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Introduction
Because the Mollusca are designated as a phylum we know that all the organisms in the taxon
must share a common body plan that they have inherited from the ancestral mollusc; its the
very definition of a phylum after all. But, if we were to put a representative from each of the
different classes, both extinct and extant, on the lab bench we would have a tough time trying
to find the characteristics of the ancestral body plan that unifies the classes of the phylum.
Thats because each class is a spectacular examples of invertebrate adaptive radiation with the
result that seemingly unrelated animals, such as snails, slugs, clams, the octopus and squids are
all related to each other and share a single common ancestor.
Whatever it looked like the ancestral Molluscs body plan was sufficiently flexible that it
allowed its descendents to adapt with a range of behaviors, and habitats. From sedentary filter
feeders to rapid swimming active predators, and just about everything in between, there is a
mollusc that does it. The adaptive potential of the ancestral molluscs is also seen in where you
find molluscs; freshwater, marine and terrestrial environments.
Their general appearance wouldnt necessarily give us any obvious clues to why these animals
are all included in one phylum but there are a variety of traits, or characters, that unify the group
and you should look for these in each of the specimens. These include the presence of a
muscular foot, involved in locomotion, and a sensory head that was also associated with food
acquisition by the unique molluscan feature of a radula. Another is the visceral mass, which is
dependent on cilia for its function, and positioned dorsally surrounded by the protective shell.
The shell is secreted by the underlying mantle extending out from the body creating a mantle
cavity inside which youll find the ctenidia, the principle respiratory structures of molluscs.
Polyplacophora
These animals are referred to as chitons and their body form is specially adapted for the rough
conditions associated with the intertidal zone of the oceans. When chitons are active they slowly
creep across the rocks feeding on encrusted algae and other organic debris. If threatened they
can roll up into a ball surrounded by the protective armor of their shell.
The large oval foot dominates the ventral surface of a chiton and along its lateral edges are the
mantle cavity includes grooves formed from a trough between the foot on the inside and the
fleshy girdle. Inside the mantle cavity you can see the multiple ctenidia used for gas exchange.
The mouth is easy to see at the anterior end but there are neither eyes nor tentacles associated
with it. At the opposite end, the anus is located on the roof of the mantle cavity, on the tip of a
small papillae. Like all molluscs cilia on the surface of the ctendia propel water through the
mantle cavity pulling it in at the anterior end surrounding the mouth, down the two mantle
cavities on each side, and over the ctenidia. At the back the left and right mantle cavities fuse to
form a single exhalent canal where the anal opening is located. If you look closely in the region
of the last few ctenidia you may also be able to see nephridiopores or gonopores that open into
this posterior part of the mantle cavity.
Gastropoda
Gastropod shells
One gastropod trait that is easy to recognize is their spiralled shell; the most effective way to
package the increasing size of the visceral mass as the snail grows. The shell consists of only
one piece, a univalve, molded into a cone wound on itself. Essentially the snail winds its shell
up and lives underneath it. A snails shell is a protective safe haven where it can pull in its whole
Examine the intact and sectioned conch shells to understand the spiralled nature of the shell. The
main parts of the body includes the head and the foot extending out of the opening of the shell,
the aperture. Not all apertures open in the same direction. Place the apex of the shell away from
you. Is the apertures opening on the left or right? If its on the right is dextral, on the left
sinistral just like right and left-handedness. One complete circle of the shell is a whorl and the
edges of each whorl are connected to the next by suture lines. These lines are often sculptured
and can form spines. Like all mollusc shells, growth lines are visible on the surface of the shell
and the oldest part of the shell is the apex. In the shells that have been cut open identify the
columella. Are there individual chambers in the shell, or is it one single continuous
chamber?
Not all gastropods have shells; slugs have discarded the cumbersome shell.
Helix
Helix is a terrestrial herbivore that prefers to feed at night, or after a rain when conditions are
damp. Many gardeners battle the nocturnal dining of snails and slugs in their gardens. They
overwinter by crawling under rocks or digging into the soil.
External Anatomy
Take a relaxed specimen; its body extends outside of the shell. Locate the apex of the shell and
the anterior and posterior ends of the animal. Only the head and foot extend from the shell, the
visceral mass remains protected inside. The mantles edge is thickest in the region of the collar.
