DevelopmentalBiologyBiology4361

EarlyDevelopmentin
Invertebrates

July8,2008
 EarlyDevelopmentConcepts
Asazygotetheorganism:
consistsofonecellofonlyonetype
consistsofunrealizedgenomicpotential
operatesusingstoredmaterials
hasrelationshipswithnoothercells
isrelativelyhomogeneous*
(*NOTEthezygoteisusuallypolarizedtosomedegree)

Earlydevelopmentinvolves:
developmentofnewrawmaterialsforbuildingembryo
establishingnewrelationshipsbetweenamongnumerouscells
furtherpolarizingtheorganism
EarlyDevelopmentOverview
Cleavage
cellcyclecontrol
rapidcelldivisions
divisionsoffertilizedeggintomanycells

Establishmentofcellfate
specification

Establishmentofbodyaxes
anteriorposterior
dorsalventral
leftright

Gastrulation
cellsundergodisplacement
cellsmovetodifferentpartsoftheembryo
cellsacquirenewneighborsnewinductiverelationships
 CellCycleControl
Initially,factorsstoredintheeggcontrolcleavage(thecellcycle)e.g.
storedproteins

storedmRNAs

..initiatethecelldivision

MitosisPromotingFactor(MPF)stimulatesthecellcyclei.e.
MPFinitiatescelldivisionduringmitosisandmeiosis

divisionstopsintheabsenceofMPF

Later,theembryotakesovercontrol(necessitatingnewproteins)
CleavageStageCellCycle
MPFmitosispromotingfactor:
CyclinBcontrolscdc2activity
cdc2=cyclindependentkinase
CDKphosphorylateshistones,etc.

CyclinBdegrades
celldivisionstops

cyclinB

mitosis mitosis mitosis

CleavageStageCellCycle
CyclinBpresence/degradation
controlledbyeggcytoplasmic
proteins

earlycleavagepreprogrammed
maternallycontrolled

aftermaternalstoresusedup:
embryoniccontrolofcellcycle=

MidBlastulaTransition(MBT)
PostCleavageCellCycle
PostMBT:
cellcycleaddstwoGphases

newmRNAtranscription

celldivisionbecomesasynchronous

cyclinA cyclinB
cyclinE

cyclinD

G1 S G2 mitosis
 CytoskeletalMechanisms
Noteorientation

Karyokinesis

Cytokinesis
 CleavagePatterns
Cleavage
rapidcelldivisions
divisionsoffertilizedeggintomanycells

zygote 2cell 4cell 8cell

(1cell)
 Holoblastic(completecleavage)Cleavage Species
Yolk
Radial echinoderms,
amphioxis

Spiral annelids,
mostmolluscs,
Isolecithal flatworms

Bilateral tunicates

Rotational mammals,
nematode

Displaced amphibians
Mesolecithal
radial
Meroblastic(incompletecleavage)

Bilateral cephalopod
molluscs
Telolecithal
Discoidal fish,reptiles,
birds

Centrolethical Superficial mostinsects

SeaUrchinCleavage

holoblasticradialcleavage
 GermLayers/FateMaps

SeaUrchinZygote

GermLayers

inner= endoderm
middle= mesoderm
outer= ectoderm
 2nd meridional

1st cleavage
meridional 3rd equatorial

1st 2nd

pronuclei

3rd 4th

4th unequalcleavage
betweenanimal
andvegetal
hemispheres
 SeaUrchinCleavage

5th 6th 7th+:at128cells,

divisionsbecome
lessregular
formsBlastula

Blastula
onecellthick
5th
tightjunctionsinepithelium
cellsadhereto:
1)hyalinelayer(outside)
2)basalmembrane(inside) proteinaceous
3)neighboringcells fluidfillscavity
osmoticpressure
ciliadevelopembryorotates expandsblastocoel
hatchinginitiatedat
animalpole
 CellFate
Cellsarespecifiedbyeither:
asymmetricdistributionofpatterning
moleculesintoparticularcells
or
cellcellinteractions

