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Chapter 16

he !olecular Basis o" Inheritance


PowerPoint Lectures for Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero


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Overview: Lifes Operating Instructions In 1953, James Watson an !rancis "ric# s$oo# t$e wor%
# Wit$ an e%egant ou&%e'$e%ica% mo e% for t$e structure of eo()ri&onuc%eic aci , or *+,

Figure 16.1
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*+,, t$e su&stance of in$eritance


# Is t$e most ce%e&rate mo%ecu%e of our time

-ere itar) information


# Is enco e in t$e c$emica% %anguage of *+, an repro uce in a%% t$e ce%%s of )our &o )

It is t$e *+, program


# .$at irects t$e eve%opment of man) ifferent t)pes of traits
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"oncept 1/01: *+, is t$e genetic materia% 1ar%) in t$e 23t$ centur)
# .$e i entification of t$e mo%ecu%es of in$eritance %oome as a ma4or c$a%%enge to &io%ogists

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The Search for the Genetic Material: Scientific Inquiry

.$e ro%e of *+, in $ere it)


# Was first wor#e out &) stu )ing &acteria an t$e viruses t$at infect t$em

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Evidence That DNA Can Transform acteria !re eric# 5riffit$ was stu )ing Streptococcus pneumoniae
# , &acterium t$at causes pneumonia in mamma%s

-e wor#e wit$ two strains of t$e &acterium


# , pat$ogenic strain an a nonpat$ogenic strain

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5riffit$ foun t$at w$en $e mi(e $eat'#i%%e remains of t$e pat$ogenic strain
# Wit$ %iving ce%%s of t$e nonpat$ogenic strain, some of t$ese %iving ce%%s &ecame pat$ogenic
EXPERIMENT 6acteria of t$e 789 :smoot$; strain of Streptococcus pneumoniae are pat$ogenic &ecause t$e) $ave a capsu%e t$at protects t$em from an anima%s efense s)stem0 6acteria of t$e 7<9 :roug$; strain %ac# a capsu%e an are nonpat$ogenic0 !re eric# 5riffit$ in4ecte mice wit$ t$e two strains as s$own &e%ow: Living 8 :contro%; ce%%s Living < -eat'#i%%e :contro%; ce%%s :contro%; 8 ce%%s =i(ture of $eat'#i%%e 8 ce%%s an %iving < ce%%s

RESULTS Mouse dies Mouse health Mouse health Mouse dies

Living 8 ce%%s are foun in &%oo samp%e0

Figure 16.!

CONCLUSION 5riffit$ conc%u e t$at t$e %iving < &acteria $a &een transforme into pat$ogenic 8 &acteria &) an un#nown, $erita&%e su&stance from t$e ea 8 ce%%s0

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5riffit$ ca%%e t$e p$enomenon transformation


# +ow efine as a c$ange in genot)pe an p$enot)pe ue to t$e assimi%ation of e(terna% *+, &) a ce%%

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Evidence That !iral DNA Can "ro#ram Cells , itiona% evi ence for *+, as t$e genetic materia%
# "ame from stu ies of a virus t$at infects &acteria

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>iruses t$at infect &acteria, &acteriop$ages


# ,re wi e%) use as too%s &) researc$ers in mo%ecu%ar genetics
Phage head

Tail Tail #i$er %N& 133 nm

Figure 16."

