Cole 1926

( 309 )
IV.—A Monograph on the general Morphology of the Myxinoid Fishes, based on a study
of Myxine. Part VI. The Morphology of the Vascular System. By F. J. Cole,
D.Sc. Oxon., Professor of Zoology, University College, Reading. Communicated by
Professor J. H. ASHWORTH, D . S C , F.R.S. (With Two Text-figures and Five Plates.)
(MS. received July 17, 1925. Read February 9, 1925. Issued separately November 6.)
CONTENTS.
PAGE PAGE
A. Thymus and Pancreas 310 10. Subcesophageal Sinus 327
B. Pericardial Cavity 310 11. Carotid Sinus . . . . 327
C. Arterial System 312 F. Red Lymphatics—Peribranchial Cycle-
D. Venous System 314 1. Cardiac Sinus . . . . 327
E. Lacunar Spaces. Red Lymphatics—Subcutaneous 2. Peribranchial Sinuses 328
Cycle— 3. Peribranchial Anastomosis 329
1. Subcutaneous Sinus 322 4. Subchordal Sinus . 329
2. Subcutaneous Anastomosis . . . . 323 G. White Lymphatics—
3. Rostral Sinus 324 1. Lateral Chordal Sinus 330
4. Dorsal Oral Sinu3 324 2. Segmental Lymphatics . 331
5. Hypophysio-velar Sinus * 324 3. Intestinal Lymphatic Trunk . 331
6. Subhypophysial Sinus 325 H. Circulation of the Blood . 332
7. Ventral Sinus 325 J. Summary 336
8. Dental Sinus 326 K. Literature 337
9. Lingual Sinus 326 L. Explanation of the Plates. 339
The first five parts of this work on the skeleton, muscles, vascular system, and visceral
organs were published in the Transactions of the Society in 1905, 1907, 1909, 1912, and 1913.
The present section deals with a difficult and puzzling system which adds to its funda-
mental incongruity a range of variation which reduces the normal status to a condition of
speculative uncertainty. The works of A. A. RETZITJS, J. MULLER, and JACKSON have resulted
in a fairly accurate knowledge of the arterial system ; but the venous system, which is much
more important as regards its morphological bearings, is only imperfectly known, and the
series of so-called veno-lymphatic spaces hardly known at all. Unhappily, material for a
study of the development of the vascular system of Myxine has still to be obtained. The
smallest individual out of many hundreds which I have collected was over 10 cm. long.
BEARD records a badly damaged 6'5 cm. specimen and MAAS one of 8 "5 cm. The structure
of BEARD'S example, which I have examined, does not differ in any noteworthy respect from
that of the adult, and hence throws no light on embryonic conditions. Consequently the
interpretation of the venous system which has been adopted in this paper awaits embryological
confirmation, and is to be regarded in the meantime as a working hypothesis only. The
spawning grounds of Myxine have not yet been discovered, and it seems likely that the
animal spawns in deep water in places remote from its summer resorts.
The terminology of the skeletal system is the same as that adopted in Parts I and III
of this work, which it would be premature to change at this stage, but it must not be regarded
as having any established morphological significance. Similarly, in the absence of develop-
mental data, it would be unwise to attempt any systematic comparison of the Myxinoid
blood-vessels with those of higher Vertebrates ; and even when this information is available,
as it is in the case of the allied Lampreys, the task is not an easy one (cf. D E BEER, Q.J.M.S.
TRANS. ROY. SOC. EDIN., VOL. LIV, PART II (NO. 4). 41
310 PROFESSOR FRANK J. COLE
68, p. 327). The terminology adopted in the following description is therefore subject to
revision.
I am indebted to my museum assistant, Mr W. E. STONEMAN, B.A., who has, in a series
of dissections and injections, checked many of the results here described.
The expenses of this research have been defrayed by a grant from the Government Grant
Committee of the Royal Society.
A. THYMUS AND PANCREAS.
A transient thymus-like organ was first described in a Marsipobranch in 1894 by

