Peter J.

Russell
AmolecularApproach2ndEdition

CHAPTER 2
DNA: The Genetic Material

editedbyYueWenWangPh.D.
Dept.ofAgronomy,NTU

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TheSearchfortheGeneticMaterial
1. Somesubstancemustberesponsibleforpassage
oftraitsfromparentstooffspring.Forasubstance
todothisitmustbe:
a.Stableenoughtostoreinformationforlongperiods.
b.Abletoreplicateaccurately.
c.Capableofchangetoallowevolution.

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TheSearchfortheGeneticMaterial
2. Intheearly1900s,chromosomeswereshownto
bethecarriersofhereditaryinformation.In
eukaryotestheyarecomposedofbothDNAand
protein,andmostscientistsinitiallybelievedthat
proteinmustbethegeneticmaterial.

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GriffithsTransformationExperiment
1. FrederickGriffiths1928experimentwith
Streptococcuspneumoniaebacteriainmice
showedthatsomethingpassedfromdeadbacteria
intonearbylivingones,allowingthemtochange
theircellsurface.
2. Hecalledthisagentthetransformingprinciple,
butdidnotknowwhatitwasorhowitworked.

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Fig. 2.2 Griffiths transformation experiment

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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AverysTransformationExperiment
Animation:DNAasGeneticMaterial:TheAveryExperiment
1.In1944,Avery,MacLeodandMcCartypublishedresultsofastudythat
identifiedthetransformingprinciplefromS.pneumoniae.Their
approachwastobreakopendeadcells,chemicallyseparatethe
components(e.g.,protein,nucleicacids)anddeterminewhichwas
capableoftransformingliveS.pneumoniaecells.
2.Onlythenucleicacidfractionwascapableoftransformingthebacteria.
3.Criticsnotedthatthenucleicacidfractionwascontaminatedwith
proteins.TheresearcherstreatedthisfractionwitheitherRNaseor
proteaseandstillfoundtransformingactivity,butwhenitwastreated
withDNase,notransformationoccurred,indicatingthatthe
transformingprinciplewasDNA.

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Fig. 2.3 Experiment that showed that DNA, not RNA, was the transforming principle

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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TheHersheyChaseBacteriophageExperiment
Animation:DNAasGeneticMaterial:TheHersheyChaseExperiment
1.MoreevidenceforDNAasthegeneticmaterialcamein1953with
AlfredHersheyandMarthaChasesworkonE.coliinfectedwith
bacteriophageT2.
2.Inonepartoftheexperiment,T2proteinswerelabeledwith 35S,andin
theotherpart,T2DNAwaslabeledwith32P.Theneachgroupof
labeledviruseswasmixedseparatelywiththeE.colihost.Afterashort
time,phageattachmentwasdisruptedwithakitchenblender,andthe
locationofthelabeldetermined.
3.The35Slabeledproteinwasfoundoutsidetheinfectedcells,whilethe
32PlabeledDNAwasinsidetheE.coli,indicatingthatDNAcarriedthe
informationneededforviralinfection.Thisprovidedadditionalsupport
fortheideathatgeneticinheritanceoccursviaDNA.

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Fig. 2.4 Bacteriophage T2

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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Fig. 2.5 Lytic life cycle of a virulent phage, such as T2

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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Fig. 2.6 Hershey-Chase experiment demonstrating DNA is genetic material

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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TheDiscoveryofRNAasViralGeneticMaterial
1. AllknowncellularorganismshaveDNAastheirgenetic
material.Someviruses,however,useRNAinstead.
2. Tobaccomosaicvirus(TMV)iscomposedofRNAand
protein;itcontainsnoDNA.In1956GiererandSchramm
showedthatwhenpurifiedRNAfromTMVisapplied
directlytotobaccoleaves,theydevelopmosaicdisease.
PretreatingthepurifiedRNAwithRNasedestroysitsability
tocauseTMVlesions(Figure2.7).
3. In1957FraenkelConratandSingershowedthatinTMV
infectionswithvirusescontainingRNAfromonestrainand
proteinfromanother,theprogenyviruseswerealwaysof
thetypespecifiedbytheRNA,notbytheprotein.

