(FIGURES 2.222 TO 2.224. Transverse sections of young Amphioxus-larvae (diagrammatic, from Ralph.) (Cf. also Figure 2.216.) In Figure 2.222 there is free communication from without with the gut-cavity (D) through the gill-clefts (K). In Figure 2.223 the lateral folds of the body-wall, or the gill-covers, which grow downwards, are formed. In Figure 2.224 these lateral folds have united underneath and joined their edges in the middle line of the ventral side (R seam). The respiratory water now passes from the gut-cavity (D) into the mantle-cavity (A). The letters have the same meaning throughout: N medullary tube, Ch chorda, M lateral muscles, Lh body-cavity, G part of the body-cavity in which the sexual organs are subsequently formed. D gut-cavity, clothed with the gut-gland layer (a). A mantle-cavity, K gill-clefts, b = E epidermis, E1 the same as visceral epithelium of the mantle-cavity, E2 as parietal epithelium of the mantle-cavity.)
At an early stage of embryonic development the structure of the Amphioxus-larva is substantially the same as the ideal picture we have previously formed of the "Primitive Vertebrate" (Figures 1.98 to 1.102). But the body afterwards undergoes various modifications, especially in the fore-part. These modifications do not concern us, as they depend on special adaptations, and do not affect the hereditary vertebrate type. When the free-swimming Amphioxus-larva is three months old, it abandons its pelagic habits and changes into the young animal that lives in the sand. In spite of its smallness (one-eighth of an inch), it has substantially the same structure as the adult. As regards the remaining organs of the Amphioxus, we need only mention that the gonads or sexual glands are developed very late, immediately out of the inner cell-layer of the body-cavity. Although we can find afterwards no continuation of the body-cavity (Figure 2.216 U) in the lateral walls of the mantle-cavity, in the gill-covers or mantle-folds (Figure 2.224 U), there is one present in the beginning (Figure 2.224 Lh). The sexual cells are formed below, at the bottom of this continuation (Figure 2.224 S). For the rest, the subsequent development into the adult Amphioxus of the larva we have followed is so simple that we need not go further into it here.
We may now turn to the embryology of the Ascidia, an animal that seems to stand so much lower and to be so much more simply organised, remaining for the greater part of its life attached to the bottom of the sea like a shapeless lump. It was a fortunate accident that Kowalevsky first examined just those larger specimens of the Ascidiae that show most clearly the relationship of the vertebrates to the invertebrates, and the larvae of which behave exactly like those of the Amphioxus in the first stages of development. This resemblance is so close in the main features that we have only to repeat what we have already said of the ontogenesis of the Amphioxus.
The ovum of the larger Ascidia (Phallusia, Cynthia, etc.) is a simple round cell of 1/250 to 1/125 of an inch in diameter. In the thick fine-grained yelk we find a clear round germinal vesicle of about 1/750 of an inch in diameter, and this encloses a small embryonic spot or nucleolus. Inside the membrane that surrounds the ovum, the stem-cell of the Ascidia, after fecundation, passes through just the same metamorphoses as the stem-cell of the Amphioxus. It undergoes total segmentation; it divides into two, four, eight, sixteen, thirty-two cells, and so on. By continued total cleavage the morula, or mulberry-shaped cluster of cells, is formed. Fluid gathers inside it, and thus we get once more a globular vesicle (the blastula); the wall of this is a single stratum of cells, the blastoderm. A real gastrula (a simple bell-gastrula) is formed from the blastula by invagination, in the same way as in the amphioxus.
Up to this there is no definite ground in the embryology of the Ascidiae for bringing them into close relationship with the Vertebrates; the same gastrula is formed in the same way in many other animals of different stems. But we now find an embryonic process that is peculiar to the Vertebrates, and that proves irrefragably the affinity of the Ascidiae to the Vertebrates. From the epidermis of the gastrula a medullary tube is formed on the dorsal side, and, between this and the primitive gut, a chorda; these are the organs that are otherwise only found in Vertebrates. The formation of these very important organs takes place in the Ascidia-gastrula in precisely the same way as in that of the Amphioxus. In the Ascidia (as in the other case) the oval gastrula is first flattened on one side—the subsequent dorsal side. A groove or furrow (the medullary groove) is sunk in the middle line of the flat surface, and two parallel longitudinal swellings arise on either side from the skin layer. These medullary swellings join together over the furrow, and form a tube; in this case, again, the neural or medullary tube is at first open in front, and connected with the primitive gut behind by the neurenteric canal. Further, in the Ascidia-larva also the two permanent apertures of the alimentary canal only appear later, as independent and new formations. The permanent mouth does not develop from the primitive mouth of the gastrula; this primitive mouth closes up, and the later anus is formed near it by invagination from without, on the hinder end of the body, opposite to the aperture of the medullary tube.
During these important processes, that take place in just the same way in the Amphioxus, a tail-like projection grows out of the posterior end of the larva-body, and the larva folds itself up within the round ovolemma in such a way that the dorsal side is curved and the tail is forced on to the ventral side. In this tail is developed—starting from the primitive gut—a cylindrical string of cells, the fore end of which pushes into the body of the larva, between the alimentary canal and the neural canal, and is no other than the chorda dorsalis. This important organ had hitherto been found only in the Vertebrates, not a single trace of it being discoverable in the Invertebrates. At first the chorda only consists of a single row of large entodermic cells. It is afterwards composed of several rows of cells. In the Ascidia-larva, also, the chorda develops from the dorsal middle part of the primitive gut, while the two coelom-pouches detach themselves from it on both sides. The simple body-cavity is formed by the coalescence of the two.
When the Ascidia-larva has attained this stage of development it begins to move about in the ovolemma. This causes the membrane to burst. The larva emerges from it, and swims about in the sea by means of its oar-like tail. These free-swimming larvae of the Ascidia have been known for a long time. They were first observed by Darwin during his voyage round the world in 1833. They resemble tadpoles in outward appearance, and use their tails as oars, as the tadpoles do. However, this lively and highly-developed condition does not last long. At first there is a progressive development; the foremost part of the medullary tube enlarges into a brain, and inside this two single sense-organs are developed, a dorsal auditory vesicle and a ventral eye. Then a heart is formed on the ventral side of the animal, or the lower wall of the gut, in the same simple form and at the same spot at which the heart is developed in man and all the other vertebrates. In the lower muscular wall of the gut we find a weal-like thickening, a solid, spindle-shaped string of cells, which becomes hollow in the centre; it begins to contract in different directions, now forward and now backward, as is the case with the adult Ascidia. In this way the sanguineous fluid accumulated in the hollow muscular tube is driven in alternate directions into the blood-vessels, which develop at both ends of the cardiac tube. One principal vessel runs along the dorsal side of the gut, another along its ventral side. The former corresponds to the aorta and the dorsal vessel in the worms. The other corresponds to the subintestinal vein and the ventral vessel of the worms.
With the formation of these organs the progressive development of the Ascidia comes to an end, and degeneration sets in. The free-swimming larva sinks to the floor of the sea, abandons its locomotive habits, and attaches itself to stones, marine plants, mussel-shells, corals, and other objects; this is done with the part of the body that was foremost in movement. The attachment is effected by a number of out-growths, usually three, which can be seen even in the free-swimming larva. The tail is lost, as there is no further use for it. It undergoes a fatty degeneration, and disappears with the chorda dorsalis. The tailless body changes into an unshapely tube, and, by the atrophy of some parts and the modification of others, gradually assumes the appearance we have already described.
