1. Introduction
At present, research on ecology is mainly based on the long-term survival perspective constructed by the interaction between predator and prey to understand population dynamics in ecosystems. Mathematical models provide quantitative insights for studying species interactions. Due to the complexity of biological survival and the limitations of constructing models, the study of populations can only be based on the construction and analysis of some features (predation fear [
1,
2], harvest [
3], prey refuge [
4], additional food [
5] and so on) through mathematical models, rather than all features.
In biological mathematics, many scholars assume that the growth rate of a biological population is a linear function of the population, so they establish logistic growth models to study the changes in species populations. However, organisms that meet the logistic growth model are limited by limited resources [
6], do not take into account individual reproduction and nutrient supply, and they are suitable for low biological populations. In 1963, Smith [
7] found that experimental data of Daphnia algae did not conform to a linear model. Through simulations of experimental data, Smith hypothesized that the relative growth rate of population density was proportional to the amount of remaining food. Therefore, population dynamics with limited growth should be based on the proportion of unused available resources, the Smith model replaces the linear function in the Logistic model with a hyperbolic function [
8]. Currently, researchers are particularly interested in the following research. The authors explored the optimum control strategy of Cydia Pomonella integrated management predator–prey model with Smith growth in [
6]. The authors proposed a partial differential predator–prey model with the Smith growth function in [
9]. Utilizing the coupled map lattice method, the model transformed into the space- and time- discrete model, and researchers examined the potential of flip bifurcation, Neimark–Sacker bifurcation and Turing bifurcation. The authors explored the stability and Hopf bifurcation of the model with Smith growth under ODE and PDE models in [
10].
To describe the interaction of prey and predator more realistic, biological intrinsic relationships have been taken into consideration in the formulation of a predator–prey model. For example, bacteria emit signals between populations to alleviate predation pressure. When the concentration of signal molecules exceeds a critical value, the bacteria will initiate an anti-predation response [
11]. In addition, researchers find that prey often gather and exhibit collective defense mechanisms to resist predation. Therefore, the Monod–Haldane functional response is proposed and demonstrated to play a role in the defense of prey populations. When the prey exhibits individual behavior at a low population size, the predation rate will monotonically increase. Once the prey population exceeds a certain threshold size, it will exhibit group behavior and the predation rate will decrease [
12]. However, some species cooperate with each other to defend themselves against prey in order to maintain their sustainable growth [
13], and the Monod–Haldane functional response fails to capture this phenomenon. Therefore, some scholars discover a new response function that can effectively capture the inhibitory effect of cooperative defense on predation in the population, and in this functional response, group defense and cooperative defense can work together [
11]. Actually, the change of the cooperative defense affects species richness by experimental analysis rather than the viewpoint of dynamic analysis. Motivated by above works, in this paper, we study a prey–predator model with the group defense and cooperative defense.
In the real world, the dynamic behaviors of a model is determined by both the internal laws and the changes in the external environment. Interestingly, the biological models presented by stochastic differential equations may exhibit a wider range of complex dynamic features than ordinary differential Equations [
1,
14]. In particular, seasonal changes in food and climate can lead to seasonal fluctuations in population numbers. Thus, the variability and randomness of the environment are taken into account in the state of the species, which results in one or more model parameters being randomly perturbed. Since these perturbations are small and independent, they can be represented in terms of white noise. Many scholars study the dynamic behavior of stochastic models by introducing environmental white noise into deterministic models [
15,
16,
17]. Based on the above description, we study a stochastic perturbation predator–prey model.
In
Section 2, we formulate a stochastic prey–predator model with Smith growth rate and the effect of defense. In
Section 3, we provide the dynamical properties of the stochastic model, including basic mathematical analysis, sufficient conditions for the stationary distribution and its ergodic. In
Section 4, we present the conditions for the extinctions of populations. In
Section 5, we carry out comprehensive numerical simulations to substantiate the mathematical results. Finally, we discuss the ecological significance of our mathematical outcomes in
Section 6.
2. The Model
In 1838, considering that the population was constrained by food and living space, Belgian mathematician P F. Verhulst introduced a density constraint factor based on the Malthus population model and proposed the famous Logistic equation [
18].
Verhulst assumed a population growth rate of , and as the population increases, the population growth slows down. When , the growth rate is infinitely close to 0. When , the population no longer changes. This indicates that the amount of remaining resources will significantly affect the growth of the population. The higher the population density, the less resources, and the population growth will slow down. Conversely, the population will increase significantly.
