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As mentioned above, Ludwig has proved that the paxillae develop in
the life-history of the individual out of ordinary plates, the axis of the
paxilla representing the plate.

Order V. Forcipulata.
This order, which includes the most highly developed members of
the class Asteroidea, is at once distinguished by the possession of
forcipulate pedicellariae which, as we have seen, possess a well-
marked basal piece with which the two plates articulate. The
pedicellariae are consequently sharply marked off from the spinelets,
and no intermediate forms occur. The first conjoined adambulacrals,
which in other orders form the "teeth" or mouth-angles, do not here
project beyond the first pairs of ambulacral plates.

Fam. 1. Asteriidae.—Forcipulata in which the tube-feet are

apparently arranged in four rows. Aboral skeleton a loose reticulum.

The general features of the family Asteriidae have been explained in

the description of Asterias rubens (p. 432). There are five well-
marked species of the genus found on the British coasts. Of these A.
glacialis is found chiefly in the south-western parts of the English
Channel. It is a large Starfish of a purplish-grey colour, with large
spines surrounded by cushions of pedicellariae arranged in one or
two rows down each arm. A. muelleri resembles the foregoing
species, but is of much smaller size, and is further distinguished by
having straight pedicellariae in the neighbourhood of the ambulacral
groove only. It is found on the east coast of Scotland, and carries its
comparatively large eggs about with it until development is
completed. A. rubens is the commonest species, and is found on
both east and west coasts. Its colour is a bright orange, but varies to
almost a straw colour. It is at once distinguished from the foregoing
species by the spines of the dorsal surface, which are small and
numerous, an irregular line of somewhat larger ones being
sometimes seen down the centre of each arm. A. murrayi is a
peculiar species restricted to the west coast of Scotland and Ireland.
It has flattened arms, with vertical sides, and only three rows of small
spines on the dorsal surface. It is of a violet colour. A. hispida is also
a western species. It is a small Starfish with short stout arms; there
are no straight pedicellariae, and only a few sharp spines on the
dorsal surface.

On the eastern coast of North America there are several species of

Asterias, of which the most noteworthy is the 6-rayed A. polaris of
the Gulf of St. Lawrence. This species exhibits a marvellous range of
colour-variation, ranging from bluish-violet through purple to red and
straw-coloured. This variation seems to show that colour, as such, is
of no importance to the animal, but probably depends on some
compound of slightly varying composition which is being carried by
the amoebocytes towards the exterior. On the Pacific coast there is a
rich fauna of Starfish, among which we may mention as members of
this family Asterias ochracea, a large violet species, so strong that it
requires a severe wrench to detach it from the rock, and Pycnopodia
with twenty-two arms.

Fam. 2. Heliasteridae.—Forcipulata allied to the Asteriidae, but with

very numerous arms and double interradial septa. Heliaster.

Fam. 3. Zoroasteridae.—Forcipulata with the tube-feet in four rows

at the base of the arm, in two rows at the tip. Aboral skeleton of
almost contiguous plates bearing small spines or flattened scales.
Zoroaster, Pholidaster.

Fam. 4. Stichasteridae.—Forcipulata with the tube-feet in four rows.

Aboral skeleton of almost contiguous plates covered with granules.
Stichaster, Tarsaster.

The Stichasteridae and Zoroasteridae have acquired a superficial

resemblance to some of the long-armed Valvata, from which they are
at once distinguished by their pedicellariae. It would be exceedingly
interesting if more could be found out concerning the normal
environment of these animals; it might then be possible to discover
what is the cause of the assumption of this uniform mail of plates.

Fam. 5. Pedicellasteridae.—Forcipulata with two rows of tube-feet.

The aboral skeleton bears projecting spines surrounded by cushions
of straight pedicellariae. Pedicellaster, Coronaster.

Fam. 6. Brisingidae.—Forcipulata with numerous arms and only

two rows of tube-feet. Aboral skeleton largely rudimentary and
confined to the base of the arms. The small blunt spines are
contained in sacs of skin covered with pedicellariae.