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Fig. 2.7 Typical tobacco mosaic virus (TMV) particle

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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Fig. 2.8 Demonstration that RNA is the genetic material in tobacco mosaic virus (TMV)

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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TheCompositionandStructureofDNAand
RNA
1.DNAandRNAarepolymerscomposedofmonomers
callednucleotides.
2.Eachnucleotidehasthreeparts:
a.Apentose(5carbon)sugar.
b.Anitrogenousbase.
c.Aphosphategroup.

3.ThepentosesugarinRNAisribose,andinDNAits
deoxyribose.Theonlydifferenceisatthe2position,
whereRNAhasahydroxyl(OH)group,whileDNAhas
onlyahydrogen.

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Figs. 2.9 Structures of deoxyribose and ribose in DNA and RNA

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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TheCompositionandStructureofDNAand
RNA
4. Therearetwoclassesofnitrogenousbases:
a.Purines(doublering,ninememberedstructures)
includeadenine(A)andguanine(G).
b.Pyrimidines(onering,sixmemberedstructures)
includecytosine(C),thymine(T)inDNAanduracil
(U)inRNA.

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Figs. 2.10 Structures of the nitrogenous bases in DNA and RNA

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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TheCompositionandStructureofDNAand
RNA
5.Thestructureofnucleotideshasthesefeatures:
a.Thebaseisalwaysattachedbyacovalentbondbetweenthe1 carbon
ofthepentosesugarandanitrogeninthebase(specifically,thenine
nitrogeninpurinesandtheonenitrogeninpyrimidines).
b.Thesugarbasecombinationisanucleoside.Whenaphosphateis
added(alwaystothe5carbonofthepentosesugar),itbecomesa
nucleosidephosphate,orsimplynucleotide.
c.NucleotideexamplesareshowninFigure2.11,andnaming
conventionsaregiveninTable2.1.

6.PolynucleotidesofbothDNAandRNAareformedbystablecovalent
bonds(phosphodiesterlinkages)betweenthephosphategrouponthe5
carbonofonenucleotide,andthe3hydroxylonanothernucleotide.
Thiscreatesthebackboneofanucleicacidmolecule.
7.Theasymmetryofphosphodiesterbondscreates35polaritywithinthe
nucleicacidchain.

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Fig. 2.11 Chemical structures of DNA and RNA

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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TheDiscoveryoftheDNADoubleHelix
1.JamesWatsonandFrancisCrickpublishedthefamous
doublehelixstructurein1953.Whentheybegantheir
work,itwasknownthatDNAiscomposedof
nucleotides,buthowthenucleotidesareassembledinto
nucleicacidwasunknown.Twoadditionalsourcesof
dataassistedWatsonandCrickwiththeirmodel:
a.ErwinChargaffsratiosobtainedforDNAderivedfromavariety
ofsourcesshowedthattheamountofpurinealwaysequalsthe
amountofpyrimidine,andfurther,thattheamountofGequals
C,andtheamountofAequalsT.
b.RosalindFranklinsXraydiffractionimagesofDNAshoweda
helicalstructurewithregularitiesat0.34nmand3.4nmalong
theaxisofthemolecule(Figure8.9).

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Fig. 2.12 X-ray diffraction analysis of DNA

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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Fig. 2.13 Molecular structure of DNA

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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TheDiscoveryoftheDNADoubleHelix
2.WatsonandCricksthree
dimensionalmodel(Figure2.13)
hasthesemainfeatures:
a.Itistwopolynucleotidechains
woundaroundeachotherina
righthandedhelix.
b.Thetwochainsareantiparallel.
c.Thesugarphosphatebackbones
areontheoutsideofthehelix,and
thebasesareontheinside,
stackedperpendicularlytothe
longaxislikethestepsofaspiral
staircase.

