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genital plates is also pierced by the madreporic pores. Some
zoologists have separated the ocular and the genital plates under
the name of "calyx" from the rest of the corona, under a mistaken
idea that they are homologous with the plates of the body or calyx of
a Crinoid.

Fig. 229.—The peristome of Echinus esculentus. × 2. 1, Tube-feet of the lower

ends of the radii; 2, gill; 3, teeth; 4, buccal tube-foot; 5, smooth peristomial
membrane. (After Kükenthal.)

The periproct (Fig. 228, 4) is covered with small plates and bears a
few pedicellariae. The peristome (Fig. 229) is covered by flexible
skin with abundant pedicellariae; it terminates in a thick lip
surrounding the mouth, from which the tips of five white teeth are just
seen projecting. There are ten short tube-feet projecting from the
peristome—one pair in each radius—and each tube-foot terminates
in an oval disc and is capable of little extension, and each has
around its base a little plate. The presence of these tube-feet shows
that in Echinus the peristome extends outwards beyond the water-
vascular ring, whereas in Asteroidea it is contained entirely within the
ring. In the primitive Cidaridae (Fig. 235) the whole peristome down
to the lip surrounding the mouth is covered with a series of
ambulacral and interambulacral plates similar to those forming the
corona, though smaller and not immovably united, and the series of
tube-feet is continued on to it. It is thus evident that the peristome is
merely part of the corona, which has become movable so as to
permit of the extension of the teeth. In Echinus the peristome is
continued in each interradius into two branched outgrowths called
gills, the relation of which to the respiratory function will be described
later. These gills (Fig. 229, 2) are situated in indentations of the edge
of the corona called "gill-clefts" (Fig. 230, g).

Fig. 230.—The dried peristome of Echinus esculentus and the surrounding

portions of the corona. × 1. amb, Ambulacral plate; b.t, buccal tube-foot; g,
gill-cleft; inter, interambulacrum; per, peristome.

The most conspicuous plates in the peristome are those surrounding

the buccal tube-feet; besides these, however, there are in Echinus
esculentus, and probably in most species, a large number of thinner
irregularly-scattered plates (Fig. 230).

The term ambulacral plate, applied to the plate pierced by the pores
for the tube-feet, conveys a misleading comparison with the
ambulacral plate of an Asteroid. In Echinoids the ambulacral groove
has become converted into a canal called the "epineural canal," and
the ambulacral plates form the floor, not the roof, of this canal; they
may perhaps correspond with the adambulacral plates of the
Starfish, which one may imagine to have become continually
approximated as the groove became narrower until they met.
 Fig. 231.—Dissection of Echinus esculentus. × 1. The animal has been opened
by a circumferential cut separating a small piece of the skeleton at the
aboral end, which is turned outwards exposing the viscera on its inner
surface. The other viscera are seen through the hole thus made. amp,
Ampullae of the tube-feet; aur, auricle; b.v, so-called "dorsal blood-vessel";
comp, "compasses" of Aristotle's lantern, often termed "radii" by English
authors; comp.elv, elevator muscles of the compasses; comp.ret, retractor
muscles of the compasses; eph, epiphyses of the jaws in Aristotle's lantern;
gon, gonad; g.rach, genital rachis; int, intestine; oe, oesophagus; prot,
protractor of Aristotle's lantern; rect, rectum; ret, retractor of Aristotle's
lantern; siph, siphon; st, stomach; stone.c, stone-canal.

The internal organs of the Urchin can best be examined by making a

horizontal incision about one-third the distance from the mouth and
pulling the two parts gently asunder. A large amount of fluid escapes
from the exceedingly spacious coelomic cavity, the alimentary canal
being comparatively narrow.

The alimentary canal commences with a short vertical tube which

has been shown to be a stomodaeum; this is surrounded by the
upper ends of the teeth and their supporting ossicles, which are
collectively termed "Aristotle's lantern." The oesophagus leads into a
baggy, flattened tube, the stomach, which runs horizontally round the
animal, supported by strings of tissue from the coelomic wall, so that
it hangs down in a series of festoons. Having encircled the animal, it
bends directly back on itself and immediately opens into the
intestine, which is also a flattened tube, which runs round the
circumference of the animal, but in the opposite direction, the
festoons of the second circle alternating with those of the first. The
intestine opens into a short rectum which ascends vertically to open
by the anus. The stomach is accompanied by a small cylindrical tube
called the "siphon" (Fig. 231, siph), which opens into it at both ends;
this represents merely a gutter which has been completely grooved
off from the main intestine; it is lined by cilia, and its function is
believed to be that of keeping a stream of fresh water flowing
through the gut, so as to subserve respiration.

