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    The Nucleus

      The nucleus is the headquarters of theThe nucleus is the headquarters of thecell. Itcell. It is the most obvious organelle in

    any eukaryotic cell and appears as aa

    large dark spot in EUKA!"TI# cells. Itlarge dark spot in EUKA!"TI# cells. Itcontrols all cell activity.controls all cell activity.

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    • The Nucleus is a membrane$enclosed organelle %hich house

    most of the genetic information

    and regulatory machineryresponsible for providing the cell

    %ith its unique characteristics. 

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     Nucleus is the most important organelle in cell.

    In mammalian cells& e'cepting (#&

    all cells else are the nucleus contained cells.

    In prokaryotic cells& there is no membraneto package the nucleic acid substance& so&

    %e call this nucleic substance enriched

    area as )Nucleoid*.

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     The ma+or structures of nucleus include,

    ① nuclear envelope.

    ② nucleolus.

    ③ nuclear matri'.

    ④ chromatin.

    ⑤ nuclear lamina.

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    The ma+or functions of nucleus,

    ① inheritance, maintain the geneticcontinuity of generation by the

    replication of -NA chromatin and the

    proliferation of cell.② development, regulate the cell

    differentiation by the regulation of

    spatiotemporal sequence of genee'pression.

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    • TE NU#/EU0,• 1UN#TI"N0

    • It stores the cell2s hereditary material& or -NA.

    • 0ite of -NA replication

    • 0ite of -NA transcription to mNA

    • ibosomal formation

     – Nucleolus, NA 3 protein required for ribosomal

    synthesis

    • It coordinates the cell2s activities& %hich include

    gro%th& intermediary metabolism& protein synthesis&and reproduction 4cell division5 by regulating gene

    e'pression.

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    http://upload.wikimedia.org/wikipedia/commons/3/38/Diagram_human_cell_nucleus.svg

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    nuclearpores

    chromatin

    nucleolus

    nuclearenvelope

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    nuclearpores

    nucleus

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    I. Nuclear envelope 4Nuclear membrane5

    Nuclear envelope is the lipid bilayer that packages

    the nucleus.

    Nuclear envelope separates the -NA from cell plasma

    and forms a stable inner environment to,

    ①  protect the -NA from damage&

      ② separate the replication of -NA from the

    translation of NA spatiotemporally&

    ③ the chromatin is anchored on to the nuclear

    envelope& that is beneficial to be despiraled&

    replicated& condensed& and distributed into ne%nuclei equally&

    ④ the pores on the envelope are the channels for

    the substance e'change.

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    Nuclear envelope is bilayer membrane,

     Nuclear envelope is composed of 

      inner nuclear membrane&

    outer nuclear membrane& andperinuclear space.

    There are nuclear pores on the membrane

    that are linked %ith plasma.

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     • The inner surface of the NE is bound to

    a thin filamentous net%ork 4lamins

     polypeptides5 called the nuclearlamina. It provides mechanical support

    to the NE and seeves as sites for

    attachment for chromatin fibers.

    • 9utations in the lamin genes are

    responsible for several distinct human

    diseases 4e.g. a rare form of muscular

    dystrophy5.

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    The nuclear pores are the channels for the

    substance transportation,

    •Nuclear proteins are synthesi:ed in plasma& then

    %ill be imported into nucleus by the pores.

    •The NAs and the ribosome subunits synthesi:ed

    in nucleus %ill be e'ported into plasma by the

    pores also.

    •In addition& it is indicated by a in+ection

    e'periment that small molecules can enter the

    nucleus by diffusion from the pores.

    TE NU#/EA ;"E

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    Nuclear pores are composed of 87 different

    nucleoporins at least& and %e call these pore

    structure as nuclear pore comple' 4N;#5.

    Usually& a mammalian nucleus contains

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    The structures of nuclear pore include

    ① cytoplasmic ring located on the cell

    plasma part of the pore comple'contains filaments e'tending into

    plasma.

