Unit 7:  Phylum Annelida and Protostomes (Annelids)

1.  Define and explain these terms:  eucoelomate, prostomia, deuterostomia, schizocoelous, enterocoelous and trochophore.
2.  Explain the etymology of Annelida and compare the general complexity of the body sustems of the annelids with previously studied phyla.
3.  Explain metamerism.  Highlight the two different forms of segmentation displayed by the remaining phyla to be studied.
4.  State the three classes within the phylum Annelida and give examples of each.
5.  Discuss the ecological impact of members of the phylum, citing the earthworm specifically and its role in maintaining good soils.
6.  List several general phylum characteristics concentrating on new developments.
7.  Identify members of the class Hirudinea and discuss differences between members of this class and the other two of the phylum.
8.  Cite differences between polychaete worms and others in the phylum.
9.  Discribe the reproductive patterns of the South Pacific Palolo worm.
10.  Discribe the digestive tract of the Oligocheata, tracing the passage of materials from mouth to anus.
11.  List and briefly describe the organs and systems which are introduced for the first time with the annelids, or which are more highly developed in the annelids than in previously studied groups.
12.  Describe how reproduction occurs in the earthworm, a representative of the class Oligochaeta.

Resources:  Text:  Animal Diversity, Chapter 11
Internet Resources:
    http://gnv.fdt.net/~windle
    http://dmoz.org/Science/Biology/Zoology/
    http://www.il-st-acad-sci.org/health/wormlink.html
    http://www.mhhe.com/hickmanad4e

Metamerism - segmentation, linear series of essentially similar units called segments,
somites or metameres

    -metamerism evolved twice in Animal Kingdom

            1. annelid/arthropod line: locomotion ---> burrowing

            2. chordate line: locomotion ---> swimming

           homonomous segmentation - metameres structurally and functionally similar,
            each  acts  as semi-independent unit, however, cannot function independently
            (ex. Annelida)

           heteronomous segmentation- metameres in different regions of body
            dissimilar, not always  obvious (ex. Arthropods, chordates)

            *Metamerism allowed development of greater complexity of structure and function;

                -made possible independent and seperate movements by seperate segments,
                    the need for fine control led to evolution of more sophisticated nervous system

                -repetition of body parts gave organism built in redundancy, safety factor, if one
                    segment fails, others can still function, thus injury to one part not necessarily fatal.

                -evolutionary significance evident in great success of arthropods and vertebrates.

Phylum Annelida - segmented worms(bristle worms) ~9000 species
    -inhabit virtually all marine environments, have successfully invaded freshwater and
        moist terrestrial

    CLASS OLIGOCHAETA- most familiar, earthworms, nearly all F.W. and Terr.

    CLASS HIRUDINEA- carnivorous, bloodsucking, leeches

    CLASS POLYCHAETA- largest class ~5000 species, marine, Nereis

Ecology/Economics
    -predators, ciliary filter feeders, saprophytes
    -burrow, build tubes, free swimming
    -Earthworms and leeches directly influence human welfare

        Charles Darwin- one m2 of ground may contain 20 earthworms, castings
                scattered over soil  surface.
            -can equate to 4.5 Kg (~10lb) of soil carried to surface/year, over 20 years
                a layer 8cm thick would be transformed.

            *in addition to soil turnover:
                1. makes soil porous, better penetration of water and air
                2. accelerates bacterial decomposition of organic matter.

        Aristotle- "intestines of soil"

       Leech used in medicine- bad blood/blood letting

            -red stripe on barbers pole indicated that barber was also blood letter (white
                stripe   signified bandage)

            -salivary gland produces HIRUDIN, anticoaggulant

            *making medical comeback, microsurgury, reattachment, leeches relieve congestion

General Phylum Characteristics:
    -segmented body, metamerism, bilateral symmetry

    -antagonizing muscle groups

    -setae, chitinous bristles (except in leeches), polychaetes have setea on fleshy
        parapodia,  setae used to anchor worm and aid in locomotion

    -coelom divided by septa (except leeches)

    -closed circulatory system; hemoglobin present in some amoebocytes present
        in blood  (similar to WBC's)

    -digestive tract complete but not segmented

    -Respiratory system absent or reduced to gills, gas exchange by diffusion through skin
        (moisture necessary)

    -excretory by nephridia; one pair per segment (why?)