'a(terial (ell

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,%fre -ers$e) an =art$a "$ase


# Performe e(periments s$owing t$at *+, is t$e genetic materia% of a p$age #nown as .2

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.$e -ers$e) an "$ase e(periment


EXPERIMENT In t$eir famous 1952 e(periment, ,%fre -ers$e) an =art$a "$ase use ra ioactive su%fur an p$osp$orus to trace t$e fates of t$e protein an *+,, respective%), of .2 p$ages t$at infecte &acteria% ce%%s0 1 =i(e ra ioactive%) %a&e%e p$ages wit$ &acteria0 .$e p$ages infecte t$e &acteria% ce%%s0 P$age 6acteria% ce%% 'at(h 1) P$ages were grown wit$ ra ioactive su%fur :35S;, w$ic$ was incorporate into p$age protein :pin#;0 *+, P$age *+, "entrifuge <a ioactive *+, Pe%%et :&acteria% ce%%s an contents; ! ,gitate in a &%en er to " "entrifuge t$e mi(ture separate p$ages outsi e so t$at &acteria forme t$e &acteria from t$e a pe%%et at t$e &ottom of &acteria% ce%%s0 t$e test tu&e0 <a ioactive 1mpt) protein protein s$e%% * =easure t$e ra ioactivit) in t$e pe%%et an t$e %i?ui <a ioactivit) :p$age protein; in %i?ui

'at(h !) P$ages were grown wit$ ra ioactive p$osp$orus :32P;, w$ic$ was incorporate into p$age *+, :&%ue;0

<a ioactivit) :p$age *+,; Pe%%et in pe%%et RESULTS P$age proteins remaine outsi e t$e &acteria% ce%%s uring infection, w$i%e p$age *+, entere t$e ce%%s0 W$en cu%ture , &acteria% ce%%s wit$ ra ioactive p$age *+, re%ease new p$ages wit$ some ra ioactive p$osp$orus0 CONCLUSION -ers$e) an "$ase conc%u e t$at *+,, not protein, functions as t$e .2 p$ages genetic materia%0

"entrifuge

Figure 16.*

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Additional Evidence That DNA Is the $enetic %ateria Prior to t$e 1953s, it was a%rea ) #nown t$at *+, # Is a po%)mer of nuc%eoti es, eac$ consisting of t$ree components: a nitrogenous &ase, a sugar, an a p$osp$ate group
Sugar+phosphate $a(,$o-e 5 en O@ 5 O P O "-2 O 1 O @ A 2 3 O O P O "-2 O O@ O O P O "-2 O O@ O 5 O P O "-2 O 1 O@ A P$osp$ate3 2 O- 8ugar : eo()ri&ose; 3 en + + Nitroge-ous $ases "-3 O O .$)mine :.; + + + + +

, enine :,; - + + + O ")tosine :"; + + +

O + *+, nuc%eoti e

Figure 16..

+ 5uanine :5;

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1rwin "$argaff ana%)Be t$e &ase composition of *+, # !rom a num&er of ifferent organisms In 19AC, "$argaff reporte # .$at *+, composition varies from one species to t$e ne(t .$is evi ence of mo%ecu%ar iversit) among species # =a e *+, a more cre i&%e can i ate for t$e genetic materia%

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Building a Structural Model of DNA: Scientific Inquiry

Once most &io%ogists were convince t$at *+, was t$e genetic materia%
# .$e c$a%%enge was to etermine $ow t$e structure of *+, cou% account for its ro%e in in$eritance

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=aurice Wi%#ins an <osa%in !ran#%in


# Were using a tec$ni?ue ca%%e D'ra) cr)sta%%ograp$) to stu ) mo%ecu%ar structure

<osa%in !ran#%in
# Pro uce a picture of t$e *+, mo%ecu%e using t$is tec$ni?ue

Figure 16.6 a2 $

/a0 Rosali-d Fra-,li-

/$0 Fra-,li-1s X+ra di##ra(tioPhotograph o# %N&

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Watson an "ric# e uce t$at *+, was a ou&%e $e%i(


# .$roug$ o&servations of t$e D'ra) cr)sta%%ograp$ic images of *+,
5 , . 1 nm 5 " , " 5 5 . " 30A nm " . ,

. .

, , , , . .

5 ,

" .