    ② nuclear ring located on the nuclear

    plasma part of the pore comple'

    e'tending filaments also.

    ③ transporter located in center of the

    pore as a plug particle.④ 0poke located on the edge of the pore

    as the spines.

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    • 

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    The nuclear pore structures on the cell plasma side after an e'traction

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    The nuclear pore structures on the nuclear plasma side after an e'traction

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    Nuclear plasma protein 4nucleoplasmin5 is

    transported by the follo%ing steps,

      ① The protein combines to the B C D dimer ofthe receptor 4imporin5.

    ② The comple' of the protein transported

    and the receptor used combines to thefilaments located on the N;# cytoplasmic

    ring.

    ③ The filaments curve to the nuclear center&

    the transporter structure %ill be changedto form a hydrophilic channel& and the

    protein passes through the channel.

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     ④ The comple' of the protein transported

    and the receptor used combines to an$

      T;& the comple' is disassembled andreleases out the protein transported.

    ⑤ The imporin D combined %ith an$T; %ill

    be e'ported out of the nucleus& the T;

    combined %ith an %ill be hydroly:ed incell plasma& and the an$-; %ill go back

    to nucleus to be transformed to an$T;

    again.

    ⑥ The imporin B %ill be transported back to

    cell plasma %ith the help from e'portin.

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    Fe kno% a little about ho% the macromolecules are

    transported to cell plasma from nucleus.

    In most of cases& the NA in nucleus iscombined %ith protein to form an N; comple'&

    then& transported into cell plasma.

    There is nuclear e'portation signal 4NE05 on the

    protein of N; comple' that can combine to the

    intracellular receptor& e'portin& to form the

    comple' of N;$e'portin$an$T;.

    In the cell plasma& this comple' %ill bedisassembled and release out the an$T;& NA&

    an$-;& e'portin& and N; protein.

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    The nucleoplasmin is transported into nucleus

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     #hromosome

    #hromatin %as named byF. 1lemming in 6=G.

    #hromosome %as named by

    Faldeyer in 6.

    #hromatin and chromosome are

    same substance %ith different shape

    presentation in different cell cyclephases.

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    The chemical components of chromatin,

      -NA&

    histone protein&

    nonhistone protein& andsome NA

    at ratio about 6,6,46$6.85,7.78.

    -NA,

      -NA is the carrier of genetic information.

    -NA sequences can be sorted as < types, 

    nonrepeated fraction&

    moderately repeated fraction 4676$6785& and

    highly repeated fraction 4H6785.

    -NA forms,

    ($-NA&

    $-NA& and

    A$-NA.

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    -NA forms 4ed color sho%s the couple backbones5

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     #hromosome -NA contains three basic sequences,

    ① autonomously replicating -NA sequence 4A05.

    A0 is the starting site of -NA replication.In yeast genome& there are J77$77 A0s

    included& and most of them contain a AT

    enriched 66bp sequence called as A0

    consensus sequence 4A#05.

    ② centromere -NA sequence 4#EN5 composed of

    a lot of repeated sequences.

    ③ telomere -NA sequence 4TE/5.

    TE/ is similar in different bio organisms&

    and composed of 8 L 67bp repeated

    sequences. uman

    TE/ repeated sequence is TTA.

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    In 6G

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    Three basic sequences of chromosome

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    I0T"NE

    istone is positively charged and contains

    arginine and lycine.istone is alkaline protein.

    istones can be sorted as t%o types,

    6. ighly conserved core histone including JA&

    J(&

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    The reasons for that may be as the follo%s,

      6. 9ost of the amino acids of core histone

    interact %ith -NA or other histones& so&any change of them %ill cause the fatal

    mutation.

      J. In all bio organisms& the -NA

    phosphodiester skeleton that interacts%ith histone is same.

    6 is easy to be mutated&

    and it is species specific and tissue

    specific.

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    N"NI0T"NE ;"TEIN,

    Nonhistone protein is

    the protein that binds to the specific -NA sequence

    of chromosome& so& %e call it as sequence specific

    -NA binding protein.