Echinus esculentus seems to feed chiefly on the brown fronds of

Laminaria and the small animals found thereon, which it chews up
with its teeth, but it may regale itself on the same diet as Brittle
Stars, as Allen[474] has shown to be the case in Plymouth Sound.
Dohrn[475] has described the Neapolitan Sphaerechinus granularis
attacking and capturing Crustacea such as Squilla.

The water-vascular system presents several features of great

interest. The ring-canal is situated at a considerable distance above
the nerve-ring, and is separated from it by the whole of the jaws and
teeth. It has five small interradial pouches on it, which apparently
correspond to Tiedemann's bodies in an Asteroid. The stone-canal
(Fig. 231) opens as usual into the ring-canal, and is accompanied by
the axial sinus and genital stolon. The name "stone-canal" is very
unsuitable in this order, for there are no calcifications in its walls; it is
a simple membranous tube of circular section. On reaching the
upper wall of the test it expands into an ampulla, into which the
numerous ciliated pore-canals traversing the madreporite open. The
radial canals, starting from the ring-canal, pursue a downward
course till they come into contact with the radial nerve-cords, and
they then bend upwards and run along the centre of the ambulacral
region, finally terminating in the small terminal tentacles. In the just
metamorphosed Echinoid these are well-developed tube-feet, each
with a well-developed sucker, in the centre of which is a conical
sensory prominence, but as development proceeds they become
enclosed in a circular outgrowth of the test, so that only the tip
projects in the adult.

The long extensible tube-feet are connected by transverse canals

with the radial canal. Instead of the pair of valves which in Asteroids
prevent the reflux of liquid into the canal, there is a perforated
diaphragm[476] with circular muscles, which by contraction close the
opening in the diaphragm, while when they are relaxed fluid can
return from the tube-foot. The ampulla is flattened, and is contracted
by muscular fibres called "trabeculae" stretching across its cavity.
These muscular strands are developed by the cells lining the
ampulla. The external portion of the tube-foot, as in Asteroids, is
provided with powerful longitudinal muscles, and there is the same
alternate filling and emptying of the ampulla as the tube-foot is
contracted and expanded. The tube-foot is connected by a double
canal with the ampulla, the object of which is to assist in respiration.
The cells lining it are ciliated, and produce a current up one side of
the tube-foot and down the other, and the double canal leading to the
ampulla separates these two currents and prevents them interfering
with one another. Thus water is continually transported from the
ampulla to the tube-foot, through the thin walls of which it absorbs
oxygen, and it is then carried back to the ampulla, and transfers its
oxygen to the fluid of the general body-cavity through the walls of the
ampulla. The disc of the tube-foot is supported by a calcareous plate
(Fig. 232, oss), a circumstance which enabled Johannes Müller to
recognise the Echinoid larva when the form of the adult was as yet
unrecognisable. Below the edge of the disc there is a well-marked
nerve-ring, from which two bundles of nerve-fibres go to the disc
itself, in the edge of which there is an abundance of sense-cells.

The buccal tube-feet (Fig. 229, 4) are much shorter than the rest,
and are provided with oval discs which are highly sensory. These
feet are not used for seizing, but for tasting food; when a piece of
food is placed near them they are thrown into the most violent
agitation.
 Fig. 232.—Diagrammatic transverse section of the radius of an Echinoid.
amb.oss, Ambulacral ossicle; amp, ampulla of the tube-foot; ep, epineural
canal; musc, muscles attaching spine to its boss; nerv, nervous ring in base
of spine; n.r, radial nerve-cord; oss, ossicle in sucker of tube-foot; ped,
tridactyle pedicellaria; perih, radial perihaemal canal; pod, tube-foot; wv.r,
radial water-vascular canal.

The nervous system has the same form as in an Asteroid, viz. that
of a ring surrounding the mouth and giving off radial nerve-cords
(Fig. 232, n.r), one of which accompanies each water-vascular canal
to the terminal tentacle, where it forms a nervous cushion in which
pigmented cells are embedded.