The Brisingidae, including Brisinga and Odinia, are a very

remarkable family, chiefly on account of the smallness of the disc
and of the extraordinary length of the arms. The arms have what we
must consider to have been the primitive arrangement, since there is
no lateral adhesion between them, and interbrachial septa are
consequently entirely absent. The reduction of the skeleton is a very
marked peculiarity and, like the tendency to the reduction of the
skeleton of deep-sea fish, may stand in some relation to the great
pressure under which the animals live.

Fig. 205.—Aboral view of Odinia. × ⅔. (After Perrier.)

 Fossil Asteroidea.
The Asteroidea occur somewhat plentifully as fossils. In the Lower
Jurassic Asterias, Astropecten, Luidia, Solaster, and Goniaster have
already made their appearance. In the Cretaceous Pentaceros
appears. In the older rocks occur a number of forms of different
character from any now existing. Of these Aspidosoma (Fig. 206),
with short lancet-shaped arms sharply distinguished from the disc
and continued along its under surface, seems to be intermediate
between Asteroidea and Ophiuroidea. The skeleton of the arm is
composed of alternating ambulacral ossicles bordered by
adambulacral ossicles, which are at the same time marginals and
sharply distinguished from the marginals forming the edge of the
disc. Palaeaster, on the other hand, is a true Asteroid; there are
marginals distinct from the adambulacrals, but the disc is reduced to
its smallest dimensions, there being only one plate on the ventral
side of each interradius. There are a number of genera
(Palaeocoma, for instance) with a large disc and very short arms and
very shallow ambulacral grooves; all have alternating ambulacral
plates. Some genera appear to have had the madreporite on the
ventral surface of an interradius. On the other hand, in the Devonian
occurs Xenaster, which was a fairly normal Asteroid, with pavement-
like marginals, deep ambulacral grooves, and broad arms.

Fig. 206.—Three views of Aspidosoma, a fossil Asteroid. A, oral view; B, aboral

view of one arm; C, enlarged view of a portion of the ambulacral groove.
adamb, Adambulacral plate; amb, ambulacral plate; marg, marginal plate;
pod, aperture for extension of tube-foot.

Thus it will be seen that already in Jurassic times the three orders,
Forcipulata, Paxillosa, and Spinulosa were differentiated from each
other, but how these are related to the older Palaeozoic forms it is at
present impossible to say.

CHAPTER XVII

ECHINODERMATA (CONTINUED): OPHIUROIDEA = BRITTLE STARS

CLASS II. OPHIUROIDEA

The second class of Eleutherozoa are familiarly known as "Brittle
Stars," on account of their tendency, when seized, to escape by
snapping off an arm, although this habit is by no means confined to
them, but is shared in a marked degree by many Asteroidea, such as
Luidia, for instance. Like the Asteroidea, they are "starfish," that is to
say, they consist of a disc and of arms radiating from it; but the
scientific name Ophiuroidea really expresses the great dominating
feature of their organisation. Literally it signifies "Snake-tail" (ὄφις,
snake; οὐρά, tail), and thus vividly describes the wriggling, writhing
movements of the long thin arms, by means of which the Ophiuroid
climbs in and out of the crevices between the stones and gravel in
which it lives. This feature, viz. the effecting of movement by means
of muscular jerks of the arms, instead of by the slow protrusion and
retraction of the tube-feet, is the key to the understanding of most of
the points wherein the Brittle Stars differ from the true Starfish.

Asteroidea and Ophiuroidea agree in the common ground-plan of

their structure, that is, they both possess arms; but the most obvious
difference in their outer appearance is that whereas in Asteroidea
the arms merge insensibly into the disc, in Ophiuroidea the disc is
circular in outline and is sharply marked off from the arms. Closer
inspection shows that in the Ophiuroid the arms are continued
inwards along grooves, which run on the under surface of the disc,
and that they finally coalesce to form a buccal framework
surrounding the mouth. In the very young Ophiuroid the arms melt
into a small central disc, as in the Starfish, but the disc of the adult is
made up of a series of interradial dorsal outgrowths which meet one
another above the arms.

Fig. 207.—Aboral view of Ophiothrix fragilis. × 1. r, Radial plate.

Fig. 208.—Oral view of the disc of Ophiothrix fragilis. g.b, Opening of the genital
bursa; m.p, madreporite; pod, podia; t.p, tooth-papillae; v.p, ventral plates of
the arms. × 1.

