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A.1Neuraldevelopment
Fillerwords
1.Outlinehowtheneuraltubeofembryonicchordatesisformedbyinfoldingof
ectodermfollowedbyelongationofthetube
Thedevelopmentofthehumancentralnervoussystembeginsintheembryoduringthefirst
12weeksafterconception.Asimilarpatternofdevelopmentcanbeobservedinall
chordates(vertebratesandotheranimalsthatpossessasupportingdorsalrodcalleda
notochord).Intheembryonicectoderm,anareaofcellscalledtheneuralplatedevelops
andbecomesaregionknownastheneuralgroove.
Itisthesecellsthateventuallybecomethebrainandspinalcord.Thecellsoftheneural
plategraduallychangeanddevelopintofolds,whichextendupfromitandeventuallymeet
andcloseovertobegintheformationofatube.Inthisway,asthegrowthofthefolds
continues,theneuralgroovedevelopsintotheneuraltube.Thetubeelongatesanditsouter
cellsformthefoundationofthenervoussystem-cellsattheanterior(front)endbecomethe
brainandthoseintheposterior(back)regionbecomethespinalcord.
neuralplate
ectoderm--------
neuraIfold--------———n]
Wneuiral'growove
groove尸7一
deepens心
i
neuralgroove
closesovertox__neural
formneuralL~JX—-4crestcells
neural,tubbeZ\
separatesfrom
ectoderm
2.Explainhowneuronsareinitiallyproducedbydifferentiationintheneuraltube
Therearebillionsofneuronsinthecentralnervoussystem(CNS),mostoftheminthebrain.
Theoriginsoftheseneuronscanbetracedbacktotheearlystagesofembryonic
developmentwhenpartoftheectodermdevelopsintoneuroectodermalcellsintheneural
plate.Althoughnotyetneurons,thedevelopmentalfateofthesecellsisnowdeterminedand
itisfromthemthatthenervoussystemisformed.
Theneuralplatedevelopsintotheneuraltube,withthecontinuedproliferationofcellsby
mitosisanddifferentiationalongthepathwaysleadingtothecellsbecomingfunctioning
neurons.ThematureCNShasfarmoreneuronsthanwhatisinitiallypresentinthe
embryonicneuraltube,socellproliferationcontinuesinboththedevelopingspinalcordand
brain.Althoughcelldivisionceasesbeforebirthinmostpartsofthenervoussystem,there
aremanypartsofthebrainwherenewneuronsareproducedduringadulthood.
3.Describehowimmatureneuronsmigratetoafinallocation
Neuron
(signalling)
StemcellGlialcell
(selfrenewing){support)
Withinthedevelopingbrain,neuronsmigratetodifferentareasandbegintoconnectwith
otherneuronstomakeupneuralnetworks.Astheembryoandthenthefetusdevelops,
neuronsthatareproducedinthedevelopingbrainmigratetodifferentpositions.Somemove
byamethodknownassomalmigrationinwhichaneuronmovesbyextendingalong
processfromitscellbodytotheouterregionofthebrain.Thisregionwilllaterbecomethe
cortex.Thelongprocessattachesitselftotheoutersurfaceofthebrainandthecellnucleus
thentravels,throughitscytoplasmtotakeupanewposition.Onceyoungneuronshave
reachedtheirtargetregion,theymustbecomepartofanetworkinordertobeableto
processinformation.
Neuronsneedtodeveloptheaxonsanddendrites(extensionsoftheircellbodies)that
enablethemtocommunicatewithotherneurons.Axonssendsignalsawayfromtheneuron
cellbody,whiledendritesreceivesignalsfromotherneurons.Eachcelldevelopsanetwork
ofmanydendritesclosetothecellbodyandasingleaxonthatcanextendforsomedistance
awayfromthecell.Everyaxonextendsfromanareaknownasa'growthcone'atthe
extremeendoftheaxon.
