CHAPTER 12: MOLLUSCA

Mollusca is an enormous taxon 2nd only to Arthropoda in number of living species Includes many familiar animals such as snails, clams, oysters, mussels, slugs, octopods and squids in seven lining classes Mollusca includes the taxa: Aplacophora consists of small, wormlike molluscs w/ numerous calcareous spicules rather than a shell. Polyplacophorans or chitons have eight shell plates Monoplacophorans have a low, conical, limpetlike shell and live in deep water. (The limpet shape appears often in molluscan evolution and includes an uncoiled, caplike, low-profile shell and a large creeping foot) Gastropods are the snails and slugs and have a one-piece, often coiled shell and are found in the sea, in fresh water, and on land Cephalopods include the squids and octopods and mostly have reduced, absent, or internalized shells. The bivalves are the clams and their relatives w/ a shell divided into 2 pieces Scaphopods are the tusk molluscs w/ a one-piece tubular shell.

GENERALIZED MOLLUSC A marine benthic animal, bilaterally symmetric, dorsoventrally depressed, and ovoid in outline The body is divided into: Head anterior, small, poorly defined Visceral Mass large dorsal Foot broad, flat, ventral Shell dorsal shieldlike; can be clamped against the substratum to protect the soft parts from predators. This animal is adapted for life on hard, rocky substratum, where it uses its rasplike radula to graze on the biofilm of microscopic algae and other small sessile organisms Attaches to the substratum by the muscular foot, w/c it uses to move slowly about in search for foot.

MANTLE Dorsally, the body wall over the visceral mass is elaborated to form the mantle or pallium w/c is characteristics of all molluscs Mantle epidermis secretes protein, calcium salts, and mucus and is also sensory.

SHELL Simple, low, conical cap covering the dorsum of the generalized mollusc Originated in the ancestral mollusk as a thick dorsal cuticle of chitin and protein, to w/c calcium salts were later added. The eumolluscan shell consists of 3 basic layers: one organic and 2 calcareous. Periostracum outermost layer, composed of the protein conchiolin. Ostracum calcareous & immediately under the periostracum. Hypostracum inside the ostracum The ostracum & hypostracum are composed of calcium carbonate deposited over an organic matrix In the gastropods, cephalopods and bivalves the hypostracum may be nacreous (pearly) MANTLE CAVITY Mantle skirt dorsal peripheral outfold of the mantle Mantle cavity a pocket that is partially enclosed by the mantle skirt; continuous w/ the surrounding seawater. - Central & characteristic feature of molluscan body and one on w/c many important processes depend. - Contains the gills, nephridiopores, anus, gonopores & sensory osphradia - Involved in feeding and when anterior may provide space for the retracted head and foot. The outer epidermis of the mantle skirt underlies the shell - Secretes the calcium carbonate required for increasing shell thickness Inner epidermis lines the mantle cavity, secretes mucus and is often ciliated. GILLS The generalized mollusks has several pairs of gills (ctenidia; ctene = comb), one pair of w/c is located in the posterior mantle cavity. Consists of a central axis, w/c is attached to the mantle along one edge. - Contains muscles, blood vessels, and nerves Gill filaments arise from opposite sides of the axis; two rows & leaflike. - Each filament has an upstream frontal margin on one side and a downstream abfrontal margin on the opposite side. Bipectinate gills two rows of filaments Monopectinate gills filaments on only one side of the axis Interfilamentary water spaces separates the filaments. The gills divide the mantle cavity into ventral inhalant (infrabranchial) and dorsal exhalant (suprabranchial) chambers The exposed gill surfaces are formed by the combined edges of the filaments are known as lamellae