A large band-like nerve is given off from the radial nerve-cord to

each tube-foot. This pedal nerve, as it is called, contains bipolar
neurons, and is really an extension of the nerve-cord itself. Beneath
the sucker it branches out to form a sensory ring. From the base of
the pedal nerve, branches are given off which run to the ectoderm
and enter into connexion with the plexus there. Romanes[477]
scraped away the radial cords and found that the spines still
converged when a point on the ectoderm was stimulated, but that,
on the other hand, if definite locomotor movements were to be
carried out, the presence of these cords was a necessity; hence he
concluded that the superficial plexus sufficed for ordinary reflexes,
but that for purposeful movements the central nervous system was
necessary.

Von Uexküll[478] has made an exhaustive study of the physiology of

the nervous system in the Echinoidea. He points out that all the
organs controlled by the nervous system, spines, pedicellariae, tube-
feet, and (see below) Aristotle's lantern, give two opposite reactions
in response to the same stimulus according as it is strong or weak,
bending away from the point of stimulation when it is strong and
towards it when it is weak. This reversal of reaction can only be due
to the action of the neuron in altering the effect of the stimulus on the
muscles, and this Uexküll regards as its fundamental property. Thus
in Preyer's[479] experiments with Starfish the strong form of
stimulation is obtained by directly applying the stimulus to the radial
cord or to the tube-feet, the weak form by stimulating the back, when
of course the stimulus has to traverse a longer path before affecting
the tube-feet, and is consequently weakened. Von Uexküll also
introduces the conception of "tone" with regard to the nervous
system. This term has been used to denote the amount of chronic
contraction in a muscle, and it is to be distinguished from the fleeting
contractions which cause movement. The more tone there is in a
muscle the less responsive it is to stimuli tending to bring about
movement. As applied to the nervous system "tone" denotes a
condition when it is not receptive to small stimuli, but when it is
maintaining a condition of tone in a muscle by which of course its
own tone is measured. Tone in a neuron can therefore be measured
by the produced tone in the muscle, and the one is to be
discriminated from the other only by using stimulants, such as
caffeine, which have no direct action on muscle. Tone can also be
measured by the amount of stimulus necessary to irritate the neuron.
When muscles are stretched the tone is lowered, and this loss of
tone extends to the neuron controlling the muscle, and vice versa.
When the spines on being gently stimulated bend towards the point
of stimulation, this is due to the contraction of the muscles on the
side towards the point of stimulus, for if the superficial plexus of
nerve-fibres be cut through so that the stimulus has to pursue a
round-about course the spine will bend towards the direction from
which the stimulus comes. The bending of the spines away from the
stronger stimulus is likewise due to the muscles on the side towards
the stimulus. It is caused by a sudden fall of tone in these muscles,
which causes them to yield to the tone of the muscles on the
opposite side, and this fall of tone is due to a fall of tone in the
neurons, for it can be produced by chemicals, and the direct action
of all chemicals applied to muscle is to raise tone.

In Arbacia this form of reaction cannot be produced; the spines

respond to stimuli of all degrees of intensity by convergence towards
the point of stimulation.

When a general skin-irritant like dilute acetic acid, or even strong

light, is applied to the skin of a Sea-urchin the spines bend
alternately to all points of the compass, or, in a word, rotate. This is
due to the fact that the weight of the inclined spine stretches the
muscles of one side and so renders them more open to the general
stimulus; these muscles in consequence, contract, and so move the
spine to a new position in which other muscles are stretched, and a
similar result follows. A continuation of this process brings about
rotation.

When a piece of glass rod or other light object is laid on the spines of
a Sea-urchin, it naturally, by its weight, presses asunder the spines
and stretches their muscles on one side, thus lowering the tone. If
now the skin be stimulated at any point the piece of rod will be rolled
by the spines towards the point of stimulation. This is caused by the
fact that the muscles of the spines holding the rod are made more
receptive by being stretched, and therefore they contract more than
do the others in response to the stimulation, and so the rod is rolled
onwards on to the next spines, which then act in the same manner.
This passage of stimulus is entirely independent of direct nervous
connexion between the bases of the spines, for it will traverse at
right angles a crack going clean through the shell; it is merely the
result of the mechanical weight of the object and of the juxtaposition
of the spines.

