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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
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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.

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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

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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.

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