 Frontal lamella faces the inhalant chamber; composed of combined frontal margins of the gill filaments Abfrontal lamella faces the exhalant chamber; composed of the combined abfrontal egdes of the filaments. The epidermis of the filaments bears 3 types of cilia: Lateral cilia generates the respiratory current through the mantle cavity. - The 2 remaining types of cilia move mucus and particles that might interfere w/ the respiratory flow of water bet. Filaments. Frontal cilia upstream frontal margin of the filament; generate a current moving away from the gill axis toward the tip of the filament. Abfrontal cilia participate in transporting pseudofeces. Secretory cells of the gill epithelium secrete mucus, w/c entangles particles from the respiratory current. The cilia move the particles along the edge of the filament to the exhalant side where they are released as pseudofeces into the exhalant water current, w/c sweeps them out of the mantle cavity. Pseudofeces are strings or pellets of mucus and sediment. Hypobranchial glands found on the mantle roof; 2 patches of mucus-secreting epithelium. - Trap sediment in the exhalant chamber. Blood (hemolymph) transports oxygen from gills to tissues. Afferent branchial vessel brings unoxygenated blood to the gill filaments Efferent branchial vessel removes oxygenated blood from the gill & transports it to the heart. Blood circulates through the filaments in an exhalant-to-inhalant direction Respiratory water current moves in an inhalant-to-exhalant direction OSPHRADIA A pair of sensory osphradia in the inhalant water current is widespread among mollusks. Receptor cells monitor the water entering the mantle cavity for chemical and perhaps for sediment. FOOT Occupies the ventral surface of the generalized body; large & muscular w/ broad, flat, ciliated creepsole. Attaches to hard substrata & moves the animal over their surfaces Abundantly supplied w/ mucous gland cells that produce a lubricating mucus to facilitate locomotion Pedal hemocoel blood sinus Pedal retractor muscles paired; dorsoventral; extends from the inner surface of the shell to the foot. - Contraction of these muscles pulls the shell toward the foot, or vice versa

 In generalized mollusk, the foot is usually attached to the substratum, & the action of the pedal retractors is to clamp the shell over the visceral mass, head, & foot to protect them. NUTRITION The generalized mollusk is a microphagous browser that uses its radula to scrape microscopic algae, other organisms & detritus from hard substrata The gut is adapted for separating & processing the mixture of fine organic particles (food) & fine mineral particles (nonfood) Many other living mollusks are macrophagous herbivores or carnivores that feed on large pieces of organic material. - Have simpler digestive systems in w/c sorting mineral & organic particles is not necessary The gut consists of: Foregut mouth, buccal cavity, pharynx Hindgut rectum & anus - Foregut & hindgut are lined by cuticle because of their ectodermal origin. Midgut esophagus, stomach, digestive ceca, intestine; endodermal & unlined. The mouth is anterior & opens into a cuticularized buccal cavity The radular apparatus lies in the floor of the buccal cavity & includes the radula itself Odontophore connective tissue; supports the radula; complex of muscles that operate the radula & odontophore - Odonto = tooth; phore = to carry - Chondroid ( cartilage-like) Radular sac ventral evagination of the floor of the buccal cavity; contains the radula Radula flexible, longitudinal ribbon of transverse rows of tiny chitinous teeth. - Unique to mollusks & is present in all taxa except the bivalves - Its teeth are composed of alpha chitin & tanned proteins that are sometimes hardened by iron or silicon compounds. Teeth -transverse rows & exhibit a variety of shapes adapted for food utilized by various taxa The odontophore & radula are equipped w/ numerous protractor & retractor muscles whose function is to move the odontophore & radula anteriorly & posteriorly During feeding, the odontophore protractor muscles pull the odontophore anteriorly, causing it to protrude slightly from the mouth in a position close to the substratum on w/c the animal wishes to feed. Odontophore retractor muscles the odontophore is returned to its resting position The radular teeth are secreted by epithelium at the posterior end of the sac. Salivary glands secrete mucuc into the buccal cavity Mucus mixes w/ food & mineral particles in the buccal cavity to form a mucous string that can be transported posteriorly by cilia Esophagus exits the posterior buccal cavity & extends to stomach Cilia lines the esophagus & stomach; moves the mucous string posteriorly into the stomach The stomach of the generalized mollusk is a pear-shaped sac in the anterior visceral mass.

