Anton S. Zadorin’s research while affiliated with ESPCI Paris and other places

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Publications (23)


Multilevel selection in the evolution of sexual dimorphism in phenotypic plasticity
  • Article
  • Publisher preview available

May 2023

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54 Reads

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2 Citations

Anton S. Zadorin

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Olivier Rivoire

Phenotypes are partly shaped by the environment, which can impact both short-term adaptation and long-term evolution. In dioecious species, the two sexes may exhibit different degrees of phenotypic plasticity and theoretical models indicate that such differences may confer an adaptive advantage when the population is subject to directional selection, either because of a systematically varying environment or a load of deleterious mutations. The effect stems from the fundamental asymmetry between the two sexes: female fertility is more limited than male fertility. Whether this asymmetry is sufficient for sexual dimorphism in phenotypic plasticity to evolve is, however, not obvious. Here, we show that even in conditions where it provides an adaptive advantage, dimorphic phenotypic plasticity may be evolutionarily unstable due to sexual selection. This is the case, in particular, for panmictic populations where mating partnerships are formed at random. However, we show that the effects of sexual selection can be counteracted when mating occurs within groups of related individuals. Under this condition, sexual dimorphism in phenotypic plasticity can not only evolve but offset the twofold cost of males. We demonstrate these points with a simple mathematical model through a combination of analytical and numerical results.

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How sexual dimorphism in phenotypic plasticity may evolve

December 2022

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20 Reads

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1 Citation

Phenotypes are partly shaped by the environment, which can impact both short- term adaptation and long-term evolution. In dioecious species, the two sexes may exhibit different degrees of phenotypic plasticity and theoretical models indicate that such differences may confer an adaptive advantage when the population is subject to directional selection, either because of a systematically varying environment or of a load of deleterious mutations. The effect stems from the fundamental asymmetry between the two sexes: female fertility is more limited than male fertility. Whether this asymmetry is sufficient for sexual dimorphism in phenotypic plasticity to evolve is, however, not obvious. Here, we show that even in conditions where it provides an adaptive advantage, dimorphic phenotypic plasticity may be evolutionarily unstable due to sexual selection. This is the case, in particular, for panmictic populations where mating partners are formed at random. However, we show that the effects of sexual selection can be counteracted when mating occurs within groups of related individuals. Under this condition, sexual dimorphism in phenotypic plasticity can not only evolve but offset the twofold cost of males. We demonstrate these points with a simple mathematical model through a combination of analytical and numerical results.



Exact travelling solution for a reaction-diffusion system with a piecewise constant production supported by a codimension-1 subspace

January 2022

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10 Reads

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1 Citation

Communications on Pure &amp Applied Analysis

A generalisation of reaction diffusion systems and their travelling solutions to cases when the productive part of the reaction happens only on a surface in space or on a line on plane but the degradation and the diffusion happen in bulk are important for modelling various biological processes. These include problems of invasive species propagation along boundaries of ecozones, problems of gene spread in such situations, morphogenesis in cavities, intracellular reaction etc. Piecewise linear approximations of reaction terms in reaction-diffusion systems often result in exact solutions of propagation front problems. This article presents an exact travelling solution for a reaction-diffusion system with a piecewise constant production restricted to a codimension-1 subset. The solution is monotone, propagates with the unique constant velocity, and connects the trivial solution to a nontrivial nonhomogeneous stationary solution of the problem. The properties of the solution closely parallel the properties of monotone travelling solutions in classical bistable reaction-diffusion systems.


Sex as information processing: Optimality and evolution

June 2021

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14 Reads

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3 Citations

PHYSICAL REVIEW E

The long-term growth rate of populations in varying environments quantifies the evolutionary value of processing the information that biological individuals inherit from their ancestors and acquire from their environment. Previous models were limited to asexual reproduction with inherited information coming from a single parent with no recombination. We present a general extension to sexual reproduction and an analytical solution for a particular but important case, the infinitesimal model of quantitative genetics which assumes traits to be normally distributed. We study with this model the conditions under which sexual reproduction is advantageous and can evolve in the context of autocorrelated or directionally varying environments, mutational biases, spatial heterogeneities, and phenotypic plasticity. Our results generalize and unify previous analyses. We also examine the proposal made by Geodakyan that the presence of two phenotypically distinct sexes permits an optimal adaptation to varying environments. We verify that conditions exists where sexual dimorphism is adaptive but find that its evolutionary value does not generally compensate for the twofold cost of males.