Thegrowthconerespondstoandisguidedbymoleculesofcertainchemicalsubstances,
whichdirectittothecorrectarea.Someofthesechemicalstimuliareattractivewhileothers
repelthegrowingaxon.Axonsdevelopwithintheareasthatwillbecomethebrainbutalso
extendbeyondtheneuraltubetoreachotherpartsofthebody.Whenanaxonhasreached
itstargetarea,itbeginstodevelopmanysynapseswithothercells.Everydeveloping
neuronformmultiplesynapsesandthesesynapsesallowforcommunicationbetweenthe
cellsofthenervoussystemvianeurotransmitters.
4.Outlinehowanaxongrowsfromeachimmatureneuroninresponseto
chemicalstimuli
Animmatureneuronconsistsofacellbodywithcytoplasmandanucleus.Anaxonisalong
narrowoutgrowthfromthecellbodythatcarriessignalstootherneurons.Onlyoneaxon
developsoneachneuron,butitmaybehighlybranched.
Manysmallerdendritesthatbringimpulsesfromotherneuronstothecellbodymayalso
develop.Chemicalstimulidetermineneurondifferentiationwhentheaxongrowsoutfrom
thecellbodyandalsothedirectioninwhichitgrowsinthedevelopingembryo.
Axonsgrowattheirtips.Insomecases,theyarerelativelyshortandmakeconnections
betweenneuronswithinthecentralnervoussystem,butotherneuronsdevelopverylong
axonswhichcanreachanypartofthebody.Despiteonlybeingoutgrowthsofasinglecell,
axonscanbemorethanametrelonginhumansandmanymetreslonginlargermammals
suchasbluewhales.
AxonGuidancebvChemicalSignals
Axonscarryimpulsestootherneuronsortocellsthatactaseffectors-eithermuscleor
glandcells.Aslongasthecellbodyofitsneuronremainsintact,itsaxonmaybeableto
regrowifseveredordamagedinotherwaysoutsidethecentralnervoussystem.
Regrowthratescanbeasrapidasvemillimetresperdaysosensationorcontrolofmuscles
cansometimesreturnovertimeafterdamage.Ofcourse,thisrecoverydependsonthe
correctconnectionsbeingre-establishedbetweenanaxonandthecellswithwhichitshould
becommunicating.
5.Statethatdevelopingneuronformsmultiplesynapses.
Thegrowthofanaxonordendriteisdirectedsothatitreachesacellwithwhichitinteracts.
Asynapseisthendevelopedbetweentheneuronandtheothercell.Theaxonsofmotor
neuronsdevelopsynapseswithstriatedmusclefibresorglandcellsforexample.Synapse
developmentinvolvesspecialstructuresbeingassembledinthemembranesoneitherside
ofthesynapseandinthesynapticcleftbetweenthem.
Thesmallestnumberofsynapsesthataneuroncouldtheoreticallyhaveistwo-oneto
bringimpulsesfromanothercellandanothertopassthemon.Inpractice,mostneurons
developmultiplesynapsesandsomeneuronsinthebraindevelophundreds,allowing
complexpatternsofcommunication.
TypesofSynapseswithintheCentralNervousSystem
Manysynapsesareformedatanearlystageofdevelopment,butnewsynapsescanbe
formedatanystageoflife.Synapsesoftendisappeariftheyarenotused.When
transmissionoccursatasynapse,chemicalmakersareleftthatcausethesynapsetobe
strengthened.Synapsesthatareinactivedonothavethesemarkerssobecomeweakerand
areeventuallyeliminated.Themaxim"useitorloseit"thereforedescribessynapsesvery
well.
6.Explaintheprocessofneuralpruning
Duringhumandevelopment,thereisanenormousamountofcellgrowth,butthedeathof
neuronsisalsoimportant.Naturaldeathofcells(apoptosis)removesabouthalfofthe
neuronsincertainregionsofthebrainand,inaddition,thesecondtypeofcellmodification
knownas'neuralpruning'removesuptohalfthesynapsesthathavedevelopedbetween
neurons.