(FIGURE 2.225. An Appendicaria (Copelata), seen from the left. m mouth, k branchial gut, o gullet, v stomach, a anus, n brain (ganglion above the gullet), g auditory vesicle, f ciliated groove under the gills, h heart, t testicles, e ovary, c chorda, s tail.)
Among the living Tunicates there is a very interesting group of small animals that remain throughout life at the stage of development of the tailed, free Ascidia-larva, and swim about briskly in the sea by means of their broad oar-tail. These are the remarkable Copelata (Appendicaria and Vexillaria, Figure 2.225). They are the only living Vertebrates that have throughout life a chorda dorsalis and a neural string above it; the latter must be regarded as the prolongation of the cerebral ganglion and the equivalent of the medullary tube. Their branchial gut also opens directly outwards by a pair of branchial clefts. These instructive Copelata, comparable to permanent Ascidia-larvae, come next to the extinct Prochordonia, those ancient worms which we must regard as the common ancestors of the Tunicates and Vertebrates. The chorda of the Appendicaria is a long, cylindrical string (Figure 2.225 c), and serves as an attachment for the muscles that work the flat oar-tail.
Among the various modifications which the Ascidia-larva undergoes after its establishment at the sea-floor, the most interesting (after the loss of the axial rod) is the atrophy of one of its chief organs, the medullary tube. In the Amphioxus the spinal marrow continues to develop, but in the Ascidia the tube soon shrinks into a small and insignificant nervous ganglion that lies above the mouth and the gill-crate, and is in accord with the extremely slight mental power of the animal. This insignificant relic of the medullary tube seems to be quite beyond comparison with the nervous centre of the vertebrate, yet it started from the same structure as the spinal cord of the Amphioxus. The sense-organs that had been developed in the fore part of the neural tube are also lost; no trace of which can be found in the adult Ascidia. On the other hand, the alimentary canal becomes a most extensive organ. It divides presently into two sections—a wide fore or branchial gut that serves for respiration, and a narrower hind or hepatic gut that accomplishes digestion. The branchial or head-gut of the Ascidia is small at first, and opens directly outwards only by a couple of lateral ducts or gill-clefts—a permanent arrangement in the Copelata. The gill-clefts are developed in the same way as in the Amphioxus. As their number greatly increases we get a large gill-crate, pierced like lattice work. In the middle line of its ventral side we find the hypobranchial groove. The mantle or cloaca-cavity (the atrium) that surrounds the gill-crate is also formed in the same way in the Ascidia as in the Amphioxus. The ejection-opening of this peribranchial cavity corresponds to the branchial pore of the Amphioxus. In the adult Ascidia the branchial gut and the heart on its ventral side are almost the only organs that recall the original affinity with the vertebrates.
The further development of the Ascidia in detail has no particular interest for us, and we will not go into it. The chief result that we obtain from its embryology is the complete agreement with that of the Amphioxus in the earliest and most important embryonic stages. They do not begin to diverge until after the medullary tube and alimentary canal, and the axial rod with the muscles between the two, have been formed. The Amphioxus continues to advance, and resembles the embryonic forms of the higher vertebrates; the Ascidia degenerates more and more, and at last, in its adult condition, has the appearance of a very imperfect invertebrate.
If we now look back on all the remarkable features we have encountered in the structure and the embryonic development of the Amphioxus and the Ascidia, and compare them with the features of man's embryonic development which we have previously studied, it will be clear that I have not exaggerated the importance of these very interesting animals. It is evident that the Amphioxus from the vertebrate side and the Ascidia from the invertebrate form the bridge by which we can span the deep gulf that separates the two great divisions of the animal kingdom. The radical agreement of the lancelet and the sea-squirt in the first and most important stages of development shows something more than their close anatomic affinity and their proximity in classification; it shows also their real blood-relationship and their common origin from one and the same stem-form. In this way, it throws considerable light on the oldest roots of man's genealogical tree.
CHAPTER 2.18. DURATION OF THE HISTORY OF OUR STEM.
Our comparative investigation of the anatomy and ontogeny of the Amphioxus and Ascidia has given us invaluable assistance. We have, in the first place, bridged the wide gulf that has existed up to the present between the Vertebrates and Invertebrates; and, in the second place, we have discovered in the embryology of the Amphioxus a number of ancient evolutionary stages that have long since disappeared from human embryology, and have been lost, in virtue of the law of curtailed heredity. The chief of these stages are the spherical blastula (in its simplest primary form), and the succeeding archigastrula, the pure, original form of the gastrula which the Amphioxus has preserved to this day, and which we find in the same form in a number of Invertebrates of various classes. Not less important are the later embryonic forms of the coelomula, the chordula, etc.
Thus the embryology of the Amphioxus and the Ascidia has so much increased our knowledge of man's stem-history that, although our empirical information is still very incomplete, there is now no defect of any great consequence in it. We may now, therefore, approach our proper task, and reconstruct the phylogeny of man in its chief lines with the aid of this evidence of comparative anatomy and ontogeny. In this the reader will soon see the immense importance of the direct application of the biogenetic law. But before we enter upon the work it will be useful to make a few general observations that are necessary to understand the processes aright.
We must say a few words with regard to the period in which the human race was evolved from the animal kingdom. The first thought that occurs to one in this connection is the vast difference between the duration of man's ontogeny and phylogeny. The individual man needs only nine months for his complete development, from the fecundation of the ovum to the moment when he leaves the maternal womb. The human embryo runs its whole course in the brief space of forty weeks (as a rule, 280 days). In many other mammals the time of the embryonic development is much the same as in man—for instance, in the cow. In the horse and ass it takes a little longer, forty-three to forty-five weeks; in the camel, thirteen months. In the largest mammals, the embryo needs a much longer period for its development in the womb—a year and a half in the rhinoceros, and ninety weeks in the elephant. In these cases pregnancy lasts twice as long as in the case of man, or one and three-quarter years. In the smaller mammals the embryonic period is much shorter. The smallest mammals, the dwarf-mice, develop in three weeks; hares in four weeks, rats and marmots in five weeks, the dog in nine, the pig in seventeen, the sheep in twenty-one and the goat in thirty-six. Birds develop still more quickly. The chick only needs, in normal circumstances, three weeks for its full development. The duck needs twenty-five days, the turkey twenty-seven, the peacock thirty-one, the swan forty-two, and the cassowary sixty-five. The smallest bird, the humming-bird, leaves the egg after twelve days. Hence the duration of individual development within the foetal membranes is, in the mammals and birds, clearly related to the absolute size of the body of the animal in question. But this is not the only determining feature. There are a number of other circumstances that have an influence on the period of embryonic development. In the Amphioxus the earliest and most important embryonic processes take place so rapidly that the blastula is formed in four hours, the gastrula in six, and the typical vertebrate form in twenty-four.
In every case the duration of ontogeny shrinks into insignificance when we compare it with the enormous period that has been necessary for phylogeny, or the gradual development of the ancestral series. This period is not measured by years or centuries, but by thousands and millions of years. Many millions of years had to pass before the most advanced vertebrate, man, was evolved, step by step, from his ancient unicellular ancestors. The opponents of evolution, who declare that this gradual development of the human form from lower animal forms, and ultimately from a unicellular organism, is an incredible miracle, forget that the same miracle takes place within the space of mine months in the embryonic development of every human being. Each of us has, in the forty weeks—properly speaking, in the first four weeks—of his development in the womb, passed through the same series of transformations that our animal ancestors underwent in the course of millions of years.
