Muscular System.
Attachment of the Body to the Shell.—The prosoma which carries the posterior thoracic segments, and in appearance the mouth, is the only part of the body which is attached to the general covering (Pl. [25], fig. [1]), namely, to the opercular valves. Except through the continuity of the lining membranes of the sack, the body lies free within the walls of the shell. The area of attachment (shown by a sinuous broken line round a and b) extends from about the middle of the two scuta down to their basal margins. As these valves lie obliquely across the orifice of the shell, the animal’s body comes to be suspended almost in the middle of the sack. The two scuta, as we have seen, have the power of opening and shutting a little; and are brought together by the adductor scutorum muscle (a), which is generally very powerful. The body is attached to these valves, round and beneath the adductor, so as to hide it until one of the valves be removed. The attachment is chiefly effected by three pairs of widely expanded, superficial muscles, two pairs of which are spread over the flanks of the prosoma, and the third pair over its rounded (properly dorsal) surface, which lies close to the rostral compartment (A, fig. [1]) of the shell. I should have stated, that my chief examination of the attachment of the body to the scutal valves, has been made on [Coronula balænaris], and less closely on [Balanus tintinnabulum]. Within these three pairs of superficial muscles, there are (besides the adductor) no less than five other pairs; of these one long pair is attached at one end to the basal margin of the labrum (e), and at the other end, to the under side, near to the basal margin of the scuta: two other, shorter, parallel pairs of muscles are attached at one end to the interspace of membrane between the basal edge of the labrum and the adductor scutorum muscle, and at the other end, to the under side of the scuta, above the attachment of the first pair: the fourth and shortest pair curls close under the adductor, and is there attached at both ends beneath it. The action of these four pairs of muscles must be to draw back, from the orifice of the shell, the mouth, and that interspace of body between the basal margin of the labrum and the adductor muscle. This movement I saw in living specimens. The last and fifth pair of muscles is small, but of considerable length; it is a diverging pair, attached at the converging end, above and exteriorly to the adductor muscle; and at the diverging end, low down on the under side of the scuta; I am very doubtful regarding the function of this pair. Altogether we have seen that round and within the fleshy pedicel, by which the body is attached to the scuta, there are no less than eight pairs of muscles. The central space between these muscles is hollow, and here many lacunal channels seem to converge. These muscles receive nerves from the supra-œsophageal ganglions. The interspace above alluded to, between the basal edge of the labrum and the adductor scutorum muscle, occupies a very different position according as the animal’s body is protruded as far as it can be, or is retracted. It is homologically part of the third cephalic segment; and consequently the mouth ought to have stood posteriorly (i. e. above, in the position figured in Pl. [25], fig. [1]) to this interspace; yet, in fact, when the animal is retracted within its shell, the mouth usually lies almost directly beneath this interspace and the adductor scutorum muscle.
Besides these muscles of attachment, the prosoma is furnished with several other muscles. There are superficial muscles running up towards the basal margin of the sides of the mouth; and other deeper muscles, to which, I presume, the movements of the mouth, as a whole, are due. The muscles moving the gnathites do not, as far as I could make out, extend beneath the basal edge of the mouth. There are, also, powerful muscles giving movement to the basal segments of the pedicel of the first pair of cirri. Again, there are superficial muscles running to the next succeeding thoracic segment; the anterior ends of which are separated by a clear interspace from the lower ends of the above-described superficial muscles, by which the prosoma is attached to the scuta. On each flank, moreover, but more deeply imbedded, are the long flexor and extensor muscles, presently to be described, running to the five posterior thoracic segments. The last muscle which I need here mention, is a deep-seated diverging pair, attached near the upper end of the stomach, on its ventral surface, and diverging from this point to the sides of the prosoma high up beneath the mouth. The probable action of this pair, as well as of the three superficial pairs of muscles by which the body is attached to the scuta, is to draw up the whole prosoma towards or from the orifice; and likewise to contract it firmly, so as to serve as a fulcrum for the movements of the five posterior thoracic segments, together with the cirri, which they carry.