    -nervous system, ventral nerve cord w/ lateral nerves in each segment (due to
        metamerism)  dorsal cerebral ganglion (paired brain)

BODY PLAN
    *head (prostomium); segmented body, terminal anus
            -growth at posterior end just in front of last segment

    *muscle system for locomotion/burrowing although coelomic fluid does serve as
        hydrostatic   skeleton (presence of septa allows greater efficiency because muscle
        contractions are   not whole body) widening and elongation can occur in localized areas

    *as segments widened (by longitudinal contraction) and body anchored to burrow wall,
        setae   aid in anchoring, anterior stretches (circular muscles contract); locomotion

    *coelom- schizocoel w/ peritoneum that forms mesentaries; covers all organs

    *segments seperated by septa

CLASS HIRUDINEA  ~520 SPECIES; leeches

    -mainly freshwater but few marine and terrestrial
    -fixed # of segments, usually 34 (similar to eutely)
    -posterior and anterior suckers

    -no parapodia; no setae (main reason for grouping)
    -no internal segmentation (coelom filled w/fluid and connective tissue
    -carnivores as well as external parasites.
    -feeding:
        mouth opens for muscular proboscis or muscular pharynx w/ 3 jaws and teeth

        blood suckers, take large quantities

        digest slowly mainly w/aid of gut bacteria

CLASS POLYCHAETA ~5000 species, largest class  Nereis and feather dusters

    -exclusively marine, inhabit every possible niche, significant part of marine food chain

    -polychaetes have well developed head
           Prostomium (head)- often has eyes, statocysts.  eyes range from eyespot to well
            developed organs.

           Peristomium- surrounds mouth, in Nereis bears chitinous jaws, in tube worms
            bears   fans for filter feeding (tentacular crown)

                -food attracted to arms (radioles), cilia move food to mouth

    -no clitella, no permanent sex organs, no permanent ducts for sex cells, dioecious,
        fertilization is external

                Palolo worm of South Pacific:  Posterior epitokes (gamete sacs)
                                                                     Anterior atoke (non-reproductive)

                    -lives in burrows among coral reefs
                    -during reproductive cycle epitoke becomes swollen w/gametes
                    -swarming occurs during last quarter of October/November moon
                    -epitokes break off and swim to surface
                    -at surface epitoke segments burst releasing gametes
                    -atoke regenerates posterior epitoke

CLASS OLIGOCHAETA (few setae) ~3000 species, most F.W. and terrestrial ***earthworm

    -no parapodia but do possess several varieties of setae

    -peristaltic movement, waves of contraction along length of body, setae anchor segments.

    DIGESTION- unsegmented tract
        food into mouth (buccal cavity) by contractions of muscular  pharynx, then to   esophagus

        -3pr calciferous glands by esophagus (excrete excess Ca+ andCO3- ions from blood)

        -in pharynx, saliva begins digestion

        -from esophagus to crop, storage organ

        -crop to gizzard, grinds food into minute particles by geing squeezed and rolled about.
            Injested grains of sand aid process

        -food then to intestine:
                Enzymes:  amylase (carbos), cellulase; pepsin and trypsin (protein); lipase (fats)

       TYPHLOSOLE- folded area of intestine, why??????

       CHLOROGOGUE TISSUE- surrounds tract and dorsal blood vessel, intermediary
            metabolism, synthesize and store glycogen and fats, synthesize urea.

CIRCULATORY SYSTEM (coelomic fluid serves as accessory system)

        -nutrients absorbed through intestine into coelom as well as blood vessels

        -5pr. of aortic arches ("hearts") at anterior end, dorsal blood vessel receives blood
            from lateral vessels, dorsal blood moves from post. to ant.; ventral takes blood
            form   ant. to post.

        -blood contains hemoglobin and amoeboid cells

EXCRETION- nephridia, pr. in each segment; nephridium occupies two successive
        segments; nephridium in segment 5 drains segment 4;

       Nephrostome funnels wastes from preceeding segment, cilia conduct liquids.

        -nitrogenous wastes eliminated through nephridiopores

NERVOUS- cerebral ganglia (bilobed "brain") located dorsan to pharynx in 3rd segment
        -connected to ventral nerve cord by connective nerve tissue around pharynx

        -ventral nerve cord extends length of body, lateral nerves branch into each segment.

       Brain + nerve cord = Central nervous system
        Lateral nerves = peripheral nervous system

REPRODUCTION
        -hermaphroditic organisms that practice cross fert.

        -during copulation worms joined by mucus secretion from clitellum and by specialized
            setae

        -sperm discharged and travels via seminal grooves to receptical of other worm

        -after cop. (sometimes days later) cocoon formed from clitellum, as cocoon moved
            along body toward head, eggs, sperm and albumin are deposited in cocoon.
            Fertilization occurs in cocoon.

        -clitellum develops when worm sexually mature

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