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The fertilized egg of any animal consists of a single cell, a little ma.s.s of protoplasm containing a nucleus and surrounded by a structureless membrane. The egg is globular. The nucleus undergoes certain very peculiar, still but little understood, changes and divides into two. The protoplasm also soon divides into two ma.s.ses cl.u.s.tering each around its own nucleus. The plane of division will be marked around the outside by a circular furrow, but the cells will still remain united by a large part of the membrane which bounds their adjacent, newly formed, internal faces.
Let us suppose that the egg lay so that the first plane of division was vertical and extending north and south. Each cell or half of the egg will divide into two precisely as before. The new plane of division will be vertical, but extending east and west. Each plane pa.s.ses through the centre of the egg, and the four cells are of the same form and size, like much-rounded quarters of an orange. The third plane will lie horizontal or equatorial, and will divide each of these quarters into an upper and lower octant. The cells keep on dividing rapidly, the eight form sixteen, then thirty-two, etc. The sharp angle by which the cells met at the centre has become rounded off, and has left a little s.p.a.ce, the segmentation cavity, filled with fluid in the middle of the embryo. The cells continue to press or be crowded away from the centre and form a layer one cell deep on the surface of the sphere.
This embryo, resembling a hollow rubber ball filled with fluid, is called a blastosphere. It corresponds in structure with the fully developed volvox, except, of course, in lacking reproductive cells.
[Ill.u.s.tration: 4. GASTRULA. HATSCHEK, FROM HERTWIG.
Outer layer is the ectoderm; inner layer, the entoderm; internal cavity, the archenteron; mouth of cavity, blastopore.]
If the rubber ball has a hole in it so that I can squeeze out the water, I can thrust the one-half into the other, and change the ball into a double-walled cup. A similar change takes place in the embryo. The cells of the lower half of the blastosphere are slightly larger than those of the upper half. This lower hemisphere flattens and then thrusts itself, or is inv.a.g.i.n.ated, into the upper hemisphere of smaller cells and forms its lining. This cup-shaped embryo is called the gastrula. The cup deepens somewhat and becomes ovoid. Take a boiled egg, make a hole in the smaller end and remove the yolk, and you have a pa.s.sable model of a gastrula. The sh.e.l.l corresponds to the ectoderm or outer layer of smaller cells; the layer of "white" represents the entoderm or lining of larger cells.
The s.p.a.ce occupied by the yolk corresponds to the archenteron or primitive digestive cavity; and the opening at the end to the primitive mouth or blastopore. Ectoderm and entoderm unite around the mouth. Both the blastosphere and gastrula often swim freely by flagella.
You can hardly have failed to notice how closely the gastrula corresponds to a hydra, and many facts lead us to believe that the still earlier ancestor of the hydra was free swimming, and that the tentacles are a later development correlated with its adult sessile life. Yet we must not forget that the hydra is even now not quite sessile, it moves somewhat. And our ancestor was almost certainly a free swimming gastraea, or hypothetical form corresponding in form and structure to the gastrula. The ancestor of man never settled down lazily into a sessile life.
But how is an adult worm or vertebrate formed out of such a gastrula? To answer this would require a course of lectures on embryology. But certain changes interest us. Between the ectoderm and entoderm of the gastrula, in the s.p.a.ce occupied by the supporting membrane of hydra, a new layer of cells, the mesoderm, appears. This has been produced by the rapid growth and reproduction of certain cells of the entoderm which have migrated, so to speak, into this new position. In higher forms it becomes of continually greater importance, until finally nearly all the organs of the body develop from it. In our bodies only the lining of the mid-intestine and of its glands has arisen from the entoderm. And only the epidermis, or outer layer of our skin, and the nervous system and parts of our sense-organs have arisen from the ectoderm. But our mid-intestine is still the greatly elongated archenteron of the gastrula.
We may therefore compare the hydra or gastrula to a little portion of the lining of the human mid-intestine covered with a little flake of epidermis. This much the hydra has attained. But our bones and muscles and blood-vessels all come from the mesoderm by folding, plaiting, and channelling, and division of labor resulting in differentiation of structure. Of all true mesodermal structures the hydra has actually none, but in the ectodermal and entodermal cells he has the potentiality of them all. We must now try to discover how these potentialities became actualities in higher forms.
