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HAL Id: hal-03013229

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Submitted on 11 Nov 2021

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Evolution of dietary diversity and a starvation driven cross-diffusion system as its singular limit

Elisabetta Brocchieri, Lucilla Corrias, Helge Dietert, Yong-Jung Kim

To cite this version:

Elisabetta Brocchieri, Lucilla Corrias, Helge Dietert, Yong-Jung Kim. Evolution of dietary diversity and a starvation driven cross-diffusion system as its singular limit. Journal of Mathematical Biology, Springer Verlag (Germany), 2021, �10.1007/s00285-021-01679-y�. �hal-03013229v2�

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Evolution of dietary diversity and a starvation driven cross-diffusion system as its singular limit

E. Brocchieri1, L. Corrias1, H. Dietert2, Y.-J. Kim3

11th November 2021

Abstract

We rigorously prove the passage from a Lotka-Volterra reaction- diffusion system towards a cross-diffusion system at the fast reaction limit. The system models a competition of two species, where one spe- cies has a more diverse diet than the other. The resulting limit gives a cross-diffusion system of a starvation driven type. We investigate the linear stability of homogeneous equilibria of those systems and rule out the possibility of cross-diffusion induced instability (Turing instability). Numerical simulations are included which are compatible with the theoretical results.

Keywords.Cross-diffusion, starvation-driven diffusion, entropy, Turing instability.

2010 Mathematics Subject Classification. Primary : 35B25, 35B40, 35K57, 35Q92, 92D25. Secondary 35B45, 35K45

1 Introduction

1.1 Problem setting

We consider a semilinear reaction-diffusion system that models a competition dy- namics when two species have partially different diets. The population densities of the two species are denoted by u =u(t, x) and v = v(t, x). The species uhas a more diverse diet and is divided into two substatesua=ua(t, x) andub=ub(t, x) so thatu=ua+ub. The system is parametrized by a small parameter ε >0 and written as

tuεa=da∆uεa+fa(uεa) +1

εQ(uεa, uεb, vε), in (0,+)×Ω,

tuεb =db∆uεb+fb(uεb, vε)1

εQ(uεa, uεb, vε), in (0,+)×Ω,

tvε=dv∆vε+fv(uεb, vε), in (0,+)×Ω,

(1.1)

where ΩRN, N1, is a bounded domain with a smooth boundary, and da, db

and dv are diffusivities for the three populations. The unknown solutions depend

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on the parameter ε and we denote it explicitly if needed. The above system is complemented with nonnegative initial data

uεa(0, x) =uina(x), uεb(0, x) =uinb(x), vε(0, x) =vin(x), x, (1.2) and zero flux boundary conditions,

dauεa·σ=dbuεb·σ=dvvε·σ= 0, on (0,+)× ∂Ω, (1.3) whereσdenotes the outward unit normal vector on the boundary∂Ω.

In this paper, we explore the effect of diet diversity in a competition context and show the emergence of cross-diffusion triggered by the different substates ua

and ub, as ε 0. The competition dynamics is given in the reaction terms. The reaction terms of order one are given by

fa(ua) :=ηaua

1ua

a , fb(ub, v) :=ηbub

1ub+v b

, (1.4)

fv(ub, v) :=ηvv

1ub+v b

,

wherea, b >0 are carrying capacities supported by two different groups of resources andηa, ηb, and ηv >0 are the intrinsic growth rates ofua, ub, andv, respectively.

The competition of the two species, u and v, is for the resource b. However, the species u has a diverse diet and can survive by consuming the other resource a without competition. To model such a competition using a Lotka-Volterra type system, the speciesu is divided into two substatesua and ub depending on their diets. In the above reaction terms,uatakes a logistic equation type reaction, andub

andv take Lotka-Volterra competition equations type reactions as given in (1.4).

Since competition exists only partially for the species u, the competition is weak foru. However, the speciesvcompetes withufor all of its resources and hence the competition is not weak in general and the competition result may depend on the parameterε(see Sections4 and5).

