The Evolution of Man, V.2 by Ernst Haeckel (comprehension books .TXT) 📕

Chapter 1.

8). In the genealogical tree of the Vertebrates this palingenetic form of segmentation has been preserved in the Amphioxus alone, all the other Vertebrates having cenogenetically modified forms of cleavage. In any case, the latter were developed from the former, and so the segmentation of the ovum in the Amphioxus has a great interest for us (cf. Figure 1.38). The outcome of this repeated cleavage is the formation of a round cluster of cells, composed of homogeneous, indifferent cells of the simplest character (Figure 2.230). This is called the morula (= mulberry-embryo) on account of its resemblance to a mulberry or blackberry.

(FIGURE 2.229. Original or primordial ovum-cleavage. The stem-cell or cytula, formed by fecundation of the ovum, divides by repeated regular cleavage first into two (A), then four (B), then eight (C), and finally a large number of segmentation-cells (D).

FIGURE 2.230. Morula, or mulberry-shaped embryo.)

It is clear that this morula reproduces for us to-day the simple structure of the multicellular animal that succeeded the unicellular amoeboid form in the early Laurentian period. In accordance with the biogenetic law, the morula recalls the ancestral form of the Moraea, or simple colony of Protozoa. The first cell-communities to be formed, which laid the early foundation of the higher multicellular body, must have consisted of homogeneous and simple amoeboid cells. The oldest Amoebae lived isolated lives, and even the amoeboid cells that were formed by the segmentation of these unicellular organisms must have continued to live independently for a long time. But gradually small communities of Amoebae arose by the side of these eremitical Protozoa, the sister-cells produced by cleavage remaining joined together. The advantages in the struggle for life which these communities had over the isolated cells favoured their formation and their further development. We find plenty of these cell-colonies or communities to-day in both fresh and salt water. They belong to various groups both of the Protophyta and Protozoa.

To have some idea of those ancestors of our race that succeeded phylogenetically to the Moraeada, we have only to follow the further embryonic development of the morula. We then see that the social cells of the round cluster secrete a sort of jelly or a watery fluid inside their globular body, and they themselves rise to the surface of it (Figure 1.29 F, G). In this way the solid mulberry-embryo becomes a hollow sphere, the wall of which is composed of a single layer of cells. We call this layer the blastoderm, and the sphere itself the blastula, or embryonic vesicle.

This interesting blastula is very important. The conversion of the morula into a hollow ball proceeds on the same lines originally in the most diverse stems--as, for instance, in many of the zoophytes and worms, the ascidia, many of the echinoderms and molluscs, and in the amphioxus. Moreover, in the animals in which we do not find a real palingenetic blastula the defect is clearly due to cenogenetic causes, such as the formation of food-yelk and other embryonic adaptations. We may, therefore, conclude that the ontogenetic blastula is the reproduction of a very early phylogenetic ancestral form, and that all the Metazoa are descended from a common stem-form, which was in the main constructed like the blastula. In many of the lower animals the blastula is not developed within the foetal membranes, but in the open water. In those cases each blastodermic cell begins at an early stage to thrust out one or more mobile hair-like processes; the body swims about by the vibratory movement of these lashes or whips (Figure 1.29 F).

We still find, both in the sea and in fresh water, various kinds of primitive multicellular organisms that substantially resemble the blastula in structure, and may be regarded in a sense as permanent blastula-forms--hollow vesicles or gelatinous balls, with a wall composed of a single layer of ciliated homogeneous cells. There are "blastaeads" of this kind even among the Protophyta--the familiar Volvocina, formerly classed with the infusoria. The common Volvox globator is found in the ponds in the spring--a small, green, gelatinous globule, swimming about by means of the stroke of its lashes, which rise in pairs from the cells on its surface. In the similar Halosphaera viridis also, which we find in the marine plancton (floating matter), a number of green cells form a simple layer at the surface of the gelatinous ball; but in this case there are no cilia.

Some of the infusoria of the flagellata-class (Signura, Magosphaera, etc.) are similar in structure to these vegetal clusters, but differ in their animal nutrition; they form the special group of the Catallacta. In September, 1869, I studied the development of one of these graceful animals on the island of Gis-Oe, off the coast of Norway (Magosphaera planula), Figures 2.231 and 2.232). The fully-formed body is a gelatinous ball, with its wall composed of thirty-two to sixty-four ciliated cells; it swims about freely in the sea. After reaching maturity the community is dissolved. Each cell then lives independently for some time, grows, and changes into a creeping amoeba. This afterwards contracts, and clothes itself with a structureless membrane. The cell then looks just like an ordinary animal ovum. When it has been in this condition for some time the cell divides into two, four, eight, sixteen, thirty-two, and sixty-four cells. These arrange themselves in a round vesicle, thrust out vibratory lashes, burst the capsule, and swim about in the same magosphaera-form with which we started. This completes the life-circle of the remarkable and instructive animal.

If we compare these permanent blastulae with the free-swimming ciliated larvae or blastulae, with similar construction, of many of the lower animals, we can confidently deduce from them that there was a very early and long-extinct common stem-form of substantially the same structure as the blastula. We may call it the Blastaea. Its body consisted, when fully formed, of a simple hollow ball, filled with fluid or structureless jelly, with a wall composed of a single stratum of ciliated cells. There were probably many genera and species of these blastaeads in the Laurentian period, forming a special class of marine protists.