- Esophagus enters at its anterior, expanded end. - Intestine exits at its narrow, tapered posterior end. - Extracellular digestion occurs. Digestive Ceca (Digestive glands, liver, digestive diverticula, or heptopancreas)paired & highly branched; connects by ducts with the anterior region of the stomach. - Absorption, phagocytosis, intracellular digestion & storage occur here. - Present in all eumolluscan taxa, but are absent in Aplacophora Digestive enzymes secreted by the gastrodermis of the digestive ceca & transported to the stomach. Gastric shield chitinous & lines the posterior end of the stomach. -protects the stomach gastrodermis from abrasion - Bears a sorting field of ciliated ridges & grooves Separates indigestible mineral particles from nutritious organic particles. Style sac found at the posterior end of the stomach; narrow & ciliated Intestinal groove long, deep, extends from the sorting field posteriorly through the style sac to the intestine Major & minor typhlosoles two ciliated ridges that extend from the sorting field to the opening of the intestine at the posterior end of the sac Protostyle or fecal rod stiff mass made from mucus and embedded fecal particles Crystalline style composed of digestive enzymes - present in some conchiferans, most notably the bivalves & some gastropod Rotation of the protostyle helps pull the mucous string through the esophagus into the stomach, where it is wound into the anterior end of the protostyle. Large organic particle tend to be sent to the digestive ceca for digestion, and smaller mineral particles become incorporated in the protostyle Contractions of circular muscles periodically pinch fecal pellets from the posterior end of the protostyle into the intestine. The stomach of macrophagous molluscs lacks the sorting equipment, style sac, gastric shield, & protostyle, and is usually a simple sac. Intestines materials entering it are wastes destined for defecation. - Modification and storage of fecal pellets. Anus located dorsally in the mantle cavity, in the exhalant current, where fecal pellets can be swept from the mantle cavity w/o fouling the gills.

COELOM Its original function as a skeleton has been assumed by the hemocoel & shell, but it continues to be an integral component of the excretory & reproductive systems It is the pericardial cavity & gonocoel, w/c contain the heart and gonad respectively.

INTERNAL TRANSPORT The hemal system of HAM consists of a heart, a dorsal aorta, blood vessels, blood, and a hemocoel. The blood contains amebocytes as well as the respiratory pigment hemocyanin. The hemal system w/c lacks a cellular lining, lies in the connective-tissue compartment in the spaces bet. The basal laminae of the major epithelia, namely the epidermis, mesothelium, & gastrodermis The heart consists of one or more pairs of atria (auricles), w/c receive oxygenated blood from the gills via efferent branchial vessels. The atria empty into the median unpaired, muscular ventricle, w/c is continuous w/ the aorta The aorta branches into smaller arteries that deliver blood to the several hemocoelic sinuses, or capillary beds. Blood is delivered to the gills by the afferent branchial vessel. - This system is sometimes characterized as open, meaning that arterial outflow & venous return are connected by large sinuses In contrast, a closed hemal system, such as that of annelids, has small capillaries connecting arteries & veins. EXCRETION It is accomplished by the heart-kidney complex, w/c is derived from & functions like hemalcoelom-metanephridial system of other coelomate animals. The kidney is a large sac w/ thick secretory & absorptive walls surrounded by blood in the visceral sinus of the hemocoel, w/ w/c it exchanges materials. The connection w/ the pericardial cavity is the renopericardial canal, whose opening from the pericardial cavity is the nephrostome Podocytes in the walls of the atria make possible the formation of an ultrafiltrate (primary urine) in the pericardial cavity. The final modified urine leaves the kidney via the nephridiopore & enters the exhalant current of the mantle cavity. NERVOUS SYSTEM & SENSE ORGANS A ganglion or neuromere consists of a cortex of cell bodies & medulla of axons known as the neuropil - Is enclosed in a connective-tissue capsule - Are connected to sensory receptors by afferent or sensory neurons and to muscles & endocrine organs by efferent or motor neurons The cerebral ganglia(brain) receive sensory nerves from the eyes, tentacles, and statocysts The buccal ganglia are located in the buccal mass & innervate the muscles of radula & odontophore. Motor nerves from the pedal ganglia innervate foot muscles

