This method of looking at the problem seems to me to be more in consonance with the facts than the reverse; for, as pointed out by Jaekel, the fishes with large plates are the oldest, and in Cyathaspis, the very oldest of all, the size of the plates is most conspicuous; he considers, therefore, this preconceived view that large plates are formed by the fusion of small ones must give way to the opposite belief.
Fig. 139.—Drepanaspis. Ventral and Dorsal Aspects.(AfterLankester.)A., anus;E., lateral eyes.
Fig. 139.—Drepanaspis. Ventral and Dorsal Aspects.(AfterLankester.)A., anus;E., lateral eyes.
Fig. 139.—Drepanaspis. Ventral and Dorsal Aspects.(AfterLankester.)
A., anus;E., lateral eyes.
So also Rohon, as quoted by Traquair, who, in his first paper accepted Lankester's view that the ridges of the pteraspidian shield were formed by the fusion of a linear arrangement of numbers of placoid scales, suggests in his second paper that these ridges may have been the most primitive condition of the dermal skeleton of the vertebrate, out of which, by differentiation, the dermal denticles (placoid scales) of the selachian, as well as their modifications in the ganoids, teleosteans, and amphibians, have arisen.
One thing is agreed upon on all sides; no sign of bone-corpuscles is to be found in this dermal covering of Pteraspis. In the deeper layers are large spaces, the so-called pulp-cavities leading into narrow canaliculi, the so-called dentine canals. The structure islooked upon as similar to that of the pulp and dentine canals of many fish-scales.
On the other hand, this dermal covering of Pteraspis has been compared by Patten with the arrangement of the chitinous structure of certain parts of the external covering of Limulus, a comparison which to my mind presents a great difficulty. The chitin-layers in Limulus areexternalto the epidermal cells, being formed by them; the layers in Pteraspis which look like chitin must have beeninternalto the epidermal layer; for each vascular canal which passes from a pulp-cavity on its way to be distributed into the dentine canals of the ridge gives off short side branches, which open directly into the groove between the ridges. If these canals were filled with blood they could not possibly open directly into the open grooves between the ridges; these openings must, therefore, have been covered over with an epithelial layer which covered over the surface of the animal, and consequently the chitin-like structure must have been internal to the epidermis, and not external, as on Patten's view. The comparison of this structure with the dentine of fish-scales signifies the same thing, for in the latter the epidermis is external to the dentine-plates, the hard skeletal structure is in the position of the cutis, not of the cuticle.
The position appears to me to be this: the dermal cranial skeleton of vertebrates, whether it takes the form of a bony skull or of the dorsal plates of a cephalaspid or a pteraspid is, in all cases, not cuticular,i.e.is not an external formation of the epidermal cells, but is formed in tissue of the nature of connective tissue underlying the epidermis. On the contrary, the hard part of the head-carapace of the palæostracan is an external formation of the epidermal cells.
If, then, this tissue of Pteraspis is not to be looked upon as chitin, how can we imagine its formation? It is certainly not bone, for there are no bone-corpuscles; it is a very regular laminated structure resembling in appearance chitin rather than anything else.
As in all cases of difficulty, turn to Ammocœtes and let us see what clue there is to be found there. The skin of Ammocœtes is peculiar among vertebrates in many respects. It consists of a number of epidermal cells, as in Fig.140, the varying function of which need not be considered here, covered over with a cuticular layer which is extraordinarily thick for the cuticle of a vertebrate skin; this cuticular layer is perforated with fine canaliculi, through which thesecretion of the underlying cells passes, as is seen in Fig.140, A and B. This cuticle corresponds to the chitinous covering of the arthropod, and like it is perforated with canaliculi, and, according to Lwoff, possibly contains chitin. The epidermal cells rest on a thick layer of most striking appearance (Fig.141), for it resembles, in an extraordinary degree, when examined superficially, a layer of chitin; it is called the laminated layer, and is characterized by the extreme regularity of the laminæ. This appearance is due, as the observations of Miss Alcock show, to alternate layers of connective tissue fibres arranged at right angles to each other, each fibre running a straight course and possessing its own nucleus. Although the fibres in each layer are packed close together, they are sufficiently apart to form with the fibres of the alternate layers a meshwork rather than a homogeneous structure, and thus the surface view of this layer shows a regular network of very fine spaces through which nerve-fibres and fluid pass. This layer is easily dissolved in a solution of hypochlorite of soda, a fluid which dissolves chitin. Any one looking at Ammocœtes would say that the only part of its skin which resembles chitin is this laminated layer, and therefore the only part of its skin which would afford an indication of the nature of the skeleton of Pteraspis is this laminated layer, which belongs to the cutis, and not to the cuticle. Yet another significant peculiarity of this layer is its entire disappearance at transformation. Miss Alcock, in a research not yet published, has shown that this layer is completely broken up and absorbed at transformation; the cutis of Petromyzon is formed entirely anew, and no longer presents any regular laminated character, but resembles rather the sub-epidermal connective tissue layer of the skin of higher vertebrates. This laminated layer, then, just like the muco-cartilage, shows, by its complete disappearance at transformation, its ancestral character.
Fig. 140.—Epithelial Cells of Ammocœtes to show the Canaliculi in the Thick Cuticle (B). A, Transverse Section through the Cuticle.
Fig. 140.—Epithelial Cells of Ammocœtes to show the Canaliculi in the Thick Cuticle (B). A, Transverse Section through the Cuticle.
Fig. 140.—Epithelial Cells of Ammocœtes to show the Canaliculi in the Thick Cuticle (B). A, Transverse Section through the Cuticle.