SCHAFFEB, who found a series of branchiomeric lymph-cell masses in the Ammoccete of
Petromyzon planeri.* In 1906, however, he stated that subsequent investigations had con-
vinced him that these masses could not be directly compared with any known thymus struc-
tures in higher fishes. In Myxine a thymus has been described by J. MtiLLER and STANNIUS,
but this so-called thymus is now identified as the pronephros. The only structure in Myxine
which may be interpreted as a thymus is an extensive lymphoid organ, exhibiting signs of
degeneration, which is found in the pharyngeal velum associated with the anterior two-thirds
of the external lateral velar -bar. It was figured in my Part V, Plate IV, fig. 10, and in the
present communication is shown in fig. 6, th.
Apart from the very small organ surrounding a portion of the bile duct, described in
Myxine and Bdellostoma by MAAS in 1896, a definitive pancreas is apparently absent in
Myxinoids. I have already shown that whatever function MAAS' organ may have, its size and
relations rule it out as a possible functional pancreas. The first point to determine, therefore,
was the function of the liver. That it is a true Vertebrate liver I showed in my last part,
and since then I have found that the secretion of the liver, when activated, digests fibrin and
coagulated white of egg. Consequently the liver of Myxine must be regarded as an hepato-
pancreas. In my last part I stated that microscopically there is only one kind of tubule in
the liver, and that I had not succeeded in finding any glandular tissue which might constitute
a histological pancreas. I had, however, noticed that the liver contained small aggregations
of tubules surrounding certain veins which contrasted sharply with the bulk of the liver
tubules, but I had not then observed the significant fact that these aggregations are invariably
associated with the branches of the portal vein in the liver and with no other vessels. This
at once recalled the condition of the pancreas in other fishes, where it is represented by
glandular acini surrounding the branches of the portal vein within the substance of the liver
parenchyma. Such a state of affairs appears to be paralleled exactly in the liver of Myxine,
and is represented diagrammatically in fig. 9.
B. PERICARDIAL CAVITY (Fig. 9).
I have nothing of importance to add to the structure of the systemic heart, but the
relations of the abdominal and pericardial coeloms must be dealt with. These two cavities
overlap each other—the abdominal coelom on the right side extending almost as far forwards
as the pericardial. The latter lies entirely dorsal to the former, the anterior lobe of the
liver almost completely separating the two cavities, which are only in contact on the extreme
right where the abdominal coelom bends over the right edge of the liver on to its dorsal sur-
face, at this point being in close contact with the pericardial coelom. It is here that the
* GIACOMINI does not confirm SCHAFFBB'S work on the thymus of Ammocuetes.
ON THE GENERAL MORPHOLOGY OF THE MYXINOID FISHES. 311
posterior wall of the latter cavity becomes continuous with the wall of the abdominal coelom
to form the pericardio-peritoneal foramen (fig. 9, p.p.f.), which is thus situated on the right
side and was described for the first time by J. MULLER in 183G. Through this, foramen the
portal vein appears to pass, but of course is morphologically outside it. The relations of the
foramen only are indicated in fig. 9, the actual position being much further back. The pos-
terior boundary of the pericardial cavity is irregular, and on the left side extends beyond the
anterior extremity of the posterior lobe of the liver. On the right side it also extends far
back, and almost entirely encloses the anterior end of the portal vein. Anteriorly it is
bounded on the right side by the last gill sac and on the left by the ductus oesophago-cutaneus.
It will be noticed on examining fig. 9, in which the pericardial cavity is represented in
black, that the heart projects into it from the ventral side and the portal heart from the
dorso-median. In the former case a wide gap is left ventrally on the left side which is
occupied by the sinus venosus, so that a ventral cardiac ligament or mesocardium, described
by JACKSON in Bdellostoma, has only an indefinite existence in Myxine. On the other hand,
the portal heart (and also the anterior portion of the portal vein) is almost completely
enclosed, forming a portal septum on the dorso-median side of these structures (cf. fig. 9).
It should be noted that the heart is pushed into the pericardial cavity from in front and
below, and the anterior extremity of the ventricle projects slightly beyond it. Like the
portal heart and vein, the common portal vein is sunk into the pericardial ccelom from above.
The distinction between the cardiac sinus (cd.s.), which is a true blood space and actually
consists of a very coarse venous plexus almost entirely surrounding the ventricle, and the
true pericardial cavity is very obvious in fig. 9. MECKEL erroneously identified the cardiac
sinus as the pericardial cavity, but the mistake was corrected by J. MULLER. JACKSON does
not mention the cardiac sinus, which he appears to have confused partly with what he calls
the " inner chamber of the right pericardial cavity."
Whilst the pericardial space is a single continuous cavity, it may be conveniently divided
into three regions. The central one encloses the ventricle of the heart but does not pass
anteriorly on to the ventral aorta, which lies in a forward extension of the cardiac sinus. The
central chamber is situated between the anterior lobe of the liver and the gut, and behind
the ventricle it becomes greatly narrowed dorso-ventrally. The right chamber is related to,
and more or less encloses, the dorsal division of the anterior section of the right posterior
cardinal vein with its included pronephros (r.a.c.1), the portal heart, portal vein, common
portal vein and systemic aorta, and communicates with the abdominal ccelom by the large
pericardio-peritoneal foramen, which forms its posterior boundary {p.p.f.). The left chamber
commences in front at the left of the ductus oesophago-cutaneus. It is related to, and more
or less encloses, the auricle of the heart, the dorsal division of the anterior section of the left
posterior cardinal vein with its included pronephros (l.a.c.1), the ventral division of the same
(l.a.c"), and sinus venosus.
It is important to note the asymmetrical disposition of the Myxinoid heart, since this is
clearly associated with other and more striking anomalies of the vascular system. The
ventricle is almost median but slightly to the right, whilst the auricle and sinus venosus are
well to the left, the auricle being entirely dorsal to the sinus venosus, which latter is lateral
and slightly ventral to the ventricle. The auricle is dorso-lateral to the ventricle. It is
obvious that this arrangement is quite different from that of the Selachian heart, and the
developmental history of the Myxinoid heart should prove very interesting. Its asymmetry
is accompanied by an asymmetry in the respiratory organs—the ductus cesophago-cutaneus,
which represents presumably a seventh gill, being restricted to the left side.
C. T H E ARTERIAL SYSTEM (Fig. l).
The arterial system of Myxinoids is well known from the publications of J. MILLER
and JACKSON, and I propose to refer to a few points only which have been overlooked.
The coeliac or gastric artery (coe.a., coe.a.') is paired, the left, the larger of the two,
usually arising behind the right from the anterior extremity of the systemic aorta, but
the left may be the more anterior of the pair. The left coeliac near its origin is generally
attached to the anterior border of the bile duct, and it gives off the hepatic artery when
close to the gall bladder, finally crossing the bile duct to be distributed to the gut and
occasionally to the posterior liver also. A somewhat different condition is figured in my
Part V, PL II, fig. 4, where the hepatic artery is given off earlier in the course of the coeliac,
and is the vessel which accompanies the bile duct. The two coeliacs form a ring round
the gut, and meet on its ventral wall. They may there fuse completely so as to form a
single vessel, but they may only be connected up by an anastomosis, or may even remain
completely distinct. The hepatic artery (h.a.) is a branch of the left coeliac, and normally
divides into three branches to the gall bladder and two lobes of the liver, but in one case
the hepatic supplied only the anterior liver and gall bladder, the posterior liver deriving its
artery from the right coeliac. The coeliacs supply the gut as far back as the level of the
posterior extremity of the liver.
The pronephros (pn.) usually receives its arterial blood from the coeliac. Generally only
one artery is present on each side, and this passes to the glomus (fig. 1, right side), but there
may be more than one, in which case the body of the pronephros may receive an artery.
The pronephros, however, may be supplied by a branch from a segmental artery arising
from the aorta behind the coeliac of its side (fig. 1, left side), or from the root of the latter.
In one case the glomus of the pronephros received branches both from the cceliac and from a
segmental artery, which latter arose from the aorta some little distance behind the coeliac of
its side. Several other variations might be recorded. The isolated Malpighian bodies with
attached fragments of pronephric duct (pn.'), which are commonly found between the
pronephros and the anterior extremity of the abdominal kidney, are always supplied by
branches from segmental arteries (fig. 1, both sides).
In Bdellostoma, according to JACKSON, the coeliac artery is not paired, does not normally
supply the pronephros. and lacks the posterior branch or branches to the wall of the gut.
J. MULLER mentions only one coeliac artery (the left), but states that the pronephros (also
presumably the left) derives its branches mostly from it, and further, that it supplies the
liver, gall bladder, and the " stomach."
The position of the heart coincides in the tranverse plane with the constitution of the
single systemic posterior aorta (fig. l).
The posterior or abdominal aorta is formed by the union of three main arteries—a
median and two lateral. The former is the anterior systemic aorta or median anterior aorta
or vertebralis impar, and lies over the gut, not necessarily in the median position, whilst the
latter are the common carotid arteries or paired anterior aortse, situated at the side of the
gut. The common carotids are sometimes regarded as commissural vessels, and in the gill
region they certainly have that appearance in Bdellostoma, as J. MULLER points out, but their
integrity is maintained for a considerable distance in front of the gill region. The median
and paired anterior aortae are connected up by four pairs of anastomoses, of which the last
pair are usually very small and may be absent. I have, however, never found less than
three. The position of the anastomoses also varies, and they are apparently not always
ON THE GENERAL MORPHOLOGY OF THE MYXINOLD FISHES. 313
homologous in different individuals. Each gill has two efferent branchial arteries coursing
respectively in front of and behind the corresponding efferent gill duct. In Bdellostoma
there are one or two efferent gill arteries according to species. The only arteries which
arise from the median aorta are segmental arteries, the gut and all muscles other than those
of the body wall being supplied from the common carotid. The segmental arteries have no
regular arrangement, and, as in the case of the segmental veins, a separate vessel is not
detached for each segment, there being about four arteries to every seven segments.
In the gill region the common carotids vary considerably. The artery widens and
narrows in a capricious manner, loops are formed, and the vascular papillse, described in my
fourth part, are more numerous and complex than elsewhere. Posteriorly the artery may
become so thin as to be almost imperforate, and the object of the anastomoses appears to be
to ensure a passage for the arterial blood to the median aorta. This applies particularly to
the posterior half of the gill region. In front the artery preserves an even calibre, and its
individuality cannot be questioned. The vascular papillae are confined to the common
carotid, and no trace of them appears on the median aorta. They occur along the whole
course of the common carotid from the point where the posterior abdominal aorta is con-
stituted to where the carotid divides to form the external and " internal" carotids. In one
specimen I counted 95 on the left carotid and 85 on the right. For further details of these
curious structures the reader is referred to my Part IV (4).
On approaching the skull, but whilst still well behind it, the median and lateral aortae
break up. The common carotid divides into external and " internal" carotids, but as the
latter amongst other things supply somatic musculature and the skin, they are obviously
something more than this name implies. These latter vessels (fig. 1, v.i.'") by their fusion
in the middle line above the gut constitute the factors of what J. MULLER calls the unpaired
vertebral artery of the head or the vertebralis impar capitis (v.i/). This vessel was first seen
by MtiLLER, and the fusion which forms it completes what he calls the circulus cephalicus of
Myxinoids. At the point where the fusion occurs an artery is despatched downwards into
the velum (vel.a.), and in this way red blood reaches the velar portion of the hypophysio-
velar sinus.
The median aorta, somewhat anterior to the point where the common carotid splits,
divides into three vessels, which, omitting certain minor complexities, behave as follows
(fig. 1) :—The median one unites with the left factor of the vertebralis impar capitis, and
the paired vessels pass upwards and forwards to the side of the notochord, the right one,
however, sending an anastomosis to the right factor of the vertebralis impar capitis as it
passes it. The paired vessels give off two segmental vessels, and also send branches to the
constrictor pharyngis, velo-spinalis, and cranio-hyoideus muscles.
The median portion of the vertebralis impar capitis, after a short course covering about
three segments over the roof of the gut behind and the hypophysial tube in front, splits into
two vessels (figs. 1 and 6, v.i.), which bend dorsally and follow the inner ventral diverging
margin of the auditory capsule dorsal to the hypophysial tube (fig. 6, h.c). They give off
the artery to the brain (cb.a.), which passes upwards median to the trabecula, and perhaps
represents the true internal carotid, and is the first conspicuous vessel to arise from the
vertebralis impar capitis counting from behind. This artery is difficult to find even in
sections, and was not seen by J. MTJLLER.* The branches of the median section, closely
attached but external to the membranous cranium, then pass upwards and outwards internal
to the trabecula, and hence do not pass through any of the lateral fenestrae of the skull.
* According to STERZI there are three encephalic arteries.
314 PROFFSSOR FRANK J. COLE
They continue their upward and outward course, bend externally over the trabecular and
palatine bar, and finally pass under the eye to their final distribution. They therefore course
obliquely forwards, upwards, and outwards through the big gap in the base of the skull
bounded by the notochord, auditory capsules, trabeculse, and hypophysial plate. They
supply the M.M. velo-quadratus, copulo-quadratus profundus, tentacularis posterior, palato-
ethmoidalis superficialis, and the segmental muscle (parietalis), and a prominent branch is also
despatched to the skin (v.i.").
The external carotids (figs. 1, 4, 5, 6, ex.c.) from their posterior origin pass forwards and
downwards round the side of the gut on to its ventro-lateral surface, where they take up a
position just external to the ventral or lower division of the second branchial arch and the
lateral margin of the sub-cesophageal sinus. They now travel straight forwards in this
position, passing internal to the first branchial arch, until they reach the lateral margin of
the middle segment of the basal plate, which they now accompany. In this way they reach
and thereafter follow the lateral margin of the external bars of the anterior segment of the
basal plate, which, owing to its greater width and spread, extends upwards at the side of the
gut, so that the arteries are now lateral to the gut. They continue to pass upwards and
forwards, and just in front of the anterior extremity of the basal plate they lie dorso-lateral
to the mouth between the copulo-ethmoidalis and palato-ethmoidalis profundus muscles.
From this point they course at first outwards and then inwards and upwards between the
lateral labial cartilage and the dorsal limb of the tentacularis posterior muscle to their final
distribution at the tip of the snout.
It is largely by the anterior terminal branches of the external carotid artery that blood
enters the subcutaneous sinus.
Posteriorly the external carotids give off segmental vessels, but in front the segmental
musculature does not exist, and hence there are no segmental arteries and veins anterior to
the nasal capsule. The external carotids otherwise supply the gut, the MM. hyo-copulo-
palatinus, copulo-palatinus, quadrato-palatinus, copulo-quadratus profundus, copulo-quadratus
superficialis, palato-ethmoidalis profundus, tentacularis posterior, nasalis, transversus oris and
ethmoideo-nasalis, and also the skin of the snout. Two important branches are those which
supply the tongue—the anterior (l.a.) and posterior (La.1) lingual arteries. The former is the
more important, and supplies the greater part of the tongue. The latter reaches the tongue
via the " tether " of the hyo-copulo-glossus muscle, and supplies the ventro-lateral region of
the posterior extremity of this organ. In this way red blood reaches the dental sinus, in all
parts of which it is found.
On the tail the aorta splits into two vessels somewhat in front of the anterior knob of
the median ventral bar of the caudal fin skeleton (fig. 10, r.c.a., l.c.a.). The right vessel
passes on to the ventro-lateral surface of the notochord on this side and becomes small and
inconspicuous (r.c.a.). Further back it enlarges again, and the two vessels balance both as
regards size and position. They are, of course, separated by the median ventral bar of the
caudal fin skeleton. Posteriorly the caudal arteries are connected by a capillary system with
the factors of the caudal vein.
D. THE VENOUS SYSTEM.
The best description to date of the Myxinoid venous system is still that of A. A.
RETZITJS, published in 1822.
The venous system varies very greatly in its details and even in certain aspects of
ON THE GENERAL MOEPHOLOGY OF THE MYXTNOID FISHES. 315
its general arrangement, and I select for description an example which exhibited the venous
system of Myxine in its most interesting and significant condition, and was investigated,
as it happened, with particular care. I have, however, confirmed the facts as described on
many other examples. I believe the Myxinoid venous system has undergone at least one
fundamental change in comparatively recent times, since individuals may from time to time
be found which depart from what may be regarded as the normal existing status in a
direction which can only be interpreted as ancestral.
The cerebral veins (figs. 2 and 6, cb.v.) arise ventro-laterally from the olfactory bulbs
as a pair of vessels, which then bend upwards and backwards on to the dorsal surface of
the brain, where they fuse at or near the middle line, the result further back becoming
the left efferent vessel of the brain. Apart from their origin, all the cerebral vessels are
confined to the dorsal surface of the brain. They are not symmetrical, the above vessel,
which we may call the left, draining the whole of the left half of the brain and also the
right side of the anterior region, the right posterior part of the brain having a special
vessel which, however, forms with the posterior section of the left a symmetrical pair of
vessels. Posteriorly the pair anastomose (fig. 2), but soon separate out again. They then
bend outwards, downwards, and backwards, and pierce the cranium by a special foramen
dorsal to the posterior region of the auditory capsule and posterior and dorsal to the vagus
foramen (indicated by.circles in fig. 2). They may now be described as the right and left
superficial anterior cardinal veins (r.s.c). The vessels at once pass outwards dorsal and
external to all skeletal parts and take up a position internal to the parietal muscle but
external to the constrictor pharyngis (fig. 7, l.a.c.s.). The superficial anterior cardinal veins
usually receive the segmental vessels of the region which they serve.
At about the transverse level of the eighth spinal ganglion the cardinal veins exhibit a
very curious arrangement which, however, I believe to be normal (fig. 2). The right super-
ficial cardinal (r.s.c.) perforates the constrictor pharyngis muscle and fuses with the right
deep anterior cardinal (r.d.c), the resulting vessel (r.d.c.') therefore passing backwards
internal to the constrictor pharyngis muscle. On the left side it is the deep cardinal which
perforates the muscle, so that the composite vessel (l.a.c.) courses external to the constrictor
pharyngis. Therefore the superficial vessel disappears posteriorly on the right side and
the deep vessel on the left, This anomalous state of affairs, which according to JACKSON
does not exist in Bdellostoma* continues for about eleven segments, i.e. up to about the
level of the anterior margin of the first gill. Consequently in this region we have vessels
which are not homologous performing the same functions. Each receives segmental vessels,
and also factors from the gut and the constrictor pharyngis, copulo-copularis, and parietal
muscles. At the level of the nineteenth spinal ganglion the right vessel swerves inwards
and becomes the median inferior jugular vein (i.j.v.).
Apart from the cerebral and cutaneous veins, no definite veins exist in front of the
auditory capsule, and the venous system of the snout is represented by the irregular so-
called veno-lymphatic spaces, which must therefore be regarded as an integral part of the
blood vascular system. The deep anterior cardinal vein (fig. 6, r.d.c.) commences as a large
sac, narrow from side to side but extensive dorso-ventrally, which is wedged in between
the hyoid arch and the velar portion of the gut. It receives its blood anteriorly by means
of a valved aperture (figs. 2 and 6, cl.h., h.v.s.') from the hypophysio-velar sinus, and some-
what further back there are other valves (fig. 2). The anterior extremity of the vein,
* In one Bdellostoma stouti dissected, the disposition of these vessels as regards the constrictor pharyngis muscle was exactly
as in Myxine.
therefore, is a large chamber guarded at each end by valves. This fact, and the disposition
of the valves, at once suggests that the chamber may act as a local heart, collecting the
blood from the large irregular spaces of the snout, where the blood-pressure may well be
a minus quantity, and driving it posteriorly towards the heart. 1 had already noticed
(Part II, pp. 686 and 721-2) that the velo-quadratus and velo-spinalis muscles closely resembled
the extrinsic muscles of the caudal heart, and thus differed from all other muscles of the
body, but was not then aware of the interesting condition of the vein. If a Myxine which
has been narcotised in warm sea water be dissected so as to expose the " hyoid" region,
the rhythmic action of the small-fibred velar muscles can be readily observed. The result
of this action is to squeeze the vein against the hyoid arch, which itself is kept stable
by the constricting action of the cranio-hyoideus muscle—a muscle the function of which
is otherwise difficult to explain (cf. Part II, PI. IV, fig. 11, c.h.). I therefore regard the enlarged
anterior extremity of the deep anterior cardinal vein as structurally and functionally similar
to the caudal heart, and have named it the cardinal heart (figs. 2 and 6, cl.h.).
Behind the cardinal heart the deep anterior cardinal vein passes outwards through the
fourth fenestra of the skull, i.e. internal to the first branchial arch and external to the
second (cf. Part III, fig. I,/*.4, br.a.1, br.a.2). The transverse level is indicated by the circles
in fig. 2 of the present communication. It now lies lateral to the gut and internal to the
constrictor pharyngis muscle (fig. 7, l.a.c.d.). Before fusing with the superficial cardinal
it may receive one or more ventral segmental vessels, and also factors from the rectus,
copulo-quadratus superficialis, copulo-glossus profundus, and constrictor pharyngis muscles.
To continue the description of the anterior cardinal system, the left anterior cardinal
is clearly nothing but the backward extension of the superficial cardinal, with which the
deep cardinal has fused (fig. 2, l.a.c). This vessel represents the anterior cardinal sinus
or jugular vein of Elasmobranchs. At first it lies external to the constrictor pharyngis
muscle, but soon bends downwards and takes up a position lateral to the copulo-copularis
muscle. It soon rises again and assumes its old position external to the constrictor pharyngis,
where it is to be found for the greater part of its course. Behind the latter muscle it
lies external to the dorsal anterior limb of the constrictor branchiarum et cardise muscle,
afterwards occupying a similar position in respect of the posterior external sheet of the
same muscle (fig. 8, l.a.c). By this time it has increased very considerably in size, and
finally, in the neighbourhood of the ductus oesophago-cutaneus, it perforates the constrictor
branchiarum muscle and fuses with the corresponding vessel of the right side by means
of the extensive and somewhat irregular cardinal anastomosis (fig. 2, <?.«.), which passes
between the systemic aorta and the dorsal surface of the gut. The position of this
anastomosis excludes the possibility of its forming any part of the systemic heart. It is
surprising that no previous observer has seen the anastomosis, the existence of which con-
siderably modifies the course of the circulation, and is possibly associated with the loss of
the right Cuvierian duct. If it were absent, for example, the blood in the right anterior
cardinal could only reach the heart by passing through the liver. The anastomosis is easily
found both in Myxine and Bdellostoma. At the point where the anterior cardinal joins
with the anastomosis there is a valved aperture on each side, by which the sixth peri-
branchial sinuses discharge their contents into the venous system (fig. 2, p.b.s.'). Posteriorly
the left anterior cardinal received a vessel formed by the union of the dorsal and ventral
divisions of the anterior section of the left posterior cardinal (fig. 2, l.a.c.', l.a.c"). The
left anterior cardinal receives the segmental veins of its region, and also vessels from the
gut and the constrictor pharyngis, copulo-copularis, and constrictor branchiarum et cardise
muscles. JACKSON described the left anterior cardinal of Bdellostoma as opening directly
into the sinus venosus. This is certainly not the case in Myxine nor was it the case in
one Bdellostoma which was dissected with special reference to this point.
On the right side the vessel which passes backwards and corresponds functionally with
the left anterior cardinal is obviously the direct continuation of the right deep cardinal
vein and is therefore not homologous with the vessel of the left side (fig. 2, r.d.c.') After
receiving the right superficial cardinal it courses backwards lateral to the gut and internal
to the constrictor pharyngis muscle. At the posterior end of the club muscle it passes
outwards and downwards under the constrictor pharyngis, and curving ventrally round the
club muscle it soon reaches the neighbourhood of the mid-ventral line, as first described
by A. A. RETZITTS. At this point the contents of the lingual sinus are discharged into it
by a single aperture with a double valve (fig. 2, l.s.'). From this point backwards the vessel
is known as the median inferior jugular vein (figs. 2 and 8, i.j.v.). It is situated internal
to the muscles of the body wall and the constrictor branchiarum et cardise somewhat to
the right side of the median plane, and is closely opposed to the ventral wall of the cardiac
sinus. Further back, owing to the crowding together of the surrounding organs, it becomes
small. In the region of the ductus oesophago-cutaneus it crosses over to the left side,
becomes greatly enlarged but retains its relations to the surrounding muscles, and passes
imperceptibly into the sinus venosus. During the lateral part of its course the vessel
receives the segmental veins of its region, and also factors from the constrictor pharyngis
and copulo-copularis muscles. I did not detect any important vessels opening into it during
the median part of its course. The inferior jugular vein may be formed by the fusion of
the deep cardinal of both sides, the anterior factor disappearing on the left side. Its
formation and variations suggest this. A small vessel is sometimes received from the
region of the first gill pouch of the left side which may represent the missing left
factor.
A vessel which is undoubtedly the morphological equivalent of the left anterior cardinal,
having precisely the same relations to the surrounding organs, arises far back at the level
of the anterior margin of the first gill, i.e. at the point where the right inferior jugular
swerves towards the mid-ventral line. It is constituted by the union of the dorsal and
ventral factors of a typical segmental vein, and is much smaller than the vessel of the left
side. It is, however, a typical right anterior cardinal (figs. 2 and 8, r.a.c), and represents
the posterior extremity of the right superficial cardinal, which on this side is interrupted
for about eleven segments (seventh to eighteenth spinal). It is a vessel named by JACKSON
the anterior portal vein on account of its posterior connection with the portal heart, but
JACKSON failed to recognise the identity of this vessel with the anterior cardinal. Posteriorly
it communicates with the left cardinal by the cardinal anastomosis (fig. 2, c.a.) as already
described. Immediately posterior to the anastomosis it receives a large vessel formed by
a factor from the portal heart fusing with the dorsal division of the anterior section of
the right posterior cardinal (fig. 2, r.a.c.'). The right anterior cardinal receives a few
segmental vessels but no others of any importance. The extraordinary fact that the right
anterior cardinal communicates with the portal heart was first observed by A. A. RETZIUS.
The caudal veins are constituted at the tip of the tail by capillary systems which
place them in direct communication with the paired caudal arteries. The vein is also
paired, the sagittal plane being occupied by the median ventral bar of the caudal fin
skeleton (figs. 2 and 11, c.v.', m.v.b.). I have seen three cases in which there was more than
one caudal vein on each side. The caudal veins pass forward and expand to form the
paired caudal hearts, of which they are the afferent vessels. Anterior to the median ventral bar
the efferent vessels of the caudal heart (fig. 2, c.v.") unite to form the median caudal vein
(c.v.), and at this point the valves are situated which prevent any posterior movement of the
blood-stream (figs. 2 and l l ) .
The posterior cardinal veins (fig. 2, r.p.c, l.p.c) are formed by the bifurcation of the
median caudal vein anterior to the cloaca. The left posterior cardinal is much larger
than the right, and the two are connected by transverse anastomoses which may be
simple or ampulliform and pass ventral to the aorta. The posterior cardinals lie
ventral to the systemic aorta, which itself is ventral to the lateral chordal sinuses. The
latter are situated immediately below the notochord. There is no renal portal system, but
the pronephros is very closely associated with (presumably) the posterior cardinal system.
In the neighbourhood of the 31st spinal segment and the posterior extremity of the
anterior lobe of the liver, the right posterior cardinal effects its last anastomosis with the
left, and the single vessel anterior to this point is known as the common posterior
cardinal vein (fig. 2, c.p.c). The fact, however, that in many, if not most examples, the
right posterior cardinal is actually continued forwards in front of this region (fig. 2, r.p.c.')
shows that the so-called common posterior cardinal is simply the anterior extension of the
left vessel. This forward continuation of the right posterior cardinal was evidently seen
(but misinterpreted) by J. MULLER, who regarded it as the efferent vessel of the right
pronephros. The common posterior cardinal now bends to the left, and passes inperceptibly
into the sinus venosus. It is not clear, judging from adult conditions, where one ends
and the other begins, and it is therefore difficult to be quite certain whether the hepatic
veins open into the sinus venosus or the common posterior cardinal, or both. Several
vessels from the gut wall (avoiding the portal system),* and a few segmentals, open into
the common posterior cardinal + sinus venosus, and isolated kidney glomeruli may occur
between the pronephros and the definitive kidney which drain into the common posterior
cardinal. I have also found efferent vessels from the pronephros which pass backwards
to open into the common posterior cardinal on the left side and the right posterior
cardinal on the right. The posterior cardinals receive only segmental and renal veins, the
genitals opening into the portal vein. It is unusual for the same segment to have both
an artery and a vein. There is no regularity or symmetry in the arrangement, but the
segmental arteries are generally more numerous than the veins.
The posterior hepatic or subintestinal vein (fig. 2, s.i.v.) arises on the ventral surface
of the gut. It may extend over the whole length of the gut as it does in the Ammoccete,
but generally it is confined to a short region behind the posterior lobe of the liver. On
reaching the posterior lobe it may divide into two large vessels—one coursing on the
concave dorsal surface, and the other, the main trunk of the vessel, on the convex ventral
surface of the posterior liver. The former is dissolved in the liver, and in my Part V I
concluded that it was an afferent hepatic vein, and that therefore the subintestinal acted
both as an afferent and an efferent vessel of the posterior lobe of the liver. This, of course,
is suggested by a knowledge of its development in other types. There are, however, no
valves anywhere in the subintestinal vein, so that the action of the portal heart must
inevitably drive the blood into the subintestinal, in spite of the fact that in the case of
the dorsal factor the blood would have to pass backwards in order to reach the main trunk
of the subintestinal. We must therefore conclude that the subintestinal is an efferent
hepatic vessel only. Assuming this to be true, as it is in Bdellostoma according to
* Both A. A. RBTZIUS and J. MULLER note that a vein from the "stomach" opens into the left posterior cardinal (see fig. 2).
JACKSON, the subintestinal cannot be regarded as an extra portal vein, since it does not
distribute any of its blood to the liver. The main trunk of the vessel is always more or
less superficial in position and of a fairly uniform calibre throughout, and as it courses over
the posterior lobe it receives from it a few large efferent vessels. The subintestinal is
always the principal, and may be the sole, hepatic vein from the posterior lobe. An
accessory posterior hepatic vein opening separately into the sinus venosus is however
usually present, and forms an anastomosis with the hepatic vein from the anterior lobe of
the liver. The subintestinal differs from all the other hepatic veins in retaining its con-
nection with the gut, but the extent to which it does so varies considerably. It opens
apparently into the sinus venosus. The subintestinal was described in Myxine by
KLINCKOWSTROM in 1890, who mentions it as extending back to the cloaca in one specimen.
It was afterwards independently found by JACKSON in Bdellostoma.
The anterior hepatic vein (fig. 2, a.h.) receives a number of efferent vessels from the
anterior lobe of the liver, and may also include a factor from the posterior lobe and an anas-
tomosis from the accessory posterior hepatic vein when present. It opens into the sinus
venosus. In addition there may be as many as ten large and small accessory anterior
hepatic veins, each having a separate opening into the sinus venosus.
At the point where the common posterior cardinal passes into the sinus venosus it
receives two vessels, which are fused in front and arise from the junction of the left
anterior cardinal with the cardinal anastomosis (figs. 2 and 9, l.a.c.', La.c."). These vessels I
have named the dorsal and ventral divisions of the anterior section of the left posterior
cardinal. Whether they in fact belong to the posterior or anterior cardinal system, or to
both, can only be determined when we know their development. They may conceivably
belong to the anterior cardinal, in which case a ductus Cuvieri is formed on the left side
only by the union of anterior and posterior cardinals. The figures which A. A. RETZIUS
gives of this region, if somewhat crude, agree essentially with my fig. 2. There are no
valves associated with the dorsal and ventral divisions, but presumably their contents will
flow backwards into the common posterior cardinal owing to the action of the cardinal
heart. Both vessels, and the fused portion, project into the dorso-median section of the
pericardial coelom (fig. 9, shown in black), and are partly surrounded by it. The dorsal one
of the two (La.c.') is the larger, and is wrapped round the pronephros which lies apparently
inside it. This member of the pair, and also the fused anterior part, may receive veins
from the gut and the constrictor branchiarum et cardiae muscle, and also segmental veins.
It is curious that in none of the many detailed memoirs on the Myxinoid pronephros which
have appeared in recent years is the true relation of the pronephros to the venous system
recognised. MAAS, for example, simply states that the pronephros is enclosed by a venous
sinus. KIRKALDY, it is true, identifies the sinus as the " post-cardinal vein," but says
nothing of the connections of this vessel.
On the right side the conditions are less different than they appear at first sight.
There is a vessel corresponding exactly to the dorsal division of the left posterior cardinal
as regards its topographical position and relations to the pronephros (figs. 2 and 9, r.a.c.').
Variations of this vessel establish its identity beyond question, since in many, if not most
examples, it develops posteriorly a definite anastomosis with the right posterior cardinal at
the point where the latter fuses with the left. This anastomosis can only represent the
forward extension of the right posterior cardinal (fig. 2, r.p.c.'), the corresponding vessel
of the left side being the so-called common posterior cardinal (c.p.c). In three dissections
the anastomosis had no connection with the portal vein behind the portal heart, and was
the direct continuation forwards of the right posterior cardinal.* The dorsal vessel there-
fore must be homologous with the dorsal division of the anterior section of the posterior
cardinal of the other side. It may, however, terminate posteriorly by passing into the
portal vein posterior to the portal heart, and this posterior section may be very small.
Anteriorly it fuses with the portal heart to form an unpaired section which opens into the
junction of the right anterior cardinal with the cardinal anastomosis, and thus behaves
exactly as the corresponding vessel of the left side (cf. fig. 2).
The portal or supra-intestinal vein of Myxinoids lies dorsal to the gut and to the
right of the median plane, and acts not only as the efferent vessel of the abdominal
section of the gut, but also of the gonad (cf. my Part II, PI. I, p.v.). These facts indicate
that it is not comparable with the portal vein of the true fishes, which develops from the
sub-intestinal vein of the embryo and has no association with the gonad. In Ceratodus,
however, according to KELLICOTT, the hepatic portal of the adult is derived from an
anterior new formation dorsal to the gut and the persisting posterior section of the sub-
intestinal vein. The portal vein of the Myxinoids therefore may represent the anterior
portion of Ceratodus. An examination of fig. 9 shows that the portal heart, as regards
its topographical relations, must represent the ventral division of the anterior section of
the posterior cardinal of its side (figs. 2 and 9, cf. p.h., l.a.c"). The course of the
afferent and efferent portal veins (p.v., cp.v.) suggests that the portal heart is an addition
to the system. The heart differs from the other accessory hearts in having an intrinsic
musculature, and is further larger and more powerful. The three openings from the right
anterior cardinal, portal vein, and common portal vein are all guarded by effective valves,
which ensure that blood passes into the heart from the first two vessels and away from it
by the third. A cystic vein (or veins) is constituted on the surface of the gall bladder
and opens into the portal vein (fig. 2, cys.v.). In one Bdellostoma stouti dissected I found
that the relations of the great veins to each other and to the heart, although differing in
certain details, were essentially the same as in Myxine. i t seems therefore that the accepted
description of the venous system of Bdellostoma might be revised with advantage.
The portal heart was first discovered by A. A. RETZIUS in Myxine in 1822. He then
described it as a spongy sac very similar in appearance to the auricle of the heart. He
noted that the right anterior cardinal vein passed into it, and also that this connection
was paralleled by the union of the anterior cardinal vein and pronephric vein of the left
side (cf. fig. 2). He discovered the pronephros, and suggested that it might be a "repre-
sentative of the kidney." He is extremely surprised that the pronephric vein should open
into the portal system on the right side and the posterior cardinal on the left. This
difference, however, as we have seen, is more apparent than real. When RETZIUS' paper was
translated into German in Isis in 1825, the translator notes that he had received a letter
from the author describing the presence of muscle fibres in the portal heart, and stating that
he had watched its pulsation in a living animal. J. MULLER appears to have overlooked this
note, for in 1841 he denied that RETZIUS' "long sac" was a portal heart, holding that it had
no valves or musculature and was capable only of elastic resistance. In 1845, however, he
corrects this statement, having observed the rhythmic pulsation of the portal heart in 1841,
but he still says nothing of the striated musculature which the heart undoubtedly possesses.
The interpretation of this unusual type of circulation can only be properly attempted
* J. MDLLER appears to have seen the vessels lettered in fig. 2 as r.p.c' and r.a.c.', but his description of them is diffi-
cult to follow. JACKSON found a vein passing from the right pronephros to the posterior cardinal in one specimen of
Bdellostoma.
when complete information as to its development is available. In the meantime, however,

