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The result showed that nearly all species have a marked predilection
one way or the other, but not all in the same way. Helix aspersa,
Arion empiricorum, six species of Limax, and three of Planorbis, are
lovers of darkness, while H. nemoralis, Succinea putris, and two
species of Limnaea are lovers of light. Physa fontinalis stands alone
in being quite indifferent.
M. Willem endeavoured further to discover whether any of the
Mollusca possessed ‘dermatoptic perception,’ or the faculty of
perceiving variation of light by means of the skin alone. He
accordingly repeated the above-mentioned experiments, having
previously extirpated the eyes in all cases. The result was
remarkable. In a few instances the experiment was not conclusive,
but H. aspersa, A. empiricorum, several species of Limax, and one
Limnaea shunned or sought the light just as they had done when
their eyes were present. A few marine Mollusca (Littorina littorea,
Trochus cinerarius, T. umbilicatus, Patella vulgata) were also shown
to be exceedingly sensitive to the impact of a shadow, whether with
or without their eyes.
Blind and Eyeless Mollusca.—In a large number of marine
Mollusca which habitually creep about half buried in wet sand (Bullia,
Sigaretus, Scaphander, Philine), eyes are altogether absent. In
some species of Natica and Sigaretus, and in Doris, eyes are
developed, but are enclosed in a thick layer of skin, through which
they can probably do little more than faintly appreciate different
degrees of light and darkness. Chiton has cephalic eyes in the
embryo, but loses them in the adult stage. The two great Auricula, A.
auris Judae and A. auris Midae, which habitually creep about in the
liquid mud of mangrove swamps, have entirely lost their eyes.
Certain pelagic Mollusca seem to have a tendency, which is not
easily explained, to lose their eyes or the power of seeing with them.
Thus Ianthina has no eyes at all. Pteropoda as a rule have no eyes,
and the few that have (Creseis, Cavolinia) possess only certain
pigmented spots placed near to the nervous centres. In the
Heteropoda, however, and the Cephalopoda, many of which are
pelagic, the eyes are unusually large.
 Fig. 91.—Sigaretus
laevigatus Lam., a
species frequenting
wet sand, and
destitute of external
eyes: F, anterior
portion of foot. (After
Souleyet.)
Eyes in Deep-sea and Underground Mollusca.—Deep-sea
Mollusca, as a rule, possess no visual organs, or possess them only
in a rudimentary state, but this rule has its exceptions. Dr. Pelseneer
found[289] no trace of eyes in two species of Pleurotoma from 1850
and 1950 fath., none in a Fossarus from 1400 f., none in a
Puncturella from 1340 f. A remarkable form of Voluta (Guivillea) from
1600 f. possessed eyes which could hardly be functional, as they
were destitute of pigment, and exhibited other changes of structure.
On the other hand, it is remarkable to notice that in three different
species of Trochus from 450 f., 565 f., and 1375 f., the eyes were
pigmented and well developed.
In land Mollusca which live beneath the surface of the ground or
in absolute darkness, the eyes are generally more or less modified.
Thus in Testacella, which usually burrows deeply in the soil, but
occasionally emerges into the open air, the eyes are very small, but
distinct and pigmented. Our little Caecilianella acicula, which is never
seen above the surface, is altogether destitute of eyes. A species of
Zospeum, a Helix, and a Bithynella from dark caves in Carniola have
suffered a similar loss. On the other hand, a small Hyalinia from a
dark cave in Utah (probably a recent addition to the cave fauna) has
the eyes normally developed.
Eyes of Onchidium.—Many species of Onchidium, a naked land
pulmonate which creeps on rocks near high-water mark, are
provided with dorsal eyes of various degrees of organisation, and in
numbers varying up to nearly one hundred. The tropical Onchidium
are the prey of a fish (Periophthalmus) which skips along the beach
by the aid of its large ventral fins, and feeds principally on insects
and Onchidium. Karl Semper suggests[290] that the eyes are of
service to Onchidium as enabling it to apprehend the shadow of the
approaching Periophthalmus, and defend itself by suddenly
contracting certain glands on the skin and expressing a liquid
secretion which flies into the air like shot and frightens the
Periophthalmus away. This theory for it is no more than theory—may
or may not be true, but it is remarkable that Onchidium with dorsal
eyes have precisely the same geographical distribution as
Periophthalmus, and that where no Periophthalmus exists, e.g. on
our own S.W. coasts, the Onchidium are entirely destitute of dorsal
eyes. In those species of Onchidium which have no dorsal eyes, the
latter are on the tips of the tentacles, as in Helix. The eyes are
developed on the head, and afterwards ascend with the growth of
the ommatophores, while in Helix the ommatophores are formed
first, and the eyes developed upon them.[291]
Dorsal Eyes in the Chitonidae.—The remarkable discoveries of
Moseley with regard to the dorsal eyes of Chiton were first published
in 1884.[292] He happened to notice, while examining a specimen of
Schizochiton incisus, a number of minute black dots on the outer
surface of the shell, which appeared to refract light as if composed of
glass or crystal. These ‘eyes,’ in all the species of Chiton yet
examined, are restricted to the outer surface of the exposed area of
the shell, never being on the laminae of insertion or on the girdle. In
certain sub-genera of Chiton the eyes are scattered irregularly over
the surface, in others they are arranged symmetrically in rows
diverging from the apex of each plate, but in old specimens the eyes
towards the apices are generally rubbed off by erosion or abrasion.
Moseley regarded the occurrence of scattered eyes as indicating an
original stage of development, when the eyes were at first disposed
irregularly all over the surface of the shell; the gathering into regular
rows showing a later stage.