One of the commonest British Ophiuroids is Ophiothrix fragilis (Figs.

207, 208), which is found in swarms in shallow water off the west
coast of England and Scotland. We may therefore select it as the
type, and, since the arm is the most characteristic organ of an
Ophiuroid, we may commence by studying it. Speaking generally, an
Ophiuroid either drags itself forward by two arms and pushes itself
by the other three (Fig. 207),[458] or else it drags itself by one and
pushes with the other four (Fig. 217). The arms during this process
are bent into characteristic curves, by the straightening of which in
the posterior arms the animal is pushed onwards, whilst the
intensification of these curves in the anterior arms causes the animal
to be dragged forwards. The grip of the arm on the substratum is
chiefly in the distal portion of the curve. The alteration of the
curvature is due to the contraction of the muscles on one side of the
arms. There is no ambulacral groove such as is found on the under
side of the arms of all Asteroidea, for the arm is completely
ensheathed by four series of plates, an upper row of dorsal plates,
an under row of ventral plates, and two lateral rows of lateral plates.
The last named, which in all probability correspond to the
adambulacral plates of Starfish, bear each a transverse row of seven
spines with roughened surfaces; these enable the animal to get a
grip on the substratum over which it moves. The podia in
Ophiuroidea are termed "tentacles"; they are totally devoid of
suckers, being simple conical papillae used as sense-organs, and
are of little, if any, service in locomotion. They issue from openings
called "tentacle-pores" situated between the edges of the ventral and
lateral plates, guarded each by a valve-like plate called the "tentacle-
scale." In Ophiothrix they are covered with sense-organs, each
consisting of a hillock-like elevation of the ectoderm, in which are
cells carrying long stiff sense-hairs. In most Ophiuroids such organs
are not present, though abundant scattered sense-cells occur, and
the outer surface of the tube-feet and the lining of certain pockets
called "genital bursae" (Fig. 208, g.b) are the only portions of the
surface where the ectoderm persists. Everywhere else, although
present in the young, it disappears, leaving as remnants a few nuclei
here and there attached to the under side of the cuticle.[459]
 Fig. 209.—Diagrammatic transverse section of the arm of an Ophiuroid. coe,
Dorsal coelomic canal; ect, ectoderm covering the tube-foot; ep, epineural
canal; gang.p, pedal ganglion; L, nerve-cord; musc, longitudinal muscles
attaching one vertebra to the next; nerv.rad, radial nerve-cord; perih, radial
perihaemal canal; pod, podium (tube-foot); sp, lateral spines; w.v.r, radial
water-vascular canal.

The greater part of the section of the arm is occupied by a disc-like

ossicle called the "vertebra." Each vertebra articulates with its
predecessor and successor by cup-and-ball joints, and it is
connected to each of them by four powerful longitudinal muscles.
Above, its outline is notched by a groove, in which lies an extension
of the coelom of the disc (Fig. 209, coe), but contains no outgrowth
of the alimentary canal, as is the case in Asteroidea. The vertebra is
also grooved below, and in this lower groove are contained the radial
water-vascular canal (Fig. 209, w.v.r), and below it perihaemal
canals as in Asteroidea; below this again the radial nerve-cord (L),
and beneath this again a canal called the "epineural canal" (ep),
which represents the missing ambulacral groove. This canal in the
very young Brittle Star is an open groove, but becomes closed by the
approximation of its edges. The vertebra, which has a double origin,
represents a pair of fused ambulacral ossicles. In Ophiohelus these
are only slightly adherent to one another (Fig. 216).
 Fig. 210.—Proximal and distal views of the three types of vertebra found
amongst Ophiuroidea. A, Ophioteresis, a type of the Streptophiurae (after
Bell), × 24; B, Astroschema, a type of the Cladophiurae (after Lyman), × 10;
C, Ophiarachna, a type of the Zygophiurae (after Ludwig), × 3. The upper
figure in all cases represents the distal aspect, the lower the proximal
aspect of the vertebra. v.g, Ventral groove.

