Research article Special Issues

Incompressible limit of a continuum model of tissue growth with segregation for two cell populations

  • This paper proposes a model for the growth of two interacting populations of cells that do not mix. The dynamics is driven by pressure and cohesion forces on the one hand and proliferation on the other hand. Contrasting with earlier works which assume that the two populations are initially segregated, our model can deal with initially mixed populations as it explicitly incorporates a repul-sion force that enforces segregation. To balance segregation instabilities potentially triggered by the repulsion force, our model also incorporates a fourth order diffusion. In this paper, we study the influ-ence of the model parameters thanks to one-dimensional simulations using a finite-volume method for a relaxation approximation of the fourth order diffusion. Then, following earlier works on the single population case, we provide formal arguments that the model approximates a free boundary Hele Shaw type model that we characterise using both analytical and numerical arguments.

    Citation: Alina Chertock, Pierre Degond, Sophie Hecht, Jean-Paul Vincent. Incompressible limit of a continuum model of tissue growth with segregation for two cell populations[J]. Mathematical Biosciences and Engineering, 2019, 16(5): 5804-5835. doi: 10.3934/mbe.2019290

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  • This paper proposes a model for the growth of two interacting populations of cells that do not mix. The dynamics is driven by pressure and cohesion forces on the one hand and proliferation on the other hand. Contrasting with earlier works which assume that the two populations are initially segregated, our model can deal with initially mixed populations as it explicitly incorporates a repul-sion force that enforces segregation. To balance segregation instabilities potentially triggered by the repulsion force, our model also incorporates a fourth order diffusion. In this paper, we study the influ-ence of the model parameters thanks to one-dimensional simulations using a finite-volume method for a relaxation approximation of the fourth order diffusion. Then, following earlier works on the single population case, we provide formal arguments that the model approximates a free boundary Hele Shaw type model that we characterise using both analytical and numerical arguments.


    Nonlinear differential equations (NDEs) play an important fundamental role not only in mathematics and physics, but also in other fields of science in describing nonlinear natural phenomena and processes. The theories and practical applications of NDEs have led to the need to develop methods for solving and investigating such equations for more than two centuries. This led to the discovery of the method of the inverse scattering transform (IST) in 1967. It is now rather developed, well established, and well known as soliton theory [1,2,3]. The main representatives of soliton theory born more than half a century ago, namely the Korteweg-de Friese equation, the nonlinear Schrödinger equation, the sine-Gordan equation, and the Heisenberg model, have become more popular due to the clarity with which they explain a wide range of complex physical phenomena and display dynamical regularities using localized wave solutions [4,5]. Integrable multidimensional extensions of soliton models have attracted considerable interest not only in theoretical and mathematical physics, but also in otsher sections of nonlinear sciences [6,7,8]. The methods of their investigation have also been extended. Thus, in [9], the (2+1)-dimensional nonlinear damped Klein-Gordon equation is studied using the classical method of Lie symmetry. Closed wave solutions of the (4+1)-dimensional fractional Davy-Stewartson-Kadomtsev-Petviashvili equation are studied using the modified auxiliary equation method and the method of elliptic Jacobi functions in [10]. The running solution of the generalized Zakharov system is obtained in [11]. The authors of [12,13,14] devote their research to equations with fractional order derivatives, which, in turn, provide a more general framework for modeling systems with nonlocality, memory, and anomalous behavior. In the recent work of the authors [15], integrable systems with self-consistent potentials were proposed. The works of the authors [16,17,18] are devoted to the study of integrable systems mentioned in [15]. One of the representatives of nonlinear Lax integrable equations (admitting a Lax pair with a non zero spectral parameter) is the Nurshuak-Tolkynay-Myrzakulov (NTM) system [15]. This system belongs to the theory of solitons and describes the dynamic interaction of different fields, where nonlinear effects lead to the formation of stable waves that are capable of propagating over large distances without changing their shape. The purpose of this paper is to give a geometric interpretation of the NTM system and to construct its soliton solutions. The task of establishing a geometric connection between the NTM system and the equation of motion of spatial curves is accomplished by applying the Serret-Frenet equation for curvature and torsion, which describes the properties of curves and surfaces in differential geometry. In it, curvature defines how a curve or surface is curved in space, and torsion is related to how a curve twists around its axis [19,20,21]. These geometric characteristics can be related to the physical properties described by the NTM equation. One-soliton solutions of the NTM system are obtained using the traveling wave method [22,23,24], which allows the efficient derivation of exact solutions in explicit form without resorting to the time-consuming analytical techniques typical of the classical methods of soliton theory.