Fig. 92.—Dorsal eyes of Chitonidae, showing the

various forms of arrangement in the first and
fourth valves of 1, 1a, Acanthopleura
spinigera Sowb., E. Indies, × 2; 2, 2a,
Tonicia suezensis Reeve, Suez, × 3; 3, 3a,
Acanthopleura granulata Gmel., W. Indies, ×
2; 4, 4a, Tonicia lineolata Fremb., Chili, × 2.
From specimens in the Museum of Zoology,
Cambridge.
The eyes appear to be invariably more numerous on the anterior
plate. Thus in Corephium aculeatum there are about 12,000 in all, of
which more than 3000 are on the anterior plate. In Schizochiton they
are arranged in very symmetrical rows, six of which are situated on
the anterior, and only two, sometimes only one, on the central plates.
In Tonicia marmorata the eyes are sunk in little cup-shaped
depressions of the shell, possibly to escape abrasion. As regards
shape and size, in Ch. incisus they are circular, and about 1/35 inch
in diameter, this being the largest size known; in Ch. spiniger and
Ch. aculeatus they are oval, measuring about 1/100 x 1/600 inch.
There are no eyes in Chiton proper, nor in Mopalia, Maugeria,
Lorica, and Ischnochiton.[293] None of our English species appear to
possess them.[294]
Eyes in Bivalve Mollusca.—Some, possibly most, of the
Pelecypoda possess, in the larval state, true paired eyes at the oral
end of the body. These become aborted as the animal develops,
since that part of the body becomes entirely screened from the light
by the growth of the shell. To compensate for their loss, numerous
ocelli, or pigmented spots sensitive to the action of light, are in many
cases developed on different parts of the mantle, functionally
corresponding to the ‘eyes’ of Chiton described above. As in Chiton,
too, we have here an interesting series of instances in which true
eyes have suffered total obliteration, through disuse, and, as if to
restore to the animal in some measure its lost sense, visual organs
of a low power have subsequently been developed and are now
observed in various stages of specialisation.
Concentration of Eyes in Special Parts of the Mantle.—Sharp
has shown[295] that in several species of Ostrea, Cardium, Anomia,
Lima, Avicula, Arca, and Tellina pigmented cells, with a highly
refractive cuticle, are scattered over a considerable portion of the
mantle. Experiment has proved the powers of ‘vision,’ i.e. of
sensitiveness to different degrees of light, possessed by these
organs. In Dreissena polymorpha, Tapes decussatus, and two
species of Venus these cells are concentrated on that particular part
of the mantle which is not always covered by the shell, i.e. the
siphon, but since the siphon can be completely retracted within the
shell, there is no special provision for their protection. A further step
is shown in the case of Mya arenaria, where the siphon is scarcely
capable of complete retraction. Here, while some of the pigment
cells are scattered about over the surface of the siphon, the majority
are placed in grooves at the base of the siphonal tentacles, forming
an intensely black band round them. A higher stage still is shown in
Solen vagina, S. ensis, and Mactra solidissima, where the cells are
situated only in the siphonal grooves, which are more or less
specialised in numbers and complexity.
Arca Noae, according to Patten, is very sensitive to any sudden
change in the amount of light falling upon its mantle-edge. A faint
shadow cast upon it by the hand is sufficient to cause it to close its
valves quickly, but always one or two seconds afterwards, the
promptitude in all cases depending upon the depth of the shadow.
Sensitiveness in this direction was found to depend greatly upon the
vitality of the animals themselves, since it always became less in
those specimens which had been kept for long in confinement. A
shadow was not always necessary to make them close. An ordinary
black pencil, if approached within two or three inches with extreme
caution, produced the same result, while a glass rod brought within
the same distance, and even moved rapidly to and fro, appeared to
cause no alarm. Sensitiveness to change in intensity of light was
experimentally noticed by the same author in the case of Ostrea,
Mactra, Avicula (to a special extent), and Cardium. It is very
remarkable to find that increased elaboration in the structure of the
eyes does not necessarily carry with it increased sensitiveness, i.e.
higher visual powers. Avicula, which is only provided with a few
scattered ommatidia, which would entirely escape the notice of any
one who had not seen them better developed elsewhere, was
considerably more sensitive to light and shade than Arca, with its
eyes of conspicuous size and much more perfect organisation,
instantly contracting the mantle upon the impact of a shadow so faint
as to be invisible to the experimenter.[296]
Visual Faculties of Solen and Ostrea.—The visual power of
Solen may be exemplified by any one who is walking along almost
any of our sandy bays at extreme low-water mark. If the day be
warm and sunny, numbers of Solen will be seen raising themselves
an inch or two out of their holes; but if you wish to catch them you
must approach very cautiously, and on no account allow your
shadow to fall upon them, or they will pop down into their burrows in
an instant, and it is vain to attempt to dig them out. ‘How sensitive,’
remarks Mr. W. Anderson Smith, with reference to oysters,[297] ‘the
creatures are to the light above them; the shadow [of the boat] as it
passes overhead is instantaneously noted, and, snap! the lips are
firmly closed.’
Ocelli of Pecten.—In Pecten and Spondylus the ocelli are
remarkably large and prominent, shining like precious stones, and
are placed along the two edges of the mantle so as to receive the
light when the shell gapes (Fig. 93). In Pecten opercularis,
jacobaeus, and maximus their number varies from 80 to 120. In
Spondylus gaederopus, a very inequivalve shell, 60 have been
counted on the right or fixed valve, and 90 on the left or upper valve.
Each ocellus is connected, by means of its optic nerve, with the large
circumpalleal nerve, and so with the branchial ganglion. They
possess a cornea, lens, choroidea, and optic nerve, and, according
to Hickson,[298] bear a considerable resemblance to the vertebrate
type of eye. In spite of this, the power of vision in these genera does
not appear at all superior to that of other Pelecypoda.

