Abstract
We study a reaction–diffusion model for population–toxin interactions on a two-dimensional torus, where avoidance is driven by a nonlocal average of toxin-related memory. The variables represent population density, toxin concentration, and a phenomenological information field generated by population–toxin encounters. This information diffuses, decays, and enters the taxis term through convolution with a perceptual kernel. We prove local well-posedness, global existence and uniform boundedness, and give sufficient conditions for exponential convergence to either the positive equilibrium or the toxin-only equilibrium. Numerical simulations with two perceptual radii yield stripe and spot patterns, illustrating that the sensing scale can alter spatial organization. These simulations do not establish an effect on population persistence and should be viewed as hypotheses for empirical study.
MSC:
35K57; 92D40
1. Introduction
1.1. The Model
Toxicants can alter reproduction, mortality, and movement, thereby changing the abundance and spatial distribution of exposed populations [1,2,3,4,5]. Their effects may propagate through interacting populations and, under sustained exposure, contribute to local population loss. Reaction–diffusion models provide a natural setting in which demographic processes, dispersal, and environmental stress can be studied together [6,7,8,9].
Movement need not depend only on conditions at an individual’s current location. Chemical, visual, and social cues can convey information acquired at other locations or earlier times, allowing animals to avoid contaminated regions or locate resources [10,11,12,13,14,15]. Such spatially distributed information motivates models in which movement responds to a nonlocal signal rather than solely to the contemporaneous local toxin concentration [16].
Reaction–diffusion and taxis models have been used to describe contaminated environments. Huang et al. [2] and Shan and Huang [17] analyzed toxin-dependent population interactions, with emphasis on the direct demographic effects of toxicants. Deng et al. [18] and Xing et al. [19] incorporated avoidance of toxicants and studied global dynamics, bistability, and pattern formation. In those systems, movement is driven by the current local toxicant field; there is no separate information state used over a spatial neighborhood.
Keller–Segel-type systems supply much of the analytical background for taxis-driven aggregation and pattern formation [7,20]. Bellomo et al. [20] reviewed mechanisms that prevent blow-up, whereas Xu et al. [21] studied finite-speed propagation under degenerate diffusion. These studies identify difficulties caused by taxis, but they do not include toxin-related memory.
Nonlocal perception and spatial memory appear in several ecological movement models. Potts and Lewis [22], Wang and Salmaniw [16], and Painter et al. [23] considered movement based on spatially averaged information and its role in aggregation or pattern formation. Song et al. [24] and Xue et al. [25] examined detection scales, stability, and bifurcation in cognitive or memory-based systems, while Bo et al. [26] studied bifurcation in a cognitive population model for a toxic environment. The closest analytical precedent is the cognitive consumer–resource model of Zhang et al. [15], for which global solvability and stability were established. Here the resource equation is replaced by an externally supplied toxin that is depleted by the population, and the analysis covers both coexistence and toxin-only equilibria under periodic boundary conditions.
The contribution is therefore specific to the population–toxin feedback considered here: exposure generates an information field, and a nonlocal average of this field drives avoidance. We establish well-posedness and uniform boundedness, derive sufficient conditions and convergence rates for two equilibria, and examine the effect of perceptual radius on the simulated spatial distribution. Periodic boundary conditions remove boundary fluxes from this analysis.
We consider the following reaction–diffusion system on a two-dimensional torus:
where is a torus of side length , with periodic boundary conditions. The variables u, v, and z denote population density, toxin concentration, and toxin-related information, respectively. The convolution is the spatially averaged information used in the taxis term. The parameters r and K are the intrinsic growth rate and carrying capacity; a is the toxin-induced mortality rate; h, , and are the toxin input, decay, and population uptake rates; and b and are the production and decay rates of toxin-related information. The diffusion coefficients of u, v, and z are , , and .
The equation for z is phenomenological. Its source assumes that information is generated when the population and toxin co-occur, with b converting exposure into shared information. The terms and describe spatial transmission and loss of information. This production–transmission–decay structure is consistent with models of memory-dependent movement and collective sensing [16,27,28,29]. Accordingly, z is an aggregate information state, not a directly measured physiological concentration.
The field averages information over a perceptual neighborhood. The term represents movement away from locations with larger perceived toxin information, following the use of nonlocal signals in collective-movement and spatial-memory models [22,30].