The individuals of the speciesumay freely change the type of food depending on the availability, which is modelled by the fast reaction term of orderε−1,

1

εQ(ua, ub, v) := 1 ε

φub+v b

ubψua

a ua

, ε >0. (1.5) In this reaction term,ǫ−1φub+v

b

is the conversion rate for individuals in the stateub which switch to the other stateua, and ǫ−1ψua

a

is the conversion rate in the other direction. The conversion rateφ ubb+v

is assumed as a function of the starvation measure ubb+v for the populationsubandv. If the resourcebdwindles or the populationub+vincreases, the resourceb becomes scarce relatively, and more individuals of populationub will convert to ua and consume the other resourcea.

Hence, we assume thatφis an increasing function of the starvation measure (see [22]

for more discussion on the starvation measure). In the same way, the conversion rate ψ is a function of the starvation measure uaa for the population ua and is assumed to be increasing. For this reason, it makes sense to call the conversion dynamics given by (1.5) a starvation-driven conversion, which eventually results in the starvation-driven cross-diffusion after taking the limitε0 (see [8,9]). More specifically, we assume the following starvation-driven conversion hypothesis

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(H1) φand ψin (1.5) are increasing functions belonging toC1([0,+)); in addi- tion, there exist strictly positive constantsδψ, δφ, Mφ, andMψ such that, for allx0,

ψ(x)δψ>0, φ(x)δφ>0, φ(x)Mφ, and ψ(x)Mψ. The main result of the paper is that, asε0, the (unique) solution (uεa, uεb, vε) of the initial boundary value problem (1.1)–(1.5) converges to a limit (ua, ub, v) and this limit is a weak solution of the reaction cross-diffusion system

(tu= ∆(daua+dbub) +fa(ua) +fb(ub, v), in (0,+)×Ω,

tv=dv∆v+fv(ub, v), in (0,+)×Ω, (1.6) whereua andub are (uniquely) determined by the nonlinear system

ua+ub=u and Q(ua, ub, v) = 0, (1.7) complemented by the initial data,

u(0, x) =uin(x) :=uina(x) +uinb(x), v(0, x) =vin(x), x, (1.8) and the zero flux boundary condition,

(daua+dbub)·σ=dvv·σ= 0, in (0,+)×∂Ω. (1.9) Note that the zero flux boundary conditions in (1.3) are equivalent to the homo- geneous Neumann boundary conditions,

uεa·σ=uεb·σ=vε·σ= 0, on (0,+)×∂Ω

(see [20] for similar diffusion operator for a single species with two phenotypes).

However, after taking the singular limit, we obtain the zero flux boundary conditions (1.9), but not the homogeneous Neumann boundary conditions.

If da = db, the diffusion for the species ugiven in (1.6) is the homogeneous linear diffusion. However, the diffusivity of a species usually depends on its food (or prey) andda 6=db in general. In that case (da 6=db), the diffusion for the total population in (1.6) contains cross-diffusion dynamics depending on the distribution of the three populations groups,ua, ub andv, through the relations in (1.7). This explains the starvation-driven diffusion for the specific case of the paper, a concept formally introduced by Cho and Kim [7]. Funaki et al.[18] derived a macroscopic cross-diffusion model from a system of two phenotypes and a signaling chemical in the context of chemotaxis.

The proof of the convergence as ε 0 is rigorously obtained via a priori estimates foruεa, uεb,andvε. The main tool is the energy (or entropy) functional

E(ua, ub, v) :=

Z

h1(ua)dx+ Z

h2(ub, v)dx , (1.10) where

h1(ua) :=

Z ua

0

ψz a

z dz, and h2(ub, v) :=

Z ub 0

φz+v b

z dz . (1.11)

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Notice here that the assumption (H1) implies thath1 is positive, increasing, and convex, and thath2is positive, increasing in both variables, and convex with respect to the first variable. Therefore, the name entropy for the function given in (1.10) is justified. We refer to [10] and [15] for the use of such entropies in the context of triangular cross-diffusion systems (that is, systems in which only one of the two equations includes a cross-diffusion term). For more general systems, we refer to [5,6,12,14,21,2] among other works.