The third stage in our ancestral series is the turbellarian. This is a little, flat, oval worm, varying greatly in size in different species, and found both in fresh and salt water. Some would deny that this worm belonged in our series at all. But, while doubtless considerably modified, it has still retained many characteristics almost certainly possessed by our primitive bilateral ancestor. The different parts of hydra were arranged like those of most flowers, around one main vertical axis; it was thus radiate in structure, having neither front nor rear, right nor left side. But our little turbellaria, while still without a head, has one end which goes first and can be called the front end. The upper or dorsal surface is usually more colored with pigment cells than the lower or ventral surface, on which is the mouth. It has also a right and left side.
It is thus bilateral.
The gastraea swam by cilia, little eyelash-like processes which urge the animal forward like a myriad of microscopic oars. In our bodies they are sometimes used to keep up a current, _e.g._, to remove foreign particles from the lungs. The turbellaria is still covered with cilia, probably an inheritance from the gastraea; for, while in smaller forms they may still be the princ.i.p.al means of locomotion, in larger ones the muscles are beginning to a.s.sume this function and the animal moves by writhing. The bilateral symmetry has arisen in connection with this mode of locomotion and is thus a mark of important progress.
In the turbellaria we find for the first time a true body-wall distinct from underlying organs. The outer layer of this is a ciliated epithelium or layer of cells. Under this an elastic membrane may occur. Then come true body muscles, running transversely, longitudinally and dorso-ventrally. Between the external transverse and the internal longitudinal layers we often find two muscular layers whose fibres run diagonally. The body is well provided with muscles, but their arrangement is still far from economical or effective.
Within the body-wall is the parenchym. This is a spongy ma.s.s of connectile tissue in which the other organs are embedded. The mouth lies in the middle, or near the front of the ventral surface. The intestine varies in form, but is provided with its own layers of longitudinal and transverse muscles, and usually has paired pouches extending out from it into the body parenchym. These seem to distribute the dissolved nutriment; hence the whole cavity is still often called a gastro-vascular cavity as serving both digestion and circulation. There is no a.n.a.l opening, but indigestible material is still cast out through the mouth.
The animal can gain sufficient oxygen to supply its muscles and nerves, which are the princ.i.p.al seats of combustion, through the external surface. It has, therefore, no special respiratory organs.
But the waste matter of the muscles cannot escape so easily, for these are becoming deeper seated. Hence we find an excretory system consisting of two tubes with many branches in the parenchym, and discharging at the rear end of the body. This again is a sign that the muscles are becoming more important, for the excretory system is needed mainly to remove their waste. These tubes maybe only greatly enlarged glands of the skin.
[Ill.u.s.tration: 5. TURBELLARIAN. LANG.
_va_ and _ha_, front and rear branches of gastro-vascular cavity; _ph_, pharynx. The dark oval with fine branches represents the nervous system.]
The nervous system consists of a plexus of fibres and cells, the cells originating impulses and the fibres conveying them. But this much was present in hydra also. Here the front end of the body goes foremost and is continually coming in contact with new conditions.
Here the lookout for food and danger must be kept. Hence, as a result of constant exercise, or selection, or both, the nerve-plexus has thickened at this point into a little compact ma.s.s of cells and fibres called a ganglion. And because this ganglion throughout higher forms usually lies over the oesophagus, it is called the supra-oesophogeal ganglion. This is the first faint and dim prophecy of a brain, and it sends its nerves to the front end of the body. But there run from it to the rear end of the body four to eight nerve-cords, consisting of bundles of nerve-threads like our nerves, but overlaid with a coating of ganglion cells capable of originating impulses. These cords are, therefore, like the plexus from which they have condensed, both nerves and centres; differentiation has not gone so far as at the front of the body.
Sense organs are still very rudimentary. Special cells of the skin have been modified into neuro-epithelial cells, having sensory hairs protruding from them and nerve-fibrils running from their bases.
