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<title>Chromatin Walkthrough</title>
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<body lang=3DEN-US style=3D'tab-interval:.5in'>

<div class=3DSection1>

<h1><span style=3D'font-family:Tahoma'>A walk through the chromatin structu=
res of
the maize and <span class=3DSpellE>arabidopsis</span> <span class=3DSpellE>=
<span
class=3DGramE>Adh</span></span> genes…<o:p></o:p></span></h1>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt;
font-family:Arial'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt;
font-family:Arial'>The chromatin research in our laboratory is primarily
focused on promoter structures of three plant genes encoding alcohol <span
class=3DGramE>dehydrogenase<span style=3D'mso-spacerun:yes'>  </span>(</spa=
n>maize <i
style=3D'mso-bidi-font-style:normal'>Adh1</i> and <i style=3D'mso-bidi-font=
-style:
normal'>Adh2</i>, and the <span class=3DSpellE><i style=3D'mso-bidi-font-st=
yle:
normal'>Adh</i></span> gene from <span class=3DSpellE>arabidopsis</span>) a=
nd the
genome organization of these plants.<span style=3D'mso-spacerun:yes'>  </sp=
an>The
following highlights a few of the things that we know thus far.<o:p></o:p><=
/span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal align=3Dcenter style=3D'text-align:center'><b style=3D=
'mso-bidi-font-weight:
normal'><span style=3D'font-size:12.0pt;font-family:Tahoma'>Maize Adh1<o:p>=
</o:p></span></b></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><o:p>&nbsp;</o:p></span></p>

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<span style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt'><o:p></o:p></spa=
n></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt;
font-family:Arial'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal style=3D'text-indent:.5in'><span style=3D'font-size:11=
.0pt;
mso-bidi-font-size:10.0pt;font-family:Arial'>The maize Adh1 promoter is div=
ided
into two regions of varying accessibility to <span class=3DSpellE>DNase</sp=
an>
I.<span style=3D'mso-spacerun:yes'>  </span>The TATA proximal region contai=
ns
hypersensitive sites that are present only when <i style=3D'mso-bidi-font-s=
tyle:
normal'>Adh1</i> is <span class=3DSpellE>transcriptionally</span> active (i=
nduced
by hypoxia) and a region from about –600 to –150 that contains constitutive=
ly
present hypersensitive sites (Paul et al.1987, PNAS <b style=3D'mso-bidi-fo=
nt-weight:
normal'><span style=3D'color:black'>84,</span></b><span style=3D'color:blac=
k'> 799</span>).<span
style=3D'mso-spacerun:yes'>   </span>A closer look at these sites with in v=
ivo
DMS <span class=3DSpellE>footprinting</span> revealed the binding sites of
several proteins (Ferl and Nick, 1987, JBC; Paul and Ferl, 1991, Plant Cell=
).
In the left panel below, the sites tagged in yellow (3-8) are constitutively
present whereas the sites tagged in green (1-2) are present only when <i
style=3D'mso-bidi-font-style:normal'>Adh1</i> is <span class=3DSpellE>trans=
criptionally</span>
active.<span style=3D'mso-spacerun:yes'>  </span>The right hand panel shows=
 the
in vivo DMS footprints from the region.<span style=3D'mso-spacerun:yes'> 
</span>Open triangles indicate protected sites and filled triangles indicate
sites of enhanced reactivity to DMS.<o:p></o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt;
font-family:Arial'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal style=3D'text-align:justify;line-height:12.0pt;tab-sto=
ps:.5in'><span
style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt;font-family:Arial'><span
style=3D'mso-tab-count:1'>            </span>The distribution of <span
class=3DSpellE>DNase</span> I hypersensitive sites clues you into another
structural feature of the maize <i style=3D'mso-bidi-font-style:normal'>Adh=
1</i>
promoter; a matrix attachment region (MAR). The 5' boundary of the <span
class=3DSpellE>DNase</span> I hypersensitive sites coincides with a region =
of the
promoter that remains attached to the nuclear matrix after <span class=3DSp=
ellE>histones</span>
have been depleted from isolated nuclei. The map below shows the positions =
of
the hypersensitive sites (again, yellow indicates those present constitutiv=
ely
and green indicates those present only when the gene is <span class=3DSpell=
E>transcriptionally</span>
active), the MAR.<o:p></o:p></span></p>

<p class=3DMsoNormal style=3D'text-align:justify;line-height:12.0pt;tab-sto=
ps:.5in'><span
style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt'><o:p>&nbsp;</o:p></spa=
n></p>

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ps:.5in'><span
style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt'><o:p>&nbsp;</o:p></spa=
n></p>

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<p class=3DMsoNormal style=3D'text-align:justify;line-height:12.0pt;tab-sto=
ps:.5in'><span
style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt;font-family:Arial'><o:p=
>&nbsp;</o:p></span></p>

