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Muller's ratchet, in its prototype version, models a haploid, asexual population whose size~N is constant over the generations. Slightly deleterious mutations are acquired along the lineages at a constant rate, and individuals carrying less mutations have a selective advantage. The classical variant considers {\it fitness proportional} selection, but other fitness schemes are conceivable as well. Inspired by the work of Etheridge et al. ([EPW09]) we propose a parameter scaling which fits well to the ``near-critical'' regime that was in the focus of [EPW09] (and in which the mutation-selection ratio diverges logarithmically as N→∞). Using a Moran model, we investigate the``rule of thumb'' given in [EPW09] for the click rate of the ``classical ratchet'' by putting it into the context of new results on the long-time evolution of the size of the best class of the ratchet with (binary) tournament selection, which (other than that of the classical ratchet) follows an autonomous dynamics up to the time of its extinction. In [GSW23] it was discovered that the tournament ratchet has a hierarchy of dual processes which can be constructed on top of an Ancestral Selection graph with a Poisson decoration. For a regime in which the mutation/selection-ratio remains bounded away from 1, this was used in [GSW23] to reveal the asymptotics of the click rates as well as that of the type frequency profile between clicks. We will describe how these ideas can be extended to the near-critical regime in which the mutation-selection ratio of the tournament ratchet converges to 1 as N→∞.
Motivated by the question of the impact of selective advantage in populations with skewed reproduction mechanims, we study a Moran model with selection. We assume that there are two types of individuals, where the reproductive success of one type is larger than the other. The higher reproductive success may stem from either more frequent reproduction, or from larger numbers of offspring, and is encoded in a measure Λ for each of the two types. Our approach consists of constructing a Λ-asymmetric Moran model in which individuals of the two populations compete, rather than considering a Moran model for each population. Under certain conditions, that we call the "partial order of adaptation", we can couple these measures. This allows us to construct the central object of this paper, the Λ−asymmetric ancestral selection graph, leading to a pathwise duality of the forward in time Λ-asymmetric Moran model with its ancestral process. Interestingly, the construction also provides a connection to the theory of optimal transport. We apply the ancestral selection graph in order to obtain scaling limits of the forward and backward processes, and note that the frequency process converges to the solution of an SDE with discontinous paths. Finally, we derive a Griffiths representation for the generator of the SDE and use it to find a semi-explicit formula for the probability of fixation of the less beneficial of the two types.
Motivated by the question of the impact of selective advantage in populations with skewed reproduction mechanims, we study a Moran model with selection. We assume that there are two types of individuals, where the reproductive success of one type is larger than the other. The higher reproductive success may stem from either more frequent reproduction, or from larger numbers of offspring, and is encoded in a measure Λ for each of the two types. Our approach consists of constructing a Λ-asymmetric Moran model in which individuals of the two populations compete, rather than considering a Moran model for each population. Under certain conditions, that we call the ``partial order of adaptation'', we can couple these measures. This allows us to construct the central object of this paper, the Λ−asymmetric ancestral selection graph, leading to a pathwise duality of the forward in time Λ-asymmetric Moran model with its ancestral process. Interestingly, the construction also provides a connection to the theory of optimal transport. We apply the ancestral selection graph in order to obtain scaling limits of the forward and backward processes, and note that the frequency process converges to the solution of an SDE with discontinous paths. Finally, we derive a Griffiths representation for the generator of the SDE and use it to find a semi-explicit formula for the probability of fixation of the less beneficial of the two types.
Motivated by the question of the impact of selective advantage in populations with skewed reproduction mechanisms, we study a Moran model with selection. We assume that there are two types of individuals, where the reproductive success of one type is larger than the other. The higher reproductive success may stem from either more frequent reproduction, or from larger numbers of offspring, and is encoded in a measure Λ for each of the two types. Λ-reproduction here means that a whole fraction of the population is replaced at a reproductive event. Our approach consists of constructing a Λ-asymmetric Moran model in which individuals of the two populations compete, rather than considering a Moran model for each population. Provided the measure are ordered stochastically, we can couple them. This allows us to construct the central object of this paper, the Λ−asymmetric ancestral selection graph, leading to a pathwise duality of the forward in time Λ-asymmetric Moran model with its ancestral process. We apply the ancestral selection graph in order to obtain scaling limits of the forward and backward processes, and note that the frequency process converges to the solution of an SDE with discontinuous paths. Finally, we derive a Griffiths representation for the generator of the SDE and use it to find a semi-explicit formula for the probability of fixation of the less beneficial of the two types.
