Part 16

The cement-glands, the trunk, and the ducts, except the two, three, or even four last-formed ones, are all filled with an apparently solid thread of transparent, brownish cement, differing in no respect from the cement under the central parts of the basal membrane. In one instance, in which the last-formed pair of glands and ducts had apparently been only just developed, they were so perfectly transparent that I could distinguish them only under certain lights, and I could not perceive any contents. The last-formed glands and ducts always appear very delicate, and include a tube of very delicate tissue, containing more or less of granular matter. The next succeeding pair of glands and ducts are always more opaque, and contain much more granular matter; which, in the next, or next but one, may be seen passing into the state of pale brown, transparent chitine. I have seen some most distinct instances, in which, in the same duct, the part towards the centre of the basis was filled with homogeneous cement, and the part towards the circumference with still plainly granular matter. In the successive circular slips of cement-tissue, lying attached under the circumferential slip and under the two or three succeeding slips of basal membrane, an exactly analogous series of changes is presented; as indeed might have been expected, as the slips of cement are absolutely continuous with the contents of the ducts. Moreover, if a vertical section be made across one of the last-formed slips of cement, it may sometimes be seen to be apparently in the act of separating into layers, with the lower layers more pulpy, elastic, and white than the upper layers, which are less coherent, and show as yet even still less the character of cement. The cement under the circumferential and adjoining slips, often presents a peculiar wrinkled appearance, in lines conformable with the outline of the basal membrane; but I do not believe that these are real wrinkles, though so like them; they seem to consist of sinuous threads, longer or shorter, sometimes slightly branched, crossing and inter-joined, and composed apparently of faintly coloured cement. I suspect that these threads are formed by the union and subsequent drawing out of aggregations of that matter, which within the ducts is first granular, and then changes into cement; for at the orifices of the ducts these wrinkled threads sweep outwards in curved lines on both sides. The cement in these early stages adheres, with very little force, to the basal membrane; and with only a little more force to the underlying layers of cement; in fact, till it assumes the brown translucent appearance, like solid glue, it hardly seems to act as cement.

How the cement reaches the skin of the whale, will be best understood by referring to the sectional diagram (Pl. 28, fig. 1 _b_). When the circumferential slip of membrane (_b_) splits, a new circumferential slip will be formed over it, together with new cement-glands and ducts, and cement (_z_, _z_) will issue from four new orifices, and will extend on both sides of these orifices, till the ends meet and become united, thus forming a narrow, 18-sided, continuous, new slip of cement, with 18 spokes proceeding from it. I have not noticed lines of union in the cement of any one slip; but the matter forming each slip, certainly has proceeded from four distinct orifices. Seeing how perfectly successive layers of cement often become blended together, lines of union or junction, could hardly be expected to be preserved in the same individual layer. When the circumferential slip of basal membrane splits, the underlying slip of cement, which we will call A, does not split, but becomes stretched, so that the newly formed slip of cement, which we will call B, does not reach the skin of the whale. As the new circumferential slip of basal membranes goes on increasing in width, A continues to be stretched, but does not split, till at least another circumferential slip of basal membrane has been formed and has been split, and till B has been also stretched. By this time, the cement-tissue A has assumed its normal structure, and has the power of adhering to the whale's skin, which it has now reached, owing to the splitting of the under and older slips of cement. At the next period of growth, A itself will split, and B will touch the whale's skin and adhere to it; and this, also, will ultimately split. It results from this action, that the cement has a stretched, and sometimes even a fibrous appearance, with the lower edges of the layers, of which each slip of cement is formed, thinning out. I have before stated, that the two or three last-formed slips of basal membrane are formed at first too narrow, and apparently have to be dragged outwards, over each other; and it is perhaps owing to this circumstance, and to globules of cement having first adhered to the under surface of the slips of basal membrane, that these slips are studded beneath with parallel little vermiform bodies, sometimes of considerable length, and furnished with heads, all directed outwards. These tapering, vermiform bodies have a considerable resemblance to the threads before mentioned, which give the wrinkled, concentric appearance to the newly-formed layers of cement, and have probably a closely analogous origin: in one case, indeed, it appeared as if some of these concentric threads were in process of being drawn out at right angles to their original course. Lastly, it should be observed, that as the exterior half of the membrane of the circumferential slip, after each splitting, is dragged down, and thus comes to invest the outer surface of the wall of the shell (the wall not being represented in the diagram, but standing where the letter (_p_) stands), so it must be with the cement, which thus likewise comes, in an unusual manner, to invest the outer surface of the folded walls of the shell, and attaches them to the skin of the whale,--which latter is always growing upwards, and tending to bury the shell.