Fig. 93.—Pecten opercularis L., showing

the ocelli on the two edges of the
mantle.
 Fig. 94.—Compound eyes (c.e) of Arca
barbata L.; m.l, mantle fold; omm,
ommatidia. (After Patten.)
According to the elaborate investigations of Patten, the ‘eyes’ in
Arca occur upon the middle or ‘ophthalmic’ fold of the mantle-edge,
which is thickened at the end to admit of their reception. Along this is
ranged a row of dark brown spots of various sizes, which are larger
at the anterior and posterior ends of the mantle-edge, but smaller
and more numerous towards the middle. These brown spots, or
‘eyes,’ are many of them compound, being made up of the fusion of
a number of ommatidia (from 10 to 80) into one large round eye,
which is generally elevated above the surface of the surrounding
epithelium. Sometimes these eyes themselves tend to fuse together.
In one specimen of Arca Noae, 133 of these faceted eyes were
counted in one mantle border, and 102 in the other.
There can be little doubt that the development of these functional
eyes, or sensitive spots, in bivalve Mollusca, is due to special needs.
They appear to be entirely absent in fresh-water bivalves (with the
exception of Dreissensia, which is obviously a marine genus recently
become fresh-water), while they are most abundant in genera living
between tide marks (Solen, Mya, Mactra), and most highly
specialised in a genus that is, for a bivalve, of singularly active habits
(Pecten). Now genera living in sand between tide marks, as the
three above-mentioned genera are in the habit of doing, and also
protruding their siphons, and occasionally a considerable portion of
their shells, out of their burrow, are manifestly very much at the
mercy of their watchful enemies the gulls, and anything which would
enable them to apprehend the approach of their enemies would be
greatly to their advantage. Here, perhaps, lies the explanation of the
greater elaboration of these pigmented spots in littoral genera, as
compared with those inhabiting deeper water. Pecten, again, a
genus distinguished by great activity, which can ‘fly’ for considerable
distances in the water by flapping its valves together and expelling
the water from the apertures at either side of the hinge, may be
greatly assisted by its ocelli in directing its flight so as to escape its
enemies.