303A nm /(0 Spa(e+#illi-g 5odel

Figure 16.3a2 (

/a0 4e #eatures o# %N& stru(ture

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!ran#%in $a conc%u e t$at *+,


# Was compose of two antipara%%e% sugar' p$osp$ate &ac#&ones, wit$ t$e nitrogenous &ases paire in t$e mo%ecu%es interior

.$e nitrogenous &ases


# ,re paire in specific com&inations: a enine wit$ t$)mine, an c)tosine wit$ guanine

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5 en O P
@

OO O .

-) rogen &on 3 en O, O "-2 O O O 5 " O "-2 O O@ O P O O@

O
@

P O

-2" O
@

O O -2 " O

O " 5 O "-2 O

O P
@

O O -2 " O

O , . O "-2 O O

P O

O@

O3 en

P O

O@

Figure 16.3$

/$0 Partial (he5i(al stru(ture

5 en

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Watson an "ric# reasone t$at t$ere must &e a itiona% specificit) of pairing
# *ictate &) t$e structure of t$e &ases

1ac$ &ase pair forms a ifferent num&er of $) rogen &on s


# , enine an t$)mine form two &on s, c)tosine an guanine form t$ree &on s

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+ + O "-3

+ 8ugar +

+ + O 8ugar Th 5i-e /T0 -

&de-i-e /&0

+ 8ugar +

+ +

+ Figure 16.7 6ua-i-e /60

8ugar C tosi-e /C0

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"oncept 1/02: =an) proteins wor# toget$er in *+, rep%ication an repair .$e re%ations$ip &etween structure an function
# Is manifest in t$e ou&%e $e%i(

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The Basic rinci!le: Base airing to a Tem!late Strand

8ince t$e two stran s of *+, are comp%ementar)


# 1ac$ stran acts as a temp%ate for &ui% ing a new stran in rep%ication

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In *+, rep%ication
# .$e parent mo%ecu%e unwin s, an two new aug$ter stran s are &ui%t &ase on &ase' pairing ru%es
& C T & 6 T 6 & T C & C T & 6 T 6 & T C & C T & 6 & T C 6
/a0 .$e parent mo%ecu%e $as two comp%ementar) stran s of *+,0 1ac$ &ase is paire &) $) rogen &on ing wit$ its specific partner, , wit$ . an 5 wit$ "0 /(0 1ac$ parenta% stran now serves as a temp%ate t$at etermines t$e or er of nuc%eoti es a%ong a new, comp%ementar) stran 0 /d0 .$e nuc%eoti es are connecte to form t$e sugar'p$osp$ate &ac#&ones of t$e new stran s0 1ac$ 7 aug$ter9 *+, mo%ecu%e consists of one parenta% stran an one new stran 0

T 6

& C T &

T 6 & T C

& C T & 6

T 6 & T C

& C T & 6

T 6 & T C

/$0 .$e first step in rep%ication is separation of t$e two *+, stran s0

Figure 16.8 a9d


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*+, rep%ication is semiconservative


# 1ac$ of t$e two new aug$ter mo%ecu%es wi%% $ave one o% stran , erive from t$e parent mo%ecu%e, an one new%) ma e stran
First Se(o-d Pare-t (ell repli(atio- repli(atio/a0 Co-ser;ati;e 5odel. .$e two parenta% stran s reassociate after acting as temp%ates for new stran s, t$us restoring t$e parenta% ou&%e $e%i(0 /$0 Se5i(o-ser;ati;e 5odel. .$e two stran s of t$e parenta% mo%ecu%e separate, an eac$ functions as a temp%ate for s)nt$esis of a new, comp%e' mentar) stran 0 /(0 %ispersi;e 5odel. 1ac$ stran of both aug$ter mo%' ecu%es contains a mi(ture of o% an new%) s)nt$esiBe *+,0

Figure 16.1: a9(

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1(periments performe &) =ese%son an 8ta$%


