Queen control of a key life-history event in a eusocial insect

In eusocial insects, inclusive fitness theory predicts potential queen–worker conflict over the timing of events in colony life history. Whether queens or workers control the timing of these events is poorly understood. In the bumble-bee Bombus terrestris, queens exhibit a ‘switch point’ in which they switch from laying diploid eggs yielding females (workers and new queens) to laying haploid eggs yielding males. By rearing foundress queens whose worker offspring were removed as pupae and sexing their eggs using microsatellite genotyping, we found that queens kept in the complete absence of adult workers still exhibit a switch point. Moreover, the timing of their switch points relative to the start of egg-laying did not differ significantly from that of queens allowed to produce normal colonies. The finding that bumble-bee queens can express the switch point in the absence of workers experimentally demonstrates queen control of a key life-history event in eusocial insects. In addition, we found no evidence that workers affect the timing of the switch point either directly or indirectly via providing cues to queens, suggesting that workers do not fully express their interests in queen–worker conflicts over colony life history

A mechanical model for biological pattern formation: A nonlinear bifurcation analysis

We present a mechanical model for cell aggregation in embryonic development. The model is based on the large traction forces exerted by fibroblast cells which deform the extracellular matrix (ECM) on which they move. It is shown that the subsequent changes in the cell environment can combine to produce pattern. A linear analysis is carried out for this model. This reveals a wide spectrum of different types of dispersion relations. A non-linear bifurcation analysis is presented for a simple version of the field equations: a non-standard element is required. Biological applications are briefly discussed

An analysis of one- and two-dimensional patterns in a mechanical model for morphogenesis

In early embryonic development, fibroblast cells move through an extracellular matrix (ECM) exerting large traction forces which deform the ECM. We model these mechanical interactions mathematically and show that the various effects involved can combine to produce pattern in cell density. A linear analysis exhibits a wide selection of dispersion relations, suggesting a richness in pattern forming capability of the model. A nonlinear bifurcation analysis is presented for a simple version of the governing field equations. The one-dimensional analysis requires a non-standard element. The two-dimensional analysis shows the possibility of roll and hexagon pattern formation. A realistic biological application to the formation of feather germ primordia is briefly discussed

A modified Oster-Murray-Harris mechanical model of morphogenesis

There are two main modeling paradigms for biological pattern formation in developmental biology: chemical prepattern models and cell aggregation models. This paper focuses on an example of a cell aggregation model, the mechanical model developed by Oster, Murray, and Harris [Development, 78 (1983), pp. 83--125]. We revisit the Oster--Murray--Harris model and find that, due to the infinitesimal displacement assumption made in the original version of this model, there is a restriction on the types of boundary conditions that can be prescribed. We derive a modified form of the model which relaxes the infinitesimal displacement assumption. We analyze the dynamics of this model using linear and multiscale nonlinear analysis and show that it has the same linear behavior as the original Oster--Murray--Harris model. Nonlinear analysis, however, predicts that the modified model will allow for a wider range of parameters where the solution evolves to a bounded steady state. The results from both analyses are verified through numerical simulations of the full nonlinear model in one and two dimensions. The increased range of boundary conditions that are well-posed, as well as a wider range of parameters that yield bounded steady states, renders the modified model more applicable to, and more robust for, comparisons with experiments

Spatial and spatio-temporal patterns in a cell-haptotaxis model

We investigate a cell-haptotaxis model for the generation of spatial and spatio-temporal patterns in one dimension. We analyse the steady state problem for specific boundary conditions and show the existence of spatially hetero-geneous steady states. A linear analysis shows that stability is lost through a Hopf bifurcation. We carry out a nonlinear multi-time scale perturbation procedure to study the evolution of the resulting spatio-temporal patterns. We also analyse the model in a parameter domain wherein it exhibits a singular dispersion relation

Fluctuation theorem for currents and Schnakenberg network theory

A fluctuation theorem is proved for the macroscopic currents of a system in a nonequilibrium steady state, by using Schnakenberg network theory. The theorem can be applied, in particular, in reaction systems where the affinities or thermodynamic forces are defined globally in terms of the cycles of the graph associated with the stochastic process describing the time evolution.Comment: new version : 16 pages, 1 figure, to be published in Journal of Statistical Physic

Spatio-temporal patterns in a mechanical model for mesenchymal morphogenesis

We present an in-depth study of spatio-temporal patterns in a simplified version of a mechanical model for pattern formation in mesenchymal morphogenesis. We briefly motivate the derivation of the model and show how to choose realistic boundary conditions to make the system well-posed. We firstly consider one-dimensional patterns and carry out a nonlinear perturbation analysis for the case where the uniform steady state is linearly unstable to a single mode. In two-dimensions, we show that if the displacement field in the model is represented as a sum of orthogonal parts, then the model can be decomposed into two sub-models, only one of which is capable of generating pattern. We thus focus on this particular sub-model. We present a nonlinear analysis of spatio-temporal patterns exhibited by the sub-model on a square domain and discuss mode interaction. Our analysis shows that when a two-dimensional mode number admits two or more degenerate mode pairs, the solution of the full nonlinear system of partial differential equations is a mixed mode solution in which all the degenerate mode pairs are represented in a frequency locked oscillation

Forces and pattern in limb morphogenesis

Prior to cartilage and bone formation in the limb bud chondroblasts condense into foci which provide the pattern for subsequent bone development. Formation of these condensations is, finally, a mechanical event, and so it is natural to ask what are the forces responsible for creating them. We have constructed a model for the process of cell aggregation during chondrogenesis which involves the following forces: (1) the passive elasticity of the extracellular matrix (ECM), (2) the osmotic swelling pressure of the ECM, which is generated principally by the hyaluronate (HA) component, (3) the active cell tractions developed by the chondroblasts. By examining the balance of forces between the cells and matrix we find that patterns of cell aggregation can spontaneously arise by an instability mechanism analogous to that which occurs in chemical pattern formation models

Towards an integrated experimental-theoretical approach for assessing the mechanistic basis of hair and feather morphogenesis

In his seminal 1952 paper, ‘The Chemical Basis of Morphogenesis’, Alan Turing lays down a milestone in the application of theoretical approaches to understand complex biological processes. His deceptively simple demonstration that a system of reacting and diffusing chemicals could, under certain conditions, generate spatial patterning out of homogeneity provided an elegant solution to the problem of how one of nature's most intricate events occurs: the emergence of structure and form in the developing embryo. The molecular revolution that has taken place during the six decades following this landmark publication has now placed this generation of theoreticians and biologists in an excellent position to rigorously test the theory and, encouragingly, a number of systems have emerged that appear to conform to some of Turing's fundamental ideas. In this paper, we describe the history and more recent integration between experiment and theory in one of the key models for understanding pattern formation: the emergence of feathers and hair in the skins of birds and mammals