It is impossible to determine even approximately, in hundreds or even thousands of years, the real and absolute duration of the phylogenetic period. But for some time now we have, through the research of geologists, been in a position to assign the relative length of the various sections of the organic history of the earth. The immediate data for determining this relative length of the geological periods are found in the thickness of the sedimentary strata—the strata that have been formed at the bottom of the sea or in fresh water from the mud or slime deposited there. These successive layers of limestone, sandstone, slate, marl, etc., which make up the greater part of the rocks, and are often several thousand feet thick, give us a standard for computing the relative length of the various periods.
To make the point quite clear, I must say a word about the evolution of the earth in general, and point out briefly the chief features of the story. In the first place, we encounter the principle that on our planet organic life began to exist at a definite period. That statement is no longer disputed by any competent geologist or biologist. The organic history of the earth could not commence until it was possible for water to settle on our planet in fluid condition. Every organism, without exception, needs fluid water as a condition of existence, and contains a considerable quantity of it. Our own body, when fully formed, contains sixty to seventy per cent of water in its tissues, and only thirty to forty per cent of solid matter. There is even more water in the body of the child, and still more in the embryo. In the earlier stages of development the human foetus contains more than ninety per cent of water, and not ten per cent of solids. In the lower marine animals, especially certain medusae, the body consists to the extent of more than ninety-nine per cent of sea-water, and has not one per cent of solid matter. No organism can exist or discharge its functions without water. No water, no life!
But fluid water, on which the existence of life primarily depends, could not exist on our planet until the temperature of the surface of the incandescent sphere had sunk to a certain point. Up to that time it remained in the form of steam. But as soon as the first fluid water could be condensed from the envelope of steam, it began its geological action, and has continued down to the present day to modify the solid crust of the earth. The final outcome of this incessant action of the water—wearing down and dissolving the rocks in the form of rain, hail, snow, and ice, as running stream or boiling surge—is the formation of mud. As Huxley says in his admirable Lectures on the Causes of Phenomena in Organic Nature, the chief document as to the past history of our earth is mud; the question of the history of past ages resolves itself into a question about the formation of mud.
As I have said, it is possible to form an approximate idea of the relative age of the various strata by comparing them at different parts of the earth's surface. Geologists have long been agreed that there is a definite historical succession of the different strata. The various superimposed layers correspond to successive periods in the organic history of the earth, in which they were deposited in the form of mud at the bottom of the sea. The mud was gradually converted into stone. This was lifted out of the water owing to variations in the earth's surface, and formed the mountains. As a rule, four or five great divisions are distinguished in the organic history of the earth, corresponding to the larger and smaller groups of the sedimentary strata. The larger periods are then sub-divided into a series of smaller ones, which usually number from twelve to fifteen. The comparative thickness of the groups of strata enables us to make an approximate calculation of the relative length of these various periods of time. We cannot say, it is true, "In a century a stratum of a certain thickness (about two feet) is formed on the average; therefore, a layer 1000 feet thick must be 500,000 years old." Different strata of the same thickness may need very different periods for their formation. But from the thickness or size of the stratum we can draw some conclusion as to the RELATIVE length of the period.
The first and oldest of the four or five chief divisions of the organic history of the earth is called the primordial, archaic, or archeozoic period. If we compute the total average thickness of the sedimentary strata at about 130,000 feet, this first period comprises 70,000 feet, or the greater part of the whole. For this and other reasons we may at once conclude that the corresponding primordial or archeolithic period must have been in itself much longer than the whole of the remaining periods together, from its close to the present day. It was probably much longer than the figures I have quoted (7 : 6) indicate—possibly 9 : 6. Of late years the thickness of the archaic rocks has been put at 90,000 feet.
SYNOPSIS OF THE PALEONTOLOGICAL FORMATIONS, OR THE FOSSILIFEROUS STRATA OF THE CRUST.
COLUMN 1 : Groups (V. down to I.).
COLUMN 2 : Systems (XIV. down to I.).
COLUMN 3 : Formations (38 down to 1).
COLUMN 4 : Synonyms of Formations.
V. Anthropolithic group, or anthropozoic (quaternary) group of strata
: XIV. Recent (alluvium) : 38. Present : Upper alluvial.
V. Anthropolithic group, or anthropozoic (quaternary) group of strata
: XIV. Recent (alluvium) : 37. Recent : Lower alluvial.
V. Anthropolithic group, or anthropozoic (quaternary) group of strata
: XIII. Pleistocene (diluvium) : 36. Post-glacial : Upper diluvial.
V. Anthropolithic group, or anthropozoic (quaternary) group of strata
: XIII. Pleistocene (diluvium) : 35. Glacial : Lower diluvial.
IV. Cenolithic group, or cenozoic (tertiary) group of strata : XII.
Pliocene (neo-tertiary) : 34. Arverne : Upper pliocene.
IV. Cenolithic group, or cenozoic (tertiary) group of strata : XII.
Pliocene (neo-tertiary) : 33. Subapennine : Lower pliocene.
IV. Cenolithic group, or cenozoic (tertiary) group of strata : XI.
Miocene (middle tertiary) : 32. Falun : Upper miocene.
IV. Cenolithic group, or cenozoic (tertiary) group of strata : XI.
Miocene (middle tertiary) : 31. Limbourg : Lower miocene.
IV. Cenolithic group, or cenozoic (tertiary) group of strata : Xb.
Oligocene (old tertiary) : 30. Aquitaine : Upper oligocene.
IV. Cenolithic group, or cenozoic (tertiary) group of strata : Xb.
Oligocene (old tertiary) : 29. Ligurium : Lower oligocene.
IV. Cenolithic group, or cenozoic (tertiary) group of strata : Xa.
Eocene (primitive tertiary) : 28. Gypsum : Upper eocene.
IV. Cenolithic group, or cenozoic (tertiary) group of strata : Xa.
Eocene (primitive tertiary) : 27. Coarse chalk : Middle eocene.
IV. Cenolithic group, or cenozoic (tertiary) group of strata : Xa.
Eocene (primitive tertiary) : 26. London clay : Lower eocene.
III. Mesolithic group, or mesozoic (secondary) group of strata : IX.
Chalk (cretaceous) : 25. White chalk. : Upper cretaceous.
III. Mesolithic group, or mesozoic (secondary) group of strata : IX.
Chalk (cretaceous) : 24. Green Sand : Middle cretaceous.
III. Mesolithic group, or mesozoic (secondary) group of strata : IX.
Chalk (cretaceous) : 23. Neocomian : Lower cretaceous.
III. Mesolithic group, or mesozoic (secondary) group of strata : IX.
Chalk (cretaceous) : 22. Wealden : Weald-formation.
III. Mesolithic group, or mesozoic (secondary) group of strata : VIII.
Jurassic : 21. Portland : Upper oolithic.
III. Mesolithic group, or mesozoic (secondary) group of strata : VIII.
Jurassic : 20. Oxford : Middle oolithic.
III. Mesolithic group, or mesozoic (secondary) group of strata : VIII.
Jurassic : 19. Bath : Lower oolithic.
III. Mesolithic group, or mesozoic (secondary) group of strata : VIII.
Jurassic : 18. Lias : Liassic.
