The Works of Francis Balfour 3-12
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Foster M. and Sedgwick A. The Works of Francis Balfour Vol. III. A Treatise on Comparative Embryology 2 (1885) MacMillan and Co., London.
- Cephalochorda | Urochorda | Elasmobranchii | Teleostei | Cyclostomata | Ganoidei | Amphibia | Aves | Reptilia | Mammalia | Comparison of the Formation of Germinal Layers and Early Stages in Vertebrate Development | Ancestral form of the Chordata | General Conclusions | Epidermis and Derivatives | The Nervous System | Organs of Vision | Auditory, Olfactory, and Lateral Line Sense Organs | Notochord, Vertebral Column, Ribs, and Sternum | The Skull | Pectoral and Pelvic Girdles and Limb Skeleton | Body Cavity, Vascular System and Glands | The Muscular System | Excretory Organs | Generative Organs and Genital Ducts | The Alimentary Canal and Appendages in Chordata
Foster M. and Sedgwick A. The Works of Francis Balfour Vol. II. A Treatise on Comparative Embryology 1. (1885) MacMillan and Co., London.
Foster M. and Sedgwick A. The Works of Francis Balfour Vol. III. A Treatise on Comparative Embryology 2 (1885) MacMillan and Co., London.
Foster M. and Sedgwick A. The Works of Francis Balfour Vol. IV. Plates (1885) MacMillan and Co., London.
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Vol. III. A Treatise on Comparative Embryology 2 (1885)
CHAPTER XII. OBSERVATIONS ON THE ANCESTRAL FORM OF THE CHORDATA
THE present section of this work would not be complete without some attempt to reconstruct, from the materials recorded in the previous chapters, and from those supplied by comparative anatomy, the characters of the ancestors of the Chordata ; and to trace as far as possible from what invertebrate stock this ancestor was derived.
The second of these questions has been recently dealt with in a very suggestive manner by both Dohrn (No. 250) and Semper (Nos. 255 and 256), but it is still so obscure that I shall refrain from any detailed discussion of it.
While differing very widely in many points both Dohrn and Semper have arrived at the view, already tentatively put forward by earlier anatomists, that the nearest allies of the Chordata are to be sought for amongst the Chaetopoda, and that the dorsal surface of the Chordata with the spinal cord corresponds morphologically with the ventral surface of the Chaetopods with the ventral ganglion chain. In discussing this subject some time ago x I suggested that we must look for the ancestors of the Chordata, not in allies of the present Chaetopoda, but in a stock of segmented forms descended from the same unsegmented types as the Chaetopoda, but in which two lateral nerve-cords, like those of Nemertines, coalesced dorsally, instead of ventrally to form a median nervous cord. This group of forms, if my suggestion as to its existence is well founded, appears now to have perished. The recent researches of Hubrecht on the anatomy of the Nemertines a have, however, added somewhat to the probability of my views, in that they shew that in some existing Nemertines the nerve-cords approach each other very closely in the dorsal line.
With reference to the characters of the ancestor of the Chordata the following pages contain a few tentative suggestions rather than an attempt to deal with the whole subject ; while the
1 Monograph on the development of Elasmobranch Fishes, pp. 170 173.
2 Hubrecht, "Zur Anat. u. Phys. d. Nervensystems der Nemertinen. " Kon. Akad. Wiss. Amsterdam; and "Researches on the Nervous System of Nemertines." Quart. Journ. of Micr. Science, 1880.
312 THE PR^iORAL LOBE.
origin of certain of the organs is dealt with in a more special manner in the chapters on organogeny which form the second part of this work.
Before entering upon the more special subject of this chapter, it will be convenient to clear the ground by insisting on a few morphological conclusions to be drawn from the study of Amphioxus, a form which, although probably in some respects degenerate, is nevertheless capable of furnishing on certain points very valuable evidence.
(1) In the first place it is clear from Amphioxus that the ancestors of the Chordata were segmented, and that their mesoblast was divided into myotomes which extended even into the region in front of the mouth. The mesoblast of the greater part of what is called the head in the Vertebrata proper was therefore segmented like that of the trunk.
(2) The only internal skeleton present was the unsegmented notochord a fact which demonstrates that the skeleton is of comparatively little importance for the solution of a large number of fundamental questions, as for example the point which has been mooted recently as to whether gill-clefts existed at one time in front of the present mouth ; and for this reason : that from the evidence of Amphioxus and the lower Vertebrata 1 it is clear that such clefts, if they ever existed, had atrophied
1 The greater part of the branchial skeleton of Petromyzon appears clearly to belong to an extra-branchial system much more superficially situated than the true branchial bars of the higher forms. At the same time-there is no doubt that certain parts of the skeleton of the adult Lamprey have, as pointed out by Huxley, striking points of resemblance to parts of a true mandibular and hyoid arches. Further embryological evidence is required on the subject, but the statements on this head on p. 84 ought to be qualified.