Sex as information processing: optimality and evolution

February 2021

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25 Reads

Models of asexually reproducing populations in varying environments have provided insights into the diversity of adaptive strategies, revealed deep analogies with information theory and stochastic thermodynamics, and inspired quantitative methods to analyze single-cell experiments. Here we show how these models can be generalized to sexually reproducing populations. The resulting framework is applied to two fundamental questions. First, what are the conditions under which sexual reproduction is advantageous over asexual reproduction? Second, what are the conditions under which sexual dimorphism, the presence of two phenotypically distinct sexes, is advantageous? We study these questions from the two standpoints of optimal information processing and evolvability. Our analysis provides a new perspective on long-standing problems as well as fresh insights into unexamined questions. More generally, it opens the door to novel interdisciplinary approaches to a central problem of evolutionary biology, the role of sex in adaptation.


Fig. 1. Convergence of IPT in the two-dimensional example (3.2). Left: In the complex λ-plane, RS perturbation theory (RS-PT) converges inside a circle of radius 1/2 (orange line) bounded by the exceptional points ±i/2 where eigenvalues have branch-point singularities and M is not diagonalizable. Dynamical perturbation theory (IPT) converges inside the domain bounded by the blue cardioid, which is larger-especially along the real axis, where there is no singularity. Outside this domain, the map can converge to a periodic cycle, be chaotic or diverge to infinity, following flip bifurcations (along the real axis) and fold bifurcations (at the singularities). The domain where the map remains bounded (black area) is a conformal transformation of the Mandelbrot set. Right: The bifurcation diagram for the quadratic map f along the real λ-axis illustrates the period-doubling route to chaos as λ increases away from 0 (in absolute value). Orange and left vertical lines indicate the boundary of the convergence domains of RS-PT and IPT respectively.
Fig. 8. The convergence domain on the λ-plane for the first column of A (the first eigenvector z 1 ) for the 3 × 3 example. The Mandelbrot-like set (domain where orbits remain bounded) of the iterative scheme is shown in black and grey. The empirical convergence domain is shown in black. Its largest component corresponds to the stability of a steady state (the applicability domain of the iterative method). Small components correspond to stability of various periodic orbits. Various shades of grey show the values of λ that lead to divergence to infinity (the darker the slower the divergence). In red are the values of λ where the matrix is non-diagonalizable.
Fig. 9. Same as Figure 8 for the second eigenvector z 2 (the second column of A). Small components correspond to stability of various periodic orbits. Various shades of grey show the values of λ that lead to divergence to infinity (the darker the slower the divergence).
Fig. 10. Same as Figure 8 for the third eigenvector z 3 (the third column of A) .
A fast iterative algorithm for near-diagonal eigenvalue problems

December 2020

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115 Reads

We introduce a novel iterative eigenvalue algorithm for near-diagonal matrices termed \textit{iterative perturbative theory} (IPT). Built upon a "perturbative" partitioning of the matrix into diagonal and off-diagonal parts, IPT has the complexity of one matrix-vector (when one eigenpair is requested) or one matrix-matrix multiplication (when all eigenpairs are requested) per iteration. Thanks to the high parallelism of these basic linear algebra operations, we obtain excellent performance on multi-core processors and GPUs, with large speed-ups over standard methods (up to 50\sim50x with respect to LAPACK and ARPACK, 5\sim5x with respect to Davidson). When off-diagonal elements are comparable to eigengaps, IPT diverges like the quadratic map outside the Mandelbrot set; for such general matrices IPT can nevertheless be used to refine low-precision eigenvectors obtained by other methods. We give sufficient conditions for linear convergence and demonstrate performance on dense and sparse test matrices.