III. Smell
The sense of smell—touch at a distance, as Moquin-Tandon has
called it—is probably the most important sense which the Mollusca
possess, and is unquestionably far more valuable to them than that
of sight. Any one who has ever enjoyed the fun of hauling up lobster
pots will recollect that part of the contents was generally a plentiful
sprinkling of Buccinum, Nassa, and Natica, attracted by the smell of
the stinking piece of fish with which the trap was baited. According to
Mr. J. S. Gibbons,[299] Bullia rhodostoma congregates in hundreds
on gigantic medusae which are stranded on the sandy bays near the
Cape of Good Hope. Dr. J. G. Jeffreys says[300] that quantities of the
common Neptunea antiqua “are procured on the Cheshire coast by
the fishermen placing a dead dog on the sands at low-water mark
during spring tides. The bait is then completely covered with stones,
which are piled up like a cairn. On the next turn of the tide the heap
of stones is visited, and the whelks are found on the surface in great
numbers, having been apparently attracted by the smell of the bait,
but unable to get at it.” Mr. W. A. Lloyd kept specimens of Nassa
reticulata in a tank in the sand, at the bottom of which they usually
remained buried. If a piece of meat of any kind were drawn over the
sand, the Nassa would appear above the surface in a few minutes.
Half-picked beef or mutton bones, if placed in the tank, were covered
in a few minutes. In fact, no animal matter, whether living or dead,
could be introduced without the Nassa smelling it, and coming up to
see what they could get.[301]
Any one can experiment for themselves on the olfactory powers of
our common snails or slugs. Moquin-Tandon records[302] two
interesting cases, one communicated to him by letter, the other
occurring to himself. His correspondent, a M. Parenteau, was one
day walking along a dusty high-road, when he noticed, near the
middle of the road, an empty bean-pod and two Arions eating it.
Attributing the meeting of feeders and food to mere chance, he was
walking on, when he noticed a second bean-pod, and, about two
yards away from it, a third Arion, hurrying straight towards it. When
the Arion had yet more than a yard to traverse, M. Parenteau picked
up the bean and put it in his pocket. The Arion stopped, raised its
head, and turned in every direction, waving its tentacles, but without
advancing. M. Parenteau then carried the bean to the other side of
the road, and put it in a small hole behind a piece of stone. The
Arion, after a moment’s indecision, started off straight for the bean.
Again the position of the precious morsel was changed, and again
the Arion made for it, this time without being further tantalised. M.
Moquin-Tandon noticed, one rainy day in the botanical gardens at
Toulouse, two Limax maximus approaching a rotten apple from
different directions. He changed the position of the apple several
times, placing it at a sufficient distance, to be sure they could not see
it, but they always hit it off correctly, after raising their heads and
moving their long tentacles in every direction. It then occurred to him
to hold the apple in the air, some centimetres above the head of the
Limax. They perceived where it was, raised their heads and
lengthened their necks, endeavouring to find some solid body on
which to climb to their food.
Several of the land Mollusca have the power of exhaling a
disagreeable smell, Hyalinia alliaria smelling strongly of garlic, and
Stenogyra decollata of laudanum; but this need not be any argument
for the sense of smell in the creatures themselves.
Position of Olfactory Organs in Pulmonata.—Most authorities
are of opinion that the olfactory organs are situated in the tentacles.
Moquin-Tandon considered that in the Helicidae and Limacidae the