a few comments may be permitted. The controlling factor in the situation appears to be
the position of the systematic heart—the sinus venosus and auricle being placed well on
the left side of the body (fig. 9, s.v., aur.). The presence of the ductus oesophago-cutaneus,
which occurs only on the left side, must, at least to some extent, affect the position of
the heart so as to produce a certain degree of asymmetry. The asymmetry of the heart
may explain the curious right-to-left swerve of the so-called inferior jugular vein
and the occasional disappearance of the right posterior cardinal in front. It is evident,
however, in the latter case, that there has been only a separation of the anterior and
posterior sections of the right posterior cardinal, since it must be concluded that the
anterior section on the right side is represented by the portal heart and the dorsal vein
lettered r.a.c' Such an assumption, which explains the connection of the portal heart
with the anterior cardinal system, is based on the state of affairs on the left side where
the two corresponding vessels l.a.c' and l.a.c." constitute the anterior section of the left
posterior cardinal, and also on the connection between the right posterior cardinal and
the vessels of the right side (fig., 2, r.p.c.'). On any interpretation, however, the course
of these vessels (p.h., r.a.c.', l.a.c.', l.a.c") is anomalous in the adult Myxine. If they
belong to the posterior cardinal system they extend too far forwards, and if to the
anterior cardinal system too far backwards. On the latter hypothesis the anterior and
posterior cardinals fuse to form a forwardly coursing ductus Cuvieri, which is absent on
the right side. The view outlined above assumes that the new section of the venous
system is represented by the obliquely coursing stretch of the inferior jugular (i.e. from
the most anterior gill to the sinus venosus), together with the sinus itself. It seems
probable that the vein associated with the pronephros belongs to the posterior cardinal
system, although according to HATTA the pronephric veins of the Ammoccete are
derived in younger larvae from the posterior extremity of the anterior cardinal, and in
older larvae from the ductus Cuvieri also. It is to be noted further that the circulation
of the blood in the pronephric system differs on the two sides owing to the interpolation
of the hepatic portal system on the right side. The evolution of the anterior cardinal
system is more obvious as regards the sequence of events, but still puzzling to explain.
It is surprising that neither anterior cardinal has a direct connection with the heart, and
that the blood in these vessels can only reach the sinus venosus by devious routes,
in one of which it may even have to pass through the liver before reaching the heart,
but this might follow from the shifting of the sinus to the left side. It is also not
clear why the course of the deep cardinal vein should be shortened on the left side and
why the superficial cardinal should be interrupted on the right (represented in fig. 2 by
the dotted line), for the right anterior cardinal undoubtedly represents the posterior
continuation of the right superficial cardinal. The absence of definite veins in front
of the auditory capsule, and the consequent establishment of a lacunar circulation in the
snout through the so-called lymphatic spaces, is commented on elsewhere.
In his recent important paper on the vascular system of the Ammoccete (Zool. Jahrb.
44, 1922), HATTA questions whether the inferior jugular of Myxinoids is homologous with
the jugularis impar of the Ammoccete, and suggests that the Myxine vessel is a derivative
of the right deep cardinal only. This is obviously partly if not wholly true. In younger
stages the two ductus Cuvieri of the Ammoccete are almost equally well developed, but
the left then becomes weaker and is obliterated in the fully developed larva. In the
Myxinoid the right ductus Cuvieri is stated to be absent in the adult, but if my inter-
pretation of the cardinal system be accepted both are absent. According to HATTA, the
anterior cardinal system of the Ammocoste is also anomalous, but not in the same way as
in Myxine. He concludes from topograpical relations and development that the anterior
cardinal vein terminates just behind the second gill arch, and that- the section of the
vein posterior to this arch is formed secondarily and does not correspond to a true
cardinal vein. This posterior part is an extension of the mandibular vein which becomes
secondarily connected with the anterior cardinal. In another respect Myxine differs from
the Ammoccete. Veins are constituted anteriorly on the tip of the snout in the usual
way, but (and in this they differ from the arteries) they are originally post-otic vessels
which afterwards extend headwards. The true head veins are thus reduced to vessels
which belong to the posterior region of the head. The Ammoccete therefore represents a
later stage in the evolution of the venous system than the adult Myxinoid.
The lacunar species of the Myxinoid, which are all lined by a tenuous flattened
endothelium, are, for reasons to be stated later, divisible into two series, which I propose
to call the red lymphatics and the white lymphatics. The former always contain red
blood in more or less greater quantity, and may be regarded provisonally as a part
of the blood vascular system interposed between arteries and veins. The red lymphatics
are grouped further into two cycles—the subcutaneous cycle and the peribranchial cycle.
The white lymphatics contain a white lymph only, which is discharged into the venous
system. They are, moreover, not a part of the blood vascular system, and therefore
do not contain red blood. They fall into a definite system by themselves. This is
schematised in fig. 3, which illustrates the relations of all the white lymphatics except
the pairs of segmental lymphatics. The white lymphatics may be considered, also
provisionally, as constituting a true lymphatic system.
E. RED LYMPHATICS. SUBCUTANEOUS CYCLE.
1. Subcutaneous Sinus (Figs. 4-11, s.c.s.).