Very suggestive is the arrangement of the different skeletaltissues in the head-region of Ammocœtes. Fig.141represents a section through the head near the pineal eye. Most internally isa, a section of the membranous cranium, then comesb, the muco-cartilaginous skeleton, thenc, the laminated layer, and finallyd, the external cuticle. If in Ammocœtes we possess an epitome of the history of the vertebrate, how would these layers be represented in the past ages, supposing they could be fossilized?
Fig. 141.—Section of Skin and Underlying Tissues in the Head-Region of Ammocœtes.a, cranial wall;b, muco-cartilage;c, laminated layer;d, external cuticular layer.
Fig. 141.—Section of Skin and Underlying Tissues in the Head-Region of Ammocœtes.a, cranial wall;b, muco-cartilage;c, laminated layer;d, external cuticular layer.
Fig. 141.—Section of Skin and Underlying Tissues in the Head-Region of Ammocœtes.
a, cranial wall;b, muco-cartilage;c, laminated layer;d, external cuticular layer.
The most internal layera, by the formation of cartilage and then bone, represents the great mass of vertebrate fossils; the next layerb, by a process of calcification, as previously argued, represents the head-shield of the Osteostracan fishes; while the cuticular layerd, no longer thin, is the remnant of the Palæostracan head-carapace. Between these two layers,bandd, lies the laminated layerc. Intermediate to the Palæostracan and the Osteostracan comes the Heterostracan, with its peculiar head-shield—a head-shield whose origin is more easily conceivable as arising from something of the nature of the laminated layer than from any other structure represented in Ammocœtes.
My present suggestion, then, is this: the transition from the skeletal covering of the Palæostracan to that of the highest vertebrates was brought about by the calcification of successive layers from without inwards, all of which still remain in Ammocœtes and show how the external chitinous covering of the arthropod was gradually replaced by the deep-lying internal bony cranium of the higher vertebrates.
In Ammocœtes the layer which represents the covering of thePalæostracan has already almost disappeared. At transformation the layers representing the stage arrived at by the Heterostracan and the Osteostracan disappear; but the stage representing the higher vertebrates, far from disappearing, by the formation of cartilage reaches a higher stage and prepares the way for the ultimate stage of all—the formation of the bony cranium.
So much for the evidence as to the nature of the structure of the head-shield of the Pteraspidæ.
It suggests that these fishes were covered anteriorly with armoured plates derived from the cutis layer of the skin, a layer which was specially thickened and very vascular, apparently, to enable respiration to be very largely, if not entirely, effected by the surface of the body. It is difficult to understand how the sea-scorpions breathed, and it is easy to see how the formation of ventral and dorsal plates enclosing the mesosomatic appendages may at the outset have hindered the action of the branchiæ. The respiratory chamber, according to my view, had at first the double function of respiration and digestion. A new digestive apparatus was the pressing need at the time; it would, therefore, be of distinct advantage to remove, as much as possible, the burden of respiration from this incipient alimentary canal.
What can be said as to the shape of these ancient forms of fishes? Certain parts of them are absolutely known, other parts are guesswork. They are known to have possessed a dorsal shield, a ventral shield formerly looked upon as belonging to a separate species, called Scaphaspis, and a spine attached to the dorsal shield. The rest of their configuration, as given in Smith Woodward's restoration (Fig.142) is guesswork; the fish-like body with its scales, the heterocercal tail, is based on the most insufficient evidence of something of the nature of scales having being found near the head-plates.
The dorsal shield is characterized by a pair of lateral eyes situated on the edge of the shield, not as in Cephalaspis near the middle line. In the middle line, where the rostrum meets the large dorsal plate, median eyes were situated. But the slightest sign of any median single nasal opening, such as is so characteristic of the head-shield of the Osteostraci and of Ammocœtes has never been discovered. The olfactory organ must have been situated on the ventral side as in the larval stage of Ammocœtes, or in the Palæostraca. Many of these head-shields are remarkably well preserved,and it is difficult to believe that an olfactory opening would not be seen if any such had existed, as it does in Thyestes.
Fig. 142.—Restoration of Pteraspis.(AfterSmith Woodward.)
Fig. 142.—Restoration of Pteraspis.(AfterSmith Woodward.)
Fig. 142.—Restoration of Pteraspis.(AfterSmith Woodward.)
The difficulty of interpreting these types is the difficulty of understanding their method of locomotion; that is largely the reason why the spine has been placed as if projecting from the back, and a fish-like body with a heterocercal tail-fin added. If, on the contrary, the spine is a terminal tail-spine, then, as far as the fossilized remains indicate, the animal consisted of a dorsal shield, a ventral shield, and a tail-spine, to which must be added two apparently lateral pieces and a few scales. If the animal did not possess a flexible body with a tail-fin, but terminated in a rigid spike after the fashion of a Limulus-like animal, then it must have moved by means ofappendages. At present we have not sufficient evidence to decide this question.
That the animal crawled about in the mud by means of free appendages is by no means an impossible view, seeing how difficult it is to find the remains of appendages in the fossils of this far-back time, even when we are sure that they existed. Thus, for many generations, the appendages of trilobites, which occur in such countless numbers, and in such great variety of form, were absolutely unknown, until at last, in consequence of a fortunate infiltration by pyrites, they were found by Beecher preserved down to the minutest detail. Even to this day no trace of appendages has been found in such forms as Hemiaspis, Bunodes, Belinurus, Prestwichia.
