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Dearspeaker
This presentation is intended as a flexible tool for scientists, science communicators and
educators. Not all the slides will be useful for any one occasion. Choose the ones most
suitable for your audience, mix them with your own slides, or just use the diagrams.
Contents
Stem cell biology basics: For school students aged 16+, or adult public with little or
no scientific knowledge
Cloning: For adult public with little or no scientific knowledge; initial slides also
suitable for students aged 16+
Stem cell biology in more detail: For informed non-specialist audiences, e.g.
clinicians, scientists working in fields other than stem cell biology.
Presenters notes
Each slide in the Basics and Cloning sections includes notes that give a simple, jargonfree explanation of the key points. The more detailed slides in the last section have much
briefer notes and assume some knowledge of stem cell science.
Alifestory
Alifestory
Humandevelopmentstartswithjust1cell thefertilizedegg.Thiscelldividesto
produce2daughtercells.Thesedaughtersdivide,andtheirdaughtersdivideagain,and
soon.Thereareagreatmanysteps neededtoformanadultbody,orevenababy.
Alongtheway,lotsofdifferenttypesofcellsmustbemade.
Whatisastemcell?
stemcell
SELFRENEWAL
(copying)
stemcell
DIFFERENTIATION
(specializing)
specializedcell
e.g.musclecell,nervecell
Whatisastemcell?
Note:Thenextslideprovidesanalternativeversionofthisdiagramthatsomeyoungeraudiencesmayfind
easiertounderstand.Itaimstoavoidthemisconceptionthatastemcellalwaysmakesonecopyof
itselfandonespecializedcellwhenitdivides(seebelow).The conceptofastemcellisverywell
explainedintheshortfilm,AStemCellStory atwww.eurostemcell.org/films
Whatthediagramshows
Stemcellsaredifferentfromothercellsofthebodybecausestemcellscanboth:
1) Selfrenew:Makecopiesofthemselves
AND
2)Differentiate:Makeothertypesofcells specializedcellsofthebody.
Specialized ordifferentiated cellsplayparticularrolesinthebody,e.g.bloodcells,nervecells,muscle
cells.Specializedcellscannotdividetomakecopiesofthemselves.Thismakesstemcellsvery
important.Thebodyneedsstemcellstoreplacespecializedcellsthatdie,aredamagedorgetused
up.
Celldivision possiblequestions
1)16+yearoldstudentsmayrememberlearningabout2kindsofcelldivision mitosisandmeiosis.They
mayhavelearntthatmitosishappensinwoundhealingortoreplaceshortlivedcells,butprobably
wonthavediscussedstemcellsinthiscontext.Youmighttherefore needtoexplainthatmost
specializedcellscannotundergomitosis.Thereareafewexceptions(e.g.livercellsorTcells)butin
generalspecializedcellscannolongerdivide.Skincells,redbloodcellsorgutliningcellscannot
undergomitosis.Stemcellsdodividebymitosisandthismakesthemveryimportantforreplacing
lostordamagedspecializedcells.
2)Shouldmitosisbediscussed,youmaywishtonotethefollowing:Inmitosis,theDNAinthedaughter
cellsisidenticaltotheDNAinthedividingcell.Thisistrue fordividingstemcells,bothinselfrenewal
andindifferentiation.Indifferentiation,thedaughtercellsaremorespecializedthantheoriginal
stemcell.So,thedaughtercellsbehavedifferentlyeventhough theyhavethesameDNAasthestem
cell.Thisisbecausetherearelotsofothermoleculesinsideandaroundthecellsthatcanchangethe
waythecellsbehave.
3)ScientiststhinkthatwhenhumanstemcellsdividetheyprobablymakeEITHERtwostemcells,ORtwo
morespecializedcells.Infruitflies,stemcellscandivideto makeonestemcellandonemore
specializedcellinasingledivision.
Whatisastemcell?
Stem cell
SELFRENEWAL
(copying)
Stem cell
DIFFERENTIATION
(specializing)
Specialized cells
Whatisastemcell?
Note:Thepreviousslideprovidesanalternativeversionofthisdiagram.Theconceptofastemcellisvery
wellexplainedintheshortfilm,AStemCellStory atwww.eurostemcell.org/films
Whatthediagramshows
Stemcellsaredifferentfromothercellsofthebodybecausestemcellscanboth:
1. Selfrenew:Makecopiesofthemselves
AND
2.Differentiate:Makeothertypesofcells specializedcellsofthebody.
