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Saturday, 26 November 2016

Lotka Volterra equation for competition and Predation



The Lotka Volterra equation are used to interpret the population dynamics in which two organisms interact in one of the two ways, (a) either compete for common resources or (b) associated in a prey-predator system.
The equation for first type of interaction i.e. competing for common resources is termed as the Competitive Lotka-Volterra equations while another type of interaction is described by predator-prey equation.
1.     Competitive Lotka-Volterra equations
The Lotka-Volterra equation for competition is based on the logistic equation. This equation is similar to Predation prey equation of Lotka-Volterra where species interact with others by one term and to itself by another term but this equation follows exponential mode rather than logistic model.
Ecologists used the equation for logistic model is given as:
dN/dt= rN (1- N/K)
·        N= population size
·        r=growth rate of population
·        K=carrying capacity

For competition between two species
In the Lotka-Volterra equation two additional terms were added to depict the species interactions between two given population N1 and N2 related logistic dynamics.
The equation is given as:
dN1/dt= r1N1 [1-(N1+a12N1/K1)] for species 1
dN2/dt= r2N2 [1-(N2+a21N2/K2)] for species 2

We all know that each organism has its own carrying capacity (K1 and K2 are different) and growth rate (r1 and r2 are different). As this equation for population dynamics is associated with interaction (competition) which are harmful to interesting species, in equation all a values are positive. In this equation a12 termed as competition coefficient depicts the competitive effect on population one (N1) by another population (N2) that’s why represent as (a12). Similarly for the equation of population of N2, a21 depicts the competitive effect on population N2 by the population N1 represented as (a21). If a12<1 it means species 1 has more effect on its own rather than the  effect of species 2 on species 1 i.e. more intense intra-specific competition.

In short the effected population comes first to the population affecting it.

Outcomes Interpretation
1.     If a12=0, this means species 1 follow logistic model of population dynamics.
2.     If a21=0, this means species 2 follow logistic model of population dynamics.
3.     If a12=1, species 1 and 2 strongly compete with equal magnitude for common resources means intraspecific competition equals interspecific competition.
4.     If a12= “-” negative, species 2 facilitates resource availability to species 1.
5.     a12/ a21 both negative indicate symbiotic relationships.
6.     If one is zero (0) among two, either a12 or a21 and the other is negative it indicates commensalism.
7.     If one is positive among two, either a12 or a21 and the other has no affect it means parasitism.
8.     If both have negative values it indicates competition.

Lotka Volterra Predation equations

The Predator prey equations given by Lotka-Volterra describe the interaction between prey and predation as a dynamic biological system.
The population follows a non-linear, first order differential equation and represented in pair of equation as
dx/dt = ax – βxy
dy/dt = δxy- γy
where “x” and “y” is the number of prey and predator population and “dx/dt” or “dy/dt” are the growth rate of prey and predator population in time “t”. The symbols α, β, γ and δ denotes the real and positive parameters related to the interaction between prey and predator species.

Assumptions
Lotka-Volterra predation equation is based on the assumptions:
1.     Predator can eat limitless.
2.     Supply of food resource (i.e. prey) depends on the prey population size.
3.     The rate of change of population directly depends on its size.
4.     Environment is constant, inconsequential genetic adaptations for both species.
5.     All time unlimited food supply for prey.
The equation is continuous and deterministic indicated continuous overlapping of prey and predator population.
Prey
The rate of growth for prey population is given by equation:
dx/dt = ax – βxy
The change in prey population follows exponential growth model represent as “ax” in the equation unless subjected to predation. The rate of predation is directly dependent on the rate of prey-predator interaction represented as “βxy”. “x” and “y” are the population size of prey and predator, if x/y is zero it clearly indicates that there is no predation.
Predator
The rate of growth for predator population is given by equation:
dy/dt = δxy – γy
The change in predator population equals to food supply mediated growth minus death of predators.
The growth of predator population is represented as “δxy”. “δ” is different from “β” as the rate of predator population growth is very much different from the rate of predation on prey. “γy” is the decay rate of predator either due to emigration or mortality. In the prey absence the predator follow an exponential decay.

The solution to equation is periodic and yields a simple harmonic motion where prey population is traced by predation population by 90 in cycle.

Figure 27: Harmonic motion of prey-predator system

Functional and numerical responses
In 1959, Holling suggested that as the prey density increases it leads to increase in Predation rates due to two effects:
(a)  Functional Response where the consumption rate of predator increases in presence of high prey population density.
(b)  Numerical Response in which increase in prey density leads to increase in predator population density.

a.     Functional Response
There are three types of functional response curves which relate density of prey population to prey consumption by single predator per unit time.

Type I Response curve
It’s a rare form of response in nature, for example in Filter feeders.
It’s an initial exponential relationship between prey and predator and its consumption by predator till reaches saturation point where predator cannot eat maximum from that rate.
Type II Response curve
It’s a common type of response in nature, for example rodents and weasels. At low population density the rate of consumption increases at decelerating rate (increases at slow rate).the rate of consumption is dependent on two factors, (a) the searching or locating a prey and (b) handling time of prey (capture, kill and eat). At low population density the searching for prey is important and predator kill prey at constant effort fashion while at high density of prey searching for prey becomes easy but handling time is limiting factor and thus the rate of consumption increases slowly. Subsequently, searching Is not required and rate of consumption levels off at maximum rate.
Type III Response curve
Type III response is also common in nature depicting logistic increase in the rate of consumption on increase in prey density.

Figure 28: Three types of Functional response curves
Numerical response curves

Numerical response means increase in predator density on increase in prey population density due to direct responses i.e. (a) on increase in prey density the fecundity rate of predator increases and (b) Aggregation response
a.     As the prey population size increases, the consumption rate of predator increases leads to high fecundity rate and low mortality rate.
b.     Aggregation response: Predators aggregation in prey hot spots is called aggregation response. This type of response increase prey-predator system stability.

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