6years
SynapticPruning
Thesetwotypesofcelldeathareessentialtoremoveunusedneuronsandtherebyhelpto
establishandstreamlinethecomplexnervenetworksinthebrain.Thetimingofthetwo
typesofcelldeathisdifferent:mostcelldeath(apoptosis)occursbeforeababyisbornbut
mostneuralpruningandsynapticmodificationoccurafterbirth.
7.Explainhowtheplasticityofthenervoussystemallowsittochangewith
experience
Brainplasticityistheabilityofthenervoussystemtochangeinbothstructureandfunction
overaperson'slife,astheyreacttothechangesintheirenvironment.Asapersonacquires
newknowledge,learnsnewskills,orhasnewexperiences,thebraincanestablishnew
neuralpathways.Throughpracticeandrevision,communicationbetweensynapsesis
enhancedandsignalstravelmoreefficiently.
Later,ifthesameneuralpathwayisusedagain,theconnectionsbetweentheneuronsare
re-establishedandeachnewattemptmeansthatmemoryandcognitionaremadefaster.
Thissynapticplasticityisestablishedinasimilarwaytothewayawalkermightlearna
pathwaythroughafieldofcorn.Ifthepathisusedeveryday,aclearpathwillsoonbecome
establishedandthewalkerwillbeabletocrossthefieldmorequicklyandefficiently.
Mostofyouguysreadingthiscanexperiencesynapticplasticityasyoureviseforyour
exams.Ontheotherhand,ifsynapticconnectionsarenotused,pathwaysmaybelostand
unusedneuronscanberemovedduringneuralpruning.
MechanismsofNeuroplasticitv
Rerouting:NewconnectionsaremadebetweenSprouting:Newaxonanddendriteextensions
activeneuronstocreatealternateneuralpathwaysallowexistingneuronstoformnewconnections
NeuralconnectionsbyCollateralsproutingof
neuronAandneuronBneuronBafterdamage
PrimarypathSecondarypathTerminatedpathpriortodegenerationtoaxonofneuronA
Synapticplasticityenhancesconnectionsbetweenneurons,butasecondexampleofthe
changesthatcanoccurinthenervoussystemisneurogenesis,thebirthandproliferationof
newneuronsinthebrain.Formanyyearsitwasbelievedthatwhenneuronsdied,theywere
neverreplaced,butstemcellshavebeenfoundincertainareasofthebrain(the
hippocampusandthedentategyrus)thatareabletoreproduceandmigratetootherareas
ofthebrainwheretheyareneeded.Thismayoccurafteratraumaticeventsuchastroke,
whichk川smanybraincells.Neurogenesisallowsthebraintoreplacecellsthathavedied
andrestorefunctionsthathavebeenlost.Alternatively,insomecases,undamagednerves
indifferentareasofthebrainareabletotakeovertherolesofcellsthathavediedand
restoresomeofthefunctionslostwhenapersonhasastroke.
TedTalkaboutneuroplasicity
8.Determinethecausesofspinabifida
Spinabifidaisacongenitaldisordercausedwhentheneuraltubefailstocloseproperly
duringembryonicdevelopment.Somevertebraedonotformcompletelyandremainunfused
andopen,sothataportionofthespinalcordmaypassthroughtheopeninginthebones.
Thisoccursmostofteninthelowerbackinthelumbarorsacralvertebrae.Spinabifidacan
betreatedbysurgerysoonafterababyisborn.
Thesurgeonplacesthespinalcordbackintothebodyandclosesthegapbetweenthe
vertebrae,buttheaffectedpartofthespinalcordw川notbeabletofunctionnormally.Spina
bifidaisoneofthemostcommonbirthdefectsandoccursinapproximately1in1000births
worldwide.Folicacid,takenasadietarysupplementduringthefirstthreemonthsof
pregnancy,hasbeenfoundtogreatlyreducetheriskofspinabifidaandotherneuraltube
defects.
TypesofSoinaBifida
cerebrospinal
fluid
vertebral
meninges
body
SpinabifidaoccultaMeningocele
9.Outlinethepromotionofthereorganizationofbrainfunction.