Specialized ordifferentiated cellsplayparticularrolesinthebody,e.g.bloodcells,nervecells,muscle
cells.Specializedcellscannotdividetomakecopiesofthemselves.Thismakesstemcellsvery
important.Thebodyneedsstemcellstoreplacespecializedcellsthatdie,aredamagedorgetused
up.
Celldivision possiblequestions
1)16+yearoldstudentsmayrememberlearningabout2kindsofcelldivision mitosisandmeiosis.They
mayhavelearntthatmitosishappensinwoundhealingortoreplaceshortlivedcells,butprobably
wonthavediscussedstemcellsinthiscontext.Youmighttherefore needtoexplainthatmost
specializedcellscannotundergomitosis.Thereareafewexceptions(e.g.livercellsorTcells)butin
generalspecializedcellscannolongerdivide.Skincells,redbloodcellsorgutliningcellscannot
undergomitosis.Stemcellsdodividebymitosisandthismakesthemveryimportantforreplacing
lostordamagedspecializedcells.
2) Shouldmitosisbediscussed,youmaywishtonotethefollowing: Inmitosis,theDNAinthedaughter
cellsisidenticaltotheDNAinthedividingcell.Thisistrue fordividingstemcells,bothinselfrenewal
andindifferentiation.Indifferentiation,thedaughtercellsaremorespecializedthantheoriginal
stemcell.So,thedaughtercellsbehavedifferentlyeventhough theyhavethesameDNAasthestem
cell.Thisisbecausetherearelotsofothermoleculesinsideandaroundthecellsthatcanchangethe
waythecellsbehave.
3)ScientiststhinkthatwhenhumanstemcellsdividetheyprobablymakeEITHERtwostemcells,ORtwo
morespecializedcells.Infruitflies,stemcellscandivideto makeonestemcellandonemore
specializedcellinasingledivision.
WhyselfrenewANDdifferentiate?
1stemcell
1stemcell
Selfrenewal maintains
thestemcellpool
4specializedcells
Differentiation replacesdeadordamaged
cellsthroughoutyourlife
WhyselfrenewANDdifferentiate?
1)Selfrenewalisneededbecauseifthestemcellsdidntcopythemselves,youwould
quicklyrunout.Itisimportantforthebodytomaintainapool ofstemcellstouse
throughoutyourlife.
2)Differentiationisimportantbecausespecializedcellsareusedup,damagedordieall
thetimeduringyourlife.Specializedcellscannotdivideandmakecopiesof
themselves,buttheyneedtobereplacedforyourbodytocarryonworking.For
example,yourbodyneeds100,000millionnewbloodcellseveryday.Ofcourse,
differentiationisalsoimportantformakingallthedifferentkindsofcellinthebody
duringdevelopmentofanembryofromasinglefertilizedegg.
Possiblequestionsormisconceptions
1)Schoolstudentsmayhavelearntsimplythatcellsundergomitosistomakecopiesof
themselvestohealwoundsorreplacebloodcells.Youmayneedtoexplainthat
specializedcellslikeskin,redbloodorgutcellscannotundergomitosis,whichiswhy
youneedstemcells.Thereareafewexceptions(e.g.livercellsorTcells)butin
generalspecializedcellscannolongerdivide.Foradultaudiences,thiscouldbe
expandedtocovertheideathatthereareintermediatecells(progenitors)between
stemcellsandspecializedcellsthatdividetoallowalargenumberofnewcellstobe
made(seeslide26onrenewingtissues)
2)Scientiststhinkthatstemcellsinthehumanbodydontgenerallydividetoproduce
onestemcellandonespecializedcellatthesametime.Theyprobablydivideto
makeEITHERtwostemcells,ORtwomorespecializedcells.Infruitflies,stemcells
candividetomakeonestemcellandonemorespecializedcell.
Wherearestemcellsfound?
embryonicstemcells
blastocyst averyearly
embryo
tissuestemcells
fetus,babyandthroughoutlife
Wherearestemcellsfound?
Therearedifferenttypesofstemcells:
Embryonicstemcells:foundintheblastocyst,averyearlystageembryothathasabout
50to100cells;
Tissuestemcells:foundinthetissuesofthebody(inafetus, baby,childoradult).