It is an interesting fact that in the plant kingdom also the simple hollow sphere is found to be an elementary form of the multicellular organism. At the surface and below the surface (down to a depth of 2000 yards) of the sea there are green globules swimming about, with a wall composed of a single layer of chlorophyll-bearing cells. The botanist Schmitz gave them the name of Halosphaera viridis in 1879.

The next stage to the Blastaea, and the sixth in our genealogical tree, is the Gastraea that is developed from it. As we have already seen, this ancestral form is particularly important. That it once existed is proved with certainty by the gastrula, which we find temporarily in the ontogenesis of all the Metazoa (Figure 1.29 J, K). As we saw, the original, palingenetic form of the gastrula is a round or oval uni-axial body, the simple cavity of which (the primitive gut) has an aperture at one pole of its axis (the primitive mouth). The wall of the gut consists of two strata of cells, and these are the primary germinal layers, the animal skin-layer (ectoderm) and vegetal gut-layer (entoderm).

The actual ontogenetic development of the gastrula from the blastula furnishes sound evidence as to the phylogenetic origin of the Gastraea from the Blastaea. A pit-shaped depression appears at one side of the spherical blastula (Figure 1.29 H). In the end this invagination goes so far that the outer or invaginated part of the blastoderm lies close on the inner or non-invaginated part (Figure 1.29 J). In explaining the phylogenetic origin of the gastraea in the light of this ontogenetic process, we may assume that the one-layered cell-community of the blastaea began to take in food more largely at one particular part of its surface. Natural selection would gradually lead to the formation of a depression or pit at this alimentary spot on the surface of the ball. The depression would grow deeper and deeper. In time the vegetal function of taking in and digesting food would be confined to the cells that lined this hole; the other cells would see to the animal functions of locomotion, sensation, and protection. This was the first division of labour among the originally homogeneous cells of the blastaea.

(FIGURE 2.231. The Norwegian Magosphaera planula, swimming about by means of the lashes or cilia at its surface.

FIGURE 2.232. Section of Magosphaera planula, showing how the pear-shaped cells in the centre of the gelatinous ball are connected by a fibrous process. Each cell has a contractile vacuole as well as a nucleus.)

The effect, then, of this earliest histological differentiation was to produce two different kinds of cells--nutritive cells in the depression and locomotive cells on the surface outside. But this involved the severance of the two primary germinal layers--a most important process. When we remember that even man's body, with all its various parts, and the body of all the other higher animals, are built up originally out of these two simple layers, we cannot lay too much stress on the phylogenetic significance of this gastrulation. In the simple primitive gut or gastric cavity of the gastrula and its rudimentary mouth we have the first real organ of the animal frame in the morphological sense; all the other organs were developed afterwards from these. In reality, the whole body of the gastrula is merely a "primitive gut." I have shown already (

Chapters

1.8 and 1.9) that the two-layered embryos of all the Metazoa can be reduced to this typical gastrula. This important fact justifies us in concluding, in accordance with the biogenetic law, that their ancestors also were phylogenetically developed from a similar stem-form. This ancient stem-form is the gastraea.

The gastraea probably lived in the sea during the Laurentian period, swimming about in the water by means of its ciliary coat much as free ciliated gastrulae do to-day. Probably it differed from the existing gastrula only in one essential point, though extinct millions of years ago. We have reason, from comparative anatomy and ontogeny, to believe that it multiplied by sexual generation, not merely asexually (by cleavage, gemmation, and spores), as was no doubt the case with the earlier ancestors. Some of the cells of the primary germ-layers probably became ova and others fertilising sperm. We base these hypotheses on the fact that we do to-day find the simplest form of sexual reproduction in some of the living gastraeads and other lower animals, especially the sponges.

The fact that there are still in existence various kinds of gastraeads, or lower Metazoa with an organisation little higher than that of the hypothetical gastraea, is a strong point in favour of our theory. There are not very many species of these living gastraeads; but their morphological and phylogenetic interest is so great, and their intermediate position between the Protozoa and Metazoa so instructive, that I proposed long ago (1876) to make a special class of them. I distinguished three orders in this class--the Gastremaria, Physemaria, and Cyemaria (or Dicyemida). But we might also regard these three orders as so many independent classes in a primitive gastraead stem.

The Gastremaria and Cyemaria, the chief of these living gastraeads, are small Metazoa that live parasitically inside other Metazoa, and are, as a rule, 1/50 to 1/25 of an inch long, often much less (Figure 2.233, 1 to 15). Their soft body, devoid of skeleton, consists of two simple strata of cells, the primary germinal layers; the outer of these is thickly clothed with long hair-like lashes, by which the parasites swim about in the various cavities of their host. The inner germinal layer furnishes the sexual products. The pure type of the original gastrula (or archigastrula, Figure 1.29 I) is seen in the Pemmatodiscus gastrulaceus, which Monticelli discovered in the umbrella of a large medusa (Pilema pulmo) in 1895; the convex surface of this gelatinous umbrella