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d.Thebasesofthetwostrands
areheldtogetherbyhydrogen
bondsbetweencomplementary
bases(twoforATpairsand
threeforGCpairs).Individual
Hbondsarerelativelyweak
andsothestrandscanbe
separated(byheating,for
example).Complementary
basepairingmeansthatthe
sequenceofonestranddictates
thesequenceoftheother
strand.

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e.Thebasepairsare0.34nmapart,andonefullturnoftheDNA
helixtakes3.4nm,sothereare10bpinacompleteturn.The
diameterofadsDNAhelixis2nm.
f.BecauseofthewaythebasesHbondwitheachother,the
oppositesugarphosphatebackbonesarenotequallyspaced,
resultinginamajorandminorgroove.ThisfeatureofDNA
structureisimportantforproteinbinding.

3.The1962NobelPrizeinPhysiologyorMedicinewas
awardedtoFrancisCrick,JamesWatsonandMaurice
Wilkins(theheadofthelabinwhichFranklinworked).
Franklinhadalreadydied,andsowasnoteligible.

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DifferentDNAStructures
XraydiffractionstudiesshowthatDNAcanexistin
differentforms(Figure2.15).
a.ADNAisthedehydratedform,andsoitisnotusuallyfoundin
cells.Itisarighthandedhelixwith10.9bp/turn,withthebases
inclined13fromthehelixaxis.ADNAhasadeepandnarrow
majorgroove,andawideandshallowminorgroove.
b.BDNAisthehydratedformofDNA,thekindnormallyfound
incells.Itisalsoarighthandedhelix,withonly10.0bp/turn,
andthebasesinclinedonly2fromthehelixaxis.BDNAhasa
widemajorgrooveandanarrowminorgroove,anditsmajorand
minorgroovesareofaboutthesamedepth.
c.ZDNAisalefthandedhelixwithazigzagsugarphosphate
backbonethatgivesititsname.Ithas12.0bp/turn,withthe
basesinclined8.8fromthehelixaxis.ZDNAhasadeepminor
groove,andaveryshallowmajorgroove.Itsexistenceinliving
cellshasnotbeenproven.

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Fig. 2.15 Space-filling models of different forms of DNA. a) A-DNA b) B-DNA c) Z-DNA

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DNAintheCell
AllknowncellularDNAisintheBform.
ADNAwouldnotbeexpectedbecauseitis
dehydratedandcellsareaqueous.
ZDNAhasneverbeenfoundinlivingcells,
althoughmanyorganismshavebeenshownto
containproteinsthatwillbindtoZDNA.

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RNAStructure
1. RNAstructureisverysimilartothatofDNA.
a. Itisapolymerofribonucleotides(thesugarisriboseratherthan
deoxyribose).
b. Threeofitsbasesarethesame(A,G,andC)whileitcontainsU
ratherthanT.

2. RNAissinglestranded,butinternalbasepairingcan
producesecondarystructureinthemolecule.
3. SomevirusesuseRNAfortheirgenomes.Insomeitis
dsRNA,whileinothersitisssRNA.Doublestranded
RNAisstructurallyverysimilartodsDNA.

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 iActivity:CrackingtheViralCode

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TheOrganizationofDNAinChromosomes
1. CellularDNAisorganizedintochromosomes.A
genomeisthechromosomeorsetof
chromosomesthatcontainsalltheDNAofan
organism.
2. Inprokaryotesthegenomeisusuallyasingle
circularchromsome.Ineukaryotes,thegenomeis
onecompletehaploidsetofnuclear
chromosomes;mitochondrialandchloroplast
DNAarenotincluded.

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ViralChromosomes
1. Avirusisnucleicacidsurroundedbyaprotein
coat.ThenucleicacidmaybedsDNA,ssDNA,
dsRNAorssRNA,anditmaybelinearor
circular,asinglemoleculeorseveralsegments.
2. Bacteriophagesarevirusesthatinfectbacteria.
ThreedifferenttypesthatinfectE.coliaregood
examplesofthevarietyofchromosomestructure
foundinviruses.

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Tevenphage
3. TheTevenphages(T2,T4andT6)havesimilar
structures;allhavedsDNAgenomescomposedof
aonelinearDNAmoleculesurroundedbya
proteincoat.