When the surface of a vertebra is examined it is found that it can be

divided into a thin border, to which are attached the four muscles by
which it is connected to its successor and predecessor, and a central
portion, on which are situated the knobs and pits, by means of which
it articulates with the next vertebra.

The simultaneous contraction of the two upper muscles causes the

arm to bend upwards. The contraction of the two lower bend it
downwards, whilst a sideward movement is effected by the
contraction of the upper and lower muscle of the same side. On the
proximal surface of the central portion of the vertebra there is a
central knob and two ventro-lateral knobs, a median ventral pit and
two dorso-lateral pits, and on the distal surface there are pits
corresponding to the knobs on the proximal side and vice versa (Fig.
210, C). These knobs and pits restrict the movement of one vertebra
on the next, so that although the arms can undergo an unlimited
amount of flexion from side to side, they cannot be rolled up in the
vertical plane. When the under surface of the vertebra is examined
there is seen on each side of the central groove two round holes, a
distal and a proximal. The distal pair are for the passage of the
canals connecting the radial water-vessel with the tentacles, these
canals traversing the substance of the vertebra for a part of their
course; the proximal pair are for nerves going to the longitudinal
muscles, which likewise perforate part of the ventral border of the
vertebra.

In order to understand the anomalous circumstance that the canals

going to the tentacles actually perforate the vertebrae, it must be
clearly borne in mind that the basis of the body-wall in all
Echinoderms is a mass of jelly with amoebocytes in it, to which we
must assign the power of secreting carbonate of lime, and all we
have to assume in the case of Ophiuroids is that calcification spread
outwards from the original ambulacral ossicles into the surrounding
jelly, enclosing any organs that happened to traverse it.

When the ossicles of the arm are followed inwards towards the
mouth, they are seen to undergo a profound modification, so as to
form, by union with the corresponding ossicles of adjacent arms, a
structure called the mouth-frame. The general character of this
modification is similar to that affecting the first ambulacral and
adambulacral ossicles in the arms of an Asteroid, but in the
Ophiuroid the change is much more profound. The first apparent
vertebra consists of two separated halves, and each is fused with the
first adambulacral (lateral) plate, which in turn is firmly united with
the corresponding plate in the adjoining arm. Thus is formed the
"jaw," as the projection is called. The extensions of the mouth-cavity
between adjacent jaws are termed "mouth-angles." To the apex of
each jaw is attached a plate bearing a vertical row of seven short
blunt spines called "teeth" (Fig. 212, p). The plate is called the "torus
angularis" (Fig. 211, T), and on its ventral edge there is a tuft of
spines which are termed "tooth-papillae" (Fig. 208, t.p). On the upper
aspect of the jaw are a pair of plates termed "peristomial plates."
These discs—of which there are two in each radius, one on each jaw
which flanks the radius—possibly represent the separated halves of
the first vertebra, the apparent first vertebra being really the second.
On the flank of the jaw there is dorsally a groove for the water-
vascular ring and nerve-ring (Fig. 212, n.r), and beneath this a
groove for the first tentacle and a pore for the second, both of which
spring directly from the ring-canal; below these, in most Ophiuroidea,
but not in Ophiothrix, there is a row of blunt triangular spines called
"mouth-papillae" (Fig. 212, p1).

Fig. 211.—Diagrams to show the modification of the ambulacral and

adambulacral ossicles to form the armature of the mouth. A, Asteroid; B,
Ophiuroid. A1-A4, the first four ambulacra ossicles; Ad1-Ad4, the first four
adambulacral ossicles; J1, the first plate of the interradius (in the Ophiuroid
the scutum buccale); P, the spines borne by the jaw (in the Ophiuroid the
teeth); T, the torus angularis; W, the water-vascular ring; Wr, the radial
water-vessel; I, II, the first two pairs of tube-feet. (After Ludwig.)