Therapy evasion – and subsequent disease progression – is a major challenge in current oncology. An important role in this context seems to be played by various forms of cancer cell dormancy. For example, therapy-induced dormancy, over short timescales, can create serious obstacles to aggressive treatment approaches such as chemotherapy, and long-term dormancy may lead to relapses and metastases even many years after an initially successful treatment. The underlying dormancy-related mechanisms are complex and highly diverse, so that the analysis even of basic patterns of the population-level consequences of dormancy requires abstraction and idealization, as well as the identification of the relevant specific scenarios.
In this paper, we focus on a situation in which individual cancer cells may switch into and out of a dormant state both spontaneously as well as in response to treatment, and over relatively short time-spans. We introduce a mathematical ‘toy model’, based on stochastic agent-based interactions, for the dynamics of cancer cell populations involving individual short-term dormancy, and allow for a range of (multi-drug) therapy protocols. Our analysis shows that in our idealized model, even a small initial population of dormant cells can lead to therapy failure under classical (and in the absence of dormancy successful) single-drug treatments. We further investigate the effectiveness of several multidrug regimes (manipulating dormant cancer cells in specific ways) and provide some basic rules for the design of (multi-)drug treatment protocols depending on the types and parameters of dormancy mechanisms present in the population.
Highlights
• We study dormancy in the ‘rare mutation’ regime of stochastic adaptive dynamics.
• We first derive the polymorphic evolution sequence, based on prior work.
• Our evolutionary branching criterion extends a result by Champagnat and Méléard.
• In a classical model dormancy can favour evolutionary branching.
• Dormancy also affects several more population characteristics.
Abstract
In this paper, we investigate the consequences of dormancy in the ‘rare mutation’ and ‘large population’ regime of stochastic adaptive dynamics. Starting from an individual-based micro-model, we first derive the Polymorphic Evolution Sequence of the population, based on a previous work by Baar and Bovier (2018). After passing to a second ‘small mutations’ limit, we arrive at the Canonical Equation of Adaptive Dynamics, and state a corresponding criterion for evolutionary branching, extending a previous result of Champagnat and Méléard (2011).
The criterion allows a quantitative and qualitative analysis of the effects of dormancy in the well-known model of Dieckmann and Doebeli (1999) for sympatric speciation. In fact, quite an intuitive picture emerges: Dormancy enlarges the parameter range for evolutionary branching, increases the carrying capacity and niche width of the post-branching sub-populations, and, depending on the model parameters, can either increase or decrease the ‘speed of adaptation’ of populations. Finally, dormancy increases diversity by increasing the genetic distance between subpopulations.
For genus g=r(r+1)2+1, we prove that via the forgetful map, the universal Prym-Brill-Noether locus Rrg has a unique irreducible component dominating the moduli space Rg of Prym curves.
For genus g=2i≥4 and the length g−1 partition μ=(4,2,…,2,−2,…,−2) of 0, we compute the first coefficients of the class of D¯¯¯¯(μ) in PicQ(R¯¯¯¯g), where D(μ) is the divisor consisting of pairs [C,η]∈Rg with η≅OC(2x1+x2+⋯+xi−1−xi−⋯−x2i−1) for some points x1,…,x2i−1 on C. We further provide several enumerative results that will be used for this computation.
For genus g=2i≥4 and the length g−1 partition μ=(4,2,…,2,−2,…,−2) of 0, we compute the first coefficients of the class of D¯¯¯¯(μ) in PicQ(R¯¯¯¯g), where D(μ) is the divisor consisting of pairs [C,η]∈Rg with η≅OC(2x1+x2+⋯+xi−1−xi−⋯−x2i−1) for some points x1,…,x2i−1 on C. We further provide several enumerative results that will be used for this computation.
For genus g=2i≥4 and the length g−1 partition μ=(4,2,…,2,−2,…,−2) of 0, we compute the first coefficients of the class of D¯¯¯¯(μ) in PicQ(R¯¯¯¯g), where D(μ) is the divisor consisting of pairs [C,η]∈Rg with η≅OC(2x1+x2+⋯+xi−1−xi−⋯−x2i−1) for some points x1,…,x2i−1 on C. We further provide several enumerative results that will be used for this computation.
Between his arrival in Frankfurt in 1922 and and his proof of his famous finiteness theorem for integral points in 1929, Siegel had no publications. He did, however, write a letter to Mordell in 1926 in which he explained a proof of the finiteness of integral points on hyperelliptic curves. Recognizing the importance of this argument (and Siegel's views on publication), Mordell sent the relevant extract to be published under the pseudonym "X".
The purpose of this note is to explain how to optimize Siegel's 1926 technique to obtain the following bound. Let K be a number field, S a finite set of places of K, and f∈oK,S[t] monic of degree d≥5 with discriminant Δf∈o×K,S. Then: #|{(x,y):x,y∈oK,S,y2=f(x)}|≤2rankJac(Cf)(K)⋅O(1)d3⋅([K:Q]+#|S|).
