Unit 6:  Eucoelomate Animals:  Phylum Mollusca  (Mollusks)

1.  Diagram and discuss the schizocoel and enterocoel hypotheses of coelom development.
2.  Define the terms protostome and deuterostome.  Discuss the differences between the two forms of development.
3.  Summarize the evidence which places the mollusks where they are in relationship to the annelids on the phylogenetic tree.
4.  Compare the size and diversity of the Phylum Mollusca with other phyla.
5.  List the seven classes of mollusks, give any common names for each class, and representatives of each class.
6.  List and briefly describe the molluscan characteristics of shell, mantle, radula and foot in each of the seven classes.
7.  Diagram and describe the three layers of a typical molluscan shell.
8.  Describe the formation of a pearl.
9.  Use the variation within the mollusks to illustrate what is meant by adaptive radiation, and relate the success of the mollusks in adaptive radiation to the new phylum characteristics and the conditions at the time the phylum evolved.
10.  Compare the mollusk circulatory system and oxygen-combining pigment with those of earlier phyla.
11.  Detail the benefits that mollusks provide man, particularly the economic benefits.  Contrast the economic benefits of this group with the economic impacts of the previous groups discussed in this class.
12.  Describe gastropod torsion and the conditions that result from this process.
13.  Define operculum and radula.
14.  Describe coiling in gastropods.  Identify the differences between conispiral and planospiral coiling.
15.  List characteristics of the three subclasses of gastropods, the Prosobranchia, the Pulmonata and the Opisthobranchia.
16.  Outline the life cycle of a typical freshwater clam.
17.  Briefly describe the anatomy and functioning of the respiratory system, digestive system, circulatory system, excretory system and reproductive system of the clam.
18.  Describe the foot in each class, and the general arrangement of the digestive tract.
19.  Name the unique organ in the mollusk mouth which is used for feeding.
20.  Detail the role of the crystalline style in the digestive system of a clam.  Trace the route of a food particle through the digestive system of a clam starting with entry into the clam and ending with exiting of waste.
21.  Briefly describe the method of movement, activities in general and the feeding method of members of each class.
22.  Summarize the ways in which cephalopods are more complex than the other kinds of mollusks.
23.  Compare the circulatory system of the cephalopods to that of the bivalves and higher organisms such as chordates.
24.   Describe the unique features of reproduction in the cephalopods.

Resources:  Text:  Animal Diversity, Chapter 10
Internet Resources:
    http://www.austmus.gov.au/
    http://www.mhhe.com/hickmanad4e

Eucoelomata

Coelom- mesentaries, mesodermal lining (peritoneum)

Advances/Problems so far.
    Advances                         Problems
multicellular                            internal transport
specialization                        respiration
diplo and triplo                      waste removal
circulatory system                 nutrient transfer
(Nemertea)
protonephridia
branching gut

    Eucoelomates:    1. peritoneum, ciliated
                                    2. body wall, visceral wall
                                    3. blood vessels in mesentaries
                                    4. excretory system and gonadoducts held in place

How coelomates are formed today.

    1.  Schizocoelomates- annelids, arthropods, mollusca
            -coelom appears as a space in mesoderm
 
 
 
 
 

    2.  Enterocoelomate- echinoderms, chordates
            -coelom appears as pockets of gut
 
 
 
 
 
 

Phylum Mollusca
        -second only to Arthropoda in named species (50,000 living; 35,000 extinct)

Classes
       Monoplacophora- one plate, Neopilina, thought to be extinct

        Polyplacophora- many plates (8), chitons  Tonicella

   ***Gastropoda- belly foot; snails, nudibranchs

   ***Bivalvia- (Pelecypoda) two doors; clams, oysters, mussel/Mytilus

   ***Cephalopoda- head foot; squid, octopus (Loligo)
 

GENERAL FEATURES OF MOLLUSKS:
Mantle-             -"skin" surrounding visceral mass
                           -outer surface secretes shell

                           -houses respiratory organs (gills in aquatic, lungs in terr.