    The features of nonhistone protein are as the follo%s,

     ① Nonhistone protein is negatively charged andacidic protein that contains a large number of

    aspartic acids and glutamic acids.

    ② Nonhistone protein can be synthesi:ed during

    the %hole cell cycle& but histone protein issynthesi:ed during the 0 phase only.

    ③ Nonhistone protein can recogni:e the specific

    -NA sequence.

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    The functions of nonhistone are as the

    follo%s,① elp -NA molecules to be pleated

    and form different structure

    domains that are beneficial to-NA replication and gene

    transcription.

    ②elp to start -NA replicationreaction.

    ③ egulate transcription and gene

    e'pression.

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    1"9 -NA T" #"9"0"9E,

    There are J< pairs of chromosomes in a human

    nucleus.• If you open and e'tend the -NA molecule in

    each chromosome& it %ill be 8 cm long.• If you link all -NA molecules in a nucleus

    together& it %ill be 6.= L J.7 m long.(ut& the diameter of nucleus is shorter

    than 67Mm.

    The primary structure formed by the po%erfulcompaction is called as nucleosome.

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    Nucleosome,

    . Kornberg figured out the model of nucleosome.

    Nucleosome is a beaded structure composed of core

    particles and linker -NA.

    Fe can describe the structure as the follo%s,

    ① Each nucleosome includes about J77bp -NA& one

    histone core& and an 6.② The octameric histone core is composed of

    molecules from JA& J(&

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    410tructures of nucleosome

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    #hromatin -NA filament,

      The -NA is compacted to be shortened by =

    folds and forms the -NA filament in 66nmdiameter %hen it %as transformed to the beaded

    nucleosome chain.

    #hromatin -NA e'ists in another style by that the

    beaded nucleosome chain is condensed by ?

    folds.

    Under electron microscope& %e can see the

    chromatin -NA filament in

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    The -NA filaments in

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      For the advanced package of the chromosome, e keep deta!" #nknonso far$ %ro&a&"', !t !s the ser!a" over"apped or p"eated "!ke the fo""os(

    From )*+ to hromosome(

    )*+ 11nm f!"ament -&eaded n#c"eosome cha!n. 30nm f!"ament

    p"eat as "oop cha!n &!nd to the s!tes on n#c"ear ske"eton here !s +/

    enr!ched assem&"' of chromosome

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    Assembly of chromosome

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    eterochromatin and euchromatin,

      n the !nter phase -1 and 2. of ce"" c'c"e, the chromat!n !n the n#c"e#s

    can &e sorted as heterochromat!n and e#chromat!n$

    #chromat!n !s the )*+ reg!ons here the transcr!pt!on !s ver' act!ve$

    #chromat!n "ooks "!ke "oose "oop and &r!ght sta!n!ng #nder e"ectron

    m!croscope$ #chromat!n !s eas' to &e c"eaved &' n#c"ease at some

    h'persens!t!ve s!tes$

    eterochromat!n !s condensed !n phase !tho#t an' transcr!pt!on, so, !t

    as named as !nact!ve chromat!n$ eterochromat!n !s the genet!c "a' reg!ons,

    and rep"!cated "ate"', condensed ear"', that !s ca""ed as heteropyknosis$

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    eterochromatin

    4dark staining5

    and euchromatin

    4bright staining5

    eterochromatin

    Euchromatin

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    #onstitutive

    hetero$

    chromatin !s

    hetero

    p'knosedchromat!n !n

    each t'pe of

    ce"" and

    "ocated !n

    centromere

    reg!on$

    /he F!g shos

    'o# the

    onst!t#t!ve

    hetero

    chromat!nd!sp"a'ed &'

    f"#orescence

    h'&r!d!at!on !n

    s!t#$

    1acultative heterochromatin 

    ! h t h t! d

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    !s heterochromat!n appeared