(Tissuestemcellsaresometimesreferredtoasadultstemcells,eventhoughtheyare
foundinthefetusandinbabies,aswellasinadults.)
embryonicstemcellstakenfrom
theinnercellmass
fluidwithnutrients
cultureinthelab
togrowmorecells
outerlayerofcells
=trophectoderm
Embryonicstemcells:Wheretheycomefrom
Embryonicstem(ES)cellsaretakenfrominsidetheblastocyst,averyearlystage
embryo.Theblastocyst isaballofabout50100cellsanditisnotyetimplantedinthe
womb.Itismadeupofanouterlayerofcells,afluidfilledspaceandagroupofcells
calledtheinnercellmass.EScellsarefoundintheinnercell mass.
Forasimple,clearexplanationofhowembryonicstemcellsareobtained,watchthe
film,AStemCellStory,atwww.eurostemcell.org/films
10
differentiation
embryonicstemcells
PLURIPOTENT
allpossibletypesofspecializedcells
Embryonicstemcells:Whattheycando
Embryonicstemcellsareexcitingbecausetheycanmakeallthedifferenttypesofcellin
thebody scientistssaythesecellsarepluripotent.
11
gr
sA
ition
d
n
o
rc
nde
ow u
embryonicstemcells
itions B
grow
und
erco
nditi
gro
onsC
w
un
de
rc
on
dit
ion
sD
skin
neurons
blood
liver
Embryonicstemcells:Challenges
Scientistsaroundtheworldaretryingtounderstandhowandwhy embryonicstemcells
produceskin,blood,nerveoranyotherparticularkindofspecializedcell.Whatcontrols
theprocesssothatthestemcellsmaketherightamountofeach celltype,attheright
time?
Thebigchallengeforscientistsistolearnhowtocontrolthesefascinatingcells.Ifwe
couldforceembryonicstemcellstomakewhateverkindofcellwewant,thenwewould
haveapowerfultoolfordevelopingtreatmentsfordisease.Forexample,perhapswe
couldgrownewinsulinproducingcellstotransplantintoapatientwithdiabetes.But
thereisagreatdealtolearnbeforesuchtherapiescanbedeveloped.Scientistsalso
wanttousestemcellsto:
Understandhowdiseasesdevelop(diseasemodelling)
Testdrugsinthelaboratory
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13
skin
testicles
brain
breast
intestines(gut)
bonemarrow
muscles
Tissuestemcells:Wherewefindthem
Weallhavestemcellsinourbodiesallthetime.Theyareessentialforkeepingusfit
andhealthy.Theyreplacecellsthataredamagedorusedup.Scientistsarestilllearning
aboutallthedifferentkindsoftissuestemcellsfoundinour bodiesandhowtheywork.
14
bloodstemcell
differentiation
foundin
bonemarrow
MULTIPOTENT
onlyspecializedtypesofbloodcell:
redbloodcells,whitebloodcells,
platelets
Tissuestemcells:Whattheycando
Tissuestemcellscanoftenmakeseveralkindsofspecializedcell,buttheyaremore
limitedthanembryonicstemcells.TissuestemcellscanONLYmakethekindsofcell
foundinthetissuetheybelongto.So,bloodstemcellscanonlymakethedifferent
kindsofcellfoundintheblood.Brainstemcellscanonlymake differenttypesofbrain
cell.Musclestemcellscan onlymakemusclecells.Andsoforth.
Scientistssaythattissuestemcellsaremultipotent becausetheycanmakemultiple
typesofspecializedcell,butNOTallthekindsofcellinyour body.
15
16
Inducedpluripotentstemcells(iPS cells)
geneticreprogramming
=addcertaingenestothecell
cellfromthebody
inducedpluripotentstem(iPS)cell
behaveslikeanembryonicstemcell
differentiation
cultureiPS cellsinthelab
Advantage:noneedforembryos!
allpossibletypesof
specializedcells
Inducedpluripotentstemcells(iPS cells)
Note:Thisslidecontainsalotofinformationandmaybetoocomplexforsome
audiencesunlessthereisplentyoftimeforexplanationsanddiscussions.
WhatareiPS cells?
In2006,scientistsdiscoveredthatitispossibletomakeanew kindofstemcellinthe
laboratory.Theyfoundthattheycouldtransformskincellsfrom amouseintocells
thatbehavejustlikeembryonicstemcells.In2007,researchers didthiswithhuman
cellstoo.Thenewstemcellsthataremadeinthelabarecalledinducedpluripotent
stemcells.Justlikeembryonicstemcells,theycanmakeallthedifferenttypesof
cellinthebody sowesaytheyarepluripotent.