Should Huxley's views on this subject be finally proved correct, it is probable that, taking into consideration the resemblance of these skeletal parts in the Tadpole to those in the Lamprey, the cartilaginous mandibular bar, before being in any way modified to form true jaws, became secondarily adapted to support a suctorial mouth, and that it subsequently became converted into the true jaws. Thus the evolution of this bar in the Frog would be a true repetition of the ancestral history, while its ontogeny in Elasmobranchii and other types would be much abbreviated. For a fuller statement on this point I must refer the reader to the chapter on the skull.
It is difficult to believe that the posterior branchial bars could have coexisted with such a highly developed branchial skeleton as that in Petromyzon, so that the absence of the posterior branchial bars in Petromyzon receives by far its most plausible explanation on the supposition that Petromyzon is descended from a vertebrate stock in which true branchial bars had not been evolved.
ON THE ANCESTRAL FORM OF THE CHORDATA. 313
completely before the formation of cartilaginous branchial bars ; so that any skeletal structures in front of the mouth, which have been interpreted by morphologists as branchial bars, can never have acted in supporting the walls of branchial clefts.
(3) The region which, in the Vertebrata, forms the oesophagus and stomach, was, in the ancestors of the Chordata, perforated by gill-clefts. This fact, which has been clearly pointed out by Gegenbaur, is demonstrated by the arrangement of the gill-clefts in Amphioxus, and by the distribution of the vagus nerve in the Vertebrata 1 . On the other hand the insertion of the liver, which was probably a very primitive organ, appears to indicate with approximate certainty the posterior limit of the branchial clefts.
With these few preliminary observations we may pass to the main subject of this section. A fundamental question which presents itself on the threshold of our enquiries is the differentiation of the head.
In the Chaetopoda the head is formed of a praeoral lobe and of the oral segment ; while in Arthropods a somewhat variable number of segments are added behind to this primitive head, and form with it what may be called a secondary compound head. It is fairly clear that the section of the trunk, which, in Amphioxus, is perforated by the visceral clefts, has become the head in the Vertebrates proper, so that the latter forms are provided with a secondary head like that of Arthropods. There remain however difficult questions (i) as to the elements of which this head is composed, and (2) as to the extent of its differentiation in the ancestors of the Chordata.
In Arthropods and Chaetopods there is a very distinct element in the head known as the procephalic lobe in the case of Arthropods, and the praeoral lobe in that of Chaetopods ; and this lobe is especially characterized by the fact that the supracesophageal ganglia and optic organs are formed as differentia 1 The extension forwards in the vertebrata of an uninterrupted body-cavity into the region previously occupied by visceral clefts presents no difficulty. In Amphioxus the true body cavity extends forwards, more or less divided by the branchial clefts, for the whole length of the branchial region, and in embryos of the lower Vertebrata there is a section of the body cavity the so-called head-cavities between each pair of pouches. On the disappearance of the pouches all these parts would naturally coalesce into a continuous whole.
314 THE PR/KORAL LOBE.
tions of part of the epiblast covering it. Is such an element to be recognized in the head of the Chordata ? From a superficial examination of Amphioxus the answer would undoubtedly be no ; but then it has to be borne in mind that Amphioxus, in correlation with its habit of burying itself in sand, is especially degenerate in the development of its sense-organs ; so that it is not difficult to believe that its praeoral lobe may have become so reduced as not to be recognizable. In the true Vertebrata there is a portion of the head which has undoubtedly many features of the praeoral lobe in the types already alluded to, viz. the part containing the cerebral hemispheres and the thalamencephalon. If there is any part of the brain homologous with the supracesophageal ganglia of the Invertebrates, and it is difficult to believe there is not such a part, it must be part of, or contain, the fore-brain. The fore-brain resembles the supraoesophageal ganglia in being intimately connected in its development with the optic organs, and in supplying with nerves only organs of sense. Its connection with the olfactory organs is an argument in the same direction. Even in Amphioxus there is a small bulb at the end of the nervous tube supplying what is very probably the homologue of the olfactory organ of the Vertebrata ; and it is quite possible that this bulb is the reduced rudiment of what forms the fore-brain in the Vertebrata.
The evidence at our disposal appears to me to indicate that the third nerve belongs to the cranio-spinal series of segmental nerves, while the optic and olfactory nerves appear to me equally clearly not to belong to this series 1 . The mid-brain, as giving origin to the third nerve, would appear not to have been part of the ganglion of the prseoral lobe.
These considerations indicate with fair probability that the part of the head containing the fore-brain is the equivalent of the praeoral lobe of many Invertebrate forms ; and the primitive position of the Vertebrate mouth on the ventral side of the head affords a distinct support for this view. It must however be admitted that this part of the head is not sharply separated in development from that behind ; and, though the fore-brain is
1 Marshall, in his valuable paper on the development of the olfactory organ, takes a very different view of this subject. For a discussion of this view I must refer the reader to the chapter on the nervous system.
ON THE ANCESTRAL FORM OF THE CHORDATA. 315
usually differentiated very early as a distinct lobe of the primitive nervous tube, yet that such differentiation is hardly more marked than in the other parts of the brain. The termination of the notochord immediately behind the fore-brain is, however, an argument in favour of the morphological distinctness of the latter structure.