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X174phage
4. X174isasmall,simpleviruswithoneshortssDNA
chromosome.In1959,RobertSinsheimerfoundthatthe
DNAofX174hasabasecompositionthatdoesnotfit
thecomplementarybasepairrules.singlestrand
DNAratherthandsDNA

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phage
5.BacteriophageissomewhatliketheTeven
phagesinstructure.However,itschromosome
changesform.AlinearmoleculeofdsDNAis
packagedinsidetheproteinhead(Fig.2.17),but
afterthevirusinfectsitshostthechromosome
becomescircularduetobasepairingof
complementary12basesinglestrandedregionsat
theendsofthelinearmolecule(Fig.2.18).

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Fig. 2.17 Bacteriophage

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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Fig. 2.18 chromosome structure varies at stages of lytic infection of E. coli

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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ProkaryoticChromosomes
1.ThetypicalprokaryoticgenomeisonecirculardsDNA
chromosome,butsomeprokaryotesaremoreexotic,with
amainchromosomeandoneormoresmallerones.When
aminorchromosomeisdispensabletothelifeofthecell,
itiscalledaplasmid.Someexamples:
a.Borreliaburgdorferi(Lymediseaseinhumans)hasa0.91Mb
linearchromosome,plusanadditional0.53MbofDNAin17
differentlinearandcircularmolecules.
b.Agrobacteriumtumefaciens(crowngalldiseaseofplants)hasa
3.0Mbcircularchromosomeanda2.1Mblinearone.

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2. Archaebacteriaalsovaryinchromosomalorganization,
butonlycircularformshavebeenfound.Examples:
a.Methanococcusjannaschiihasthreechromosomesof1.66Mb,
58kband16kb.
b.Archaeoglobusfulgidushasone2.2Mbcircularchromosome.

3.BothEubacteriaandArchaebacterialackamembrane
boundednucleus,hencetheirclassificationas
prokaryotes.TheirDNAisdenselyarrangedina
cytoplasmicregioncalledthenucleoid.
4. InanexperimentwhereE.coliisgentlylysed,itreleases
one4.6Mbcircularchromosome,highlysupercoiled
(Figures2.19and2.20).A4.6Mbdoublehelixisabout
1mminlength,about103timeslongerthananE.coli
cell.DNAsupercoilinghelpsitfitintothecell.
Animation:DNAsupercoiling

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Fig. 2.20 Illustration of DNA supercoiling

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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5. AmoleculeofBDNA,with10bp/turnofthehelix,isinrelaxed
conformation.Ifturnsofthehelixareremovedandthemolecule
circularized,theDNAwillformsuperhelicalturnstocompensateforthe
addedtension.
6. AnickinsupercoiledDNAwillallowittoreturntoarelaxedDNAcircle
(Figure2.21).
7. EitheroverwindingorunderwindingDNAwillcreateastructurewhere
10bp/turnofthehelixisnotthemostenergeticallyfavoredconformation,
andsupercoilswillbeinduced.Bothpositiveandnegativesupercoilswill
condensetheDNA.
8. AllorganismscontaintopoisomeraseenzymestosupercoiltheirDNA.
9. ProkaryotesalsoorganizetheirDNAintoloopeddomains,withtheendsof
thedomainsheldsothateachissupercoiledindependently(Figure2.22).
10.Thecompactionfactorforloopeddomainsisabout10fold.InE.colithere
areabout100domainsofabout40kbeach.

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Fig. 2.22 Model for the structure of a bacterial chromosome

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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EukaryoticChromosomes
1.Thegenomeofmostprokaryotesconsistsofone
chromosome,whilemosteukaryoteshavea
diploidnumberofchromosomes.
2.Agenomeistheinformationinonecomplete
haploidchromosomeset.Thetotalamountof
DNAinthehaploidgenomeofaspeciesisitsC
value(Table2.4).Thestructuralcomplexityand
theCvalueofanorganismarenotrelated,
creatingtheCvalueparadox.