_Platylepas decorata._--This genus is closely allied to Coronula, and the cementing apparatus is essentially similar. In one specimen, I counted no less than forty-nine slips of basal membrane, each of which, of course, had its pair of cement-glands, and each of the latter its two ducts. The glands consists of a transverse enlargement of the trunk, as in the early stages of Coronula. Neither the glands nor the duct, when old, become filled up with cement, but only the main-trunk. The ducts are very delicate and thin; the larger ones being only 3/10,000 of an inch in diameter. The glands stand some way apart on the two cement trunks; and the latter, instead of being straight as in Coronula, are deeply serpentine; the glands are formed on each bend, so that, though all on one side are connected on the same trunk, they form a double row on each side of the basal membrane. The basal membrane (in the centre of which I distinctly saw the antennæ of the pupa) has six deep bays or excisions, corresponding with the midribs (see Pl. 17, 1 _a_, 1 _d_) of the six compartments; and the two ducts from each gland, on the right and left sides, debouch at the heads of the four lateral excisions, exactly opposite the midribs of the lateral and carino-lateral compartments. The later-formed glands, owing to all of them being situated some way apart from each other on the two cement-trunks, lie further from the centre of the basis than do the orifices of their ducts; hence the later-formed ducts are directed a little backwards, or from near the circumference towards the heads of the deep excisions.

_Tubicinella._--The cementing apparatus is here less symmetrical; but this, I believe, is chiefly owing to the basal membrane being formed of successively larger discs (not slips) of membrane, thrown down not quite concentrically one over the other; each new disc of membrane seems to cover the last-formed cement-glands and ducts; and there are as many ducts and glands as there are discs of membrane, all adhering together. In one specimen, it appeared that normally there were four sets of cement-ducts, as in the allied genera of Coronula and Platylepas; but in other specimens, the ducts were distributed very irregularly. In one case the two cement-trunks extended parallel and close together, one of them terminating long before the other. I have given a figure (Pl. 28, fig. 3) of three of the cement-glands, removed from the basal membrane, together with their ducts. The cement-trunk (_f f_) seemed to be a little enlarged, and to be crossed by septa, as it entered the glands (_h_), but I could not make out this structure clearly enough to be represented. Whilst young, the cement-glands stand some little way apart from each other; and in the figure given of some of the last-formed glands, they are hardly separate enough. Each gland gives out obliquely, on one side, a cement-duct (_c_) which I traced in several cases to the margin of one of the discs of basal membrane, where cement issued from it; and on the opposite side, a tapering spur (_b_), varying in length, which may be called, and I believe really is, a rudimentary duct. Of these spurs we shall meet many instances in other genera. The duct (_c_) and the spur (_b_), close to where they entered the gland, in some specimens gave off, at about right angles, short blunt points, or rudimentary branches. This duct and spur correspond, I believe, with the two ducts in Coronula; but besides these, a duct (_a_) is given off from one end of the gland, from the surface opposite to that at which the cement-trunk enters. This duct (_a_) is very singular, from always forming a loop (_a′_), with two spurs projecting from it: these two spurs occasionally spring from a common point: I have seen nothing like this structure in any other Cirripede. This duct (_a_) runs, like the duct (_c_), to the margin of its own disc of basal membrane, where it debouches. Besides these ducts, in the best specimen which I examined, there were two other sets of ducts, which were slightly zig-zag, and at each angular bend, a mere knob or point, or at most a short branch, was given off; but this branch seemed never to run to the margin of the basal membrane or to give out cement; whereas the main branch did give out cement. I was not able to trace these ducts to their glands. In these zig-zag ducts, and in the rudimentary points sometimes observed at the base of the duct (_c_), and likewise at the base of the spur (_b_), we see the first indication of that tendency to bifurcation, so strongly characteristic of the cement-ducts in all the genera, excepting those already described, which are allied to Coronula.

