Cited by W.B. Ransom, 'On the Cardiac Rhythm of Invertebrata', Journal of Physiology 5 (4-6) (1885), pp. 261-372.
Tags: alcohol, haemotoxylin, osmic acid, gold chloride, picrocarmine
Description:

Explanation of Plate VII, figs. 1-9:

'Fig. 1. Diagram of the Cardiac apparatus and gills of Octopus, with the cardiac and respiratory nerves.
Vc. = Vena Cava. KY. = Kidney Vein. BH.= Branchial Heart. G. = Gill. LA. = Left auricle. RA. = Right auricle. V. = Ventricle. CA. = Cephalic Aorta. Ga. = Genital artery. Va. = Visceral artery. SF. = Veins from sides. P. = Pleural ganglia. S. = Stellate ganglia. Vg. = Visceral nerve. C. = 1st Cardiac Ganglion. Cr. = 2nd Cardiac Ganglion. B. = Branchial Ganglion.

The dotted lines indicate the parts covered by glands.

Fig. 2. a. Muscle fibre of the ventricle of Octopus. (x 500.) Alcohol and Hematoxylin.

Fig. 2. b. Portion of a muscle fibre of the ventricle of Sepiola. (x 500.) Gold chloride.

Fig. 3. a. and 3. b. Plasma cells from the julletion of the kidney and auricle of Pterotrachea. (x 500.) Osmic acid and Picrocarmine.

Fig. 4. Ganglion-cell from the cerebral ganglia of Pterotrachea. (x 500.) Osmic acid and Picrocarmine.

Fig. 5. Plasma cell from the cutis of Pterotrachea. (x 500.) Osmic acid and Picrocarmine.

Fig. 6. Connective-tissue cells; a. from the auricle, b. from the cutis of Pterotrachea. (x 500.) Osmic acid and Picrocarmine.

Fig. 7. Diagram of the branchial and cardiac nerves of Aplysia.

LV., RV. = Left and Right Visceral nerves. Vg. = Visceral ganglion. Gen. = Genital nerve. O. = "Olfactory organ." rn. = Left branch of the nerve to the gill. nb. = Branch to the pericardium. G. = Gill. P. = Pericardium. A. = Auricle. V. = Ventricle. a. = Aorta.

Fig. 8. a. Plasma cell on auricular muscle of Helix.

Fig. 8. b. Plasma cell from the connective tissue membrane between the loops of the genital duct of Helix. (Zeiss. D. Oc. 4.)

Fig. 9. Diagram of the cardiac nerves of Helix.

SO. = Supracesophageal ganglia. Sub. O. = Subcesophageal ganglia. Lv. = Left visceral nerve. Os. = Ovisperm duct. K. = Kidney. P. = Pericardium. A. = Auricle. V. = Ventricle. a. = Aorta.' (338-339)

 

Fig. 1 in text:

'The species chosen as most convenient for study was the common Poulp, Octopus vulgaris, and to it the following pages will be almost exclusively confined.

The cardiac apparatus of Octopus (Fig. 1, Plate VII.) consists of a pre-branchial or venous, and a post-branchial or arterial section. The former collects the blood from the veins and drives it through the gills; the latter receives the aerated blood fromn the gills and propels it through the bodv. The former section may be divided into the following parts:-

Vena Cava.

Kidney Veinis.

Branchial Hearts.

The latter consists of :-

Auricles.

Ventricles.

The Vena Cava (Fig. 1. VC.) is a straight thlin-walled tube, which runs down the ventral side of the liver, parallel to the intestine, towards the heart.

At the level of the front end of the ventricle it divides into two branches, which being covered with glands considered to have a renal function, may be conveniently distinguished as the Kidney Veins (KV.)

Each Kidney Vein, curving out towards a gill, is continued into a bulged oval body known as the Branchial Heart (BH.), their cavities being separated by a pair of valves at the junction. The Branchial Hearts have been usually held to be muscular organs devoted to propelling the blood through the gills; but that this view is not entirely correct will be shewn by the description of their minute structure given below. From each Branchial Heart a thick-walled "branchial artery" leads to the gill (G). The efferent vessels of each gill form a number of veins which soon unite to form a tuLbular auricle (LA. RA.), thin-walled but stouter than the Vena Cava. The two auricles open into the medianly situated, fleshy, somewhat globular ventricle (V), from the cavity of which the blood is prevented from returning by a pair of valves at each auriculo-ventricuilar orifice. 