The evidence at our disposal appears to indicate that the posterior part of the head was not differentiated from the trunk in lower Chordata ; but that, as the Chordata rose in the scale of development, more and more centralizing work became thrown on the anterior part of the nervous cord, and part passu this part became differentiated into the mid- and hind-brain. An analogy for such a differentiation is supplied in the compound subcesophageal ganglion of many Arthropods ; and, as will be shewn in the chapter on the nervous system, there is strong embryological evidence that the mid- and hind-brains had primitively the same structure as the spinal cord. The head appears however to have suffered in the course of its differentiation a great concentration in its posterior part, which becomes progressively more marked, even within the limits of the surviving Vertebrata. This concentration is especially shewn in the structure of the vagus nerve, which, as first pointed out by Gegenbaur, bears evidence of having been originally composed of a great series of nerves, each supplying a visceral cleft. Rudiments of the posterior nerves still remain as the branches to the oesophagus and stomach 1 .
The atrophy of the posterior visceral clefts seems to have taken place simultaneously with the concentration of the neural part of the head ; but the former process did not proceed so rapidly as the latter, so that the visceral region of the head is longer in the lower Vertebrata than the neural region, and is dorsally overlapped by the anterior part of the spinal cord and the anterior muscle-plates (vide fig. 47).
On the above view the posterior part of the head must have been originally composed of a series of somites like those of the
1 The lateral branch of the vagus nerve probably became differentiated in connection with the lateral line, which seems to have been first formed in the head, and subsequently to have extended into the trunk (vide section on Lateral Line).
Till. MEDULLARY CANAL.
trunk, but in existing Vertebrata all trace of these, except in so far as they are indicated by the visceral clefts, has vanished in the adult. The cranial nerves however, especially in the embryo, still indicate the number of anterior somites ; and an embryonic segmentation of the mesoblast has also been found in many lower forms in the region of the head, giving rise to a series of cavities known as head-cavities, enclosed by mesoblastic walls which afterwards break up into muscles. These cavities correspond with the nerves, and it appears that there is a praemandibular cavity corresponding with the third nerve (fig. 193, \pp) and a mandibular cavity (2pp) and a cavity in each of the succeeding visceral arches. The fifth nerve, the seventh nerve, the glossopharyngeal nerve, and the successive elements of the vagus nerve correspond with the posterior head-cavities.
The medullary canal. The general history of the medullary plate seems to point to the conclusion that the central canal of the nervous system has been formed by a groove having appeared in the ancestor of the Chordata along the median dorsal line, which caused the sides of the nervous plate, which was placed immediately below the skin, or may perhaps at that stage not have been distinctly differentiated from the skin, to be bent upwards ; and that this groove subsequently became converted into a canal. This view is not only supported by the actual development of the central canal of the nervous system (the types of Teleostei, Lepidosteus and Petromyzon being undoubtedly secondary), but also (i) by the presence of cilia in the epithelium lining the canal, probably inherited from cilia coating the external skin, and (2) by
FIG. 193. TRANSVERSE SECTION THROUGH THE FRONT PART OF THE HEAD OF A YOUNG PRISTIURUS
The section, owing to the cranial flexure, cuts both the fore- and the hind-brain. It shews the prsemandibular and mandibular head-cavities ipp and ipp, etc.
fb. fore-brain; /. lens of eye; /. mouth ; pt. upper end of mouth, forming pituitary involution; iao.. mandibular aortic arch; ipp. and ipp. first and second head-cavities ; ivc. first visceral cleft ; V. fifth nerve ; aun. ganglion of auditory nerve ; VII. seventh nerve ; aa, dorsal aorta ; acv. anterior cardinal vein; ^..notochord.
ON THE ANCESTRAL FORM OF THE CHORDATA.
the posterior roots arising from the extreme dorsal line (fig. 194), a position which can most easily be explained on the supposition that the two sides of the plate, from which the nerves originally proceeded have been folded up so as to meet each other in the median dorsal line 1 .
The medullary plate, before becoming folded to form the medullary groove, is (except in Amphibia) without any indication of being composed of two halves. In both the embryo and adult the walls of the tube have however a structure which points to their having arisen from the coalescence of two lateral, and most probably at one time independent, cords ; and as already indicated this is the view I am myself inclined to adopt ; vide pp. 303 and
304 The origin and nature of the mouth. The most obvious point connected with the development of the mouth is the fact that in all vertebrate embryos it is placed ventrally, at some little distance from the front end of the body. This feature is retained in the adult stage in Elasmobranchii, the Myxinoids, and some Ganoids, but is lost in other vertebrate forms. A mouth, situated as is the embryonic vertebrate mouth, is very ill adapted for biting ; and though it acquires in this position a distinctly biting character in the Elasmobranchii, yet it is almost certain that it had not such a character in the ancestral Chordata, and that its terminal position in higher types indicates a step in advance of the Elasmobranchii.
On the structure of the primitive mouth there appears to me
FIG. 194. TRANSVERSE SECTION THROUGH THE TRUNK OF AN EMBRYO SLIGHTLY OLDER THAN FIG. 28 E.
nc. neural canal ; pr. posterior root of spinal nerve ; x. subnotochordal rod ; ao. aorta ; sc. somatic mesoblast ; sp. splanchnic mesoblast ; mp. muscle-plate ; mp'. portion of muscle-plate converted into muscle ; Vv. portion of the vertebral plate which will give rise to the vertebral bodies ; al. alimentary tract.