_Chelonobia patula._--The cementing apparatus is here chiefly remarkable for the thinness and straightness of the main trunk, (_f f_, Pl. 28, fig. 2), and from the great distance at which the glands stand apart; had another gland been drawn, it would, on the scale here used, have stood exactly under the two upper, (_c′ c′_) in fig. 1 _c_. We here see that the trunk (_f_), before entering the gland (_h_), has an enlarged portion (_g_); this, I suspect, is a very general structure. Each gland gives out, on opposite sides, two ducts (_a a_, _b b_), larger even than the main trunk; and these ducts bifurcate repeatedly, and inosculate. By this inosculation it is not improbable that all four ducts, proceeding from the two glands of the same age, may be connected together; certainly the bifurcating branches from the same duct thus become repeatedly connected. For the first two or three bifurcations the ducts decrease very little or not at all in diameter; but nearer the circumference they become smaller. The ducts, also, proceeding from the younger and smaller glands, are, of course, proportionably smaller. In one case I was able to count four bifurcations in the duct between the gland and the edge of the basal membrane. It follows from this structure, that the basal membrane, at each period of growth, is cemented down by cement issuing from several orifices; but we shall presently find that in other genera the cement proceeds from many more orifices. In fig. 2 there is represented, by the aid of the camera, a small portion (from the outer (_a_) to the outer (_b_) being 12/100ths of an inch in length) of the basal membrane, with all the several cement-ducts adhering to it, which I could distinguish, and drawn of their proper relative sizes; this figure also shows some of the bifurcations, but no inosculation happened to be included in the space here given; the basal membrane itself has not been represented. In taking a view of a considerable portion of the basal membrane, especially towards the circumference, some parallelism in the branches could be perceived; one set of branches tending to run in the direction of the ray of the circle, and the other set in the line of the circumference.

_Elminius Kingii._--The cement-glands here resemble those of Chelonobia, but the trunk does not seem to be enlarged before entering the gland. The glands are situated rather far apart; and the chief peculiarity is, that the trunk connecting the glands is as tortuous as the track of a worm. Each gland gives out two ducts, which bifurcate repeatedly, and often inosculate, making, in parts, an hexagonal mesh-work: some of the branches do not debouch on the basal membrane, but terminate in blunt points. So numerous are the ducts, that the basal membrane may be compared to pieces of paper with the fine fibrous branching roots of some plant dried and heaped on it. Some of these ducts are very regularly jointed, and resemble a conferva,--an appearance which I believe is owing to divisions in the contained cement; other ducts are partially marked by little wrinkles, as presently to be described under Balanus. The cement, instead of, as heretofore, invariably forming a slip round and beneath the circumference of the basal membrane, here often forms little, independent, circular, and irregularly-shaped discs, each of which has issued from a single orifice. I may here add that in two species of _Tetraclita_ I saw the cement-ducts repeatedly bifurcating, with some of the branches inosculating, as in Elminius and Chelonobia.

In _Balanus balanoides_, which, like all the Cirripedes hitherto mentioned, has a membranous basis, I could only make out an amazing number of bifurcating and inosculating cement-ducts, of very various diameters. The cement-tissue, on the under side of the basal membrane, generally consisted of little circular discs, on an average from 2 to 5/1000ths of an inch in diameter; but there were also globules and short tortuous threads of cement. In some very much elongated and crowded specimens--in which, during the downward growth of the walls, the basal membrane had ceased to reach the surface of attachment, and being thus suspended had become, as viewed from the outside, deeply concave--the cement had apparently continued to try to reach the rock, and now hung down in the form of two thickish roots, some tenths of an inch in length. These roots were round, and tapered either to a fine or blunt point; one was doubled on itself, and so had become united; in the other, I could perceive five layers or sheaths of the cement-tissue, one within the other; the innermost of these layers, which once, no doubt, formed the outside surface of the root, was only a quarter of its present length.