Makinginducedpluripotentstem(iPS)cellsisabitliketurningbacktime.Scientistsadd
particulargenestocellsfromthebodytomakethembehavelike embryonicstem
cells.Genesgivecellsinstructionsabouthowtobehave.So,thisprocessisabitlike
changingtheinstructionsinacomputerprogramme tomakethecomputerdoanew
task.ScientistscalltheprocesstheyusetomakeiPS cellsgeneticreprogramming.
Whyaretheyexciting?
ResearchershopethatonedaytheymightbeabletouseiPS cellstohelptreatdiseases
likeParkinsonsorAlzheimers.Theyhopeto:
1) Takecellsfromthebody likeskincells fromapatient
2) MakeiPS cells
3) UsethoseiPS cellstogrowthespecializedcellsthepatientneedstorecoverfrom
thedisease,e.g.certainbraincells.Thesecellswouldbemade fromthepatients
ownskincellssothebodywouldnotrejectthem.
Thereisalongwaytogobeforescientistscandothis,butiPS cellsareanexciting
discovery.
17
Inducedpluripotentstemcells(iPS cells)
geneticreprogramming
pluripotentstemcell
(iPS)
cellfromthebody(skin)
differentiation
Inducedpluripotentstemcells(iPS cells)
Thisisanalternativerepresentationofthesameinformationas onthepreviousslide.
Pleaseseethepreviousexplanatorynotes.
18
Stemcelljargon
Potency
Ameasureofhowmanytypesofspecializedcellastemcell
can make
Pluripotent
Canmakeall typesofspecializedcellsinthebody
Embryonicstemcellsarepluripotent
Stemcelljargon
Scientistsusethewordspluripotent andmultipotent tohelpthemdescribestemcells.
ALLstemcellscanbothselfrenewanddifferentiate,BUTsomestemcellscanmake
morekindsofspecializedcellsthanothers.Thetermsontheslidearethekeyonesto
remember.Therearealsostemcellsthatare:
TOTIPOTENT:candifferentiateintoalltypesofspecializedcellsinthebodyPLUScells
thatareneededduringdevelopmentoftheembryoonly:placenta, yolksac,umbilical
cord.
UNIPOTENT:canonlydifferentiateintoonetypeofspecializedcell.Forexample,
spermatogonial stemcells(foundinthetesticles)areunipotent becausetheycanonly
formspermcells.
Ausefulplacetolookupotherwordsandphrasestodowithstemcellsisthe
EuroStemCell onlineglossary:www.eurostemcell.org/glossary
19
Cloning
20
Cloning
TherearetwoVERYdifferenttypesofcloning:
Reproductivecloning
Molecularcloning
gene1
gene2
Usetomaketwoidenticalindividuals
Usetostudywhatagenedoes
Verydifficulttodo
Routineinthebiologylabs
Illegaltodoonhumans
Cloning
Whenmostpeoplethinkofcloning,theythinkoftheideaofmakingacopyofan
individual ananimalorevenaperson.Thisiscalledreproductivecloning.Ithitthe
headlinesinthelate1990swhenDollythesheep wascloned.Shewasthefirst
mammalevertobecloned.
Infact,thiskindofcloningisverydifficulttodoanditisillegaleventotrytoclonea
humanbeing.
Thereisanothertypeofcloningthatmanybiologistsdoeveryday:molecularcloning.
Thisisatechniqueusedtohelpscientistsinvestigatewhatparticulargenesdoandhow
theywork.
Thefollowingslidesexplaintheseprocessesinmoredetail.
21
Reproductivecloning
cellfromthebody
egg
removenucleus
andtakethe
restofthecell
takethenucleus
(containingDNA)
Clone
identicaltotheindividual
thatgavethenucleus
Dollythesheep
Reproductivecloning
Dollythesheepwasthefirstclonedmammal.TomakeDolly,scientiststookthenucleus
outofanormalcellfromasheep.Theyputthatnucleusintoan eggcellthathadno
nucleus. Theythenhadanewcell.Tomakethenewcellstarttodivideandgrow,they
gaveitanelectricshock.Thenitstartedtodivideanddevelopintoan embryo.Whenit
hadgrownintoaveryearlystageembryocalledablastocyst aballofjust50100cells
itwasimplantedintothewombofanothersheepsothatitcouldgrowintoalamb
andbeborn.Thenewsheepisacloneofthesheepfromwhichthenucleuswastaken
atthestartoftheprocess.BothsheephavethesameDNA.