Three efferent vessels issue from the ventricle, the most important being the great cephalic aorta (CA.), which starting from the posterior dorsal right region of the ventricle curls round to run to the head. Medianly and posteriorly is given off a small genital artery (G a.); and from the anterior ventral edge a small bulb gives rise laterally to a fine artery to each auricle and to a somewlhat larger visceral artery (V a.) medianly to the intestine.' (263-264)

'The whole cardiac system is supplied by a pair of nerves from the pleural ganglia in thie head... Their relations to the neighbouring organs and directions for finding them have been accurately given by Fredericq... Their branches and ganglia need however a more detailed description (Fig. 1). On issuing from the skull, each nerve appears as a double cord from which various small branches are given off. The cords on either side end in a small ganglion lying under the liver, from which go nerves to the body wall and the columnar muscle inserted into the mantle, while a single main trunk is continued downwards towards the heart. Just in front of the auricle it dilates into a ganglion, which may be called the 1st Cardiac Ganglion (C). From this issue a fine nerve to the generative duct, a nerve which enters the auricle, gives off a branch there and then passes through to the ventricle, and lastly a stout nerve which runs dorsally to the auricle down to the branchial heart, where it is connected with a ganglion-the 2nd Cardiac Ganglion (Cr). From this ganglion go nerves to the substance of the branchial heart and a short way into the kidney vein, but the main trunk proceeds to the gill, at the base of which it expands into the Branchial Ganglion (B). No other ganglia are revealed by dissection on the visceral nerves or their branches...

From the pleural ganglia there also runs on either side a stout nerve ending in tlle stellate ganglion (s) and containing the motor fibres for the mantle; so that the gills and heart are connected by nervous structures with the motor organs of respiration.' (267-268)

 

Fig. 2 in text:

'The fibres of the ventricle are of an elongated fusiform shape, with long tails, but are shorter and thicker than those of the auricle. Like the others they have no sarcolemma. Thus far they resemble the ordinary smooth muscle cells of vertebrata, but in addition they possess a fine but regular transverse striation very like that of the branchial heart, the fibres of which they closely resemble. Careful focussing also shows in the fibre indications of a granular core of a different nature to the outer zone. Both of these features are best seen in osmic acid or alcohol preparations (Fig. 2a) [note: This axial column of apparently less differentiated substance has been noticed by most observers of molluscan muscles. Thus Ranvier (Traite technique d'Histologie, p. 851), figures a gold chloride preparation of the retractor muscle of Helix pomatia showing distinctly the "cordon protoplasmique"; Leydig and Kölliker saw it in the buccal muscles of Gasteropoda, and H. Muller and F. Boll in the branchial hearts of Cephalopoda. (See F. Boll, Arch. f. mikr. Anat. Bd. v. Suppl. Hft. p. 28).'].' (266)

'while the ventricular fibres of Eledone resemble those of Octopus, those of Sepia present a bolder striation, and in Sepiola the difference is still greater. In this animal the fibres are much larger, and the average distance between the striations is 3.3m.; the dark bands are very sharp, but narrow; and the margin of the fibre is bulged out opposite each broad clear disc (Fig. 2. b). The central core is much more distinct than in Octopus, occupying about one third of the diameter of the fibre, and staining very deeply with gold chloride, while the outer zone remains clear. It also shows a beaded outline corresponding with the striations. [note: 'Transverse striation has also been noticed - chiefly in hearts or the buccal mass - by many observers, a list of whom is given by Boll (loc. cit.) and by F. Darwin (Journ. of Anat. a. Phys. Vol. x. Part Ii. April 1876, p. 506). Dogiel (Arch. f. mikr. Anat. Bd. 14) has also seen it in various hearts, (Pecten, Anodon, Aplysia, Helix) and Haller (Gegenbaur's Jahrbuch, 1883) in the hearts of Fissurella and Haliotis. Margo (Wien. Sitzusngsb, Bd. 39) has described the black bands in the striated shell muscle of Anodon as doubly refracting.'].' (266 [note 266-267])

 

Figs. 3-5 in text:

