Neutral Evolution

&
Molecular Clock

Presented By
Miss Aditi Patil Under the
M.Sc. Part I Guidance of
24/01/2009 Dr. Swati
Bhavsar
Neutral Theory Of Molecular
Evolution
• Motoo Kimura (1960-1970)
• Does not deny natural selection
• Genetic polymorphism
• Neutral alleles with respect to
fitness
• Random fixation of selectively
neutral mutants
• Mutation and genetic drift
Relative importance of drift
and selection

• Molecular
evolution
• Evolution at
phenotypic
Figure 1 : Balance
between natural
selection and random
level
Mathematical Expression Of The
Neutral Theory
• If population size is N and µ is
neutral mutation rate per gamete
per generation at a locus.
No. of new mutations = µ X 2N
(E.g.: mutation in protein ‘x’ in
experimental rat)
- given µ for the rat is 2.5 X 10 -9
- consider population size N = 50
No. of new mutations = 1.25X 10 -7
• According to drift theory,
P = 1/(2N)
where P = probability that a mutation will be fixed
by drift

Thus, P for protein ‘x’ ,

P = 1/(2X50) = 0.01

• No. of mutations fixed per generation is,

= µ X 2N X 1/(2N) = µ
Where µ = Neutral mutation rate per gamete
per generation at a locus
• No. of mutations in protein ‘x’ fixed
per generation is,
= 2.5 X 10 -9 X100 X 1/100
=2.5X10 -9 mutation per gamete
per generation.

• Therefore, the rate of neutral

evolution is theoretically
constant and is equal to the rate of
mutation.
Applications Of Neutral
Theory
• The faster rates

• The degrees of polymorphism

• Constancy of molecular
evolution
 The Molecular Clock
Hypothesis

• Zuckerkandl and Pauling

(1965)
• The rate of molecular
evolution
• Molecular clock
• Species divergences and
constructing phylogenies
 Molecular Clock And Neutral
Theory
• The molecular clock
hypothesis
• The neutral theory prediction
• Rate of evolution per
generation
• Rate of mutation
• Rate variation among
lineages
Figure 2 : Proteins evolving at different rates
 Causes of Rate Variation
Among Lineages
• The Generation-time Effect
• Nucleotide substitution rates
• Substitution fixation rates
• species with shorter generation
times
• Higher rates of evolution
• Effect of near-neutrality
• offset the generation time effect
Limitations Of Molecular
Clock
• Changing generation times
• Population size
• Species-specific differences
• Changes in the intensity of
natural selection
Applications Of Molecular
Clock
• Molecular systematics
• Correct scientific classification
of organisms
• Dates of phylogenetic events

• Divergence of living taxa

• Formation of the phylogenetic

tree
 References
• M. Kimura, The neutral theory of molecular evolution: a
review of recent evidence, Japanese Journal of Genetics
(1991), 66: 367-86.
• http://understandingevolution.com/evosite/evo101/IIIE5b
Neutraltheory.shtml
• http://plato.stanford.edu/entries/evolutionary-
genetics/figure2.gif
• www.pbs.org/wgbh/evolution/library/05/1/pdf/l_051_06.p
df
• http://en.wikipedia.org/wiki/Neutral_theory_of_molecular
_evolution
• http://en.wikipedia.org/wiki/Molecular_clock
Thank you