The Dynamics of Head Activation Changes during Proportioning inHydra oligactiswith Altered Head–Body Ratios

Normal hydra head–body proportions were altered by axially grafting a second head in place of the lower body column. The resulting animals had double the head tissue and one-quarter the normal body column. Changes in the head activation potential of tissue subjacent to both heads were monitored by assaying the ability of these animals to regenerate heads. The host head, the grafted head, or both heads were removed at varying times following graft construction and the animals were scored for head regeneration and/or the ability to express a head-specific antigen recognized by monoclonal antibody, CP8. In the presence of the grafted head, tissue subjacent to the host head lost the ability to regenerate a head or express the head-specific antigen over a 48-hr period. In the presence of the host head, tissue subjacent to the grafted head regenerated heads at a very low frequency and lost the ability to express the head-specific antigen over the same 48-hr period. Following simultaneous removal of both heads, animals initially regenerated both heads for the first 48 hr after graft construction. Then, both head regeneration and expression of the head-specific antigen declined gradually over the next 3 days, though not to the very low levels observed when one head remained. These data, especially the loss of CP8 labeling, support the hypothesis that loss of regeneration ability was due to a loss of head activation potential in tissues subjacent to the heads. We propose that this reflected the attempt of grafted animals to compensate for the altered head–body proportions through reproportioning. In keeping with this hypothesis, feeding the grafted animals to stimulate growth of body column tissue and a shift toward more normal head–body proportions prior to decapitation resulted in animals which were capable of regenerating heads when decapitated. Several interpretations of the results based on the Gierer–Meinhardt reaction-diffusion model of pattern formation are discussed.