III. Mesolithic group, or mesozoic (secondary) group of strata : VII.
Triassic : 17. Keuper : Upper triassic.
III. Mesolithic group, or mesozoic (secondary) group of strata : VII.
Triassic : 16. Muschelkalk : Middle triassic.
III. Mesolithic group, or mesozoic (secondary) group of strata : VII.
Triassic : 15. Bunter : Lower triassic.
II. Paleolithic group, or paleozoic (primary) group of strata : VIb.
Permian : 14. Zechstein : Upper permian.
II. Paleolithic group, or paleozoic (primary) group of strata : VIb.
Permian : 13. Neurot sand : Lower permian.
II. Paleolithic group, or paleozoic (primary) group of strata : VIa. Carboniferous (coal-measures) : 12. Carboniferous sandstone : Upper carboniferous.
II. Paleolithic group, or paleozoic (primary) group of strata : VIa. Carboniferous (coal-measures) : 11. Carboniferous limestone : Lower carboniferous.
II. Paleolithic group, or paleozoic (primary) group of strata : V.
Devonian : 10. Pilton : Upper devonian.
II. Paleolithic group, or paleozoic (primary) group of strata : V.
Devonian : 9. Ilfracombe : Middle devonian.
II. Paleolithic group, or paleozoic (primary) group of strata : V.
Devonian : 8. Linton : Lower devonian.
II. Paleolithic group, or paleozoic (primary) group of strata : IV.
Silurian : 7. Ludlow : Upper silurian.
II. Paleolithic group, or paleozoic (primary) group of strata : IV.
Silurian : 6. Wenlock : Middle silurian.
II. Paleolithic group, or paleozoic (primary) group of strata : IV.
Silurian : 5. Llandeilo : Lower silurian.
I. Archeolithic group, or archeozoic (primordial) group of strata :
III. Cambrian : 4. Potsdam : Upper cambrian.
I. Archeolithic group, or archeozoic (primordial) group of strata :
III. Cambrian : 3. Longmynd : Lower cambrian.
I. Archeolithic group, or archeozoic (primordial) group of strata :
II. Huronian : 2. Labrador : Upper laurentian.
I. Archeolithic group, or archeozoic (primordial) group of strata : I.
Laurentian : 1. Ottawa : Lower laurentian.
The primordial period falls into three subordinate sections—the Laurentian, Huronian, and Cambrian, corresponding to the three chief groups of rocks that comprise the archaic formation. The immense period during which these rocks were forming in the primitive ocean probably comprises more than 50,000,000 years. At the commencement of it the oldest and simplest organisms were formed by spontaneous generation—the Monera, with which the history of life on our planet opened. From these were first developed unicellular organisms of the simplest character, the Protophyta and Protozoa (paulotomea, amoebae, rhizopods, infusoria, and other Protists). During this period the whole of the invertebrate ancestors of the human race were evolved from the unicellular organisms. We can deduce this from the fact that we already find remains of fossilised fishes (Selachii and Ganoids) towards the close of the following Silurian period. These are much more advanced and much younger than the lowest vertebrate, the Amphioxus, and the numerous skull-less vertebrates, related to the Amphioxus, that must have lived at that time. The whole of the invertebrate ancestors of the human race must have preceded these.
The primordial age is followed by a much shorter division, the paleozoic or Primary age. It is divided into four long periods, the Silurian, Devonian, Carboniferous, and Permian. The Silurian strata are particularly interesting because they contain the first fossil traces of vertebrates—teeth and scales of Selachii (Palaeodus) in the lower, and Ganoids (Pteraspis) in the upper Silurian. During the Devonian period the "old red sandstone" was formed; during the Carboniferous period were deposited the vast coal-measures that yield us our chief combustive material; in the Permian (or the Dyas), in fine, the new red sandstone, the Zechstein (magnesian limestone), and the Kupferschiefer (marl-slate) were formed. The collective depth of these strata is put at 40,000 to 45,000 feet. In any case, the paleozoic age, taken as a whole, was much shorter than the preceding and much longer than the subsequent periods. The strata that were deposited during this primary epoch contain a large number of fossils; besides the invertebrate species there are a good many vertebrates, and the fishes preponderate. There were so many fishes, especially primitive fishes (of the shark type) and plated fishes, during the Devonian, and also during the Carboniferous and Permian periods, that we may describe the whole paleozoic period as "the age of fishes." Among the paleozoic plated fishes or Ganoids the Crossopterygii and the Ctenodipterina (dipneusts) are of great importance.
During this period some of the fishes began to adapt themselves to living on land, and so gave rise to the class of the amphibia. We find in the Carboniferous period fossilised remains of five-toed amphibia, the oldest terrestrial, air-breathing vertebrates. These amphibia increase in variety in the Permian epoch. Towards the close of it we find the first Amniotes, the ancestors of the three higher classes of Vertebrates. These are lizard-like animals; the first to be discovered was the Proterosaurus, from the marl at Eisenach. The rise of the earliest Amniotes, among which must have been the common ancestor of the reptiles, birds, and mammals, is put back towards the close of the paleozoic age by the discovery of these reptile remains. The ancestors of our race during this period were at first represented by true fishes, then by dipneusts and amphibia, and finally by the earliest Amniotes, or the Protamniotes.
The third chief section of the organic history of the earth is the Mesozoic or Secondary period. This again is subdivided into three divisions Triassic, Jurassic, and Cretaceous. The thickness of the strata that were deposited in this period, from the beginning of the Triassic to the end of the Cretaceous period, is altogether about 15,000 feet, or not half as much as the paleozoic deposits. During this period there was a very brisk and manifold development in all branches of the animal kingdom. There were especially a number of new and interesting forms evolved in the vertebrate stem. Bony fishes (Teleostei) make their first appearance. Reptiles are found in extraordinary variety and number; the extinct giant-serpents (dinosauria), the sea-serpents (halisauria), and the flying lizards (pterosauria) are the most remarkable and best known of these. On account of this predominance of the reptile-class, the period is called "the age of reptiles." But the bird-class was also evolved during this period; they certainly originated from some division of the lizard-like reptiles. This is proved by the embryological identity of the birds and reptiles and their comparative anatomy, and, among other features, from the circumstance that in this period there were birds with teeth in their jaws and with tails like lizards (Archeopteryx, Odontornis).
Finally, the most advanced and (for us) the most important class of the vertebrates, the mammals, made their appearance during the mesozoic period. The earliest fossil remains of them were found in the latest Triassic strata—lower jaws of small ungulates and marsupials. More numerous remains are found a little later in the Jurassic, and some in the Cretaceous. All the mammal remains that we have from this section belong to the lower promammals and marsupials; among these were most certainly the ancestors of the human race. On the other hand, we have not found a single indisputable fossil of any higher mammal (a placental) in the whole of this period. This division of the mammals, which includes man, was not developed until later, towards the close of this or in the following period.
The fourth section of the organic history of the earth, the Tertiary or Cenozoic age, was much shorter than the preceding. The strata that were deposited during this period have a collective thickness of only about 3,000 feet. It is subdivided into four sections—the Eocene, Oligocene, Miocene, and Pliocene. During these periods there was a very varied development of higher plant and animal forms; the fauna and flora of our planet approached nearer and nearer to the character that they bear to-day. In particular, the most advanced class, the mammals, began to preponderate. Hence the Tertiary period may be called "the age of mammals." The highest section of this class, the placentals, now made their appearance; to this group the human race belongs. The first appearance of man, or, to be more precise, the development of man from some closely-related group of apes, probably falls in either the miocene or the pliocene period, the middle or the last section of the Tertiary period. Others believe that man properly so-called—man endowed with speech—was not evolved from the non-speaking ape-man (Pithecanthropus) until the following, the anthropozoic, age.