Mechanismsforestablishingasymmetry:
1.Patterningmoleculesboundtoeggcytoskeleton

2.Moleculesactivelytransportedalongthecytoskeleton

3.Moleculesbecomeassociatedwithonecentrosome,andthen
followthatcentrosomeintooneofthetwomitoticsistercells

Onceasymmetryisestablished,onecellcanspecifyanother
(andparticipateinreciprocalinductions)
 SeaUrchinCellFate

mesenchymelooselypacked,unconnectedmesoderm,neuralcrestorigin
coelominternalbodywallmesodermorigin
 SeaUrchinCellFate

60cellstage

specified,

butnotcommitted

blastomeresundergo
conditionalspecification
(mostly..)
 BlastomereSpecification

micromeresaretheonlyblastomerestoundergo
autonomousspecification

Allothercelltypesundergoconditionalspecification
mostearlyinductioneventstriggeredbymicromeres
 MicromereSpecification
cateninplaysthemajorrolein
micromerespecification

catenintranscriptionfactor
oftenactivatedbyWntsignal
transductionpathway

catenin

cateninalsoaccumulatesincellsfatedto
becomeendodermandmesoderm
Endoderm&MesodermSpecification
catenininveg2 cellsbiasesthemtobecomeendomesoderm

NOTE Lithiumchloride(LiCl)causesaccumulationofcatenin
ineverycelltransformspresumptiveectodermintoendoderm

Micromeresignalearlyveg2(?)amplifiesendomesodermspecification

endomesoderm
catenin:

endomesoderm earlyveg2:
SecondaryMesenchymeSpecification
MicromereDeltaproteinactivatesNotchpathwayinadjacentveg2 cells

micromere Notchpathwayactivationresultsin
secondarymesenchymefate
(ratherthanendoderm)

veg2

Notch

Delta
 SpecificationofEndoderm
Wnt8ismadebymicromeresandendodermcells
(i.e.endomesodermcellsnotreceivingDeltasignal)

Wnt8actsasanautocrinesignal
boostsspecificationofbothveg2 endodermandmicromeres

Wnt8
AxisFormation
 UrchinAxes

JeanMarieCavanihac

Dorsal/ventral?
anterior/posterior?
left/right?

JeanMarieCavanihac
 AxisSpecification

Anterior

Aboral Oral

Posterior
 AxisSpecification
Anteriorposterior
Cellfateslineupalongananimalvegetalaxisestablishedinthe
eggcytoplasmpriortofertilization.
animalvegetalaxisappearstostructurefutureanteriorposterioraxis
thevegetalregionsequestersmaternalcomponentsnecessary
forposteriordevelopment
Anterior
Dorsalventral
Dorsalventral(oralaboral)axis
Aboral Oral
establishedafterfertilization
approximately45 clockwise
fromthe1st cleavageplane
oralfatespromotedbyNodal
aboralfatespromotedbyBMP2/4 Posterior
*
Leftright
Leftrightaxisestablishedafteroralaboral
asymmetricexpressionofaNodalgene
 CellMovements
Invagination Ingression

Infoldingregionofcells Migrationofindividualcellsfrom
theinternalsurfacelayerinto
theinterioroftheembryo
cellsbecomemesenchymal
Involution migrateindependently

Inturningorinwardmovementofan
expandingouterlayerspreadsoverthe
internalsurfaceoftheexternalcelllayer
 CellMovements,cont.
Delamination

Splittingonecellularsheetintotwoparallel
sheets(resemblesingression,butresults
inanewlayerofcells)

Epiboly

Epithelialsheets(usuallyectoderm)spreadasa
unittoenclosedeeperlayersoftheembryo
Occursbycellsdividing,changingshape,orby
severallayersofcellsintercalatingintofewerlayers

Intercalation
UrchinGastrulation

ingression
skeletogenic
mesenchyme
 LarvalStages
prism pluteus
 PrimaryMesenchymeCellIngression
adhesion
 SyncitialCables

syncytialcables
InvaginationoftheVegetalPlate
ConvergentExtension
 MidGastrula

secondarymesenchyme

filopodialextensions
 ImaginalRudiment

5foldradialsymmetry