The pleural ganglia innervate the mantle Commisure connects any given pair of ganglia to each other; transverse Connectives connects the ganglia of different pairs; longitudinal The 2 cerebral ganglia are connected by the cerebral commisure whereas the cerebral & pedal ganglia are linked by the cerebropedal connective The pleural & pedal ganglia are joined by a pair of connectives Another pair of connectives, the visceral nerve cords, exit the pleural ganglia & extend posteriorly to the visceral ganglia, w/c innervate the organs of the visceral mass A pair of esophageal( intestinal or parietal) ganglia occur on the visceral cords - Supply the gills, osphradium & mantle The cerebral & pedal ganglia along w/ their connectives & commisures form the circumesophageal nerve ring A pair of pedal nerve cords extends from the pedal ganglia to the foot & the 2 visceral nerve cords run from the nerve ring to the visceral ganglion Tetraneuous a system consists of 4 nerve cords The typical molluscan sense organs include chemomechanoreceptive cephalic tentacles on the head, a pair of eyes on the head, a pair of statocysts in the foot, & a pair of osphradia in the inhalant chamber of the mantle cavity

REPRODUCTION The generalized mollusk is gonochoric w/ external fertilization in the sea Gametes are shed by the gonads into the coelom - They enter the nephrostomes of the nephridia, w/c serves as gonoducts, & exit via the nephridiopore into the exhalant chamber of the mantle cavity Fertilization is internal in many molluscs, & some are hermaphroditic DEVELOPMENT Spiral, holoblastic cleavage is a feature of the generalized mollusk The developing blastula bears the characteristic molluscan cross of cells at its animal pole Gastrulation is by invagination in species w/ small, microlecithal eggs and by epiboly in species w/ large, yolky eggs The blastopore becomes the mouth Mesoderm arises from the 4d mesentoblast cell, w/c divides to produce right & left mesodermal bands & these cavitate to produce the coelom by schizocoely The gastrula develops into a trochophore larve, w/c in primitive molluscs is the hatching stage In most species, the trocophore is suppressed & a later stage, often a veliger hatches Trochophore larva The trochophore is the characteristic larva of molluscs, annelids, sipunculans, echiurans, & many other protostomes

 Prototroch conspicuous preoral girdle of cilia, that encircles the body near its middle, but anterior to the mouth In many molluscs, polychaetes and sipunculans a second ciliary girdle that is post-oral, the metatroch, develops just posterior to the mouth. In some taxa a third ring, the telotroch, is found at the posterior end An apical tuft of cilia is located at the anterior end The adult mesodermal structures develop from a pair of ventrolateral mesodermal bands The rosette cells of the molluscan cross differentiate into an anterior ectodermal apical plate, from w/c arises the apical tuft The cerebral ganglia develop from the apical plate A fully developed trochophore consists of 3 regions: Pretrochal region consists of the apical plate, prototrich, & the area surrounding the mouth Pygidium the region consisting of the telotroch & the area around the anus Growth zone lies between the mouth & the telotroch Planktotrophic larvae usually planktonic & feed on tiny suspended particles, living or dead; have long planktonic lives Lecithotropic larvae depend on yolk & usually have a short planktonic life The prototroch is the swimming organ & in planktotrophic species, is also responsible for collecting food particles A food groove lies between the prototroch, w/c is preoral and the metatroch w/c is post-oral Veliger larva a more advanced stage that follows the trochophore larva; found in gastropods, bivalves & scaphopods. The hatching stage may be the trochophore or veliger if the trochophore is suppressed in the egg. The veliger has a foot, shell, & 2 lateral ciliated lobes together called the velum, w/ w/c it swims & feeds Some molluscs such as the cephalopods & many gastropods, have macrolecithal eggs & direct development , w/o a larva