The muscles of these five thoracic segments are numerous and powerful; they are also complicated, chiefly owing to the segments on their dorsal and dorso-lateral surfaces being driven, like wedges, one into the other. As far as I could make out, there are on each side three, superficial, dorso-lateral and lateral muscles (generally, if not always, destitute of striæ), which do not cross the articulations, but extend merely from articulation to articulation; and of which the function can be only to contract each separate segment, and consequently to open out the intermediate infolded articulations; the effect of this would be to separate slightly the cirri from each other,—more especially those borne on the two or three anterior segments, between which the infolded articulations are deeper or broader. There are other more deeply imbedded, powerful, long, dorso-lateral extensor, and ventri-lateral flexor muscles, attached at one end within the flanks of the prosoma, and at the other end to the successive segments of the thorax. The action of the former is to straighten and stretch out the thorax; of the latter, or ventri-lateral muscles, to retract it. In tracing these muscles, a fascia could be seen to become attached to a segment, and then this same fascia would run on to the next succeeding segment: the effect of this must be, that each segment can be retracted and protracted either from the prosoma as a fulcrum, or from the antecedent segment as a fulcrum: we have, also, seen that each segment can, by the agency of the superficial, non-striated muscles, contract itself. Hence these thoracic segments are capable of diverse movements, as was very evident when the shell of a living specimen was opened. By one movement in common, the whole five posterior segments could be drawn back, so as to become even partly imbedded in the prosoma: lateral, twisting or wriggling movements were also quite distinct: the three posterior segments seemed to be capable of less independent movements than the anterior segments; and I observed that the more powerful flexor and extensor muscles did not run into these three posterior segments. The cirri, of course, partake of the movements of the thorax; and in watching, in an uninjured specimen, the alternate, protruding, gracefully sweeping and retracting movements of the posterior pairs of cirri, it was evident that the thorax was the chief agent in their movement. Besides the muscles now mentioned, there are some immediately to be noticed, which extend from within the thoracic segments to within the pedicels of the cirri.
Movements and Muscles of the Cirri.
Although the cirri have not been described, it will be most convenient here to treat shortly of their muscles. Each cirrus consists of a pedicel, having a long basal and a short upper segment, supporting two multiarticulate rami. The lower segment of the pedicel can be drawn forward by an adductor muscle, attached low down within this segment, and crossing at right angles (at least in the case of the anterior cirri) the corresponding muscle of the opposed cirrus, on the central, ventral surface of the thorax. This segment can also be drawn back by a muscle springing from the dorso-lateral surface of the thorax, and running only a little way within the segment: I am far from sure that the lower segment does not possess other muscles. The short upper segment of the pedicel can be moved backwards and forwards, as I saw in living specimens, independently of the lower segment; this movement being best seen in the anterior cirri, which are much more often moved independently of each other than are the posterior cirri. The rami are capable, I believe, of being moved backwards and forwards as a whole, by the movement of the few lower segments, which are generally more or less confluent. They can, also, be curled up and uncurled by the combined movement of each separate segment. The uncurling seems to separate the two rami a little laterally. Each ramus, at least in the two or three anterior pairs, can be moved to a certain extent, independently of the other ramus of the same cirrus; and the few terminal segments, either of both rami or of one ramus, are often a little moved and curled (and this is especially the case with the long anterior ramus of the first pair), without the lower segments or the pedicel being moved.
The flexor and extensor muscles, which, as I believe, move the upper segment of the pedicel (a and b, Pl. [29], fig. [1]), are attached at their upper ends to its basal margin, and are thus enabled to draw it a little way down within the lower segment, and so move it. The short flexor muscle (c), which is attached at its lower end within the upper segment of the pedicel, and the longer extensor (d), also, attached within this same lower segment, serve, I believe, to move the lower, partially confluent segments of each ramus as a whole. In the case of these muscles, and of those last mentioned, I am surprised that the extensors (b) and (d) are not attached nearer to the exterior and dorsal surface. Other muscles (e, f) attached at their lower ends within the upper segment of the pedicel, run up each of the two rami to their tips, with some of the fasciæ terminating within each segment: of these muscles, the outer one (f, f) appears to be the extensor, and the inner one (e, e) the flexor. But besides these, there are other short flexor muscles (g, g) which run on the anterior face,[31] from segment to segment, serving to pull the front edge of one segment within the edge of the next lower segment. These muscles differ much in plainness in the several genera: they are very distinct in [Coronula]. In some specimens of this genus, a few of the articulations between the basal segments of the rami having been obliterated, the short muscles (g, g) running from articulation to articulation were absent, and their presence and nature in the upper segments thus rendered the plainer. The muscular system in the several pairs of cirri seems to be the same, with the exception of the first pair, in which the muscle answering, as I suppose, to (a), namely, the flexor of the upper segment of the pedicel, is much spread out at its lower end, and is there attached to the exterior surface of the lower segment.
[31] For a considerable time I thought that there were muscles going to the spines, especially to those which arise from the upper dorsal edge of each segment; but I have since ascertained that these are the cases within which new spines, with their lower ends doubled like the fingers of a glove hastily pulled off, are in process of formation.
The backward and forward movements of the segments, both in the rami and in the pedicels of the cirri, are apparently effected, as already noticed, by the outer or inner (as the movement may be) basal edge of one segment being drawn a little way down within the next succeeding lower segment. If, at the same time, both the inner and outer margins of all the segments were drawn one within the other, the whole limb would necessarily be shortened; and I distinctly saw a shortening action, with very slight movement in any other direction, in the first and second pairs of cirri; and I think it almost certain that this movement might be performed by the other cirri. If I correctly understand a statement of Milne Edwards,[32] this is an important fact, as he asserts that only the higher Crustaceans possess the power of shortening their limbs.
[32] ‘Annales des Sciences Naturelles,’ tom. xviii, 1852, p. 121.