Abstract
The compartmentalization of distinct templates in protocells and the exchange of templates between them (migration) are key elements of a modern scenario for prebiotic evolution. Here we use the diffusion approximation of population genetics to study analytically the steady-state properties of such a prebiotic scenario. The coexistence of distinct template types inside a protocell is achieved by a selective pressure at the protocell level (group selection) favoring protocells with a mixed template composition. In the degenerate case, where the templates have the same replication rate, we find that a vanishingly small migration rate suffices to eliminate the segregation effect of random drift and so to promote coexistence. In the nondegenerate case, a small migration rate greatly boosts coexistence as compared with the situation where there is no migration. However, increase of the migration rate beyond a critical value leads to the complete dominance of the more efficient template type (homogeneous regime). In this case, we find a continuous phase transition separating the homogeneous and the coexistence regimes, with the order parameter vanishing linearly with the distance to the transition point.
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Acknowledgements
The research of J.F.F. was supported in part by Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) and the research of M.S. was partially supported by PRIN 2009 protocollo n.2009TA2595.02.
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M. Serva is on leave of absence from Dipartimento di Ingegneria e Scienze dell’Informazione e Matematica, Università dell’Aquila, I-67010 Coppito, L’Aquila, Italy.
Appendices
Appendix A: Local Fixation Probability for the M=0 Nonergodic Segregation Regime
As shown by Fontanari and Serva (2013), setting M=0 in Eq. (14) yields two possible steady-state solutions: the solution corresponding to the ergodic coexistence phase, which is a combination of two Delta functions and a regular function, ϕ(x)=A 0 δ(x)+A 1 δ(x−1)+Bϕ r (x) with A 0+A 1+B=1, and the solution corresponding to the nonergodic segregation phase, which is a combination of two Delta functions, ϕ(x)=A 0 δ(x)+A 1 δ(x−1) with A 0+A 1=1. The nonergodic regime, which is our focus here, occurs for C<π 2+S 2/4. Note that in both regimes, A 1 may be interpreted as the probability that the type 1 template fixates in a given protocell and a similar interpretation holds for A 0 as well. However, the result \(\bar{x} = A_{1}\), which we used to draw the curve for M=0 in Fig. 4, holds in the segregation regime only. In Fontanari and Serva (2013) we have calculated the dependence of the weight A 1 on the initial probability density ϕ(x,0) for S=0 only, and in this appendix we generalize that calculation for S≥0.
We begin by rewriting Eq. (9) for M=0,
and introducing the abbreviation \(\langle f (x) \rangle_{\tau}= \int_{0}^{1} f (x) \phi ( x , \tau ) \,dx\) for the expected value of a regular function f(x) at time τ. Hence,
with \(\bar{C} ( \tau ) = C \langle x ( 1- x ) \rangle_{\tau}\). The idea is to choose a function f(x) such that the first three terms of the right-hand side of Eq. (34) cancel out. This choice depends on the value of the parameter Γ≡C−S 2/4, as discussed next. We note that Γ<π 2 in the nonergodic regime.
Region 0<Γ<π 2
In this region we choose \(f (x) = e^{S x/2} \sin ( \sqrt{\varGamma} x + \theta ) \), where θ is an arbitrary constant. Then Eq. (34) rewrites
which has the formal solution
As the right-hand side of this equation does not depend on θ, neither does the ratio in its left-hand side. Hence, equating the ratios evaluated at θ=0 and \(\theta = \pi/2 - \sqrt{\varGamma}/2\) yields
In the limit as τ→∞ we have
and
which leads to
where
In the limit as Γ→π 2, we have Ξ 0→2 regardless of the initial probability density ϕ(x,0), and so \(A_{1} \to A_{1}^{c} = 1/ (1 + e^{S/2} )\). In addition, for the initial probability density ϕ(x,0)=δ(x−1/2) used to calculate \(\bar{x}\) at M=0 in Fig. 4, the dependence on Γ (and hence on C) disappears, and so \(A_{1} = A_{1}^{c}\).
Region −S 2/4<Γ<0
In this region the choice f(x)=e Sx/2(e ux+θe −ux) with \(u = \sqrt{-\varGamma}\) and θ arbitrary leads to the canceling of the first three terms of the right-hand side of Eq. (34), yielding
The same argument used in the analysis of the Γ>0 region allows us to equate the ratio that appear in the left-hand side of this equation for θ=−1 and θ=0,
Finally, taking the limit as τ→∞ yields
where
By taking the limit as u→0 we can easily verify that A 1 is continuous at the boundary of the two regions. In addition, in the limit as C→0, i.e., u→S/2, we recover the classical formula for the fixation of an allele with selective disadvantage S (Crow and Kimura 1970),
Similarly to our finding in the analysis of the previous region, the initial probability density ϕ(x,0)=δ(x−1/2) yields A 1=1/(1+e S/2) regardless of the value of C, as shown in Fig. 4.
Appendix B: Critical Line for the Limit as M→0
Setting M=0 in Eq. (25) yields
with the condition
In the region C−S 2/4>0 its solution is
where \(\gamma= \sqrt{C- S^{2} /4}\), and θ=θ(γ,S) is fixed by condition (48) as
We note that the critical value C c (S) is in the region C−S 2/4>0 (see Fig. 5). To evaluate Eq. (28), we use the equality
which follows directly from the definition \(y_{c} (x) = \int_{0}^{x} z_{c} ( \xi ) d \xi\) with z c given by (49). Now the integrals in Eq. (28) can be readily evaluated, yielding
This equation can be further simplified using the equalities \(\sin ( \gamma + \theta )= S/ ( 2\sqrt{C} )\) and \(\cos (\gamma + \theta )=\gamma/\sqrt{C}\), which follow from Eq. (50). The final result is simply
Finally, we rewrite Eq. (50) as
in order to make clear that Eq. (53) yields a relation C=C c (S), which is the critical line M→0 depicted in Fig. 5.
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Fontanari, J.F., Serva, M. Effect of Migration in a Diffusion Model for Template Coexistence in Protocells. Bull Math Biol 76, 654–672 (2014). https://doi.org/10.1007/s11538-014-9937-7
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DOI: https://doi.org/10.1007/s11538-014-9937-7