In this fifth and last section of the organic history of the earth we have the full development and dispersion of the various races of men, and so it is called the Anthropozoic as well as the Quaternary period. In the imperfect condition of paleontological and ethnographical science we cannot as yet give a confident answer to the question whether the evolution of the human race from some extinct ape or lemur took place at the beginning of this or towards the middle or the end of the Tertiary period. However, this much is certain: the development of civilisation falls in the anthropozoic age, and this is merely an insignificant fraction of the vast period of the whole history of life. When we remember this, it seems ridiculous to restrict the word "history" to the civilised period. If we divide into a hundred equal parts the whole period of the history of life, from the spontaneous generation of the first Monera to the present day, and if we then represent the relative duration of the five chief sections or ages, as calculated from the average thickness of the strata they contain, as percentages of this, we get something like the following relation:—
I. Archeolithic or archeozoic (primordial) age : 53 : 6.
II. Paleolithic or paleozoic (primary) age : 32 : 1.
III. Mesolithic or mesozoic (secondary) age : 11 : 5.
IV. Cenolithic or cenozoic (tertiary) age : 2 : 3
V. Anthropolithic or anthropozoic (quaternary) age : 0 : 5.
Total : 100 : 0.
In any case, the "historical period" is an insignificant quantity compared with the vast length of the preceding ages, in which there was no question of human existence on our planet. Even the important Cenozoic or Tertiary period, in which the first placentals or higher mammals appear, probably amounts to little over two per cent of the whole organic age.
Before we approach our proper task, and, with the aid of our ontogenetic acquirements and the biogenetic law, follow step by step the paleontological development of our animal ancestors, let us glance for a moment at another, and apparently quite remote, branch of science, a general consideration of which will help us in the solving of a difficult problem. I mean the science of comparative philology. Since Darwin gave new life to biology by his theory of selection, and raised the question of evolution on all sides, it has often been pointed out that there is a remarkable analogy between the development of languages and the evolution of species. The comparison is perfectly just and very instructive. We could hardly find a better analogy when we are dealing with some of the difficult and obscure features of the evolution of species. In both cases we find the action of the same natural laws.
All philologists of any competence in their science now agree that all human languages have been gradually evolved from very rudimentary beginnings. The idea that speech is a gift of the gods—an idea held by distinguished authorities only fifty years ago—is now generally abandoned, and only supported by theologians and others who admit no natural development whatever. Speech has been developed simultaneously with its organs, the larynx and tongue, and with the functions of the brain. Hence it will be quite natural to find in the evolution and classification of languages the same features as in the evolution and classification of organic species. The various groups of languages that are distinguished in philology as primitive, fundamental, parent, and daughter languages, dialects, etc., correspond entirely in their development to the different categories which we classify in zoology and botany as stems, classes, orders, families, genera, species, and varieties. The relation of these groups, partly co-ordinate and partly subordinate, in the general scheme is just the same in both cases; and the evolution follows the same lines in both.
When, with the assistance of this tree, we follow the formation of the various languages that have been developed from the common root of the ancient Indo-Germanic tongue, we get a very clear idea of their phylogeny. We shall see at the same time how analogous this is to the development of the various groups of vertebrates that have arisen from the common stem-form of the primitive vertebrate. The ancient Indo-Germanic root-language divided first into two principal stems—the Slavo-Germanic and the Aryo-Romanic. The Slavo-Germanic stem then branches into the ancient Germanic and the ancient Slavo-Lettic tongues; the Aryo-Romanic into the ancient Aryan and the ancient Greco-Roman. If we still follow the genealogical tree of these four Indo-Germanic tongues, we find that the ancient Germanic divides into three branches—the Scandinavian, the Gothic, and the German. From the ancient German came the High German and Low German; to the latter belong the Frisian, Saxon, and modern Low-German dialects. The ancient Slavo-Lettic divided first into a Baltic and a Slav language. The Baltic gave rise to the Lett, Lithuanian, and old-Prussian varieties; the Slav to the Russian and South-Slav in the south-east, and to the Polish and Czech in the west.
We find an equally prolific branching of its two chief stems when we turn to the other division of the Indo-Germanic languages. The Greco-Roman divided into the Thracian (Albano-Greek) and the Italo-Celtic. From the latter came the divergent branches of the Italic (Roman and Latin) in the south, and the Celtic in the north: from the latter have been developed all the British (ancient British, ancient Scotch, and Irish) and Gallic varieties. The ancient Aryan gave rise to the numerous Iranian and Indian languages.
This "comparative anatomy" and evolution of languages admirably illustrates the phylogeny of species. It is clear that in structure and development the primitive languages, mother and daughter languages, and varieties, correspond exactly to the classes, orders, genera, and species of the animal world. In both cases the "natural" system is phylogenetic. As we have been convinced from comparative anatomy and ontogeny, and from paleontology, that all past and living vertebrates descend from a common ancestor, so the comparative study of dead and living Indo-Germanic tongues proves beyond question that they are all modifications of one primitive language. This view of their origin is now accepted by all the chief philologists who have worked in this branch and are unprejudiced.
But the point to which I desire particularly to draw the reader's attention in this comparison of the Indo-Germanic languages with the branches of the vertebrate stem is, that one must never confuse direct descendants with collateral branches, nor extinct forms with living. This confusion is very common, and our opponents often make use of the erroneous ideas it gives rise to for the purpose of attacking evolution generally. When, for instance, we say that man descends from the ape, this from the lemur, and the lemur from the marsupial, many people imagine that we are speaking of the living species of these orders of mammals that they find stuffed in our museums. Our opponents then foist this idea on us, and say, with more astuteness than intelligence, that it is quite impossible; or they ask us, by way of physiological experiment, to turn a kangaroo into a lemur, a lemur into a gorilla, and a gorilla into a man! The demand is childish, and the idea it rests on erroneous. All these living forms have diverged more or less from the ancestral form; none of them could engender the same posterity that the stem-form really produced thousands of years ago.
It is certain that man has descended from some extinct mammal; and we should just as certainly class this in the order of apes if we had it before us. It is equally certain that this primitive ape descended in turn from an unknown lemur, and this from an extinct marsupial. But it is just as clear that all these extinct ancestral forms can only be claimed as belonging to the living order of mammals in virtue of their essential internal structure and their resemblance in the decisive anatomic characteristics of each ORDER. In external appearance, in the characteristics of the GENUS or SPECIES, they would differ more or less, perhaps very considerably, from all living representatives of those orders. It is a universal and natural procedure in phylogenetic development that the stem-forms themselves, with their specific peculiarities, have been extinct for some time. The forms that approach nearest to them among the living species are more or less—perhaps very substantially—different from them. Hence in our phylogenetic inquiry and in the comparative study of the living, divergent descendants, there can only be a question of determining the greater or less remoteness of the latter from the ancestral form. Not a single one of the older stem-forms has continued unchanged down to our time.
