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Josef Hofbauer

Publications and source records attributed to Josef Hofbauer.

At least 19 recordsLinked to original sources

Bifurcations in planar, quadratic mass-action networks with few reactions and low molecularity

In this paper we study bifurcations in mass-action networks with two chemical species and reactant complexes of molecularity no more than two. We refer to these as planar, quadratic networks as they give rise to (at most) quadratic differential equations on the nonnegative quadrant of the plane. Our aim is to study bifurcations in networks in this class with the fewest possible reactions, and the lowest possible product molecularity. We fully characterise generic bifurcations of positive equilibria in such networks with up to four reactions, and product molecularity no higher than three. In these networks we find fold, Andronov--Hopf, Bogdanov--Takens and Bautin bifurcations, and prove the non-occurrence of any other generic bifurcations of positive equilibria. In addition, we present a number of results which go beyond planar, quadratic networks. For example, we show that mass-action networks without conservation laws admit no bifurcations of codimension greater than $m-2$, where $m$ is the number of reactions; we fully characterise quadratic, rank-one mass-action networks admitting fold bifurcations; and we write down some necessary conditions for Andronov--Hopf and cusp bifurcations in mass-action networks. Finally, we draw connections with a number of previous results in the literature on nontrivial dynamics, bifurcations, and inheritance in mass-action networks.

math.DS

The inheritance of local bifurcations in mass action networks

We consider local bifurcations of equilibria in dynamical systems arising from chemical reaction networks with mass action kinetics. In particular, given any mass action network admitting a local bifurcation of equilibria, assuming only a general transversality condition, we list some enlargements of the network which preserve its capacity for the bifurcation. These results allow us to identify bifurcations in reaction networks from examination of their subnetworks, extending and complementing previous results on the inheritance of nontrivial dynamical behaviours amongst mass action networks. A number of examples are presented to illustrate applicability of the results.

math.DS

Stability and Fluctuations in Complex Ecological Systems

From 08-12 August, 2022, 32 individuals participated in a workshop, Stability and Fluctuations in Complex Ecological Systems, at the Lorentz Center, located in Leiden, The Netherlands. An interdisciplinary dialogue between ecologists, mathematicians, and physicists provided a foundation of important problems to consider over the next 5-10 years. This paper outlines eight areas including (1) improving our understanding of the effect of scale, both temporal and spatial, for both deterministic and stochastic problems; (2) clarifying the different terminologies and definitions used in different scientific fields; (3) developing a comprehensive set of data analysis techniques arising from different fields but which can be used together to improve our understanding of existing data sets; (4) having theoreticians/computational scientists collaborate closely with empirical ecologists to determine what new data should be collected; (5) improving our knowledge of how to protect and/or restore ecosystems; (6) incorporating socio-economic effects into models of ecosystems; (7) improving our understanding of the role of deterministic and stochastic fluctuations; (8) studying the current state of biodiversity at the functional level, taxa level and genome level.

q-bio.PE

Oscillations in three-reaction quadratic mass-action systems

It is known that rank-two bimolecular mass-action systems do not admit limit cycles. With a view to understanding which small mass-action systems admit oscillation, in this paper we study rank-two networks with bimolecular source complexes but allow target complexes with higher molecularities. As our goal is to find oscillatory networks of minimal size, we focus on networks with three reactions, the minimum number that is required for oscillation. However, some of our intermediate results are valid in greater generality. One key finding is that an isolated periodic orbit cannot occur in a three-reaction, trimolecular, mass-action system with bimolecular sources. In fact, we characterise all networks in this class that admit a periodic orbit; in every case all nearby orbits are periodic too. Apart from the well-known Lotka and Ivanova reactions, we identify another network in this class that admits a center. This new network exhibits a vertical Andronov--Hopf bifurcation. Furthermore, we characterise all two-species, three-reaction, bimolecular-sourced networks that admit an Andronov--Hopf bifurcation with mass-action kinetics. These include two families of networks that admit a supercritical Andronov--Hopf bifurcation, and hence a stable limit cycle. These networks necessarily have a target complex with a molecularity of at least four, and it turns out that there are exactly four such networks that are tetramolecular.

