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

(FIGURE 2.360. Transverse section of the trunk of a chick-embryo, forty-five hours old. (From Balfour.) A ectoderm (horny-plate), Mc medullary tube, ch chorda, C entoderm (gut-gland layer), Pv primitive segment (episomite), Wd prorenal duct, pp coeloma (secondary body-cavity). So skin-fibre layer, Sp gut-fibre layer, v blood-vessels in latter, ao primitive aortas, containing red blood-cells.)

The lymph-cells (leucocytes), commonly called the "white corpuscles" of the blood, are phylogenetically older and more widely distributed in the animal world than the red. The great majority of the Invertebrates that have acquired an independent vascular system have only colourless lymph-cells in the circulating fluid. There is an exception in the Nemertines (Figure 2.358) and some groups of Annelids. When we examine the colourless blood of a cray-fish or a snail (Figure 2.358) under a high power of the microscope, we find in each drop numbers of mobile leucocytes, which behave just like independent Amoebae (Figure 1.17). Like these unicellular Protozoa, the colourless blood-cells creep slowly about, their unshapely plasma-body constantly changing its form, and stretching out finger-like processes first in one direction, then another. Like the Amoebae, they take particles into their cell-body. On account of this feature these amoeboid plastids are called "eating cells" (phagocytes), and on account of their motions "travelling cells" (planocytes). It has been shown by the discoveries of the last few decades that these leucocytes are of the greatest physiological and pathological consequence to the organism. They can absorb either solid or dissolved particles from the wall of the gut, and convey them to the blood in the chyle; they can absorb and remove unusable matter from the tissues. When they pass in large quantities through the fine pores of the capillaries and accumulate at irritated spots, they cause inflammation. They can consume and destroy bacteria, the dreaded vehicles of infectious diseases; but they can also transport these injurious Monera to fresh regions, and so extend the sphere of infection. It is probable that the sensitive and travelling leucocytes of our invertebrate ancestors have powerfully co-operated for millions of years in the phylogenesis of the advancing animal organisation.

The red blood-cells have a much more restricted sphere of distribution and activity. But they also are very important in connection with certain functions of the craniote-organism, especially the exchange of gases or respiration. The cells of the dark red, carbonised or venous, blood, which have absorbed carbonic acid from the animal tissues, give this off in the respiratory organs; they receive instead of it fresh oxygen, and thus bring about the bright red colour that distinguishes oxydised or arterial blood. The red colouring matter of the blood (haemoglobin) is regularly distributed in the pores of their protoplasm. The red cells of most of the Vertebrates are elliptical flat disks, and enclose a nucleus of the same shape; they differ a good deal in size (Figure 2.358). The mammals are distinguished from the other Vertebrates by the circular form of their biconcave red cells and by the absence of a nucleus (Figure 1.1); only a few genera still have the elliptic form inherited from the reptiles (Figure 1.2). In the embryos of the mammals the red cells have a nucleus and the power of increasing by cleavage (Figure 1.10).

The origin of the blood-cells and vessels in the embryo, and their relation to the germinal layers and tissues, are among the most difficult problems of ontogeny--those obscure questions on which the most divergent opinions are still advanced by the most competent scientists. In general, it is certain that the greater part of the cells that compose the vessels and their contents come from the mesoderm--in fact, from the gut-fibre layer; it was on this account that Baer gave the name of "vascular layer" to this visceral layer of the coeloma. But other important observers say that a part of these cells come from other germinal layers, especially from the gut-gland layer. It seems to be true that blood-cells may be formed from the cells of the entoderm before the development of the mesoderm. If we examine sections of chickens, the earliest and most familiar subjects of embryology, we find at an early stage the "primitive-aortas" we have already described (Figure 2.360 ao) in the ventral angle between the episoma (Pv) and hyposoma (Sp). The thin wall of these first vessels of the amniote embryo consists of flat cells (endothelia or vascular epithelia); the fluid within already contains numbers of red blood-cells; both have been developed from the gut-fibre layer. It is the same with the vessels of the germinative area (Figure 2.361 v), which lie on the entodermic membrane of the yelk-sac (c). These features are seen still more clearly in the transverse section of the duck-embryo in Figure 1.152. In this we see clearly how a number of stellate cells proceed from the "vascular layer" and spread in all directions in the "primary body-cavity"--i.e. in the spaces between the germinal layers. A part of these travelling cells come together and line the wall of the larger spaces, and thus form the first vessels; others enter into the cavity, live in the fluid that fills it, and multiply by cleavage--the first blood-cells.

