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    Origin of Metazoa

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    Sambitsabyasachi Hota
    1. Most zoologists agree that multicellular animals (metazoans) evolved from colonial protozoans. The colonial theory proposes that metazoans descended from spherical colonies of flagellated cells held together by an extracellular matrix. 2. Choanoflagellates are the closest living relatives to metazoans. Their colonies resemble what the ancestral metazoan colony may have looked like, with flagellated cells embedded in an extracellular matrix. 3. Establishing polarity along an anterior-posterior axis and internalizing growth cells allowed for both locomotion and growth, avoiding conflicts between these functions. This set the stage for the evolution of cell specialization in metazoans.

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    Origin of Metazoa

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    Sambitsabyasachi Hota
    2K views12 pages
    1. Most zoologists agree that multicellular animals (metazoans) evolved from colonial protozoans. The colonial theory proposes that metazoans descended from spherical colonies of flagellated cells held together by an extracellular matrix. 2. Choanoflagellates are the closest living relatives to metazoans. Their colonies resemble what the ancestral metazoan colony may have looked like, with flagellated cells embedded in an extracellular matrix. 3. Establishing polarity along an anterior-posterior axis and internalizing growth cells allowed for both locomotion and growth, avoiding conflicts between these functions. This set the stage for the evolution of cell specialization in metazoans.

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    1. Most zoologists agree that multicellular animals (metazoans) evolved from colonial protozoans. The colonial theory proposes that metazoans descended from spherical colonies of flagellated cells held together by an extracellular matrix. 2. Choanoflagellates are the closest living relatives to metazoans. Their colonies resemble what the ancestral metazoan colony may have looked like, with flagellated cells embedded in an extracellular matrix. 3. Establishing polarity along an anterior-posterior axis and internalizing growth cells allowed for both locomotion and growth, avoiding conflicts between these functions. This set the stage for the evolution of cell specialization in metazoans.

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    © All Rights Reserved
    Download as DOCX, PDF, TXT or read online from Scribd
    Download as docx, pdf, or txt
    2K views12 pages

    Origin of Metazoa

    Uploaded by

    Sambitsabyasachi Hota
    1. Most zoologists agree that multicellular animals (metazoans) evolved from colonial protozoans. The colonial theory proposes that metazoans descended from spherical colonies of flagellated cells held together by an extracellular matrix. 2. Choanoflagellates are the closest living relatives to metazoans. Their colonies resemble what the ancestral metazoan colony may have looked like, with flagellated cells embedded in an extracellular matrix. 3. Establishing polarity along an anterior-posterior axis and internalizing growth cells allowed for both locomotion and growth, avoiding conflicts between these functions. This set the stage for the evolution of cell specialization in metazoans.

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    © All Rights Reserved
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    of 12

    1

    ORIGIN OF METAZOA
    Most zoologists agree that metazoans share a common
    ancestrywithsorneunicellularorganism.Theclassical
    colonial theory, in which Metazoa is derived from a
    colony of flagellated protozoa, is the most widely
    acceptedhypothesisamongcontemporaryzoologists(Fig.
    412). An alternative hypothesis, the syncytial theory,
    proposesthatmetazoansevolvedfromamultinucleatebut
    unicellularplasmodiumsimilartoaslimemoldorperhaps
    aciliateprotozoan.Later,membranesevolvedtoproduce
    acellboundaryaroundeachofthenuclei.Thesyncytial
    theory receives sorne support from the development of
    organismssuchasslimemoldsorinsects(Drosophila),in
    which an early multinucleated stage is followed by
    cellularization to form a multicellular body. However,
    phylogenetic analysis based on morphology and gene
    sequencesasweilasthedevelopmentalpatternsofmost
    animais contradict the syncytial theory and favor the
    colonialtheory.Forthesereasons,weconsideronlythe
    colonialtheoryhere.
    Amodernversionofthecolonialtheorystatesthatthe
    premetazoan(aprotozoan)consistedofasmallspherical
    colony bearing a surface layer of flagellated cells that
    was usedforlocomotionandfeeding(Fig.412C).The

    colonyoriginatedfromacellthatdividedrepeatedlyby
    mitosis,butthedaughtercellsdidnotseparateaftercell
    division.Thosedaughtercellsweresurroundedandheld
    together by a proteinaceous ECM in which they were
    deeply embedded. The gelatinous ECM also occupied
    much of the interior of the sphere. Similar to extant
    choanoflagellates, the cells each bore a single collared
    flagellum.Afewnonflagellatedcells,capableofgivingriseto
    flagellatedcellsandgametes,werescatteredinthesubsurface
    ECM.