# 8upporte t$e semiconservative mo e% of *+, rep%ication
EXPERIMENT =att$ew =ese%son an !ran#%in 8ta$% cu%ture E. coli &acteria for severa% generations on a me ium containing nuc%eoti e precursors %a&e%e wit$ a $eav) isotope of nitrogen, 15+0 .$e &acteria incorporate t$e $eav) nitrogen into t$eir *+,0 .$e scientists t$en transferre t$e &acteria to a me ium wit$ on%) 1A+, t$e %ig$ter, more common isotope of nitrogen0 ,n) new *+, t$at t$e &acteria s)nt$esiBe wou% &e %ig$ter t$an t$e parenta% *+, ma e in t$e 15+ me ium0 =ese%son an 8ta$% cou% istinguis$ *+, of ifferent ensities &) centrifuging *+, e(tracte from t$e &acteria0
1

6acteria cu%ture in me ium containing 15 +

6acteria transferre to me ium containing 1A +

RESULTS
"

*+, samp%e centrifuge after 23 min :after first rep%ication;

*+, samp%e centrifuge after A3 min :after secon rep%ication;

Less ense =ore ense

.$e &an s in t$ese two centrifuge tu&es represent t$e resu%ts of centrifuging two *+, samp%es from t$e f%as# Figure 16.11 in step 2, one samp%e ta#en after 23 minutes an one after A3 minutes0
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CONCLUSION =ese%son an 8ta$% conc%u e t$at *+, rep%ication fo%%ows t$e semiconservative mo e% &) comparing t$eir resu%t to t$e resu%ts pre icte &) eac$ of t$e t$ree mo e%s in !igure 1/0130 .$e first rep%ication in t$e 1A+ me ium pro uce a &an of $)&ri :15+@1A+; *+,0 .$is resu%t e%iminate t$e conservative mo e%0 , secon rep%ication pro uce &ot$ %ig$t an $)&ri *+,, a resu%t t$at e%iminate t$e ispersive mo e% an supporte t$e semiconservative mo e%0 First repli(atioSe(o-d repli(atio-

Co-ser;ati;e 5odel

Se5i(o-ser;ati;e 5odel

%ispersi;e 5odel

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DNA Re!lication: A Closer &oo' .$e cop)ing of *+,


# Is remar#a&%e in its spee an accurac)

=ore t$an a oBen enB)mes an ot$er proteins


# Participate in *+, rep%ication

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$ettin# Started( )ri#ins of Replication .$e rep%ication of a *+, mo%ecu%e


# 6egins at specia% sites ca%%e origins of rep%ication, w$ere t$e two stran s are separate

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, eu#ar)otic c$romosome
# =a) $ave $un re s or even t$ousan s of rep%ication origins
Origin of rep%ication Parenta% :temp%ate; stran *aug$ter :new; stran 3025 Em 1 <ep%ication &egins at specific sites w$ere t$e two parenta% stran s separate an form rep%ication &u&&%es0 ! .$e &u&&%es e(pan %atera%%), as *+, rep%ication procee s in &ot$ irections0 " 1ventua%%), t$e rep%ication &u&&%es fuse, an s)nt$esis of t$e aug$ter stran s is comp%ete0

6u&&%e

<ep%ication for#

.wo aug$ter *+, mo%ecu%es

/a0 In eu#ar)otes, *+, rep%ication &egins at man) sites a%ong t$e giant *+, mo%ecu%e of eac$ c$romosome0

Figure 16.1! a2 $
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/$0 In t$is micrograp$, t$ree rep%ication &u&&%es are visi&%e a%ong t$e *+, of a cu%ture "$inese $amster ce%% :.1=;0

Elon#atin# a Ne* DNA Strand 1%ongation of new *+, at a rep%ication for#


# Is cata%)Be &) enB)mes ca%%e *+, po%)merases, w$ic$ a nuc%eoti es to t$e 3 en of a growing stran
Ne< stra-d 5 en 8ugar P$osp$ate Te5plate stra-d 3 en 5 en 3 en & 6ase C 6 T 6 C &
P P P