Then, by invoking the Aubin-Lions Lemma, we pass to the limit along a sub- sequence and conclude that the limit is a weak solution of (1.6)–(1.9). To use the energy estimate, we take initial values with bounded energy, which is our second hypothesis

(H2) uina L1+(Ω), uinb L1+(Ω),vinL+(Ω), andE(uina, uinb , vin)<.

Remark 1. Under Hypothesis (H2), the initial datauina, uinb , vinfor the reaction dif- fusion system(1.1)do not satisfy a priori the nonlinear equationQ(uina , uinb , vin) = 0 in (1.7). Thus, the appearance of an initial layer is expected (see also Section5).

We conclude this introduction proposing a formal derivation of (1.1) out of a microscopic system. We shall consider problems left open here (such as regularity, uniqueness, stability and long time asymptotic behaviour of the macroscopic solu- tions) in a forthcoming paper, where a more general class of cross-diffusion system is analysed.

The rest of the paper is organised as follows.Section2is devoted to the state- ment of the existence result. InSection 3.1, we prove a priori estimates, which are the preliminary ingredients for the proof of the existence result obtained in Sec- tion 3.2. The paper concludes with the existence and linear stability analysis of trivial and non-trivial spatially homogeneous steady states, inSection 4 and Ap- pendix A, with a particular emphasis put on the coexistence state. Some numerical tests inSection5illustrate the linear stability analysis. The discussion inSection6 completes the article.

1.2 Formal derivation of the reaction-diffusion system with fast switching

We explain here how the mesoscopic scale model (1.1) is obtained at a formal level from a microscopic scale model in which the resources inducing the competition explicitly appear. Consider

ts1= 1 δ

hr1s1

1 s1

A1

p1s1U1

i

ts2= 1 δ

hr2s2

1 s2

A2

p2s2U2pVs2Vi

tU1=D1∆U1+k1p1s1U1+1 ε

hΦp2U2+pVV s2

U2Ψp1U1

s1

U1

i

tU2=D2∆U2+k2p2s2U21 ε

hΦp2U2+pVV s2

U2Ψp1U1

s1

U1

i

tV =DV∆V +kVpVs2V,

(1.12)

whereδ >0 is the microscopic reaction time scale andεis the mesoscopic one (hence δε1). These equations describe the time evolution of a small ecosystem with

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two prey population densities (or vegetal resources),s1 and s2, and two predator population densities (or harvesters of the vegetal resources),UandV. Moreover, the populationU is composed of two subpopulationsU1andU2depending on the prey they consume, i.e., s1 ands2, respectively. The prey speciessi follows the logistic dynamics with a carrying capacityAiand an intrinsic growth rateri. The predator species consume a certain amount of preys which is proportional to the prey density with proportionality factorsp1, p2andpV. The harvested prey mass is converted to the predator mass with conversion ratesk1, k2 andkV. The subpopulationsU1and U2convert to each other depending on the availability of the prey. The two functions Φ and Ψ are the conversion rates which are respectively increasing functions of the starvation measures p2U2s+p2 VV and p1sU11. The other species V consumes only the second preys2. Hence, the active competition is only betweenV andU2, whileU1

competes with V passively (via conversion). Finally, since the dispersal rate of a predator species usually depends on the nature of its prey,D16=D2in general.

Remark 2. The expression (1.12) has no diffusion terms for the prey species s1

and s2. Since growth is the dominant factor for plant species and their dispersal is negligible, this is especially relevant when the prey species are vegetal resources.

Mathematically, this choice yields an explicit form in the singular limitsδ0, see (1.13). Adding diffusion terms in the prey species equations could result in a less explicit formulas.