'Two kinds of elements however occur [in Pterotrachea] which might be mistaken for nerve cells, and which I believe have been taken for such in other Molluscs. The first of these are large, moderately granular, often roundish or oval cells, with a not very distinct oval nucleus placed near the periphery of the cell, and in which a nucleolus is sometimes visible. A distinct capsule is not present. These cells occur in great abundance on the walls of the rhythmically contractile kidney, and a few are usually to be found in the auricle [note: 'Although in teased preparations of auricle these cells were often found, I am inclined to doubt whether they occur normally in the auricle itself, and to think that in these cases they have appeared from adhering fragments of the closely connected kidney, which owing to the transparency of the tissues it is difficult to cut clearly off at the boundary. In preparations where the auricle was cut through at a distance from the kidney no such cells were found. At the same time it is possible that they may occasionally wander from the kidney to the auricle. They are also found in the pericardial wall.']. I have not observed them in the ventricle. Although the approximately oval form (Fig. 3 (a) Pl.VII.) is perhaps the commonest shape, yet it is by no means constant. In many cells short blunt processes or pseudo-podia are seen (b) which suggest the power of amoeboid movement, and occasionally a single such process may occur, causing some resemblance to a ganglion cell with the stump of a nerve fibre. The number and variations of the processes however oppose this idea, and a comparison of such a cell with a true ganglion cell (Fig. 4) at once shews a number of differences in appearance. Further these cells are never found in connection by fibres with either nerve or muscle, and are shewn to be very loosely applied to the tissues by the fact that in teased preparations large numbers of them become entirely detached and float like the blood corpuscles in the fluid in which they are mounted. Lastly, they occur not only in the parts mentioned, but in the connective tissue all over the body, as is well seen in preparations of the cutis, where they (Fig. 5) abound. These facts point to the conclusion that they are wandering connective tissue cells; and we may almost certainly identify them with the "plasma cells" which have been shewn by Brock [note: 'J. Brock. " Untersuch. ii. d. interstitiellen Bindesubstafiz der Mollusken." Zeit. f. Wios. Zool. Bd. 39. 1883.'] to be so characteristic of the connective tissue of some other Molluscs (Aplysiadae, Pulmonatae).' (321-322)

 

Fig. 6 in text:

'The second form of element which might be considered nervous [in Pterotrachea] is of much smaller size and more delicate structure. These cells again I found only where connective tissue was abundant. They are granular branched cells with a usually indistinct nucleus. Occasionally the branches may be reduced to two, and there then appears some resemblance to a small bipolar ganglion cell (Fig. 6 (a)). These cells however are nothing more than the ordinary connective tissue cells, which give off processes to form fibres and are found everywhere throughout the connective tissue (Fig. 6 (b)). A comparison of the two figures makes plain the identity of those of the heart with those of the cutis. These also are described and figured by Brock in the connective tissue of Aplysia as stellate connective tissue cells, and very similar cells by Haller as occurring 'in small quantity in the auricle of Fissurella and as forming a second type of ganglion cell.' (323)

 

Fig. 7 in text:

'Apart from the improbability of the existence of apolar ganglion-cells in the heart, the incorrectness of Dogiel's drawing of the heart and the adjacent nerves causes great doubt as to the accuracy of his interpretation of these cells.

In his figure (Taf. V. a. Fig. 15) of the "branchial" ("visceral," Spengel) ganalion he entirely overlooks the olfactory organ to which the right nerve runs, and he represents the left nerve as of equal thickness and as running straight to the auricle. But instead of such a short thick left nerve, there come off from the left half of the double "visceral " ganglion two distinct long nerves. Of these one supplies the generative duct, while the other runs on to the gill. Here it divides into two, the left one running on straight towards the hind end of the gill while the right curls round anteriorly and appears to end in the pericardium near the origin of the auricle (Fig. 7).' (325)

 

Fig. 8 in text:

'In the auricle of the Snail may be found upon the muscle bundles [note: 'Both the auricle and ventricle are formed of a meshwork of bundles of striated muscle fibres, that of the ventricle being denser and thicker.'] a number of plasma cells which are, there is little doubt, identical with Dogiel's "apolar!" ganglion cells. Some of these cells of an oval or pear-shaped form present a considerable resemblance to ganglion cells, and if only these were observed might be considered such. But when others of varying shapes, and others in process of division are seen, such an idea becomes untenable; and when the striking similarity of these cells with the plasma cells of the connective tissue is noticed, no doubt can remain of the identity of the two (Fig. 8).' The various stages of division also which are met with, from a group of two or three contiguous cells to a nest of closely packed small ones, point clearly to the true nature of these elements. True ganglion cells I believe do not exist in the Snail's heart.' (327)

 

Fig. 9 in text:

'The heart... receives nerves both at the auricular and aortic ends [note: 'A similar double nerve supply has been described by Haller, loc. cit. as existing in Muricidae and Fissurella.']. (Fig. 9.) The discovery of this nerve to the heart at once necessitated a reconsideration of the results obtained by previous workers, and the question of its function became important.' (328)