1 Vidf for further details the chapter on the nervous system.
318 PRIMITIVE SUCTORIAL MOUTH.
to be some interesting embryological evidence, to which attention has already been called in the preceding chapters. In a large number of the larvae or embryos of the lower Vertebrates the mouth has a more or less distinctly suctorial character, and is connected with suctorial organs which may be placed either in front of or behind it. The more important instances of this kind are (i) the Tadpoles of the Anura, with their posteriorly placed suctorial disc, (2) Lepidosteus larva (fig. 195) with its anteriorly placed suctorial disc, (3) the adhesive papillae of the larvae of the Tunicata. To these may be added the suctorial mouth of the Myxinoid fishes 1 .
All these considerations point to the conclusion that in the ancestral Chordata the mouth had a more or less definitely suctorial character 2 , and was placed on the ventral surface immediately behind the praeoral lobe; and that this mouth has become in the higher types gradually modified for biting purposes, and has been carried to the front end of the head.
The mouth in Elasmobranchii and other Vertebrates is originally a wide somewhat rhomboidal cavity (fig. 28 G) ; on the development of the mandibular and its maxillary (pterygoquadrate) process the opening of the mouth becomes narrowed to a slit. The wide condition of the mouth may not improbably be interpreted as a remnant of the suctorial state. The fact that no more definite remnants of the suctorial mouth are found in so primitive a group as the Elasmobranchii is probably to be explained by the fact that the members of this group undergo an abbreviated development within the egg.
1 The existing Myxinoid P'ishes are no doubt degenerate types, as was first clearly pointed out by Dohrn ; but at the same time (although Dohrn does not share this view) it appears to me almost certain that they are the remnants of a large and very primitive group, which have very likely been preserved owing to their parasitic or semiparasitic habits ; much in the same way as many of the Insectivora have been preserved owing to their subterranean habits. I am acquainted with no evidence, embryological or otherwise, that they are degraded gnathostomatous forms, and the group probably disappeared as a whole from its incapacity to compete successfully with Vertebrata in which true jaws had become developed.
3 I do not conceive that the existence of suctorial structures necessarily implies parasitic habits. They might be used for various purposes, especially by predaceous forms not provided with jaws.
ON THE ANCESTRAL FORM OF THE CHORDATA. 319
While the embryological data appear to me to point to the existence of a primitive suctorial mouth, very different conclusions have been put forward by other embryologists, more especially by Dohrn, which are sufficiently striking and suggestive to merit a further discussion.
As mentioned above, both Dohrn and Semper hold that the Vertebrata are descended from Chastopod-like forms, in which the ventral surface has become the dorsal. In consequence of this view Dohrn has arrived at the following conclusions : (i) that primitively the alimentary canal perforated the nervous system in the region of the original cesophageal nerve-ring ; (2) that there was therefore an original dorsal mouth (the present ventral mouth of the Cheetopoda) ; and (3) that the present mouth was secondary and derived from two visceral clefts which have ventrally coalesced.
A full discussion of these views 1 is not within the scope of this work ; but, while recognizing that there is much to be said in favour of the interchange of the dorsal and ventral surfaces, I am still inclined to hold that the difficulties involved in this view are so great that it must, provisionally at least, be rejected; and that there are therefore no reasons against supposing
FIG. 195. VENTRAL VIEW OF THE HEAD OF A LEPIDOSTEUS EMBRYO SHORTLY
BEFORE HATCHING, TO SHEW THE LARGE SUCTORIAL DISC.
m. mouth ; op. eye ; sd. suctorial disc.
the present vertebrate mouth to be the primitive mouth. There is no embryological evidence in favour of the view adopted by Dohrn that the present mouth was formed by the coalescence of two clefts.
If it is once admitted that the present mouth is the primitive mouth, and is more or less nearly in its original situation, very strong evidence will be required to shew that any structures originally situated in front of it are the remnants of visceral clefts ; and if it should be proved that such remnants of visceral clefts were present, the views so far arrived at in this section would, I think, have to be to a large extent reconsidered.
The nasal pits have been supposed by Dohrn to be remnants of visceral
1 For a partial discussion of this subject I would refer the reader to my Monograph on Elasmobranch Fishes, pp. 165 172.
320 FORMATION OF THE JAWS.
clefts, and this view has been maintained in a very able manner by Marshall. The arguments of Marshall do not, however, appear to me to have any great weight unless it is previously granted that there is an antecedent probability in favour of the presence of a pair of gill-clefts in the position of the nasal pits ; and even then the development of the nasal pits as epiblastic involutions, instead of hypoblastic outgrowths, is a serious difficulty which has not in my opinion been successfully met. A further argument of Marshall from the supposed segmental nature of the olfactory nerve has already been spoken of.