The words "jaw" and "tooth" are misleading. There is no evidence

that the jaws of a Brittle Star are ever used for crushing food, but by
means of the muscles attaching them to the first complete vertebra
in the arm they can be rotated downwards so as greatly to enlarge
the mouth, and again rotated upwards and inwards, when they form
an excellent strainer to prevent the entrance of coarse particles. To
permit this extensive movement the articulatory facets on the
proximal surface of the first vertebra have been much modified; the
median knob and pit have disappeared, and the dorso-lateral pits are
raised on to the surface of processes, so that there are in all four
processes, two of which articulate with one half of a jaw.
 Fig. 212.—Lateral view of mouth-frame of Ophiarachna incrassata. × 4. A1?,
peristomial plate, possibly the half of the first vertebra; A2, the half of the
second vertebra; A3, the third vertebra; F1, pores for pair of tentacles; gen,
genital scale lying beside opening of genital bursa; musc, longitudinal
muscles connecting vertebrae; n.r, groove for nerve-ring; p, tooth; p1,
mouth-papilla; t, torus angularis. (After Ludwig.)

The mouth can be narrowed and the jaws forced inwards towards
the centre by the simultaneous contraction of five muscles (musc. tr,
Fig. 213) each, which unite the two halves of a jaw.

Turning now to the skeleton of the disc, we notice that dorsally it

consists of a closely-fitting mosaic of small plates, which are usually
concealed from view by a covering of minute spines. Opposite the
insertion of each arm there are, however, a pair of large triangular
plates ("radials"), which extend outwards to the periphery and
strengthen it, much as the ribs do in an umbrella. These radial plates
are always exposed, in Ophiothrix, even when the rest of the dorsal
plates are concealed by spines. On the under surface there is a
similar plating; but adjoining the jaws are five large, more or less
rhomboidal, plates termed "scuta buccalia" (Fig. 211, J1), on one of
which open the few madreporic pores which the animal possesses.
Attached to the sides of the scuta buccalia are the "lateral mouth
shields," which are in fact the adambulacral plates belonging to the
second pair of ambulacral plates which form the main mass of the
jaws. Further out, on the under side of the disc, there is, on each
side of each arm, a long narrow slit—the opening of the genital bursa
(Fig. 208, g.b), so that there are ten genital bursae. The "genital
bursa" (Fig. 214) is a sac lined by ciliated ectoderm projecting into
the interior of the disc. It is called genital because the openings of
the genital organs are situated on its surface; its main function,
however, is respiratory, the cilia bringing about a constant inward
current of fresh sea-water, the oxygen contained in which diffuses
through the thin wall of the sac into the coelomic fluid. The opening
of the bursa is strengthened on its radial side by a rod-like ossicle,
the "genital plate," and on its interradial side by an ossicle called the
"genital scale" (Fig. 212, gen), and in Ophiothrix the outer end of the
radial plate articulates with the outer end of the genital plate.
Muscles connect the two plates running on either side of the
articulation.

Observations on Ophiothrix[460] show that in this species at any rate

the radial plates can be raised or lowered. When they are raised the
centre of the disc is lifted into a cone and water is sucked into the
genital bursae, whereas when they are lowered the bursae are
compressed and water is expelled. This forced respiration appears
to come into play when the supply of oxygen is getting scanty.

The alimentary canal of Ophiothrix is a simple flattened sac (Fig.

213). It is devoid of an anus and cannot be everted through the
mouth. There is a horizontal pouch given off into each interradial
lobe of the disc. The sac is attached to the dorsal wall of the coelom
by numerous mesenteries, fibrous cords traversing the coelomic
cavity and clothed on the outer side by coelomic epithelium. To the
mouth-frame it is attached by a circular membrane, which we have
reason for believing is a functionless remnant of the retractor
muscles of the stomach of Asteroidea. In the young Asteroid there is
a similar sheet of membrane, which later becomes resolved into the
ten retractor bands.

The simple structure of the alimentary canal appears to be correlated

with the exceedingly simple character of the food. Ophiothrix feeds
on the most superficial layer of mud at the bottom of the sea. This
deposit consists partly of microscopic Algae and partly of decaying
organic matter, and is much more easily disposed of than the living
animals on which the Starfish preys. The food is shovelled into the
mouth by the first two or "buccal" pairs of tube-feet in each ray.

