1. The theory that the medusa is simply an organ, which has become
detached and has acquired a certain degree of independence, like the
well-known instance of the hectocotyle of the cuttle-fish. On this
view, put forward by E. van Beneden and T. H. Huxley, the sporosac is
the starting-point of an evolution leading up through the various
types of gonophores to the free medusa as the culminating point of a
phyletic series. The evidence against this view may be classed under
two heads: first, comparative evidence; hydroids very different in
their structural characters and widely separate in the systematic
classification of these organisms may produce medusae very similar, at
least so far as the essential features of medusan organization are
concerned; on the other hydroids closely allied, perhaps almost
indistinguishable, may produce gonophores in the one case, medusae in
the other; for example, _Hydractinia_ (gonophores) and _Podocoryne_
(medusae), _Tubularia_ (gonophores) and _Ectopleura_ (medusae),
_Coryne_ (gonophores) and _Syncoryne_ (medusae), and so on. If it is
assumed that all these genera bore gonophores ancestrally, then medusa
of similar type must have been evolved quite independently in a great
number of cases. Secondly, there is the evidence from the development,
namely, the presence of the entocodon in the medusa-bud, a structure
which, as explained above, can only be accounted for satisfactorily by
derivation from a medusan type of organization. Hence it may be
concluded that the gonophores are degenerate medusae, and not that the
medusae are highly elaborated gonophores, as the organ-theory
requires.
2. The theory that the medusa is an independent individual, fully
equivalent to the polyp in this respect, is now universally accepted
as being supported by all the facts of comparative morphology and
development. The question still remains open, however, which of the
two types of person may be regarded as the most primitive, the most
ancient in the race-history of the Hydromedusae. F. M. Balfour put
forward the view that the polyp was the more primitive type, and that
the medusa is a special modification of the polyp for reproductive
purposes, the result of division of labour in a polyp-colony, whereby
special reproductive persons become detached and acquire organs of
locomotion for spreading the species. W. K. Brooks, on the other hand,
as stated above, regards the medusa as the older type and looks upon
both polyp and medusa, in the Hydromedusae, as derived from a
free-swimming or floating actinula, the polyp being thus merely a
fixed nutritive stage, possessing secondarily acquired powers of
multiplication by budding.
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