We find just the same thing in comparing the various dead and living languages that have developed from a common primitive tongue. If we examine our genealogical tree of the Indo-Germanic languages in this light, we see at once that all the older or parent tongues, of which we regard the living varieties of the stem as divergent daughter or grand-daughter languages, have been extinct for some time. The Aryo-Romanic and the Slavo-Germanic tongues have completely disappeared; so also the Aryan, the Greco-Roman, the Slavo-Lettic, and the ancient Germanic. Even their daughters and grand-daughters have been lost; all the living Indo-Germanic languages are only related in the sense that they are divergent descendants of common stem-forms. Some forms have diverged more, and some less, from the original stem-form.
This easily demonstrable fact illustrates very well the analogous case of the origin of the vertebrate species. Phylogenetic comparative philology here yields a strong support to phylogenetic comparative zoology. But the one can adduce more direct evidence than the other, as the paleontological material of philology—the old monuments of the extinct tongue—have been preserved much better than the paleontological material of zoology, the fossilised bones and imprints of vertebrates.
We may, however, trace man's genealogical tree not only as far as the lower mammals, but much further—to the amphibia, to the shark-like primitive fishes, and, in fine, to the skull-less vertebrates that closely resembled the Amphioxus. But this must not be understood in the sense that the existing Amphioxus, or the sharks or amphibia of to-day, can give us any idea of the external appearance of these remote stem-forms. Still less must it be thought that the Amphioxus or any actual shark, or any living species of amphibia, is a real ancestral form of the higher vertebrates and man. The statement can only rationally mean that the living forms I have referred to are COLLATERAL LINES that are much more closely related to the extinct stem-forms, and have retained the resemblance much better, than any other animals we know. They are still so like them in regard to their distinctive internal structure that we should put them in the same class with the extinct forms if we had these before us. But no direct descendants of these earlier forms have remained unchanged. Hence we must entirely abandon the idea of finding direct ancestors of the human race in their characteristic EXTERNAL FORM among the living species of animals. The essential and distinctive features that still connect living forms more or less closely with the extinct common stem-forms lie in the internal structure, not the external appearance. The latter has been much modified by adaptation. The former has been more or less preserved by heredity.
Comparative anatomy and ontogeny prove beyond question that man is a true vertebrate, and, therefore, man's special genealogical tree must be connected with that of the other Vertebrates, which spring from a common root with him. But we have also many important grounds in comparative anatomy and ontogeny for assuming a common origin for all the Vertebrates. If the general theory of evolution is correct, all the Vertebrates, including man, come from a single common ancestor, a long-extinct "Primitive Vertebrate." Hence the genealogical tree of the Vertebrates is at the same time that of the human race.
Our task, therefore, of constructing man's genealogy becomes the larger aim of discovering the genealogy of the entire vertebrate stem. As we now know from the comparative anatomy and ontogeny of the Amphioxus and the Ascidia, this is in turn connected with the genealogical tree of the Invertebrates (directly with that of the Vermalia), but has no direct connection with the independent stems of the Articulates, Molluscs, and Echinoderms. If we do thus follow our ancestral tree through various stages down to the lowest worms, we come inevitably to the Gastraea, that most instructive form that gives the clearest possible picture of an animal with two germinal layers. The Gastraea itself has originated from the simple multicellular vesicle, the Blastaea, and this in turn must have been evolved from the lowest circle of unicellular animals, to which we give the name of Protozoa. We have already considered the most important primitive type of these, the unicellular Amoeba, which is extremely instructive when compared with the human ovum. With this we reach the lowest of the solid data to which we are to apply our biogenetic law, and by which we may deduce the extinct ancestor from the embryonic form. The amoeboid nature of the young ovum and the unicellular condition in which (as stem-cell or cytula) every human being begins its existence justify us in affirming that the earliest ancestors of the human race were simple amoeboid coils.
But the further question now arises: "Whence came these first amoebae with which the history of life began at the commencement of the Laurentian epoch?" There is only one answer to this. The earliest unicellular organisms can only have been evolved from the simplest organisms we know, the Monera. These are the simplest living things that we can conceive. Their whole body is nothing but a particle of plasm, a granule of living albuminous matter, discharging of itself all the essential vital functions that form the material basis of life. Thus we come to the last, or, if you prefer, the first, question in connection with evolution—the question of the origin of the Monera. This is the real question of the origin of life, or of spontaneous generation.
We have neither space nor occasion to go further in this Chapter into the question of spontaneous generation. For this I must refer the reader to the fifteenth chapter of the History of Creation, and especially to the second book of the General Morphology, or to the essay on "The Monera and Spontaneous Generation" in my Studies of the Monera and other Protists.* (* The English reader will find a luminous and up-to-date chapter on the subject in Haeckel's recently written and translated Wonders of Life.—Translator.) I have given there fully my own view of this important question. The famous botanist Nageli afterwards (1884) developed the same ideas. I will only say a few words here about this obscure question of the origin of life, in so far as our main subject, organic evolution in general, is affected by it. Spontaneous generation, in the definite and restricted sense in which I maintain it, and claim that it is a necessary hypothesis in explaining the origin of life, refers solely to the evolution of the Monera from inorganic carbon-compounds. When living things made their first appearance on our planet, the very complex nitrogenous compound of carbon that we call plasson, which is the earliest material embodiment of vital action, must have been formed in a purely chemical way from inorganic carbon-compounds. The first Monera were formed in the sea by spontaneous generation, as crystals are formed in the mother-water. Our demand for a knowledge of causes compels us to assume this. If we believe that the whole inorganic history of the earth has proceeded on mechanical principles without any intervention of a Creator, and that the history of life also has been determined by the same mechanical laws; if we see that there is no need to admit creative action to explain the origin of the various groups of organisms; it is utterly irrational to assume such creative action in dealing with the first appearance of organic life on the earth.
This much-disputed question of "spontaneous generation" seems so obscure, because people have associated with the term a mass of very different, and often very absurd, ideas, and have attempted to solve the difficulty by the crudest experiments. The real doctrine of the spontaneous generation of life cannot possibly be refuted by experiments. Every experiment that has a negative result only proves that no organism has been formed out of inorganic matter in the conditions—highly artificial conditions—we have established. On the other hand, it would be exceedingly difficult to prove the theory by way of experiment; and even if Monera were still formed daily by spontaneous generation (which is quite possible), it would be very difficult, if not impossible, to find a solid proof of it. Those who will not admit the spontaneous generation of the first living things in our sense must have recourse to a supernatural miracle; and this is, as a matter of fact, the desperate resource to which our "exact" scientists are driven, to the complete abdication of reason.
A famous English physicist, Lord Kelvin (then Sir W. Thomson), attempted to dispense with the hypothesis of spontaneous generation by assuming that the organic inhabitants of the earth were developed from germs that came from the inhabitants of other planets, and that chanced to fall on our planet on fragments of their original home, or meteorites. This hypothesis found many supporters, among others the distinguished German physicist, Helmholtz. However, it was refuted in 1872 by the able physicist, Friedrich Zollner, of Leipzig, in his work, On the Nature of Comets. He showed clearly how unscientific this hypothesis is; firstly in point of logic, and secondly in point of scientific content. At the same time he pointed out that our hypothesis of spontaneous generation is "a necessary condition for understanding nature according to the law of causality."