    This paper is organized as follows: Section 2 presents the NTM equation, its various reductions, and their Lax representations (LRs). The geometric formulation of this equation in terms of space curves is presented in Section 3. In Section 4, soliton solutions of the NTM equation are obtained. We discuss and draw conclusions from our results in Section 5.

    In this paper, the object of study is a previously unstudied system of nonlinear equations called the NTM system. The authors present, for the first time, a geometrical interpretation of this system. In particular, a method of finding the Lax pair for this nonlinear system based on the equation of motion of spatial curves is proposed.

    The NTM equation has the form

    qt+qxxtvq(wq)x=0, (2.1)
    rt+rxxt+vr(wr)x=0, (2.2)
    vx+2(rxtqrqxt)=0, (2.3)
    wx2(rq)t=0, (2.4)

    where q(x,t) and r(x,t) are complex functions.

    The corresponding Lax representation of Eqs (2.1)–(2.4) are given by

    Φx=UΦ, (2.5)
    Φt=VΦ, (2.6)

    where

    U=iλσ3+Q,V=114λ2{λB1+B0}.

    Here

    B1=iwσ3+2iσ3Qt,Q=(0qr0),B0=12vσ3+(0qxt+wqrxt+wr0).

    The compatibility condition of the systems (2.5) and (2.6)

    UtVx+[U,V]=0

    gives the NTM equations (2.1)–(2.4).

    Next, we present some special cases of the NTM equations (2.1)–(2.4).

    The NTM equations (2.1)–(2.4) admit some exact integrable reductions, such as the following cases: r=ϵˉq, r=ν, and r=ϵq, where ϵ=±1.

    First, we consider the reduction r=ϵˉq, where ϵ=±1. Then the NTME equations (2.1)–(2.4) take the form

    qt+qxxtvq(wq)x=0, (3.1)
    vx+2ϵ(ˉqxtqˉqqxt)=0, (3.2)
    wx2ϵ(|q|2)t=0. (3.3)

    Let q(x,t) be a complex function. We now consider the reduction r=ϵq, where ϵ=±1. Then the NTME equations (2.1)–(2.4) take the form

    qt+qxxt(wq)x=0,wx2ϵ(q2)t=0.

    Now we present the case when q(x,t) is a real function and the reduction r=ϵq, where ϵ=±1. Then the NTM equations (2.1)–(2.4) will take the same form as in the case when q(x,t) is a complex function.

    Let q(x,t) be a real function and assume the reduction r=ν, where ν is a constant. Then the NTM equations (2.1)–(2.4) take the form

    qt+qxxtvq(wq)x=0, (3.4)
    vx2νqxt=0, (3.5)
    wx2νqt=0. (3.6)

    Finally, we note that the NTM-III equations (3.4)–(3.6) can be rewritten in the following equivalent forms:

    qt+qxxt+(2νk1qt+μ2)q(wq)x=0,wx2νk2qt=0,

    or

    qt+qxxt+(2νμ1qt+μ2)qwqx=0,wx2νk2qt=0,

    where μj represents constants. Note that for the NTM-III equation, the q(x,t) is a real function.

    All the special cases of the NTM equations (2.1)–(2.4) above admit Lax pairs.

    The aim of this section is to present the geometric formulation of the NTM equation in terms of curves and to find its geometrical equivalent counterpart.

    We start from the differential geometry of space curves [25,26,27]. In this section, we consider the integrable motion of space curves induced by the NTM equation. As usual, we use a smooth space curve γ(x,t):[0,X]×[0,T]R3 in R3. Let x be the arc length of the curve at each time t. In differential geometry, such a curve is given by the Serret-Frenet (SF) equation. The SF equation and its temporal counterpart look like

    (e1e2e3)x=C(e1e2e3),(e1e2e3)t=G(e1e2e3), (4.1)

    where ej represents the unit tangent vector (j=1), the principal normal vector (j=2), and the binormal vector (j=3) which are given by e1=γx,e2=γxx|γxx|,ande3=e1×e2, respectively. Here, × means the vector product. In this case

    C=(0κ1κ2κ10τκ2τ0)=τL1+κ2L2κ1L3so(3),G=(0ω3ω2ω30ω1ω2ω10)=ω1L1+ω2L2ω3L3so(3),

    where τ, κ1,κ2 are the torsion, geodesic curvature, and normal curvature of the curve, respectively; ωj are some functions. Here, we must note that in the standard case, τ,κj,ωj are some real functions of (x,t). However, in this paper, we assume that, in general, these functions are complex.

