Supplementary MaterialsS1 Appendix: Exact expression of the Laplace operator and noise. applied on an R8 cell pattern generated by the mathematical model. The mathematical model is evaluated on an elongated cell grid of 200 vs 44 cells in the case of no noise with the parameters in S1 Table. The order measure used is the fidelity of the nearest neighbor distance distributions. In this case, the fidelity order measure uses as a reference the pattern of the first 0-40 cells in the direction of the morphogenetic furrow (chosen as the x-direction in this paper) and uses as a target the pattern of the 40-80, 80-120 and 160-200 cells as shown in the x-axis of the plot. Nelotanserin The first data point starts at 1 and then saturates to a value of 9.85 for the rest of the slices. The plot supports the claim that, in the zero noise case, the resulting pattern retains its degree of order as it propagates. The reason that the first data point is higher than the other Nelotanserin is that the first column of cells is placed manually as an initial condition and it is placed slightly to the right to avoid the effects of reflective boundary conditions. Details on the quantitative measure of order can be found in the Methods section of the Nelotanserin main text.(PDF) pone.0210088.s004.pdf (32K) GUID:?64DBA03C-36C7-41DA-A436-F29C51FC0756 S2 Fig: Parametric variation leads to a threshold response when it is applied to a variety of parameters and in different combinations. (a) Parametric variation is applied in equal degree for all parameters except the ones that determine propagation. (b) Parametric variation is applied in variable amplitude for the pair and as in [7]. The order measure used is the fidelity of the nearest neighbor distance distributions. Parametric variation was not applied on the elements that define as it is approximated analytically and the pattern exhibited great sensitivity to such variations.(PDF) pone.0210088.s005.pdf (1008K) GUID:?518D53B5-A934-4160-8054-2CBFB6E01F87 S3 Fig: Probability distance order measures lead to threshold response when uniform distribution is used as a reference. (a)-(b) are generated using the fidelity, F, and (c)-(d) using the Kolmogorov distance, K, for nearest neighbor distances and angles respectively. The probability distance measures are applied on the R8 point pattern Mouse monoclonal to PTH1R with noise added in the model for the parameters and as a local inhibitor. (PDF) pone.0210088.s008.pdf (46K) GUID:?2493C5DD-1076-45BC-93E7-5AC2DC461754 S6 Fig: Fidelity order measure as a function of position on the simulated eye-disc. In this plot, noise was introduced in Du and Ds. Similarly to Fig 12, the plot shows the order of the pattern as a function of position from anterior to posterior. The x-axis refers to regions of the simulated eye disc 0-40, 40-80, 80-120 respectively, whereas the y-axis refers to the fidelity probability distance measure applied to nearest neighbor distance distributions. The conclusion is that the pattern saturates to a value of F. As the noise is increased, this saturation happens earlier in the eye disc.(PDF) pone.0210088.s009.pdf (51K) GUID:?E8BEC689-8674-4FC2-B4FE-3E188BB52A70 Data Availability StatementAll relevant data are within the paper and its Supporting Information files. Abstract During development of biological organisms, multiple complex structures are formed. In many instances, these structures need to exhibit a high degree of order to be functional, although many of their constituents are intrinsically stochastic. Hence, it has been suggested that biological robustness ultimately must rely on complex gene regulatory networks and clean-up mechanisms. Here we explore developmental processes that have evolved inherent robustness against stochasticity. In the context of the Drosophila eye disc, multiple optical units, ommatidia, develop into crystal-like patterns. During the larva-to-pupa stage of metamorphosis, the centers of the ommatidia are specified initially through the diffusion of morphogens, followed by the specification of R8 cells. Establishing the R8 cell is crucial in setting up the geometric, and functional, relationships of cells within an ommatidium and among neighboring ommatidia. Here we study an PDE mathematical model of these spatio-temporal processes in the presence of parametric stochasticity, defining and applying measures that quantify order within the resulting spatial patterns. We observe a universal sigmoidal response to increasing transcriptional noise. Ordered patterns persist Nelotanserin up to a threshold noise level in the model parameters. In accordance with prior qualitative observations, as the noise is further increased past a threshold point of no return, these ordered patterns rapidly become disordered. Such robustness in development allows for the accumulation of genetic variation without any observable changes in phenotype. We argue that the observed sigmoidal dependence.