I repeat that we must call in the aid of the hypothesis only as regards the Monera, the structureless "organisms without organs." Every complex organism must have been evolved from some lower organism. We must not assume the spontaneous generation of even the simplest cell, for this itself consists of at least two parts—the internal, firm nuclear substance, and the external, softer cellular substance or the protoplasm of the cell-body. These two parts must have been formed by differentiation from the indifferent plasson of a moneron, or a cytode. For this reason the natural history of the Monera is of great interest; here alone can we find the means to overcome the chief difficulties of the problem of spontaneous generation. The actual living Monera are specimens of such organless or structureless organisms, as they must have boon formed by spontaneous generation at the commencement of the history of life.
CHAPTER 2.19. OUR PROTIST ANCESTORS.
Under the guidance of the biogenetic law, and on the basis of the evidence we have obtained, we now turn to the interesting task of determining the series of man's animal ancestors. Phylogeny us a whole is an inductive science. From the totality of the biological processes in the life of plants, animals, and man we have gathered a confident inductive idea that the whole organic population of our planet has been moulded on a harmonious law of evolution. All the interesting phenomena that we meet in ontogeny and paleontology, comparative anatomy and dysteleology, the distribution and habits of organisms—all the important general laws that we abstract from the phenomena of these sciences, and combine in harmonious unity—are the broad bases of our great biological induction.
But when we come to the application of this law, and seek to determine with its aid the origin of the various species of organisms, we are compelled to frame hypotheses that have essentially a DEDUCTIVE character, and are inferences from the general law to particular cases. But these special deductions are just as much justified and necessitated by the rigorous laws of logic as the inductive conclusions on which the whole theory of evolution is built. The doctrine of the animal ancestry of the human race is a special deduction of this kind, and follows with logical necessity from the general inductive law of evolution.
I must point out at once, however, that the certainty of these evolutionary hypotheses, which rest on clear special deductions, is not always equally strong. Some of these inferences are now beyond question; in the case of others it depends on the knowledge and the competence of the inquirer what degree of certainty he attributes to them. In any case, we must distinguish between the ABSOLUTE certainty of the general (inductive) theory of descent and the RELATIVE certainty of special (deductive) evolutionary hypotheses. We can never determine the whole ancestral series of an organism with the same confidence with which we hold the general theory of evolution as the sole scientific explanation of organic modifications. The special indication of stem-forms in detail will always be more or less incomplete and hypothetical. This is quite natural. The evidence on which we build is imperfect, and always will be imperfect; just as in comparative philology.
The first of our documents, paleontology, is exceedingly incomplete. We know that all the fossils yet discovered are only an insignificant fraction of the plants and animals that have lived on our planet. For every single species that has been preserved for us in the rocks there are probably hundreds, perhaps thousands, of extinct species that have left no trace behind them. This extreme and very unfortunate incompleteness of the paleontological evidence, which cannot be pointed out too often, is easily explained. It is absolutely inevitable in the circumstances of the fossilisation of organisms. It is also due in part to the incompleteness of our knowledge in this branch. It must be borne in mind that the great majority of the stratified rocks that compose the crust of the earth have not yet been opened. We have only a few specimens of the innumerable fossils that are buried in the vast mountain ranges of Asia and Africa. Only a part of Europe and North America has been investigated carefully. The whole of the fossils known to us certainly do not amount to a hundredth part of the remains that are really buried in the crust of the earth. We may, therefore, look forward to a rich harvest in the future as regards this science. However, our paleontological evidence will (for reasons that I have fully explained in the sixteenth chapter of the History of Creation) always be defective.
The second chief source of evidence, ontogeny, is not less incomplete. It is the most important source of all for special phylogeny; but it has great defects, and often fails us. We must, above all, clearly distinguish between palingenetic and cenogenetic phenomena. We must never forget that the laws of curtailed and disturbed heredity often make the original course of development almost unrecognisable. The recapitulation of phylogeny by ontogeny is only fairly complete in a few cases, and is never wholly complete. As a rule, it is precisely the earliest and most important embryonic stages that suffer most from alteration and condensation. The earlier embryonic forms have had to adapt themselves to new circumstances, and so have been modified. The struggle for existence has had just as profound an influence on the freely moving and still immature young forms as on the adult forms. Hence in the embryology of the higher animals, especially, palingenesis is much restricted by cenogenesis; it is to-day, as a rule, only a faded and much altered picture of the original evolution of the animal's ancestors. We can only draw conclusions from the embryonic forms to the stem-history with the greatest caution and discrimination. Moreover, the embryonic development itself has only been fully studied in a few species.
Finally, the third and most valuable source of evidence, comparative anatomy, is also, unfortunately, very imperfect; for the simple reason that the whole of the living species of animals are a mere fraction of the vast population that has dwelt on our planet since the beginning of life. We may confidently put the total number of these at more than a million species. The number of animals whose organisation has been studied up to the present in comparative anatomy is proportionately very small. Here, again, future research will yield incalculable treasures. But, for the present, in view of this patent incompleteness of our chief sources of evidence, we must naturally be careful not to lay too much stress in human phylogeny on the particular animals we have studied, or regard all the various stages of development with equal confidence as stem-forms.
In my first efforts to construct the series of man's ancestors I drew up a list of, at first ten, afterwards twenty to thirty, forms that may be regarded more or less certainly as animal ancestors of the human race, or as stages that in a sense mark off the chief sections in the long story of evolution from the unicellular organism to man. Of these twenty to thirty stages, ten to twelve belong to the older group of the Invertebrates and eighteen to twenty to the younger division of the Vertebrates.
In approaching, now, the difficult task of establishing the evolutionary succession of these thirty ancestors of humanity since the beginning of life, and in venturing to lift the veil that covers the earliest secrets of the earth's history, we must undoubtedly look for the first living things among the wonderful organisms that we call the Monera; they are the simplest organisms known to us—in fact, the simplest we can conceive. Their whole body consists merely of a simple particle or globule of structureless plasm or plasson. The discoveries of the last four decades have led us to believe with increasing certainty that wherever a natural body exhibits the vital processes of nutrition, reproduction, voluntary movement, and sensation, we have the action of a nitrogenous carbon-compound of the chemical group of the albuminoids; this plasm (or protoplasm) is the material basis of all vital functions. Whether we regarded the function, in the monistic sense, as the direct action of the material substratum, or whether we take matter and force to be distinct things in the dualistic sense, it is certain that we have not as yet found any living organism in which the exercise of the vital functions is not inseparably bound up with plasm.
The soft slimy plasson of the body of the moneron is generally called "protoplasm," and identified with the cellular matter of the ordinary plant and animal cells. But we must, to be accurate, distinguish between the plasson of the cytodes and the protoplasm of the cells. This distinction is of the utmost importance for the purposes of evolution. As I have often said, we must recognise two different stages of development in these "elementary organisms," or plastids ("builders"), that represent the ultimate units of organic individuality. The earlier and lower stage are the unnucleated cytodes, the body of which consists of only one kind of albuminous matter—the homogeneous plasson or "formative matter." The later and higher stage are the nucleated cells, in which we find a differentiation of the original plasson into two different formative substances—the caryoplasm of the nucleus and the cytoplasm of the body of the cell (cf. Chapter 1.6.)
(FIGURE 2.226. Chroococcus minor (Nageli), magnified 1500 times. A phytomoneron, the globular plastids of which secrete a gelatinous structureless membrane. The unnucleated globule of plasm (bluish-green in colour) increases by simple cleavage (a to d).