    Theorem 1. If the elements of the matrices U,Vsu(2) are related to the elements of the matrices C,Dso(3) of the SF in Eq (4.1) in the form

    κ1=2λ,κ2=rq,τ=i(r+q),

    and

    ω1=i(rxt+qxtwrwq),ω2=rxt+qxt+wrwq,ω3=iv,

    then the matrices U,V are the Lax pair of the NTM system (2.1)–(2.4) in the case where λ=0.

    Proof. We underline Lj are basic elements of so(3) algebra and have the forms

    L1=(000001010),L2=(001000100),L3=(010100000).

    They satisfy the following commutation relations:

    [L1,L2]=L3,[L2,L3]=L1,[L3,L1]=L2.

    Further, we also need the basic elements of algebra su(2)

    e1=12i(0110),e2=12i(0ii0),e3=12i(1001),

    where the Pauli matrices have the form

    σ1=(0110),σ2=(0ii0),σ3=(1001).

    These elements satisfy the following conditions

    [e1,e2]=e3,[e2,e3]=e1,[e3,e1]=e2.

    Note that the Pauli matrices obey the following commutation relations:

    [σ1,σ2]=2iσ3,[σ2,σ3]=2iσ1,[σ3,σ1]=2iσ2,

    or

    [σi,σj]=2iϵijkσk.

    The well-known isomorphism between the Lie algebras su(2) and so(3) implies the following correspondence between their basic elements Ljej. Using this isomorphism, let us construct the following two matrices:

    U=τe1+κ2e2κ1e3=12i(κ1τ+iκ2τiκ2κ1)=(u11u12u21u11),V=ω1e1+ω2e2ω3e3=12i(ω3ω1+iω2ω1iω2ω3)=(v11v12v21v11).

    Hence, we obtain

    κ1=2iu11,κ2=(u12u21),τ=i(u12+u21),ω1=i(v12+v21),ω2=(v12v21),ω3=2iv11.

    The compatibility condition of the Eq (4.1) reads as

    CtGx+[C,G]=UtVx+[U,V]=0,

    or, in elements

    κ1tω3xκ2ω1+τω2=0, (4.2)
    κ2tω2x+κ1ω1τω3=0, (4.3)
    τtω1xκ1ω2+κ2ω3=0. (4.4)

    Now, we suggest that the functions κ1,κ2,τ,ω1,ω2,ω3 have the forms

    κ1=2λ,κ2=rq,τ=i(r+q), (4.5)

    and

    ω1=i14λ2(2iλqt2iλrtrxtqxt+wr+wq), (4.6)
    ω2=114λ2(2iλrt2iλqtrxt+qxt+wrwq), (4.7)
    ω3=2i14λ2(0.5viλw), (4.8)

    respectively. Here, λ is a complex constant, and r,q,v,w are some functions of (x,t). In the case when λ=0, Eqs (4.5)–(4.8) take the forms

    κ1=0,κ2=rq,τ=i(r+q), (4.9)

    and

    ω1=i(rxt+qxtwrwq), (4.10)
    ω2=rxt+qxt+wrwq, (4.11)
    ω3=iv. (4.12)

    Substituting the expressions (4.9)–(4.12) into Eqs (4.2)–(4.4) give us the following equations for the functions r,q,v,w:

    qt+qxxtvq(wq)x=0,rt+rxxt+vr(wr)x=0,vx+2(rxtqrqxt)=0,wx2(rq)t=0.

    It is well-known that this is the NTM equation.

    Thus, the SF equation for curvature and torsion (4.1) depends on the functions r,q,v, and w, which are solutions of the NTM equation. Parametrization of curvature and torsion through these functions shows that changes in the dynamics described by the NTM equation directly affect the geometric properties of curves and surfaces.

    As an integrable equation, the NTM equation has all ingredients of integrable systems like Lax representation, conservation laws, bi-Hamiltonian structure, soliton solutions, and so on [28,29,30]. In particular, it admits the N-soliton solutions. In this section, we want to find the 1-soliton solution of the NTM equation. Namely, we will look for the 1-soliton solution of the following NTM-I equations (3.1)–(3.3):

    qt+qxxtvq(wq)x=0, (5.1)
    vx+2ϵ(ˉqxtqˉqqxt)=0, (5.2)
    wx2ϵ(|q|2)t=0. (5.3)