By extension, the Lotka–Volterra equation was established, which takes into account the interactions between species and establishes the initial multi group prey–predator model.
The functional response function
represents the mutual constraint relationship between populations, which affects the stability of biological system dynamics and maintains the dynamic balance of community structure. Based on the above, we consider a general predator–prey model.
where
u,
v denote the density of prey and predator populations, respectively.
is the growth rate of prey in the absence of any predators.
is the functional response.
d is the natural death rate of prey.
b and
m represent the intrinsic growth rate and intra-specific competition of the predator population, respectively.
is environmental capacity of predator.
e is the conversion rate of prey.
The growth pattern of the prey population that we previously studied in the absence of a predator is logical growth, and the average growth rate is a linear function of population density. However, this is not realistic for certain species with limited food. Smith [
7] found that the average growth rate of large water fleas is non-linear. Through experimental analysis, Smith assumed that the relative rate of population density at the moment of
t was proportional to the amount of food remaining at that time. Therefore, incorporating the growth of prey based on the proportion of unused available resources into prey reproduction makes the model more realistic from an ecological point of view. In recent years, many mathematical models have been developed to investigate the influence of growth restrictions based on the proportion of available resources on the population structure and ecological diversity [
9,
10]. Thus,
the function of
in these ecological models has replaced the classical logical term, where
c represents the resource constraint parameter of the population, and
represents the mass substitution rate of the population at
k.
Some commonly used forms of function response
are Holling type I-IV, Beddington DeAngelis, ratio-dependent and Monod–Haldane. In particular, the Monod–Haldane type functional response
indicates that when the prey density is high enough, the predation rate decreases relative to the prey density, and the prey usually forms group defense to better protect themselves and resist predators. In fact, defense functions also include group defense and collaborative defense. By observing the collective phenomenon of bacteria, it was recognized that cooperative defense plays an important role in bacterial survival [
11]. Therefore, the combined effects of group defense and cooperative defense is reflected in the following functional reaction [
11],
where
T is the total time required for the predator. The number of prey consumed by the predator is linearly proportional to the search period
and prey density
u.
M is the total number of prey captured by the predator. Then,
,
with the proportion
. By deformation, the equation becomes
where
s is the strength of the group defense, and
is the cooperative defense. In addition,
Figure 1 shows the variation of prey with changes in group defense parameter intensity
s and collaborative defense parameter
.
This indicates that when the prey density is greater than , the prey exhibits a defensive effect. That is, is the critical density for activating group defense.
When the value of
is
,
will reach minimum. This indicates that when the cooperation of prey is low, the group defense is demonstrated by obtaining a high population size. On the contrary, as cooperation increases, the threshold population will decrease, meaning that the prey can exhibit group defense at a low population size, see
Figure 2.
To achieve the influence of this function response under the Smith growth pattern of prey, the proposed model takes the following form:
In fact, ecosystems are inevitably affected by changes in their living environment [
14,
19]. Environmental variables can serve as sources of random noise, and their random fluctuations lead to the stochastic dynamics of biological models. Therefore, we use white noise to represent random fluctuation in the natural world and propose the following stochastic predator–prey model:
where
are independent standard Brownian motions with
,
denote the intensities of white noises.
Throughout this paper, the processes and are defined on a complete probability space with a filtration . Denote . and almost surely is replaced by a.s.
3. Stationary Distribution and Ergodicity
In this section, we investigate the stochastic dynamics of model Equation (10) including the existence and uniqueness of the solution and the existence of ergodic stationary distribution.
Theorem 1. For any initial value , there is a unique positive solution of model Equation (10) and will remain in for all almost surely.
Proof. The proof is divided into two steps.
- Step 1.
Let
and
, then model Equation (10) is transformed into the following equation.
with initial values
,
. Obviously, the coefficients of Equation (11) satisfy local Lipschitz continuous, and there is a unique local solution of Equation (11) on
, where
is the explosion time. By Itô’s formula,
and
is the unique local positive solution of model Equation (10) with initial value
.
- Step 2.
Now, we prove
. Take a sufficiently large number
such that
,
. For each integer
, we define the stopping time:
Obviously, is increasing as . Set , where a.s.
Let us prove that . If this statement is false, there is a pair of constants and such that . Thus, there exists an integer such that , .
Define a
-function
by
By Itô’s formula,
where
with
. Thus,
where
.
