The works of Francis Maitland Balfour, Volume 2 (of 4) : $b A treatise on comparative embryology: InvertebrataBalfour, Francis M. (Francis Maitland)
Science
The works of Francis Maitland Balfour, Volume 2 (of 4) : $b A treatise on comparative embryology: Invertebrata
Balfour, Francis M. (Francis Maitland)
Embryology; Zoology
Numerous descriptions have been given during the last few years of the
internal phenomena of segmentation. The most recent contribution on
this head is that of Fol (No. 87). He appears to have been more
successful than other observers in obtaining a complete history of the
changes which take place, and it will therefore be convenient to take
as type the ovum of _Toxopneustes (Echinus) lividus_, on which he
made his most complete series of observations. The changes which take
place may be divided into a series of stages. The ovum immediately
after the fusion of the male and female pronucleus contains a central
segmentation nucleus.
In the first stage a clear protoplasmic layer derived from the plasma
of the cell is formed round the nucleus, from which there start
outwards a number of radial striæ, which are rendered conspicuous by
the radial arrangement of the yolk granules between them. The
nucleus during this process remains perfectly passive.
In the second stage the nucleus becomes less distinct and somewhat
elongated, and around it the protoplasmic layer of the earlier stage
is arranged in the form of a disc-shaped ring, compared by Fol to
Saturn's ring. The protoplasmic rays still take their origin from the
perinuclear protoplasm. This stage has a considerable duration (20
minutes).
In the third stage the protoplasm around the nucleus becomes
transported to the two nuclear poles, at each of which it forms a
clear mass surrounded by a star-shaped figure formed by radial striæ.
The nucleus is hardly visible in the fresh condition, but when brought
into view by reagents is found to contain many highly refractive
particles, and to be still enveloped in a membrane.
In the fourth stage the nucleus when treated by reagents has assumed
the well-known spindle form. The striæ of which it is composed are
continuous from one end of the spindle to the other and are thickened
at the centre. The central thickenings constitute the so-called
nuclear plate. The clear protoplasmic masses and stars are present as
before at the apices of the nucleus, and the rays of the latter
converge as if they would meet at the centre of the clear masses, but
stop short at their periphery. There is no trace of a membrane round
either the nuclear spindle or the clear masses; and in the centre of
the latter is a collection of granules. The striæ of the polar stars
are very fine but distinct.
Between the stage with a completely formed spindle and the previous
one the intermediate steps have not been made out for Toxopneustes;
but for Heteropods Fol has been able to demonstrate that the striæ of
the spindle and their central thickenings are formed, as in the case
of the spindle derived from the germinal vesicle, _from the
metamorphosis of the nuclear reticulum_. They commence to be formed
at the two poles, and are then (in Heteropods) in immediate contiguity
with the striæ of the stars. The striæ gradually grow towards the
centre of the nucleus and there meet.
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