sense of smell is confined to the little knob or elevation at the end of
the longer tentacles, close to the eye. He found that when he cut off
these tentacles both in Limax and Arion, the creatures were quite
unable to discover the whereabouts even of strongly-scented food.
The same author believed that in the Basommatophora the sense of
smell was present in the whole of the tentacle, which is covered with
an exceedingly sensitive ciliated epithelium. Lacaze-Duthiers,
however, places the olfactory sense in this group at the outer side of
the base of the tentacles, near to the eyes. Some authorities[303]
deny that the Helicidae have the olfactory organ at the tip of the
tentacles, and locate it in a pedal gland near the mouth, which
contains conspicuous sensitive cells. A Helix whose tentacles had
been removed manifested its repulsion to the smell of spirits of
turpentine, while another Helix, which was unmutilated, did not
object to the turpentine being held between its tentacles. Altogether,
then, the exact position of the smell-organ in the Helicidae must be
considered as not yet thoroughly determined. Simroth holds that the
sense of smell is distributed over the whole soft integument, and is
especially concentrated in the feelers, and in the neighbourhood of
the respiratory orifice.[304]
In nearly all marine Mollusca yet examined, the organ of smell or
osphradium is in situation intimately connected with the breathing
organs, being generally placed near their base, with the object,
apparently, of testing the quality of the water before it passes over
the branchiae. It consists of a patch of the epithelium, modified in a
special manner, and connected by its own nerve with one of the
visceral ganglia.
An osphradium does not necessarily occur in all genera; for
instance, it has not been detected in Fissurella. It is most highly
specialised in the Conidae, and in the carnivorous Gasteropoda
generally. In Buccinum undatum, for instance, it is very large indeed,
and, from its plumed form, has sometimes been mistaken for an
accessory branchia (Fig. 95). In Haliotis it is paired, one lying in
close proximity to each of the two branchiae, but in Turbo it is single,
corresponding to the single branchia. In Chiton there is an
osphradium at the base of each separate gill filament, making a total
of twenty or more on each side. Its position in Physa and in
Cyclostoma will be seen by reference to Figs. 103 and 104 (p. 205).
In the Pelecypoda the osphradia are paired, and lie adjacent to the
posterior adductor muscle, close to the hinder end of the axis of the
branchiae. In the Tetrabranchiate Cephalopoda there are two
osphradia, placed between the bases of the two pairs of gills. In the
Dibranchiates on the other hand, a groove above the eyes has been
regarded as the seat of the organ of smell. This groove contains
sensory and ciliated cells, and appears to be connected with a
special nerve centre of its own, which ultimately is derived from the
cerebral ganglion.

Fig. 95.—Buccinum undatum L.,

deprived of its shell, showing the
relative position of branchia (br)
and osphradium (os); m, mucous
glands; s, siphon. The portion of
the mantle covering the
osphradium has been removed.
Scarcely any instances of the exercise of the sense of smell on
the part of bivalve Mollusca have been recorded. Something of the
sort, however, seems to have been present in a case related by Mr.
R. L. King.[305] A skull of a fox had been placed in a small ditch in
order to soak, and after a few days, when taken out, was found to be
covered with Pisidium pusillum to the number of at least two
hundred, which had been probably attracted from the water in the
immediate neighbourhood by the smell of the decaying flesh.