APLACOPHORAC
Consists of 2 taxa, Neomeniomorpha & Chaetodermomorpha, of about 300 species of small, strange, wormlike marine molluscs Presence of calcareous spicules instead of a shell Absence or reduction of the foot Cylindrical & wormlike shape of the body The mantle cavity is posterior A radula is present in most species Exhibit many characteristics of the generalized molluscs, but in some respects are specialized

FORM Mantle dorsal body wall; extends laterally to cover the entire animal except for the foot, if present The mantle epidermis secretes a glycoprotein cuticle that contains embedded calcareous (aragonite) spicules of various shapes A shell is absent The body cavity or hemocoel is adjacent to the innermost (longitudinal) muscle layer The foot, when present, is a longitudinal fold filled w/ the pedal hemocoel - Its epithelium is ciliated - Locomotion is ciliary The foot lies in the pedal groove w/ is lined by cuticle The mantle cavity is posterior & contains the anus & the openings of the gonoducts (coelomoducts) In chaetodermomorphs, it contains a [pair of bipectinate gills In neomeniomorphs, the mantle epithelium is folded to form 2ndary gills (respiratory papillae) that are not true gills The gut consists of the mouth, cuticularized buccal cavity, esophagus, midgut, intestine, rectum, anus The buccal cavity contains the radula The anus is in the mantle cavity The hemal system includes a posterior heart similar to that described for the HAM The ventricle forms as an invagination of the dorsal pericardium & usually remains attached to it by a membrane The hemocoel is divided into a dorsal perivisceral hemocoel & a ventral pedal hemocoel by a longitudinal, muscular, horizontal septum The pedal hemocoel is present in all Aplacophora The nervous system is ladderlike - A pair of cerebral ganglia dorsal to the anterior gut extend the length of the body and are connected w/ each other by commisures The cords of the right & left sodes commect via a suprarectal commisure that arches over the posterior guts. A posterior chemoreceptive dorsoterminal sense organ is a characteristic feature of the aplacophoran sensory system In neomeniomorphs the mucus-secreting ciliated pedal pit has a sensory function and there is also a dorsofrontal sensory pit above the mouth The gonads are paired & release their gametes directly into the pericardial cavity (coelom) via gonopericardial ducts The pericardial cavity is connected w/ the mantle cavity by a pair of elaborate coelomoducts(gonoducts) Most aplacophorans are hermaphroditic, but chaetodermomorphs are gonochoric

 Eggs may be retained , in w/c case development is direct or spawned, whereupon a lecithotropic larva latches from the egg DIVERSITY OF APLACOPHORA NeomeniomorphasC (Solenogastres) - larger taxon - reduced foot - Gills lost - Carnivores feed on the cnidarians on w/c they live - Hermaphroditic - Copulation w/ internal fertilization - 25 families - Eleutheromenia, Lyratoherpia, Neomenia ChaetodermomorphasC (Caudofoveata) - Small taxon of 3 families - Foot completely lost - Have a pair of gills - Oral shield partly or completely encircles the mouth - Selective deposit feeders or carnivore burrowing in soft substrates - Gut w/ a digestive cecum - Gonochoric - External fertilization - Chaetoderma, Epimenia, Falcidens, Limifossor, Scutopus "To be beautiful means to be yourself. You don't need to be accepted by others. You need to accept yourself." - Thich Nhat Hanh

POLYPLACOPHORAc
Chitons or coat-of-mail mollusks Found only in the sea Chitons are highly adapted for a life adhering to rocks & shells