& C 6 T OP)rop$osp$ate 3 en
P P

T 6 C & C

O-

Figure 16.1"

+uc%eosi e trip$osp$ate

2 P 5 en 5 en

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Antiparallel Elon#ation -ow oes t$e antipara%%e% structure of t$e ou&%e $e%i( affect rep%icationF

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*+, po%)merases a

nuc%eoti es

# On%) to t$e free 3 en of a growing stran

,%ong one temp%ate stran of *+,, t$e %ea ing stran


# *+, po%)merase III can s)nt$esiBe a comp%ementar) stran continuous%), moving towar t$e rep%ication for#

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.o e%ongate t$e ot$er new stran of *+,, t$e %agging stran


# *+, po%)merase III must wor# in t$e irection awa) from t$e rep%ication for#

.$e %agging stran


# Is s)nt$esiBe as a series of segments ca%%e O#aBa#i fragments, w$ic$ are t$en 4oine toget$er &) *+, %igase

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8)nt$esis of %ea ing an %agging stran s uring *+, rep%ication


1 *+, po% I%% e%ongates *+, stran s on%) in t$e 5 3 irection0 3 5 Pare-tal %N& O#aBa#i fragments 2 1 3 5 5 3 ! One new stran , t$e %ea ing stran , can e%ongate continuous%) 5 3 as t$e rep%ication for# progresses0 " .$e ot$er new stran , t$e %agging stran must grow in an overa%% 3 5 irection &) a ition of s$ort segments, O#aBa#i fragments, t$at grow 5 3 :num&ere $ere in t$e or er t$e) were ma e;0

*+, po% III .emp%ate stran

Lea ing stran Lagging stran 2 1

* *+, %igase 4oins O#aBa#i fragments &) forming a &on &etween t$eir free en s0 .$is resu%ts in a continuous stran 0

.emp%ate stran

*+, %igase

Figure 16.1*

Overa%% irection of rep%ication

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"rimin# DNA Synthesis *+, po%)merases cannot initiate t$e s)nt$esis of a po%)nuc%eoti e
# .$e) can on%) a nuc%eoti es to t$e 3 en

.$e initia% nuc%eoti e stran


# Is an <+, or *+, primer

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On%) one primer is nee e for s)nt$esis of t$e %ea ing stran
# 6ut for s)nt$esis of t$e %agging stran , eac$ O#aBa#i fragment must &e prime separate%)

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1 Primase 4oins <+, nuc%eoti es


into a primer0

3 5 3

.emp%ate stran
3

! *+, po% III a <+, primer


5
1

s *+, nuc%eoti es to t$e primer, forming an O#aBa#i fragment0


5

3 ,fter reac$ing t$e ne(t


<+, primer :not s$own;, *+, po% III fa%%s off0
3 5
O#aBa#i fragment 1

3 5

,fter t$e secon fragment is prime 0 *+, po% III a s *+, nuc%eoti es unti% it reac$es t$e first primer an fa%%s off0 5
3
2 1

3 5

. *+, po% 1 rep%aces t$e


<+, wit$ *+,, a ing to t$e 3 en of fragment 20
3 5
2 1

6 *+, %igase forms a &on


&etween t$e newest *+, an t$e a 4acent *+, of fragment 10 5
3
2 1

3 .$e %agging stran

in t$is region is now comp%ete0


5

Figure 16.1.