IV. Hearing
Experiments made with a view to ascertain whether the Mollusca
are sensitive to noises have usually led to the conclusion that they
are deaf to very loud sounds. This is the more curious, because an
undoubted auditory apparatus has been discovered in a large
number of genera. In the case of an experiment, it is not easy to be
sure that the animal is not affected, at least in part, by the shock or
jar, rather than by the actual sound. In some experiments, however,
conducted at the Plymouth Marine Biological Laboratory, Mr.
Bateson found[306] that Anomia could be made to shut its shell by
smearing the glass of the tank with the finger in such a way as to
make a creaking sound. It was evident that the cause of alarm was
not the jarring of the solid framework of the tank, for the same result
occurred when the object on which the Anomia were fixed was
suspended in the water by a thread. It was found that the sound had
to be of a particular pitch to excite the attention of the mollusc.
As a rule the organ of hearing is nothing more than a small vesicle
or sac (the otocyst), filled with a fluid secretion, in which are
suspended one or usually more calcareous concretions known as
otoliths. By means of cilia, which connect with sense-cells, these
otoliths are given a peculiar movement or oscillation in the medium
in which they are suspended. The number of the otoliths varies in
different genera and species; there are several hundreds in Arion
and Limax, about a hundred in Helix pomatia, nemoralis, hispida,
arbustorum, rotundata, Succinea putris, and Limnaea stagnalis;
about fifty in Planorbis contortus and Physa fontinalis, only one in
Cyclostoma elegans. The number increases with age. In young
specimens of Limn. stagnalis as few as ten, nine, and seven have
been noticed.[307]
The otocysts are always paired, and, in Gasteropoda, are placed
close to the pedal ganglia. The acoustic nerve, however, has been
shown by Lacaze-Duthiers to connect with the cerebral ganglia in
certain cases. The otocysts are never on the surface of the body and
are rarely connected with it by any passage or tube; it is probable
therefore that sound reaches them simply through the medium of the
tissues.
In the Pelecypoda the otocyst is similarly situated near the pedal
ganglion, and is probably (though this has not yet been proved)
similarly connected with the cerebral. There is only a single otolith.
Pelseneer finds[308] in Nuculidae alone a free communication
between the otocyst and the exterior. Anodonta has been
observed[309] to withdraw its foot into the shell at the noise of an
opening door, a loud voice, or a shrill whistle, whether in a basin of
water or lying on a study table.

Fig. 96.—Illustrating the otocyst in A, Anodonta, B, Cyclas; ot,

otolith; a, b, c, c´, cellular layers surrounding the chamber;
ci, cilia on interior walls of chamber: C, an otolith crushed.
(After Simroth.)
Delage extirpated the otocysts in certain Octopoda, and obtained
some unexpected results. He found that remarkable effects were
produced upon the animal’s powers of locomotion, so that it was
unable to preserve its proper balance in the water when in rapid
motion, but its body was forced to undergo a form of rotation more or
less pronounced. He concluded that the otocysts must possess,
besides their auditory functions, a power which stands in some
relation to the proper orientation of the body in locomotion, a power
which is not wholly supplied by sight and touch alone. The otocysts
may thus regulate locomotion by stimulating muscular acts which
tend to keep the body in the straight line during the process of
movement.[310]

The Foot
 One of the most characteristic organs of the Mollusca is the foot,
which, under one form or another, occurs throughout the whole
phylum. The foot is a thickening, on the ventral side, of a portion of
the integument of the animal, modified to serve different forms of
motion. It attains its maximum relative area in the Chitonidae, many
Nudibranchs, and the slugs generally, in nearly all of which there is
no portion of the body which is not subtended by the foot. Here too it
presents the form of a regular disc or ellipse, which is more or less
produced. In many cases, however, the foot becomes modified in
such a way that we are enabled to recognise well-marked anterior
and posterior portions, which have received the name of propodium
and metapodium respectively, while the intervening central portion is
termed the mesopodium.