    Although it is unrelated to metazoans, Volvox is an


    analogforanancestralmetazoanbecauseitdemonstrates
    howamulticellularorganismevolvedfromaunicellular
    ancestor,inthiscaseaChlamydomonaslikecell.Volvox
    isnottheancestorofMetazoa,butratherisanautotrophic
    organism with plantlike cells. rRNA sequence data
    indicatethatmulticellularityin Volvox evolved50to75
    million years ago, far too late for it to have been a
    progenitorofmetazoans,whichhadtheiroriginat!east
    600millionyearsinthepast.Thus,thevolvocidsevolved
    multicellularity inparallel withthemetazoans(andwith
    at!eastfourothergroups:fungi,brownalgae,redalgae,
    andgreenplants).

    The actual sister taxon of Metazoa is most likely


    Choanoflagellata. Among the. extant choanoflagellates,
    coloniesofProterospongiahaeckelicloselyresemblethe
    hypothetical premetazoan. The colony consists of
    flagellated collar cells embedded in the surface of a
    gelatinousECM.Cellsundergoingdivisionandlacking
    flagellaoccurdeeperintheECM.Asurfacesheetlike
    layerofcellsandanECMcontainingindividualfreecells

    foreshadowthetwoprimarymetazoantissues,epiti).elial
    andconnective,describedearlierinthischapter.Atthe
    cellular leve!, choanoflagellate collar cells are
    virtually.identical tocollar cells(choanocytes) foundin
    themetazoansponges.Achoanoflagellatecellandsponge
    choanocytebothhaveasingleflagellumsurroundedby a
    collarofmicrovilliandaflagellarshaftwithabilateral
    ~nlike vane.Inboth,theflagellumisanchored
    10
    the cell by microtubules, which radiate from the
    flagellarbasalbody.
    .
    Thehypotheticalfirstmetazoan,theprotometazoanmay
    havedifferedfromthepremetazoaninsevera!ways.
    First,thesurfacecellsprobablyclosly adjoinedorwere
    incontactwitheachother,thus
    faCihtatingintercellularcommunicationandprovidinga
    gulatory barrier between the external environment and
    ECM from the cells, thus separating the ECM into
    external and Internal layers, each of which could then
    adopt indenent functions . Third, the body was
    polarizedizedalongtheanteriorposterioraxis.Fourth,the
    separation of layers and body polarity promoted cell
    specialization.

    ORIGIN OF POLARITY AND


    CELL SPECIALIZATION

    Most motile protozoans are polarized cells that have


    leading(anterior)andtrailing(posterior)endsor,ifsessile
    and attached, they have oral free ends and aboral
    attached ends. Metazoans are similarly polarized,

    dependingonwhethertheyaremotileorsessile,buthow
    did the polarity of the multicellular metazoan body
    evolve from the unicellular polarity of protozoans? A
    cluetotheanswerisfoundintheeggsofsevera!groups
    ofmetazoans.Duringoogenesisinthesegroups,theeggs
    expressarudimentaryflagellumandacollarofmicrovilli
    atasiteonthecellsurfacethatcorrespondstotheanimal
    poleofthezygote.Incertaininvertebratetaxa,theanimal
    polecorrespondstotheanteriorendofthelarva(although
    itistheposteriorendinothers).Additionalresearchis
    needed, but the current evidence suggests a line of
    descentfromthepolarityofachoanocytetotheprimary
    anteriorposteriorpolarityofthemetazoanbody.
    Theprotometazoanprobablywaspolarizedalongan
    anteriorposterioraxis,butwhatenvironmentalconditions
    mighthaveselectedfortheevolutionofsuchpolarity?
    Foranaquaticmetazoan,theenvironmentpresentsitself
    ingradientsoflight,temperature,oxygen,andfood
    availability.If,forselectionmayhavefavoredany
    organismcapableoftrackingaresourceconcentration
    graclient.Altematively,developmentaibiologistLewis
    Wolpertsuggeststhatbodypolarityevolvedfrom
    attachmenttoasubstrat~.Attachmenttoarockin
    water,forexample,placesanorganismataninterface,a
    verysteepgradi.ent.Onceattached,variantswouldbe
    favoredthatadherewellattheattachedendandperform

    othertasks,suchasfeeding,attheoppositeend.This
    againleadstoapolarizedbody.