Notonlysheephavebeencloned.Scientistshavenowclonedmany differentanimals,
includingmice,cats,dogs,frogs,goats,horses,pigs,rabbitsandothers.However,itisa
difficultprocessanddoesnotalwayswork.Itisillegaltocloneahumanbeinginthis
way.
22
gene2
cell2
2)MakeanewpieceofDNA
gene1
gene1
gene2
gene2
3)Put newDNAintoatestcellandgrowcopies
gene1
Daughtercells
containsameDNA:
gene2
insertnewDNA
celldivides
Genes1and2have
beencloned
Molecularcloning:Principles
Molecularcloningisaprocessusedbyscientiststomakecopies ofaparticulargeneor
genesinsideacell.Theyusethetechniquetofindoutmoreaboutwhatcertain
genesdoorhowtheywork.Molecularcloningisdoneroutinelyinlaboratories
today.Itinvolvesseveralsteps:
1) TaketheDNAoutofacell.
2) Cutoutthegeneyouareinterestedin(gene2inthisexample). Insertitintoastrand
ofDNAtakenfromanothercell.Thegeneisnotliterallycutoutwithaknifeor
scissors carefullychosenenzymesbreaktheDNAchainatparticularpoints.More
enzymesareusedtoinsertthegeneintoanotherpieceofDNAat exactlytheright
place(inthisexample,nexttogene1).
3) OnceyouhavemadeapieceofDNAcontainingthegeneyouwanttostudy,put
yournewDNAintoatestcell.Whenthecelldivides,itmakescopiesofitself.Each
newdaughtercellcontainsanexactcopyoftheDNAinyourtest cell,including
genes1and2.Thegeneshavethereforebeencopiedandwesaytheyhavebeen
cloned.
Thisisasimplifieddescriptionofthetechnique.Therearesomeintermediatesteps
involvedandthedetailsofthetechniquecanvary,butthisschemeillustratesthe
keyprinciple,i.e.weareabletomakecellscontainingparticulargenesinorderto
findoutwhatthosegenesdo.Someexamplesofhowthistechniquecanbeusedare
givenonthenextslide.
23
Reportergene
Lineagetracing
removeagenetoseeif
anythingworksdifferently
addagenethatshows
uswhenanothergeneis
working
markagroupofcellsto
seewheretheir
daughtercellsendup
geneisactiveinblue
areasonly
geneispassedon
tocellsalloverthebody
eye
Normal mouse
embryo
geneAmissing
geneisinvolvedin
givingtheeyeitscolour
Molecularcloning:Applications
Molecularcloningisanimportanttoolusedbyscientiststolearnmoreabouttheroles
ofgenesindevelopmentanddisease.Someexamplesofhowmolecularcloningcanbe
usedinthelabare:
Lossoffunction(oftencalledgeneknockout): acommontechniquethathasbeen
veryusefulinhelpingscientistsunderstandhowparticulargenesareinvolvedindisease.
Ageneisremoved orblockedsothatitdoesnotwork,andthenscientistswatchtosee
whathappens.Thishasbeenofsuchwidebenefitforscienceandmedicinethat the
scientistswhodevelopedthistechnologywereawardedtheNobelPrizeforMedicinein
2007.
Reportergene: thisgenerallyinvolvesusingcolour tohelpscientistseasilyseewhena
particulargeneisworking.Areportergene isaddedtotheDNAofcells.Thisreporter
genemakesthecellsproduceacoloured protein forexample,ablueprotein.The
reportergeneisputintothecells DNArightnexttoanothergene(genex)that
scientistsreallywanttoinvestigate. Wherevergenexisactive(orswitchedon)ina
cell,thereportergeneisalsoactive.Thismeansthecellmakestheblueproteinand
looksblue.So,itiseasytoseewhichcells haveanactivegenexbecausethosecellsare
blue.
Lineagetracing: thisinvolveslookingtoseewhathappenstoa cellsdaughtercells,and
theirdaughters,inadevelopinganimal.First,somecellsaremarkedbygivingthem a
genethatscientistscaneasilyseeworking,e.g.agenetomakeaproteinthatisa
fluorescentgreencolour.Thismakesthecellslookgreen.Everytimethecellsdivide,
theirdaughtercellsinheritthegeneforthegreenprotein,sothedaughtercellsare
greentoo. Thisallowsustoseewhentheirmarkedcellsdivideandwheretheyendup
asananimaldevelops.