Overa%% irection of rep%ication

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)ther "roteins That Assist DNA Replication -e%icase, topoisomerase, sing%e'stran &in ing protein
# ,re a%% proteins t$at assist *+, rep%ication

Ta$le 16.1
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, summar) of *+, rep%ication


Overa%% irection of rep%ication

1 -e%icase unwin s t$e parenta% ou&%e $e%i(0


! =o%ecu%es of sing%e' stran &in ing protein sta&i%iBe t$e unwoun temp%ate stran s0 " .$e %ea ing stran is s)nt$esiBe continuous%) in t$e 5 3 irection &) *+, po% III0

Lagging Lea ing stran Origin of rep%ication stran

*+, po% III

Lagging stran Lea ing stran

O>1<>I1W

Lea ing stran

5 3 Parenta% *+, * Primase &egins s)nt$esis of <+, primer for fift$ O#aBa#i fragment0

<ep%ication for# Primase *+, po% III Primer A

*+, po% I Lagging stran 3 2

*+, %igase 1 3 5

. *+, po% III is comp%eting s)nt$esis of t$e fourt$ fragment, w$en it reac$es t$e <+, primer on t$e t$ir fragment, it wi%% issociate, move to t$e rep%ication for#, an a *+, nuc%eoti es to t$e 3 en of t$e fift$ fragment primer0

Figure 16.16

6 *+, po% I removes t$e primer from t$e 5 en of t$e secon fragment, rep%acing it wit$ *+, nuc%eoti es t$at it a s one &) one to t$e 3 en of t$e t$ir fragment0 .$e rep%acement of t$e %ast <+, nuc%eoti e wit$ *+, %eaves t$e sugar' p$osp$ate &ac#&one wit$ a free 3 en 0

3 *+, %igase &on s t$e 3 en of t$e secon fragment to t$e 5 en of t$e first fragment0

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The DNA Replication %achine as a Stationary Comple+

.$e various proteins t$at participate in *+, rep%ication


# !orm a sing%e %arge comp%e(, a *+, rep%ication 7mac$ine9

.$e *+, rep%ication mac$ine


# Is pro&a&%) stationar) uring t$e rep%ication process

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roofreading and Re!airing DNA *+, po%)merases proofrea new%) ma e *+,


# <ep%acing an) incorrect nuc%eoti es

In mismatc$ repair of *+,


# <epair enB)mes correct errors in &ase pairing

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In nuc%eoti e e(cision repair


# 1nB)mes cut out an rep%ace amage stretc$es of *+,
1 , t$)mine imer istorts t$e *+, mo%ecu%e0 ! , nuc%ease enB)me cuts t$e amage *+, stran at two points an t$e amage section is remove 0 +uc%ease

*+, po%)merase

" <epair s)nt$esis &) a *+, po%)merase fi%%s in t$e missing nuc%eoti es0 *+, %igase

Figure 16.13
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* *+, %igase sea%s t$e !ree en of t$e new *+, .o t$e o% *+,, ma#ing t$e stran comp%ete0

Re!licating the "nds of DNA Molecules .$e en s of eu#ar)otic c$romosoma% *+,


# 5et s$orter wit$ eac$ roun of rep%ication
1n of parenta% *+, stran s 5 Lea ing stran Lagging stran 3 Last fragment <+, primer Lagging stran 5 <emova% of primers an rep%acement wit$ *+, w$ere a 3 en is avai%a&%e 5 8econ roun of rep%ication 5 +ew %ea ing stran 3 +ew %agging stran 5 3 Primer remove &ut cannot &e rep%ace wit$ *+, &ecause no 3 en avai%a&%e for *+, po%)merase 3 Previous fragment

3
!urt$er roun s of rep%ication
8$orter an s$orter aug$ter mo%ecu%es

Figure 16.17

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1u#ar)otic c$romosoma% *+, mo%ecu%es


# -ave at t$eir en s nuc%eoti e se?uences, ca%%e te%omeres, t$at postpone t$e erosion of genes near t$e en s of *+, mo%ecu%es

Figure 16.18
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1 Em

If t$e c$romosomes of germ ce%%s &ecame s$orter in ever) ce%% c)c%e


# 1ssentia% genes wou% eventua%%) &e missing from t$e gametes t$e) pro uce

,n enB)me ca%%e te%omerase


# "ata%)Bes t$e %engt$ening of te%omeres in germ ce%%s

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