Factors determining persistent asymmetry and egg shape in birds: A hypothesis

The shape of birds' eggs has fascinated scientists for many years. It is now possible mathematically to describe shape accurately, allowing exploration of the physical and ecological factors driving the evolution of egg shape. However, there has been relatively little consideration of how egg shape is established in the oviduct or, given that even without an external calcitic layer eggs retain their shape, how shape is fixed in the isthmus. This paper proposes a hypothesis that attempts to explain how egg shape is established and fixed in the oviduct. The hypothesis suggests that as the egg mass (i.e. yolk and albumen) moves from the magnum into the isthmus, it is squeezed by the physical restriction imposed by the isthmus lumen and cannot easily move into the isthmus. As the leading edge of the egg mass enters the isthmus, the egg mass in the distal magnum is forced to bulge outwards, resulting in an asymmetrical shape. The various egg shapes observed in birds are, hence, produced by the interaction between the size of the egg mass relative to female body mass, and the degree of the restriction of the isthmus. Thus, a large egg mass, i.e. relative to female body mass, entering a narrow isthmus will produce a pointed egg shape. If the egg mass is relatively small, and the isthmus lumen wide, more of the egg mass could enter the isthmus and the degree of asymmetry would be reduced. It is further proposed that egg shape is fixed during the formation of the shell membranes in the isthmus because the constituent protein fibres permanently stick together as they are deposited. For the first time this hypothesis helps explain the pattern of deposition and characteristics of the calcitic egg in relation to the diversity of egg shapes in birds and reptiles.