Mechanical Design of Fiber-Wound Hydraulic Skeletons: The Stiffening and Straightening of Embryonic Notochords

The notochord can play an important mechanical role in shapechanges during early morphogenesis of vertebrates. For example,osmotic inflation of notochords elongates and straightens theaxis of frog early tail-bud embryos. In Xenopus laevis, thesheath of cross-helically arranged fibers around the notochordlimits the shape changes it undergoes when inflating, causingthe notochord to stiffen and straighten (Adams et al., 1990;Koehl et al., 1990). We used physical models of stage 24 X.laevis notochords to explore the mechanical consequences ofdifferent arrangements of the sheath fibers on the behaviorof such curved hydraulic cylinders. All the models straightenedupon inflation regardless of initial fiber angle ( = angle ofthe fibers to long axis of the cylinder). Notochord models with > 54° lengthened and narrowed as they straightened;although they could push, the forces they exerted were limitedby their tendency to buckle, which increased the greater the. In contrast, models with < 54° shortened and widenedas they straightened and showed pronounced increases in flexuralstiffness. The mean of X. laevis early tail-bud notochordsis 54°, a fiber angle that permits an increase in the end-to-enddistance of the model (along the anterior-posterior axis ofthe embryo) as it straightens and pushes when pressurized, butthat is less prone to Euler and local buckling than are modelswith higher 's. Nonetheless, a of 54° in notochords maysimply be the result of osmotic swelling.