While most of the structures supposed to be remains of gill-clefts in front of the mouth do not appear to me to be of this nature, there is one organ which stands in a more doubtful category. This organ is the so-called choroid gland. The similarity of this organ to the pseudo-branch of the mandibular or hyoid arch was pointed out to me by Dohrn, and the suggestion was made by him that it is the remnant of a praemandibular gill which has been retained owing to its functional connection with the eye 1 . Admitting this explanation to be true (which however is by no means certain) are we necessarily compelled to hold that the choroid gland is the remnant of a gill-cleft originally situated in front of the mouth ? I believe not. It is easy to conceive that there may originally have been a praemandibular cleft behind the suctorial mouth, but that this cleft gradually atrophied (for the same reasons that the mandibular cleft shews a tendency to atrophy in existing fishes, &c.), the rudiment of the gill (choroid gland) alone remaining to mark its situation. After the disappearance of this cleft the suctorial mouth may have become relatively shifted backwards. In the meantime the branchial bars became developed, and as the mouth was changed into a biting one, the
1 The probability of the choroid gland having the meaning attributed to it by Dohrn is strengthened by the existence of a praemandibular segment as evidenced by the presence of a pnemandibular head-cavity, the walls of which as shewn by Marshall and myself give rise to the majority of the eye-muscles and of a nerve (the third nerve, cf. Marshall) corresponding to it; so that these parts together with the choroid gland may be rudiments belonging to the same segment. On the other hand the absence of the choroid gland in Ganoidei and Elasmobranchii, where a mandibular pseudo-branch is present, coupled with the absence of a mandibular pseudo-branch in Teleostei where alone a choroid gland is present, renders the above view about the choroid gland somewhat doubtful. A thorough investigation of the ontogeny of the choroid gland might throw further light on this interesting question, but I think it not impossible that the" choroid gland may be nothing else but the modified mandibtddr pseudo-branch, a view which fits in very well with the relations of the vessels of the Elasmobranch mandibular pseudo-branch to the choroid. For the relations and structure of the choroid gland vide F. Miiller, Vergl. Anal. Myxinoiden, Part in. p. 82.
It is possible that the fourth nerve and the superior oblique muscle of the eye which it supplies may be the last remaining remnants of a second praemandibular segment originally situated between the segment of the third nerve and that of the fifth nerve (mandibular segment).
ON THE ANCESTRAL FORM OF THE CHORDATA. 321
bar (the mandibular arch) supporting the then first cleft became gradually modified and converted into a supporting apparatus for the mouth, and finally formed the skeleton of the jaws. In the hyostylic Vertebrata the hyoid arch also became modified in connection with the formation of the jaws.
The conclusions arrived at may be summed up as follows : The relations which exist in all jaw-bearing Vertebrates between the mandibular arch and the oral aperture are secondary, and arose paripassu with the evolution of the jaws 1 .
The cranial flexure and the form of the head in vertebrate embryos. All embryologists who have studied the embryos of the various vertebrate groups have been struck with the remarkable similarity
FIG. 196. THE HEADS OF ELASMOBRANCH EMBRYOS AT TWO STAGES VIEWED
AS TRANSPARENT OBJECTS.
A. Pristiurus embryo of the same stage as fig. 28 F. B. Somewhat older Scyllium embryo.
///. third nerve; V. fifth nerve; VII. seventh nerve; au.n. auditory nerve; gl. glossopharyngeal nerve; Vg. vagus nerve; fb. fore-brain; pn. pineal gland; mb. midbrain; hb. hind-brain; iv.v. fourth ventricle; cb. cerebellum; ol. olfactory pit; op. eye; au.V. auditory vesicle; m. mesoblast at base of brain; ch. notochord; ht. heart; Vc. visceral clefts ; eg. external gills ; //. sections of body cavity in the head.
1 I do not mean to exclude the possibility of the mandibular arch having supported a suctorial mouth before it became converted into a pair of jaws.
B. III. 21
322 POST-ANAL GUT.
which exists between them, more especially as concerns the form of the head. This similarity is closest between the members of the Amniota, but there is also a very marked resemblance between the Amniota and the Elasmobranchii. The peculiarity in question, which is characteristically shewn in fig. 196, consists in the cerebral hemispheres and thalamencephalon being ventrally flexed to such an extent that the mid- brain forms the termination of the long axis of the body. At a later period in development the cerebral hemispheres come to be placed at the front end of the head ; but the original nick or bend of the floor of the brain is never got rid of.
It is obvious that in dealing with the light thrown by embryology on the ancestral form of the Chordata the significance of this peculiar character of the head of many vertebrate embryos must be discussed. Is the constancy of this character to be explained by supposing that at one period vertebrate ancestors had a head with the same features as the embryonic head of existing Vertebrata ?
This is the most obvious explanation, but it does not at the same time appear to me satisfactory. In the first place the mouth is so situated at the time of the maximum cranial flexure that it could hardly have been functional ; so that it is almost impossible to believe that an animal with a head such as that of these embryos can have existed.
Then again, this type of embryonic head is especially characteristic of the Amniota, all of which are developed in the egg. It is not generally so marked in the Ichthyopsida. In Amphibia, Teleostei, Ganoidae and Petromyzontidae, the head never completely acquires the peculiar characteristic form of the head of the Amniota, and all these forms are hatched at a relatively much earlier phase of development, so that they are leading a free existence at a stage when the embryos of the Amniota are not yet hatched. The only Ichthyopsidan type with a head like that of the Amniota is the Elasmobranchii, and the Elasmobranchii are the only Ichthyopsida which undergo the major part of their development within the egg.
