In Branchipus and other primitive Crustacea, Berger also finds the same
retinal layers, but is unable to distinguish in the brain the rest of the
optic ganglion. Judging from Berger's description of Branchipus, and
Bellonci's of Sphæroma, it would almost appear as though the cerebral part
of the retina in the higher forms originated from two ganglionic
enlargements, an external and internal enlargement, as Bellonci calls them.
The external ganglion (_Op. g. I._, Fig. 39) may be called the ganglion of
the retina, the cells of which form the nuclear layer of the higher forms,
and the internal ganglion (_Op. g. II._, Fig. 39), from which the optic
nerve-fibres to the brain arise, may therefore be called the ganglion of
the optic nerve. Bellonci describes how in this latter ganglion cells are
found very different to the small ones of the external ganglion or ganglion
of the retina. So also in Branchipus, judging from the pictures of Berger,
Claus, and from my own observations (_cf._ Fig. 46, in which the double
nature of the retinal ganglion is indicated), the peripheral part of the
optic ganglion--_i.e._ the retinal ganglion--may be spoken {91}of as
composed of two ganglia. The external of these is clearly the ganglion of
the retina; its cells form the nuclear layer, the striking character of
which, and close resemblance to the corresponding layer in vertebrates, is
shown by Claus' picture, which I reproduce (Fig. 40). The internal ganglion
with which the optic nerve is in connection contains large ganglion cells,
which, together with smaller ones, form the ganglionic layer of Berger.
The most recent observations of the structure of the compound retina of the
crustacean eye are those of Parker, who, by the use of the methylene blue
method, and Golgi's method of staining, has been able to follow out the
structure of the compound retina in the arthropod on the same lines as had
already been done for the vertebrate. These two methods have led to the
conclusion that the arthropod central nervous system and the vertebrate
central nervous system are built up in the same manner--viz. by means of a
series of ganglia connected together, each ganglion being composed of
nerve-cells, nerve-fibres, and a fine reticulated substance called by
Leydig in arthropods 'Punktsubstanz,' and known in vertebrates and in
invertebrates at the present time as 'neuropil.' A further analysis
resolves the whole system into a combination of groups of neurones, the
cells and fibres of which form the cells and fibres of the ganglia, while
their dendritic connections with the terminations of other neurones,
together with the neuroglia-cells form the 'neuropil.' As is natural to
expect, that part of the central nervous system which helps to form the
compound retina is built up in the same manner as the rest of the central
nervous system.
[Illustration: FIG. 40.--BIPOLAR CELLS OF NUCLEAR LAYER IN RETINA OF
BRANCHIPUS. (After CLAUS.)
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