[Ill.u.s.tration: 6. CROSS-SECTION OF TURBELLARIAN. HATSCHEK, FROM JIJIMA.
_e_, external skin; _rm_, lateral muscles; _la_ and _li_, longitudinal muscles; _mdv_, dorso-ventral muscles; _pa_, parenchyma; _h_, t.e.s.t.i.c.l.e; _ov_, oviduct; _dt_, yolk-gland; _n_, ventral nerve; _i_, gastro-vascular cavity.]
In a very few turbellaria we find otolith vesicles. These are little sacks in the skin, lined with neuro-epithelial cells and having in the middle a little concretion of carbonate of lime hung on rather a stiffer hair, like a clapper in a bell. Such organs serve in higher animals as organs of hearing, for the sensory hairs are set in vibration by the sound-waves. It is quite as probable that they here serve as organs for feeling the slightest vibrations in the surrounding water, and thus giving warning of approaching food or danger. The animal has also eyes, and these may be very numerous. They are not able to form images of external objects, but only of perceiving light and the direction of its source. A little group of these eyes lies directly over the brain, near the front end of the body; the others are distributed around the front or nearly the whole margin of the body.
The turbellaria, doubtless, have the sense of smell, although we can discover no special olfactory organ. This sense would seem to be as old as protoplasm itself.
This distribution of the eyes around a large portion of the margin, and certain other characteristics of the adult structure and of the embryonic development, are very interesting, as giving hints of the development of the turbellaria from some radiate ancestor. The mouth is in a most unfavorable position, in or near the middle of the body, rarely at the front end, as the animal has to swim over its food before it can grasp it. The animal only slowly rids itself of old disadvantageous form and structure and adapts itself completely to a higher mode of life.
By far the most highly developed system in the body is the reproductive. It is doubtful whether any animal, except, perhaps, the mollusk, has as complicated and highly developed reproductive organs. By markedly higher forms they certainly grow simpler.
And here we must notice certain general considerations. We found that reproduction in the amoeba could be defined as growth beyond the limit normal to the individual. This form of growth benefits especially the species. The needs and expenses of the individual will therefore first be met and then the balance be devoted to reproduction. Now the income of the animal is proportional to its surface, its expense to its ma.s.s, and activity. And the ratio of surface to ma.s.s is most favorable in the smallest animals.[A] Hence, smaller animals, as a rule, increase faster than larger ones; and this is only one ill.u.s.tration of the fact that great size in an animal is anything but an unmixed advantage to its possessor. But muscles and nerves are the most expensive systems; here most of the food is burned up. Hence energetic animals have a small balance remaining. Now the turbellarian is small and sluggish, with a fair digestive system. With a great amount of nutriment at its disposal the reproductive system came rapidly to a high development, and relatively to other organs stands higher than it almost ever will again.
[Footnote A: Cf. p. 35.]
It is only fair to state that good authorities hold that so primitive an animal could not originally have had so highly developed a system, and that this characteristic must be acquired, not ancestral.
That certain portions of it may be later developments may be not only possible but probable. But anyone who has carefully studied the different groups of worms, will, I think, readily grant that in the stage of these flat worms reproduction was the dominant function, which had most nearly attained its possible height of development.
From this time on the muscular and nervous systems were to claim an ever-increasing share of the nutriment, and the balance for reproduction is to grow smaller.
At the close of this lecture I wish to describe very briefly a hypothetical form. It no longer exists; perhaps it never did. But many facts of embryology and comparative anatomy point to such a form as a very possible ancestor of all forms higher than flat worms, viz., mollusks, arthropods, and vertebrates.
It was probably rather long and cylindrical, resembling a small and short earthworm in shape. The skin may have been much like that of turbellaria. Within this the muscles run in only two-directions--longitudinally and transversely. Between these and the intestine is a cavity--the perivisceral cavity--like that of our own bodies, but filled with a nutritive fluid like our lymph. This cavity seems to have developed by the expansion and cutting off of the paired lateral outgrowths of the digestive system of some old flat worm. But other modes of development are quite possible. The intestine has now an a.n.a.l opening at or near the rear end of the body. The food moves only from front to rear, and reaches each part always in a certain condition. Digestion proper and absorption have been distributed to different cells, and the work is better done.