This improves bounds of Evertse-Silverman and Bombieri-Gubler from 1986 and 2006, respectively.
The main point underlying our improvement is that, informally speaking, we insist on "executing the descents in the presence of only one root (and not three) until the last possible moment".
For genus g=2i≥4 and the length g−1 partition μ=(4,2,…,2,−2,…,−2) of 0, we compute the first coefficients of the class of D¯¯¯¯(μ) in PicQ(R¯¯¯¯g), where D(μ) is the divisor consisting of pairs [C,η]∈Rg with η≅OC(2x1+x2+⋯+xi−1−xi−⋯−x2i−1) for some points x1,…,x2i−1 on C. We further provide several enumerative results that will be used for this computation.
For genus g=2i≥4 and the length g−1 partition μ=(4,2,…,2,−2,…,−2) of 0, we compute the first coefficients of the class of D¯¯¯¯(μ) in PicQ(R¯¯¯¯g), where D(μ) is the divisor consisting of pairs [C,η]∈Rg with η≅OC(2x1+x2+⋯+xi−1−xi−⋯−x2i−1) for some points x1,…,x2i−1 on C. We further provide several enumerative results that will be used for this computation.
We prove that the projectivized strata of differentials are not contained in pointed Brill-Noether divisors, with only a few exceptions. For a generic element in a stratum of differentials, we show that many of the associated pointed Brill-Noether loci are of expected dimension. We use our results to study the Auel-Haburcak Conjecture: We obtain new non-containments between maximal Brill-Noether loci in Mg. Our results regarding quadratic differentials imply that the quadratic strata in genus 6 are uniruled.
Affine Bruhat--Tits buildings are geometric spaces extracting the combinatorics of algebraic groups. The building of PGL parametrizes flags of subspaces/lattices in or, equivalently, norms on a fixed finite-dimensional vector space, up to homothety. It has first been studied by Goldman and Iwahori as a piecewise-linear analogue of symmetric spaces. The space of seminorms compactifies the space of norms and admits a natural surjective restriction map from the Berkovich analytification of projective space that factors the natural tropicalization map. Inspired by Payne's result that the analytification is the limit of all tropicalizations, we show that the space of seminorms is the limit of all tropicalized linear embeddings ι:Pr↪Pn and prove a faithful tropicalization result for compactified linear spaces. The space of seminorms is in fact the tropical linear space associated to the universal realizable valuated matroid.
We use recent results by Bainbridge–Chen–Gendron–Grushevsky–Möller on compactifications of strata of abelian differentials to give a comprehensive solution to the realizability problem for effective tropical canonical divisors in equicharacteristic zero. Given a pair (Γ,D) consisting of a stable tropical curve Γ and a divisor D in the canonical linear system on Γ, we give a purely combinatorial condition to decide whether there is a smooth curve X over a non-Archimedean field whose stable reduction has Γ as its dual tropical curve together with an effective canonical divisor KX that specializes to D.
We show that the non-Archimedean skeleton of the d-th symmetric power of a smooth projective algebraic curve X is naturally isomorphic to the d-th symmetric power of the tropical curve that arises as the non-Archimedean skeleton of X. The retraction to the skeleton is precisely the specialization map for divisors. Moreover, we show that the process of tropicalization naturally commutes with the diagonal morphisms and the Abel-Jacobi map and we exhibit a faithful tropicalization for symmetric powers of curves. Finally, we prove a version of the Bieri-Groves Theorem that allows us, under certain tropical genericity assumptions, to deduce a new tropical Riemann-Roch-Theorem for the tropicalization of linear systems.
Using the notion of a root datum of a reductive group G we propose a tropical analogue of a principal G-bundle on a metric graph. We focus on the case G=GLn, i.e. the case of vector bundles. Here we give a characterization of vector bundles in terms of multidivisors and use this description to prove analogues of the Weil--Riemann--Roch theorem and the Narasimhan--Seshadri correspondence. We proceed by studying the process of tropicalization. In particular, we show that the non-Archimedean skeleton of the moduli space of semistable vector bundles on a Tate curve is isomorphic to a certain component of the moduli space of semistable tropical vector bundles on its dual metric graph.
In this article we provide a stack-theoretic framework to study the universal tropical Jacobian over the moduli space of tropical curves. We develop two approaches to the process of tropicalization of the universal compactified Jacobian over the moduli space of curves -- one from a logarithmic and the other from a non-Archimedean analytic point of view. The central result from both points of view is that the tropicalization of the universal compactified Jacobian is the universal tropical Jacobian and that the tropicalization maps in each of the two contexts are compatible with the tautological morphisms. In a sequel we will use the techniques developed here to provide explicit polyhedral models for the logarithmic Picard variety.