The whole question of the evidence of any prosomatic appendages in these ancient fishes is one of very great interest, and of late years has been investigated by Patten. It has long been known that forms such as Pterichthys and Bothriolepis possessed two large, jointed locomotor appendages, and Patten has lately obtained better specimens of Bothriolepis than have ever been found before, which show not only the general configuration of the fish, but also the presence of mandibles or gnathites in the mouth-region resembling those of an arthropod. These mandibles had been seen before (Smith Woodward), but Patten's specimens are more perfect than any previously described, and cause him to conclude that these ancient fish were of the nature of arthropods rather than of vertebrates.
Patten has also been able to obtain some excellent specimens of the under surface of the head of Tremataspis, which, as evident in Fig.143, show the presence of a series of holes, ranging on each side from the mouth-opening, in a semicircular fashion towards the middle line. He considers that these openings indicate the attachments of appendages, in opposition to other observers, such as Jaekel, who look upon them as gill-slits. To my mind, they are not in the right position for gill-slits; they are certainly in a prosomatic rather than in a mesosomatic position, and I should not be at all surprised if further research justified Patten's position. So convinced is he of the presence of appendages in all these old forms, that he considers them to be arthropods rather than vertebrates, although, at the same time, he looks upon them as indicating the origin of vertebrates from arthropods. Here, perhaps, it is advisable to say a few words on Patten's attitude towards this question.
Two years after I had put forward my theory of the derivation of vertebrates from arthropods, Patten published, in theQuarterly Journal of Microscopical Science, simultaneously with my paper in that journal, a paper entitled "The Origin of Vertebrates from Arachnids." In this paper he made no reference to my former publications, but he made it clear that there was an absolutely fundamental difference between our treatment of the problem; for he took the old view that of necessity there must be a reversal of surfaces in order that the internal organs should be in the same relative positions in the vertebrate and in the invertebrate. He simply, therefore, substituted Arachnid for Annelid in the old theory. Because of this necessity for the reversal of surfaces he discarded the terms dorsal and ventral as indicative of the surfaces of an animal, and substituted hæmal and neural, thereby hopelessly confusing the issue and making it often very difficult to understand his meaning.
Fig. 143.—Under-Surface of Head-Region in Tremataspis.(AfterPatten.)
Fig. 143.—Under-Surface of Head-Region in Tremataspis.(AfterPatten.)
Fig. 143.—Under-Surface of Head-Region in Tremataspis.(AfterPatten.)
He still holds to his original opinion, and I am still waiting to find out when the reversal of surfaces took place, for his investigations lead him, as must naturally be the case, to compare the dorsal (or, as he would call it, the hæmal) surface of Bothriolepis, of the Cephalaspidæ, and of the Pteraspidæ with the dorsal surface of the Palæostraca.
All these ancient fishes are, according to him, still in the arthropod stage, have not yet turned over, though in a peculiarly unscientific manner he argues elaborately that they must have swum on their back rather than on their front, and so indicated the coming reversal. Because they were arthropods they cannot have had afrontal nose-organ; therefore, Patten looks upon the nose and the two lateral eyes of the Osteostraci as a complex median eye, regardless of the fact that the median eyes already existed.
Every atom of evidence Patten has brought forward, every new fact he has discovered, confirms my position and makes his still more hopelessly confused. Keep the animal the right side uppermost, and the evidence of the rocks confirms the transition from the Palæostracan to the Cyclostome; reverse the surfaces, and the attempt to derive the vertebrate from the palæostracan becomes so confused and hopelessly muddled as to throw discredit on any theory of the origin of vertebrates from arthropods. For my own part, I fully expect that appendages will be found not only in the Cephalaspidæ but also in the Pteraspidæ, and I hope Patten will continue his researches with increasing success. I feel sure, however, his task will be much simplified if he abandons his present position and views the question from my standpoint.
Summary.
The shifting of the nasal tube from a ventral to a dorsal position, as seen in Ammocœtes, is, perhaps, the most important of all clues in connection with the comparison of Ammocœtes to the Palæostracan on the one hand, and to the Cephalaspid on the other; for, whereas the exact counterpart of the opening of such a tube is always found on the dorsal head-shield in all members of the latter group, nothing of the kind is ever found on the dorsal carapace of the former group.The reason for this difference is made immediately evident in the development of Ammocœtes itself, for the olfactory tube originates as a ventral tube—the tube of the hypophysis—in exactly the same position as the olfactory tube of the Palæostracan, and later on in its development takes up a dorsal position.In fact, Ammocœtes in its development indicates how the Palæostracan head-shield became transformed into that of the Cephalaspid.In another most important character Ammocœtes indicates its relationship to the Cephalaspidæ, for it possesses an external skeleton or head-shield composed of muco-cartilage, which is the exact counterpart of the so-called bony head-shield of the latter group; and still more strikingly the structure of the cephalaspidian head-shield is remarkably like that of muco-cartilage. In the one case, by the deposition of calcium salts, a hard external skeleton, capable of being preserved as a fossil, has been formed; in the other, by the absence of the calcium salts, a soft chondro-mucoid matrix, in which the characteristic cells and fibrils are embedded, distinguishes the tissue.The recognition that the head-shields of these most primitive fishes were not composed of bone, but of muco-cartilage, the precursor of both cartilage and bone, immediately clears up in the most satisfactory manner the wholequestion of their derivation from elasmobranch fishes; for the main argument in favour of the latter derivation is the exceedingly strong one that bone succeeds cartilage—notvice versâ—therefore, these forms, since their head-shield is bony, must have arisen from some other fishes with a cartilaginous skeleton, most probably of an elasmobranch nature. Seeing, however, that the structure of their shields resembles muco-cartilage much more closely than