This is by far the largest of the vascular spaces in Myxine. That it contains red
blood in the living animal is obvious from the fact that the red colour of the contents
of the sinus can be seen through the skin. The sinus consists of three portions situated
immediately under the skin—two lateral and one ventral. The lateral portions are
separated at the mid-dorsal line by the strand of tissue or septum which transmits the
nerves and blood-vessels to and from the skin {cf. Part II, PL I). Laterally they are
continued downwards to the row of slime sacs, and longitudinally they extend from the
nasal tentacles to the tip of the tail. Anteriorly on the head the dorsal septum is
wanting, so that the lateral portions are here in open communication (fig. 4). Between
the lateral and ventral portions in the neighbourhood of the ducts of the slime glands is
a kind of coarse plexus which places the lateral and ventral portions in communication
with each other. In front of the region of the slime sacs the latter communication
becomes an open one. The ventral portion of the sinus lies between the two rows
of slime sacs. It is comparatively small, not well defined, and much broken up by
strands of connective tissue. Almost the whole body, therefore, is surrounded by a
voluminous vascular space.
The connection between the subcutaneous sinus and the caudal hearts may now
be described {cf. figs. 2, 10 and 11). The caudal part of the sinus (s.c.s.) is connected
with a paired coarse plexus confined to the caudal fin (fig. l l ) , into which the contents
of the sinus flow quite freely. This plexus, which is easily injected from the subcutaneous
sinus, and also the sinus itself (fig. 10), is drained by a marginal vessel (s.c.s.1), which,
starting on the dorsal side, passes backwards round the tip of the tail and then forwards
to open by a valved aperture into the antero-ventral angle of the hearts. Valves prevent
any regurgitation from the hearts to the afferent vessels. At the same place the lateral
chordal sinus also discharges into the heart (l.c.s/). The marginal vessel is situated
at the roots of the fin rays, and may be regarded either as a single vessel which bifur-
cates to surround the base of every fin ray, or as a double vessel which fuses between
the fin rays (cp. fig. 10). It does not completely split to open into the paired caudal
hearts, but sends an extension upwards for this purpose on each side of the median
ventral bar of the caudal fin skeleton, at the ventral edge of which it is situated during
the whole of its forward course. In front of the opening into the caudal heart the marginal
vessel may be continued forwards for a short distance. In addition to the above,
ALLEN describes a lateral lymphatic which passes dorsal to the series of slime sacs
anterior to the caudal hearts. I have frequently found this vessel in Myxine also.
KLINCKOWSTROM, in a brief note (1891), mentions the plexus on the caudal fin and the
marginal 'vessel, which he regards as paired, and states that it opens into the caudal
hearts. He missed, however, the connection with the lateral chordal sinus. FAVARO
describes the caudal vein (c.v.') as arising by the splitting of the ventral marginal vessel
(s.c.s.'). I doubt whether this ever occurs, especially as these vessels are related in such
a way that it is easy to understand how such an error could arise. FAVARO'S account of
the caudal circulation in Myxine is otherwise incomplete.
Connections—direct.—1. Hypophysio-velar sinus. 2. Rostral sinus. 3. Dorsal oral
sinus. 4. Ventral sinus. 5. Subcutaneous anastomosis. 6. Segmental lymphatics.
Connections—indirect.—7. Hypophysio-velar sinus, ventral sinus, and lateral chordal
sinus—all via the subcutaneous anastomosis. 8. Lateral chordal sinus via the segmental
lymphatics. 9. Caudal vein and posterior cardinals via the caudal hearts.
2. Subcutaneous Anastomosis (Figs. 3 and 6, s.c.a., s.c.a.'~'").

This opens into the subcutaneous sinus by a well-marked aperture bounded by the
ventro-posterior extremity of the tentacularis posterior muscle and the second myotome—
forming a notch in the latter muscle (cf. Part II, PL IV, figs. 8 and 9). The channel
connected with this aperture is at first large, and passes inwards and backwards dorsal
and lateral to the fused trabecular and pterygo-quadrate cartilages. It then travels
obliquely through the second fenestra of the skull (cf. fig. 6) so as to communicate with
that section of the hypophysio-velar sinus situated in the neighbourhood of the third
fenestra of the skull (figs. 3 and 6, s.c.a.'). There appears to be a valve at this point,
and if so its function is not clear, as the flow would naturally be an outward one, i.e.
from the hypophysio-velar sinus into the subcutaneous sinus, the pressure in the latter
sinus being the lower of the two. The anastomosis continues to pass backwards, and in
the region of the auditory capsule splits into two sections—a dorsal one which passes
internal, and a ventral one which passes external to the origin of the cranio-hyoideus
muscle. The dorsal section (s.c.a.'") takes up a lateral position in the skeletogenous layer
of the sheath of the notochord, and, as confirmed by injections, fuses with the origin of
the lateral chordal sinus. The ventral section (s.c.a.") opens into the latero-dorsal
extension of the ventral sinus.
Connections.—1. Subcutaneous sinus. 2. Hypophysio-velar sinus. 3. Ventral sinus.

4. Lateral chordal sinus.
3. Rostral Sinus (Fig. 4, rs.s.).
Extends backwards from near the tip of the snout to the level of the base of the
median dorsal tooth. It is a paired sinus closely associated with the inner surface of
the tentacularis posterior muscle, and also in front with the posterior external surfaces of
the tentaculo-ethmoidalis and transversus oris muscles. Anteriorly it merges gradually
into the subcutaneous sinus, with which further back it has several additional and varied
connections. It communicates also with the median portion of the hypophysio-velar sinus
lying below the nasal tube. The posterior extremity of the sinus is blind.
Connections.—1. Subcutaneous sinus. 2. Hypophysio-velar sinus.
4. Dorsal Oral Sinus (Fig. 4, d.o.s.).

A short median sinus lying immediately over the gut and extending from the mouth
to the transverse level of the tip of the subnasal bar. In front it merges gradually into
the subcutaneous sinus, of which it is the posterior extension produced by the invagina-
tion of the skin to form the lining of the mouth. Further back it despatches outwards
a blind lateral wing which is closely associated with the palato-ethmoidalis profundus
muscle. Behind this region it communicates with a diverticulum from the hypophysio-
velar sinus, which arises from the latter sinus just behind the posterior extremity of the
subnasal bar and passes forwards under the bar to fuse with the dorsal oral sinus.
Posteriorly the sinus bifurcates owing to the intervention of the median dorsal tooth, and
terminates blindly on each side shortly afterwards.
Connections.—1. Subcutaneous sinus. 2. Hypophysio-velar sinus.
5. Hypophysio-velar Sinus (Figs. 4-7, h.v.s., h.v.s.', h.v.s.").

This is an extensive and irregular sinus associated with the following three structures :—
(l) Nasal tube and nasal organ. The main part of this section of the sinus is situated
below the nasal tube external to the nasal skeleton, but in places it may surround the
tube (fig. 4), here lying internal to the skeleton. In front it is in wide communication with
the subcutaneous sinus and the rostral sinus. Below, and posterior to the hinder
extremity of the subnasal bar, it receives a backwardly coursing anastomosis from the
dorsal oral sinus (fig. 4), which courses below the subnasal bar and is wedged between
the two palato-ethmoidalis profundus muscles. Factors are received from the olfactory
laminae of the nose (fig. 5—median lamina, unlettered). (2) Behind the olfactory organ
the sinus is paired, and is situated immediately lateral to the naso-pharyngeal tube
(fig 6). Further back it establishes a connection with the subhypophysial sinus, and
receives important factors which surround the velo-spinalis and the whole of the velo-
quadratus muscles and also penetrate between the divisions and bundles of the latter
muscle (fig. 6). It is this factor which receives dorso-laterally, via the third fenestra of the
skull, a vessel from the subcutaneous anastomosis which connects the hypophysio-velar
sinus with the subcutaneous sinus (figs. 3 and 6, s.c.a.'). The factor also communicates
by means of a large valved aperture with the expanded blind venous sinus or cardinal
heart, which, in the absence of a capillary system, forms the origin of each deep anterior
cardinal vein (figs. 2 and 6, h.v.s.', cl.h.). The disposition of the valves, and numerous
injection experiments, demonstrate that the contents of the sinus can pass into the veins
ON THE GENERAL MORPHOLOGY OP THE MYXINOID FISHES. 325
but not vice versa. (3) Behind the velo-spinalis muscle the remainder of the sinus is
constituted within the velum or pharyngeal valve by a coarse but extensive plexus, which
forms the velar lymphatic system and arises blindly at the posterior extremity of the
valve. A special section of this plexus lies alongside the lymphoid organ of the velum
(? thymus) which is associated with the anterior half of the external lateral velar bar.
Red blood reaches the velar section of the sinus from the capillaries of a median artery,
which arises from the point where the right and left factors of the vertebralis impar
capitis fuse to form the median portion (fig. 1, vel.a.).
Connections.—1. Subcutaneous sinus (direct connections anteriorly). 2. Rostral sinus.
3. Dorsal oral sinus. 4. Subhypophysial sinus. 5. Subcutaneous sinus, ventral sinus, and
lateral chordal sinus (indirectly via the subcutaneous anastomosis). 6. Deep anterior
cardinal veins.
6. Subhypophysial Sinus (Figs. 5, 6, s.h.s.).

This is the sinus of the sheet of tissue, stiffened by the hypophysial plate and having
a free posterior border, which separates the naso-pharyngeal tube from the mouth region
(cf. Part V, PI. I, fig. l). It commences blindly behind at the free border mentioned
above, and passes forwards under the hypophysial plate moulding itself to the V shape
of the broad section of the cartilage. Dorsally it communicates with the sinus immediately
above it (hypophysio-velar sinus), and laterally on each side with the latero-dorsal extension
of the dental sinus (fig. 5, d.s.'). In the region of the nasal chamber the sinus becomes
more centralised and its cavity increases considerably in area (fig. 5). Anteriorly it
narrows and flattens again, and terminates blindly underneath the palatine commissure
between the latter and the base of the median dorsal tooth.
Connections.—1. Hypophysio-velar sinus. 2. Dental sinus (latero-dorsal extensions of).
7. Ventral Sinus (Figs. 5, 6, 7, vn.s.).

Arises blindly posteriorly at the mid-ventral line immediately below the posterior
segment of the basal plate. It soon expands laterally up the sides of the cartilage, and
is now a wide, shallow, U-shaped sinus wedged in between the basal plate and the median
head of the copulo-glossus profundus muscle. This constitutes the median section of the
sinus. Just behind the region of the second branchial arch a dorso-lateral extension of the
sinus arises blindly, and passes upwards internal to the parietal muscle until it reaches
and surrounds the anterior extremity of the superior lateral cartilage, at which point it
fuses with the ventral section of the subcutaneous anastomosis, and thus establishes an
indirect connection with the subcutaneous, hypophysio-velar, and lateral chordal sinuses
(fig. 6). The dorso-lateral extension passes forwards and downwards internal to the
parietal and obliquus muscles, and fuses with the median section near the junction of the
middle and posterior segments of the basal plate. The latter section now extends laterally
and ventrally so as to enclose completely the copulo-glossus superficialis and the whole of
the copulo-glossus profundus muscles of both sides for the remainder of their course
forwards (fig. 6). Hence the tendons of these muscles also come to lie apparently in the
cavity of the ventral sinus. The sinus now fuses dorsally and indistinguishably with the
dental sinus, and finally splits into two, each half becoming continuous laterally with the
subcutaneous sinus just behind the mouth.
Connections.—1. Lateral chordal sinus, hypophysio-velar sinus and subcutaneous sinus via
$26 PROFESSOR FRANK J. COLE
the subcutaneous anastomosis. 2. Subcutaneous sinus (direct wide communication in front).