If the stimulation be too violent the first spines affected diverge wildly
and strike their neighbours with vehemence, so arousing into activity
the block musculature of these. This causes them to stand rigidly up,
and so the path of the stimulus is barred.
Now the escape movements of the animal under strong stimulation
which Romanes[480] alludes to are just an example of this handing
on of stimulation from spine to spine, not by nervous connexion but
by mechanical touch only; the object in this case is the substratum
on which the animal lies, which is, so to speak, rolled towards the
point of stimulation, or putting it otherwise, the animal is rolled away
from it. Righting when upset is another example of the same
phenomenon; the aboral spines are stretched by the weight of the
animal, and the animal acts as if it were stimulated in the region of
the periproct. When a Sea-urchin is in its normal position and is
stimulated in the periproct (as for instance by a strong light), it would,
according to this rule, tend to move downwards, which is of course
impossible; but as the stimulus never affects all sides quite alike the
result is that the Urchin rotates, turning itself ever away from the
point of strongest stimulation. In the case of Strongylocentrotus
lividus when living on limestone, as on the west coast of Ireland, this
results in the animal excavating for itself holes in the rock, where it is
safe from the action of the breakers.[481]

But it may be objected that no account is taken in the above

description of the action of the "central nervous system," i.e. of the
ring and the radial cords, and yet Romanes found that when they
were removed the escape movements could not be carried out. The
answer is that the central nervous system is a store-house of tone,
not, as in higher animals, a controlling centre for co-ordinating the
movements of the spines. When it is removed at first the escape
movements can be carried out, but in a day or two all tone in the
spine-muscles is lost, and then, since the tone of all is equally low,
there is no tendency in those that are stretched to be more
responsive than others, and hence the escape movements cannot
be carried out. Sea-urchins kept in the tanks of an aquarium are apt
to lose the tone of their spines owing to the poisoning of the nervous
system.

The central nervous system is, however, the system which controls
the movements of the tube-feet. As we have seen, extensions of the
radial nerves run to the tip of each podium. Tube-feet are chiefly
used in ordinary progression; when this is quickened the spines
come into play exclusively. The extent to which these two organs of
locomotion are used varies from genus to genus. Thus
Centrostephanus uses its spines a good deal, Echinus and
Strongylocentrotus very little. The last-named genus sometimes
walks on its tube-feet entirely without touching the ground with its
spines.

The faculty of vision in its simplest form may be defined as

sensitiveness to light and shade. Now strong light acts on all Sea-
urchins as a general skin irritant. They fly from it towards the darkest
corner, and then if it continues the spines rotate. A number of little
violet spines on the aboral pole of Centrostephanus longispinosus
are especially sensitive to light, and hence are almost constantly in
rotation. This is due, according to Uexküll,[482] to a pigment of a
purple colour, which can be extracted by means of alcohol and which
is decomposed by light, the products of decomposition being
supposed to irritate the nerves. Centrostephanus when exposed to
light becomes darker in colour. This is due to the migration outwards
of amoebocytes, which carry a pigment which acts as a screen in
order to prevent the valuable visual purple being too rapidly
decomposed. Not all Sea-urchins, in fact very few of those living in
northern waters, give a reaction to shadow. C. longispinosus is one
of the few; it reacts to a shadow by converging its spines towards it.
A much larger number of species inhabiting tropical waters show this
reaction. It is entirely stopped if the radial nerve-cords be removed,
whereas the reaction to strong light continues. The reaction to shade
is strongest after a long previous exposure to light, hence Uexküll
has given the following explanation of it. The continued irritation due
to light, having spread to all the spines, eventually reaches the radial
cords and is there stored in the bipolar nerve-cells as tone. When the
light-stimulus is interrupted some of the stored tone spreads
upwards to the spines, causing the weak form of spine reaction, and
the spines converge.
 Fig. 233.—To show character and distribution of the sphaeridia in
Strongylocentrotus droëbachiensis. A, a portion of a radius, with sphaeridia,
and the adjoining edge of the peristome. p, Pair of pores for a tube-foot; per,
peristome; t, primary tubercle. B, an isolated sphaeridium. (After Lovén.)