ComparisonofVertebrae
NOFUSION
(foramenexposed)
-0、
Transverse
Body
process
(centrum)
NormalHumanVertebrae(Thoracic)SpinaBifidaVertebrae(Thoracic)
Anischemicstrokeisadisruptionofthesupplyofbloodtoapartofthebrain.Moststrokes
arecausedbyabloodclotblockingoneofthesmallvesselsinthebrain,butbleedingfroma
bloodvesselisanothercause.Duringastrokepartofthebrainisdeprivedofsufficient
oxygenandglucose.Ifcellrespirationceasesinneurons,theybecomeirreparablydamaged
anddie.Strokesvarygreatlyinseverity.
Manyaresominorthatthepatienthardlynotices.Aboutone-thirdofsufferersfrommajor
strokesmakeafullrecoveryandanotherthirdsurvivebutareleftwithadisability.Inmany
cases,recoveryfromstrokesinvolvespartsofthebraintakingonnewfunctionsto
supplementthedamagedareas.Mostrecoveryhappensoverthefirstsixmonthsaftera
majorstrokeandmayinvolverelearningaspectsofspeechandwriting,regainingspatial
awarenessandtheabilitytocarryoutskilledphysicalactivitiessuchasdressingor
preparingfood.
10.AnnotateadiagramofembryonictissuesinXenopus,usedasananimal
model,duringneurulation.
Xenopusareagenusoffrogthatrobustembryosthatcantolerateextensivemanipulation.
Thismakesthemsuitableanimalmodelsforinvestigatingthedevelopmentalstagesof
embryogenesis.Duringneurulation,thefollowingembryonictissuesshouldbeeasily
identifiable:
1.Threegermlayers(outer=ectoderm;middle=mesoderm;inner=endoderm)
2.Ahollowcavitycalledthearchenteron(willdevelopintothedigestivetract)
3.Notochord(theflexiblerodthatstimulatesneurulation)
4.Neuraltube(developedfromtheunfoldingoftheneuralplate)
StructureandfunctionrelationshipbetweenthetissuelayersofaXenopusembryo
Transversecross-sectionofaXenopusembryo(left:micrograph;right:schematic)
DorsalDorsal
-endoderm-
VentralVentral
A.2Thehumanbrain
1.Explainhowtheanteriorpartoftheneuraltubeexpandstoformthebrain
Duringthedevelopmentofvertebrateembryos,aneuraltubeformsalongthewholeofthe
dorsalside,abovethegut,nearthesurface.Mostoftheneuraltubebecomesthespinal
cord,buttheanteriorendexpandsanddevelopsintothebrainaspartofaprocesscalled
cephalization,thedevelopmentofahead.
Thehumanbraincontainsapproximately86billionneurons.Thebrainactsasthecentral
controlcentreforthewholebody,bothdirectlyfromcranialnervesandindirectlyviathe
spinalcordandnumeroussignalmoleculescarriedbytheblood.Theadvantageofhavinga
brainisthatcommunicationbetweenthebillionsofneuronsinvolvedcanbemorerapidthan
ifcontrolcentresweremoredispersed.Themajorsensoryorgansarelocatedattheanterior
endofvertebrates:theeyes,ears,nose,andtongue.
TypesofStrokes
Hemorrhage/blood
leaksintobraintissue
EmbryonicBrainRegionsDevelopedBrainStructures
Forebrain
Midbrain••MesencephalonPartsofbrainstem
MetencephalonPonsCerebellum
Hindbrain—
MyelencephalonMedullaoblongata
Embryoat3weeksEmbryoat7weeksEmbryoat11weeks
2.Describethedifferentpartsofthebrainandtheirspecificroles
Thebrainhasregionsthataredistinguishablebytheirshape,colourorbyamicroscopic
structure.Theseregionshavedifferentroles,identifiedbyphysiologicalresearchinhumans
andothermammals.Eachpartofthebrainhasaparticularfunction,someregulating
automaticprocesses,suchasheartbeatandbalance,whileotherscontrolourphysical
coordination,speech,andabilitytoreason.