    To find soliton solutions, we use the following complex transformation:

    q=ϕ(y)eiθ,v=χ(y),w=ρ(y),

    where

    y=βx+νt+y0,θ=αx+γt+θ0,

    and α,β,ν,γ,y0,θ0 are some real constants. We have

    qx=(βϕ+iαϕ)eiθ, (5.4)
    qt=(νϕ+iγϕ)eiθ, (5.5)
    qtx=[β(νϕ+iγϕ)+iα(νϕ+iγϕ)]eiθ, (5.6)
    qtxx=[β2(νϕ+iγϕ)+iαβ(νϕ+iγϕ)+iαβ(νϕ+iγϕ)α2(νϕ+iγϕ)]eiθ, (5.7)
    ˉqqxt=ϕ[β(νϕ+iγϕ)+iα(νϕ+iγϕ)]=ϕ[βνϕαγϕ]+iϕ[βγϕ+ανϕ], (5.8)
    ˉqqxtˉqxtq=2iϕ[βγϕ+ανϕ], (5.9)
    vx=βv=4iϵϕ[βγ+αν]ϕ, (5.10)
    wx=βw=2ϵν(ϕ2). (5.11)

    Hence, we obtain

    v=2iϵβ(βγ+αν)ϕ2+v0, (5.12)
    w=2ϵνβϕ2+w0, (5.13)

    where v0 and w0 are some constants. Below, we assume that v0=w0=0. Inserting Eqs (5.4)–(5.13) into Eq (5.1) and splitting the real and imaginary parts yields

    νϕ+β2νϕ2αβγϕα2νϕ6ϵνϕ2ϕ=0,γϕ+β2γϕ+αβνϕ+αβνϕα2γϕ2ϵναβϕ32ϵβ(βγ+αν)ϕ3=0,

    or

    β2νϕ+(ν2αβγα2ν)ϕ2ϵν(ϕ3)=0, (5.14)
    (β2γ+2αβν)ϕ+γ(1α2)ϕ2ϵβ(βγ+2αν)ϕ3=0. (5.15)

    Differentiating Eq (5.15), we obtain

    (β2γ+2αβν)ϕ+γ(1α2)ϕ2ϵβ(βγ+2αν)(ϕ3)=0. (5.16)

    Comparing Eqs (5.14) and (5.16), we obtain the following constraints for real constants of the solutions:

    ν2αβγα2νβ2ν=γ(1α2)β2γ+2αβν,2ϵβ2=2ϵβ(β2γ+2αβν)(βγ+2αν).

    Hence, we get

    βγ=ν(α±1),γ=νβ(α±1).

    Note that we have four α,β,ν,γ real constants of the solutions. We can define one of them via the last algebraic equations. The other constants remain free.

    Let us rewrite Eq (5.15) as

    a2ϕ+a1ϕ+a3ϕ3=0, (5.17)

    where

    a2=β2γ+2αβν,a1=γ(1α2),a3=2ϵβ(βγ+2αν).

    Thus, the system of NTM-I equations (5.1)–(5.3) has been reduced to a second-order nonlinear differential equation represented as (5.17). In the following subsections, different types of analytic solutions of the NTM-I equations (5.1)–(5.3) obtained under different conditions of the constant coefficients a1,a2,a3 of Eq (5.17) are found.

    The 1-soliton solution of the NTM-I equations (5.1)–(5.3) is as follows:

    q(x,t)=kcosh(y)eiθ,v(x,t)=iϵk2(βγ+αν)cosh2(y),w(x,t)=2ϵk2βcosh2(y),

    where k is some constant, which is expressed by the four constants α,β,ν,γ as

    a2=a1,k2=2a1a3.

    Figures 13 present the plots of the obtained solution of the NTM-I equations (5.1)–(5.3) for traveling waves that have steady speeds and shapes, where the velocity is V=νβ.

    Figure 1.  (a) The 3D plot and (b) contour plot of the complex function q(x,t) with the parameters ϵ=1,α=0.2,β=0.2,ν=0.3, andγ=0.3.
    Figure 2.  (a) The 3D plot and (b) contour plot of iv(x,t). Parameters: ϵ=1, α=0.2, β=0.2,ν=0.3, and γ=0.3.
    Figure 3.  (a) The 3D plot and (b) contour plot of w(x,t). Parameters: ϵ=1,α=0.2, β=0.2, ν=0.3, and γ=0.3.

    To find the elliptic solutions, let us rewrite Eq (5.17) as

    ϕ+b1ϕ+b3ϕ3+b0=0,

    where

    b1=a1a2,b3=a3a2,b0=const.

    Let us rewrite this equation as

    ϕ+(1+m)ϕ2mϕ3=0, (5.18)

    where

    m=b11=0.5b3.