More precisely, , where weights information at displacement . We impose the following assumptions on the kernel:
- (H1)
- , with a constant such that .
- (H2)
- There exists such that for all smooth , , or equivalently, with total variation , where is the space of functions of bounded variation.
- (H3)
- , , and for .
- (H4)
- G is compactly supported.
We specifically consider the perceptual kernel
where is the Euclidean distance and is the perceptual range. For some , the initial data satisfy
1.2. Objectives and Challenges
We establish local well-posedness and global boundedness, and then derive sufficient conditions for the global asymptotic stability of the positive and toxin-only equilibria. Numerical simulations are used to examine how the perceptual range changes spatial organization. The analytical results concern the proposed model; claims about persistence or management would require empirical validation.
The principal estimate couples u and z. The population equation contains , while the information equation contains the source . A bound for u therefore requires control of derivatives of , which in turn depends on the regularity available for z. The taxis term also prevents a direct application of the scalar maximum principle to the u-equation. These features require joint energy and semigroup estimates rather than separate estimates for the three components [31].
Two points require particular care:
- (i)
- The taxis term contains , the gradient of a convolution. Higher-order estimates must accommodate the limited regularity , rather than assuming a smooth kernel such as [32].
- (ii)
- Global continuation requires uniform control of the norms in the extensibility criterion. Taxis may concentrate the population even when movement is directed away from toxin, so the proof must combine diffusion, logistic damping, and decay to rule out loss of regularity [24].
The resulting argument combines energy inequalities, heat-semigroup estimates, interpolation, and parabolic regularity.
1.3. Main Results
The first theorem gives global existence and boundedness for (1).
Theorem 1.
The asymptotic analysis concerns the homogeneous equilibria and . Their components satisfy
Eliminating v from the first two equations gives
If , then and . Hence, f has a unique root , namely
The remaining components are
The stability result is stated next.
Theorem 2.
Section 2 proves local existence and uniqueness. The a priori estimates in Section 3 yield global boundedness and Theorem 1. Lyapunov arguments in Section 4 establish Theorem 2, after which Section 5 examines the effect of perceptual range in numerical simulations. The final section summarizes the results and their limitations.
2. Local Existence and Uniqueness of Solutions
We first obtain a local classical solution of (1) by a fixed-point argument and record the corresponding continuation criterion.
Lemma 1.
Proof.
We construct a fixed point, verify that the defining map is contractive for small time, and then give an energy proof of uniqueness.
(i) Existence. Write the nonlinear terms in (1) as
where .
We define the solution space for a time interval as
equipped with the norm
where is determined by the regularity of the initial data . Define the closed subset
Here , and is chosen below the initial infima so that positivity is retained for short time.
We construct the mapping , defined by , where solves
with . We show that when is sufficiently small.
We begin with the u-equation. Since have norms bounded by M, the convolution satisfies
where . In particular, . For the taxis term,
the Hölder product estimates give
It follows that . The reaction terms satisfy
Hence, . The parabolic Schauder estimate gives
Choose and such that , ensuring .
The estimates for v and z are similar. For v,
Thus, . For and sufficiently small T, we ensure .
For z,
The Schauder estimate for yields
Choose and then choose T so that . This gives . The constant may depend on the fixed coefficient .
For small T, continuity from the positive initial data and the strong maximum principle give ; the source in the z-equation is nonnegative.
Hence, for the choices of M and T above.
(ii) Contraction property of . Take two inputs , , with outputs , and define
We estimate the difference of the corresponding outputs in .
Product estimates in Hölder spaces give
The Schauder estimate gives
For v and z, the same argument gives
and the Schauder estimate gives
For z,
and by Schauder estimates,
Combining these estimates gives
Choose T so that . Then is a contraction and has a unique fixed point in . Standard continuation gives a maximal interval and the blow-up criterion (6).
(iii) Uniqueness. Assume that two classical solutions and in have the same initial data. Define
The difference equations are
Multiplying the U-equation by U and integrating by parts gives
For the taxis term,
For the reaction terms,
Young’s inequality gives
Consequently,
Apply Young’s inequality to the final term:
Thus,
For V,
For Z,
Define
Adding the three inequalities yields
Because , Gronwall’s inequality gives on . Hence, , and uniqueness follows. □
We next record the periodic heat-semigroup estimates used in the continuation and boundedness arguments.