The pneustome, the opening to the mantle cavity, is located on the surface of the collar. One
of the best ways to find the pneustome in preserved specimens is to gently squeeze the body of
the snail and watch the collar for where preservative comes out. These are terrestrial animals
and the mantle cavity has been modified as a lung that we will see later when we remove the
shell.
The anterior position of the opening to the mantle cavity is a consequence of torsion, another
is the anal opening positioned in front of the animal instead of behind. Locate the anus just
beneath the pneustome, and the mouth with its three lips. All three openings are located on the
anterior part of the body, and when you see this in a mollusc you know you have a gastropod.
Where are the openings to the anus and mantle cavity usually located in molluscs? The
Circulatory and excretory system Submerse your specimen under water and to one side
of the dissecting dish so that you will be able to look at it under the dissecting scope. Use a pair
of fine scissors and cut through the collar in the region of the pneustomes. Cut along the edge
of the collar towards the rectum and then along the length of the mantle on the inner edge of the
whorl. Pin aside the roof of the mantle cavity and identify the blood vessels; efferent blood
Later, when you have finished your observations of the reproductive system, open up the
pharynx to see the radula inside. Below the radula locate the jaw. The radula works against the
jaw to grind up the ingested meal.
Nervous system The main nerve ring may be apparent over the buccal bulb. The remaining
part of the nervous system is difficult to trace as a consequence of torsion and there really isnt
any reason to try and find it.
Reproductive system Snails are monoecious and youll be able to see most of the
structures of the male and female reproductive systems. Be sure to spend a bit of time unwinding
things so that you can see the main features. When you cut through the floor of the mantle cavity
to expose the digestive system you will have also exposed the different parts of the reproductive
system. We will need to find a few landmarks from which we can map the remainder of the
system. The mucus glands, with their filamentous components and the dart sac are the easiest
to find. As its name suggests the dart sac contains a dart that is jabbed into the side of this snails
partner during copulation. Is the dart a part of the male or female reproductive system?
Gastropod diversity
Examine the different gastropod specimens. How have they adapted to the problems associated
with torsion? To answer this question you might want to follow how water moves through the
mantle cavity and note the position of the anal or excretory openings. Specimens if available
include: limpets, the garden slug, Limax, conch, and nudibranch
Scaphopoda
There are two plastic mounts that show the internal anatomy of the tusk shelled Scaphopods.
These animals have a characteristic tooth-like shell and the mantle cavity runs the length of the
shell. Compare these specimens to the diagrams showing the internal anatomy of these animals.
Bivalvia
Clams and their bivalve relatives are one of only a few mollusc classes found in both freshwater
and marine environments. In both places they're specialized as filter feeders, and like most filter
feeders they're essentially sessile. To accommodate their way of life, bivalves have made some
major modifications to the mollusc body plan. The most obvious is that the body is compressed
laterally with the mantle folded down over the sides in two lobe-like sheets. The shell matches
the shape of the mantle's two parts. The shell has two valves that cover each side of the animal
and an elastic hinge ligament connects the sides across the top. The two valves give the class
their name, the bivalvia. But, be careful. These are univalve molluscs because the original single
valve is only folded!
The lateral compression of the bivalve body not only changes the shape of the shell but also the
organs of the visceral mass. Instead of sitting above the muscular foot, the visceral mass is
partially buried in the foot. The only part still visible above the foot is the pericardial cavity on
the dorsal side. Another modification for the filter-feeding lifestyle has resulted in the loss of
the radula and head.
Examine the inner surface of the shell you removed. Near each end are the large adductor
muscle scars. These muscles run from the left to right sides connecting the two valves to each
other. Contraction of the shell adductor muscles closes the shell and the hinge teeth insure that
the two shells align properly as it closes. How does a clam open its shell? Next to each of the
adductor muscle scars is a scar from the attachment of the foot retractor muscle. The anterior
muscle scar lies a bit above and behind the anterior adductor. The posterior retractor lays next
the posterior shell adductor. As their name implies the foot retractor mucles pull the foot into
the shell. How does a clam extend its foot? The pallial line extends from the adductor muscle
scars and runs along the edge of the shell. This scar is created by the attachment of muscle along
the mantle edge.
Break off a piece of shell from the valve you have removed and identify its three layers. These
include an inner mother of pearl layer, or nacreous layer; a middle prismatic layer and the outer
periostracum.