bone, and that Ammocœtes forms a head-shield of muco-cartilage closely resembling theirs, there is no longer any necessity to derive the jawless fishes from the gnathostomatous; but, on the contrary, we may look with certainty upon the Agnatha as the most primitive group from which the others have been derived.The history of the rocks shows that the group of fishes, Pteraspis and Cyathaspis, are older than the Cephalaspidæ—come, therefore, phylogenetically between the Palæostraca and the latter group. In this group the head-shields are of a very different character, without any sign of any structure comparable with that of bone, and although they possessed both lateral and median eyes, there is never in any case any trace of a dorsal nasal orifice. Their olfactory passage, like that of the Palæostraca, must have been ventral.The remarkable comparison which exists between the head-shields of Ammocœtes and Cephalaspis, enables us to locate the position of the brain and cranium of the latter with considerable accuracy, and so to compare the segmental markings found in many of these fossils with the corresponding markings, found either in fossil Palæostraca or on the head-carapaces of living scorpions and spiders, such as Phrynus and Mygale. In all cases the cranial region was covered with a median plate, often especially hard, which corresponded to the glabellum of the trilobite; the growth of the cranium can be traced from its beginnings as the upturned lateral flanges of the plastron to the membranous cranium of Ammocœtes.From such a comparison it follows that the segments, found in the antero-lateral region of the head-shield, were not segments of the cranium, but of parts beyond the region of the cranium, and from their position must have been segments supplied by the trigeminal nerve, and not by the vagus group; segments, therefore, which did not indicate gills and gill-slits, but muscles, innervated by the trigeminal nerve; muscles which, as indicated by the corresponding markings on the carapace of Phrynus, Mygale, etc., were the tergo-coxal muscles of the prosomatic appendages.The discovery of the nature of these appendages in the Pteraspidæ and Cephalaspidæ, as well as in the Asterolepidæ (Pterichthys and Bothriolepis), is a problem of the future, though in the latter, not only have the well-known oar-like appendages been long since discovered, but Patten has recently found specimens of Bothriolepis which throw light on the anterior masticating gnathite-like appendages which these ancient forms possessed.
The shifting of the nasal tube from a ventral to a dorsal position, as seen in Ammocœtes, is, perhaps, the most important of all clues in connection with the comparison of Ammocœtes to the Palæostracan on the one hand, and to the Cephalaspid on the other; for, whereas the exact counterpart of the opening of such a tube is always found on the dorsal head-shield in all members of the latter group, nothing of the kind is ever found on the dorsal carapace of the former group.
The reason for this difference is made immediately evident in the development of Ammocœtes itself, for the olfactory tube originates as a ventral tube—the tube of the hypophysis—in exactly the same position as the olfactory tube of the Palæostracan, and later on in its development takes up a dorsal position.
In fact, Ammocœtes in its development indicates how the Palæostracan head-shield became transformed into that of the Cephalaspid.
In another most important character Ammocœtes indicates its relationship to the Cephalaspidæ, for it possesses an external skeleton or head-shield composed of muco-cartilage, which is the exact counterpart of the so-called bony head-shield of the latter group; and still more strikingly the structure of the cephalaspidian head-shield is remarkably like that of muco-cartilage. In the one case, by the deposition of calcium salts, a hard external skeleton, capable of being preserved as a fossil, has been formed; in the other, by the absence of the calcium salts, a soft chondro-mucoid matrix, in which the characteristic cells and fibrils are embedded, distinguishes the tissue.
The recognition that the head-shields of these most primitive fishes were not composed of bone, but of muco-cartilage, the precursor of both cartilage and bone, immediately clears up in the most satisfactory manner the wholequestion of their derivation from elasmobranch fishes; for the main argument in favour of the latter derivation is the exceedingly strong one that bone succeeds cartilage—notvice versâ—therefore, these forms, since their head-shield is bony, must have arisen from some other fishes with a cartilaginous skeleton, most probably of an elasmobranch nature. Seeing, however, that the structure of their shields resembles muco-cartilage much more closely than bone, and that Ammocœtes forms a head-shield of muco-cartilage closely resembling theirs, there is no longer any necessity to derive the jawless fishes from the gnathostomatous; but, on the contrary, we may look with certainty upon the Agnatha as the most primitive group from which the others have been derived.
The history of the rocks shows that the group of fishes, Pteraspis and Cyathaspis, are older than the Cephalaspidæ—come, therefore, phylogenetically between the Palæostraca and the latter group. In this group the head-shields are of a very different character, without any sign of any structure comparable with that of bone, and although they possessed both lateral and median eyes, there is never in any case any trace of a dorsal nasal orifice. Their olfactory passage, like that of the Palæostraca, must have been ventral.
The remarkable comparison which exists between the head-shields of Ammocœtes and Cephalaspis, enables us to locate the position of the brain and cranium of the latter with considerable accuracy, and so to compare the segmental markings found in many of these fossils with the corresponding markings, found either in fossil Palæostraca or on the head-carapaces of living scorpions and spiders, such as Phrynus and Mygale. In all cases the cranial region was covered with a median plate, often especially hard, which corresponded to the glabellum of the trilobite; the growth of the cranium can be traced from its beginnings as the upturned lateral flanges of the plastron to the membranous cranium of Ammocœtes.
From such a comparison it follows that the segments, found in the antero-lateral region of the head-shield, were not segments of the cranium, but of parts beyond the region of the cranium, and from their position must have been segments supplied by the trigeminal nerve, and not by the vagus group; segments, therefore, which did not indicate gills and gill-slits, but muscles, innervated by the trigeminal nerve; muscles which, as indicated by the corresponding markings on the carapace of Phrynus, Mygale, etc., were the tergo-coxal muscles of the prosomatic appendages.