These considerations appear to shew that the peculiar characters of the embryonic head above alluded to are in some way connected with an embryonic as opposed to a larval development ; and for reasons which are explained in the section on larval forms, it is probable that a larval development is a more faithful record of ancestral history than an embryonic development. The flexure at the base of the brain appears however to be a typical vertebrate character, but this flexure never led to a conformation of the head in the adult state similar to that of the embryos of the Amniota. The form of the head in these embryos is probably to be explained by supposing that some advantage is gained by a relatively early development of the brain, which appears to be its proximate cause ; and since these embryos had not to lead a free existence (for which such a form of the head would have been unsuited) there was nothing to interfere with the action of natural selection in bringing about this form of head during fcetal life.
Post-anal gut and neurenteric canal. One of the most
ON THE ANCESTRAL FORM OF THE CHORDATA. 323
remarkable structures in the trunk is the post-anal gut (fig. 197). Its structure is fully dealt with in the chapter on the alimentary tract, but attention may here be called to the light which it appears to throw on the characters of the ancestor of the Chordata.
In face of the facts which are known with reference to the post-anal section of the alimentary tract, it can hardly be doubted that this portion of the alimentary tract must have been at one time functional. This seems to me to be shewn (i) by the constancy and persistence of this obviously now functionless rudiment, (2) by its greater development in the lower than in the higher forms, (3) by its relation to the formation of the notochord and subnotochordal rod.
If the above position be admitted, it is not permissible to shirk the conclusions which seem necessarily to follow, however great the difficulties may be which are involved in their accept
FIG. 197. LONGITUDINAL SECTION THROUGH AN ADVANCED EMBRYO OF BOM BINATOR. (After Gotte.)
m. mouth; an. anus; /. liver; ne. neurenteric canal; me. medullary canal; ch. notochord; pn. pineal gland.
ance. These conclusions have in part already been dealt with by Dohrn in his suggestive tract (No. 250). In the first place the alimentary canal must primitively have been continued to the end of the tail ; and if so, it is hardly credible that the existing anus can have been the original one. Although, therefore, it is far from easy, on the physiological principles involved in the Darwinian theory, to understand the formation of a new anus 1 ; it is nevertheless necessary to believe that the present
1 Dohrn (No. 250, p. 25) gives an explanation of the origin of the new anus which does not appear to me quite satisfactory.
vertebrate anus is a formation acquired within the group of the Chordata, and not inherited from some older group. This involves a series of further consequences. The opening of the urinogenital ducts into the cloaca must also be secondary, and it is probable that the segmental tubes were primitively continued along the whole post-anal region of the vertebrate tail, opening into the body cavity which embryology proves to have been originally present there. They are in fact continued in many existing forms for some distance behind the present anus. If the present anus is secondary, there must have been a primitive anus, which was probably situated behind the post-anal vesicle ; and therefore in the region of the neurenteric canal. The neurenteric canal is, however, the remnant of the blastopore (vide p. 277). It follows, therefore, that tJie vertebrate blastopore is probably almost, if not exactly identical in position with the primitive aims. This consideration may assist in explaining the remarkable phenomenon of the existence of the neurenteric canal. The attempt has already been made to shew that the central canal of the nervous system is really a groove converted into a tube and lined by the external epidermis. This tube (as may be concluded from embryological considerations) was probably at first open posteriorly, and no doubt terminated at the primitive anus. On the closure of the primitive anal opening, the terminal portions of the post-anal gut and the neural tube, may conceivably have been so placed that both of them opened into a common cavity, which previously had communication with the exterior by the anus. Such an arrangement would necessarily result in the formation of a neurenteric canal. It seems not impossible that a dilated vesicle, often present at the end of the post-anal gut (vide fig. 28*, p. 58), may have been the common cavity into which both neural and alimentary tubes opened 1 .
1 As pointed out in Vol. II. p. 255, there is a striking similarity between the history of the neurenteric canal in Vertebrates, and the history of the blastopore and ventral groove as described by Kowalevsky in the larva of Chiton. Mr A. Sedgwick has pointed out to me that the ciliated ventral groove in Protoneomenia, which contains the anus, is probably the homologue of the groove found in the larva of Chiton, and not, as usually supposed, simply the foot. Were this groove to be converted into a canal, on the sides of which were placed the nervous cords, there would be formed a precisely similar neurenteric canal to that in Vertebrata, though I do not mean to suggest that there is any homology between the two (vide Hubrecht, Zool. Anzeigcr, 1880, p. 589).
ON THE ANCESTRAL FORM OF THE CHORDATA. 325
Till further light is thrown by fresh discoveries upon the primitive condition of the posterior continuation of the vertebrate alimentary tract, it is perhaps fruitless to attempt to work out more in detail the 'above speculation.