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ps:.5in'><o:wrapblock><v:shape
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<span style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt;font-family:Arial=
'>Looking
at the map, it is easy to see that the MAR marks the edge of the constituti=
ve
hypersensitive area, and in fact, hypersensitive site #7 coincides with the
major OsO4 hypersensitive site (-589) that is evident when <span class=3DSp=
ellE>torsional</span>
stress is imposed on the promoter in a <span class=3DSpellE>supercoiled</sp=
an>
plasmid.<span style=3D'mso-spacerun:yes'>  </span>This type of hypersensiti=
vity
is absent when the promoter is not under <span class=3DSpellE>torsional</sp=
an>
stress, as with a <span class=3DSpellE>linearized</span> plasmid (Paul and =
Ferl,
1994). <o:p></o:p></span></p>

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    <td><![endif]>
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    <p class=3DMsoNormal><span style=3D'font-family:Arial;mso-bidi-font-fam=
ily:
    "Times New Roman"'>Linear / Supercoiled<o:p></o:p></span></p>
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    <![if !mso]></td>
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</v:shape><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt'><o:p>&=
nbsp;</o:p></span></p>

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<span style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt'><o:p>&nbsp;</o:p=
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<p class=3DMsoNormal style=3D'text-align:justify;line-height:12.0pt;tab-sto=
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  <![if !mso]>
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    <div>
    <p class=3DMsoNormal><span class=3DGramE><span style=3D'font-family:Ari=
al;
    mso-bidi-font-family:"Times New Roman"'>OsO<sub>4</sub><span
    style=3D'mso-spacerun:yes'>  </span>hyperreactivity</span></span><span
    style=3D'font-family:Arial;mso-bidi-font-family:"Times New Roman"'><o:p=
></o:p></span></p>
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    <![if !mso]></td>
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</v:shape><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt'><o:p>&=
nbsp;</o:p></span></p>

<p class=3DMsoNormal style=3D'text-align:justify;line-height:12.0pt;tab-sto=
ps:.5in'><span
style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt'><o:p>&nbsp;</o:p></spa=
n></p>

<br style=3D'mso-ignore:vglayout' clear=3DALL>

<p class=3DMsoNormal style=3D'text-align:justify;line-height:12.0pt;tab-sto=
ps:.5in'><span
style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt;font-family:Arial'>The =
5’ MAR
may play a role in creating a <span class=3DSpellE>transcriptionally</span>
competent Adh1 promoter, but it does not appear to be involved in the higher
order organization of the maize genome. <o:p></o:p></span></p>

<p class=3DMsoNormal style=3D'text-align:justify;line-height:12.0pt;tab-sto=
ps:.5in'><span
style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt'><o:p>&nbsp;</o:p></spa=
n></p>

<p class=3DMsoBodyText><o:p>&nbsp;</o:p></p>

<p class=3DMsoBodyText><o:p>&nbsp;</o:p></p>

<p class=3DMsoBodyText><o:p>&nbsp;</o:p></p>

<p class=3DMsoBodyText><o:p>&nbsp;</o:p></p>

<p class=3DMsoBodyText><o:wrapblock><v:shape id=3D"_x0000_s1046" type=3D"#_=
x0000_t75"
  style=3D'position:absolute;left:0;text-align:left;margin-left:221.75pt;
  margin-top:6.55pt;width:206pt;height:165.1pt;z-index:11'>
  <v:imagedata src=3D"adhchromatinsummaryforwebpage_files/image005.wmz" o:t=
itle=3D""/>
  <w:wrap type=3D"topAndBottom"/>
 </v:shape></o:wrapblock><br style=3D'mso-ignore:vglayout' clear=3DALL>
<v:shape id=3D"_x0000_s1050" type=3D"#_x0000_t202" style=3D'position:absolu=
te;left:0;
 text-align:left;margin-left:75.3pt;margin-top:174.85pt;width:78.75pt;heigh=
t:24.75pt;
 z-index:12' stroked=3D"f">
 <v:textbox>
  <![if !mso]>
  <table cellpadding=3D0 cellspacing=3D0 width=3D"100%">
   <tr>
    <td><![endif]>
    <div>
    <p class=3DMsoNormal><span style=3D'font-size:12.0pt;font-family:Tahoma=
'>Maize<o:p></o:p></span></p>
    </div>
    <![if !mso]></td>
   </tr>
  </table>
  <![endif]></v:textbox>
</v:shape><v:shape id=3D"_x0000_s1051" type=3D"#_x0000_t202" style=3D'posit=
ion:absolute;
 left:0;text-align:left;margin-left:295.3pt;margin-top:171.1pt;width:78.75p=
t;
 height:24.75pt;z-index:13' stroked=3D"f">
 <v:textbox>
  <![if !mso]>
  <table cellpadding=3D0 cellspacing=3D0 width=3D"100%">
   <tr>
    <td><![endif]>
    <div>
    <p class=3DMsoNormal><span style=3D'font-size:12.0pt;font-family:Tahoma=
'>Arabidopsis<o:p></o:p></span></p>
    </div>
    <![if !mso]></td>
   </tr>
  </table>
  <![endif]></v:textbox>
</v:shape><o:wrapblock><v:shape id=3D"_x0000_s1036" type=3D"#_x0000_t75" st=
yle=3D'position:absolute;
  left:0;text-align:left;margin-left:-6.25pt;margin-top:1.75pt;width:203.5p=
t;
  height:172.15pt;z-index:7'>
  <v:imagedata src=3D"adhchromatinsummaryforwebpage_files/image006.wmz" o:t=
itle=3D""/>
  <w:wrap type=3D"topAndBottom"/>
 </v:shape></o:wrapblock><br style=3D'mso-ignore:vglayout' clear=3DALL>
<o:p>&nbsp;</o:p></p>