Lemma 2.
Consider the heat semigroup on a periodic domain U, where denotes the first nonzero eigenvalue of under periodic boundary conditions and is a constant. There exist constants , depending only on U, such that:
- (i)
- For and any with ,
- (ii)
- For and any , the gradient satisfies:
- (iii)
- For and any , the gradient satisfies:
- (iv)
- For and any , the divergence satisfies:
Remark 1.
The corresponding estimates for the Neumann heat semigroup are proved in [7]. The periodic case follows by the same argument, so we omit the proof.
Lemma 3.
Proof.
Consider the auxiliary equation
with initial condition . The strong maximum principle implies that
The comparison principle gives , and therefore,
□
The next lemma replaces the Hölder-norm criterion in Lemma 1 by bounds on u and the gradients of v and z.
Lemma 4.
Proof.
Suppose, to the contrary, that while the following bounds hold on :
where . We show that these assumptions yield uniform -bounds for , for some . This contradicts the blow-up criterion in Lemma 1:
We use a bootstrap argument.
Testing the z-equation by z gives
Therefore,
Gronwall’s inequality gives
Because , integration over yields
Since by (13), and , the Sobolev embedding for any implies that
for any . The same reasoning, together with the -bound for v, gives
We now improve the regularity of v and z. Set
Since , for every , and , it follows that
For the z-equation, set
The bounds for imply
For , parabolic -regularity [33] implies
By Sobolev embeddings in , for (for large q), so
Similarly, for , we obtain
It remains to improve the regularity of u. Its source term is
The reaction terms are bounded in . For the taxis term,
Since and , the identity holds in the distributional sense. Young’s convolution inequality gives
Thus,
For the remaining product, (H2) gives . Hence,
Together these estimates imply
Duhamel’s formula for u reads
Taking the gradient in (15), applying Lemma 2, and estimating in , we obtain
The bounds for u, v, and z, together with Sobolev embedding, yield
The source terms are therefore Hölder continuous:
since .
For , the reaction terms belong to . The taxis term involves , , and . Since , these quantities have the required Hölder regularity, and
Applying parabolic Schauder estimates [34] to the system (1) with initial data and source terms in , we obtain
for any , with
These uniform -bounds contradict Lemma 1. Therefore, .
□
3. Boundedness and Global Existence
Starting from the continuation criterion in Section 2, we derive the a priori estimates needed to prove that and that the solution remains uniformly bounded.
Lemma 5.
Assume that the hypotheses of Lemma 1 hold. Then there are constants such that
and
where .
Proof.
Integrating the u-equation over the torus gives
Hence,
Therefore,
Let . Then
It follows that
Integrating (21) over gives
Consequently,
The two asserted bounds follow. □
The mass bound for u controls the source in the next estimate for .
Lemma 6.
Proof.
Multiplying the v-equation by v and integrating over U gives
since (from Lemma 3). Integrating over , we get
Using , we obtain
The preceding bounds also control the information variable z.
Lemma 7.
Under the hypotheses of Lemma 1, there are constants such that
Proof.
Integrating the z-equation over U gives
since (Lemma 4), (Lemma 3), and . Define . Then
Gronwall’s inequality gives
Multiplying the z-equation by z gives
where we used . Young’s inequality yields
Consequently,
The time-averaged bound (17) and the uniform Gronwall lemma give
□
Lemma 8.
Proof.
We multiply the third equation of (1) by and integrate over U. Using integration by parts and the periodic boundary conditions, we obtain the first equality below; Young’s inequality gives the subsequent estimate:
Thus
Let and . Then (29) implies
By (17),
The variation-of-constants estimate for (30) yields
since , is finite.
The Gagliardo–Nirenberg inequality then gives
□
We now use the bounds for z to estimate u in .
Lemma 9.
Assume that the initial data satisfy (3) and that the hypotheses of Lemma 1 hold. Then there are constants such that
and
Proof.
For the taxis term, Young’s inequality gives
Hence,
Gronwall’s inequality gives a constant such that
□
An -to- semigroup estimate now yields the gradient bound for v required in Lemma 4.
