Modifications of foot in Mollusca

Meaning and Origin of Foot:

Phylum Mollusca is characterised by the pronounced development of musculature known as the
foot. It is the locomotory organ in Molluscs. This organ is quite uncommon and strange to
others.
Origin:
In Mollusca, the foot originates at first as the ventral, later the mesodermal cells incorporate to
give it a definite shape.
Innervation:
The foot and its associated structures are innervated by the pedal ganglia and pedal nerve cord.

Depending on different modes of locomotion and living in varying environment, the foot in
Molluscs varies greatly in shape and form (Fig. 16.63). Variation of foot is primarily due to
various physiological activities like creeping or crawling, burrowing, leaping, looping, swimming,
reproduction, etc.

Structure of Foot:

In most typical gastropods, particularly in prosobranchs the foot is usually differentiated into:

(1) A small anterior propodium,

(2) A large middle mesopodium and
(3) A posterior metapodium (e.g., Natica, Fig. 16.65D).
Primitive foot:
The primitive and simplest form of foot in Mollusca is considered to be a broad ventral flat sole
having the above mentioned three regions (e.g., Polyplacophora, Gastropoda, Bivalvia
[Protobranchia]). Beside these they adapt certain structures as parapodia and epipodia.

Parapodia:
Lobe-like lateral extentions given out from below upwards from the edge of the ventral sole
(e.g., Aplysia), and act as fins.

Epipodia:
Projecting paired ridges or folds developing from the sides or base of the foot along its entire
length. These may be beset with papillae or tentacles (e.g., Fisurella).

These parts are variously modified in different species of gastropods.

Modification of Foot:

Foot-as the Creeping or Crawling Organ:

Class Aplacophora:
True molluscan foot is absent in Aplacophorans but some structure may be regarded to be the
starting point. In Neomenia (Subclass Neomeniomorpha), a mid-ventral groove from mouth to
anus with non-muscular ciliated ridge is believed to be homologous with the foot of other
molluscs and serves as locomotory organ (Fig. 16.65U). The foot helps to glide or to creep over
the substratum with a mucous trail.
 Class Polyplacophora:
In Polyplacophora, the foot of Chiton is broad, muscular and flattened that extends the entire
ventral surface of the body (Fig. 16.65A). In Chitonellus and Crypsoplax the foot is narrow. In
Ischnochiton the anterior portion of the foot is elongated. The foot helps to creep or glide on the
rocky substratum by the waves of muscular activity which is lubricated by mucus glands.

Class Monoplacophora:
In Neopilina and Vema the foot is centrally placed, broad, flattened and almost circular in
outline. The foot helps in creeping by muscular movement.

Class Gastropoda:
In most gastropods the foot is an elongated, flat creeping sole that contains numerous mucus-
producing gland cells. In the members of the subclass pulmonata the foot is undivided with a
very large flat lobe containing a large pedal gland. In these cases the foot is used for creeping
on a mucous trail. The terrestrial pulmonates retain the primitive type of foot.

In prosobranchs, Patella has a well-developed ventral foot with a flat creeping sole which is
adapted for clinging or moving over the rocks (Fig. 16.65C). The creeping foot may be
contractile as in Triton. In some cases foot shows partial regional modification.

Bivalvia:
The foot in Nucula and Area are considered as primitive type, which possess a flat, ventral
surface of sole on which the animal creeps.

Foot—as the burrowing organ:

In some molluscs foot becomes greatly deviated to act as burrowing organ.

Class Scaphopoda:
In Dentalium the foot is conical, trilobed and protrusible (Fig. 16.65H). In Siphonodentalium, the
foot terminates in a retractile disc with papillated margins. The foot of the Scaphopoda is
adapted to burrowing habit in sand and the co foot is buried into sand and the foot lateral,
epipodial lobes assist in burrowing.

Class Gastropoda:
In Terebra, the extremity of foot with flow of blood is extended and acts as anchor. In Natica
(Fig. 16.65D), Polinices, Sigaretus (Fig. 16.65P), the propodium is demarcated from the rest of
the body by deep transverse grooves as a semicircular flap and the metapodium is provided
with lateral parapodium. The animals have adapted for burrowing on soft bottom habitat. The
propodium acts like a plough and anchor, and a dorsal flap-like fold of the foot protective shield.
In Harpa the propodium is separated by a constriction.

Class Bivalvia:
In Anodonta and Unio, the foot is triangular and plough like. The foot can perform as effective
burrowing organ in addition to acting as a creeping organ. In Solemya, Yoldia the foot has a
flattened sole and two sides of the sole can be folded to form a blade-like edge which can
penetrate into the mud or sand and act as soft bottom burrowers.
In most bivalves, the foot is laterally compressed and blade-like, and the anterior part of the foot
acts as a burrowing organ in the soft substratum where they live.

Foot—as the leaping organ:

This type of modification is very characteristic in certain forms of Molluscs. In Cardium (Fig.
16.65L), the foot is bent upon itself as a leaping organ. In Trigonia it is compressed antero-
posteriorly as an elongated keel. In Poromya, the foot is curved with well- formed protractor and
retractor muscles. In Mytilus, the cylindrical foot acts as spring- tail. A median triangular
outgrowth in Birabia acts as leaping organ.