math.DS

The smallest bimolecular mass-action system with a vertical Andronov-Hopf bifurcation

We present a three-dimensional differential equation, which robustly displays a degenerate Andronov-Hopf bifurcation of infinite codimension, leading to a center, i.e., an invariant two-dimensional surface that is filled with periodic orbits surrounding an equilibrium. The system arises from a three-species bimolecular chemical reaction network consisting of four reactions. In fact, it is the only such mass-action system that admits a center via an Andronov-Hopf bifurcation.

math.DS

Permanence via invasion graphs: Incorporating community assembly into Modern Coexistence Theory

To understand the mechanisms underlying species coexistence, ecologists often study invasion growth rates of theoretical and data-driven models. These growth rates correspond to average per-capita growth rates of one species with respect to an ergodic measure supporting other species. In the ecological literature, coexistence often is equated with the invasion growth rates being positive. Intuitively, positive invasion growth rates ensure that species recover from being rare. To provide a mathematically rigorous framework for this approach, we prove theorems that answer two questions: (i) When do the signs of the invasion growth rates determine coexistence? (ii) When signs are sufficient, which invasion growth rates need to be positive? We focus on deterministic models and equate coexistence with permanence, i.e., a global attractor bounded away from extinction. For models satisfying certain technical assumptions, we introduce invasion graphs where vertices correspond to proper subsets of species (communities) supporting an ergodic measure and directed edges correspond to potential transitions between communities due to invasions by missing species. These directed edges are determined by the signs of invasion growth rates. When the invasion graph is acyclic (i.e. there is no sequence of invasions starting and ending at the same community), we show that permanence is determined by the signs of the invasion growth rates. In this case, permanence is characterized by the invasibility of all -i communities, i.e., communities without species i where all other missing species having negative invasion growth rates. We show that dissipative Lotka-Volterra models satisfy our technical assumptions and computing their invasion graphs reduces to solving systems of linear equations. We provide additional applications of the results and discuss open problems.

q-bio.PE

Adding species to chemical reaction networks: preserving rank preserves nondegenerate behaviours

We show that adding new chemical species into the reactions of a chemical reaction network (CRN) in such a way that the rank of the network remains unchanged preserves its capacity for multiple nondegenerate equilibria and/or periodic orbits. One consequence is that any bounded nondegenerate behaviours which can occur in a CRN can occur in a CRN with bounded stoichiometric classes. The main result adds to a family of theorems which tell us which enlargements of a CRN preserve its capacity for nontrivial dynamical behaviours. It generalises some earlier claims, and complements similar claims involving the addition of reactions into CRNs. The result gives us information on how ignoring some chemical species, as is common in biochemical modelling, might affect the allowed dynamics in differential equation models of CRNs. We demonstrate the scope and limitations of the main theorem via several examples. These illustrate how we can use the main theorem to predict multistationarity and oscillation in CRNs enlarged with additional species; but also how the enlargements can introduce new behaviours such as additional periodic orbits and new bifurcations.

math.DS

Some minimal bimolecular mass-action systems with limit cycles

We discuss three examples of bimolecular mass-action systems with three species, due to Feinberg, Berner, Heinrich, and Wilhelm. Each system has a unique positive equilibrium which is unstable for certain rate constants and then exhibits stable limit cycles, but no chaotic behaviour. For some rate constants in the Feinberg--Berner system, a stable equilibrium, an unstabe limit cycle, and a stable limit cycle coexist. All three networks are minimal in some sense. By way of homogenising the above three examples, we construct bimolecular mass-conserving mass-action systems with four species that admit a stable limit cycle. The homogenised Feinberg--Berner system and the homogenised Wilhelm--Heinrich system admit the coexistence of a stable equilibrium, an unstable limit cycle, and a stable limit cycle.