Body cavity and mesoblastic somites. The Chordata, or at least the most primitive existing members of the group, are characterized by the fact that the body cavity arises as a pair of outgrowths of the archenteric cavity. This feature 1 in the development is a nearly certain indication that the Chordata are a very primitive stock. The most remarkable point with reference to the development of the two outgrowths is, however, the fact that the dorsal part of each outgrowth becomes separated from the ventral. Its walls become segmented and form the mesoblastic somites, which eventually, on the obliteration of their cavity, give rise to the muscle-plates and to the tissue surrounding the notochord. It is not easy to understand the full significance of the processes concerned in the formation of the mesoblastic somites (vide p. 296). The mesoblastic somites have no doubt a striking resemblance to the mesoblastic somites of the Chsetopods, and most probably the segmentation of the mesoblast in the two groups is a phenomenon of the same nature ; but the difference in origin between the two types of mesoblastic somites is so striking, and the development of the muscular system from them is so dissimilar in the two groups, as to render a direct descent of the Chordata from the Chsetopoda very improbable. The ventral parts of the original outgrowth give rise to the permanent body cavity, which appears originally to have been divided into two parts by a dorsal and a ventral mesentery.
The notochord. The most characteristic organ of the Chordata is without doubt the notochord. The ontogenetic development of this organ probably indicates that it arose as a differentiation of the dorsal wall of the archenteron ; at the same time it is not perhaps safe to lay too much stress upon its mode of development. Embryological and anatomical evidence demonstrate, however, in the clearest manner that the early Chordata were provided with this organ as their sole axial skeleton ;
1 Vide the chapter on the Germinal Layers.
and no invertebrate group can fairly be regarded as genetically related to the Chordata till it can be shewn to possess some organ either derived from a notochord, or capable of having become developed into a notochord. No such organ has as yet been recognized in any invertebrate group 1 .
Gill-clefts. The gill-clefts, which are essentially pouches of the throat opening externally, constitute extremely characteristic organs of the Chordata, and have always been taken into consideration in any comparison between the Chordata and the Invertebrata.
Amongst the Invertebrata organs of undoubtedly the same nature are, so far as I know, only found in Balanoglossus, where they were discovered by Kowalevsky. The resemblance in this case is very striking ; but although it is quite possible that the gill-clefts in Balanoglossus are genetically connected with those of the Chordata, yet the organization of Balanoglossus is as a whole so different from that of the Chordata that no comparison can be instituted between the two groups in the present state of our knowledge.
Other organs of the Invertebrata have some resemblance to the gill-clefts. The lateral pits of the Nemertines, which appear to grow out as a pair of oesophageal diverticula, which are eventually placed in communication with the exterior by a pair of ciliated canals (vide Vol. II. pp. 200 and 202), are such organs.
Semper (No. 256) has made the interesting discovery that in the budding of Nais and Chaetogaster two lateral masses of cells, in each of which a lumen may be formed, unite with the oral invagination and primitive alimentary canal to form the permanent cephalic gut. The lateral masses of cells are regarded by him as branchial passages homologous in some way with those in the Chordata. The somewhat scanty observations on this subject which he has recorded do not appear to me to lend much support to this interpretation.
It is probable that the part of the alimentary tract in which gill- clefts are present was originally a simple unperforated tube provided with highly vascular walls ; and that respiration was carried on in it by the alternate introduction and expulsion of sea water. A more or less similar mode of respiration has been recently shewn by Eisig 2 to take place in the fore part
1 In the Chaetopods various organs have been interpreted as rudiments of a notochord, but none of these interpretations will bear examination.
2 " Ueb. d. Vorkommen eines schwimmblasenahnlichen Organs bei Anneliden." Mittheil. a. d. zoo!. Station zu Neapel, Vol. n. 1881.
ON THE ANCESTRAL FORM OF THE CHORDATA. 327
of the alimentary tract of many Chastopods. This part of the alimentary tract was probably provided with paired cascal pouches with their blind ends in contiguity with the skin.
Perforations placing these pouches in communication with the exterior must be supposed to have been formed ; and the existence of openings into the alimentary tract at the end of the tentacles of many Actinias and of the hepatic diverticula of some nudibranchiate Molluscs (Eolis, &C. 1 ) shews that such perforations may easily be made. On the formation of such perforations the water taken in at the mouth would pass out by them ; and the respiration would be localized in the walls of the pouches leading to them, and thus the typical mode of respiration of the Chordata would be established.
Phylogeny of the Chordata. It may be convenient to shew in a definite way the bearing of the above speculations on the phylogeny of the Chordata. For this purpose, I have drawn up the subjoined table, which exhibits what I believe to be the relationships of the existing groups of the Chordata. Such a table cannot of course be constructed from embryological data alone, and it does not fall within the scope of this work to defend its parts in detail.
L- T J
. PROTO-AMNIOTA AMPHIBIA
Cephafochorda PROTOCHORDATA Uroc/iorda
In the above table the names printed in large capitals are hypothetical groups. The other groups are all in existence at the present day, hut those printed in Italics are probably degenerate.
The ancestral forms of the Chordata, which may be called the Protochordata, must be supposed to have had (i) a
1 The openings of the hepatic diverticula through the sacks lined with thread cells are described by Hancock and Embleton, Ann. and Mag. of Nat. History, Vol. xv. 1845, p. 82. Von Jhering has also recently described these openings (Zool. Anzeiger, No. 23) and apparently attributes their discovery to himself.