Fig. 97.—Sigaretus laevigatus Lam., showing excessive

development of the propodium (pr) and metapodium (met)
in a mollusc living in sand (the shell, which covers only the
liver and adjacent parts, has been removed); l, liver; s.ap,
aperture of proboscis, here deflected from the median line;
t, t, tentacles. (After Quoy and Gaimard.)
The propodium is most strongly developed in genera which crawl
about in wet sand, e.g. Natica, Sigaretus, Oliva, Harpa, Scaphander
(Figs. 97 and 98, and compare Fig. 91). In such cases it seems to
serve as a sort of fender or snow-plough, to push the sand away on
both sides of the path the animal is traversing. In some species of
Sigaretus the propodium becomes as it were banked up against the
head and proboscis, which are thus unnaturally elevated, or tend to
disappear altogether. Bullia (Fig. 62), which crawls about rapidly on
wet sand, appears to attain its object by a wide extension of the foot
on all sides, and so slides over the sand instead of ploughing
through it; the little lappets at the end of the ‘tail’ probably serve as
rudders.
In Melampus and Pedipes the propodium is marked off by a
groove across the ventral surface. When the animal is in motion it
first advances the propodium and then pulls the rest of the foot after
it with the looping gait of certain caterpillars. In many
Cyclostomatidae this groove, instead of being transverse, is
longitudinal, and the animal advances first the right and then the left
segment of the foot, which gives it a swaying motion from side to
side.
Upon the metapodium lies the operculum, when it occurs. As a
rule the metapodium is not sharply marked off from the rest of the
foot. In Strombus (Fig. 99) it becomes erected into a sort of hump or
column, on the top of which the operculum is situated.

Fig. 98.—Oliva textilina Lam., showing how the front part

of the foot (f) is developed into a sort of fender, the
propodium (pr); e, e, eyes; m.ap, front appendage of
mantle; m.ap´, hinder appendage of mantle, folded
into the suture when the animal is at rest; si, siphon; t,
t, tentacles. (After Quoy and Gaimard.)
The epipodium is a prominent fold or border, which occurs upon
the upper edge of the foot in most Diotocardia. In Haliotis it is of
considerable breadth, and is covered by a number of lobes which
spring from a moss-like prolongation of the skin. From the epipodium
are developed the lateral tentaculae of Monodonta (Fig. 82, p. 178),
and of other sub-genera of the Trochidae.[311]
In the Opisthobranchiata the lateral edges of the foot (the
parapodia) are frequently produced into broad folds or wing-like
extensions, which in many cases tend to fold over the shell, and, in
conjunction with the mantle, eventually imbed it altogether. By the
wavy motion of the parapodia the animal is enabled to progress
through the water. The paired natatory lobes of the Pteropoda are
simply the parapodia of the Tectibranchs modified for swimming
purposes.