The discovery of the nature of these appendages in the Pteraspidæ and Cephalaspidæ, as well as in the Asterolepidæ (Pterichthys and Bothriolepis), is a problem of the future, though in the latter, not only have the well-known oar-like appendages been long since discovered, but Patten has recently found specimens of Bothriolepis which throw light on the anterior masticating gnathite-like appendages which these ancient forms possessed.
CHAPTER XI
THE EVIDENCE OF THE AUDITORY APPARATUS AND THE ORGANS OF THE LATERAL LINE
Lateral line organs.—Function of this group of organs.—Poriferous sense-organs on the appendages in Limulus.—Branchial sense-organs.—Prosomatic sense organs.—Flabellum.—Its structure and position.—Sense-organs of mandibles.—Auditory organs of insects and arachnids.—Poriferous chordotonal organs.—Balancers of Diptera.—Resemblance to organs of flabellum.—Racquet-organs of Galeodes.—Pectens of scorpions.—Large size of nerve to all these special sense-organs.—Origin of parachordals and auditory capsule.—Reason why VIIth nerve passes in and out of capsule.—Evidence of Ammocœtes.—Intrusion of glandular mass round brain into auditory capsule.—Intrusion of generative and hepatic mass round brain into base of flabellum.—Summary.
Lateral line organs.—Function of this group of organs.—Poriferous sense-organs on the appendages in Limulus.—Branchial sense-organs.—Prosomatic sense organs.—Flabellum.—Its structure and position.—Sense-organs of mandibles.—Auditory organs of insects and arachnids.—Poriferous chordotonal organs.—Balancers of Diptera.—Resemblance to organs of flabellum.—Racquet-organs of Galeodes.—Pectens of scorpions.—Large size of nerve to all these special sense-organs.—Origin of parachordals and auditory capsule.—Reason why VIIth nerve passes in and out of capsule.—Evidence of Ammocœtes.—Intrusion of glandular mass round brain into auditory capsule.—Intrusion of generative and hepatic mass round brain into base of flabellum.—Summary.
When speaking of the tripartite arrangement of the cranial nerves, an arrangement which gave the clue to the meaning of the cranial segments, I spoke of the trigeminal as supplying the sensory nerves to the skin in the head-region, and I compared this dorsal system of afferent nerves to the system of epimeral nerves in Limulus which supply the prosomatic and mesosomatic carapaces of Limulus with sensory fibres. I compared the ventral system of eye-muscle nerves with the system of nerves supplying the segmental dorso-ventral somatic muscles of the prosomatic region, and I compared the lateral system of mixed nerves with the nerves supplying the prosomatic and mesosomatic appendages of Limulus. I compared, also, the optic nerves and the olfactory nerves with the corresponding nerves in the same invertebrate group. My readers will see at once that one well-marked group of nerves—the auditory and lateral line system—has been entirely omitted up to the present, it has not even been mentioned in the scheme of the cranial segments; I have purposely reserved its consideration until now, because the organs these nerves supply, though situated in the skin, are of such a special characteras to form a category by themselves. These nerves cannot be classed among the afferent nerves of the skin any more than the nerves of the optic and olfactory apparatus; they require separate consideration. A very extensive literature has grown up on the subject of this system of lateral line sense-organs and their innervation, the outcome of which is decisively in favour of this system being classed with the sense-organs supplied by the auditory nerve, so that in endeavouring to understand the position of the auditory nerve, we must always bear in mind that any theory as to its origin must apply to the system of lateral line nerves as well.
Now, although the auditory apparatus is common to all vertebrates, the lateral line system is not found in any land-dwelling animals; it belongs essentially to the fishes, and is, therefore, an old system so far as concerns the vertebrate group. Its sense-organs are arranged along the lateral line of the fish, and, in addition, on the head-region in three well-marked lines known as the supra-orbital, infra-orbital, and mandibular line systems. These sense-organs lie in the skin in a system of canals, and are innervated by a special nervous system different to that innervating adjacent skin-areas. The great peculiarity of their innervation consists in the fact that their nerves all belong to the branchial system of nerves; no fibres arise in connection with the trigeminal, but all of them in connection with the facial, glossopharyngeal and vagus nerves. In other words, although organs in the skin, their nerve-supply belongs to the lateral nervous system which supplies splanchnic and not somatic segments, a system which, according to the theory advanced in this book, originated in the nerves supplying appendages. The conclusion, therefore, is that in order to obtain some clue as to the origin of the sense-organs of this system in the assumed palæostracan ancestor, we must examine the mesosomatic appendages and see whether they possess any special sense-organs of similar function.
Further, considering that the auditory organ is to be regarded as a specially developed member of this system, we must especially look for an exceptionally developed organ in the region supplied by the auditory nerve.
The question of the origin of this system of lateral line sense-organs possesses a special interest for all those who attempt to obtain a solution of the origin of vertebrates, for the upholders of the view that the vertebrates have descended from annelids have alwaysfound its strongest support in the similarity of two sets of segmental organs found in annelids and vertebrates. On the one hand, great stress was laid upon the similarity of the segmental excretory organs in the two groups of animals, as will be discussed later; on the other, of the similarity of the segmentally arranged lateral sense-organs.
