The deeper white muscle fibers fishes show diversity in arrangement. These fibers are organized into cone-shaped structures and attach to connective tissue sheets known as myosepta; each fiber shows a characteristic dorsoventral (α) and mediolateral (φ) trajectory. The segmented architecture theory predicts that, εx > εf. This phenomenon results in an architectural gear ratio, determined as longitudinal strain divided by fiber strain (εx / εf), greater than one and longitudinal velocity amplification; furthermore, this emergent velocity amplification may be augmented by variable architectural gearing via mesolateral and dorsoventral shape changes, a pattern seen in pennate muscle contractions. A red-to-white gearing ratio (red εf / white εf) captures the combined effect of the longitudinal red muscle fiber and oblique white muscle fiber strains. Simple bending behavior in homogenous beams suggests ε increases with distance from the neutral axis (z). This poses a problem to animals, such as fishes and salamanders, which undergo undulatory movement. Muscle fibers are constrained by the length-tension and force-velocity curves. Furthermore, it has been hypothesized that muscle fibers recruited for a particular task must operate within an optimal range of strains (ε) and contractile velocities to generate peak force and power respectively. Non-uniform ε generation during undulatory movement would force differing muscle fibers recruited for the same task to operate on differing portions of the length-tension and force-velocity curves; performance would not be optimal. Alexander predicted that the dorsoventral (α) and mediolateral (φ) orientation of the white fibers of the fish axial musculature may allow more uniform strain across varying mesolateral fiber distances. Unfortunately, the white muscle fiber musculature of fishes is too complex to study uniform strain generation; however, Azizi et al. studied this phenomenon using a simplified salamander model. Siren lacertian, an aquatic salamander, utilizes swimming motions similar to the aforementioned fishes yet contains hypaxial muscle fibers (which generate bending) characterized by a simpler organization. The hypaxial muscle fibers of S. lacertian are obliquely oriented, but have a near zero mediolateral (φ) trajectory and a constant dorsolateral (α) trajectory within each segment. Therefore, the effect of dorsolateral (α) trajectory and the distance between a given hypaxial muscle layer and the neutral axis of bending (z) on muscle fiber strain (ε) can be studied. Azizi et al. found that longitudinal contractions of the constant volume hypaxial muscles were compensated by an increase in the dorsoventral dimensions. Bulging was accompanied by fiber rotation as well as an