<p class=3DMsoBodyText><o:p>&nbsp;</o:p></p>

<p class=3DMsoBodyText><span style=3D'font-family:Arial'><o:p>&nbsp;</o:p><=
/span></p>

<br style=3D'mso-ignore:vglayout' clear=3DALL>

<p class=3DMsoBodyText><span style=3D'font-family:Arial'>Limited digestion =
of maize
and <span class=3DSpellE>arabidopsis</span> nuclei with <span class=3DSpell=
E>DNase</span>&nbsp;I
results in the conversion of the greater than 800&nbsp;kb genomic DNA to an
accumulation of fragments that may represent a collection of individual dom=
ains
of the genome created by preferential cleavage at super-hypersensitive regi=
ons.
The median size of these fragments is about 45&nbsp;kb in maize and about
25&nbsp;kb in Arabidopsis (Paul and Ferl, 1998, Plant Cell).<span
style=3D'mso-spacerun:yes'>  </span><o:p></o:p></span></p>

<p class=3DMsoNormal style=3D'text-align:justify'><span style=3D'font-size:=
11.0pt;
mso-bidi-font-size:10.0pt'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal style=3D'text-align:justify'><span style=3D'font-size:=
11.0pt;
mso-bidi-font-size:10.0pt;font-family:Arial'>Hybridization analyses with
specific gene probes revealed that individual genes occupied discrete domai=
ns
within the distribution created by <span class=3DSpellE>DNase</span> I. The=
 maize
<i style=3D'mso-bidi-font-style:normal'>alcohol dehydrogenase</i> - <i
style=3D'mso-bidi-font-style:normal'>Adh1</i> gene occupied a domain of
90&nbsp;kb and the maize<i style=3D'mso-bidi-font-style:normal'> general
regulatory factor</i> - <i style=3D'mso-bidi-font-style:normal'>GRF1</i> ge=
ne
occupied a domain of 100&nbsp;kb in length.<span style=3D'mso-spacerun:yes'=
> 
</span>Arabidopsis <span class=3DSpellE><i style=3D'mso-bidi-font-style:nor=
mal'>Adh</i></span>
was found within two distinct domains of 8.3 and 6.1&nbsp;kb, whereas an <s=
pan
class=3DGramE>Arabidopsis<span style=3D'mso-spacerun:yes'>  </span><i
style=3D'mso-bidi-font-style:normal'>GRF</i></span> gene occupied a single =
domain
of 27&nbsp;kb.<span style=3D'mso-spacerun:yes'>  </span>The domains created=
 <span
class=3DGramE>by<span style=3D'mso-spacerun:yes'>  </span><span class=3DSpe=
llE>topoisomerase</span></span>
II-mediated cleavage were identical in size to those created by <span
class=3DSpellE>DNase</span> I. These results imply that the genome is not
packaged by means of a random gathering of the genome into domains of indis=
criminate
length, but rather, that the genome is gathered into specific domains and a
gene consistently occupies a discrete physical section of the genome. Our
proposed model is that these large organizational domains represent the
fundamental structural loop domains created by attachment of chromatin to t=
he
nuclear matrix at loop basements. These loop domains may be distinct from t=
he
domains created by the matrix attachment regions that typically flank small=
er,
often functionally distinct sections of the genome.<o:p></o:p></span></p>

<p class=3DMsoNormal style=3D'text-align:justify;line-height:12.0pt;tab-sto=
ps:.5in'><span
style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt'><o:p>&nbsp;</o:p></spa=
n></p>

<p class=3DMsoNormal style=3D'text-align:justify;line-height:12.0pt;tab-sto=
ps:.5in'><span
style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt'><o:p>&nbsp;</o:p></spa=
n></p>