Lemma 10.
Under the hypotheses of Lemma 1, let solve (1) on the two-dimensional torus U. There is a constant such that
Proof.
Rewrite the v-equation as
where
By (32) and Lemma 3,
The solution to the v-equation is given by the variation-of-constants formula
Applying the gradient estimate in Lemma 2 to the semigroup gives
where is the Gamma function. Thus, (36) holds with
□
We next derive an -bound for u, which is used in the subsequent -estimate.
Lemma 11.
Proof.
Multiplying the first equation of (1) by and applying Young’s inequality and the Gagliardo–Nirenberg inequality, we obtain
which implies
where .
The -bound allows us to estimate u in .
Lemma 12.
Proof.
Rewrite the first equation of (1) as
The variation-of-constants formula gives
Consequently,
By Lemma 2 and (40),
and
Because is dense in for , the – estimate in Lemma 2, together with (40), yields
□
Lemma 13.
Let solve (1) on , with initial data satisfying the hypotheses of Lemma 1. Then there is a constant such that
Proof.
Write the u-equation as
where . Lemmas 3 and 12 give
The variation-of-constants formula and Lemma 2 give
Hölder’s inequality and interpolation yield
Thus, the taxis contribution satisfies
where
Combining the estimates gives
□
Lemma 14.
Assume that the hypotheses of Lemma 1 hold and that satisfies (3). Then there is a constant such that
Proof.
Set . Differentiating the z-equation in space gives
Testing this equation by gives
The diffusion term satisfies
Using , (50), and Hölder’s inequality, we estimate by
Likewise, and Hölder’s inequality give
Combining the estimates yields
Let . Then is a positive constant, and
Young’s inequality gives a constant such that
Substitution gives
Gronwall’s inequality gives
Therefore,
This proves the asserted bound. □
We can now prove Theorem 1.
Proof of Theorem 1.
By Lemma 1, a unique classical solution exists on . According to Lemma 4, it is enough to bound , , and on this interval.
By Lemma 12,
By Lemmas 3 and 10,
Finally, Lemma 14 gives
These estimates satisfy the continuation criterion, so . The solution is therefore global and uniformly bounded in the stated norms. □
4. Stabilization and Convergence Rate
We use Lyapunov functionals to study the positive equilibrium and the toxin-only equilibrium . Barbalat’s lemma gives asymptotic convergence, and comparison of the Lyapunov functionals with suitable norms yields exponential rates.
Lemma 15.
(Barbălat’s lemma [35]). Let be uniformly continuous. If exists, then .
4.1. Stabilization
For the positive equilibrium, define
Let and denote the uniform -bounds from Lemma 12 and Lemma 3, respectively.
Lemma 16.
Assume that
Then there exist constants and such that
where
Proof.
Let
For the term , integration by parts and Young’s inequality give
For ,
For ,
Hence, condition (56) yields constants and such that
□
The dissipation estimate and Barbalat’s lemma first give convergence in ; interpolation then upgrades it to .
Lemma 17.
Proof.
Inequality (57) gives
where
and . Since , integration from to gives
Thus,
Because , the preceding bound implies . Thus, .
Since is uniformly continuous, Barbălat’s lemma (Lemma 15) gives
To obtain -convergence, use the Gagliardo–Nirenberg inequality on the two-dimensional torus:
The relevant -norms are uniformly bounded, while the three -norms tend to zero. Hence,
□
Lemma 18.
Suppose that
Then there are and such that
satisfies
where
Proof.
Differentiating along solutions gives
where
□
Lemma 19.
Proof.
By Lemma 18,
where
where and . Integration from to gives
Since ,
Letting and using , we obtain
so .
The derivative of is bounded, so is uniformly continuous. Barbălat’s lemma therefore yields
The Gagliardo–Nirenberg inequality on the two-dimensional torus gives
Together with the uniform regularity bounds, this implies
Thus, the constant equilibrium is globally asymptotically stable. □
4.2. Convergence Rate
The next lemma strengthens the convergence in Lemma 17 to exponential decay.
Lemma 20.
Under the hypotheses of Lemma 16, there are constants and , independent of t, such that
Proof.