It will be seen therefore that the so-called central nervous system of

Echinus does not act in any sense as a brain, as indeed might have
been guessed from the absence of any differentiation in it. As
Uexküll points out, when an animal is covered all over with similar
organs, such as spines and pedicellariae, capable of acting
automatically, a brain is not needed. The object of a brain is to direct
organs which are in a certain place to a danger which may come
from any quarter, but in the Sea-urchin any spine is as good as any
other spine, and such orientation is not needed. "In a dog the animal
moves its legs, in a Sea-urchin the legs move the animal." What the
Sea-urchin does need is a means to prevent its pedicellariae
attacking its own organs with which they may come into contact.
Thus it possesses an "autodermin," a chemical contained in the
ectoderm which paralyses the muscles of the pedicellariae, as may
be seen by offering to them a spine of the same animal. If, however,
the spine be treated with boiling water, and then offered, it is
viciously seized, showing that this substance can be dissolved out.

Just as in the case of the Starfish, when the nerve-ring is cut

through, the tube-feet in the various radii are no longer co-ordinated
with one another.

Besides the tips of the tube-feet the Urchin possesses another kind
of sense-organ, the sphaeridia (Fig. 233). These are minute glassy
spheres of calcareous matter attached by connective tissue to
equally minute bosses on the plates of the ambulacra, generally near
the middle line. They are in fact diminutive spines, and like the latter
are covered with a thick layer of ectoderm, beneath which is a
particularly well-developed cushion of nerve-fibrils. Only the layer of
muscles which connects a normal spine with its boss is wanting.
Although definite experimental proof is lacking, the whole structure of
the sphaeridia shows that they belong to the category of "balancing
organs." As the animal sways from side to side climbing over uneven
ground, the heavier head of the sphaeridia will incline more to one
side or to another, and thus exercise a strain on different parts of the
sheath, and in this way the animal learns its position with regard to
the vertical.

Intervening between the radial nerve-cord and the radial vessel is a

single radial perihaemal canal (Fig. 232, perih), representing the
two parallel canals found in the same position in the Asteroid. The
five perihaemal canals lead downwards to a space called the
lantern-coelom, surrounding the oesophagus.[483] Since the
skeleton of the corona is composed of plates immovably connected
together, muscles corresponding to the ambulacral muscles of the
Asteroids would be useless, and so the wall of the perihaemal canal
remains thin and the side of it turned towards the general coelom
develops no muscles, and that turned towards the nerve-cord no
nerve-cells. Where, however, the radial nerve enters the nerve-ring,
and on the ring itself, an inner layer of nerve-cells is developed from
the lantern-coelom which represents the lower or oral portions of the
radial perihaemal canals. These cells control the muscles moving the
teeth. These canals are originally parts of the lantern-coelom, but in
the adult they become closed off from it.
 Fig. 234.—Echinus esculentus dissected in order to display Aristotle's lantern, ×
2. The whole upper part of the shell has been cut away. 1, Upper growing
end of tooth; 2, outer forked end of one "compass"; 3, muscle joining
adjacent compasses and acting as elevator of these ossicles; 4, depressor
of the compasses; 5, lower end of jaw; 6, retractor of the whole lantern; 7,
protractor of the whole lantern; 8, auricle; 9, ampullae of the tube-feet; 10,
interambulacral plate; 11, lower part of tooth; 12, water-vascular ring; 13,
meeting-point of a pair of epiphyses; 14, so-called Polian vesicle, really
equivalent to Tiedemann's body in an Asteroid; 15, oesophagus; 16, so-
called ventral blood-vessel; 17, genital stolon; 18, stone-canal; 19, rectum;
20, aboral sinus. (Partly after Chadwick.)

In the outer wall of this space are developed the calcareous rods
forming Aristotle's lantern. These are first: five teeth (Fig. 234, 11),
chisel-shaped ossicles of peculiarly hard and close-set calcareous
matter, the upper ends (1) pushing out projections of the upper wall
of the lantern-coelom. These projections are the growing points of
the teeth, whose lower ends pierce the ectoderm and project into the
lower end of the oesophagus. Each tooth is firmly fixed by a pair of
ossicles inclined towards one another like the limbs of a V and
meeting below. Each ossicle is called an "alveolus," and taken
together they form a "jaw." Their upper ends are connected by a pair
of ossicles called "epiphyses" (13). These two epiphyses meet in an
arch above. The jaws and their contained teeth are situated
interradially. Intervening between successive alveoli are radial pieces
called "rotulae," which extend directly inwards towards the
oesophagus. Above the rotulae are pieces termed "radii" or
"compasses" (2), which are not firmly attached to the other pieces
but lie loosely in the flexible roof of the lantern-coelom.