Figure S1. The convergence domain on the λ-plane for the first line of A (the first eigenvector z 1 ) for the 3 × 3 example. The Mandelbrot-like set (domain where orbits remain bounded) of the iterative scheme is shown in black and grey. The empirical convergence domain is shown in black. Its largest component corresponds to the stability of a steady state (the applicability domain of the iterative method). Small components correspond to stability of various periodic orbits. Various shades of grey show the values of λ that lead to divergence to infinity (the darker the slower the divergence). In red are the values of λ where the matrix is non-diagonalizable.
Figure S2. Same as Fig. S1 for the second eigenvector z 2 (the second line of A). Small components correspond to stability of various periodic orbits. Various shades of grey show the values of λ that lead to divergence to infinity (the darker the slower the divergence).
Figure S3. Same as Fig. S1 for the third eigenvector z 3 (the third line of A) .
Dynamical perturbation theory for eigenvalue problems

February 2020

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355 Reads

Many problems in physics, chemistry and other fields are perturbative in nature, i.e. differ only slightly from related problems with known solutions. Prominent among these is the eigenvalue perturbation problem, wherein one seeks the eigenvectors and eigenvalues of a matrix with small off-diagonal elements. Here we introduce a novel iterative algorithm to compute these eigenpairs based on a reformulation of the eigenvalue problem as an algebraic equation in complex projective space. We show from explicit and numerical examples that our algorithm outperforms the usual Rayleigh-Schr\"odinger expansion on three counts. First, since it is not defined as a power series, its domain of convergence is not a priori confined to a disk in the complex plane; we find that it indeed usually extends beyond the standard perturbative radius of convergence. Second, it converges at a faster slower rate than the Rayleigh-Schr\"odinger expansion, i.e. fewer iterations are required to reach a given precision. Third, the (time- and space-) algorithmic complexity of each iteration does not increase with the order of the approximation, allowing for higher precision computations. Because this complexity is merely that of matrix multiplication, our dynamical scheme also scales better with the size of the matrix than general-purpose eigenvalue routines such as the shifted QR or divide-and-conquer algorithms. Whether they are dense, sparse, symmetric or unsymmetric, we confirm that dynamical diagonalization quickly outpaces LAPACK drivers as the size of matrices grows; for the computation of just the dominant eigenvector, our method converges order of magnitudes faster than the Arnoldi algorithm implemented in ARPACK.


Figure 2. Expected new fraction p as a function of initial fraction p for various replication/selection functions f and for two values of λ. The exact analytical results derived from (1) are shown by solid lines. The linear approximation is shown by a dotted line for the cases both with and without sharing. ∆, the frequency jump at very small initial fraction of active variant, corresponds graphically to the intercept of the curves with the vertical axis. As explained in the text, the theoretical value of ∆ does not depend on the replication function f, allowing the derivation of general equations applicable to many protocols.
Figure 3. SQI calculated for the data in Figure 1. Lines connect data points originating from the same manuscript and done with a single value of λ (indicated next to the line). Disks are used for protocols using screening and bulk (non-monodisperse) emulsions; diamonds, for protocols using screening and microfluidic (monodisperse) emulsion; stars, for protocols using selections in bulk emulsion. The dotted line at SQI = 1 represents the theoretical maximum performance, once the initial fraction and random partitioning have been taken into account. Note that the CSR (Compartmentalized Self-Replication) experiment [8] is off the scale here (see Figure 4).
Figure 4. Compartmentalized self-replication (CSR) protocols are robust against co-encapsulation. (a): SQI calculated for CSR protocols at various λ. Red: original CSR report [8], in polydisperse emulsion. Blue and green: this work, two independent experimental replicates in monodisperse emulsions. The dotted line at SQI = 1 represents the theoretical maximum efficiency. (b): Change in expected frequency jump assuming that droplets containing more than n bacteria are poisoned and do not participate in the reaction. (c): The SQI is recomputed with a cutoff set to n = 2, corresponding to the experimental observation that PCR is quickly poisoned by excess lysate.
Figure 5. Non-Poissonian distributions in compartments. (a): The effect of aggregation of the mutants into pairs or triplets on the theoretical value of ∆ (the achievable frequency jump in one round, valid for low p). Full lines correspond to sharing, while dotted lines indicate no sharing. Red; encapsulation of discrete individuals; blue, encapsulation in pairs; green, encapsulation in triplets. To show the effect of λ relative to the best possible frequency jump in a given aggregation condition (which happens at λ = 0), we plot in inset, the rescaled curves ∆/∆ λ=0 . (b): The effect of polydispersity of the compartments in the case of screenings. Here, we assume that the volumes are Gamma-distributed with mean one and various shape parameters α. The corresponding distributions are shown in the inset.
Quantifying the Performance of Micro-Compartmentalized Directed Evolution Protocols