By (57),
where
Define
so (69) implies
The Lyapunov functional is the same one used in Lemma 16:
Since as by Lemma 17, the function is quadratically comparable to for all sufficiently large t. Indeed,
Thus, there exist and constants such that, for ,
Combining the upper bound in (70) with (69) gives
Integration of this differential inequality gives
The lower bound in (70) then gives
On the two-dimensional torus, the Gagliardo–Nirenberg inequality reads
Since are bounded (Lemmas 3 and 12), let . Then,
The same estimate applies to v and z, and hence,
□
Lemma 21.
Under the hypotheses of Lemma 18, there exist constants and , independent of t, such that
Proof.
By (67),
It follows that
which implies
The uniform bounds established above and standard parabolic smoothing [34,36] yield, for ,
Combining this estimate with (77) and the two-dimensional Gagliardo–Nirenberg inequality gives
Equation (77) therefore implies (75), for example with any after increasing the constant C.
□
We now prove Theorem 2.
Proof of Theorem 2.
The conclusion follows from Lemmas 16–21. □
5. Numerical Simulations
We use numerical simulations to compare two perceptual ranges. The kernel is the top-hat function
where is the Euclidean distance on the torus. We take and
The initial data are , , and . These are small perturbations of spatially homogeneous states.
For , the three fields have a striped configuration at (Figure 1). With this shorter averaging range, the population responds mainly to nearby toxin information and forms elongated regions of high and low density. This interpretation is limited to the simulated fields and does not show that stripes improve survival.
For , with all other parameters and initial data unchanged, the fields have a spotted configuration at (Figure 2). The wider kernel incorporates information from a larger neighborhood, and the simulated population is concentrated in separated patches. This comparison shows a change in spatial organization for the chosen parameter set, but it does not establish greater persistence or stability of the spotted state.
A possible interpretation is a lake with localized contamination, where animals use recent or socially transmitted information to avoid polluted regions. The simulations suggest that the spatial scale of this information may affect whether high-density regions form corridors or isolated patches. Testing this interpretation would require field estimates of R, b, and , together with observations linking the predicted patterns to exposure or demographic outcomes.
6. Conclusions
The model couples a population and an environmental toxin to an information field generated by their local encounters. A convolution of this field drives nonlocal avoidance. On the two-dimensional torus, the system is locally well posed and admits a global uniformly bounded solution. Under the stated sufficient conditions, solutions converge exponentially to either the positive equilibrium or the toxin-only equilibrium.
With all other parameters fixed, increasing the perceptual radius from to changes the computed pattern from stripes to spots. This result concerns spatial organization in the model. It does not demonstrate improved persistence, biodiversity, or management effectiveness, and the lake scenario remains an interpretation to be tested empirically.
The information variable and its parameters have not been calibrated against movement or exposure data. Further work could estimate perceptual and memory scales, test the predicted spatial transition, and consider heterogeneous domains, time-dependent kernels, or multispecies interactions [37].
Author Contributions
Conceptualization, X.Z. (Xinyan Zhang) and X.Z. (Xuebing Zhang); methodology, X.Z. (Xinyan Zhang) and X.Z. (Xuebing Zhang); software, X.Z. (Xinyan Zhang); validation, X.Z. (Xinyan Zhang) and X.Z. (Xuebing Zhang); formal analysis, X.Z. (Xinyan Zhang) and X.Z. (Xuebing Zhang); investigation, X.Z. (Xinyan Zhang); writing—original draft preparation, X.Z. (Xinyan Zhang); writing—review and editing, X.Z. (Xinyan Zhang) and X.Z. (Xuebing Zhang); visualization, X.Z. (Xinyan Zhang); supervision, X.Z. (Xuebing Zhang); project administration, X.Z. (Xuebing Zhang); funding acquisition, X.Z. (Xuebing Zhang) All authors have read and agreed to the published version of the manuscript.
Funding
This work was supported by the National Natural Science Foundation of China (No. 12271261) and the National Undergraduate Innovation Training Program (No. 202510300072), “Cognition-Driven Modeling of Population Dynamics and Ecological Regulation Mechanisms.”
Institutional Review Board Statement
Not applicable.
Informed Consent Statement
Not applicable.
Data Availability Statement
The original contributions presented in this study are included in the article. Further inquiries can be directed to the corresponding author.
Conflicts of Interest
The authors declare no conflicts of interest.
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