Three portions can be readily distinguished: fore-intestine with the mouth, mid-intestine, as the seat of digestion and absorption, and hind-intestine, or r.e.c.t.u.m, with the a.n.a.l opening. The front and hind-intestine are lined with infolded outer skin.
The nervous system consists of a supra-oesophageal ganglion with four posterior nerve-cords--one dorsal, two lateral, and one (or perhaps two) ventral. There were probably also remains of the old plexus, but this is fast disappearing. The excretory system consists of a pair of tubes discharging through the sides of the body-wall, and having each a ciliated, funnel-shaped opening in the perivisceral cavity. These have received the name of nephridia.
Through these also the eggs and spermatozoa are discharged. The reproductive organs are modified patches of the peritoneum, or lining of the perivisceral cavity.
The number of muscles or muscular layers has been reduced in this animal. But such a reduction in the number of like parts in any animal is a sign of progress. And the longitudinal muscles have increased in size and strength, and the animal moves by writhing.
Such a worm has the general plan of the body of the higher forms fairly well, though rudely, sketched. Many improvements will come, and details be added. But the rudiments of the trunk of even our own bodies are already visible. Head, in any proper sense of the term, and skeleton are still lacking; they remain to be developed.
And yet, taking the most hopeful view possible concerning the animal kingdom, its prospects of attaining anything very lofty seem at this point poor. Its highest representative is a headless trunk, without skeleton or legs. It has no brain in any proper sense of the word, its sense-organs are feeble; it moves by writhing. Its life is devoted to digestion and reproduction. Whatever higher organs it has are subsidiary to these lower functions. And yet it has taken ages on ages to develop this much. If _this_ is the highest visible result of ages on ages of development, what hope is there for the future? Can such a thing be the ancestor of a thinking, moral, religious person, like man? "That is not first which is spiritual, but that which is natural (animal, sensuous); and afterward that which is spiritual." First, in order of time, must come the body, and then the mind and spirit shall be enthroned in it. The little knot of nervous material which forms the supra-oesophageal ganglion is so small that it might easily escape our notice; but it is the promise of an infinite future. The atom of nervous power shall increase until it subdues and dominates the whole ma.s.s.
CHAPTER III
WORMS TO VERTEBRATES: SKELETON AND HEAD
In tracing the genealogy of any American family it is often difficult or impossible to say whether a certain branch is descended from John Oldworthy or his cousin or second cousin. In the latter cases to find the common ancestor we must go back to the grandfather or great-grandfather. The same difficulty, but greatly enhanced, meets us when we try to make a genealogical tree of the animal kingdom. Thus it seems altogether probable that all higher forms are descended from an ancestor of the same general structure and grade of organization as the turbellaria, although probably free swimming, and hence with somewhat different form and development, especially of the muscular system. It seems to me altogether probable that all, except possibly Mollusca, are descended from a common ancestor closely resembling the schematic worm last described. Some would, however, maintain that they diverged rather earlier than even the turbellaria; others after the schematic worm, if such ever existed.
As far as our argument is concerned it makes little difference which of these views we adopt.
From our turbellaria, or possibly from some even more primitive ancestor, many lines diverged. And this was to be expected. The coelenterata, as we saw in hydra, had developed rude digestive and reproductive systems. The higher groups of this kingdom had developed all, or nearly all, the tissues used in building the bodies of higher animals--muscular, reproductive, connectile, glandular, nervous, etc. But these are mostly very diffuse. The muscular fibrils of a jelly-fish are mostly isolated or parallel in bands, rarely in compact well-defined bundles. The tissues have generally not yet been moulded into compact ma.s.ses of definite form.
There are as yet very few structures to which we can give the name of organs. To form organs and group them in a body of compact definite form was the work pre-eminently of worms. The material for the building was ready, but the architecture of the bilateral animal was not even sketched. And different worms were their own architects, untrammelled by convention or heredity, hence they built very different, sometimes almost fantastic, structures.
We must remember, too, the great age of this group. They are present in highly modified forms in the very oldest palaeozoic strata, and probably therefore came into existence as the first traces of continental areas were beginning to rise above the primeval ocean.
