The works of Francis Maitland Balfour, Volume 2 (of 4) : $b A treatise on comparative embryology: InvertebrataBalfour, Francis M. (Francis Maitland)
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The works of Francis Maitland Balfour, Volume 2 (of 4) : $b A treatise on comparative embryology: Invertebrata
Balfour, Francis M. (Francis Maitland)
Embryology; Zoology
In some cases, _e.g._ Limax (Gegenbaur), Neritina (Claparède),
Pterotrachæa (Gegenbaur), the larva is stated to be coated by an
uniform covering of cilia before the formation of the velum, but the
researches of Fol have thrown very considerable doubt on these
statements. In some cases amongst the Nudibranchiata (Haddon) and
Pteropoda there are one or two long cilia in the middle of the velar
area. In many Nudibranchiata (Haddon) there is present a more or less
complete _post-oral_ ring of small cilia, which belongs to the velum.
The cilia on the velum cause a rotation of the larva within the
egg-capsule. Cilia are in most cases (Paludina, etc.) developed on the
foot and on a small anal area.
The shell-gland arises as an epiblastic thickening on the posterior
and dorsal side. In this thickening a deep invagination (fig. 101,
_shs._) is soon formed, in which a chitinous plug may become developed
(Paludina, Cymbulia? etc.), and in abnormal larvæ such a chitinous
plug is generally formed.
The foot is a simple prominence of epiblast on the ventral surface, in
the cavity of which there are usually a number of mesoblast cells
(fig. 101, _f_). The larval form just described has been named by
Lankester the trochosphere larva.
Before considering the further external changes which the larva
undergoes, it will be well to complete the history of the invaginated
hypoblast.
[FIG. 102. EMBRYO OF A HETEROPOD. (From Gegenbaur; after Fol.)
_o._ mouth; _v._ velum; _g._ archenteron; _p._ foot; _c._ body
cavity; _s._ shell-gland.]
The hypoblast has after its invagination either the form of a sack
(fig. 102) or of a solid mass (fig. 101). Whether the mouth be the
blastopore or no, the permanent oesophagus is formed of epiblast
cells, so that the oesophagus and buccal cavity are always lined by
epiblast. When the blastopore remains permanently open the outer part
of the oesophagus grows as a prominent ridge round the opening.
The mesenteric sack itself becomes differentiated into a stomach
adjoining the oesophagus, a liver opening immediately behind this,
and an intestine. The cells forming the hepatic diverticula and
sometimes also those of the stomach may during larval life secrete in
their interior peculiar albuminous products, similar to ordinary
food-yolk.
The proctodæum, except when it is the blastopore, arises later than
the mouth. It is frequently developed from a pair of projecting
epiblast cells symmetrically placed in the median ventral line behind
the foot. It eventually forms a very shallow invagination meeting the
intestine. Its opening is the anus. The anus, though at first always
symmetrical and ventral, subsequently, on the formation of the pallial
cavity, opens into this usually on the right and dorsal side.
In the cases where the hypoblast is not invaginated in the form of a
sack the formation of the mesenteron is somewhat complicated, and is
described in the sequel.
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