                           -into cavity is dumped products from digestive, excretory,
                                reproductive systems
                           -water currents bring food, flush water (bivalves)
                           -adapted for swimming in cephalopods

Foot-                 -major characteristic used for classification
                           -adapted for locomotion/attachment

                           -usually ventral, broad, flat, muscular

Radula-             (absent in bivalves)
                            -rasping tongue
                            -teeth backwards on conveyor belt, scrape food into mouth/pharynx

Shell-                 -bivalve (clam, univalve (snail), 8 plates in chitons, rod (pen) in
                                squid; divided in Nautilus, absent in slugs and nudibranchs

        3 Layers
                  Periostracum- outer layer, protects against erosion and boring animals,
                        secreted by mantle only at outer edge.  UMBO oldest part of shell.

                 Prismatic layer- calcium carbonate, thick, secreted by mantle at outer edges

                 Nacreous layer- calcium carbonate in thin sheets, secreted continuously
                        by mantle surface, grows w/animal.  Responsible for pearl formation.

ALL BODY SYSTEMS PRESENT!!
    Respiratory- diffusion still occurs (through mantle) but specialized gills, lungs
            present   (derived from mantle)

    Circulatory- pumping heart, vessels and sinuses; open system in all but cephalopods;
            Hemocyanin- "blood"- oxygen combining pigment, contains
                    copper ---> bluish  when combined w/O2

    Digestive- complete and complex

    Excretory- 2 kidneys (nephridia) empty into mantle cavity

    Nervous- ganglia and connecting nerve cord; specialized eye in ceph.

    Reproductive- most dioecious, some gastropods monoecious

HUMAN IMPORTANCE (economic benefit rather than liability)
    -food source; pearls

    -damage to warves; snails/slugs damage greens; parasite host
 

Class Gastropoda
        most successful class ~35,000 living (named):  marine, freshwater,
                terrestrial (snails/slugs)

        -bilaterally symmetric, basically, However. . . .very early and abruptly in
                larval (veliger) stage TORSION occurs

            *twisting of internal organs at least 180o because of uneven development
                of right and left muscles that attach the head/foot complex to shell
            *outcomes of torsion:
                -head can be withdrawn into shell; OPERCULUM
                -right becomes left and visa versa

                -anus is moved anterior and dorsal
                -reduces visceral volume

                -foot large, ventral, (slugs- slime trail aids movement)
                -head well developed w/eyes, sensory tentacles and a RADULA

                -respiration by gills except for terrestrial, due to torsion usually only one
                        gill present
                    *pulmonate snails hava a lung, mantle cavity functions in this capacity by
                        trapping air in pneumostome.

                -because of torsion, usually only one auricle, one nephridia

                -both mono & dioecious, insemination often preceeded by courtship; sperm
                    exchanged using spermatophores , self fertilization is avoided.

        COILING (not torsion), winding of shell
            primitive- planospiral, coiling in one plane, not compact

                           conispiral , shell tapers, more compact but  unbalanced.  Shell axis
                                perpendicular to foot  axis.

            shell eventually shifted up and back w/ shell axis not perp (or parallel) to axis of foot.

            -weight pressed on right side effectively reducing organs on that side leading to
                bilateral asymmetry.

THREE SUBCLASSES:

    Prosobranchia                           Pulmonata                                    Opisthobranchia
    -largest group                               -land and freshwater                      -undergo detorsion
    -anus remains forward                -lack gills, mantle                           -gills and anus
                                                              cavity acts as lung                         return to posterior
                                                           -aquatic 1pr of                                 still only have one
                                                               tentacles                                         gill, one nephridia
                                                           -terrestrial 2pr of                            -nudibranch no shell
                                                                    tentacles                                -naked gill
                                                                                                                    -use nematocysts
                                                                                                                      from cnidarians for
                                                                                                                             defense

Class Bivalvia (Pelecypoda = hatchet foot)  `20,000 named

    -2 lateral shells (valves), hinged dorsally
    -foot typically bladelike, adapted for burrowing, operated by muscles working against
        hydrostatic pressure of blood within the hemocoel.