1.Thecerebralhemispheresarethelargestpartofthebrainandformthe
coordinatingcentreforlearning,memory,language,andreasoning.Theseregions
receiveinformationfromthesenseorgansandcoordinateandorganisemotor
functions.
2.Thehypothalamuscontrolstheautonomicnervoussystem.Itcoordinatesthe
endocrineandnervoussystemsbyregulatingthesecretionsofthepituitarygland.
3.Thecerebellumcoordinatesmovement,posture,andbalance.
4.Themedullaoblongata(brainstem)controlsautomaticandhomeostaticactivities
suchasbreathing,swallowing,digestion,andheartrate.
5.Thepituitaryglandhastwoparts-theposteriorlobestoresreleasethehormones
oxytocinandADHfromthehypothalamus,whiletheanteriorlobeproducesand
secretessevenhormones,includingFSHandgrowthhormone,whichregulatemany
ofthebody'sfunctions.
3.Identifypartsofthebraininaphotograph,diagram,orscanofthebrain,
includingthemedullaoblongata,cerebellum,hypothalamus,pituitarygland,
andcerebralhemispheres
rightcerebralhemisphere
----------(lefthemispherenot
hypothalamus/(
pituitary、一1
glandcerebellum
、\、medullaoblongata
|干spinalcord
CerTebralHemisphere----------/
呼二
PituitaryGland/
CerebralHemisphere—4——j-
/
屋-****</f)\\」CorpusCallosum
Hypothalamus-------
v____Cerebellum
"1.4/Ponsi
PituitaryGland/
Brainstem
•------MedullaOblongata
4.Explainhowtheautonomicnervoussystemcontrolsinvoluntaryprocessesin
thebody
Theperipheralnervoussystemcomprisesallthenervesoutsidethecentralnervoussystem.
Itisdividedintotwoparts:thevoluntaryandtheautonomicnervoussystems.Involuntary
processesarecontrolledbytheautonomicnervoussystem,usingcentresinthemedulla
oblongata.
Theautonomicnervoussystemhastwoparts:sympatheticandparasympathetic.These
oftenhavecontraryeffectsonaninvoluntaryprocess.Forexample,parasympatheticnerves
causeanincreaseinbloodowtothegutwallduringdigestionandabsorptionoffood.
Sympatheticnervescauseadecreaseinbloodowduringfastingorwhenbloodisneeded
elsewhere.
5.Outlinehowcerebralcortexformsalargerproportionofthebrain
Thecerebralcortexistheouterlayerofthecerebralhemispheres.Althoughitisonlytwoto
fourmillimetresthick,uptosixdistinctivelydifferentlayersofneuronscanbeidentifiedin
sectionsstudiedunderamicroscope.Itishavingahighlycomplexarchitectureofneurons
andprocessesthemostcomplextasksinthebrain.
Onlymammalshaveacerebralcortex.Birdsandreptileshaveregionsofthebrainthat
performasimilarrangeoffunctionsbuttheyarestructurallydifferent,withcellsarrangedin
clustersratherthanlayers.Amongthemammals,thecerebralcortexvariesinsize
considerably.Inhumans,itformsalargerproportionofthebrainthaninanyothermammal.
AutonomicControlofBodySystems
PARASYMPATHETICNERVESSYMPATHETICNERVES
**Rest&digest"/"Feed&breed''••Fightorflight,
ConstrictpupilsDilatepupils
SlowheartbeatIncreaseheartbeat
ConstrictairwaysRelaxairways
StimulatestomachactivityInhibitstomachactivity,J
InhibitglucosereleaseStimulateglucoserelease
Secrete
StimulatesexualarousalPromoteejaculation
(bothmanandwoman)&vaginalcontractions
frontallobe
occipital
lobe
temporal
lobemedulla
oblongata
cerebellum
6.Thehumancerebralcortexhasbecomeenlargedprincipallybyanincreasein
totalareawithextensivefoldingtoaccommodateitwithinthecranium
Thecerebralcortexhasbecomegreatlyenlargedduringhumanevolution,andnowcontains
moreneuronsthanthatofanyotheranimal.Therehasbeenamodestincreaseinthickness,
butthecortexisstillonlyafewmillimetresthick.Theincreaseisdueprincipallytoan
increaseintotalareaandthatnecessitatesthecortexbecomingextensivelyfoldedduring
development.