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Cvalueparadox

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3. Theformofeukaryoticchromosomeschangesthrough
thecellcycle:
a. InG1,eachchromosomeisasinglestructure
b. InS,chromosomesduplicateintosisterchromatidsbutremain
joinedatcentromeresthroughG2
c. AtMphase,sisterchromatidsseparateintodaughter
chromosomes

4. InG1eukaryoticchromosomesarelineardsDNA,and
containabouttwiceasmuchproteinasDNAbyweight.
TheDNAproteincomplexiscalledchromatin,anditis
highlyconservedinalleukaryotes.

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ChromatinStructure
1.Bothhistonesandnonhistonesareinvolvedinphysicalstructureofthe
chromosome.
2.Histonesareabundant,smallproteinswithanet(+)charge.Thefive
maintypesareH1,H2A,H2B,H3andH4.Byweight,chromosomes
haveequalamountsofDNAandhistones.
3.Histonesarehighlyconservedbetweenspecies(H1lessthantheothers).
4.HistonesorganizeDNA,condensingitandpreparingitforfurther
condensationbynonhistoneproteins.Thiscompactionisnecessaryto
fitlargeamountsofDNA(2m/6.5ftinhumans)intothenucleusof
acell.
5.NonhistoneisageneralnameforotherproteinsassociatedwithDNA.
Thisisabiggroup,withsomestructuralproteins,andsomethatbind
onlytransiently.Nonhistoneproteinsvarywidely,evenindifferent
cellsfromthesameorganism.Mosthaveanet()charge,andbindby
attachingtohistones.

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6. Chromatinformationinvolveshistones,and
condensestheDNAsoitwillfitintothecell.
Chromatinformationhastwocomponents:
a. TwomoleculeseachofhistonesH2A,H2B,H3and
H4associatetoformanucleosomecore,andDNA
wrapsaroundit134timesfora7foldcondensation
factor.Nucleosomecoresareabout11nmindiameter

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Fig. 2.24 Basic eukaryotic chromosome structure

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b. H1furthercondensestheDNAbyconnecting
nucleosomestocreatechromatinwithadiameterof
30nm,foranadditional6foldcondensation.The
solenoidmodelproposesthatthenucleosomesforma
spiralwith6nucleosomesperturn(Figures2.24and
2.25).

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Fig. 2.25 the 30-nm chromatin fiber

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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7. Beyondthe30nmfilamentstage,electron
microscopyshows3090loopsofDNAattached
toaproteinscaffold(Figure2.26).Eachloopis
180300nucleosomesofthe30nmfiber.SARs
(scaffoldassociatedregions)bindnonhistone
proteinstoformloopsthatradiateoutinspiral
fashion(Figure2.27).
8. Fullycondensedchromosomeis10,000fold
shorterand400foldthickerthanDNAalone.

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Fig. 2.26 , 2.27

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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EuchromatinandHeterochromatin
1.ThecellcycleaffectsDNApacking,withDNAcondensingformitosis
andmeiosis,anddecondensingduringinterphase.Chromosomesare
mostcondensedatmetaphase,whentheloopeddomainsarefurther
coiledandthechromatichasadiameterofabout700nm.Nonhistone
proteinsformthescaffoldforthisadditionalcondensation.
2.Stainingofchromatinrevealstwoforms:
a.Euchromatincondensesanddecondenseswiththecellcycle.Itis
activelytranscribed,andlacksrepetitivesequences.Euchromatin
accountsformostofthegenomeinactivecells.
b.Heterochromatinremainscondensedthroughoutthecellcycle.It
replicateslaterthaneuchromatin,andistranscriptionallyinactive.
Therearetwotypesbasedonactivity:
i.Constituitiveheterochromatinoccursatthesamesitesinboth
homologouschromosomesofapair,andismostlyrepetitiveDNA
(e.g.,centromeres).
ii.Facultativeheterochromatinvariesbetweencelltypesor
developmentalstagesorevenbetweenhomologouschromosomes.
Itcontainscondensed,andthusinactive,euchromatin(e.g.,Barr
bodies)