3. Dental sinus.
8. Dental Sinus (Figs. 5, 6, d.s., d.s.').
A large and somewhat indefinite sinus closely associated with the wall of the gut in
the region of the tongue. Anteriorly it fuses with the ventral sinus immediately in front
of the anterior extremity of the internal bar of the anterior segment of the basal plate.
The basal plate in fact separates the dental and ventral sinuses throughout. On each
side the sinus despatches upwards into the tongue two long narrow extensions, separated
by a narrow strip of tissue, in which the dental plate is situated. In the region, there-
fore, where the dental plate extends across the mid-ventral line, the dental sinus is for
the time being divided into two portions—an inner one internal to the dental plate,
concerned with the vascular supply of the greater part of the tongue and teeth, and an
outer one associated with the more external parts of the latero-ventral wall of the gut
(cf. fig. 5). The inner section, however, is in free communication with the outer at both
extremities. The dental sinus possesses also a latero-dorsal extension, associated with the
outer wall of the gut, which passes on to the dorsal surface of the latter and completes
the circle by a definite fusion on both sides with the median dorsal subhypophysial sinus
(fig. 5, d.s.'). Posteriorly the dental sinus fuses extensively with the lingual sinus, and
is also connected with the factors of the latter sinus which accompany the paired tendons
of the longitudinalis linguae muscle (fig. 6). Behind the fusion with the lingual sinus the
dental sinus fuses with and becomes indistinguishable from the anterior suboesophageal
sinus. Blood reaches the dental sinus via the capillaries of the anterior and posterior lingual
arteries (fig. 1. La., La.'), both arising from the external carotid. The former passes at
first outwards, forwards, and downwards between the internal and external heads of the
copulo-ethmoidalis muscle. It then bends sharply inwards under the anterior extremity
of the ventral sinus on to the tongue, in which it courses backwards. The latter reaches
the tongue via the "tether" of the hyo-copulo-glossus muscle.
Connections.—1. Ventral sinus. 2. Subhypophysial sinus. 3. Lingual sinus. 4. Sub-
cesophageal sinus (anterior).
9. Lingual Sinus (Figs. 2, 6, 7, Ls., Ls.').

This is the sinus of the club-shaped muscle. Anteriorly it fuses with the dental
sinus (fig. 6), and is connected with the suboesophageal sinus. It then passes backwards,
at first partly, but soon entirely, surrounding the complex tendon of the longitudinalis
linguae muscle. For this tendon compare Part II, p. 713, and PL III. Before surrounding
this tendon, however, it receives a pair of factors which are closely associated with the
paired central portions of the above tendon (fig. 6). These factors originate blindly as a
single vessel on the dorsal surface of the anterior extremity of the central unsplit
tendon of the muscle. When the tendon splits into two the lymphatic also becomes paired.
The contents of this section of the sinus therefore must pass forwards to discharge into
the main portion of the lingual sinus. The latter now passes backwards, completely
surrounding the outer or tubular portion of the tendon of the longitudinalis linguse (fig. 7),
and later the belly of the muscle itself. For the greater part of its course, therefore, the
sinus has a tubular structure, and is in its turn surrounded by the copulo-copularis
muscle except at the posterior extremity of the club muscle, where the longitudinalis
linguae extends beyond the copulo-copularis. As it approaches the posterior extremity of
the latter muscle a break appears first dorsally and then ventrally, thus dividing it
into two portions. These, however, at the extreme posterior end of the club muscle join
up again, after which the sinus rapidly diminishes in bulk, curves ventrally to the right
of the cardiac aorta but to the left of the inferior chondroidal bar, bends to the left of and
below the tip of the perpendicular muscle, and opens by a well-marked valved aperture into
the inferior jugular vein (fig. 2). The passage between the vein and the sinus in an
average adult forms a vessel about 5 mm. long. The structure of the valve and the
results of numerous injection experiments establish conclusively that the contents of
the sinus can pass into the vein but not vice versa.
Connections.—1. Dental sinus. 2. Subcesophageal sinus. 3. Inferior jugular vein.
10. Subcesophageal Sinus (Figs. 7, 8, s.o.s., s.o.s.').

This was first described by J. MULLER in 1836. He says it is "probably a lymph sac,"
and describes its connection with the subchordal sinus based on inflations.
The suboesophageal sinus is in two sections—an anterior and a posterior, which should
perhaps be regarded as two distinct sinuses. The anterior section commences as an irregular
very coarse plexus on the ventral wall of the gut in the region of the third gill. In front of
the gill region, however, it forms a large, wide, but shallow sinus covering the entire ven-
tral wall of the gut (fig. 7). As it terminates blindly behind its contents must course
forwards. There are several wide communications at the side of the gut between the
anterior section of this sinus and the subchordal sinuses. After a long course the sinus
narrows down anteriorly, and, after establishing several connections with the lingual sinus,
it fuses with the posterior extremity of the dental sinus just posterior to the hyoid arch.
The posterior section of the sinus is much shorter and less extensive, and is separated
from the anterior section by a distinct interval. Its anterior extremity is very indefinite,
and is situated in the region of the fifth gill. As it passes backwards, it expands so as to
cover more or less the ventro-lateral wall of the gut. At this point there are connections
at the side of the gut with the subchordal sinus (fig. 8). Posteriorly the section breaks
up and dies away at about the level of the external aperture of the ductus oesophago-
cutaneus. Both extremities of the section being blind, its contents can only escape via
the subchordal sinus, and its included blood must also come from the subchordal.
Connections.—1. Dental sinus. 2. Lingual sinus (anterior section of subcesophageal).
3. Subchordal sinus (both sections).
11. Carotid Sinus.
A short inconspicuous but clearly defined sinus, blind at each extremity, associated
with the external carotid artery throughout the region corresponding to the posterior half
of the nasal tube. No connections could be traced.
F. RED LYMPHATICS. PERIBRANCHIAL CYCLE.
1. Cardiac Sinus (Figs. 8, 9, cd.s.).

Described by J. MULLER in 1836, who traced its connection with the peribranchial
sinuses, and its extension on to the ventricle of the heart between the wall of the
ventricle and the pericardial cavity (fig. 9).
The cardiac sinus is the sinus of the ventricle of the heart and the cardiac aorta. It
arises blindly on the ventricle as an elaborate plexus of coarse vessels which ramify and
anastomose over the surface of the ventricle. This plexus abuts on the sinus venosus
and auricle, and is itself surrounded on all sides except the median sagittal by the
pericardial ccelom (cf. fig. 9). In front the plexus fuses to form a conspicuous sinus
which completely surrounds the cardiac aorta throughout the whole of its length (fig. 8).
On each side the latter part of the sinus gives off wide channels which transmit the
afferent branchial arteries and place it in communication with all the peribranchial
sinuses. The first pair of these channels to the first pair of peribranchial sinuses is
formed by the bifurcation of the anterior extremity of the cardiac sinus. The portion
of the sinus surrounding the ventricle was known to MECKEL, who erroneously regarded
it as the pericardial cavity. J. MtiLLER. himself failed to observe that the ventricular part
of the sinus is really a coarse plexus, and was therefore unable to understand why
it could not fully be inflated. He considered whether the cardiac "and peribranchial
sinuses were lymph spaces, and decided that they were not, holding that they corresponded
rather with the pleural cavities of higher vertebrates.
Connections.—Peribranchial sinuses.
2. Peribranchial Sinuses (Figs. 2, 8, p.b.s., p.b.s.').

First described by J. MULLER in 1836. These sinuses, otherwise known as the
pleural sacs, consist of six large bags on each side enclosing the gills, and include
always a considerable quantity of red blood. It is clear, however, that the gills are
morphologically outside the sacs, and are related to them in the same way as the
thoracic and abdominal viscera are to their respective cavities. Owing to the way the
gills overlap the sinuses also overlap, so that between any two adjacent gills there
is a double partition which is normally imperforate. Each sac consists of a visceral
layer, closely opposed to the gill itself, and a parietal layer which forms the external
wall of the sac. There is an internal and an external invaginated area for the entry
and exit of the afferent and efferent gill ducts and vessels (fig. 8). There is no sinus
associated with the ductus oesophago-cutaneus, as mentioned by J. MULLER. Each sac,
except usually the first, opens freely dorsally into the peribranchial anastomosis, which
itself opens posteriorly into the subchordal sinus at the sixth peribranchial sinus. In fig. 8
the latter connection is shown, but its actual position is posterior to the region of this
section.
Each peribranchial sinus is evaginated on to the external surface of its efferent gill
duct (fig. 8). This evagination is considerable except in the case of the first and second
gills. Each of these duct extensions also opens widely into the extension in front and
the sinus behind, so that duct extension 3, for example, is derived from peribranchial
sinus 3, and in addition communicates with duct extension 2 and peribranchial sinus 4.
Some of the sinuses, however, may communicate directly with each other. They are
therefore not independent units, but are linked up, at least indirectly, by the dorsal
peribranchial anastomosis, the lateral duct extensions, and the ventral cardiac sinus.
Posteriorly the last or sixth peribranchial sinus, at the place where it fuses with the
posterior extremity of the peribranchial anastomosis, opens by a large valved aperture into
the anterior cardinal vein, or rather into the large venous anastomosis connecting the
two anterior cardinal veins (fig. 2, p.b.s.'). An injection mass passes readily from the
sinus into the veins, but the valve prevents a flow in the opposite direction.
Connections.—1. Subchordal sinus via the peribranchial anastomosis. 2. Cardiac sinus.
3. Anterior cardinal venous system via the sixth peribranchial sinus + the peribranchial
anastomosis.
3. Peribranchial Anastomosis (Fig. 8, p.b.a.).
This is a longitudinal somewhat small paired sinus situated dorsal to the peri-
branchial sinuses and attached to the internal surface of the dorsal anterior limb of the
constrictor branchiarum et cardise muscle (cf. Part II, PI. II, figs. 2 and 3). It commences
blindly in front and has no connection in most cases with the first peribranchial sinus, but it
may at this point be in communication with the sub-chordal sinus. The anastomosis
opens freely into all the peribranchial sinuses behind the first, and appears to fuse com-
pletely with the last of them. The point, however, where the sixth peribranchial sinus
opens into the subchordal sinus is the place where the anastomosis has fused with it. Also
the place where the sixth peribranchial sinus opens into the anterior cardinal anastomosis
(fig. 2, p.b.s.') is where the peribranchial anastomosis has fused with it, so that the latter
may be described as the vessel which connects up the sixth peribranchial and subchordal
sinuses and discharges into the vein. Thus the anastomosis opens at each end into the
subchordal sinus, and in the intervening region into the peribranchial sinuses.
Connections.—1. Subchordal sinus. 2. Peribranchial sinuses. 3. Anterior cardinal
venous system.
4. Subchordal Sinus (Figs. 7, 8, s.ch.s).
First described by J. MULLER in 1836, as "probably a lymph sac." He describes its
connection with the subcesophageal sinus, but appears to confuse it with the lateral chordal
sinus.
A large extensive median sinus constituted anteriorly and coursing backwards. It
lies for the most part immediately below the notochord, and stretches from the region just
behind the skull to the level of the external branchial opening. At each extremity it
consists of an irregular and variable coarse plexus, which has no connection with any part
of the blood vascular system. This is established by injection experiments. Posteriorly
the sinus dips ventrally under the anterior median edge of the constrictor branchiarum et
cardise muscle (fig. 8), and it then lies ventral to the median unpaired section of the lateral
chordal sinus on the dorsal surface of the gut, where it terminates in a plexus as above
mentioned. This posterior region is connected at the side of the gut with the posterior
section of the suboesophageal sinus (fig. 8). The peribranchial anastomosis opens posteriorly
into the subchordal sinus in the region of the sixth peribranchial sinus (fig. 8), and at
its anterior extremity may also open into the subchordal. This places the latter sinus in
indirect communication with all the peribranchials (except usually the first).
In a letter to me written in 1906 KLINCKOWSTROM mentioned that on one occasion he
was able to inject the portal heart as well as the veins entering and leaving it from the
subchordal lymph sac, but that he could never succeed in injecting the lymph sac from
the veins, from which .he concluded that the connection between the two series of vessels
must be by a valved aperture. In the experiment in question the injection would pass
from the subchordal sinus into the peribranchial anastomosis, and from thence through the
valved opening into the anterior cardinal venous anastomosis. From this point there is
nothing to prevent it extending into the veins in any direction (figs. 8 and 2).
Connections.—1. Suboesophageal sinus (anterior)—frequent lateral communications.
2. Suboesophageal sinus (posterior). 3. Peribranchial sinuses via the peribranchial anastomosis.
G. W H I T E LYMPHATICS.
1. Lateral Chordal Sinus (Figs. 3, 7, 8, 9, 10, 11, l.c.s., l.c.s.', l.c.s.").

