If the embryo is set free as a free-swimming, so-called planula-larva, in the blastula, parenchymula, or gastrula stage, then a free actinula stage is not found; if, on the other hand, a free actinula occurs, then there is no free planula stage.
Formation of archenteron and blastopore may, however, be deferred till a later stage (actinula or after).
- Free Actinula rhiza from which a polyp appears to be of Tubularia.
In Trachylinae the polyp-stage is passed over, and is represented only by the actinula as a transitory embryonic stage.
In Cunina parasitica, however, the ovum develops into an actinula, which buds actinulae as before, but only the daughter-actinulae develop into medusae, while the original, parent-actinula dies off; here, therefore, larval budding has led to a true alternation of generations.
Hydra must, in short, be a living representative of the ancestor of which the actinula-stage is a transient reminiscence in the development of higher forms. It may be pointed out in this connexion that the fixation of Hydra is only temporary, and that the animal is able at all times to detach itself, to move to a new situation, and to fix itself again.
There is no difficulty whatever in regarding Hydra as bearing the same relation to the actinula-stage of other Hydromedusae that a Rotifer bears to a trochophore-larva or a fish to a tadpole.
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.
If a polyp, such as Hydra, be regarded simply as a sessile actinula, we must certainly consider the polyp to be the older type, and it may be pointed out that in the Anthozoa only polyp-individuals occur.
This must not be taken to mean, however, that the medusa is derived from a sessile polyp; it must be regarded as a direct modification of the more ancient free actinula form, without primitively any intervening polyp-stage, such as has been introduced secondarily into the development of the Leptolinae and represents 'a revival, so to speak, of an ancestral form or larval stage, which has taken on a special role in the economy of the species.
Without a hydroid phase; the medusa develops directly from the actinula larva, which may, however, multiply by budding.
The development of the Trachomedusae, so far as it is known, shows an actinula-stage which is either free (larval) or passed over in the egg (foetal) as in Geryonia; in no case does there appear to be a free planula-stage.
The actinula, when free, may multiply by larval budding, but in all cases both the original actinula and all its descendants become converted into medusae, so that there is no alter nation of generations.
In Gonionemus the actinula becomes attached and polyp-like and reproduces by budding.
The two ends of the planula become greatly lengthened and give rise to the two primary tentacles of the actinula, of which the mouth arises from one side of the planula.
In these species the actinula is parasitic upon another medusa; for instance, Cunoctantha octonaria upon Turritopsis, C. proboscidea upon Liriope or Geryonia.
The parasitic actinula is found attached to the proboscis of the medusa; it thrusts its greatly elongated hypostome into the mouth of the medusa and nourishes itself upon the food in the digestive cavity of its host.
In such cases the original parent-actinula does not itself become a medusa, but remains arrested in development and ultimately dies off, so that a true alternation of generations is brought about.
The gastrula has now become an actinula, which may be termed the distinctive larva of the Cnidaria, and doubtless represents in a transitory manner the common ancestor of the group. In no case known, however, does the actinula become the adult, sexually mature individual, but always undergoes further modifications, whereby it develops into either a polyp or a medusa.
- Diagram showing the change of the Actinula (A) into a Polyp (B); a-b, principal (vertical) axis; c-d, horizontal axis.
The ectoderm loses entirely the ciliation which it had in the planula and actinula stages and commonly secretes on its external surface a protective or supporting investment, the perisarc. Contrasting with this, the anthopolyp is generally of s q uat form, the diameter often exceeding the height; the peristome is wide, a hypostome is lacking, and the ectoderm, or so much of it as is exposed, i.e.
- Diagram showing the change of the Actinula into a Medusa.
A, Vertical section of the actinula; a-b and c-d as in fig.
In short, the hydropolyp is characterized by a more simple type of organization than the anthopolyp, and is in most respects less modified from the actinula type of structure.
Returning now to the actinula, this form may, as already stated, develop into a medusa, a type of individual found only in the Hydrozoa, as here understood.
To become a medusa, the actinula grows scarcely at all in the direction of the principal axis, but greatly along a plane at right angles to it.
Thus the development of the two types of individual seen in the Hydrozoa may be summarized as follows: - Egg Free Blastula "Planula" Parenchymula Stage I Gastrula Actinula 1 Polyp Medusa This development, though probably representing the primitive sequence of events, is never actually found in its full extent, but is always abbreviated by omission or elimination of one or more of the stages.
(For other anatomical variations see Hydromedusae and Scyphomedusae.) In development the medusa can be derived easily by a process of differential growth, combined with concrescence of cell-layers, from the actinula-larva.
(For figures see Hydrozoa.) The actinula is polyp-like, with a sack-like or rounded body; a crown of tentacles surrounds a wide peristome, in the centre of which is the mouth, usually raised on a conical process termed the hypostome.
To produce a medusa the actinula grows greatly along a plane at right angles to the vertical axis of the body, whereby the aboral surface of the actinula becomes the exumbrella, and the peristome becomes the subumbrella.
On the other hand, the parenchymula may develop directly into the actinula or even into the polyp, with suppression of the intervening steps.
Great apparent differences may also be brought about by variations in the period at which the embryo is set free as a larva, and since two free-swimming stages, planula and actinula, are unnecessary, one or other of them is always suppressed.
In Cordylophora the embryo is set free at the parenchymula stage as a planula which fixes itself and develops into a polyp, both gastrula and actinula stages being suppressed.
In Tubularia, on the other hand, the parenchymula develops into an actinula within the maternal tissues, and is then set free, creeps about for a time, and after fixing itself, changes into a polyp; hence in this case the planula-stage, as a free larva, is entirely suppressed.
The Hydrozoa may be defined, therefore, as Cnidaria in which two types of individual, the polyp and the medusa, may be present, each type developed along divergent lines from the primitive actinula form.