Internal Anatomy
If it isnt already, fold the mantle back over the body of the clam lying inside the left shell. The
mantle is fused to the dorsal surface of the animal and on the ventral side it hangs free and
encloses the mantle cavity. With the mantle still covering the body look at the posterior edge.
The incurrent and excurrent openings are only visible when the two sides of the mantle lie next
Cut a piece of gill from your specimen and look at its edge under the dissecting microscope to
see the paired lamella, the connections between them and the water tubes inside. Examine the
side of the same piece under low power to see the grid of ridges and connectives that form the
demibranchs. Blood vessels are embedded in the tissue and as the water flows through, the
blood is oxygenated.
Excretory system The metanephridia are located just underneath the pericardial cavity
and in front of the posterior adductor muscle. They appear as dark glandular structures and the
opening in the wall of the pericardial cavity is difficult to see. The metanephridium is composed
of two parts the glandular part near the heart, and the duct that carries the urine to the
nephridiopores opening on the side of the visceral mass near the base of the inner demibranchs.
This opening is also hard to see. Fold the gills up and see if you can locate it. How do metabolic
wastes pass from the circulatory system to the metanephridia?
Digestive system The same water used to aerate the gills also contains particulate food.
Cilia on the surface of the gill move the particulate food to the ventral margin of the
demibranchs and cilia along the grooved edge pass it forward to the labial palps. There are
two pairs of labial palps, inner and outer, and these fuse above the mouth. The labial palps are
also covered in cilia and their beating propel food particles into the mouth. Weve already made
a note of the intestine running through the pericardial cavity; follow it towards the anus that
empties into the suprabranchial chamber.
The rest of the digestive tract is contained in the visceral mass embedded in the foot. To be able
to see this youll have to carefully remove the muscular tissue of the foot. This is best done by
using a sharp scalpel and gradually slicing away tissue starting in the region of the labial palps.
The slices should be from dorsal to ventral and parallel to the main axis of the body. With each
slice youll expose a bit more of the underlying digestive system starting with the mouth and
Reproductive system Although the sexes are separate there is no easy way to tell the sex
of your specimen because morphologically the gonads are the same in both sexes. If youre
specimen has the modified gill with glochidia inside then youve got a female. The tissue of the
gonads spreads throughout the foot and is often difficult to distinguish from the spongy
musculature of the foot. Youll need to use your dissection scope and a well-flooded specimen
to be able to see the difference between the two.
Nervous system The nervous system is difficult to see and in fresh specimens the various
ganglia will have a pinkish color. It consists of three pairs of ganglia; cerebral, pedal, and
visceral ganglia. The visceral ganglion is located on the surface of the posterior adductor
muscle. The brain is located just above the mouth and behind the anterior adductor muscle.
Three major nerves radiate from the brain and connect to the visceral ganglion, the pedal
ganglion and the mantle, and the pallial nerve. The pedal ganglion is located in the base of the
foot
Cephalopoda
A cephalopods visceral mass has been stretched along the dorsoventral axis above the foot,
bringing the head and foot closer together on the ventral side. Thats how they got their name,
Cephalopoda (head, foot). The mantle surrounds the visceral mass, and ancestrally a hard shell
surrounded all of this to form an elongated cone-shaped shell with the head and foot poking out
the open end. It was easier to point the tip of the shell in the direction that the animal was
moving. Cephalopods swim with what was their original dorsal surface pointing in the direction
they move, rather than up. In most animals the surface of the body facing into the direction of
movement would be the functionally anterior surface and this is usually the same side of the
body where the mouth and head are located. In cephalopods the anterior head and mouth has
now become the new dorsal side although the squeezing together of the foot compressed the
anterior/posterior axis and the oral opening and head still have a functionally anterior position
but its not facing in the direction that they swim. The result of this is that cephalopods swim
backwards!
With a larger visceral mass on top of a muscular foot, cephalopods were faced with the same
problem as gastropods had for finding a way to make a more compact body. They used the same
solution, although they wound their shells in a different way.
The fossil record includes cephalopods with unwound, wound, and even partially wound shells,
and the diversity of fully wound ammonite fossils is good evidence of the successful body plan.
At one time these cephalopod predators ruled the ancient oceans, but now only Nautilus remains
to give a hint to what these ancient animals looked like.