Fig. 99.—Strombus lentiginosus Lam., showing

the modified form of the foot (f): e, e, eyes on
their pedicels; mp, metapodium; op,
operculum; p, penis; pr, proboscis; t, t,
tentacles. (After Quoy and Gaimard.)
It is in the Heteropoda, Pteropoda, and most of all, the
Cephalopoda, groups which have, for the most part, exchanged a
crawling for a swimming life, that the modifications of the foot are
most considerable. In Oxygyrus and Atlanta, for instance, the
propodium and metapodium are sharply distinguished from the
mesopodium, and no doubt have acquired, as a means of
propulsion, the power of separate movement, the animal swimming
with these portions of the foot uppermost. In Carinaria and
Pterotrachea the metapodium has probably become continuous with
the long axis of the body, while the so-called ‘foot’ with its sucker
represents only the original propodium. In the Cephalopoda the arms
and funnel represent the modified foot, the sides of which are
prolonged into a number of very long specialised tentaculae. In the
adult Cephalopod some of the arms have assumed a position in
advance of the mouth, the latter being in fact surrounded by a circle
of arms. But in the Cephalopod embryo the mouth opens as in the
Gasteropoda, i.e. in advance of the arms, and it is only gradually that
it becomes encircled by them. Arms and funnel alike are found to be
innerved from the pedal ganglion.[312]
The pointed axe-shaped foot, which is characteristic of the
majority of Pelecypoda, is doubtless derived from a form more akin
to the flattened ‘sole’ of the Gasteropoda. A foot with something of
this disc-shaped base actually occurs in some of the Nuculidae, the
parapodia being furnished with pleats which recall similar formations
in other Orders (Fig. 100). The principal modifications of the foot are
due to its employment as a burrowing organ. In genera which burrow
but slightly it is small and feebly developed, while in genera which
habitually excavate, it becomes the largest and strongest organ of
the body. At the same time it has a tendency to shift its position from
the ventral to the anterior margin, accompanied by a corresponding
narrowing of the shell, until it arrives at the position seen in Mollusca
of the shape of Mya, Pholas, and Solen. In sedentary or attached
genera, e.g. Pecten, Chama, Ostrea, the foot tends to become
aborted.
 Fig. 100.—Yoldia limatula Say, Greenland, showing
the short plumed branchiae (br, br), the
gasteropodous foot (f), and the large labial palps
(l.p, l.p): A, as seen from the ventral margin; B,
from the left side, with the mantle turned back;
a.m, position of anterior adductor muscle; i,
intestine; l, liver; m, m, mantle.
The byssus gland, secreting a byssus of horny threads, is
characteristic of many Pelecypoda, and may be observed by any
one in the common mussel. It occurs in the larvae of many species
which do not possess a byssus in the adult stage. The pedal gland of
many Prosobranchiates, which secretes a tough and almost thread-
like slime, is possibly homologous with the byssus gland of bivalves.

Nervous System

The Mollusca possess a nervous system, which usually consists

of a number of nerve centres or ganglia, linked together by bands
(the commissures) and sending out thread-like nerves which ramify
into the various organs. The character of the nervous system varies
greatly in different groups, ranging as it does from a condition of
extreme complexity, in which the ganglia are numerous and the
commissures equally so, to that of considerable simplicity, in which
ganglia are almost entirely absent.
 The most important ganglia are (1) the cerebral,[313] which are
always placed above or on either side of the mouth, and from which
proceed the nerves of the eyes and tentacles; (2) the pedal, which in
Gasteropoda are situated below the oesophagus, in Pelecypoda at
the base of the foot, and from which the nerves of the foot and
sometimes the acoustic nerve arise; (3) the pleural,[314] whose
position varies considerably, but is always below the oesophagus
and slightly above the pedal ganglia; these innervate the mantle,
branchiae, heart, and viscera generally.
Gasteropoda.—The simplest form of nerve system as thus
understood occurs in the Amphineura, and more particularly in the
Chitons. Here we find four longitudinal nerve-cords, parallel to one
another for nearly the whole length of the mollusc. The two exterior
cords probably represent the pleural, the two interior the pedal
nervous system. There being no head or tentacles, but simply a
mouth at the anterior end, the cerebral ganglia do not exist, but they
are represented by the curved ring formed by the massing together
of the two nerve-cords on each side. The only distinct ganglia are a
pair of buccal ganglia (which are developed on a pair of
commissures which pass forward from the cerebral mass and
innervate the lips and buccal region), and a much smaller group, the
sublingual. The two pedal cords are united by a number of
transverse parallel connectives, which recall similar modes of
connexion in the Chaetopod worms and in Arthropoda.
This quadruple set of nerve-cords is characteristic of all the
Amphineura, but the absence of ganglia is most marked in the
Chitons. In Proneomenia and Neomenia there is a distinct cerebral
ganglion, formed by the massing of the two ganglia into one, while in
Proneomenia the lateral cords are joined to the pedal, as well as the
pedal to one another, by connectives. In Chaetoderma the cerebral
ganglia, though adjacent, are distinct, and both the pedal and lateral
cords connect directly with them, while there are no transverse
connectives.
The remaining three great divisions of Gasteropoda, namely, the
Prosobranchiata, Opisthobranchiata, and Pulmonata, may be