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CentromericandTelomericDNA
1.Centromeresandtelomeresareeukaryoticchromosomal
regionswithspecialfunctions.
2.Centromeresarethesiteofthekinetochore,wherespindle
fibersattachduringmitosisandmeiosis.Theyare
requiredforaccuratesegregationofchromatids.
3.Yeast(Saccharomycescerevisiae)centromeresarewell
studied.CalledCENregions,theirsequenceand
organizationaresimilar,butnotidentical,betweenthe
chromosomes.Othereukaryoteshavedifferent
centromeresequences,sowhilefunctionisconserved,it
isnotduetoasingletypeofDNAsequence.(Fig.228)

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Fig. 2.28 Consensus sequence for centromeres of the yeast Saccharomyces cerevisiae

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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4.Proteinsinteractwiththecentromereandthe
spindlemircrotubuletoformthekinetchore
structure(Fig.229)

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Fig. 2.29 Hypothetical model for the kinetochore of yeast, showing the relationship of
the spindle fiber microtubule to the centromere DNA elements

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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5.Telomeresareneededforchromosomalreplicationandstability.
Generallycomposedofheterochromatin,theyinteractwithboththe
nuclearenvelopeandeachother.Alltelomeresinaspecieshavethe
samesequence.
a.Simpletelomericsequencesareshort,speciesspecificandtandemly
repeated.(Examples:Tetrahymenais5TTGGGG3,andhumanis
5TTAGGG3.)Anewmodelsuggeststhatthesinglestrandedend
ofthechromosomefoldsbacktoformatloopandtheninvadesthe
doublestrandedregiontoformaDloop(Figure2.30).
b.Telomereassociatedsequencesareinternaltothesimpletelomeric
sequences.Thesecomplexrepetitivesequencesmayextendmanykb
intothechromosome.
c.Drosophilaisunusualinhavingtelomerescomposedoftransposons,
ratherthantheshortrepeatsseeninmosteukaryotes.

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Fig. 2.30 Telomeres

Peter J. Russell, iGenetics: Copyright Pearson Education, Inc., publishing as Benjamin Cummings.

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UniqueSequenceandRepetitiveSequence
DNA
1.Sequencesvarywidelyinhowoftentheyoccurwithina
genome.Thecategoriesare:
a.UniquesequenceDNA,presentinoneorafewcopies.
b.ModeratelyrepetitiveDNA,presentinafewto105copies.
c.HighlyrepetitiveDNA,presentinabout105107copies.

2.ProkaryoteshavemostlyuniquesequenceDNA,with
repeatsonlyofsequenceslikerRNAsandtRNAs.
Eukaryoteshaveamixofuniqueandrepetitivesequences.
3.UniquesequenceDNAincludesmostofthegenesthat
encodeproteins,aswellasotherchromosomalregions.
HumanDNAcontainsabout65%uniquesequences.

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4.RepetitivesequenceDNAincludesthemoderatelyandhighlyrepeated
sequences.Theymaybedispersedthroughoutthegenome,orclustered
intandemrepeats.
5.Dispersedrepetitivesequencesoccurinfamiliesthathavea
characteristicsequence.Oftenthesamefewsequencesarehighly
repeated,andcomprisemostofthedispersedrepeatsinthegenome.
Littleisknownoftheirfunction,orindeedwhethertheyactuallyservea
function.Therearetwotypesofinterspersionpatternsfoundinall
eukaryoticorganisms:
a.SINEs(shortinterspersedrepeatedsequences)with100500bp
sequences.AnexampleistheAlurepeatsfoundinsomeprimates,
includinghumans,wherethese200300bprepeatsmakeup9%ofthe
genome.
b.LINEs(longinterspersedrepeatedsequences)withsequencesof5kbor
more.ThecommonexampleinmammalsisLINE1,withsequencesup
to7kbinlength.

6.Tandemlyrepetitivesequencesarecommonineukaryoticgenomes,
rangingfromveryshort(110bp)sequencestogenesandevenlonger
sequences.Thisgroupincludescentromereandtelomeresequences,and
rRNAandtRNAgenes.

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