February 2020

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101 Reads

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6 Citations

Life

High-throughput, in vitro approaches for the evolution of enzymes rely on a random micro-encapsulation to link phenotypes to genotypes, followed by screening or selection steps. In order to optimise these approaches, or compare one to another, one needs a measure of their performance at extracting the best variants of a library. Here, we introduce a new metric, the Selection Quality Index (SQI), which can be computed from a simple mock experiment, performed with a known initial fraction of active variants. In contrast to previous approaches, our index integrates the effect of random co-encapsulation, and comes with a straightforward experimental interpretation. We further show how this new metric can be used to extract general protocol efficiency trends or reveal hidden selection mechanisms such as a counterintuitive form of beneficial poisoning in the compartmentalized self-replication protocol.


Natural selection in compartmentalized environment with reshuffling

September 2019

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76 Reads

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3 Citations

Journal of Mathematical Biology

The emerging field of compartmentalized in vitro evolution, where selection is carried out by differential reproduction in each compartment, is a promising new approach to protein engineering. From practical point of view, it is important to know the effect of the increase in the average number of genotype bearing agents per compartment. This effect is also interesting on its own in the context of primordial evolution in the hypothetical RNA world. The question is important as genotypes with different phenotypes in the same compartment share their fitness (the number of produced copies) rendering the selection frequency-dependent. We carried out a theoretical investigation of this problem in the context of selection dynamics for a simple model with an infinite population that is periodically redistributed among infinite number of identical compartments, inside which all molecules are copied without distinction with the success rate as a function of the total genomic composition in the compartment. Using the obtained update equation, we demonstrated that, for the linear additive fitness function, the best genotype is still selected regardless of the average number of individuals per compartment. Furthermore, the selection process is slowed down approximately inversely proportional to this number. We also derived more general expressions that cover nonadditive fitness and non-Poissonian distribution among compartments.


Citations (12)


... Such interdependent structures, composed of interacting differentiated elements where the survival of the parts necessitates the existence of the whole, have been observed in many complex biological systems. Examples include the interdependency between replicating molecules (9), that between gametes (10), and that between social insects (11). Their origins are discussed by focusing on the interactions between elements, and the adaptivity of the entire system. ...

Reference:

Formation of human kinship structures depending on population size and cultural mutation rate
Multilevel selection in the evolution of sexual dimorphism in phenotypic plasticity

... To address this question, we may extend our model to treat strategies as variables that are themselves subject to evolution (Exercise 0.5). For the model discussed in this chapter, the results show that strategies that optimize the long-term growth rate are indeed evolutionarily stable (but this is no longer necessarily the case when considering, for instance, sexually reproducing populations [19]). ...

How sexual dimorphism in phenotypic plasticity may evolve
  • Citing Preprint
  • December 2022

... This refinement algorithm also leads to a mixed precision nonsymmetric eigensolver. There is another recent algorithm called iterative perturbative theory (IPT) developed by Kenmoe, Kriemann, Smerlak, and Zadorin in [18] that works for both symmetric and nonsymmetric eigenvalue problems. ...

