Nonlinear dynamical model and response of avian cranial kinesis.

All modern birds have kinetic skulls in which the upper bill can move relative to the braincase, but the biomechanics and motion dynamics of cranial kinesis in birds are poorly understood. In this paper, we model the dynamics of avian cranial kinesis, such as prokinesis and proximal rhynchokinesis in which the upper jaw pivots around the nasal-frontal (N-F) hinge. The purpose of this paper is to present to the biological community an approach that demonstrates the application of sophisticated predictive mathematical modeling tools to avian kinesis. The generality of the method, however, is applicable to the advanced study of the biomechanics of other skeletal systems. The paper begins with a review of the relevant biological literature as well as the essential morphology of avian kinesis, especially the mechanical coupling of the upper and lower jaw by the postorbital ligament. A planar model of the described bird jaw morphology is then developed that maintains the closed kinematic topology of the avian jaw mechanism. We then develop the full nonlinear equations of motion with the assumption that the M. protractor pterygoideus and M. depressor mandibulae act on the quadrate as a pure torque, and the nasal frontal hinge is elastic with damping. The mechanism is shown to be a single degree of freedom device due to the holonomic constraints present in the quadrate-jugal bar-upper jaw-braincase-quadrate kinematic chain as well as the quadrate-lower jaw-postorbital ligament-braincase-quadrate kinematic chain. The full equations are verified via simulation and animation using the parameters of a Grey Heron (Ardea cinerea). Next we develop a simplified analytical model of the equations by power series expansion. We demonstrate that this model reproduces the dynamics of the full model to a high degree of fidelity. We proceed to use the harmonic balance technique to develop the frequency response characteristics of the jaw mechanism. It is shown that this avian cranial kinesis model exhibits the characteristics of a hardening Duffing oscillator. Beyond the identification of the characteristics of the underlying dynamics, which provides insight into the behavior of the system, the model and methodology presented here provides other potential benefits. A framework has been developed that could be utilized to study the biomechanics of feeding and bite force as well the effects of cranial kinesis on the frequency and modulation of bird songs.