Substituting into the above equation, we derive
where
. Integrating both sides of Equation (17) from 0 to
and taking the expectation yield
Set
,
. For every
,
or
equals either
or
,
is no less than
which implies
where
denotes the indicator function of
. Letting
, we obtain
, which yields a contradiction. Hence, we have
This completes the proof. □
Below, we will investigate whether prey and predator populations will continue to survive by proving the unique ergodic stationary distribution of model Equation (10). First, we give the following important lemma.
Lemma 1 ([
20]).
Assume there exists a bounded open set with a smooth boundary, satisfying the following conditions:- (B.1)
There is a positive number such that , , ;
- (B.2)
There exists a nonnegative -function and a positive constant C such that for .
For any integrable function in regard to the measure π, we have Then, the Markov process has a unique ergodic stationary distribution , and , is the diffusion matrix.
Theorem 2. If and hold, then model Equation (10) has a unique ergodic stationary distribution.
Proof. At first, we verify the condition (B.1) of Lemma 1. The diffusion matrix of model Equation (10) is
Obviously, there is a
such that
Thus, condition (B.1) of Lemma 1 holds.
Construct a
-function
as follows:
Select a suitable constant
such that
, where
It is apparent that is the unique stationary point of .
Define the following Lyapunov function
,
Utilizing Itô’s formula,
and
Therefore, for a sufficiently small
, we know that
where
and condition (B.2) holds. It follows from Lemma 1 that model Equation (10) is ergodic and has a unique stationary distribution. □
As a summary, we can clearly see from the theoretical results obtained above that the prey and predator populations tend to coexist for a long time, and the model is weakly stable.
5. Numerical Simulations
In this section, we validate our theoretical results by simulating stochastic model Equation (10) with the help of MATLAB R2021b software and possible parameters. Select initial values as , for all figures.
Case I. Choose the intensities of white noises as
,
and the other parameters values are taken as
,
,
,
,
,
,
,
,
,
,
. By calculation, the conditions of Theorem 2 are all satisfied. Then, we conclude that model Equation (10) is a stationary distribution and ergodic (see
Figure 3). The left subgraphs of
Figure 3 represent the densities of prey and predator; meanwhile, the right subgraphs give the frequency histograms for
and
. In addition, we decrease the intensities of white noises to
,
in
Figure 4, and they also present a stationary distribution. By comparing
Figure 3 and
Figure 4, we can see that with the decrease in white noises, the fluctuation regions of the solutions become smaller.
Case II. To verify the effects of Smith growth rate on the stochastic model, we keep other coefficients to be the same as Case I. We only vary the size of the resource constraint parameter
,
. By observing
Figure 5, it can be concluded that a smaller
c will lead to the extinction of prey, and when
, that is, prey growth follows a logical pattern, prey can sustain survival. In addition, the size change of
c may lead to the extinction and survival of prey, but it will not affect the long-term survival of predators.
Case III. To further clarify the implications of defense, we first change cooperative defense
and represent it numerically through time series solutions, as illustrated in
Figure 6. In addition, we change the strength of the group defense to
, as shown in
Figure 7, indicating that the ecosystem undergoes the extinction of prey species with a decrease in defense.
Case IV. Now, we increase the intensities of white noise to
,
, and keep all other parameter values as above. In this case, we observe that both the prey and predator are extinct (see
Figure 8).
6. Conclusions
The growth of species in an ecosystem depends on various interactions between species, which increases the complexity of analyzing the dynamics of mathematical models. As far as we know, few people study prey–predator dynamics, taking into account the Smith growth rate, cooperative defense and random environmental disturbance. Therefore, the main purpose of this paper is to investigate how the Smith growth rate and cooperative defense affect the prey–predator dynamics in the stochastic fluctuation environments.
Mathematically, we provide the stochastic dynamics including the existence and uniqueness, the existence of a unique ergodic stationary distribution, and the sufficient conditions for the stochastic extinction. Via numerical simulations, keeping moderate cooperation between prey, when the strength of group defense is low, the prey population moves towards extinction. In that case, if there is a decrease in their cooperation level, the survival time of prey becomes shorter and eventually disappears. Our results established that the increasing defense eventually forces the model to stay in a coexisting state. Therefore, by observing this ecosystem, defense of prey against a predator can ensure their survival, and this behavior indirectly benefits other organisms in the ecosystem, which may have a chain effect on the entire food web. Additionally, increasing the intensities of white noise can cause the coextinction of both predator and prey populations. Due to the sudden disturbance of the ecological environment caused by natural disasters, it is necessary to consider studying Smith growth rate population models with Lévy noise in the future.