math.DS

Limit cycles in mass-conserving deficiency-one mass-action systems

We present some simple mass-action systems with limit cycles that fall under the scope of the Deficiency-One Theorem. All the constructed examples are mass-conserving and their stoichiometric subspace is two-dimensional. Using the continuation software MATCONT, we depict the limit cycles in all stoichiometric classes at once. The networks are trimolecular and tetramolecular, and some exhibit two or even three limit cycles. Finally, we show that the associated mass-action system of a bimolecular reaction network with two-dimensional stoichiometric subspace does not admit a limit cycle.

math.DS

Oscillations in planar deficiency-one mass-action systems

Whereas the positive equilibrium of a mass-action system with deficiency zero is always globally stable, for deficiency-one networks there are many different scenarios, mainly involving oscillatory behaviour. We present several examples, with centers or multiple limit cycles.

math.DS

On the bijectivity of families of exponential/generalized polynomial maps

We start from a parametrized system of $d$ generalized polynomial equations (with real exponents) for $d$ positive variables, involving $n$ generalized monomials with $n$ positive parameters. Existence and uniqueness of a solution for all parameters and for all right-hand sides is equivalent to the bijectivity of (every element of) a family of generalized polynomial/exponential maps. We characterize the bijectivity of the family of exponential maps in terms of two linear subspaces arising from the coefficient and exponent matrices, respectively. In particular, we obtain conditions in terms of sign vectors of the two subspaces and a nondegeneracy condition involving the exponent subspace itself. Thereby, all criteria can be checked effectively. Moreover, we characterize when the existence of a unique solution is robust with respect to small perturbations of the exponents or/and the coefficients. In particular, we obtain conditions in terms of sign vectors of the linear subspaces or, alternatively, in terms of maximal minors of the coefficient and exponent matrices. Finally, we present applications to chemical reaction networks with (generalized) mass-action kinetics.

math.AG

Planar S-systems: Permanence

We characterize permanence of planar S-systems. Further, we construct a planar S-system with three limit cycles.

math.DS

Planar S-systems: Global stability and the center problem

S-systems are simple examples of power-law dynamical systems (polynomial systems with real exponents). For planar S-systems, we study global stability of the unique positive equilibrium and solve the center problem. Further, we construct a planar S-system with two limit cycles.

math.DS

On global stability of the Lotka reactions with generalized mass-action kinetics

Chemical reaction networks with generalized mass-action kinetics lead to power-law dynamical systems. As a simple example, we consider the Lotka reactions with two chemical species and arbitrary power-law kinetics. We study existence, uniqueness, and stability of the positive equilibrium, in particular, we characterize its global asymptotic stability in terms of the kinetic orders.

math.DS

Genetic Recombination as a Chemical Reaction Network

The process of genetic recombination can be seen as a chemical reaction network with mass-action kinetics. We review the known results on existence, uniqueness, and global stability of an equilibrium in every compatibility class and for all rate constants, from both the population genetics and the reaction networks point of view.

q-bio.MN

Robust permanence for interacting structured populations

The dynamics of interacting structured populations can be modeled by $\frac{dx_i}{dt}= A_i (x)x_i$ where $x_i\in \R^{n_i}$, $x=(x_1,\dots,x_k)$, and $A_i(x)$ are matrices with non-negative off-diagonal entries. These models are permanent if there exists a positive global attractor and are robustly permanent if they remain permanent following perturbations of $A_i(x)$. Necessary and sufficient conditions for robust permanence are derived using dominant Lyapunov exponents $λ_i(μ)$ of the $A_i(x)$ with respect to invariant measures $μ$. The necessary condition requires $\max_i λ_i(μ)>0$ for all ergodic measures with support in the boundary of the non-negative cone. The sufficient condition requires that the boundary admits a Morse decomposition such that $\max_i λ_i(μ)>0$ for all invariant measures $μ$ supported by a component of the Morse decomposition. When the Morse components are Axiom A, uniquely ergodic, or support all but one population, the necessary and sufficient conditions are equivalent. Applications to spatial ecology, epidemiology, and gene networks are given.

q-bio.PE