The uses of the various components of this structure can be made

out from an inspection of the muscles which connect them together.

Overarching each radial perihaemal canal where it leaves the lantern

is a bridge of calcareous matter called the "auricula" (Fig. 234, 8).
This arises as two rods which meet each other in a pent-house over
the canal. It is the only part of the skeleton which can be compared
to the ambulacral ossicles of the Asteroidea, and like them it serves
as the point of insertion for important muscles. Thus we find (1)
protractor (Fig. 234, 7) muscles which arise from the upper ends of
the alveoli and are inserted in the auricula; when these contract they
tend to push the whole "lantern" outwards so as to expose the tips of
the teeth. (2) The retractor muscles (Fig. 234, 6) extend from the
auriculae to the lower ends of the jaws and restore the lantern when
it has been extruded to its original position. (3) The comminator
muscles connect adjacent jaws with one another: these on
contraction approximate the pair of jaws into which they are inserted,
and it will easily be seen that by the successive contraction of the
five comminator muscles a rotating movement of the teeth would be
produced which would cause them to exert an action something like
that of an auger; by their simultaneous contraction the teeth are
brought to a point. (4) The internal and external rotula muscles:
these are small muscles which connect the outer side of the
epiphysis with the rotula. There are two facets on the epiphysis,
which permit it to rock to and fro on the rotula under the action of
these muscles. This rocking action must greatly increase the cutting
power of the tooth. These muscles are controlled by the nerve-ring
and the incipient portions of the radial nerves, which, as we have
seen, have an inner layer of nerve-cells. If the nerve-ring be gently
stimulated on one side the upper end of the lantern bends away from
the spot, causing the lower end, i.e., the teeth, to move towards it;
but a stronger stimulation produces the opposite effect, just as is the
case with spines. But besides these masticatory muscles there are
others which have nothing to do with moving the teeth. These
muscles are attached to the rods called radii or compasses (Fig.
234, 2),[484] which lie in the upper wall of the lantern-coelom, and
may be termed the compass muscles. There are two sets:—(1) The
elevator muscles (Fig. 234, 3), which connect the inner ends of the
compasses with one another. When these contract, the radii tend to
bend upwards at the inner ends and thus raise the roof of the
coelom. (2) The depressor muscles (Fig. 234, 4), which run
downwards from the forked outer ends of the compasses to the
auriculae. Uexküll[485] has shown that the function of these muscles
and of the rods to which they are attached is respiratory. These
muscles are also controlled by the nerve-ring. If this be stimulated by
passing a pin-head into the oesophagus, the roof of the lantern
cavity is raised by the contraction of the elevator muscles. This is
followed by contraction of the depressor muscles lowering it; the
same result may be brought about by placing the animal in water
with excess of carbonic acid. The ten branched gills described on p.
514 are outgrowths of the lantern-coelom. When the roof of this
cavity is depressed the fluid contents are driven out into the gills,
which are thus expanded and then absorb oxygen from the
surrounding sea water. When, on the other hand, the roof is raised
the aerated water is sucked back into the lantern cavity, and the
oxygen passes easily through the thin walls of the lantern into the
fluid filling the main coelomic cavity. There are thus two independent
respiratory mechanisms in the Sea-urchin, the one being the
compass muscles, the other the cilia lining the interior of the tube-
feet.

The function of excretion is performed, as in Asteroidea, by the

amoebocytes floating in the general coelomic cavity. These in part
escape through the thin bases of the gills. In other parts of the body
they seem not to succeed in reaching the exterior at all, but to
degenerate and to form masses of pigment; the colour of the animal
is largely due to these excrementitious substances.