This sinus and its associated segmental lymphatics are briefly described by J. MULLER.
He investigated them by inflation methods, and mentions that he had not discovered any
connections with the venous system.
The lateral chordal sinus may be regarded as the main axis of the white lymphatic
system, stretching as it does practically from one end of the body to the other, and
having connections with several other important sinuses and also with the blood-stream
(cf. fig. 3). It begins anteriorly immediately behind the auditory capsule lateral to the
notochord, and is here continuous with the subcutaneous anastomosis (fig. 3, s.c.a."') which
leads into the large subcutaneous sinus. This connection has been demonstrated by injec-
tions. The lateral chordal sinuses then pass backwards ventro-lateral to the notochord and
dorsal to the subchordal sinus, occupying the triangular space bounded by the myotomes,
notochord, and (in front) the subchordal sinus (fig. 7). During their passage backwards
and throughout their whole length they receive segmental vessels, which are, however, not
strictly segmental but arranged somewhat irregularly and differently on the two sides.
The sinus splits at intervals to allow the dorsal sections of the segmental vessels to pass
upwards at the side of the notochord. At about the level of the second and third gills
the two lateral chordal sinuses unite to form a median sinus lying immediately under the
notochord and dorsal to the subchordal sinus. Apart from certain irregularities of struc-
ture it now passes backwards as a single median sinus until it fuses with the intestinal
sinus in the region of the heart (figs. 3, 9, i.s., l.c.s.").
The intestinal sinus (i.s.) more or less surrounds the gut in the region of the heart and
the anterior lobe of the liver (fig. 9). It may be regarded as an extension of the median
portion of the lateral chordal sinuses, and has indefinite anterior and posterior boundaries.
On the right side it is associated with the common portal vein, and receives the products
of a well-marked and elaborate plexus on the surface of the gall bladder. Posteriorly it
receives the intestinal lymphatic trunk which accompanies the portal vein (fig. 3, i.l.t.).
Behind the intestinal sinus the lateral chordal sinuses separate out again as a paired
structure, and traverse the whole length of the abdominal cavity and a part of the tail
(fig. 3). Just in front of the anus the left lateral chordal despatches ventrally one or more
anastomoses to the rectal sinus, which latter is formed by the expansion of the intestinal
lymphatic trunk (fig. 3, r.s.). The latter sinus resembles the intestinal sinus in its relations
to its own part of the gut. In front of the median ventral bar of the caudal fin skeleton
the lateral chordals for a brief space fuse again. The fused section, now median in position,
deepens dorso-ventrally, and gives off ventrally on each side the anastomosis opening into
the caudal heart (figs. 3, 10, 11, l.c.s.'), which passes internal to the caudal vein and heart
of its side, but external to the forwardly projecting knob of the median ventral bar of the
caudal fin skeleton (fig. 11). In this way the contents of the lateral chordal sinus reach
the caudal vein and posterior cardinals via the caudal heart. The sinus now loses the
ventral deepening, and the remaining dorsal portion, which is situated immediately under
the notochord, splits for the last time, and is continued backwards, one division on each
side of the median ventral bar, to the level of the posterior extremity of the caudal heart,
where it terminates blindly.
Connections.—1. Subcutaneous sinus via the subcutaneous anastomosis. 2. Subcutaneous
sinus via the segmental lymphatics (fig. 7). 3. Ventral and hypophysio-velar sinuses via
the subcutaneous anastomosis. 4. Intestinal lymphatic trunk via the series of abdominal
anastomoses and the direct passage anteriorly of the intestinal lymphatic trunk into the
intestinal sinus (fig. 3). 5. Caudal vein via the caudal hearts. 6. Segmental lymphatics
of the whole body.
2. Segmental Lymphatics (Fig. 7, seg.l., seg.l.').
The segmental lymphatic has the typical dorsal and ventral rami of a segmental vessel
with the striking exception that both are double, each pair opening into the lateral chordal
sinus of its side (l.c.s.). Between the double segmental lymphatic vessels of both rami
courses the segmental artery or vein—each segment, whilst possessing a segmental nerve,
having either a segmental artery or vein, and only rarely both. Lymphatics and blood-
vessels pass anterior to their corresponding segmental nerve. The occurrence of the lym-
phatics is not regular; in some cases they occur in almost every segment, in others in alternate
segments more or less. On the tail of Bdellostoma ALLEN finds that there is a lymphatic
in every segment accompanied by either a segmental artery or vein. Immediately above
the dorsal edge of the obliquus muscle in the case of the trunk vessels, the posterior
member of each lymphatic doublet gives off an anastomosis which passes straight through
the parietal muscle to open intersegmentally into the subcutaneous sinus {seg.l.'). An
injection thrown into the latter sinus reaches the lateral chordal sinus via these anastomoses
and the segmental lymphatics, and may subsequently pass also into the intestinal lymphatic
trunk. On the tail the connection between the segmental lymphatics and the sinus is
intersegmental and dorsal to the slime sacs, and may be of a more complex nature than
the one just described. In some injections the segmental lymphatics were connected up
by a longitudinal anastomosing vessel just dorsal to the slime sacs.
Connections.—1. Subcutaneous sinus via the segmental anastomoses. 2. Lateral chordal
sinus.
3. Intestinal Lymphatic Trunk (Fig. 3, i.l.t., i.l.t.').
This is a longitudinal vessel which accompanies the portal vein, and is situated
immediately to the left of the latter. Into this vessel an elaborate anastomosing plexus
of lymph capillaries discharges. These capillaries arise blindly in the wall of the gut in
the usual way, and are not the vessels described by MAWAS (i.l.t.'). The plexus is situated
on the outer surface of the gut immediately under the serosa, and lies superficial to the
blood capillary system. If the portal vein and the lymphatic trunk are injected with
different coloured media, the two systems of capillaries, one lying over the other and
quite distinct from each other, are strikingly shown. The lymphatic trunk discharges in
front into the posterior margin of the intestinal (lateral chordal) sinus, and behind into
the anterior margin of the rectal sinus, which embraces the gut and is situated just
anterior to the cloacal opening. It may be regarded as an expansion of the trunk itself
(fig. 3). The intestinal lymphatic trunk, as demonstrated by numerous injections, is con-
nected at intervals by means of anastomoses with the left lateral chordal sinus, these anas-
tomoses accompanying the branches from the aorta to the intestine. There is also a special
larger anastomosis connecting the rectal sinus with • the left lateral chordal which passes
downwards between the two posterior cardinal veins just in front of their formation by the
bifurcation of the caudal vein. A successful injection of the lymphatic plexus in the wall of
the gut can be made from the left lateral chordal sinus, and even from the subcutaneous
sinus via the segmental lymphatics (cf. fig. 7).
There can be no question that the intestinal lymphatic trunk is a true lymphatic
vessel for the following reasons: it originates as blind capillaries in the wall of the
intestine, its contents never exhibit any admixture of red blood, and it discharges
ultimately into the blood-stream.
The lymph from the gut probably enters the blood-stream via the lateral chordal
sinus, which communicates directly with the caudal heart (fig. 3). It might, however,
pass from the lateral chordal sinus via the segmental lymphatics into the subcutaneous
sinus, and again reach the blood-stream via the caudal heart.
In some specimens I have found a ventral lymphatic on the gut corresponding with
the subintestinal vein. This also opened into the intestinal sinus.
Connections.—1. Left lateral chordal sinus by numerous anastomoses. 2. At its two
extremities it passes into sinuses—the intestinal and rectal sinuses—the former of which
may be regarded as an expansion of the fused lateral chordals at this region, the latter
being also connected with the same sinus on the left side.
H. CIRCULATION OF THE BLOOD.

We have now to deal with the question of the circulation of the blood between
the so-called lymphatic spaces and the definite blood-vessels. The occurrence of red blood
in at least one of the lacunar spaces has been known since 1824, when A. A. RETZIUS
published the following statement relating to Myxine: "Thus the animal lies in its
skin as in a loose sac; the intervening space contains, both in newly dead animals and in
those which have been kept in spirit, a considerable amount of blood. In the newly
captured animal the blood flows here and there, its red colour showing through the skin,
so that, according as there is an increased flow to one or the other place, either the head
or the tail becomes redder." This observation remained without a meaning until 1890,
when G. RETZIUS discovered the caudal heart of Myxine. He described its pulsation and
believed that it pumped the blood-like fluid from the subcutaneous sinus into the caudal
vein, but was unable to establish the latter point by injection experiments. He mentions,
however, that KLINCKOWSTROM had succeeded in injecting the caudal veins from the
subcutaneous sinus. In 1891 KLINCKOWSTROM, who had previously in 1890 concluded
that the subcutaneous sinus was a blood vascular as well as a lymphatic space, published
a brief note, in which he claimed that the contents of the sinus passed first into the
caudal heart, and from thence into the caudal vein. This was the first definite assertion
of the existence of a connection between the lacunar spaces and the venous system, and
since that time no others have been described, nor has any suggestion been made as to
how the red blood reached the lymphatic spaces. It is true KLINCKOWSTROM also found
that in one case an injection passed from the peribranchial sinuses, not only into the
portal heart but also into a part of the venous system; but he was unable to explain how
this came about. RATHKE in 1825, in his work on the Lamprey, was the first to describe
a lacunar circulation in a Vertebrate. He says : " If I can interpret what my investigation
of the backflow of the blood out of the sexual parts has disclosed, the arterial blood over-
flows into the tissue of the sexual parts without being enclosed in special vessels, flows
round the eggs or sperm spheres as it were entering into a sponge, and is collected finally
into certain vessels, the veins." These statements escaped comment probably because it was
then an article of faith that the Vertebrate circulation was a closed one. The discoveries
of G. RETZIUS and KLINCKOWSTROM re-opened the question, and in a letter to me written in
1908, the former remarked : " I have, of course, also always seen that blood was present in
the spaces of the captured living animals \Myxine\; but as they were taken from a great
ON THE GENERAL MORPHOLOGY OF THE MYXIN01D FISHES. 333
depth (60 fathoms or more, or a little less), I could not exclude or refute the objection that
the blood was derived from burst small blood-vessels in the walls of the subcutaneous
spaces. We have often here discussed this question, but could never solve the- problem.
Now more and more I have been convinced that the blood naturally belongs to the
lacunar spaces already in the fresh living animals." The belief, however, that the blood
in the subcutaneous sinus of Myxinoids was due to extravasation was held until quite
recent times, and JACKSON, in his paper on the vascular system of Bdellostoma published
in 1901, still believed that there were no red corpuscles in the lymphatic spaces in life
and in uninjected specimens. That this statement is not correct is now well known,
although, according to ALLEN, the lymphatic spaces of Bdellostoma contain great quantities
of red corpuscles in the embryonic stages, but only a few in the adult.
That red blood occurs normally in the so-called lymphatic spaces of Myxinoids
is beyond question. At four widely separated points (cf. fig. 2) definite provision is made
for the return of the contents of the spaces to the blood-stream. This indicates clearly
that they constitute a part of the general vascular system, and that there must be
a constant and considerable flow from the so-called lymphatics to the veins, but it
does not necessarily involve the presence of red blood in the spaces. The fact, however,
that they contain red blood in the earliest developmental stages (ALLEN) is an important
piece of evidence. Against the extravasation hypothesis, which is dependent on a deep-
sea habitat, we have the fact that red blood is, nevertheless, found in the spaces of the
shallow-water Myxinoids and in the corresponding cavities of the fresh-water Lampreys.
Further, I have never seen a trace of red blood in the pericardial and abdominal
cavities, where we should certainly expect to find it if extravasation had occurred. We
must therefore conclude that those lymphatics which contain red blood form an integral
part of the blood vascular system of the animal, and that any scheme of the circulation
of the blood must not only include the red lymphatics, but must also explain how
the red contents are acquired and dismissed.
An examination of the contents of the lacunar spaces reveals the fact that most
of them invariably contain red blood in more or less considerable quantity, but that
others exhibit only a very few isolated corpuscles or (normally) none at all. We must
therefore separate these two groups, and, as explained above, they may be referred
to provisionally as the red and white lymphatics. Further, the latter fall into an
independent system which is complete in itself, and corresponds, at least in part and
physiologically, with the true lymphatics of higher Vertebrates. The intestinal lymphatic
trunk, for example (fig. 3, i.l.t.), arises as an elaborate blind plexus in the wall of
the gut, and discharges finally into the blood-stream, and therefore may be compared
physiologically with a true lymphatic vessel. The white and the red lymphatics are only
connected with each other by the subcutaneous anastomosis (fig. 3), the segmental
lymphatics (fig. 7), and at one point of the caudal circulation (figs. 10 and l l ) . The
occurrence of a few red corpuscles in the white lymphatics is paralleled by the occasional
presence of red blood in the true lymphatics of higher animals. The white lymphatics
are connected with, and discharged into, the red blood-stream, as elsewhere described,
their contents entering the circulation especially by the caudal hearts—directly from the
lateral chordal sinus (figs. 3 and l l ) , and indirectly via the subcutaneous blood sinus
(cf. text-fig, l). The latter sinus is fed with white lymph by the subcutaneous anasto-
mosis and the segmental lymphatics, and it is returned to the blood-stream via the
caudal hearts. In the case of the caudal hearts there can be no question that the direction
of the flow is from the white lymphatics into the true blood-vessels, and in other cases it
is a legitimate inference, since, for example, the blood-pressure in the red subcutaneous
sinus is a negative quantity, and therefore the contents of the small white segmental
lymphatics would naturally pass into it.
Inferior Jugular l/em
Cardinal Heart
Rostral Dorsal Oral
Subcutaneous
Subhypophysial
Anastomosis
*L ateral
Chorda I
*lntestmal
Lymphatic
Lingual
Suboesophageal
(anterior)
Inferior Jugular Vein
Posterior Cardinals Subchordal

v/aCaudal Heart
FIG. 1.—Scheme to illustrate the interrelationships of the sinuses constituting the subcutaneous cycle.
The true or white lymphatics are also included, and are distinguished by asterisks.
An analysis of the anatomical relations of the red lymphatics shows that they fall
into two cycles—an anterior subcutaneous cycle and a posterior peribranchial cycle. The
two cycles are largely but not entirely independent of each other, and are connected up
via the anterior suboesophageal sinus. The dominating feature of the former cycle is the
extensive subcutaneous sinus, with which a number of the other members of the cycle are
connected. Red blood enters the cycle from the arteries of the snout.* What happens
• ALLEN was unable to find in Bdellostoma any direct connection between the arterial system and the "lymphatics."
then is almost impossible to determine, and the direction of the flow indicated in the
scheme is based on anatomical relationships only. The blood, however, is undoubtedly
returned to the veins of the definitive blood vascular stream by valved openings at the
following three points (cf. fig. 2): (l) via the hypophysio-velar sinus and cardinal heart
into the anterior section of the jugular system (h.v.s.', cl.h., r.d.c.); (2) via the lingual sinus
into the middle section of the jugular system (l.s.', i.j.v.); (3) via the subcutaneous sinus
of the tail region and caudal hearts into the caudal vein and posterior cardinals (fig. 11,
s.c.s., s.c.s.', cd.h., c.v.).
The central sinus of the peribranchial cycle is the subchordal sinus, but the largest
members of the cycle, in fact the largest sinuses in the body apart from the subcutaneous
sinus, are the peribranchials. Red blood enters the cycle via the afferent and efferent
branchial arteries and also _ the dental sinus of the subcutaneous cycle, and is returned to
the veins of the posterior section of the jugular system via the peribranchial anastomosis
(fig. 8, p.b.a.) and the anterior cardinal venous anastomosis (fig. 2, p.b.s.', c.a.).
Cardiac
Peribranchials
Peribranchial Anterior
-cardinal
anastomosis anastomosis
Subchordal
Suboesophageal Subbesophageal
(anterior) (posterior)
Dental
FIG. 2.—Scheme to illustrate the interrelationships of the sinuses constituting
the peribranchial cycle.
I think it must be admitted, assuming the statements of fact in this paper to be

correct, that the red lymphatics of the adult Myxinoid are not true lymphatics, but belong
rather to the blood vascular system; and in the places where they exist, a closed capillary
system being absent, act as the means of communication between the arteries and veins.*
* According to MCCLURE (Mem. JVistar Inst., 1915), the lymphatics of fish are not transformed veins, but true lymphatics,
which may subsequently become connected up with the veins, from which latter they derive their red blood. After the
formation of the lymphatico-venous valves blood does not pass into the lymphatics. These conclusions are not applicable to
the adult Myxinoid; and ALLEN, who has investigated the development of the lymphatics of Bdellostoma, concludes that " the
most primitive form of a lymphatic system are veins that function for both lymphatics and veins," and that, instead of having
their origin directly from veins, the lymphatics " would begin exactly as the veins did, namely, by the vacuolation of the original
mesenchyme."
There thus exists in Myxine a typical lacunar circulation. In a letter to me KLINCKOWSTROM

