Consistent left-right asymmetry in a population requires a special Kin
d of developmental specification. Models of this process must account
for the complex phenotypes seen when left-right is abnormal. In some h
uman and mouse mutations, there appears to be a randomisation of globa
l situs to either solitus or inversus, whereas in the inv/inv mouse es
sentially all are inversus. However, in these and all additional examp
les there are frequently other anomalies, including symmetry (isomeris
m) and discordance between organs (heterotaxia). Current models invoke
a molecular chiral referent as the source of the primary handed infor
mation. We describe a new two-gradient model which accommodates all de
fects of situs whilst caving the primary asymmetry unaltered.