise through
microevolutionary processes results in
macroevolution.
Significance of Developmental Biology in
Modern Synthesis
At the time when the concept of modern synthesis
was formulated, developmental biology and
developmental genetics was at a naive stage. Sig-
nificance of embryology was neglected and was
kept out of modern synthesis by most of the
geneticists and evolutionary biologists
(Hamburger 1980; Gottlieb 1992; Dietrich 1995;
Gilbert et al. 1996). It was earlier assumed that
population genetics is enough to explain the evo-
lution; hence, developmental biology seemed to
contribute almost negligible in the modern evolu-
tionary theory (Adams 1991). However, it is not
well accepted that “developmental genetics
approach to evolution concerns more the arrival
of the fittest than the survival of the fittest”
(Gilbert 2000). Even the critics of modern syn-
thetic theory agreed that macroevolution, which
refers to the large morphological changes that are
seen between species, classes, and phyla, is based
on mutation and recombination. Nevertheless,
these macroevolutionary variations occur in
developmental regulatory genes in the embryo
and larvae, and not in adults competing for repro-
ductive success (Waddington 1953; Gilbert 1998).
Developmental biology has brought a new
platform to understand the relationship between
genotype and phenotype in the light of evolution-
ary biology. Also, it enables us to understand the
genetic relationships between diverse organisms.
Developmental biology thus provides a popula-
tion genetic approach to understand gene regula-
tion (Arthur 2000; Macdonald and Goldstein
1999; Zeng et al. 1999). It opens questions as to
how various paracrine factors, transcription fac-
tors, and signal transduction pathways were mod-
ified during the evolution.
Evolutionary developmental biology also pro-
vides answers to many classical evolutionary
questions such as those of the examples of mim-
icry and industrial melanism. Now that the genes
4 Modern Synthesis
responsible for such phenomena have been iden-
tified, the mechanism can be very well explained
(Koch et al. 1998; Brakefield 1998). Thus, to
explain the mechanism of evolution, both the
population and developmental genetics should
be taken under consideration.
The Extended Synthesis
The concept of modern synthesis emerged in first
half of the twentieth century and flourished for
many years. However, as the evolutionary biology
research progresses, several research subjects
emerged that transcended the boundaries of mod-
ern synthesis. Major thrust areas that challenged
the theories and concepts of modern synthesis
included epigenetics, phenotypic plasticity, evo-
lutionary developmental biology, and systems
biology. This new conceptual framework of evo-
lution was termed as extended synthesis, as it was
an extension, rather than contradiction, to the
modern synthesis (Pigliucci 2007). An excellent
description of this is provided in the book “Evo-
lution—The Extended Synthesis” (Pigliucci and
Müller 2010).
Epigenetics has been one of the major thrust
areas in extended synthesis. The term “epige-
netics” was first introduced by Conrad
Waddington in the early 1940s (Waddington
2011). He defined epigenetics as “the branch of
biology which studies the causal interactions
between genes and their products which bring
the phenotype into being”. However, as the field
of genetics expanded, it is now defined as, “the
study of changes in gene function that are mitot-
ically and/or meiotically heritable and that do not
entail a change in DNA sequence” (Morris 2001).
Epigenetic mechanisms include interactions
among various genetic, physiological, and mor-
phological systems and serve as an important
constituent of organism-environment interactions
(Angers et al. 2010; Richards et al. 2010; Schrey
et al. 2012).
The significance of epigenetics at molecular
level has been well elucidated and appreciated
for, e.g., its function in cell determination and
cell recognition; however the role of epigenetics
in evolution is recently under attention. Some
epigenetic characters are known to stably transmit
across generations, which highlights that it
includes some mechanisms of heredity which
was not considered in the framework of modern
synthesis.
Epigenetic mechanisms play critical roles and
several mechanisms such as in phenotypic plas-
ticity, invasive species biology, soft inheritance,
response to environmental variations, and conser-
vational biology. Epigenetic variations such as
DNA methylation provide an evolutionarily sig-
nificant source of phenotypic variation among
individuals. DNA methylation is the most studied
molecular epigenetic mechanism. It is actively
involved in mechanisms such as DNA imprinting,
X-chromosome inactivation, silencing transpos-
able elements, and response to environmental
stressors. DNA methylation is a basis for
interindividual variation and is studied to regulate
various phenomena such as phenotypic variation
in shape of flower and fruit pigmentation, shape of
mouse tail, body size and coat color, and numer-
ous traits differentiating queen and worker honey-
bees. For example, Viola cazorlensis shows a high
level of interindividual DNA methylation varia-
tion that differentiated populations (Schrey
et al. 2012).
Nevertheless, DNA methylation serves as the
most comprehensively studied mechanism in
ecology and evolutionary perspectives; several
studies have described other epigenetic mecha-
nisms also. Modifications of histone, small and
long noncoding RNAs, and genome structure are
shown to regulate gene expression and contribute
to phenotypic variation in diverse taxa.
Epigenetics continues to prove as a remarkable
phenomenon in evolutionary biology. A great deal
of work, however, remains to be done. Pragmatic
studies that established the significance of epige-
netic variation in evolution will likely explain
some of the empirical questions of evolutionary
epigenetics, which will further help in develop-
ment and enhancement of a foundational theory of
evolutionary epigenetics. Thus, epigenetics stud-
ies seem to be very promising in providing infor-
mation about the individual and population
processes both at the ecological and evolutionary
timescales.
Modern Synthesis
Cross-References
▶ Epigenesist
▶ Evolution
▶ Genetic Variation
▶ Heritability
▶ Population
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