Foot—as a looping organ:

In Pedipus, the propodium of the foot is sharply marked off from the rest by a groove and helps
in looping movement.

Foot in sessile forms:

A number of sessile bivalves such as sea mussels (e.g., Mytilus, Perna, Modiolus) or oysters
(e.g., Ostrea, Crassostrea) are attached to the hard substrates (e.g., rock, corals, shells, wood,
sea walls, jetties and pilings, etc.) either by byssus or by the one valve to the substrates.
Byssus is a bunch of fine silky threads by which the bivalves are attached to the objects.

The byssal threads are proteinaceous secretions from byssus gland of the foot. The
members of mytilids use the byssus for attachment to other objects. The foot is elongated
and not used for the lead of sessile life.

Pen shells (e.g., Pinna, Atrina) are attached to the underlying solid substrate by byssus and
partly remain buried in sandy sediment. Oysters are firmly cemented to the rock and other
substrates. They lack foot and byssal threads.

Crepidula (class Gastropoda) is a sedentary animal which possesses the reduced foot, and the
ventral sole acts as an effective sucker for attachment. Vermicularia (Gastropoda) a worm-shell
snail is a sessile gastropoda and its foot is reduced. The shell is attached to solid substrates
such as rocks and other shells, or entangled in sponges.

Foot—as the swimming organ:

In many Molluscs the foot becomes variously modified to swim in water. The development of
parapodia is observed in many forms. Carinaria (Fig. 16.65N), Atlanta and Pterotrachea swims
by a flat ventral fin which bears a small sucker representing the original foot. The parapodia is
fan-like in Aplysia (Fig. 16.65J) and used for pelegic existence.

It may have wing-like processes as in Pteropoda (Fig. 16.65Q). In Oxygurus the parapodia are
hollow and possess fin-like outgrowth. In Atlanta, the metapodium is produced into laterally
flattened swimming lobe. In Clione (Fig. 16.65E) the parapodia are well-developed and are
placed on the lateral sides of the anterior end.

The most notable transformation of the foot as swimming organ is observed in cephalopods.
The foot is partly modified into muscular arms and tentacles around head for food capturing,
adhesion and locomotion, and partly into a ventral siphon for jet propulsion.

The cuttle fish and squids move backwards and not forwards on the basis of jet-propelled
mechanism. The siphon (funnel) squirts water forward, pressure of the action squirts the animal
simultaneously backwards.
The siphon (also called funnel or infundibulum) is a ventral, muscular, sub-conical tube helps in
swimming via jet propulsion by expelling water from the mantle cavity.

The expulsion of water is achieved by muscular contraction of the siphon or funnel. In Nautilus
(Subclass Nautiloidea) the funnel consists of two separate lateral muscular lobes that fold
together to form a tube-like structure which serves for jet propulsion. The funnel of Nautilus is
not a complete tube.

In subclass Coleoidea (e.g., cuttle fish, squids) the siphon is a complete one and is made up of
two muscular lobes which are completely fused. In both groups the funnel is used for swimming.

Foot in Boring molluscs:

Bivalves that bore into mud (Ark family), wood (Barnea, Bankia, Martensia, Teredo), limestone
(Lithophaga, Saxicola), sand stone (Pholas), rock (Petricola), and also coral and coralline rock,
possess elongated and cylindrical shells. The foot of the boring species is not well developed.

Mechanism of boring:
The boring is done by the anterior end of the bivalve shells. The anterior end of the valves is
usually serrated and has abrading surfaces. The drilling is a mechanical process and is done by
the anterior ends of the valves which are rotated back and forth by which the species can bore
into the hard substrates. The boring species can attach to the substrate with the help of foot
which has developed a sucker-like ventral surface.

Foot—as an organ helping reproduction:

One or more arms in some male cephalopods during breeding season are modified for the
transference of spermatophores to the female, called hectocotylus. In Sepia the fourth left arm
of the male becomes hectocotylus in which some basal rows of suckers are suppressed. In
Loligo and Octopus the third right arm is modified to form hectocotylus. The modifications may
involve suckers, calimus and ligula, etc.

Accessory glands associated with the foot:

The foot in Molluscs is also a highly glandular structure. Some glands become intimately
associated with the foot to help in locomotion. The secretion of the glands lubricates the
passage during movement. In Gastropods the pedal glands and the unpaired sole gland are the
typical instances.

In some Bivalves, in adult or in larval stages the byssus apparatus helps in the process of
adhesion. The Organ of Valenciennes in some Cephalopod females, Van der Hoeven in
Nautilus males are the other notable accessory organs associated with the foot.

Foot in Parasitic forms:

The foot of the parasitic forms of molluscs like Entoconcha, Entocolax, Enteroxemes, Thyone
(live within sea cucumbers echinoderms), Eulima, Stylifer (endoparasites of echinoderms), is
either reduced to small appendages or completely absent.