The mescoscopic system with fixedε >0 is obtained in the limitδ0. This is to say that the time-scale for the reaction of the resourcess1ands2is much faster than all other processes. In simple predator-prey models this corresponds to a fast dynamics of the prey which has been studied more carefully in [30,24,28].

In this formal limit δ0, we find s1

r1r1s1

A1 p1U1

= 0 = s1= 0 or s1=A1

1p1U1

r1

, and

s2

hr2 1 s2

A2

p2U2pVVi

= 0 = s2= 0 or s2=A2

1p2U2+pVV r2

. Only the nontrivial case, s1 6= 0 6= s2, is meaningful (since s1 = 0 and s2 = 0 correspond to unstable equilibria), and we obtain two relations

p1U1

s1

=r1

s1 r1

A1

and p2U2+pVV s2

=r2

s2 r2

A2

. Therefore, the last three equations in (1.12) turn into

tU1=D1∆U1+A1k1p1U1 1p1rU11

+1 ε

ΦU2ΨU1

tU2=D2∆U2+A2k2p2U2 1p2U2r+p2 VV

1 ε

ΦU2ΨU1

tV =DV∆V +A2kVpVV 1p2U2r+p2 VV

,

(1.13)

where the conversion rates Φ and Ψ read as Φ = Φr2

s2 r2

A2

and Ψ = Ψr1

s1 r1

A1

,

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and the Lotka-Volterra reaction dynamics of competition type naturally appears.

Now we consider the relationship between the variables in (1.1) and in (1.13).

First, we define

uεa :=U1, uεb:=U2, vε:= pV

p2

V , and keep the same diffusivity coefficients

da :=D1, db:=D2, dv:=DV .

Then, the coefficients in the Lotka-Volterra type competition dynamics,fa, fb and fv, are given as

ηa:=p1A1k1, ηb:=p2A2k2, ηv :=pVA2kV, a:= r1

p1

, b:= r2

p2

. (1.14) Finally, the mesoscopic conversion rates are as follows

φ(x) := Φr2

A2

x 1x

, ψ(x) := Ψr1

A1

x 1x

. (1.15)

After replacing variables, coefficients and functions with the above new ones, system (1.13) becomes our system (1.1).

Remark 3. (i)The conversion rates of the microscopic model, ΦandΨ, are func- tions of the starvation measures p2U2s+p2 VV and p1sU11, instead of simply U2s+V2 and

U1

s1, in order to take into account the difference in the harvesting rates p2 andpV. (ii) The mesoscopic conversion rates φ and ψ in (1.15) are increasing functions, sinceΦandΨare chosen to be increasing functions.(iii)It is worth noticing that the carrying capacitiesaandbfor the predator species are proportional to the growth ratesri’s of the prey species and that the prey carrying capacitiesAi’s are also in- volved in deciding φ and ψ (see (1.14) and (1.15)). (iv) The macroscopic system reduces to the classical Lotka-Volterra system of competition type with linear diffu- sion, whenever the conversion ratesφandψare both constant, (see the discussion section6).

2 Statement of the main result

Before stating our main result in Theorem2.2below, we introduce some notations that will be used in the sequel, and the definition of the very weak solutions of (1.6)–(1.9), with the reaction terms in (1.4).

We denote

Cck:=Cck([0,+)×Ω)¯ :=n

u=u(t, x) :T >0 s.t. uCk [0, T)ׯ

and suppu[0, T)ׯo , and, for allp[1,+),

Lploc:=Lploc((0,+)×Ω) :=n

u=u(t, x) :T >0, uLp(ΩT)o , with ΩT := (0, T)×Ω. Similarly, forp= +,

Lloc:=Lloc((0,+)×Ω) :=n

u=u(t, x) :T >0, ess sup

(t,x)∈ΩT

|u(t, x)|<+o .

