Von Foerster Equation
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The McKendrick–von Foerster equation is a linear first-order
partial differential equation In mathematics, a partial differential equation (PDE) is an equation which imposes relations between the various partial derivatives of a Multivariable calculus, multivariable function. The function is often thought of as an "unknown" to be sol ...
encountered in several areas of
mathematical biology Mathematical and theoretical biology, or biomathematics, 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 a ...
– for example,
demography Demography () is the statistics, statistical study of populations, especially human beings. Demographic analysis examines and measures the dimensions and Population dynamics, dynamics of populations; it can cover whole societies or groups ...
and
cell proliferation Cell proliferation is the process by which ''a cell grows and divides to produce two daughter cells''. Cell proliferation leads to an exponential increase in cell number and is therefore a rapid mechanism of tissue growth. Cell proliferation re ...
modeling; it is applied when age structure is an important feature in the
mathematical model A mathematical model is a description of a system using mathematical concepts and language. The process of developing a mathematical model is termed mathematical modeling. Mathematical models are used in the natural sciences (such as physics, ...
. It was first presented by
Anderson Gray McKendrick Lt Col Anderson Gray McKendrick DSc FRSE (8 September 1876 – 30 May 1943) was a Scottish military physician and epidemiologist who pioneered the use of mathematical methods in epidemiology. Irwin (see below) commented on the quality of his wor ...
in 1926 as a deterministic limit of lattice models applied to
epidemiology Epidemiology is the study and analysis of the distribution (who, when, and where), patterns and determinants of health and disease conditions in a defined population. It is a cornerstone of public health, and shapes policy decisions and evidenc ...
, and subsequently independently in 1959 by biophysics professor
Heinz von Foerster Heinz von Foerster (German spelling: Heinz von Förster; November 13, 1911 – October 2, 2002) was an Austrian American scientist combining physics and philosophy, and widely attributed as the originator of Second-order cybernetics. He was twice ...
for describing cell cycles.


Mathematical formula

The mathematical formula can be derived from first principles. It reads:where the population density n(t,a) is a function of age ''a'' and time ''t'', and m(a) is the death function. When m(a) = 0, we have: :\frac = - \frac It relates that a population ages, and that fact is the only one that influences change in population density; the negative sign shows that time flows in just one direction, that there is no birth and the population is going to die out.


Derivation

Suppose that for a change in time dt and change in age da, the population density is:n(t+dt,a + da) = -m(a)dt(t,a)That is, during a time period dt the population density decreases by a percentage m(a)dt. Taking a
Taylor series In mathematics, the Taylor series or Taylor expansion of a function is an infinite sum of terms that are expressed in terms of the function's derivatives at a single point. For most common functions, the function and the sum of its Taylor serie ...
expansion to order dt gives us that:n(t+dt,a + da) \approx n(t,a) + dt + daWe know that da/dt = 1, since the change of age with time is 1. Therefore, after collecting terms, we must have that: + = -m(a)n


Analytical solution

The von Foerster equation is a
continuity equation A continuity equation or transport equation is an equation that describes the transport of some quantity. It is particularly simple and powerful when applied to a conserved quantity, but it can be generalized to apply to any extensive quantity. S ...
; it can be solved using the
method of characteristics In mathematics, the method of characteristics is a technique for solving partial differential equations. Typically, it applies to first-order equations, although more generally the method of characteristics is valid for any hyperbolic partial d ...
. Another way is by
similarity solution In the study of partial differential equations, particularly in fluid dynamics, a self-similar solution is a form of solution which is similar to itself if the independent and dependent variables are appropriately scaled. Self-similar solutions ap ...
; and a third is a numerical approach such as finite differences. To get the solution, the following boundary conditions should be added: : n(t,0)= \int_0^\infty b (a)n(t,a) \, da which states that the initial births should be conserved (see Sharpe–Lotka–McKendrick’s equation for otherwise), and that: : n(0,a)= f(a) which states that the initial population must be given; then it will evolve according to the partial differential equation.


Similar equations

In Sebastian Aniţa, Viorel Arnăutu, Vincenzo Capasso. ''An Introduction to Optimal Control Problems in Life Sciences and Economics'' (Birkhäuser. 2011), this equation appears as a special case of the Sharpe–Lotka–McKendrick’s equation; in the latter there is inflow, and the math is based on directional derivative. The McKendrick’s equation appears extensively in the context of cell biology as a good approach to model the eukaryotic cell cycle.


See also

* Finite difference method *
Partial differential equation In mathematics, a partial differential equation (PDE) is an equation which imposes relations between the various partial derivatives of a Multivariable calculus, multivariable function. The function is often thought of as an "unknown" to be sol ...
*
Renewal theory Renewal theory is the branch of probability theory that generalizes the Poisson process for arbitrary holding times. Instead of exponentially distributed holding times, a renewal process may have any independent and identically distributed (IID) ho ...
*
Continuity equation A continuity equation or transport equation is an equation that describes the transport of some quantity. It is particularly simple and powerful when applied to a conserved quantity, but it can be generalized to apply to any extensive quantity. S ...
*
Volterra integral equation In mathematics, the Volterra integral equations are a special type of integral equations. They are divided into two groups referred to as the first and the second kind. A linear Volterra equation of the first kind is : f(t) = \int_a^t K(t,s)\,x(s) ...


References

{{Reflist Diffusion Parabolic partial differential equations Stochastic differential equations Transport phenomena Equations of physics Mathematical and theoretical biology Ecology Demography Epidemiology