They are literally "older than the hills." They were exposed to a host of rapidly changing conditions, very different in different areas. This prepares us for the fact that the worms represent a stage in animal life corresponding fairly well to the Tower of Babel in biblical history. The animal kingdom seems almost to explode into a host of fragments. Our genealogical tree fairly bristles with branches, but the branches do not seem to form any regular whorls or spirals. Few of them have developed into more than feeble growths.
They now contain generally but few species. Many of them are largely or entirely parasitic, and in connection with this mode of life have undergone modifications and degeneration which make it exceedingly difficult to decipher their descent or relationships.
Four of these branches have reached great prominence in numbers and importance. One or two others were formerly equally numerous and have since become almost extinct; so the brachiopoda, which have been almost entirely replaced by mollusks. The same may very possibly be true of others. For of the amount of extinction of larger groups we have generally but an exceedingly faint conception.
Indeed in this respect the worms have been well compared to the relics which fill the shelves of one of our grandmother's china-closets.
The four great branches are the echinoderms, mollusks, articulates, and vertebrates. The echinoderms, including starfishes, sea-urchins, and others straggled early from the great army. We know as yet almost nothing of their history; when deciphered it will be as strange as any romance. The vertebrates are of course the most important line, as including the ancestors of man. But we must take a little glance at mollusks, including our clams, snails, and cuttle-fishes; and at the articulates, including annelids and culminating in insects. The molluscan and articulate lines, though divergent, are of great importance to us as throwing a certain amount of light on vertebrate development; and still more as showing how a certain line of development may seem, and at first really be, advantageous, and still lead to degeneration, or at best to but partial success.
When we compare the forms which represent fairly well the direction of development of these three lines, a snail or a clam with an insect and a fish, we find clearly, I think, that the fundamental anatomical difference lies in the skeleton; and that this resulted from, and almost irrevocably fixed, certain habits of life.
We may picture to ourselves the primitive ancestor of mollusks as a worm having the short and broad form of the turbellaria, but much thicker or deeper vertically. A fuller description can be found in the "Encyclopaedia Britannica," Art., Mollusca. It was hemi-ovoid in form. It had apparently the perivisceral cavity and nephridia of the schematic worm, and a circulatory system. In this latter respect it stood higher than any form which we have yet studied. Its nervous system also was rather more advanced. It had apparently already taken to a creeping mode of life and the muscles of its ventral surface were strongly developed, while its exposed and far less muscular dorsal surface was protected by a cap-like sh.e.l.l covering the most important internal organs. But the integument of the whole dorsal surface was, as is not uncommon in invertebrates, hardening by the deposition of carbonate of lime in the integument. And this in time increased to such an extent as to replace the primitive, probably h.o.r.n.y, sh.e.l.l.
Into the anatomy of this animal or of its descendants we have no time to enter, for here we must be very brief. We have already noticed that the most important viscera were lodged safely under the sh.e.l.l. And as these increased in size or were crowded upward by the muscles of the creeping disk, their portion of the body grew upward in the form of a "visceral hump." Apparently the animal could not increase much in length and retain the advantage of the protection of the sh.e.l.l; and the sh.e.l.l was the dominating structure. It had entered upon a defensive campaign. Motion, slow at the outset, became more difficult, and the protection of the sh.e.l.l therefore all the more necessary. The sh.e.l.l increased in size and weight and motion became almost impossible. The snail represents the average result of the experiment. It can crawl, but that is about all; it is neither swift nor energetic. Even the earthworm can outcrawl it. It has feelers and eyes, and is thus better provided with sense-organs than almost any worm. It has a supra-oesophageal ganglion of fair size.
The clams and oysters show even more clearly what we might call the logical results of molluscan structure. They increased the sh.e.l.l until it formed two heavy "valves" hanging down on each side of the body and completely enclosing it. They became almost sessile, living generally buried in the mud and gaining their food, consisting mostly of minute particles of organic matter, by means of currents created by cilia covering the large curtain-like gills. Their muscular system disappeared except in the ploughshare-shaped "foot"