observes : "If I call these blood-carrying cavities lymph spaces it is only because I have
not yet found a direct communication between them and the vascular system by which
blood could enter them. Only after a demonstration of such a connection can the question
be solved and the cavities called lacunae with correctness. I am, however, firmly of opinion
that such a connection exists in the arterial part of the capillary network." I hope to
have shown that KLINCKOWSTROM'S conjectures were remarkably close to the truth.
The absence of valves in the veins, except where connections are effected with the
lacunar spaces, sets up a number of alternative routes when the dead animal is injected,
and the direction of flow depends on variations in pressure of the circulating liquid at
different points. If an injection be thrown into the left posterior cardinal, it naturally
passes forwards into the sinus venosus and through the heart into the cardiac aorta. But
it also passes forwards beyond the sinu-auricular aperture into the inferior jugular vein.
Further, it travels forwards along the by-pass represented by the dorsal and ventral
divisions of the anterior section of the left posterior cardinal (fig. 2, l.a.c.', l.a.c"), and so
reaches the cardinal anastomosis, from which it passes forwards into the two anterior car-
dinals, and backwards into the portal heart and veins and the right pronephric vessel (r.a.c.').
In fact, almost the whole venous system can be injected in this way. Now a precisely
similar result is obtained by injecting the portal vein forwards, in this case the injection
passing backtvards through the by-pass mentioned above, and then forwards to reach the
heart and inferior jugular vein (cf. fig. 2). The unique connection of the portal and car-
dinal systems, and the asymmetrical position of the heart, both tend to produce this result,
and it would be difficult to predict, in the case of certain vessels in the neighbourhood of
the heart, whether' the blood in the living animal flowed forwards or backwards or if
indeed it followed a definite direction at all.
J. SUMMARY.
1. A degenerating lymphoid or thymus-like organ occurs in the pharyngeal velum.
2. The liver is an hepato-pancreas. Special glandular tubules of a pancreatic nature
are associated with the branches of the portal vein within the liver parenchyma, as in
some other fishes.
3. There are two cavities associated with the heart. The inner one is a blood sinus
and surrounds the ventricle only. The outer one is related to the ventricle, auricle, and
portal heart, and is a true pericardial cavity. It communicates with the abdominal coelom
on the right side by the large pericardio-peritoneal foramen.
4. The heart is asymmetrical. The auricle and sinus venosus are situated on the
left side, and the sinus venosus is ventral to the auricle. This differs from the Selachian
arrangement.
5. The coeliac artery is paired, and encircles the gut. The hepatic artery arises from
the left member of the pair.
6. Vascular papillae occur throughout the whole length of the common carotid artery,
but on no other vessel.
7. A vertebralis impar capitis artery is formed by the union of the "internal" carotids
8. The arteries differ from the veins in being continued forwards to the extremity of
the snout, where their contents are discharged principally into the blood sinuses of the
snout and tongue.
9. The general characters of the venous system are represented in fig. 2. Attention
may be directed to the asymmetry of the anterior cardinal system, the presence of a heart-
like structure posterior to the auditory capsules, the existence of an anterior cardinal
anastomosis, and the relations of the pronephros to the cardinal system and of the anterior
and posterior cardinals to each other. The right posterior cardinal does not terminate by
fusing with the left, but is usually continued forwards symmetrically with the left. The
portal heart is probably a modified part of the cardinal system, and if so its connection
with the anterior cardinal is explained.
10. Apart from the skin and brain there are no definite veins anterior to the
auditory capsule, arteries and veins being connected by irregular blood sinuses instead of
capillaries.
11. The caudal artery and vein are connected by a capillary system. The relations
of the caudal hearts to the blood vascular and lymphatic systems are shown in figs. 10
and 11.
12. The lacunar system is lined by endothelium, and is divisible into red lymphatics,
which constitute a part of the blood vascular system proper and always contain red blood,
and the white lymphatics, which represent a true lymphatic system. The red lymphatics
fall into two cycles, largely but not entirely separate from each other. The red lymphatics
communicate with the venous system by seven valved apertures at four widely separated
points (fig. 2). The white lymphatics are connected indirectly with the veins by the red
subcutaneous sinus, but have a special direct communication via the caudal hearts. Ked
blood enters the red lymphatics directly from the arteries. Therefore the blood circulates
through these spaces and a lacunar circulation is established. A true lymphatic duct with
an associated capillary plexus in the wall of the gut accompanies the portal vein.
K. LITERATURE.
[In this list only those works are included which contain original observations or generalisations
on the vascular system of Myxinoids.]
(1) ALLEN. Q.J.M.S., 59, 1913.

Caudal hearts and veno-lymphatics of tail region of Bdellostoma.
(2) COLE. Anat. Anz., 27, 1905.
Afferent branchial arteries and lymphatic spaces in Myxine.
(3) COLE. Trans. Roy. Soc. Edin., 45, 1907.
Anatomy of segmental artery and vein in Myxine.
Afferent and efferent branchial arteries of Myxine.
Lymphatic spaces and circulation through the gills in Myxine.
(6) COLE. Anat. Anz., 46, 1914.
Venous system and lymphatic spaces in Myxine.
(7) COLE. Brit. Ass. Reports, Liverpool, 1923.
Veno-lymphatic and true lymphatic systems in Myxine
(8) FAVARO. Atti e Mem. R. Accad. Sci. Padova, 21, 1905.
Preliminary to following.
(9) FAVARO. Atti R. 1st. Veneto Sci., 65, 1906.
Caudal circulation in Myxine.
(10) FAVARO.In Bronn's Tier-Reichs, 1908.

General account of blood vascular and lymphatic systems of Myxinoids.
(11) GEACOMINI. Monit. Zool. Ital., 13, 1902. Rend. R. Accad. Set. 1st. Bologna, N.S., 8, 1904.
Notes on lymphatics of Myxinoids.
(12) GOODRICH. In LANKESTER'S Treatise on Zoology, part ix, 1909.
General account of Myxinoid circulation.
(13) GRHBNB. Amer. Jour. Phys., 3, 1900. Abstracts in Amer. Nat., 34, and J. R. Micros. Soc, 1900.
Anatomy and physiology of caudal heart of Bdellostoma.
(14) GREKNB. Science, 15, 1902.
Note on the three hearts of Bdellostoma.
(15) GREENE. Amer. Jour. Phys., 6, 1902.
Nerves of systemic heart of Bdellostoma.
(16) JACKSON. Jour. Cincinnati Soc. Nat. Hist., 20, 1901. Reprinted: Bull. Univ. Cincinnati, 1, 1901. Abstracts:
Amer. Nat, 36, and J. R. Micros. Soc, 1902.
Vascular system of Bdellostoma.
(17) KLINCKOWSTROM. Biol. Foren. Forhand. Stockholm, 2 and 4, 1890, 1891.
Gut and liver veins, caudal heart, and lymphatic system of Myxine.
(18) MAAS. Zool. Jahrb. Ait. Anat., 10, 1897.
Vascular supply of kidney system in Myxine.
(19) MAAS. Kupffer's Festschrift, 1899.
Structure and origin of blood corpuscles of Myxine.
(20) MAWAS. C. R. Acad. Set., Paris, 174, 1922.
Lymphatic tissue of the gut of Myxinoids.
(21) MECKEL. System d. Vergleich Anat., v, 1831. French translation, ix, 1837.
Pericardium of Myxine.
(22) MULLER, J. Abh. Ah. Berlin for 1834. Published 1836.
Lymphatic spaces of Myxinoids.
(23) MULLER, J. Abh. Ak. Berlin for 1839. Published 1841. Issued separately 1842. Abstracts: Ber. Ah. Berlin,
1839, and Muller's Archiv, 1840.
Vascular and lymphatic systems of Myxinoids.
(24) MULLBR, J. Abh. Ak. Berlin for 1843. Published 1845. Issued separately 1845.
Blood, portal heart, and vascular supply of the kidneys in Myxinoids.
(25) PRICE. Zool. Jahrb. Abt. Anat., 10, 1897. Amer. Jour. Anat, 4, 1904. Jour. Exper. Zool., 9, 1910.
Vascular supply of excretory organs of Bdellostoma.
(26) RAUTHER. In Bronn's Tier-Reichs, 1924.
Development of vascular system of Myxinoids.
(27) RETZIUS, A. A. K. vet. Akad. Handlgr. Stockholm, 1822 and 1824. German translations, Isis, 1825, and
Meckel's Archiv, 1826. French translation, Ann. Sci. Nat., 14, 1828.
Vascular system of Myxine.
(28) RETZIUS, G. Biol. Unters., 1, 1890.
Caudal heart of Myxine.
(29) RETZIUS, G. In FKIES, EKSTROM, and SUNDERVALL'S Scandinavian Fishes, 1895.
Vascular and lymphatic systems of Myxine.
(30) STERZI. 11 sistema nervoso centrale d. vertebrati, Padua, 1907.
Vascular supply of Myxinoid brain and cord.
(31) THOMPSON, D'ARCY. Ann. Mag. Nat. Hist. (5), 20,1887. Anat. Anz., 2, 1887. Abstract: J. R. Micros. Soc. 1887.
Blood corpuscles of Myxine.
OJST THE GENERAL MORPHOLOGY OF THE MYXINOID FISHES. 339
L. EXPLANATION OF PLATES.
REFERENCE LETTERS.
1, 1/ 2, 3. Anterior and posterior cusps of tooth 1 and cp.t.c' Posterior) divisions of the copulo-tentaculo-
teeth 2 and 3 of outer row of ventral teeth. cp.t.c." Anterior J coronarius muscle.
S. Third tentacle with skeleton. c.p.v. Common portal vein splitting into two branches
3.' Skeleton of third tentacle just prior to junction which pass to the anterior and posterior lobes
with the lateral labial cartilage. of the liver.
a.d.p. Anterior arch of the dental plate. c.q.p. Copulo-quadratus profundus muscle.
a.h. Anterior hepatic vein. e.q.s. Copulo-quadratus superficialis muscle.
a.s.ao. Anterior portion of the dorsal or systemic aorta cr. Membranous cranium.
( = median anterior aorta or vertebralis c.v. Median section of caudal vein.
impar). C.V.' Paired factor of caudal vein expanding to form
au.c. Auditory capsule. In fig. 6 with contained the caudal heart = afferent vein of caudal
labyrinth. heart.
aw. Auricle of systemic heart. c.v. Efferent vein of caudal heart which joins at
b.p.l~s Anterior, middle, and posterior segments of once with its fellow of the opposite side to
the basal plate, with guiding rails on dorsal form the median caudal vein.
surface of middle segment (fig. 6) directing cys.v. Cystic vein.
the antero-posterior motion of the dental d.o.s. Dorsal oral sinus.
apparatus. d.s. Dental sinus. In fig. 6 fused with lingual sinus
c.a. Anterior cardinal anastomosis. ventrally, and on right side communicating
c.ao. Ventral or cardiac aorta. dorsally with the subhypophysial sinus.
cb.a. Cerebral artery (= internal carotid). In fig. 5 note that the dental sinus is divided
c.h.c. Branchial portion of the constrictor branch- by the dental plate into external and in-
iarum et cardise muscle. ternal portions.
c.h.c' Ventral longitudinal tract of ditto. d.s.' Latero-dorsal extension of dental sinus com-
c.h.c. municating with the subhypophysial sinus.
) Loops 3 and 4 of ditto (cf. Part II, Plate II).
c.b.cM e.b.p.' External bar of the anterior segment of basal
cb.v. Cerebral veins constituting the right and left plate.
superficial anterior cardinal veins. e.c.a. External carotid artery.
c.c. Cornual cartilage. e.l.b. External lateral velar bar.
c.car. Common carotid artery( = paired anterior aorta). ex.c. External carotid artery.
c.car.' Anastomosis between the paired and median f-r. "Fin rays" of the caudal fin.
anterior aortse. h.a. Hepatic artery.
cd.c. Cordis caudalis muscle. h.c. Naso - pharyngeal or hypophysial canal just
cd.h. Caudal heart. anterior to its junction with the pharynx.
cd.s. Cardiac sinus surrounding the ventricle and On the right it has already fused with the
cardiac aorta. velar portion of the pharyngeal gut.
c.e. Copulo-ethmoidalis muscle. h.c.g. Hyo-copulo-glossus muscle.
e.g.p.' Lateral \ heads of the copulo-glossus profundus h.c.g.' Anterior end of " tendon" of the hyo-copulo-
c.g.p." Median J muscle. glossus muscle.
c.g.s. Copulo-glossus superficial muscle. h.c.p. Hyo-copulo-palatinus muscle.
cl.h. Cardinal heart. In fig. 6 connected by a h.p. Anterior process of hypophysial plate.
valved aperture with the hypophysis-velar h.v. Factors of hepatic veins in the liver.
sinus. h.v.s. Hypophysio-velar sinus. In fig. 5 note the
CO. Coronarius muscle. forward extension of this sinus into the
coB.a.
coe.a.' J coeliac or gastric arteries.
median olfactory lamina.
h.v.s.' Portion of hypophysio-velar sinus which
cod. Abdominal coelome. communicates by valved aperture with the
c.p.' First division of the constrictor pharyngis cardinal heart.
muscle. h.v.s." Section of hypophysio-velar sinus which arises
c.p.c. " Common " posterior cardinal vein (= left in association with the lymphoid organ of
posterior cardinal). the velum. The two halves have fused at
c.pl. Copulo-palatinus muscle. the middle line.
hy. Hyoid arch. obi. Obliquus muscle.