Fig. 213.—Longitudinal section through the disc of a young Ophiuroid passing

along one arm and the middle of the opposite interradius. (Diagrammatised
from an actual section of Amphiura squamata.) ab, Aboral sinus (dorsal in
the radius, ventral in the interradius); ax, axial sinus; coe, dorsal coelomic
canal of the arm; ep, epineural canal; gang.rad, ganglion of the radial nerve;
gen.r, genital rachis contained in the aboral sinus; gen.st, genital stolon;
mp, madreporic pore; musc.long, longitudinal muscle of the arm; musc.tr,
transverse muscle uniting the two halves of each jaw; mv, madreporic
vesicle; nerv.r, nerve-ring; p.c, pore-canal; perih, perihaemal canal; vert,
vertebra; w.vr, radial water-vessel.

The water-vascular system has undergone a most interesting set

of modifications, which can be explained by noticing the fact that the
tube-feet have almost, if not quite, lost their locomotor function and
are now used as tactile organs. The ampulla, or swollen inner end of
the tube-foot, has disappeared, and the upper end of the organ is
directly connected with the radial canal by means of a curved canal,
which traverses the outermost flange of the vertebra, appearing on
its surface in a groove on the outer side of the dorsal lateral knob on
the distal side of the ossicle. As in Asteroidea there are valves,
which regulate the entrance of fluid into the tube-foot. The stone-
canal is a curved tube of simple circular section and excessively
narrow bore which extends from the water-vascular ring downwards
to the madreporite (Fig. 213, mp) situated on one of the scuta
buccalia. The madreporite, in Ophiothrix as in most Brittle Stars, is
an exceedingly rudimentary structure, consisting of one or two pores
leading into as many pore-canals. From each interradius, except that
in which the stone-canal lies, a large Polian vesicle hangs down from
the water-vascular ring into the coelom.
We saw that in the Asteroid the ampulla was used like the bulb of a
pipette to force the fluid in the tube-foot down into the tip, so as to
press the sucker against the substratum. But when the tube-foot is
used as a sense-organ, a few circular fibres around its upper end
suffice to bring about all the extension that is needed. Since the
extension is no longer a very vigorous act, the loss of fluid by
transudation has probably been rendered insignificant, and hence
the stone-canal and madreporite, whose function it is to repair the
loss, have been reduced in size. The curious ventral curvature of the
stone-canal is, however, due to another cause. In the very young
Ophiuroid the madreporite is on the edge of the disc, and the stone-
canal extends horizontally outwards; and in some Asteroidea there is
a similar outward direction in its course. As development proceeds
the dorsal interradial areas of the disc of the young Ophiuroid grow
out into lobes, building up the conspicuous adult disc and forcing the
madreporite, and with it the stone-canal, downwards towards the
ventral surface.

The pores of the madreporite in Ophiothrix, like some of those in the

Asteroid, open not directly into the stone-canal but into the axial
sinus (Fig. 213, ax). This is a large ovoid sac, lined with thin
epithelium, lying between the stone-canal and the mouth-frame,
since of course it has shared in the ventral rotation of the stone-
canal. Its open connexion with the stone-canal was easily
recognised by Ludwig, who termed it, on this account, the "ampulla."
[461] The name "axial sinus" was bestowed mistakenly on another
cavity, which will be mentioned in connexion with the genital organs.

The radial perihaemal spaces of the arms open into a "perihaemal

ring" representing the outer perihaemal ring of Asteroids; but the
axial sinus does not have any such extension as constitutes the
inner perihaemal ring in Starfish. So-called oral circular and radial
blood strands are to be found in similar positions to the
corresponding structures in Asteroidea.
The nervous system might have been expected to have become
very much modified, since the activities of the Brittle Stars are so
different from those of the Starfish. It is indeed a universal rule in the
Animal Kingdom that, concomitantly with the increase in size and
activity of a muscle, there is a corresponding increase in the number
of ganglion-cells which control it. An accurate radial section of an
arm shows that there is, corresponding to the interspaces between
the two vertebrae, a ganglionic swelling of the nerve-cord. As in
Asteroids, there are not only ectodermic ganglion-cells on the under
surface of the cord abutting on the epineural canal, but also coelomic
ganglion-cells derived from the floor of the radial perihaemal canal.
Both these categories of cells are largely increased in number in the
ganglion. From the dorsal-cells arise a pair of large nerves which
pass directly up and supply the great intervertebral muscles. From
the interspace between the ganglia a direct prolongation of the
ventral part of the nerve-cord, the so-called pedal nerve, extends out
along the side of the tentacle, as in Asteroids. In Ophiuroids it swells
out into a ganglion, completely surrounding the tentacle and giving
off nerves to the surfaces of the arm which terminate in the cuticle.