These lateral sense-organs of the annelids have been specially described by Eisig in the Capitellidæ, and, according to Lang, "there are many reasons for considering these lateral organs to be homologous with the dorsal cirri of the ventral parapodia of other Polychæta, and in the family of the Glyceridæ we can follow, almost step by step, the transformation of the cirri into lateral organs." Eisig describes them in the thoracic prebranchial region as slightly different from those in the abdominal branchial region; in the latter region, the ventral parapodia are gill-bearing, so that these lateral organs are in the branchial region closely connected with the branchiæ, just as is also the case in the vertebrates. It is but a small step from the gill-bearing ventral parapodia of the annelid to the gill-bearing appendages of the phyllopod-like protostracan; so that if we assume that this is the correct line along which to search for the origin of the vertebrate auditory apparatus, then, on my theory of the origin of the vertebrates from a group resembling the Protostraca, it follows that special sense-organs must have existed either on or in close connection with the branchial and prebranchial appendages of the protostracan ancestor of the vertebrates, which would form an intermediate link between the lateral organs of the annelids and the lateral and auditory organs of the vertebrates.
Further, these special sense-organs could not have been mere tactile hairs, but must have possessed some special function, and their structure must have been compatible with that function. Can we obtain any clear conception of the original function of this whole system of sense-organs?
A large amount of experimental work has been done to determine the function of the lateral line organs in fishes, and they have been thought at one time or another to be supplementary organs for equilibration, organs for estimating pressure, etc. The latest experimental work done by Parker points directly to their being organs for estimating slow vibrations in water in contradistinction to the quicker vibrations constituting sound. He concludes that surface wave-movements, whether produced by air moving on the water orsolid bodies falling into the water, are accompanied by disturbances which are stimuli for the lateral line organs.
One of these segmental organs has become especially important and exists throughout the whole vertebrate group, whether the animal lives on land or in water—this is the auditory organ. Throughout, the auditory organ has a double function—the function of hearing and the function of equilibration. If, then, this is, as is generally supposed, a specialized member of the group, it follows that the less specialized members must possess the commencement of both these functions, just as the experimental evidence suggests.
In our search, then, for the origin of the auditory organ of vertebrates, we must look for special organs for the estimation of vibrations and for the maintenance of the equilibrium of the animal, situated on the appendages, especially the branchial or mesosomatic appendages; and, further, we must specially look for an exceptional development of such segmental organs at the junction of the prosomatic and mesosomatic regions.
Throughout this book the evidence which I have put forward has in all cases pointed to the same conclusion, viz. that the vertebrate arose by way of the Cephalaspidæ from some arthropod, either belonging to, or closely allied to, the group called Palæostraca, of which the only living representative is Limulus. If, then, my argument so far is sound, the appendages of Limulus, both prosomatic and mesosomatic, ought to possess special sense-organs which are concerned in equilibration or the appreciation of the depth of the water, or in some modification of such function, and among these we might expect to find that somewhere at the junction of the prosoma and mesosoma such sense-organs were specially developed to form the beginning of the auditory organ.
Now, it is a striking fact that the appendages of Limulus do possess special sense-organs of a remarkable character, which are clearly not simply tactile. Thus Gegenbaur, as already stated, has drawn attention to the remarkable branchial sense-organs of Limulus; and Patten has pointed out that special organs, which he considers to be gustatory in function, are present on the mandibles of the prosomatic appendages. I myself, as mentioned in my address to the British Association at Liverpool in 1896, searched for some special sense-organ at the junction of the prosoma and mesosoma, and was rewarded by finding that that extraordinary adjunct to thelast locomotor appendage, known as the flabellum, was an elaborate sense-organ. I now propose to show that all these special sense-organs are constructed on a somewhat similar plan; that the structure of the branchial sense-organs suggests that they are organs for the estimation of water pressures; that among air-breathing arthropods sense-organs, built up on a somewhat similar plan, are universally found, and are considered to be of the nature of auditory and equilibration organs; and, what is especially of importance, in view of the fact that the most prominent members of the Palæostraca were the sea-scorpions, that the remarkable sense-organs of the scorpions known as the pectens belong apparently to the same group.
The Poriferous Sense-Organs of the Appendages in Limulus.
On all the branchial appendages in Limulus, special sense-organs are found of a most conspicuous character. They form in the living animal bluish convex circular patches, the situation of which on the appendages is shown in Fig.58. These organs are not found on the non-branchial operculum. Gegenbaur, who was the first to describe them, has pointed out how the surface of the organ is closely set with chitinous goblets shaped as seen in Fig.144, A, which do not necessarily project free on the surface, but are extruded on the slightest pressure. Each goblet fits into a socket in the chitinous covering, and is apparently easily protruded by variations of pressure from within. The whole surface of the organ on the appendage is slightly bulged in the living condition, and the chitin is markedly softer here than in the surrounding part of the limb. Each of these organs is surrounded by a thick protection of strongly branching spines. On the surface of the organ itself no spines are found, only these goblets, so that the surface-view presents an appearance as in Fig.144, B. Each goblet possesses a central pore, which is the termination of a very fine, very tortuous, very brittle chitinous tubule (ch.t.), which passes from the goblet through the layers of the chitin into the subjacent tissue. The goblets vary considerably in size, a few very large ones being scattered here and there. The fine chitinous tubule is especially conspicuous in connection with these largest goblets. In the smaller ones there is the same appearance of a pore and a commencing tube, but I have not been able to trace the tube through the chitinous layers, as in the case of the larger goblets.
Fig. 144.—A, A Goblet from one of the Branchial Sense-organs of Limulus(ch.t., chitinous tubule);B, Surface View of a Portion of a Branchial Sense-Organ.