     (exceptions:  Mytilus is sessile, attaches to substrate, oysters secrete cement,
            scallops capable of swimming)

    -mantle modified posterior to incurrent & excurrent siphons, control  flow of water into shell
    -filter feeders:  mucus on ciliated gill, grooves, palps seperate particles
 
 
 
 
 
 
 
 
 
 

    -basis for classification:  hinge teeth, adductor muscles, gills

    -digestion in stomach, long style sac containing a crystaline style composed of
        solidified mucus and enzymes
            *style rotated, reels in food laden mucus, style projects into stomach where it is
                worn away liberating digestive enzymes.

            *stomach contains ciliated sorting area, large particles to intestine, smaller to
                digestive gland

    -dorsal heart pumps blood from gills (oxygenated) out to anterior and to kidney for
                waste removal.

    -ganglia, 3prs.
    -sense organs include eyes (scallops) w/ cornea, lens and retina
            *many bivalves have a gravity sensing statocyst embedded in foot

    -most bivalves dioecious, gametes shed externally through excurrent

            *freshwater fert. internal, larva develop into glochidium that becomes parasitic
                on gills of fish until it can resist being washed away
            *marine embryo have three free swimming stages; trochophore, veliger,
                spat ---> adult

Class Cephalopods (head foot)  squid, octopus, Nautilus
            foot becomes siphon (funnel) ~650 living species

            -highly mobile and most advanced invertebrate
            -giant squid up to 16m in length.  delicacies for sperm whale which usually
                bear scars ~25cm   in diameter inflicted by sucker that has a chitinous claw.

       SHELL
            -present in Nautiloids- series of gas chambers allow bouyancy septa divide
                chambers  (diff from gastro) animal inhibits newest chamber, all others
                connected by siphuncle
            -reduced to chitinous pen in squid, absent in octopus

       HEAD
            -mouth ringed with tentacles (90+ in Nautilus, 8 in octopus, 10 in squid)

            -mouth equipped w/ chitinous beak and radula, saliva more or less venomous

            -eyes large and closely resemble vertebrate eyes even to the extent of being
                able to adjust pupil and change focal length
            -movement by jet propulsion, siphon directs, octopus crawls

            -water movement multipurpose (waste expulsion, oxygenate, locomotion,
                gamete    expulsion)

            -brain highly developed w/millions of nerve cells, one lobe of brain dedicated entirely
                to bringing about color changes.

            CHROMATOPHORES are pigment sacs, expanding or contracting changes color,
                defensive or behavioral w/ courtship

    All cephalopods are predacious

       DIGESTION
            -Y shaped tract, large ceacum (cecum) occupying apex of Y, mouth and anus
                located    anterior

            -digestion extracellular in stomach; enzymes secreted by "pancreas" and "liver"

            -cecum folded and sorts out indigestible particles for elimination

       CIRCULATORY SYSTEM
            -closed system w/3 hearts

            -median systemic heart pumps blood through body through arteries into capillary
                beds,    vein delivers deoxygenated blood to pair of branchial hearts at base of
                each gill

            -hemocyanin respiratory pigment

       RESPIRATORY SYSTEM
            -gills not ciliated, therefore, cannot create currents

            -mantle muscle contraction draws water in and out

       REPRODUCTION
            -most dioecious
            -mating preceeded by courtship
            -sperm packaged into bundles ( spermatophores) and stored

            -during mating male transfers spermatophores w/ hectocotylized  arm to edge of
                females mantle or directly into genital duct

            -eggs fertilized as they leave oviduct and attach to fixed surface to develop
                (octopus   tends eggs)

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