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It is worth noticing here that, due to hypothesis (H1), the function q(ub, u, v) :=Q(uub, ub, v) =φub+v

b

ubψuub

a

(uub), (2.1) defined for (ub, u, v)[0, u]×(0,+)×(0,+), satisfies (for givenu >0, v >0)

ubq(ub, u, v) =φub+v b

+ub

b φub+v b

uub

a

+uub

a ψuub

a

>0 and

q(0, u, v)<0, q(u, u, v)>0.

Hence, for any given (u, v)R2+, there exists a uniqueub(u, v)(0, u) zero ofq, and thus a unique solution of the nonlinear system (1.7) is well-defined. Furthermore, the implicit function theorem guarantees the continuity (and even theC1character) ofub with respect to (u, v).

Definition 2.1. Let be a smooth bounded domain of RN,N1. Assumeuin L1+(Ω), and vinL+(Ω) be nonnegative initial densities. We say that the pair of nonnegative functions(u, v)is avery weak solutionof (1.6)–(1.9)over(0,+)× Ω, with reaction terms (1.4), if the following conditions are satisfied

(u, v) belongs toL2loc×Lloc,

for all test functionsξ1, ξ2Cc2,withξ1·σ=ξ2·σ= 0on[0,+)×∂Ω, and forua, ub defined as the unique solution of (1.7), a.e. on (0,+)×Ω, it holds

Z +∞

0

Z

(∂tξ1)u dxdt Z

ξ1(0,·)uindx Z +∞

0

Z

∆ξ1 daua+dbub dxdt

= Z +∞

0

Z

ξ1 fa(ua) +fb(ub, v)

dx dt , (2.2) and

Z +∞

0

Z

(∂tξ2)v dxdt Z

ξ2(0,·)vindxdv

Z +∞

0

Z

∆ξ2v dxdt

= Z +∞

0

Z

ξ2fv(ub, v)dxdt . (2.3)

We observe that all terms in (2.2)–(2.3) are well-defined thanks to the as- sumptions (H2) on the initial densitiesuin, vin, to the L2 integrability of the sub- population densitiesua, uband to theL bound forv. Remember that the logistic structure of the reaction functionsfa, fb, fv involves at most quadratic nonlinear- ities.

Theorem 2.2. Letbe a smooth bounded domain of RN,N1. Assume (H1) and (H2) on parameters and initial datauina , uinb ,vin, respectively. We denote(uεa, uεb, vε) the unique global strong (for t >0) solution of system (1.1)–(1.3) with those ini- tial data. Then, the triplet (uεa, uεb, vε) converges a.e. (t, x)(0,+)× (up to extraction of a subsequence) towards a nonnegative triplet (ua, ub, v), as ε 0.

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Moreover, the functionsu:=ua+ub,v satisfy the nonlinear system (1.7), for a.e.

(t, x) (0,+)×Ω, and the following bounds: u Lq(ΩT) for q = 2 + 2/N if N 3, q < 3 if N = 2 and q = 3 if N = 1; v L(ΩT); |∇u| ∈ L2(ΩT); and for the same previous q, |∇v| ∈ L2q(ΩT); xi,xjv , ∂tv Lq(ΩT), i, j = 1, . . . ,N.

Finally, (u, v) is a very weak solution of the macroscopic system (1.6)–(1.9) with the reaction terms (1.4), in the sense of Definition 2.1.

3 Proof of the main Theorem

We first recall that for any ε >0, there exists a unique global strong (for t >0) solution (uεa, uεb, vε) solution to system (1.1)–(1.3), under the assumption on the initial data of Theorem 2.2. We refer for example to [13, 29] for obtaining such a result.

3.1 A priori estimates

In this section we shall obtain a priori estimates on the subpopulation densities uεa, uεb, on the total population densitiesuε:=uεa+uεbandvε, and onQ(uεa, uεb, vε).