The reproductive system, as in the two preceding orders, consists

of a vertical pillar, the "genital stolon," and a circular "genital rachis"
giving off interradial branches from which the genital organs bud.
The genital stolon is developed from the wall of the general coelom
near the upper end of the axial sinus; it attains a great development
and ultimately completely surrounds the axial sinus, which then
appears like the cavity of a glandular tube, the walls of which are
constituted by the genital stolon. The compound structure consisting
of stolon and axial sinus was actually described as a nephridium by
the Sarasins[486] in the case of Asthenosoma. Its true nature,
however, is shown when the upper end is examined; it is then seen
to open into the stone-canal and to be in communication with the
ampulla, into which the pore-canals open. Lying alongside the upper
end of the axial sinus is the somewhat elongated "madreporic
vesicle," or right hydrocoele, which was described by Sarasin as the
accessory kidney (Nebenniere), since like the axial sinus it is partly
enveloped by the genital stolon. Leipoldt,[487] however, showed
clearly that it is a completely closed space.

The genital rachis springs from the upper end of the stolon, and as in
Asteroids, it lies in the outer wall of a space called the "aboral sinus"
(Fig. 234, 20) intervening between it and the test. In adult specimens
it seems to degenerate. The genital organs are situated at the ends
of five interradial branches of the rachis (Fig. 231, gon). Each is an
immense tree-like structure consisting of branching tubes, which are
lined by the sexual cells. So enormous do they become in the
breeding season that they form an article of food among fishermen.
The term esculentus is derived from this circumstance. Other
species are regularly sold for food as Frutta di Mare (Fruit of the
Sea) at Naples, and as "sea eggs" in the West Indian Islands. One
female Echinus esculentus will produce 20,000,000 eggs in a
season.
The so-called blood system is more distinctly developed in
Echinoidea than in Asteroidea and Ophiuroidea. There is an oral ring
of lymphoid tissue surrounding the oesophagus below the water-
vascular ring. From this are given off two strands, the so-called
"dorsal" (Fig. 231, b.v), and "ventral" vessels (Fig. 234, 16), which
run along the two opposite sides of the stomach or first coil of the
alimentary canal. The position of these strands suggests that like the
lacteals of the human intestine they are channels along which the
products of digestion exude from the stomach. The dorsal strand is
situated on the same side as the genital stolon, and from it branches
are given off which ramify on the surface of the stolon, on account of
which this organ, as in Asteroidea, was at one time regarded as a
"heart," but the distinction of the stolon from the strands is easily
made out. An aboral ring enclosing the genital rachis lies embedded
in the septum dividing the aboral sinus (Fig. 234, 20) from the
general coelom.

Classification of Echinoidea.
The Echinoidea are sharply divided into three main orders, which
differ from each other profoundly in their habits and structure. These
are: (1) The Endocyclica or Regular Urchins, of which the species
just described may be taken as the type. (2) The Clypeastroidea or
Cake-urchins, which are of extremely flattened form, and in which
the periproct is shifted from the apical pole so that it is no longer
surrounded by the genital plates, while some of the tube-feet of the
dorsal surface are flattened so as to serve as gills. (3) The
Spatangoidea or Heart-urchins, in which the outline is oval: the
periproct is shifted, as in the Cake-urchins, and the dorsal tube-feet
are similarly modified; but the Heart-urchins have totally lost
Aristotle's lantern, whilst the Cake-urchins have retained it. This
strongly-marked cleavage of the group was primarily due, as in all
such cases, to the adoption of different habits by different members
of the same group. Were we to term the three orders Rock-urchins,
Sand-urchins, and Burrowing-urchins, it would not be entirely true,
for secondary invasions of the other's territory on the part of each
order have undoubtedly taken place; but still the statement would
remain roughly true, and would give a fair idea of the differences in
habitat which have led to the differentiation of the group.

Order I. Endocyclica (Regular Urchins).

The principal variations concern (1) the peristome, (2) the periproct,
(3) the corona, (4) Aristotle's lantern and its appendages, (5) the
spines, (6) the pedicellariae, and lastly, (7) the tube-feet. We shall
consider these points in order.

Peristome.—In the vast majority of species this region is covered

only with flexible skin in which ten small plates are embedded,
pierced by pores for the buccal tube-feet; besides these there are
irregularly arranged thin plates. In the Cidaridae both the ambulacral
and the interambulacral series of plates are continued on it; these
plates differ from those of the corona in being movable on one
another. In Echinothuriidae only the ambulacral series of plates is
continued on to the peristome. In the case of both these families
there are a considerable number of tube-feet within the region of the
peristome which may be classed as buccal.