Thisissolargethatthebraincanonlybeaccommodatedinsideagreatlyenlargedcranium,
formingthedistinctiveshapeofthehumanskull.Mostofthesurfaceareaofthecerebral
cortexisinthefoldsratherthanontheoutersurface.Incontrast,miceandratshavean
unfoldedsmoothcortex,butincats,therearesomefoldsandelephantsanddolphinshave
more.Amongtheprimates,monkeysandapesshowarangeofcortexsizeanddegreeof
folding,withlargersizesinprimatesthataremorecloselyrelatedtohumans.
BrainComparison—HumanversusRat(NottoScale)
HumanBrain
smoothcerebralcortexhighlyfoldedcerebralcortex
ElephantMarmosetRhesusGorillaChimpanzeeHuman
monkey
7.Thecerebralhemispheresareresponsibleforhigher-orderfunctions.
Thecerebralhemispherescarryoutthemostcomplexofthebrain'stasks.Theseareknown
ashigher-orderfunctionsandincludelearning,memory,speech,andemotions.These
higher-orderfunctionsinvolveassociationofstimulifromdifferentsourcesincludingtheeye
andearandalsofrommemories.
Theyrelyonverycomplexnetworksofneuronsthatarestillonlypartiallyunderstoodby
neurobiologists.Themostsophisticatedthoughtprocessessuchasreasoning,decision
makingandplanningoccurinthefrontalandprefrontallobesofthecerebralcortex.Using
thesepartsofthebrainwecanorganizeouractionsinalogicalsequence,predicttheir
outcomes,developasenseofrightandwrongandbeawareofourownexistence.
LateralizationofBrainFunction
LeftHemisphereRightHemisphere
•Sensorystimulus•Sensorystimulus
fromrightsideofbodyfromleftsideofbody
•Motorcontrolof•Motorcontrolof
rightsideofbodyleftsideofbody
•Speech,languageand•Creativity
comprehension
•Spatialability
•Analysisandcalculations
•Context/perception
•Timeandsequencing
•Recognitionoffaces,
•Recognitionofwords,placesandobjects
lettersandnumbers
8.Theleftcerebralhemispherereceivessensoryinputfromsensoryreceptorsin
therightsideofthebodyandtherightsideofthevisualfieldinbotheyesand
viceversafortherighthemisphere
Thecerebralhemispheresreceivesensoryinputsfromallthesenseorgansofthebody.For
example,signalsfromtheearpasstotheauditoryareainthetemporallobe.Signalsfrom
theleftearpasstothelefthemisphereandfromtherighteartotherighthemisphere.Inputs
fromtheskin,musclesandotherinternalorganspassviathespinalcordtothe
somatosensoryareaoftheparietallobe.
Perhapssurprisingly,theimpulsesfromeachsidecrossinthebaseofthebrainsothatthe
lefthemispherereceivesimpulsesfromtherightsideofthebodyandviceversa.Inputsfrom
theeyepasstothevisualareaintheoccipitallobe,knownasthevisualcortex.Impulses
fromtherightsideoftheeldofvisionineacheyearepassedtothevisualcortexintheleft
hemisphere,whileimpulsesfromtheleftsideoftheeldofvisionineacheyepasstothe
righthemisphere.Thisintegrationofinputsenablesthebraintojudgedistanceand
perspective.
ContralateralProcessing
VisualProcessingSensorimotorProcessing
Regionsineachofthecerebralhemispherescontrolstriated("voluntary")muscles.Themain
regionisintheposteriorpartofthefrontallobeandiscalledtheprimarymotorcortex.Inthis
region,thereisaseriesofoverlappingareasthatcontrolmusclesthroughoutthebody,from
themouthatoneendoftheprimarymotorcortextothetoesattheotherend.Theprimary
motorcortexinthelefthemispherecontrolsmuscle
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