There is a large ganglion where the radial cord joins the nerve-ring,
and, owing to the more specialised condition of the nervous system,
a severed arm in an Ophiuroid is much more helpless than an arm of
an Asteroid. It will not carry out "escape movements," and is for a
long time rigid under the shock of section; at last it simply gives
reflex movements on stimulation.

Preyer[462] endeavoured to test the "intelligence" of Ophiuroids by

observing how they would adapt themselves to circumstances which
it might be fairly assumed they had never encountered in their
ordinary experience. To this end he passed over the arm of a
specimen a piece of indiarubber tubing, which clung to it tightly. He
found that the animal first tried walking off, pressing the encumbered
arm against the ground, so that the piece of tubing was rubbed off. It
was then replaced more tightly than before; the animal, having tried
the first method without result, waved the arm to and fro in the water
till the rubber floated off. In a third experiment the animal held the
rubber against the ground by a neighbouring arm, and drew the
encumbered arm out. When the rubber was replaced a fourth time,
the animal kicked it off by alternately pressing neighbouring arms
against it. Finally, when the rubber was put on so firmly that all the
above-mentioned methods failed, the arm was broken off. Preyer
concludes from this that Ophiuroids have a high degree of
intelligence; but this may be doubted, and the reader is referred to
the account of Uexküll's experiments given in the next chapter.
There is, however, no doubt at all that Ophiuroidea are by far the
most active of all Echinoderms, and one would naturally correlate
this with higher psychic development.

The radial nerve ends in a terminal tentacle sheltered by a median

plate at the end of the arm; but eyes, such as are found in Asteroids,
are wanting, and the animal does not appear to be sensitive to light.

The reproductive system in Ophiuroids consists of a genital stolon

giving rise at its distal end to a genital rachis, which extends in a
circular course round the disc, ensheathed in an "aboral sinus" (Fig.
213, ab) and swelling out so as to form the gonads (testes or
ovaries), where it passes over the inner side of the genital bursae.
The genital stolon (Fig. 213, gen.st) is a compact ovoid organ, often
termed on account of its shape the "ovoid gland." It is situated close
to the stone-canal, and, as in Starfish, it indents the outer wall of the
axial sinus; but, unlike the stolon of the Asteroid, it is separated from
the general coelom by a space, of which it forms the inner wall, but
whose outer wall is formed by a sheet of membrane. This cavity
must be carefully distinguished from the axial sinus of Asteroidea, to
which it was supposed at one time to correspond; it is really formed
by a pocket-like ingrowth of the general coelom into the septum
dividing it from the axial sinus. The cells forming the inner side of this
pocket form the primitive germ-cells, which constitute the main mass
of the ovoid gland; those of the outer side remain thin. The cavity of
the ingrowth is shut off from the general coelom, but persists
throughout life. In Asteroids a similar ingrowth takes place, but both
walls thicken and become converted into germ cells, and the cavity
disappears, and, as in Asteroidea, a considerable number of the
germ-cells in the stolon degenerate.

Fig. 214.—Diagram of a tangential section through the edge of the disc of an

Ophiuroid to show the relations of the disc, arm, and genital bursae. ep,
Epineural canal; musc, longitudinal muscle of the arm; nerv.rad, radial nerve
cord; ov, ovary; perih, radial perihaemal canal; w.v.r, radial water-vessel.