Periproct.—This area, which represents the whole dorsal surface of

Asteroidea, is very large in the Cidaridae, where, as in Echinus, it is
covered with leathery skin in which small plates are embedded. In
the Saleniidae it is covered with a single large sur-anal plate, in the
edge of which the anus is excavated; in the Arbaciidae it is covered
with four valve-like plates; whilst in the remaining species its
condition is similar to that described in the case of Echinus
esculentus.

Corona.—In Echinothuriidae all the plates are separated by slips of

membranous skin, so that the test is flexible. In all other families it is
an unyielding cuirass. In the Cidaridae the pore-plates remain
separate throughout life, and are therefore identical with the
ambulacral plates. These are small and placed in two vertical rows,
and so the ambulacra are exceedingly narrow. In Echinothuriidae
there is some tendency to adhesion amongst the pore-plates; these
are of different sizes, and usually one larger and one smaller adhere
to one another. In all other species regular ambulacral plates are
formed at least in the lower part of the radii near the peristome by
the adhesion of the pore-plates in groups of two, three, or more.
Sometimes as many as nine pore-plates may thus adhere.

When adhesion takes place between the pore-plates it is of course

preceded by crowding, and this interferes with their equal
development. Some which extend so far horizontally as to meet their
fellows of the opposite side of the radius are called primary plates;
others which are small and wedged in between the larger ones are
called demi-plates. Systems of classification have been built up
(chiefly by palaeontologists) in which great stress has been laid on
how the primaries and secondaries enter into the constitution of the
compound plate, but it does not seem to the present author as if this
were at all a satisfactory basis for classification. All the pore-plates
are primarily equivalent, and the question as to which are interfered
with in their growth so as to become secondary is trivial. The so-
called Arbacioid type consists of one primary with a secondary on
each side; the Diadematoid type of three primaries, with occasionally
a secondary between the aboral and the middle primary; and finally
the Triplechinoid type of two primaries, with one or more secondaries
between them.

Aristotle's Lantern.—Under this head we may consider the

auriculae and gills as well as the jaws and teeth. In Cidaridae
external gills appear to be absent, but from the lantern coelom large
radial pouches project upwards into the general coelom cavity.
These pouches are supposed to be respiratory, and are termed
internal gills or Stewart's organs.[488] They co-exist with external
gills in Echinothuriidae and in Diadematidae, though in the last family
they are present only in a vestigial form, two being found in each
radius. The auricular arch both in Cidaridae and in Arbaciidae is
composed of two pillars which do not meet, but in the last-named
family they are based, as in Echinidae, generally on the ambulacral
plates, whereas in Cidaridae they arise from the interambulacral
plates (the ambulacral plates being here very narrow). The
epiphyses are absent in Cidaridae and Arbaciidae, and are imperfect
in Diadematidae.

Spines.—These organs are extraordinarily variable, and usually

differ very much in species of the same genus. In the vast majority of
species there is a limited number of long spines called "primaries,"
amongst the bases of which a large number of much shorter
"secondaries" are distributed. In Cidaridae the primaries are very
long and thick and blunt at the ends, and the secondaries form small
circles around their bases. The primaries in Cidaridae and the tips of
the primaries in Arbaciidae and Echinothuriidae are covered with a
special investment of extremely close, hard, calcareous matter very
different from the loosely fenestrated material out of which the
bodies of the spines of all species are composed. In Colobocentrotus
and Heterocentrotus the primaries are very thick and triangular in
section, whilst the secondaries on the aboral surface have expanded
outer ends, which form a close-set pavement protecting the
ectoderm from the shocks of the breakers. In Echinothuriidae the
primaries are short and so delicate as to be termed silky.

Pedicellariae.—In Cidaridae only gemmiform and tridactyle

pedicellariae are found. In the gemmiform the glands lie inside the
grooved blades instead of outside as normally, and they are covered
internally by ingrowths of calcareous matter from the edges. In
Echinothuriidae only tridactyle and trifoliate are found in most
species, but rudimentary gemmiform are found in one species and
well-developed ophicephalous in another. In some species
(Centrostephanus longispinosus) there are found gemmiform
pedicellariae which have lost the jaws but retained the glands. These
are termed "globiferae." Mortensen[489] uses minute details in the
structure of the pedicellariae to discriminate species and even
genera, but in this the present author is not prepared to follow him.