The Monera are permanent cytodes. Their whole body consists of soft, structureless plasson. However carefully we examine it with our finest chemical reagents and most powerful microscopes, we can find no definite parts or no anatomic structure in it. Hence, the Monera are literally organisms without organs; in fact, from the philosophic point of view they are not organisms at all, since they have no organs. They can only be called organisms in the sense that they are capable of the vital functions of nutrition, reproduction, sensation, and movement. If we were to try to imagine the simplest possible organism, we should frame something like the moneron.
The Monera that we find to-day in various forms fall into two groups according to the nature of their nutrition—the Phytomonera and the Zoomonera; from the physiological point of view, the former are the simplest specimens of the plant (phyton) kingdom, and the latter of the animal (zoon) world. The Phytomonera, especially in their simplest form, the Chromacea (Phycochromacea or Cyanophycea), are the most primitive and the oldest of living organisms. The typical genus Chroococcus (Figure 2.226) is represented by several fresh-water species, and often forms a very delicate bluish-green deposit on stones and wood in ponds and ditches. It consists of round, light green particles, from 1/7000 to 1/2500 of an inch in diameter.
(FIGURE 2.227. Aphanocapsa primordialis (Nageli), magnified 1000 times. A phytomoneron, the round plastids of which (bluish-green in colour) secrete a shapeless gelatinous mass; in this the unnucleated cytodes increase continually by simple cleavage.)
The whole life of these homogeneous globules of plasm consists of simple growth and reproduction by cleavage. When the tiny particle has reached a certain size by the continuous assimilation of inorganic matter, it divides into two equal halves, by a constriction in the middle. The two daughter-monera that are thus formed immediately begin a similar vital process. It is the same with the brown Procytella primordialis (formerly called the Protococcus marinus); it forms large masses of floating matter in the arctic seas. The tiny plasma-globules of this species are of a greenish-brown colour, and have a diameter of 1/10,000 to 1/5000 of an inch. There is no membrane discoverable in the simplest Chroococcacea, but we find one in other members of the same family; in Aphanocapsa (Figure 2.227) the enveloping membranes of the social plastids combine; in Gloecapsa they are retained through several generations, so that the little plasma-globules are enfolded in many layers of membrane.
Next to the Chromacea come the Bacteria, which have been evolved from them by the remarkable change in nutrition which gives us the simple explanation of the differentiation of plant and animal in the protist kingdom. The Chromacea build up their plasm directly from inorganic matter; the Bacteria feed on organic matter. Hence, if we logically divide the protist kingdom into plasma-forming Protophyta and plasma-consuming Protozoa, we must class the Bacteria with the latter; it is quite illogical to describe them—as is still often done—as Schizomycetes, and class them with the true fungi. The Bacteria, like the Chromacea, have no nucleus. As is well-known, they play an important part in modern biology as the causes of fermentation and putrefaction, and of tuberculosis, typhus, cholera, and other infectious diseases, and as parasites, etc. But we cannot linger now to deal with these very interesting features; the Bacteria have no relation to man's genealogical tree.
We may now turn to consider the remarkable Protamoeba, or unnucleated Amoeba. I have, in the first volume, pointed out the great importance of the ordinary Amoeba in connection with several weighty questions of general biology. The tiny Protamoebae, which are found both in fresh and salt water, have the same unshapely form and irregular movements of their simple naked body as the real Amoebae; but they differ from them very materially in having no nucleus in their cell-body. The short, blunt, finger-like processes that are thrust out at the surface of the creeping Protamoeba serve for getting food as well as for locomotion. They multiply by simple cleavage (Figure 2.228).
(FIGURE 2.228. A moneron (Protamoeba) in the act of reproduction. A The whole moneron, moving like an ordinary amoeba by thrusting out changeable processes. B It divides into two halves by a constriction in the middle. C The two halves separate, and each becomes an independent individual. (Highly magnified.))
The next stage to the simple cytode-forms of the Monera in the genealogy of mankind (and all other animals) is the simple cell, or the most rudimentary form of the cell which we find living independently to-day as the Amoeba. The earliest process of inorganic differentiation in the structureless body of the Monera led to its division into two different substances—the caryoplasm and the cytoplasm. The caryoplasm is the inner and firmer part of the cell, the substance of the nucleus. The cytoplasm is the outer and softer part, the substance of the body of the cell. By this important differentiation of the plasson into nucleus and cell-body, the organised cell was evolved from the structureless cytode, the nucleated from the unnucleated plastid. That the first cells to appear on the earth were formed from the Monera by such a differentiation seems to us the only possible view in the present condition of science. We have a direct instance of this earliest process of differentiation to-day in the ontogeny of many of the lower Protists (such as the Gregarinae).
The unicellular form that we have in the ovum has already been described as the reproduction of a corresponding unicellular stem-form, and to this we have ascribed the organisation of an Amoeba (cf. Chapter 1.6). The irregular-shaped Amoeba, which we find living independently to-day in our fresh and salt water, is the least definite and the most primitive of all the unicellular Protozoa (Figure 1.16). As the unripe ova (the protova that we find in the ovaries of animals) cannot be distinguished from the common Amoebae, we must regard the Amoeba as the primitive form that is reproduced in the embryonic stage of the amoeboid ovum to-day, in accordance with the biogenetic law. I have already pointed out, in proof of the striking resemblance of the two cells, that the ova of many of the sponges were formerly regarded as parasitic Amoebae (Figure 1.18). Large unicellular organisms like the Amoebae were found creeping about inside the body of the sponge, and were thought to be parasites. It was afterwards discovered that they were really the ova of the sponge from which the embryos were developed. As a matter of fact, these sponge-ova are so much like many of the Amoebae in size, shape, the character of their nucleus, and movement of the pseudopodia, that it is impossible to distinguish them without knowing their subsequent development.
Our phylogenetic interpretation of the ovum, and the reduction of it to some ancient amoeboid ancestral form, supply the answer to the old problem: "Which was first, the egg or the chick?" We can now give a very plain answer to this riddle, with which our opponents have often tried to drive us into a corner. The egg came a long time before the chick. We do not mean, of course, that the egg existed from the first as a bird's egg, but as an indifferent amoeboid cell of the simplest character. The egg lived for thousands of years as an independent unicellular organism, the Amoeba. The egg, in the modern physiological sense of the word, did not make its appearance until the descendants of the unicellular Protozoon had developed into multicellular animals, and these had undergone sexual differentiation. Even then the egg was first a gastraea-egg, then a platode-egg, then a vermalia-egg, and chordonia-egg; later still acrania-egg, then fish-egg, amphibia-egg, reptile-egg, and finally bird's egg. The bird's egg we have experience of daily is a highly complicated historical product, the result of countless hereditary processes that have taken place in the course of millions of years.
The earliest ancestors of our race were simple Protophyta, and from these our protozoic ancestors were developed afterwards. From the morphological point of view both the vegetal and the animal Protists were simple organisms, individualities of the first order, or plastids. All our later ancestors are complex organisms, or individualities of a higher order—social aggregations of a plurality of cells. The earliest of these, the Moraeada, which represent the third stage in our genealogy, are very simple associations of homogeneous, indifferent cells—undifferentiated colonies of social Amoebae or Infusoria. To understand the nature and origin of these protozoa-colonies we need only follow step by step the first embryonic products of the stem-cell. In all the Metazoa the first embryonic process is the repeated cleavage of the stem-cell, or first segmentation-cell (Figure 2.229). We have already fully considered this process, and found that all the different forms of it may be reduced to one type, the original equal or primordial segmentation (cf. 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.