A Spatial Filtering Approach to Biological Patterning

Interactions between neighboring cells are essential for generating or refining patterns in a number of biological systems. We propose a discrete filtering approach to predict how networks of cells modulate spatially varying input signals to produce more complicated or precise output signals. The interconnections between cells determine the set of spatial modes that are amplified or suppressed based on the coupling and internal dynamics of each cell, analogously to the way a traditional digital filter modifies the frequency components of a discrete signal. We apply the framework to two systems in developmental biology: the Notch-Delta interaction that shapes \textit{Drosophila} wing veins and the Sox9/Bmp/Wnt network responsible for digit formation in vertebrate limbs. The latter case study demonstrates that Turing-like patterns may occur even in the absence of instabilities. Results also indicate that developmental biological systems may be inherently robust to both correlated and uncorrelated noise sources. Our work shows that a spatial frequency-based interpretation simplifies the process of predicting patterning in living organisms when both environmental influences and intercellular interactions are present.