Scarcely any exterior formative stimuli are responsible for animal
organisation; and one would hardly be wrong in saying that this
morphogenetic independence in animals is due to their comparatively
far-reaching functional independence of those external agents which
have any sort of direction. But many organogenetic relations are known
to exist between the single parts of animal germs, each of these parts
being in some respect external to every other; and, indeed, it might
have been expected already *a priori*, that such formative relations
between the parts of an animal embryo must exist, after all we have
learned about the chief lines of early embryology. If differentiation
does not go on after the scheme of Weismann, that is, if it is not
carried out by true “evolutio” from within, how could it be effected
except from without? Indeed, every embryonic part may in some respect be
a possible cause for morphogenetic events, which are to occur on every
other part: it is here that the very roots of epigenesis are to be found.
Heliotropism and geotropism are among the well-known physiological
functions of plants: the roots are seen to bend away from the light and
towards the ground; the branches behave just in the opposite way. It now
has been supposed by Herbst that such “directive stimuli” may also be
at work among the growing or wandering parts of the embryo, that their
growth or their migration may be determined by the typical character of
other parts, and that real morphogenetic characters can be the result of
some such relation; a sort of “chemotropism” or “chemotaxis” may be at
work here. Herbst himself has discussed theoretically several cases of
organogenesis in which the action of directive stimuli is very probable.
What has become actually known by experiment is not very much at
present: the mesenchyme cells of Echinus are directed in their migration
by specified places in the ectoderm, the pigment cells of the yolk-sac
of the fish fundulus are attracted by its blood vessels, and nerves
may be forced to turn into little tubes containing brain substance;
but of course only the first two instances have any bearing on typical
morphogenesis.
The first case of an “internal formative stimulus” in the proper sense,
that is, of one embryonic part causing another to appear, was discovered
by Herbst himself. The arms of the so-called pluteus of the sea-urchin
are in formative dependence on the skeleton--no skeleton, no arms; so
many skeleton primordia,[42] in abnormal cases, so many arms; abnormal
position of the skeleton, abnormal position of the arms: these three
experimental observations form the proof of this morphogenetic relation.
[42] I use the word “primordia” for the German “Anlage”; it is better
than the word “rudiment,” as the latter may also serve to signify
the very last stage of a certain formation that is disappearing
(phylogenetically).
Public-domain text, read in full here on John Shaqi.
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