    !n some spec!a" ce"" t'pe or

    deve"op!ng stage$ /he

    chromosome of fema"e

    mamma"!ans !s the fac#"tat!ve

    heterochromat!n$ s#a""',

    fema"e mamma"!an ce""

    conta!ns do#&"e

    chromosomes, and one of

    them !s heterochromat!n

    ca""ed &arr &od'$ hen ah#man em&r'o !s deve"oped

    after 16 da's, one

    chromosome !"" &e

    transformed as &arr &od' !th

    dark sta!n!ng$ o, e can

    !dent!f' the se: of a h#manem&r'o &' check!ng the &arr

    &od' of the em&r'o ce""s !n the

    amn!ot!c f"#!d$

    The barr body like a drumstick in a %hite cell

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    The structure of chromosome,

      n the ; phase of ce"" c'c"e, chromat!n !"" &e transformed as chromosomes

    &' the poerf#" condensat!on$ hromosomes are st!ck shape !th d!fferent "ength$

    /he metaphase chromosome !s the &est stage to o&serve and n#m&er them&eca#se the morpho"og' of chromosome !s sta&"e at th!s t!me$

    /he n#m&er of chromosome !s same !n the same t'pe of ce""s from d!fferent

    !nd!v!d#a"s of one spec!es$ /he chromosomes of se: ce""s are hap"o!d, e mark !t

    as n$ /he chromosomes of other ce""s are d!p"o!d, e mark !t as 2n$ /he

    chromosomes of some ce""s of some spec!es are po"'p"o!d, s#ch as, 4n, 6n, and8n$

    /he d!fferent ce""s from same !nd!v!d#a" can &e d!fferent chromosome t'pes$

    For e:amp"e, &od' ce""s of rat are 2n, t !ts "!ver ce""s can &e 4n, 8n, and 16n$ /he

    chromosome n#m&er of h#man endometr!a" ce"" !s var!a&"e from 2n

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    The terms used to the structure of chromosome,

    6. #hromatid, ;etaphase chromosome !s composed of to chromat!ds !th a

     =#nct!on at the centromere s!te$ ach chromat!d !s formed &' the over"apped and

    p"eated )*+ do#&"e strands$ hen the ce"" !s d!v!d!ng the chromat!ds can &e

    separated !nto to ne ce""s$J. #hromonema, n the or phase ce""s, each chromonema !nd!cates a

    chromat!d$

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    The terms of chromosome structure

    ? N l l i i i 4N" 5 /h th h th

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    ?. Nucleolar organi:ing regions 4N"s5, /he' are the areas here the genes

    for r!&osome >*+ are "ocated$ /he' can s'nthes!e the 28, 18, and 5$8 r>*+

    for r!&osome$ *?>s can e:!st !n secondar' constr!ct!on$

    =.

    0atellite, t !s a &a"" part "ocated at the term!na" of chromosome, and "!nked tothe ma!n part of chromosome &' secondar' constr!ct!on$ /he sate""!te "ocated at

    term!na" of chromosome !s ca""ed as term!na" sate""!te, and "ocated &eteen to

    secondar' constr!ct!ons !s ca""ed as !ntermed!ate sate""!te$

    . Telomere, t !s the spec!a"!ed part "ocated at the term!na" of chromosome$

    /he f#nct!on of te"omere !s ma!ntenance of the sta&!"!t' of chromosome$ /e"omere!s composed of the h!gh"' repeated fract!ons, and !t !s so conserved that !t !s

    s!m!"ar &eteen the tota""' d!fferent "!fe &e!ngs$ The component of human

    telomere is TTA$ /e"omere !s assoc!ated !th ag!ng$ +fter each rep"!cat!on of

    te"omere )*+, the te"omere !"" &e shortened &' 50 – 100&p$ /he rep"!cat!on of

    te"omere !s droved &' te"omerase that has reverse transcr!ptase act!v!t'$ /h!s

    en'me "acks !n norma" ce""s, so, te"omere !"" &ecome short !th the ce""pro"!ferat!on$ o, ce"" !"" &e ag!ng d#r!ng th!s act!on$