Fig. 144.—A, A Goblet from one of the Branchial Sense-organs of Limulus(ch.t., chitinous tubule);B, Surface View of a Portion of a Branchial Sense-Organ.
Fig. 145.—The Endognaths of Limulus pushed out of the way on one side in order to show the position of the Flabellum(fl.)projecting towards the crack between the Prosomatic and Mesosomatic Carapaces.
Fig. 145.—The Endognaths of Limulus pushed out of the way on one side in order to show the position of the Flabellum(fl.)projecting towards the crack between the Prosomatic and Mesosomatic Carapaces.
Gegenbaur, in his picture, draws a straight tubule passing from every goblet among the fine canaliculi of the chitin. He says they are difficult to see, except in the case of the larger goblets. The tubule from the larger goblets is most conspicuous, and is in my sections always tortuous, never straight, as represented by Gegenbaur. A special branch of the appendage-nerve passes to these organs, and upon the fine branches of this nerve groups of ganglion-cells are seen, very similar in appearance to the groups described by Patten on the terminal branches of the nerves which supply the mandibular organs. At present I can see no mechanism by which the goblets are extruded or returned into place. In the case of the Capitellidæ, Eisig describes retractor muscles by means of which the lateral sense-organs arebrought below the level of the surface, and he imagines that the protrusion is effected by hydraulic means, by the aid of the vascular system. In the branchial sense-organs of Limulus there are no retractor muscles, and it seems to me that both retraction and protrusion must be brought about by alterations of pressure in the vascular fluids. Certainly the cavity of the organ is very vascular. If this be so, it seems likely enough that such an organ should be a very delicate organ for estimating changes in the pressure of the external medium, for the position of the goblets would depend on the relation between the pressure of the fluid inside the organ and that on the surface of the appendage. Whether the chitinous tubule contains a nerve-terminal or not I am unable to decide from my specimens, but, judging from Patten's description of the similar chitinous tubules belonging to the mandibular organs, it is most highly probable that these tubules also contain a fine terminal nerve-fibre.
These organs, then, represent segmental branchial sense-organs, of which it can be said their structure suggests that they may be pressure-organs; but the experimental evidence is at present wanting.
Passing now from the branchial to the prosomatic region, the first thing that struck me was the presence of that most conspicuous projection at the base of the last locomotor appendage, which is usually called the flabellum, and has been described by Lankester as an exopodite of this appendage. It is jointed on to the most basal portion of the limb (cf.Fig.155), and projects dorsally from the limb into the open slit between the prosomatic and mesosomatic carapace, as is seen in Fig.145(fl.). Of its two surfaces, the undermost is very convex and the uppermost nearly flat from side to side, the whole organ being bent, so that when the animal is lying half buried in sand, entirely covered over by the prosomatic and mesosomatic carapaces except along this slit between the two, the upper flat or slightly convex surface of the flabellum is exposed to any movement of water through this slit, and owing to its possessing a joint, the direction of the whole organ can be altered to a limited extent. The whole of this flat upper surface is one large sense-organ of a striking character, thus forming a great contrast to the convex under surface, which is remarkably free from tactile spines or special sense-organs.
The nerve going to the flabellum is very large, almost as large as the nerve to the rest of the appendage, and the very large majorityof the nerve-fibres turn towards the flat, uppermost side, where the sense-organ is situated. Between the nerve-fibres (n.) and the chitinous surface containing the special sense-tubes masses of cells (gl.) are seen, as in Fig.146, apparently nerve-cells, which form a broad border between the nerve-fibres and the pigmented chitinogenous layer (p.). On the opposite side, nothing of the sort intervenes between the pigmented layer and the blood-spaces and nerve-fibres which constitute the central mass of the flabellum.
Fig. 146.—Section through Flabellum.ch., chitinous layers;s.o., sense-organs;sp., spike-organ;p., pigment layer;gl., ganglion cell layer;bl.andn., blood-spaces and nerves.
Fig. 146.—Section through Flabellum.
ch., chitinous layers;s.o., sense-organs;sp., spike-organ;p., pigment layer;gl., ganglion cell layer;bl.andn., blood-spaces and nerves.
Fig. 147.—Section parallel to the Surface of Flabellum, showing the Porous Terminations of the Sense-Organs and the Arrangement of the Canaliculi round them.
Fig. 147.—Section parallel to the Surface of Flabellum, showing the Porous Terminations of the Sense-Organs and the Arrangement of the Canaliculi round them.
At present I am inclined to look upon this mass of cells as constituting a large ganglion, which extends over the whole length and breadth of the upper surface of the flabellum. At the sametime, my preparations are not sufficiently clear to enable me to trace out the connections of these cells, especially their connections with the special sense-organs.
Fig. 148.—Section through the three Sense-Organs of Flabellum.bl., blood-spaces;n., nerve;gl., layer of ganglion-cells;p., pigment layer;ch., 1, 2, 3, the three layers of chitin;ch.t., chitinous tubule in large tube of sense-organ;cap., capitellum or swollen extremity of large tube;can., very fine porous canals or canaliculi of chitin.
Fig. 148.—Section through the three Sense-Organs of Flabellum.bl., blood-spaces;n., nerve;gl., layer of ganglion-cells;p., pigment layer;ch., 1, 2, 3, the three layers of chitin;ch.t., chitinous tubule in large tube of sense-organ;cap., capitellum or swollen extremity of large tube;can., very fine porous canals or canaliculi of chitin.