i.b.p.' Internal bar of the anterior segment of the o.c. Oral cavity with " tongue " and teeth.
basal plate. par. Parietalis muscle (forming the myotomes).
i.e.a. "Internal carotid artery" or posterior paired p.b.a. Peribranchial anastomosis. On the left side
factor of the vertebralis irapar (= v.i.'" of the communications with the subchordal
%. 1); and peribranchial sinuses are shown.
i.j.v. Median inferior jugular vein. p.b.s. Peribranchial sinus with extension on to the
i.l.b. Internal lateral velar bar. second, third, and fourth efferent gill ducts.
i.l.t. Intestinal lymphatic trunk. p.b.s.' Point where the sixth peribranchial sinus
i.l.t.' Small portion of capillary plexus in the wall communicates by valved apertures with the
of the gut which discharges into intestinal anterior cardinal system.
lymphatic trunk. p.c' Parachordal cartilage fused with the auditory
i.s. Intestinal sinus = expanded portion of the capsule with embedded anterior extremity
median section of the lateral chordal sinus. of notochord.
La. Anterior lingual artery. p.e.p. Palato-ethmoidalis profundus muscle.
La.' Posterior lingual artery reaching the tongue p.e.s. Palato-ethmoidalis superficialis muscle.
via the tether of the hyo-copulo-glossus ph. Pharynx just anterior to junction with hypo-
muscle. physial canal.
l.a.c. Left anterior cardinal vein. p.h. Portal heart.
l.a.c' Dorsal division of the anterior section of pi. " Palatine " bar.
the left posterior cardinal (with included pl.c. Palato-coronarius muscle.
pronephros). pn. Pronephros.
l.a.c." Ventral division of same. pn.' Isolated malpighian bodies with or without
l.a.c.d. Left deep anterior cardinal vein (= inferior attached ductules in the zone between the
jugular). pronephros and the abdominal renal organ.
l.a.c.s. Left superficial anterior cardinal vein (= in- p.p.f. Pericardio-peritoneal foramen {i.e. relations of
ternal jugular). —its actual position is further back).
Lea. Left caudal artery. processes of pterygo-quadrate carti-
l.c.s. Lateral chordal sinus. lage. Between these two processes
l.c.s.' Branch of lateral chordal sinus by which it p.q.1 Superior is the third fenestra of the skull,
discharges into the caudal heart. p.q.2 Inferior | and between the auditory capsule
l.c.s." Fused median portion of lateral chordal sinus a,ndp.q.i is the second fenestra of
expanding below into the intestinal sinus. the skull.
l.l.c. Lateral " labial" cartilage. p.v. Portal or supra-intestinal vein.
l.lg.' Tendon of longitudinalis linguae muscle. p.v.' Branches of portal vein in the liver surrounded
l.lg." Dorsal paired and median ventral tendons of by pancreatic tubules.
the longitudinalis linguee muscle. q.p.' Tendon of quadrato-palatinus muscle.
l.p. Lateral plate of the nasal capsule. r.a.c. Right anterior cardinal vein.
l.p.c. Large left posterior cardinal vein connected r.a.e.' Dorsal division of the anterior section of the
with the small right one by anastomoses. right posterior cardinal vein with included
I.s. Lingual sinus. In fig. 6 note dorsal extension pronephros.
associated with the paired tendon of the r.c.a. Right caudal artery.
longitudinalis linguse muscle, and the com- r.d.c. Right deep anterior cardinal vein.
munication with the dental sinus. r.d.c' Posterior extension of above, which is con-
I.s.' Posterior extremity of lingual sinus com- tinued backwards as the median inferior
municating by a valved aperture with the jugular vein. Not present on the left side.
inferior jugular vein. red. Rectus muscle.
m.d.b. Median dorsal bar of the caudal fin skeleton. rect.' Attachment of tendon of rectus muscle to
mt.e. Metencephalon. middle segment of basal plate.
m.v.b. Median ventral bar of the caudal fin skeleton. r.p.e. Smaller right posterior cardinal vein.
m.v.b.' Anterior knob of same. r.p.o.' Reduced portion of the right posterior cardinal,
nas. Nasalis muscle. which may be absent.
n.c. Nasal capsule. r.s. Rectal sinus, surrounding rectum, formed by
n.rA. Fourth ring of nasal tube skeleton. expansion of the posterior extremity of the
nt. Notochord. intestinal lymphatic trunk. Receives a con-
n.tb. Nasal tube. nection from the left lateral chordal sinus.
ON THE GENERAL MOEPHOLOGY OP THE MYXINOID FISHES. 341
r.s.c. Right superficial anterior cardinal vein. s.o.s.' Posterior suboesophageal sinus and communica-
rs.s. Kostral sinus. tion with the sub-chordal sinus.
s.a.a. Position of sinu-aurioular aperture of heart. sp.c. Membranous neural tube.
s.ao. Dorsal or systemic aorta. s.s. Slime sacks numbered from before backwards.
s.c.a. Subcutaneous anastomosis. s.v. Sinus venosus.
s.c.a.' Connection of same with hypophysio-velar sinus. sy. Sympathetic nerve or ft. intestinalis vagi.
s.c.a." Connection of same with ventral sinus. i.e. Tentaculo-ethmoidalis muscle.
s.c.a.'" Connection of same with lateral chordal sinus th. Lymphoid organ of the pharyngeal velum
forming the anterior extremity of the latter (1 thymus).
sinus. thy. Thyroid vesicle.
s.ch.s. Subchordal sinus. t.o. Transversus oris muscle.
ts.c.s. Subcutaneous sinus. t.p. Tentacularis posterior muscle.
s.c.s.' Marginal vessel which drains the subcutaneous vel. Pharyngeal velum.
sinus and discharges into the caudal hearts. vel.a. Velar artery.
seg.a. Segmental artery. vent. Ventricle of systemic heart.
seg.l. Segmental lymphatic. The posterior member v.i. Eight and left anterior branches of the "un-
of the pair, which communicates with the paired vertebral artery of the head" or
subcutaneous sinus, is in black. vertebralis impar capitis.
seg.l.' Connection of above with the subcutaneous v.i.' Unpaired median or fused portion of same.
sinus. v.i." Branch of same to skin.
seg.v. Segmental vein. v.i.'" Paired posterior factor of same or "internal
s.g.i. First spinal ganglion. carotid."
s.h.s. Subhypophysial sinus, on right side com- vn.s. Ventral sinus, fn fig. 6 it communicates on
municating with the dental sinus. the right side with the subcutaneous anas-
s.i.v. Subintestinal or posterior hepatic vein. tomosis.
sn.b. Subnasal bar. v.q." Dorsal division of the velo-quadratus muscle.
s.o.s. Anterior suboesophageal sinus. v.s. Velo-spinalis muscle.
PLATE I.
Fig. 1. Scheme of the arterial circulation from the region of the heart to the tip of the snout as seen from
above. The lateral series of dots represent the position of the spinal ganglia, counting as number one the first
spinal nerve which possesses both dorsal and ventral roots. The longitudinal lines in the neighbourhood of the
first spinal ganglion indicate the antero-posterior position and extent of the auditory capsule (not its position
relative to the arteries). The vascular papillae on the common carotids are shown as black dots, but only some of
them are figured. In the specimen in question there were 95 on the left common carotid and 85 on the right.
The position of the gills in the vicinity of their respective pairs of efferent branchial arteries is marked.
Fig. 2. Scheme of the entire venous system and its connections with the lymphatic spaces as seen from
below. The four breaks represent more or less long stretches in which the vessels course unchanged. The figures
on the left mark the position of the respective spinal ganglia (cf. fig. 1), and the lines anteriorly indicate the
transverse level of the auditory capsules, but not the relative position of the latter to the veins. The small circles
in l.a.c.' and r.a.c! mark the position and extent of the left and right pronephros. The position of the cloacal
aperture is behind the place where the caudal vein splits to form the two posterior cardinals. The points where
the deep anterior cardinal veins pass through the fourth fenestra of the skull, and where the superficial anterior
cardinals pierce the cranium, are indicated by circles.
Fig. 3. Scheme to illustrate the entire course of the paired axial portion of the true or white lymphatic
system, i.e. the lateral chordal sinus, and its relations with other sinuses, the intestinal lymphatic trunk and the
venous system. Ventral view. Long stretches in front of and behind the systemic heart, where the vessels course
unchanged, are represented by breaks. The segmental lymphatics which open into the lateral chordal sinus throughout
its whole length are not shown. Only a small portion of the true lymphatic plexus in the wall of the gut is
represented. Caudal hearts displayed laterally.
PLATE II.
Fig. 4. Section 150 of my large series through the anterior margin of the mouth to illustrate the relations of
the hypophysio-velar sinus with the rostral and dorsal oral sinuses. Nerves and ligaments in black. The relations
342 PROF. FRANK J. COLE OK THE GENERAL MORPHOLOGY OF THE MYXlNOID FISHES.
of the anastomosis between the hypophysio-velar and dorsal oral sinuses are shown, but the actual connection is
posterior to this region.
Fig. 5. Section 390 through the nasal capsule and olfactory laminae. Illustrates the connection between the
subhypophysial sinus and the dental sinus via the latero-dorsal extension of the latter sinus, and the splitting up of
the dental sinus by the dental plate (a.d.p.) and the "tendon" of the hyo-copulo-glossus muscle (h.c.g.'). Olfactory
nerves, tendons, horny caps of the teeth, and a portion of the anterior arch of the dental plate shown in black.
PLATE III.
Fig. 6. Section 660 through the cardinal heart and the base or root of the pharyngeal velum. Somewhat
diagrammatic. Naso-pharyngeal tube (h.c.) is about to fuse with the pharynx, and has already fused on the right
side with that portion of the pharyngeal gut associated with the velum. Several sections have been telescoped
in order to illustrate the interrelationships of the various sinuses.
Fig. 7. Section 900. Segmental lymphatic (double) and its connection with the subcutaneous and lateral
chordal sinuses figured on left side. The posterior member of the pair is shown in black. Velar portion of
hypophysio-velar sinus and the left lateral chordal sinus also in black. The velum or pharyngeal valve with its
skeleton projects into the cavity of the gut from its dorsal wall.
PLATE IV.
Fig. 8. Section 2830. On the right side the complete course of an afferent branchial artery and on the left of
an efferent branchial artery is shown. On the right also the anastomosis between the cardiac and peribranchial
sinuses is illustrated. Efferent gill ducts one to four numbered from before backwards. On the left the connection
of the peribranchial anastomosis with the peribranchial and subchordal sinuses is figured, as also is the connection
of the latter sinus with the posterior section of the subcesophageal sinus.
Fig. 9. Section 3220 illustrating the relations of the pericardial cavity (in black) to the systemic heart, portal
heart, and abdominal ccelome. • The nature and position of the connection between the abdominal coelome and
pericardial ccelome are indicated, but the actual pericardio-peritoneal foramen occurs posterior to this region. The
branches of the portal vein in the liver are shown surrounded by pancreatic tubules.
PLATE V.
•
Fig. 10. Transverse section through the tail, passing through the narrowing anterior extremity of the caudal
hearts. Slightly diagrammatic. The dorsal portion of the marginal vessel of the subcutaneous sinus (s.c.s.') is here
double so as to allow the fin ray to fuse with the median dorsal bar of the caudal fin skeleton. The ventral
portion is shown as between two fin rays, and at these points the vessel is single. Note the asymmetry of the
caudal arteries at this point. They become symmetrical further back. The lateral chordal sinus and subcutaneous
sinus are in communication with the caudal heart on the right side. Behind this point the dorsal portion only of
the lateral chordal sinus is continued backwards as a paired vessel, the left one being already detached. Note the
mode of connection between the subcutaneous sinus and the dorsal and ventral marginal vessels.
Fig. 11. Dissection of right side of tip of tail to show the relations of the caudal heart with the subcutaneous
sinus. Injected from the latter sinus. Above on the right the skin only has been removed, which exposes the
cavity of the subcutaneous sinus and also a portion of the right coarse plexus on the caudal fin connected with
the sinus. On the left and below, the myotomes and fin plexus have been dissected away so as to uncover the
caudal heart and its vessels and the marginal channel which drains the subcutaneous sinus both directly and
indirectly through the plexus on the fin, and finally discharges into the caudal heart. The posterior extremity of
the lateral chordal sinus and its connection with the caudal heart are shown, as also is the cartilaginous knob
which projects forwards from the anterior margin of the median ventral bar of the caudal fin skeleton. Remainder
of caudal fin skeleton not shown.
Trans. Roy. Soc. Edin. VOL. LIV.
Professor FRAXK J. COLE: "A Monograph on the general Morphology of the Myxinoid Fishes."—PLATE I.
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right left
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FIG. 1.
F. J. Cole, del.
Professor FRANK J. COLE : " A Monograph on the general Morphology of the Myxinoid Fishes."—PLATE II.
F. J. Cole, del.
Professor FKAXK J. COLE : " A Monograph on the general Morphology of the Myxinoid Fishes."—PLATE III.
F. J. Cole, del.
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Trans. Roy. Soc. Edin.
VOL. L1V.
Professor FRANK J. COLE: " A Monograph on the general Morphology of the Myxinoid Fishes."—PLATE V.
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F. J. Cole, del.
FIG. 11.

Cole 1926

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( 309 )

A. THYMUS AND PANCREAS.

A transient thymus-like organ was first described in a Marsipobranch in 1894 by

B. PERICARDIAL CAVITY (Fig. 9).

C. T H E ARTERIAL SYSTEM (Fig. l).

D. THE VENOUS SYSTEM.

when complete information as to its development is available. In the meantime, however,

E. RED LYMPHATICS. SUBCUTANEOUS CYCLE.

1. Subcutaneous Sinus (Figs. 4-11, s.c.s.).

2. Subcutaneous Anastomosis (Figs. 3 and 6, s.c.a., s.c.a.'~'").

Connections.—1. Subcutaneous sinus. 2. Hypophysio-velar sinus. 3. Ventral sinus.

4. Dorsal Oral Sinus (Fig. 4, d.o.s.).

5. Hypophysio-velar Sinus (Figs. 4-7, h.v.s., h.v.s.', h.v.s.").

6. Subhypophysial Sinus (Figs. 5, 6, s.h.s.).

7. Ventral Sinus (Figs. 5, 6, 7, vn.s.).

the subcutaneous anastomosis. 2. Subcutaneous sinus (direct wide communication in front).

9. Lingual Sinus (Figs. 2, 6, 7, Ls., Ls.').

10. Subcesophageal Sinus (Figs. 7, 8, s.o.s., s.o.s.').

F. RED LYMPHATICS. PERIBRANCHIAL CYCLE.

1. Cardiac Sinus (Figs. 8, 9, cd.s.).

2. Peribranchial Sinuses (Figs. 2, 8, p.b.s., p.b.s.').

1. Lateral Chordal Sinus (Figs. 3, 7, 8, 9, 10, 11, l.c.s., l.c.s.', l.c.s.").

H. CIRCULATION OF THE BLOOD.

Rostral Dorsal Oral

Posterior Cardinals Subchordal

I think it must be admitted, assuming the statements of fact in this paper to be

There thus exists in Myxine a typical lacunar circulation. In a letter to me KLINCKOWSTROM

(1) ALLEN. Q.J.M.S., 59, 1913.

(10) FAVARO.In Bronn's Tier-Reichs, 1908.

hy. Hyoid arch. obi. Obliquus muscle.

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