    So that we have

    b1=10.5b3.

    Equation (5.18) has the following solution:

    ϕ(y)=sn(y,m), (5.19)

    where sn(y,m) is the Jacobi elliptic function. We can rewrite this solution in terms of the Weierstrass elliptic function ϕ(y)=±e1e3(y)e3, where e1=(ω12) and e3=(ω1+ω22). If m=0 and m=1, we obtain particular solutions, specifically, the periodic and kink, respectively. This is the elliptic soliton solution. In Figure 4, we plot this solution for the parameters ϵ=1, α=0.2, β=0.2, ν=0.3, and γ=0.3.

    Figure 4.  (a) The 3D plot and (b) contour plot of the elliptic solution ϕ(y). Parameters: ϵ=1,α=0.2,β=0.2,ν=0.3, and γ=0.3.

    In the case when m=0, we get the periodic solution via Eq (5.19)

    ϕ(y)=sin(y).

    This is the periodic soliton solution. In Figure 5, we plot this solution for the parameters ϵ=1, α=2,β=1,γ=1,μ=1, and m=0.

    Figure 5.  (a) The 3D plot and (b) contour plot of the periodic solution ϕ(y).Parameters: ϵ=1,α=2,β=1,γ=1,μ=1, and m=0.

    In the case when m=1, we get the kink solution via Eq (5.19)

    ϕ(y)=tanh(y). (5.20)

    This is the kink soliton solution. In Figure 6, we plot this solution for the parameters ϵ=1, α=2, β=0.5,γ=1,μ=1, and m=1.

    Figure 6.  (a) The 3D plot and (b) contour plot of the kink solution ϕ(y). Parameters: ϵ=1, α=2,β=0.5,γ=1,μ=1, and m=1.

    It is not difficult to verify that the Eq (5.18) has the following rational solution:

    ϕ(y)=ky,

    where k is a real constant. This is the rational soliton solution. In Figure 7, we plot this solution for the parameters ϵ=1, α=0.2,β=0.2,ν=0.3, and γ=0.3.

    Figure 7.  (a) The 3D plot and (b) contour plot of the rational solution ϕ(y). Parameters: ϵ=1,α=0.2,β=0.2,ν=0.3, and γ=0.3.

    In the present work, the one-soliton solutions of the NTM system are obtained using the traveling wave method. The advantage of this approach is the possibility of obtaining the one-soliton solution of integrable systems in explicit form without the need for complex analytical methods.

    In conclusion, we note that the integrability of nonlinear partial differential equations is an important topic that combines both theoretical and practical aspects, and plays a key role in understanding and describing complex dynamic systems.

    It is known that the SF equation for curvature and torsion describes the properties of curves and surfaces in differential geometry. Thus, curvature determines how a curve or surface is curved in space, and torsion is related to how the curve twists around its axis. These geometric characteristics can be related to the physical properties described by the NTM equation.

    Both types of equations (NTM and SF equations) exhibit nonlinear interactions. In the NTM equation, the nonlinear terms can be related to changes in curvature and torsion, indicating that the dynamics of solitons can affect the geometry of space.

    Thus, the NTM system not only describes the dynamics of waves, but also allows us to understand how these waves interact with the geometry of space, which is an important aspect of many physical processes.

    In this paper, we propose a method of constructing a Lax pair for a system of nonlinear partial differential equations based on consideration of the equation of motion of a spatial curve. The key link between these two lines of research is the isomorphism between the Lie algebras so(3) and su(2).

    The NTM system (2.1)–(2.4) is integrable in the sense of Lax, which guarantees the existence of n-soliton solutions. In this paper, one-soliton solutions are obtained by means of traveling waves, but this method turns out to be insufficient for constructing multidimensional or n-soliton solutions. To obtain more general solutions of the NTM system (2.1)–(2.4), classical methods of soliton theory, such as Hirota's direct method, the Darboux transform, and other approaches based on the integrable structure of the system, can be applied in the future.

    The formulation of the problems and the derivation of integrable reductions of the equation in the study were carried out by R. Myrzakulov; G. Nugmanova and A. Azhikhan contributed to establishing the connection between the NTM and SF equations. Exact solutions of the equations were found by A. Myrzakul.

    The authors declare they have not used Artificial Intelligence (AI) tools in the creation of this article.

    We would like to thank the anonymous reviewers for their valuable comments and suggestions, which significantly improved the manuscript.

    This work was supported by the Ministry of Science and Higher Education of the Republic of Kazakhstan, Grant No. AP22683222.

    The authors declare no conflict of interest.



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