    Oncepolaritywasestablished,movementwouldcreate
    an environmental graclient along the locomotory axis
    that would favor differentiai expression of traits . For
    example, enhanced membrane sensitivity to
    environmental stimuli might befavoredincellsat the
    anterior end of the body because they are the first to
    encounter changes in environmental conclitions.
    Similarly,enhancedflagellargrowth,density,oractivity
    mightbefavoredincellsattheequator,orwidestpartof

    the body, since those locations best contribute to


    locomotion.Cellswithacapacityfordivision,leadingto
    growth,mightbefavoredatthe.posterlorend,because
    inthatpositiontheycontributetoandinterfere!eastwith
    locomotion.Thus,motilityalongapolaraxismayitself
    promotecellularspecializationbecausethecellsoccupy
    differentfixedpositionsinanenvironmentalgraclient.
    Accordingtoahypothesisbydevelopmentalbiologist
    LeoBuss,theoriginofrnetazoancellspecializationmay
    berelatedtoaconflictbetweenthedemandsforgrowth
    andlocomotion.Volvox,Proterospongia,andplanktonic
    blastulastagesofmetazoansrequireflagellatedsurface
    cellsforlocomotion,butmostflagellatedcellscannat
    dividebymitosisbecausethecentriolesneededtoform
    themitoticspindlesarealreadyinuseastheflagellar
    basalbodies.Ametazoanflagellatedcellcandivideonly
    aftertheflagellumregressesanditsbasalbodiesarefreed
    toformthemitoticapparatus.Thus,theoptionsfor
    growthinapremetazoancomposedsolelyofflagellated
    cells(forexample,thespeciesofProterospongia)are:
    (1)enlargementofexistingcells;(2)disassemblyof
    flagella,cellclivision,andthenflagellarreassembly;(3)
    clivisionofafewcellsscatteredthroughouttheembryo
    whileathersretainflagella;or(4)divisionofafew
    localizedcellssetasideforgrowth.Option1,becauseof
    therestrictednumberofcells,limitsultimatebodysize,

    althoughVolvoxdaughtercoloniesandafew
    postembryonicmicrometazoansgrowbycell
    enlargementonly.Option2,byrequiringtheregressionof
    flagella,compromiseslocomotion.Options3and4both
    permitgrowthandlocomotion,butmostinvertebrate
    metazoanshaveadoptedoption4.Thesettingasideof
    mitoticallyactivecellsenablesmetazoangrowthwithout
    compromisinglocomotion.(Arelatedevolutionary
    eventwasthesettingasideofgermcellscapableof
    meiosis.Surfaceflagellaareafunctionalnecessityfor
    locomotionintheprotometazoan,butwhatistheoptimal
    positionforthesetasidegrowthcells?Busssuggeststhat
    placementatthesurfacemighthaveresultedin
    overgrowthoflocomotorycellsorgrowthofatumorlike
    appendage,eitherofwhichwouldnegativelyaffect
    motility.Intemalizingthegrowthcells,however,would
    neitherdistortthebodysurfacenorintefferewiththe
    locomotorycells.Perhapsforthesereasonsthe
    protometazoandidintemalizeitsgrowthcellsandthe
    processwaspreserved,asgastrulation,initsdescendants.
    (Based on ideas in Buss, L. W 1987. The Evolution of
    lndividuality. Princeton University Press,Princeton, NJ
    201 pp.

    10

    ORIGINOFCOMPLEXITY
    Evidencesupportsthehypothesisthatmetazoansevolved
    from protozoan coloniesinwhichinitiallysimilar cells
    became specialized for different functions. If so, the
    evolutionofMetazoacanbedescribedasareplicationof
    similarunits(cells)followedbyunitspecializationand
    integration into an organism at a new, higher level of
    complexity.Thisreplicationspecializationintegrationof

    11

    units. sequence is a general pattern in the evolution of


    largebodysizeandcomplexity.Anexample,aswehave
    seen already, is the replication, specialization, and
    integrationofciliaonthebodyofmanyciliates,which
    areamongthelargest,mostdiverse,mostactive,andmost
    complexprotozoancells.Amongmetazoans,oneexample
    isabodycorriposedofaseriesofsimilarsegments,asin
    earthwormsorcrustaceans.Later,thesesegmentsbecome
    specializedandintegratedintoregions,suchasthehead,
    thorax,andabdomen.Examplesatalilevelsofbiological
    complexityareillustratednddescribed.

    References :

    Modern Text Book of Zoology: Invertebrates(Prof. R.L.Kotpal)

    https://www.wikipedia.org
    http://www.authorstream.com

    http://ebooks.cambridge.org/chapter.jsf?
    bid=CBO9780511623547&cid=CBO9780511623547
    A043

    CONCLUSION:
    From the whole topic, on the basis of different theories proposed
    by the different scientists, we can say metazoan originated from
    different types of organisms of protozoan.

    12

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