    III N l l

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    III. Nucleolus

      *#c"eo"#s ma' &e v!s!&"e !n phase n#c"e#s$ /he' are spher!ca" and 1 – 2 for

    each ce"" #s#a""'$ /he n#m&er and s!e of n#c"eo"#s are depended on the ce"" t'pe

    and f#nct!on$ /he more prote!ns s'nthes!s and the faster pro"!ferat!on the ce""

    takes, the more and &!gger n#c"eo"! the ce"" has$ *#c"eo"#s d!sappears &efore the

    ce"" d!v!s!on, and appears !n the end of d!v!s!on$ /he ma=or f#nct!ons of n#c"eo"#s

    are r>*+ transcr!pt!on and r!&osome assem&"'$

    0tructure of nucleolus,

    *o an' mem&rane packages n#c"eo"#s area$ /here are three spec!a" areas

    can &e !dent!f!ed #nder e"ectron m!croscope( f!&r!""ar centers -F. that are①s#rro#nded &' dense f!&ers, and "o e"ectr!c dens!t'$ F conta!ns >*+ po"'merase

    and r)*+ that !s naked mo"ec#"e$ dense f!&r!""ar component -)F. that !s a "oop②

    or ha"f "oop to s#rro#nd F$ /ranscr!pt!on !s carr!ed o#t !n the &order reg!on of F

    and )F$ gran#"ar component -. composed of 1520nm part!c"es that are③

    the >*%s !n d!fferent man#fact#red steps$ >*% means the >*+ com&!ned !th

    prote!n$*#c"eo"#s chromat!ns can &e sorted as to t'pes( heterochromat!n and

    e#chromat!n$ /he n#c"eo"#s heterochromat!n !s a"a's "ocated aro#nd the

    n#c"eo"#s, so e ca"" them as n#c"eo"#s per!phera" chromat!n$ /he n#c"eo"#s

    e#chromat!n !s "ocated !n n#c"eo"#s, and n#c"eo"#s organ!!ng reg!on !n that the

    r)*+ !s "ocated$

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    0tructure of nucleolus

    I ib

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    I. ibosome

      >!&osome !s the man#fact#r!ng shop to s'nthes!e prote!ns$ /here are a&o#t

    20,000 r!&osomes !n an act!ve"' gro!ng &acter!#m$ >!&osome prote!ns are 10@ of

    tota" prote!ns of ce"", and !ts >*+ !s 80@ of ce"" tota" >*+$

    0tructure of ribosome,

      /he rat!os of prote!n and >*+ to r!&osome components are 40@ and 60@$

    /he r!&osome s#n!ts are composed of the com&!nat!on of the prote!n and >*+$

    /he cata"'t!c act!v!t!es needed &' the trans"at!on are presented &' r!&osome

    prote!n, r>*+ and other he"per factors$

    /he r!&osomes can &e sorted as to t'pes$ 70 r!&osome e:!sts !n &acter!a,m!tochondr!on, and ch"orop"ast$ 80 r!&osome e:!sts !n the p"asma of e#kar'ot!c

    ce""s$

    >!&osome !s composed of a "arge s#n!t and a sma"" s#n!t$ /he &oth

    s#n!ts !"" &e com&!ned together hen the r!&osome s'nthes!es prote!n !th

    m>*+ as temp"ate$ +fter the trans"at!on, the r!&osome !"" &e separated as to

    parts aga!n$ hen a prote!n !s trans"ated on an m>*+, man' r!&osomes can &!ndto the m>*+ to s'nthes!e the prote!n$ e ca"" these r!&osomes for one prote!n