More specifically, we take advantage of the triangular structure of the system that give usa prioriestimates on the densityvεand its derivatives (see Lemma3.1). The reaction functionsfa andfbof competition type allow us to control the total mass R

uε(t)dx, and to get anL2(ΩT) estimate onuε(see Lemma3.2). The latter will be employed in Lemma3.3to obtain estimates onuεa,uεbandQ(uεa, uεb, vε), through the use of the energy functional (1.10)–(1.11). In addition, the triplet (uεa, uεb, vε) will be shown to have finite energyE(T) as well, for allT >0.

Hereafter, all constantsC andCT are strictly positive and may depend on Ω, the initial datauina, uinb, vin, the coefficients in system (1.1), the transition functions φ, ψ and on T, but never on ε. They may change also from line to line in the computations.

Lemma 3.1. Under the hypothesis of Theorem 2.2, the following statements hold:

(i) there exists a constantC >0 such that for all ε >0

kvεkL((0,+∞)×Ω)6C; (3.1) (ii) for all q(1,+)there exists a constantC(q)>0 such that, for all ε >0,

T >0 and all i, j= 1, ..,N,

ktvεkLq(ΩT)+kxi,xjvεkLq(ΩT)6C(q)(1 +kuεbkLq(ΩT)) ; (3.2) (iii) for all q (1,+)there exist C(q,N)>0 andC(q)>0 such that, for all

ε >0and all T >0,

k∇vεk2qL2q(ΩT)C(q,N)(1 +kuεbkqLq(ΩT)) +C(q)T. (3.3) Remark 4.

In the sequel, the value of q in (3.2)–(3.3) will be first chosen equal to 2 (see Lemma3.2), and then to a different number after Corollary 3.4.

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Proof. It is easily seen that 06vε(t, x)6 K:= max

kvinkL(Ω); b , for a.e. (t, x) (0,+)×. (3.4) Indeed, by the existence result of strong solution for (1.1), we know that the nonneg- ativity ofvεis preserved in time. Concerning the upper bound in (3.4), it is obtained by multiplying the equation forvε in (1.1) by (vεK)+ := max{0, vεK} and integrating over Ω, to obtain for allt >0,

Z

(vε(t)K)2+dx Z

(vin,εK)2+dx= 0.

Next, by the maximal regularity property of the heat equation (see [25] and the references therein), for allq(1,+) there exists a strictly positive constantC, which only depends on Ω andq, such that for alli, j= 1, ..,N,

ktvεkLq(ΩT)+kxi,xjvεkLq(ΩT)C(kfv(uεb, vε)kLq(ΩT)+kvinkLq(Ω)

C 1 +kuεbkLq(ΩT)

, (3.5)

so that estimate (3.2) holds. Then, thanks to the Gagliardo-Nirenberg inequality [27], for all q (1,+), there exists C(q) > 0, such that, for all t > 0 and i= 1, . . .N, we have

kxivε(t)kL2q(Ω)C(q)

N

X

j=1

kxi,xjvε(t)k1/2Lq(Ω)kvε(t)k1/2L(Ω)+C(q)kvε(t)kL(Ω). Integrating the above inequality over (0, T) and using (3.1) and (3.5), we get es- timate (3.3).

Lemma 3.2. Under the hypothesis of Theorem 2.2, for all T > 0 there exists CT >0such that for all ε >0 the following estimates hold:

sup

t∈[0,T]

Z

(uεa+uεb)(t)dxCT and kuεa+uεbkL2(ΩT)CT. (3.6) Proof. Adding the first two equations in (1.1) and using the positivity ofuεa, uεb, vε, we get

t(uεa+uεb) da∆uεa+db∆uεb+ηauεa

1uεa a

+ηbuεb

1uεb

b

(3.7)

da∆uεa+db∆uεb+1

4(aηa+b). (3.8)

Then, integrating (3.8) over Ω, the inequality becomes d

dt Z

uεa+uεb

(t)dxC , implying, for allt in [0, T], that

kuεa(t) +uεb(t)kL1(Ω)≤ kuina +uinbkL1(Ω)+C T . (3.9)

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