Fig. 148.—Section through the three Sense-Organs of Flabellum.
bl., blood-spaces;n., nerve;gl., layer of ganglion-cells;p., pigment layer;ch., 1, 2, 3, the three layers of chitin;ch.t., chitinous tubule in large tube of sense-organ;cap., capitellum or swollen extremity of large tube;can., very fine porous canals or canaliculi of chitin.
In Fig.148I give a magnified representation of a section through three of these flabellar sense-organs. As is seen, the section divides itself into four zones: (1) the chitinous layer (ch.); (2) the layer of pigment (p.) and hypodermal cells; (3) the layer of ganglion-cells (gl.); and (4) the layer of nerve-fibres (n.) and blood-spaces (bl.). The chitinous layer is composed of the usual three zones of the Limulus surface—externally (Fig.148), a thin homogeneous layer, followed by a thick layer of chitin (3), in which the fine vertical tubules or canaliculi are well marked; the external portion (2) of this layer is differentiated from the rest by the presence of well-marked horizontal layers in addition to the canaliculi.
In this chitinous layer the special sense-organs are found. They consist of a large tube which passes through all the layers of the chitin except the thin homogeneous most external layer.This tube is conical in shape, its base, which rests on the pigmented layer, being so large and the organs so crowded together that a section of the chitin across the base of the tubes gives the appearance of a honeycomb, the septa of which is all that remains of the chitin. This large tube narrows down to a thin elongated neck as it passes through the chitin, and then, at its termination, bulges out again into an oval swelling (cap.) situated always beneath the homogeneous most external layer of chitin. Within this tube a fine chitinous tubule (ch. t.) is situated similar to that seen in the branchial sense-organs; it lies apparently free in the tube, not straight, but sinuous, and it passes right through all the chitinous layers to open at the surface as a pore; in the last part of its course, where it passes through the most external layer (1) of chitin, it lies always at right angles to the surface.
If the flabellum be stained with methylene blue and acid fuchsin, then all the canaliculi in the chitin show up as fine red lines, and present the appearance given in Fig.148, and it is seen that each of the terminations of the tubules is surrounded in the homogeneous layer of chitin by a thick-set circular patch of canaliculi which pass to the very surface of the chitin, while the canaliculi in other parts terminate at the commencement of the homogeneous layer and do not reach the surface. Further, the contents of the oval swelling, and, indeed, of the tube as a whole, are stained blue, the chitinous tubule being either unstained or slightly pink in colour. We see, then, that the chitinous tubule alone reaches the surface, while the large tube, which contains the tubule, terminates in an oval swelling, which often presents a folded or wrinkled appearance, as in Fig.149(see also Patten's Fig. 1, Plate I.). This terminal bulging of the tube is reminiscent of the bulging in the chitinous tubes of the lyriform organs of the Arachnida, as described by Gaubert, and of the poriferous chordotonal organs in insects, as described by Graber (see Fig.150). This terminal swelling is filled with a homogeneous refringent mass staining blue with methylene blue, in which I have seen no trace of a nucleus; through this the chitinous tubule makes its way without any sign of bulging on its part. Patten, in his description of the sense-organs on the mandibles of Limulus, which are evidently the same in structure as those on the flabellum, refers to this homogeneous mass as a coagulum. I doubt whether this is an adequate description; it appears to me to stain rather morereadily than a blood-coagulum, yet in the sense of being structureless it resembles a coagulum.
The enormous number of these organs crowded together over the whole flat surface of the flabellum produces a very striking appearance when viewed on the surface. Such a view presents an appearance resembling that of the surface-view of the branchial sense-organs; in both cases the surface is covered with a great number of closely set circular plaques, in the centre of each of which is seen a well-marked pore. The circular plaques in the case of the flabellum are much smaller than those of the branchial sense-organs, and clearly are not protrusible as in the latter organs, the appearance as of a plaque being due to the ring of thickly-set canaliculi round the central tubule, as already described. When stained with methylene blue, the surface view of the flabellum under a low power presents an appearance of innumerable circular blue masses, from each of which springs a fine bent hair, terminating in a pore at the surface. The blue masses are the homogeneous substance (cap.) of the bulgings seen through the transparent external layer of chitin, and the hairs are the terminal part of the chitinous tubules. Patten has represented their appearance in the mandibles in his Fig. 2, Plate I.
The large tubes in the chitin alter in shape according to their position. Those in the middle of the sensory surface of the flabellum, in their course through the chitinous layers, are hardly bent at all; as they approach the two lateral edges of this surface, their long thin neck becomes bent more and more, the bending always being directed towards the middle of the surface (see Fig.146); in this way the chitinous tubules increase more or less regularly in length from the centre of the organ to the periphery. The large basal part of the conical tube contains, besides the chitinous tubule, a number of nuclei which are confined to this part of the tube; some of these nuclei look like those belonging to nerve-fibres, others are apparently the nuclei of the chitinogenous membrane lining the tube. I have never seen any sign of nerve-cells in the tube itself.
The only other kind of sense-organ I have found in connection with these sense-organs are a few spike-like projections, the appearance of which is given in Fig.149. I have always seen these in the position given in Fig.146(sp.),i.e.at the junction of the surface which contains the sense-organs and the surface which is free from them. They are, so far as I have seen, not very numerous; I havenot, however, attempted to examine the whole sense-organ for the purpose of estimating their number and arrangement.
As is seen in Fig.149, they possess a fine tubule of the same character as that of the neighbouring sense-organs, which apparently terminates at the apex of the projecting spike. They appear to belong to the same group as the other poriferous sense-organs, and are of special interest, because in their appearance they form a link between the latter and the poriferous sense-organs which characterize the pecten of the scorpion (cf.Fig.152, C).