In _Balanus tintinnabulum_, the basis is calcareous: when its upper surface is cleaned, dried, and examined under a good light, the numerous larger cement-ducts can be seen, even by the naked eye, or under a weak lens, and present an elegant appearance. These larger ducts run in parallel lines from the two chains of glands towards the circumference. They are all encrusted with calcareous matter, and in the more central parts are hidden under it; at each period of growth, when the basis is added to round the circumference, it would appear that a layer of excessive tenuity of shell is thrown down over the whole surface, just in the same way as in Tubicinella, at each period, a new and larger disc of membrane was thrown down over the pre-existing membranes with their cementing apparatus. The cement-glands, in the middle of the basal plate, seem often to give rise to small abnormal depositions of calcareous matter. When the basis (it is best to take a young specimen) is slowly dissolved in acid, all the cementing apparatus is left uninjured, adhering to the delicate tissue which before existed in a calcified condition. Near the middle I saw the two antennæ of the pupa; and from them the two cement-trunks extended about half-way towards the circumference. These two chains of glands are often placed very irregularly, but they tend to form, as in Coronula, a large angle, open towards the rostral end of the shell. The glands, close to the old antennæ, commence abruptly, of rather large size: the later-formed glands, with their ducts, are in regular order larger than the younger ones, and stand much closer together. After immersion in acid all the glands and ducts appeared empty, instead of the older ones being, as in Coronula, filled with cement. In one case I counted on each trunk twenty-five glands, besides some smaller obscure ones close to the centre.

In Pl. 28, fig. 4 _b_, I have given a drawing of two of the cement glands: the cement-trunk (_f f_) is smooth and apparently cylindrical: it becomes enlarged (at _g_) before entering the gland: it seems even to be prolonged across the gland under the form of a narrow bar (not represented), which apparently serves to keep the two ends of the trunk, on the two sides of the gland, in their proper relative places and distances. The gland itself is an elongated bag (_h_), which properly lies exactly over the enlarged portion (_g_) of the trunk, but in the drawing has been purposely displaced: it gives rise, in the later-formed glands, to a sort of neck (see the upper gland), which is either so long as to deserve rather to be called a duct and which soon bifurcates, or is quite short (see the lower gland) and gives rise to two separate ducts. On the opposite side of these glands, there is a spur (_m_), of greater or shorter length, which is evidently a rudimentary duct, for in the younger glands it existed as a perfect duct. Moreover, the first-mentioned duct often gives off branches (_t′_), having an exactly similar appearance with the spur (_m_). The membrane of which the cement-trunk (_f_), with the enlargements (_g_), is composed, is smooth, but that of the glands and of all the ducts, presents a very peculiar appearance, which at first would be called scaled, but more properly perhaps notched,--each notch being apparently formed by a line of thickened membrane, extending obliquely round only a short portion of the tube, and indenting it. The ducts, which I measured, were between 1/3000th and 4/3000ths of an inch in diameter.

In fig. 4 _a_, I have given a drawing of the two chains of glands, but with only those ducts figured which proceeded from the last-formed pair of glands. The specimen here drawn was old; and it is rare to find the structure of the ducts so simple. From both glands[72] a neck or thick duct arises, which soon bifurcates; one branch runs direct into the circumferential duct, and the other (_t_) bifurcates again; of the latter, one branch unites with its fellow from the opposite gland, and then forming a single duct (_t′_) enters, as do the two other branches, the circumferential duct. Thus, into the latter, five main ducts enter: the position of their points of entrance, with respect to the shell, varies considerably; but I think the five points tend to face the middle of the rostrum, and middle of the two lateral compartments on each side. In some other specimens, in which the ducts were nearly as simple, I observed that the neck or main duct at once divided into three branches, instead of into two, with one soon bifurcating; and on one side a rudimentary branch or spur was given off (above _t_), indicating a tendency to an additional bifurcation. In the later-formed glands, the ducts proceed only from the outer sides and form the ends of the glands furthest from the centre; but in the earlier-formed and smaller glands of the same individual, other ducts proceed from the inner sides, where in the older glands the spurs (_m_) are situated: moreover, in the younger glands, all the ducts bifurcate much oftener (how often I was not able to ascertain), before entering the circumferential duct; many of the branches, however, terminating in spur-like points. Now if we imagine twenty or thirty repetitions of the ducts given in fig. 4 _a_, (independently of the greater complication of the ducts of the younger glands), each a very little smaller than the other, and placed, with the main branches parallel, one over and within the other, we shall gain some insight into the wonderfully complicated structure of the cementing apparatus in this and many other species of Balanus.

[72] It should be observed that fig. 4 _b_ ought to have been drawn with its present upper end downwards, to make it correspond in position with fig. 4 _a_.