Mathematical and theoretical biology

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Mathematical and theoretical biology is a branch of biology which employs theoretical analysis,
mathematical models and abstractions of the living organisms to investigate the principles that
govern the structure, development and behavior of the systems, as opposed to experimental
biology which deals with the conduction of experiments to prove and validate the scientific theories.
[1]
The field is sometimes called mathematical biology or biomathematics to stress the
mathematical side, or theoretical biology to stress the biological side.[2] Theoretical biology focuses
more on the development of theoretical principles for biology while mathematical biology focuses on
the use of mathematical tools to study biological systems, even though the two terms are sometimes
interchanged.[3][4]
Mathematical biology aims at the mathematical representation and modeling of biological processes,
using techniques and tools of applied mathematics. It has both theoretical and practical applications
in biological, biomedical and biotechnology research. Describing systems in a quantitative manner
means their behavior can be better simulated, and hence properties can be predicted that might not
be evident to the experimenter. This requires precise mathematical models.
Mathematical biology employs many components of mathematics,[5] and has contributed to the
development of new techniques.

Contents

 1History

o 1.1Early history

o 1.2Recent growth

 2Areas of research

o 2.1Evolutionary biology

o 2.2Computer models and automata theory

o 2.3Molecular set theory

o 2.4Mathematical methods

o 2.5Mathematical biophysics

 2.5.1Deterministic processes (dynamical systems)

 2.5.2Stochastic processes (random dynamical systems)

 2.5.3Spatial modelling

o 2.6Organizational biology
 o 2.7Algebraic biology

o 2.8Computational neuroscience

 3Model example: the cell cycle

 4Societies and institutes

 5See also

 6Notes

 7References

 8Further reading

 9External links

History[edit]
Early history[edit]
Mathematics has been applied to biology as early as the 12th century, when Fibonacci used the
famous Fibonacci series to describe a growing population of rabbits. In the 18th century Daniel
Bernoulli applied mathematics to describe the effect of smallpox on the human population. Thomas
Malthus' 1789 essay on the growth of the human populution was based on the concept of
exponential growth. Pierre Francois Verhulst formulated the logistic growth model in 1836.
Fritz Müller described the evolutionary benefits of what is now called Müllerian mimicry in 1879, in an
account notable for being the first use of a mathematical argument in evolutionary ecology to show
how powerful the effect of natural selection would be, unless one includes Malthus's discussion of
the effects of population growth that influenced Charles Darwin: Malthus argued that growth would
be exponential (he uses the word "geometric") while resources (the environment's carrying capacity)
could only grow arithmetically.[6]
The term "theoretical biology" was first used by Johannes Reinke in 1901. One founding text is
considered to be On Growth and Form (1917) by D'Arcy Thompson,[7] and other early pioneers
include Ronald Fisher, Hans Leo Przibram, Nicolas Rashevsky and Vito Volterra.[8]
Recent growth[edit]
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Interest in the field has grown rapidly from the 1960s onwards. Some reasons for this include:

 The rapid growth of data-rich information sets, due to the genomics revolution, which are
difficult to understand without the use of analytical tools

 Recent development of mathematical tools such as chaos theory to help understand

complex, non-linear mechanisms in biology

 An increase in computing power, which facilitates calculations and simulations not previously
possible
 An increasing interest in in silico experimentation due to ethical considerations, risk,
unreliability and other complications involved in human and animal research

Areas of research[edit]
Several areas of specialized research in mathematical and theoretical biology [9][10][11][12][13] as well as
external links to related projects in various universities are concisely presented in the following
subsections, including also a large number of appropriate validating references from a list of several
thousands of published authors contributing to this field. Many of the included examples are
characterised by highly complex, nonlinear, and supercomplex mechanisms, as it is being
increasingly recognised that the result of such interactions may only be understood through a
combination of mathematical, logical, physical/chemical, molecular and computational models.
Evolutionary biology[edit]
Ecology and evolutionary biology have traditionally been the dominant fields of mathematical
biology.
Evolutionary biology has been the subject of extensive mathematical theorizing. The traditional
approach in this area, which includes complications from genetics, is population genetics. Most
population geneticists consider the appearance of new alleles by mutation, the appearance of
new genotypes by recombination, and changes in the frequencies of existing alleles and genotypes
at a small number of gene loci. When infinitesimal effects at a large number of gene loci are
considered, together with the assumption of linkage equilibrium or quasi-linkage equilibrium, one
derives quantitative genetics. Ronald Fisher made fundamental advances in statistics, such
as analysis of variance, via his work on quantitative genetics. Another important branch of population
genetics that led to the extensive development of coalescent theory is phylogenetics. Phylogenetics
is an area that deals with the reconstruction and analysis of phylogenetic (evolutionary) trees and
networks based on inherited characteristics[14] Traditional population genetic models deal with alleles
and genotypes, and are frequently stochastic.
Many population genetics models assume that population sizes are constant. Variable population
sizes, often in the absence of genetic variation, are treated by the field of population dynamics. Work
in this area dates back to the 19th century, and even as far as 1798 when Thomas
Malthus formulated the first principle of population dynamics, which later became known as
the Malthusian growth model. The Lotka–Volterra predator-prey equations are another famous
example. Population dynamics overlap with another active area of research in mathematical
biology: mathematical epidemiology, the study of infectious disease affecting populations. Various
models of the spread of infections have been proposed and analyzed, and provide important results
that may be applied to health policy decisions.
In evolutionary game theory, developed first by John Maynard Smith and George R. Price, selection
acts directly on inherited phenotypes, without genetic complications. This approach has been
mathematically refined to produce the field of adaptive dynamics.