    s'nthes!s as polyribosome$ /he "onger m>*+ !s #sed, the more r!&osomes are

    com&!ned$ /he po"'r!&osome enhances the eff!c!enc' of prote!n s'nthes!s$

    %rokar'ot!c 5 r>*+ and e#kar'ot!c 5$8 r>*+ are ver' conserved for the!r

    str#ct#res, so, the' can &e #sed to research the &!oevo"#t!on$

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    Assembly of ribosome,

    /he )*+ fragment encod!ng r>*+ !s ca""ed as r>*+ gene$ /here are a&o#t

    200 cop!es of th!s gene !n a h#man ce""$ r)*+ conta!ns no h!stone core, so, !t !s a

    naked )*+$/o transcr!pt r>*+, the >*+ po"'merase moves ahead a"ong the )*+

    mo"ec#"e$ /he s'nthes!ed r>*+ mo"ec#"es e:tend o#t the!r mo"ec#"es from the

    comp"e: of po"'merase and )*+, and form a feather"!ke str#ct#re #nder

    m!croscope$

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    r>*+ transcr!pt!on

    /he f!"aments are the

    ne s'nthes!ed 45

    r>*+ that com&!nes to

    prote!n to form >*%

    comp"e:$ /hemeth'"ated 45 r>*+

    can &e c"eaved as the

    to parts &' >*ase(

    18 r>*+ and 32

    r>*+, the "atter !s

    c"eaved as 28 r>*+and 5$8 r>*+$ /he

    s'nthes!ed 5 r>*+

    !"" &e transported !nto

    n#c"eo"#s to =o!n the

    assem&"' of the "arge

    s#n!ts of r!&osome$

    There is a ?7bp non$transcription -NA

    fragment bet%een ad+acent rNA genes

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    Assembly of ribosome

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    . Nuclear matri'

      *#c"ear matr!: !s ca""ed as n#c"eoske"eton that !s a meshork !n e#kar'ot!c

    ce""s, that !s hat to"d 'o# &efore$ Beca#se n#c"ear matr!: !s assoc!ated !th

    )*+ rep"!cat!on, >*+ transcr!pt!on and mod!f!cat!on, chromosome assem&"', andv!r#s rep"!cat!on, n#c"ear matr!: !s no pa!d more attent!ons to$

    #omponents of nuclear matri',

      ① *onh!stone f!"aments at rat!o of 96@$ /he n#c"eoske"eton conta!ns three

    scaffo"d prote!ns( , , and $Ⅰ Ⅱ Ⅲ

      ② + "!tt"e >*+ and )*+( /he >*+ !s !mportant to ma!nta!n the ske"eton

    str#ct#re$ /he )*+ !s ca""ed as matr!: Ascaffo"d assoc!ated reg!on -;+> or +>.

    here the +/ !s enr!ched to form the heterochromat!n &!nd!ng s!tes$

      ③ + "!tt"e phospho"!p!ds -1$6@. and s#gars -0$9@.$

    *#c"ear ske"eton – n#c"ear "am!na – !nter f!"aments – pore comp"e: !s a

    meshork s'stem !th ver' good sta&!"!t'$

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    The function of nuclear skeleton,

    1$ %resent the scaffo"ds for )*+ rep"!cat!on$ )*+ can &e anchored on to the

    scaffo"d !th a rep"!cat!on "oop$ /he en'mes needed &' )*+ rep"!cat!on are

    "ocated on the ske"eton, s#ch as )*+ po"'merase C, )*+ pr!merase, )*+

    topo!somerase $

    2$ s the p"ace here gene can &e transcr!pted and mod!f!ed$ /here are >*+

    po"'merase &!nd!ng s!tes on the ske"eton$ *e s'nthes!ed >*+ !s com&!nedto the ske"eton for f#rther mod!f!cat!on$

    3$ s assoc!ated !th the assem&"' of chromosome$ /he n#c"ear ske"eton ma'

    &e same th!ng to chromosome ske"eton$ 30nm chromat!n f!&ers are

    com&!ned to n#c"ear ske"eton to form "oops that !"" &e packaged f#rther !n ;

    phase to &e assem&"ed as chromosome$

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    #hromatin bound on nuclear skeleton or chromosome skeleton