A Fast Iterative Algorithm for Near-Diagonal Eigenvalue Problems
  • Citing Article
  • November 2022

SIAM Journal on Matrix Analysis and Applications

... This kind of model also occurs in the cases when the productive part of the reaction happens only on a surface in space or on a line on plane, but the degradation and the diffusion happen in bulk are important for modeling various biological processes (see [2] and more recently [22]). ...

Exact travelling solution for a reaction-diffusion system with a piecewise constant production supported by a codimension-1 subspace
  • Citing Article
  • January 2022

Communications on Pure &amp Applied Analysis

... Similarly, a meta-analysis of data on sex-specific plasticity in response to thermal adaptation revealed that of seven categories of traits, one showed a significant difference in plasticity between the sexes, namely cold resistance, which appears to be more plastic in females [4]. Only recently, however, has the question of the adaptive value of these sex differences started to be studied mathematically [4,5]. Remarkably, these studies show that sexual dimorphism in phenotypic plasticity can enhance population growth and thus prevent the extinction of populations subject to directional selective pressures. ...

Sex as information processing: Optimality and evolution
  • Citing Article
  • June 2021

PHYSICAL REVIEW E

... Their massive generation rate (≈100 Hz-10MHz (ref. 1 and 2)), their monodispersity (a few %) and their minute volume (≈1-1000 pL) make droplets ideal reactors for applications that need highthroughput, low consumption of reagents and quantitativeness. Droplets microfluidics has become routine in single-cell analysis, [3][4][5][6] digital Polymerase Chain Reaction (PCR) [7][8][9] or directed evolution, [10][11][12][13][14] where it is used to digitally encapsulate oligonucleotides, plasmids or cells. Droplets are also finding applications in other fields, ranging from chemical synthesis 2,15,16 and nonlinear chemistry, [17][18][19][20] enzymology, 14 drug screening 21 and toxicology, 22 to microbiology, 23 cell biology, and tissue engineering. ...

Quantifying the Performance of Micro-Compartmentalized Directed Evolution Protocols

Life

... Nevertheless, RCR coupled with recombination still results in a mixture of different DNA products with only a trace amount of monomeric circular DNA [46][47][48] . The dominance of (possibly non-clonal) concatemers results in a reduced enrichment efficiency 49 leading to the accumulation of inactive variants in the replicating DNA population 50,51 . Alternatively, additional DNA processing steps would be required between each round of evolution to restore the original DNA structure. ...

Selection strategies for randomly partitioned genetic replicators
  • Citing Article
  • June 2019

PHYSICAL REVIEW E

... In particular, this means that a genotype always produces the same phenotype, whatever the other genotype it may be encapsulated with. Non-additive phenotype effects, which could arise, for example, from competition between co-compartmentalized genotypes for the phenotype-expression machinery, are explored in [21]. ...

Natural selection in compartmentalized environment with reshuffling

Journal of Mathematical Biology

... Such robots can take the form of DNA origami that selfassemble into complex 3D nanostructures, able to connect to each other or change configuration depending on biochemical cues [72,73,74,75,76,77,78,79]. Simpler structures can also be programmed to move on tracks [80] and sort cargoes at the nanoscale [81]. Coating beads with DNA allows to create micro-robots with higher computing capabilities, with reaction-diffusion serving to form both controllers and signals [82,83,84]. Another emerging field is controllable active matter, where self-propelled agents process chemical signals locally, leading to self-organization [85,86,87,88,89,90,91]. ...

Microscopic agents programmed by DNA circuits
  • Citing Article
  • January 2017

Nature Nanotechnology

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A S Zadorin

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J.-C. Galas

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[...]

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Y Rondelez

... This is crucial for them to adopt fates that are appropriate for their location. Beyond biological systems, Wolpert's idea has also been realised using synthetic soft materials [5][6][7][8] where individual components such as bistable networks read out the morphogen gradient in a microchannel. This has facilitated improved controllability in experiments, making it suitable for testing new theoretical concepts. ...

Synthesis and materialization of a reaction-diffusion French flag pattern
  • Citing Article
  • January 2017

Nature Chemistry