We present a model to explain how the neurosecretory system of aquatic mollusks generates their diversity of shell structures and pigmentation patterns. The anatomical and physiological basis of this model sets it apart from other models used to explain shape and pattern. The model reproduces most known shell shapes and patterns and accurately predicts how the pattern alters in response to environmental disruption and subsequent repair. Finally, we connect the model to a larger class of neural models. neurosecretory ͉ mathematical model ͉ bifurcation S eashells display a remarkable variety of ornate pigmentation patterns. Accumulating evidence now indicates clearly that shell growth and patterning are under neural control and that shell growth and pigmentation is a neurosecretory phenomenon. Most of this evidence has been gained by detailed studies of the mantle, a tongue-like protrusion of the mollusk that wraps around the edge of the shell and deposits new shell material and pigment at the shell's growing edge (see Fig. 1 A and B) (1, 2). The shell itself is composed of crystal structures of calcium carbonate interspersed with associated proteins and other organic compounds, some of which are pigmented and arrayed in intricate patterns (3,4). This hard shell is covered by a thin organic layer of proteinaceous secretions, believed to function in regulating calcium crystallization (5). Early EM studies of the mantle recorded an extensive distribution of nerve fibers among the secretory cells (1, 6, 7). These fibers were later shown to have active synapses with secretory gland cells and synaptic inputs from other sensory organs in the mantle. From this evidence, it was proposed that neural-stimulated secretion controls shell growth (4, 7). The original evidence from gastropods has been extended to other mollusk taxa, including bivalves (8) and cephalopods, where improved experimental methods confirm clearly the role of neural control (9, 10). Neural recordings and neural cell ablation experiments have further verified the role of neural control in shell growth and repair (6,11,12).This experimental data on mollusk shell construction and pigmentation allow us to formulate a new neural network model and develop a unified explanation for the generation of both shell shapes and patterns. Unlike the purely geometric representations proposed earlier to model shell shape (13-18), our model links shape generation directly to the dynamics of the underlying neural network. Recent experimental work describes how differential growth patterns can lead to shell-like structures (19-21) but does not explain the biological origin or mechanism of these growth patterns. The neural model presented here closes this gap by explaining how the mantle neural net can encode the appropriate information required for shell growth as well as pigment deposition.Early attempts to reproduce shell patterns used cellular automata models, in which arbitrary rules determine the pigmentation of cells on a grid (22-24). Although they can reproduce som...