The genital rachis (Fig. 213, gen.r) is an outgrowth of the distal end
of the genital stolon, which extends in a complete circle round the
disc. The rachis does not, however, lie everywhere in the same
plane, but by its undulating course bears witness to the distortion
which the disc has undergone. In the radii it is, as in the Asteroid,
dorsal; but in the interradii it is ventral, this ventral portion having,
like stone-canal and axial sinus, been carried down by the
preponderant growth of the dorsal parts of the disc. It is everywhere
ensheathed by the aboral sinus, which, as in Asteroids, is an
outgrowth of the coelom. The rachis is embedded in a strand of
modified connective tissue, to which we may (as in the case of
Asterias) apply the name "aboral blood-ring." Both on the central and
peripheral sides of this sinus are vertical muscles connecting the
genital and the radial plates, which bring about the respiratory
movements already referred to. Just above the madreporite, at the
end of the genital stolon, is a small, completely closed space, which
by its position corresponds with the madreporic vesicle of Asteroids
and represents the right hydrocoel (Fig. 213, mv). As the rachis
passes over the genital bursa it gives off branches, which swell up to
form the genital organs. In Ophiothrix there is one such organ on
each side of each bursa, but in other genera (cf. Ophiarachna) a
large number of small ones. The genital products are shed into the
water through the bursae.
 Classification of Ophiuroidea.
Before proceeding to study the classification of Brittle Stars, it is
necessary to give some account of the range of structure met with in
the group.

Number of Radii.—The number of arms is rarely increased, and

hardly ever exceeds six; a few species (each an isolated one in its
genus) have six arms, and in one case (Ophiactis virens), at any
rate, this is associated with the power of transverse fission. In many
Cladophiurae the arms fork repeatedly, so that although there are
only five radii, there is quite a crowd of terminal branches.

Vertebrae.—The vertebrae differ in the manner in which they

articulate with one another. In Ophiothrix fragilis taken as the type,
which in this respect resembles the vast majority of species
(Zygophiurae), the knobs and pits on the faces of the vertebrae
prevent the arms from being coiled in the vertical plane. In
Ophioteresis (Fig. 210, A) and some allied genera (Streptophiurae)
the knobs are almost obsolete, and the arms are free to coil in the
vertical plane; whilst in Gorgonocephalus and Astrophyton
(Cladophiurae) the arms are repeatedly branched and the vertebrae
have saddle-shaped articulating surfaces, so that they have quite a
snake-like capacity for coiling themselves round external objects. In
Ophiohelus (Fig. 216) each vertebra consists of two rod-like plates
placed parallel with the long axis of the arm and fused at both ends,
but divergent in the middle, leaving a hole between them.

Covering Plates of the Arms.—The upper arm-plates are the most

variable. They may be surrounded by small supplementary plates
(Ophiopholis) or double (Ophioteresis). In all (?) Cladophiurae and
most Streptophiurae they are absent, being replaced by minute
calcareous granules. Under arm-plates are absent in Ophioteresis
and in the distal portion of the arms in many Cladophiurae. Side arm-
plates are constantly present, and in most Cladophiurae meet in the
middle line below.
Arm-Spines.—The spines borne by the lateral covering plates of the
arms vary greatly in character. In Ophiura and its allies they are short
and smooth, and are borne by the hinder edge of the arm and
directed backwards; but in the larger number of genera they are
borne nearer the centre of the plate, and are directed outwards at
right angles to the arm. They may be covered by small asperities, as
in Ophiothrix (Fig. 215, C), when they are said to be rough; or these
asperities may become secondary spines, as in Ophiacantha (Fig.
215, B), when they are said to be thorny. In Ophiopteron all the
spines borne by a single plate are united by a web of skin so as to
constitute a swimming organ. The small plates guarding the ends of
the tentacles (tentacle-scales) may be absent, or more rarely double.
In Cladophiurae there is a regular transition from tentacle-scale to
arm-spine; the tentacle-scale being merely the smallest of the series
of lateral spines.

True pedicellariae are unknown amongst Ophiuroidea, since there

is no longer a soft ectoderm to protect, but in some cases, as for
instance in Ophiohelus, small hooks movable on a basal piece
attached to the arms are found which may represent the vestiges of
such organs (Fig. 216). Similar hooks are found in the young
Ophiothrix fragilis just after metamorphosis and in all Cladophiurae,
replacing in the latter case the arm-spines in the distal portion of the
arm.

Fig. 215.—Three types of mouth-frame found in Zygophiurae. A, Ophioscolex, ×

10; B, Ophiacantha, × 6; C, Ophiothrix, × 6. (After Lyman.)