328 PHYLOGENY OF THE CHORDATA.
notochord as their sole axial skeleton, (2) a ventral mouth, surrounded by suctorial structures, and (3) very numerous gill-slits. Two degenerate offshoots of this stock still persist in Amphioxus (Cephalochorda), and the Ascidians (Urochorda).
The direct descendants of the ancestral Chordata, were probably a group which may be called the Proto-vertebrata, of which there is no persisting representative. In this group, imperfect neural arches were probably present ; and a ventral suctorial mouth without a mandible and maxillae was still persistent. The branchial clefts had, however, become reduced in number, and were provided with gill-folds ; and a secondary head (vide p. 313), with brain and organs of sense like those of the higher Vertebrata, had become formed.
The Cyclostomata are probably a degenerate offshoot of this group.
With the development of the branchial bars, and the conversion of the mandibular bar into the skeleton of the jaws, we come to the Proto-gnathostomata. The nearest living representatives of this group are the Elasmobranchii, which still retain in the adult state the ventrally placed mouth. Owing to the development of food-yolk in the Elasmobranch ovum the early stages of development are to some extent abbreviated, and almost all trace of a stage with a suctorial mouth has become lost.
We next come to an hypothetical group which we may call the Proto-ganoidei. Bridge, in his memoir on Polyodon 1 , which contains some very interesting speculations on the affinities of the Ganoids, has called this group the Pneumatoccela, from the fact that we find for the first time a full development of the air-bladder, though it is possible that a rudiment of this organ, in the form of a pouch opening on the dorsal side of the stomachic extremity of the oesophagus, was present in the earlier type.
Existing Ganoids are descendants of the Proto-ganoidei. Some of them at all events retain in larval life the suctorial mouth of the Proto-vertebrata ; and the mode of formation of their germinal layers, resembling as it does that in the Lamprey
1 Phil. Trans. 1878. Part II.
ON THE ANCESTRAL FORM OF THE CHORDATA. 329
and the Amphibia, probably indicates that they are not descended from forms with a large food-yolk like that of Elasmobranchii, and that the latter group is therefore a lateral offshoot from the main line of descent.
Of the two groups into which the Ganoidei may be divided it is clear that certain members of the one (Teleostoidei), viz. Lepidosteus and Amia, shew approximations to the Teleostei, which no doubt originated from the Ganoids ; while the other (Selachoidei or Sturiones) is more nearly related to the Dipnoi. Polypterus has also marked affinities in this direction, e.g. the external gills of the larva (vide p. 1 18).
The Teleostei, which have in common a meroblastic segmentation, had probably a Ganoid ancestor, the ova of which were provided with a large amount of food-yolk. In most existing Teleostei, the ovum has become again reduced in size, but the meroblastic segmentation has been preserved. It is quite possible that Amia may also be a descendant of the Ganoid ancestor of the Teleostei ; but Lepidosteus, as shewn by its complete segmentation, is clearly not so.
The Dipnoi as well as all the higher Vertebrata are descendants of the Proto-ganoidei.
The character of the limbs of higher Vertebrata indicates that there was an ancestral group, which may be called the Proto-pentadactyloidei, in which the pentadactyle limb became established ; and that to this group the common ancestor of the Amphibia and Amniota belonged.
It is possible that the Plesiosauri and Ichthyosauri of Mesozoic times may have been more nearly related to this group than either to the Amniota or the Amphibia. The Proto-pentadactyloidei were probably much more closely related to the Amphibia than to the Amniota. They certainly must have been capable of living in water as well as on land, and had of course persistent branchial clefts. It is also fairly certain that they were not provided with large-yolked ova, otherwise the mode of formation of the layers in Amphibia could not be easily explained.
The Mammalia and Sauropsida are probably independent offshoots from a common stem which may be called the Protoamniota.
(249) F. M. Balfour. A Monograph on the development of Elasmobranch Fishes, London, 1878.
(250) A. Dohrn. Der (Jrsprung d. Wirbelthiere und d. Princip. d. Functionswechsel. Leipzig, 1875.
(251) E. Haeckel. Sch'dpfungsgeschichte. Leipzig. Vide also Translation. The History of Creation. King and Co. , London. 1876.
(252) E. Haeckel. Anthropogenie. Leipzig. Vide also Translation. Anthropogeny. Kegan Paul and Co., London, 1878.
(253) A. Kowalevsky. " Entwicklungsgeschichte d. Amphioxus lanceolatus." Mem. Acad. d. Scien. St Petersbourg, Ser. VII. Tom. XI. 1867, and Archivf. mikr. Anat., Vol. xin. 1877.
(254) A. Kowalevsky. "Weitere Stud. lib. d. Entwick. d. einfachen Ascidien." Archivf. mikr. Anat., Vol. VII. 1871.
(255) C. Semper. "Die Stammesverwandschaft d. Wirbelthiere u. Wirbellosen." Arbeit, a. d. zool.-zoot. Instit. Wurzburg, Vol. II. 1875.
(256) C. Semper. "Die Verwandschaftbeziehungen d. gegliederten Thiere." Arbeit, a. d. zool.-zoot. Instit. Wurzburg, Vol. III. 1876 1877.