24
25
26
Stemcell
stemcell:
selfrenew
dividerarely
highpotency
rare
committedprogenitors:
transientamplifyingcells
multipotent
dividerapidly
noselfrenewal
specializedcells:
work
nodivision
Tissuestemcells:Principlesofrenewingtissues
Theslideshowsthetypicalhierarchyofcellsfromtissuestemcelltospecializedcell.
Stemcellsgiverisetocommittedprogenitors.Thesearenotfullydifferentiatedcellsbut
havedifferentpropertiesfromstemcells theyareanintermediatecelltype.
Committedprogenitorswilldividemanytimesandwillgiverisetofullydifferentiated
andfunctionalcellsviaaseriesofsteps.
Thistypicalhierarchyisapplicabletomanytypesoftissuestemcell(someexamplesare
giveninthefollowingslidestoillustratethisprinciple).
27
dendritic cell
megakaryocyte
HSC
platelets
erythrocytes
macrophage
neutrophil
bonemarrow
eosinophil
basophil
committedprogenitors
specializedcells
Tissuestemcells:Haematopoietic stemcells(HSCs)
HSCs =bloodstemcells
HSCs areisolatedfromthebonemarrow.Theygiverisetocommittedprogenitors,
whichthengiverisetoallspecializedbloodcelltypes.
28
Interneurons
Oligodendrocytes
NSC
Type2Astrocytes
Type1Astrocytes
brain
committedprogenitors
specializedcells
Tissuestemcells:Neuralstemcells(NSCs)
NSCs areisolatedfromspecificareasofthebrain.Theygiverisetocommitted
progenitors,whichthengiverisetoallspecializedbraincelltypes.
29
Gobletcells
GSC
Endocrinecells
Columnarcells
Smallintestine
committedprogenitors
specializedcells
Tissuestemcells:Gutstemcells(GSCs)
GSCs =intestinalstemcells
GSCs arepresentinthesmallintestine.Theygiverisetocommitted progenitors,which
thengiverisetoallspecializedintestinalcelltypes.
30
Cartilage(chondrocytes)
MSC
bonemarrow
Fat(adipocytes)
committedprogenitors
specializedcells
Tissuestemcells:Mesenchymal stemcells(MSCs)
MSCs areisolatedfromthebonemarrow.Theygiverisetocommittedprogenitors,
whichthengiverisetoallspecializedmesenchymal celltypes(bone,cartilage,fat).
31
32
Stemcellniches
Niche
stemcell
Microenvironmentaroundstemcellsthatprovides
supportandsignalsregulatingselfrenewaland
differentiation
Direct contact
niche
Soluble factors
Intermediate cell
Stemcellniches
Thestemcellnicheisamajorconceptinstemcellbiology.Understandingthe
microenvironmentaroundstemcellsisasimportantasunderstandingstemcells
themselves.Themicroenvironmentregulatesthebehaviorofstemcellsandthuscan
teachushowtocontrolstemcellsinculture.
Thenichecanactonastemcellbyvariousmechanisms:
Directcontactbetweenthestemcellandthenichecells
Solublefactorsreleasedbythenichethattraveltothestemcell
Intermediatecellsthatcommunicate betweenthenicheandthestemcell
Scientistsarestillworkingtounderstandexactlyhownicheswork,andmoreisknown
aboutthenichesofsomekindsofstemcellsthanothers.
33
Credits
Picture credits
Many thanks to the following people for permission to reproduce images:
Slide 17, iPS cells: Keisuke Kaji, University of Edinburgh, UK
Slide 27, blood cell diagrams: Jonas Larsson, Lund Univeristy, Sweden
Slide 29, intestinal cell diagrams: Hans Clevers and Nick Barker, Hubrecht
Institute, The Netherlands
Should you wish to re-use any of the images listed above, please contact the
owner. All other images in this presentation can be re-used freely.
Acknowledgements
Particular thanks to Dr Christele Gonneau for creating these slides and
working tirelessly to help ensure the notes are correct.
Thanks also to Freddy Radtke of EPFL, Switzerland, whose slide we copied to
make slide 27 on tissue stem cells.
34