Nautilus shell
The only living cephalopod with a shell is Nautilus and even in this animal it differs from that
of the other living Molluscs. In Nautilus the pearly shell consists of a series of closed chambers
separated from each other by septa and connected by the small openings created by the
siphuncle. This animal lives only in the very last chamber and the body does not extend back
into the shell. How does this differ, for example, from the spiral shell of a Gastropod?
Changes in the ionic composition of the fluid in the siphuncle move fluids in and out of the
chambers. This affects the levels of gas in the chambers and has the overall effect of altering the
buoyancy of Nautilus. In the other Cephalopods the shell has become reduced and internal. It
may be large such as the cuttle bone in cuttle fish or just a thin pencil or strengthening rod, or
pen, as in the squid Loligo.
Loligo
Squids in the genus Loligo are found around the world from warm shallow waters to the deep
abyssal depths of the oceans where they feed on a variety of small crustaceans and fish not fast
enough to escape from these predators.
External anatomy
To better understand how cephalopods have modified the mollusc body plan its important to
orient yourself by locating the head, foot and dorsal visceral mass. The squids body is divided
into two main regions. The first is the elongate, and somewhat conical visceral mass surrounded
by the mantle. Below this the head and foot that have fused. The last region includes the arms,
and tentacles surrounding the mouth. The mouth is the original anterior part of the body, the
funnel the posterior. Nautilus and the ancient cephalopods wound this visceral mass up, modern
cephalopods just tip over and swim with the ancestral dorsal surface pointing in the direction
that they travel. The result; the anterior side is functionally dorsal, the posterior is functionally
ventral, the ventral functionally anterior, and the dorsal surface functionally ventral and as was
Take a close look at the suckers of the arms under the microscope. Examine a sucker near the
base of an arm; those at the tip are the youngest and as a result much smaller. The cup of the
sucker is surrounded by a chitinous ring with a central muscular suction cup. Each sucker is
attached by stalk or pedicle. If you have a male, arm L5 will be modified for sperm transfer and
is referred to as the hectocotylus. Its suckers are small and located on the end of much longer
pedicles. The hectocotylus arm is used in mating and sperm, contained in a spermatophore, are
attached to these modified suckers before being passed to the female. In some species the tip of
the arm and its package of sperm breaks off. Its no great loss to the male; in cephalopods
damaged arms can be regenerated.
Lets turn our attention to the mouth region. Bend back the arms and tentacles attached to each
other by a muscular membrane surrounding the central mouth. Inside this is a second
membrane, the ruffle-edged peristomial membrane. In female squids the peristomial
membrane is modified into a horseshoe shaped seminal receptacle in the middle and below the
mouth. Sticking out from the mouth you should be able to see the beak-like chitinous teeth
used to tear apart captured prey. Well take a closer look at the buccal bulb later in the
dissection.
A pair of eyes on the head are remarkably similar to mammalian eyes; an excellent example of
convergent evolution. What is convergent evolution? Identify the cornea, iris, pupal and
lens. How does this eye differ from the mammalian eye? Just behind the eye, and near the
base of the arms is the aquiferous pore that stabilizes fluid pressure on the eye as the squid
dives. In front is a crest of tissue referred to as the olfactory crests and next to them the
olfactory grooves. They are chemosensory and positioned in the incurrent flow of the mantle
cavity.
The cone shaped funnel, siphon, is located on the ventral surface of the head and water is forced
out of the funnel for jet propelled locomotion.
Mantle cavity Like all soft bodied animals its going to be easier to see the various systems
if they are supported by water. Place you specimen in a dissection tray, pin the sides of the
mantle back and flood the specimen with water. By opening the mantle cavity on this side youll
see that the visceral mass seems to be floating inside the mantle cavity. Move it to either side
and youll see its attached to the mantle by a fine ligament on the underside of the body. Its
on that side where youll find the pen, the remnants of the shell. On the inside of the mantle
Fig. 16. Lateral view of the head and mantle in the squid. BIODIDAC
Split open the funnel and trace the way that water flows through the mantle cavity. Water enters
through the open collar. This is sealed against the body when the circular muscles of the mantle
contract, and force the water out through the funnel. The large funnel retractor muscles,
combined with the circular and longitudinal muscles in the funnel, direct the jet of water from
the funnel; controlling the direction the squid swims. Inside the funnel is a muscular valve that
prevents water from entering through the funnel. Underneath the funnel retractors are the
cephalic retractor muscles. The mantle is primarily circular muscle, but there is a smaller set
of longitudinal muscles that enlarge the mantle cavity. Why are there more circular muscles
compared to the longitudinal? The visceral mass is covered by a thin, transparent membrane;
the body wall.