But, besides these mesodermic cells of the "vascular layer" proper, other travelling cells, of which the origin and purport are still obscure, take part in the formation of blood in the meroblastic Vertebrates (especially fishes). The chief of these are those that Ruckert has most aptly denominated "merocytes." These "eating yelk-cells" are found in large numbers in the food-yelk of the Selachii, especially in the yelk-wall--the border zone of the germinal disk in which the embryonic vascular net is first developed. The nuclei of the merocytes become ten times as large as the ordinary cell-nucleus, and are distinguished by their strong capacity for taking colour, or their special richness in chromatin. Their protoplasmic body resembles the stellate cells of osseous tissue (astrocytes), and behaves just like a rhizopod (such as Gromia); it sends out numbers of stellate processes all round, which ramify and stretch into the surrounding food-yelk. These variable and very mobile processes, the pseudopodia of the merocytes, serve both for locomotion and for getting food; as in the real rhizopods, they surround the solid particles of food (granules and plates of yelk), and accumulate round their nucleus the food they have received and digested. Hence we may regard them both as eating-cells (phagocytes) and travelling-cells (planocytes). Their lively nucleus divides quickly and often repeatedly, so that a number of new nuclei are formed in a short time; as each fresh nucleus surrounds itself with a mantle of protoplasm, it provides a new cell for the construction of the embryo. Their origin is still much disputed.

(FIGURE 2.361. Merocytes of a shark-embryo, rhizopod-like yelk-cells underneath the embryonic cavity (B). (From Ruckert.) z two embryonic cells, k nuclei of the merocytes, which wander about in the yelk and eat small yelk-plates (d), k smaller, more superficial, lighter nuclei, k apostrophe a deeper nucleus, in the act of cleavage, k asterisk chromatin-filled border-nucleus, freed from the surrounding yelk in order to show the numerous pseudopodia of the protoplasmic cell-body.)

Half of the twelve stems of the animal world have no blood-vessels. They make their first appearance in the Vermalia. Their earliest source is the primary body-cavity, the simple space between the two primary germinal layers, which is either a relic of the segmentation-cavity, or is a subsequent formation. Amoeboid planocytes, which migrate from the entoderm and reach this fluid-filled primary cavity, live and multiply there, and form the first colourless blood-cells. We find the vascular system in this very simple form to-day in the Bryozoa, Rotatoria, Nematoda, and other lower Vermalia.

The first step in the improvement of this primitive vascular system is the formation of larger canals or blood-conducting tubes. The spaces filled with blood, the relics of the primary body-cavity, receive a special wall. "Blood-vessels" of this kind (in the narrower sense) are found among the higher worms in various forms, sometimes very simple, at other times very complex. The form that was probably the incipient structure of the elaborate vascular system of the Vertebrates (and of the Articulates) is found in two primordial principal vessels--a dorsal vessel in the middle line of the dorsal wall of the gut, and a ventral vessel that runs from front to rear in the middle line of its ventral wall. From the dorsal vessel is evolved the aorta (or principal artery), from the ventral vessel the principal or subintestinal vein. The two vessels are connected in front and behind by a loop that runs round the gut. The blood contained in the two tubes is propelled by their peristaltic contractions.

(FIGURE 2.362. Vascular system of an Annelid (Saenuris), foremost section. d dorsal vessel, v ventral vessel, c transverse connection of two (enlarged in shape of heart). The arrows indicate the direction of the flow of blood. (From Gegenbaur.)

The earliest Vermalia in which we first find this independent vascular system are the Nemertina (Figure 2.244). As a rule, they have three parallel longitudinal vessels connected by loops, a single dorsal vessel above the gut and a pair of lateral vessels to the right and left. In some of the Nemertina the blood is already coloured, and the red colouring matter is real haemoglobin, connected with elliptical discoid cells, as in the Vertebrates. The further evolution of this rudimentary vascular system can be gathered from the class of the Annelids in which we find it at various stages of development. First, a number of transverse connections are formed between the dorsal and ventral vessels, which pass round the gut ring-wise (Figure 2.362). Other vessels grow into the body-wall and ramify in order to convey blood to it. In addition to the two large vessels of the middle plane there are often two lateral vessels, one to the right and one to the left; as, for instance, in the leech. There are four of these parallel longitudinal vessels in the Enteropneusts (Balanoglossus, Figure 2.245). In these important Vermalia the foremost section of the gut has already been converted into a gill-crate, and the vascular arches that rise in the wall of this from the ventral to the dorsal vessel have become branchial vessels.

We have a further important advance in the Tunicates, which we have recognised as the nearest blood-relatives of our early vertebrate ancestors. Here we find for the first time a real heart--i.e. a central organ of circulation, driving the blood into the vessels by the regular contractions of its muscular wall, it is of a very rudimentary character, a spindle-shaped tube, passing at both ends into a principal vessel (Figure 2.221). By its original position behind the gill-crate, on ventral side of the Tunicates (sometimes more, sometimes less, forward), the head shows clearly that it has been formed by the local enlargement of a section of the ventral vessel. We have already noticed the remarkable alternation of the direction of the blood stream, the heart driving it first from one end, then from the other (

Chapter 2.

16). This is very instructive, because in most of the worms (even the Enteropneust) the blood in the dorsal vessel travels from back to front, but in the Vertebrates in the opposite direction. As the Ascidia-heart alternates