Random Walks In Biology Pdf

[Inell Krolick]

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Mathematical modelling of the movement of animals, micro-organisms and cells is of great relevance in the fields of biology, ecology and medicine. Movement models can take many different forms, but the most widely used are based on the extensions of simple random walk processes. In this review paper, our aim is twofold: to introduce the mathematics behind random walks in a straightforward manner and to explain how such models can be used to aid our understanding of biological processes. We introduce the mathematical theory behind the simple random walk and explain how this relates to Brownian motion and diffusive processes in general. We demonstrate how these simple models can be extended to include drift and waiting times or be used to calculate first passage times. We discuss biased random walks and show how hyperbolic models can be used to generate correlated random walks. We cover two main applications of the random walk model. Firstly, we review models and results relating to the movement, dispersal and population redistribution of animals and micro-organisms. This includes direct calculation of mean squared displacement, mean dispersal distance, tortuosity measures, as well as possible limitations of these model approaches. Secondly, oriented movement and chemotaxis models are reviewed. General hyperbolic models based on the linear transport equation are introduced and we show how a reinforced random walk can be used to model movement where the individual changes its environment. We discuss the applications of these models in the context of cell migration leading to blood vessel growth (angiogenesis). Finally, we discuss how the various random walk models and approaches are related and the connections that underpin many of the key processes involved.

In mathematics, a random walk, sometimes known as a drunkard's walk, is a stochastic process that describes a path that consists of a succession of random steps on some mathematical space.

A popular random walk model is that of a random walk on a regular lattice, where at each step the location jumps to another site according to some probability distribution. In a simple random walk, the location can only jump to neighboring sites of the lattice, forming a lattice path. In a simple symmetric random walk on a locally finite lattice, the probabilities of the location jumping to each one of its immediate neighbors are the same. The best-studied example is the random walk on the d-dimensional integer lattice (sometimes called the hypercubic lattice) Z d \displaystyle \mathbb Z ^d .[3]

If the state space is limited to finite dimensions, the random walk model is called a simple bordered symmetric random walk, and the transition probabilities depend on the location of the state because on margin and corner states the movement is limited.[4]

To answer the question of how many times will a random walk cross a boundary line if permitted to continue walking forever, a simple random walk on Z \displaystyle \mathbb Z will cross every point an infinite number of times. This result has many names: the level-crossing phenomenon, recurrence or the gambler's ruin. The reason for the last name is as follows: a gambler with a finite amount of money will eventually lose when playing a fair game against a bank with an infinite amount of money. The gambler's money will perform a random walk, and it will reach zero at some point, and the game will be over.

The central limit theorem and the law of the iterated logarithm describe important aspects of the behavior of simple random walks on Z \displaystyle \mathbb Z . In particular, the former entails that as n increases, the probabilities (proportional to the numbers in each row) approach a normal distribution.

As a direct generalization, one can consider random walks on crystal lattices (infinite-fold abelian covering graphs over finite graphs). Actually it is possible to establish the central limit theorem and large deviation theorem in this setting.[7][8]

The heterogeneous random walk draws in each time step a random number that determines the local jumping probabilities and then a random number that determines the actual jump direction. The main question is the probability of staying in each of the various sites after t \displaystyle t jumps, and in the limit of this probability when t \displaystyle t is very large.

In higher dimensions, the set of randomly walked points has interesting geometric properties. In fact, one gets a discrete fractal, that is, a set which exhibits stochastic self-similarity on large scales. On small scales, one can observe "jaggedness" resulting from the grid on which the walk is performed. The trajectory of a random walk is the collection of points visited, considered as a set with disregard to when the walk arrived at the point. In one dimension, the trajectory is simply all points between the minimum height and the maximum height the walk achieved (both are, on average, on the order of n \displaystyle \sqrt n ).

To visualize the two-dimensional case, one can imagine a person walking randomly around a city. The city is effectively infinite and arranged in a square grid of sidewalks. At every intersection, the person randomly chooses one of the four possible routes (including the one originally travelled from). Formally, this is a random walk on the set of all points in the plane with integer coordinates.

To answer the question of the person ever getting back to the original starting point of the walk, this is the 2-dimensional equivalent of the level-crossing problem discussed above. In 1921 George Plya proved that the person almost surely would in a 2-dimensional random walk, but for 3 dimensions or higher, the probability of returning to the origin decreases as the number of dimensions increases. In 3 dimensions, the probability decreases to roughly 34%.[9] The mathematician Shizuo Kakutani was known to refer to this result with the following quote: "A drunk man will find his way home, but a drunk bird may get lost forever".[10]

Another variation of this question which was also asked by Plya is: "if two people leave the same starting point, then will they ever meet again?"[13] It can be shown that the difference between their locations (two independent random walks) is also a simple random walk, so they almost surely meet again in a 2-dimensional walk, but for 3 dimensions and higher the probability decreases with the number of the dimensions. Paul Erdős and Samuel James Taylor also showed in 1960 that for dimensions less or equal than 4, two independent random walks starting from any two given points have infinitely many intersections almost surely, but for dimensions higher than 5, they almost surely intersect only finitely often.[14]

The asymptotic function for a two-dimensional random walk as the number of steps increases is given by a Rayleigh distribution. The probability distribution is a function of the radius from the origin and the step length is constant for each step. Here, the step length is assumed to be 1, N is the total number of steps and r is the radius from the origin.[15]

A Wiener process is a stochastic process with similar behavior to Brownian motion, the physical phenomenon of a minute particle diffusing in a fluid. (Sometimes the Wiener process is called "Brownian motion", although this is strictly speaking a confusion of a model with the phenomenon being modeled.)

A Wiener process is the scaling limit of random walk in dimension 1. This means that if there is a random walk with very small steps, there is an approximation to a Wiener process (and, less accurately, to Brownian motion). To be more precise, if the step size is ε, one needs to take a walk of length L/ε2 to approximate a Wiener length of L. As the step size tends to 0 (and the number of steps increases proportionally), random walk converges to a Wiener process in an appropriate sense. Formally, if B is the space of all paths of length L with the maximum topology, and if M is the space of measure over B with the norm topology, then the convergence is in the space M. Similarly, a Wiener process in several dimensions is the scaling limit of random walk in the same number of dimensions.

A random walk is a discrete fractal (a function with integer dimensions; 1, 2, ...), but a Wiener process trajectory is a true fractal, and there is a connection between the two. For example, take a random walk until it hits a circle of radius r times the step length. The average number of steps it performs is r2.[citation needed] This fact is the discrete version of the fact that a Wiener process walk is a fractal of Hausdorff dimension 2.[citation needed]

In two dimensions, the average number of points the same random walk has on the boundary of its trajectory is r4/3. This corresponds to the fact that the boundary of the trajectory of a Wiener process is a fractal of dimension 4/3, a fact predicted by Mandelbrot using simulations but proved only in 2000by Lawler, Schramm and Werner.[16]

A Wiener process enjoys many symmetries a random walk does not. For example, a Wiener process walk is invariant to rotations, but the random walk is not, since the underlying grid is not (random walk is invariant to rotations by 90 degrees, but Wiener processes are invariant to rotations by, for example, 17 degrees too). This means that in many cases, problems on a random walk are easier to solve by translating them to a Wiener process, solving the problem there, and then translating back. On the other hand, some problems are easier to solve with random walks due to its discrete nature.

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