The paired ctenidia are anchored to the wall of the mantle cavity and where they connect with
the body you will see the paired branchial hearts. The rectum and anus are located next to the
funnel and the opening is shared with the ink sac that lies along side this part of the digestive
tract. What is the function of ink? Underneath the rectum, and between the retractor muscles
is the large liver, a modified portion of the digestive gland.
Reproductive system Identify the sex of your specimen, and be sure to see both. Well
make some preliminary observations now. Some of the ducts of the system will be easier to see
once observations of the circulatory system are complete and the branchial hearts have been
removed.
If you have a reproductive female enlarged, paired nidamental glands lie on top of the viscera
near the center of the body, and the single ovary, filled with granular eggs is located at that
posterior tip of the body. The single oviduct runs along the left side of the body and opens into
the mantle cavity through the oviducal funnels. In the area where the oviduct passes under the
gills it enlarges forming an oviducal gland. Gently remove the nidamental gland to reveal the
oviduct and the opening of the oviduct underneath. Removing the nidamental gland will also
expose the accessory nidamental glands underneath. The oviducal gland adds the shell to the
eggs and the nidamental glands add the gelatinous capsule that covers the egg case.
In the male the single testis appears as a tubular structure just off center and near the posterior
end of the squid. The testis lies inside a thin membranous sac and the twisted sperm duct drains
the capsule. There is no direct connection between the two and sperm pass into the space of the
capsule and then into the sperm duct. A larger convoluted, spermatophoric gland may be
visible now, or after the branchial hearts have been removed. The spermatophoric gland
Open the pericardial cavity surrounding the systemic heart that pumps oxygenated blood from
the ctenidia through the anterior aorta to the front of the body. The posterior aorta supplies
the posterior part of the body and divides into three mantle arteries that disappear into the wall
of the mantle. Smaller branches of the anterior aorta may be visible supplying the rectum and
reproductive structures.
Digestive system The digestive system is complex and its worth remembering the general
mollusc digestive plan. The simplest description is a long tubular gut with a blind ended sac, the
gastric or digestive gland. The same applies with the squid, the difference being the digestive
Carefully remove the heart and metanephridia without destroying the underlying digestive
organs. Underneath the heart is the U shaped pancreas with a granular appearance. The
pancreas is part of the duct that leads to the liver. The liver occupies most of the anterior viscera
and embedded on its side are salivary glands. Loosen and remove the connective tissue
surrounding the liver and gently lift it up to reveal the esophagus and anterior aorta that pass
through the region. An equally large structure, this time forming most of the posterior end of the
viscera is the caecum, or stomach pouch. This thin walled structure fills most of the posterior
part of the mantle cavity and is filled with recently ingested food and secretions of the liver.
Other obvious features are the anus and the intestine leading to it.
Free the funnel from underlying tissue by cutting the two small siphon protractor muscles
between head and funnel and lift the siphon out of the way. Cut through the head between the
two arms immediately underneath the funnel to expose the pharynx modified into the buccal
bulb. The large, tough, interlocking jaws are easy to identify. Pry them apart and inside you will
find that the ancestral radula is still present. Under the radua is the ligula and the two structures
are referred to as the odontophore.
The esophagus leads out of the bulb, extends through the liver, and connects with the stomach.
The stomach has a muscular, thick wall and connects to the caecum, or stomach pouch and its
own diverticulum. The pouch in turn is connected to the liver through the pancreas. The
intestine is connected to the stomach near the entrance of the esophagus.
Nervous system The generalized mollusc body plan has a variety of ganglia positioned in
different parts of the body. In the squid these have all become fused and encased in a
cartilaginous brain case, or skull. The only exception to this are the stellate ganglia on the
inside of the mantle at the tips of the gills. To see the brain cut lengthwise through the head. It
will be hard to distinguish the differences but the supraesophagial ganglion is a single paired
ganglia and the mass of nerve tissue underneath the esophagus, the subesophagial ganglia, is
a fusion of the pedal and visceral ganglia. Circumesophagial connectives connect ganglia
above and below the esophagus. The large eyes are connected to the central nervous system
through the optic nerves attached to the ganglia underneath the esophagus.