The Development of Amphioxus

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The Development of Amphioxus

Section 12. The development of amphioxus, studied completely, is at once one of the most alluring and difficult tasks in the way of the zoologist; but certain of its earlier and most obvious fasts may very conveniently be taken into consideration now.

Section 13. The phenomena of the extrusion of polar bodies and fertilization are treated of later, and will, therefore, not be considered now. We will start our description with an egg-cell, which has escaped, of course, since there are no genital ducts, by rupture of the parent, has been fertilized by the male element, and is about to develop into a young amphioxus. It is simply a single cell, with some power of amoeboid motion, a single nucleus and nucleolus; and in amphioxus its protoplasm is clear and transparent. Frequently ova are loaded with granules of food store (yolk), which enable the young animal to go far with its development before it is hatched and has to begin fending for itself. Such an ovum as that of our present type, however being devoid of such yolk (alecithal = without yolk), necessitates a very early start in life, and, for reasons too complicated to state fully here, the development in such a case is considered particularly instructive and primitive by zoologists.

Section 14. The first thing to be seen in the developing cell is a deepening circular groove (Figure 1, Sheet 21), which divides the ovum into two parts. Another groove then cuts at right angles to this subdividing the two into four (Figure 2). Another groove, at right angles to both the former, follows, making the four eight (Figure 3). And so subdivision goes on. The whole process is called segmentation or cleavage.

Section 15. At the end of segmentation we get a hollow sphere of small cells, the cells separating from one another centrally and enclosing a cavity as the process proceeds. This is the blastosphere, shown diagrammatically in Figure 4, and of which an internal view, rather truer to the facts of the case as regards shape, is given as Figure 5. The central cavity is the segmentation cavity (s.c.).

Section 16. Invagination follows (Figure 6). In this process a portion of the blastosphere wall is the tucked into the rest, as indicated by the arrow, so that a two-layered sack is formed. The space ar. is the archenteron, the primordial intestine, and its mouth is called, the blastopore (bp.). The outer layer of this double-walled sac is called the epiblast. For the present we will give the inner lining no special term. The young amphioxus has, at this stage, which is called the gastrula stage, a curious parallelism with such a lowly form as the Hydra of our ditches. This latter creature, like the gastrula, consists essentially of two layers of cells, an outer protective and sensory layer, and an inner digestive one; it has a primordial intestine, or archenteron, and its mouth is sometimes regarded as being a blastopore. All animals that have little yolk, and start early in life for themselves, pass through a gastrula stage, substantially the same as this of amphioxus.

Section 17. The anus is perforated later near the region occupied at this stage by the blastopore. Hence the anterior end of the future amphioxus, the head end, is pointing towards the Figure 6, and the letters ep. are marked on the side which will be dorsal.

Section 18. Figure 7 i. is a dorsal view of the gastrula at a somewhat later stage, and here indications of distinctly vertebrate relationships already appear. Figure 7 ii. is a cross-section, its position, being shown by cross-lines in 7 i. and 6. Note first that the epiblast along the mid-dorsal line is sinking in to form what is called the neural plate (n.p.), and simultaneously on either side of it rise the neural folds (n.f.). Now, at Figure 8, a slightly later stage is represented, and at 9 i. the inturned part is separated from the general external epiblast as the spinal cord. The remainder of the epiblast constitutes the epidermis.

Section 19. Reverting to Figure 7 ii., along the dorsal side of the archenteron a thickening of its wall appears, and is gradually pinched off from it to form a cellular rod, lying along under the nervous axis and above the intestine. This is the notochord (compare Figures 8 and 9).

Section 20. Finally, we note two series of buds of cells, one on either side of the archenteron in Figure 7 ii. In 8 these buds have become hollow vesicles, growing out from it, the coelomic pouches. They are further developed in 9; and in 9 ii., which is a diagrammatic figure, they are indicated by dotted lines. They finally appear to (? entirely) obliterate the segmentation cavity-- they certainly do so throughout the body; and their cavities are in time cut off from the mesenteron, by the gradual constriction of their openings. In this way the coelom (body cavity) arises as a series of hollow "archenteric" outgrowths, and ms. becomes the alimentary canal. mt.c., the metapleural canals, probably arise subsequently to, and independently of, the general coelomic space, by a splitting in the body-wall substance.

Section 21. Hence, in considering the structure of amphioxus, we have three series of cells from which its tissues are developed:--

1. The epiblast.

2. Walls of the coelomic pouches, which form (a) an inner lining to the epiblast, (b) an outer coating to the hypoblast, and (c) the mesentery (m.), by which the intestine is supported. This is the mesoblast.

3. The lining of the mesenteron, or hypoblast.

From the epiblast the epidermis (not the dermis), the nervous system (including the nerves), and the sensory part of all sense organs are derived. From the mesoblast the muscles, the dermis genital and excretory organs, circulatory fluid and apparatus, any skeletal structures; and all connective tissue are derived. The mass of the body is thus evidently made of mesoblast. The hypoblast is the lining of the intestine and of the glands which open into it; and the material of the notochord is also regarded, as hypoblast.

Section 22. Figure 9 ii. shows all the essential points of the structure of amphioxus. Epiblast is indicated by a line of dashes, mesoblast by dots, and hypoblast, dark or black. The true mouth is formed late by a tucking-in of epiblast, the stomodaeum (s.d.), which meets and fuses with the hypoblast, and is then perforated. The position of this mouth is at the velum. The formation of the atrium has been described. The metapleural folds run forward in front of the velum, as the epipleurs (ep. in Sections 1 and 2), and form an oral hood (b.c.), around which the tentacles appear, and which is evidently not equivalent to the vertebrate mouth cavity, but in front of and outside it. The anus is formed by a tucking in, the proctodaeum, similar to the stomodaeum.

Section 23. The formation of the respiratory slits is complicated, and difficult to describe, but, since investigators have still to render its meaning apparent, it need not detain the elementary student.*

* See Balfour's Embryology, Volume 2, and Quarterly Journal of Microscopical Science March, 1891.

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