<p class=3DMsoNormal style=3D'text-align:justify;line-height:12.0pt;tab-sto=
ps:.5in'><span
style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt'><o:p>&nbsp;</o:p></spa=
n></p>

<p class=3DMsoNormal style=3D'text-align:justify;line-height:12.0pt;tab-sto=
ps:.5in'><span
style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt'><o:p>&nbsp;</o:p></spa=
n></p>

<p class=3DMsoNormal style=3D'text-align:justify;line-height:12.0pt;tab-sto=
ps:.5in'><span
style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt'><o:p>&nbsp;</o:p></spa=
n></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><span
style=3D'mso-tab-count:1'>            </span><o:p></o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><span
style=3D'mso-spacerun:yes'> </span><o:p></o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal align=3Dcenter style=3D'text-align:center'><b style=3D=
'mso-bidi-font-weight:
normal'><span style=3D'font-size:12.0pt;font-family:Tahoma'>Arabidopsis <sp=
an
class=3DSpellE><span class=3DGramE>Adh</span></span><o:p></o:p></span></b><=
/p>

<p class=3DMsoNormal align=3Dcenter style=3D'text-align:center'><o:wrapbloc=
k><v:shape
  id=3D"_x0000_s1028" type=3D"#_x0000_t75" style=3D'position:absolute;left:=
0;
  text-align:left;margin-left:2.55pt;margin-top:21.25pt;width:424.6pt;heigh=
t:318.15pt;
  z-index:4'>
  <v:imagedata src=3D"adhchromatinsummaryforwebpage_files/image007.wmz" o:t=
itle=3D""/>
  <w:wrap type=3D"topAndBottom"/>
 </v:shape></o:wrapblock><br style=3D'mso-ignore:vglayout' clear=3DALL>
<b style=3D'mso-bidi-font-weight:normal'><span style=3D'font-size:12.0pt;
font-family:Tahoma'><o:p></o:p></span></b></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal style=3D'text-indent:.5in'><span style=3D'font-size:11=
.0pt;
mso-bidi-font-size:10.0pt;font-family:Arial'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal style=3D'text-indent:.5in'><span style=3D'font-size:11=
.0pt;
mso-bidi-font-size:10.0pt;font-family:Arial'>The TATA proximal region of th=
e <span
class=3DSpellE>arabidppsis</span> <span class=3DSpellE><span class=3DGramE>=
Adh</span></span>
promoter also contains hypersensitive sites that are present only when <span
class=3DSpellE><i style=3D'mso-bidi-font-style:normal'>Adh</i></span> is <s=
pan
class=3DSpellE>transcriptionally</span> active (induced by hypoxia) (Vega-<=
span
class=3DSpellE>Palas</span> and Ferl 1995 Plant Cell; Paul and Ferl 1998, D=
ev
Genet).<span style=3D'mso-spacerun:yes'>   </span>A closer look at these si=
tes
with in vivo DMS <span class=3DSpellE>footprinting</span> revealed the bind=
ing
sites of several proteins (Ferl and Laughner, 1989; Lu et al 1996,). In the
left panel below, the sites tagged in green are present only when <span
class=3DSpellE><span class=3DGramE><i style=3D'mso-bidi-font-style:normal'>=
Adh</i></span></span>
is <span class=3DSpellE>transcriptionally</span> active.<span
style=3D'mso-spacerun:yes'>  </span>The right hand panel shows the in vivo =
and in
vitro DMS footprints from the region.<span style=3D'mso-spacerun:yes'> 
</span>Open triangles indicate protected sites and filled triangles indicate
sites of enhanced reactivity to DMS.<o:p></o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><o:p>&nbsp;</o:p></span></p>

<span style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt;font-family:"Time=
s New Roman";
mso-fareast-font-family:"Times New Roman";mso-ansi-language:EN-US;mso-farea=
st-language:
EN-US;mso-bidi-language:AR-SA'><br clear=3Dall style=3D'page-break-before:a=
lways'>
</span>

<p class=3DMsoNormal align=3Dcenter style=3D'text-align:center'><b style=3D=
'mso-bidi-font-weight:
normal'><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt;font-fami=
ly:
Tahoma'>Maps of chromatin features of the Arabidopsis <span class=3DSpellE>=
<span
class=3DGramE>Adh</span></span> gene<o:p></o:p></span></b></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt;
font-family:Tahoma'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt;
font-family:Tahoma'>The region around the start of transcription in Arabido=
psis
<span class=3DSpellE><span class=3DGramE>Adh</span></span> is hypersensitiv=
e to <span
class=3DSpellE>DNase</span>. The sites correspond to regions of DNA-protein
interactions (</span><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt;
font-family:Arial'>Vega-<span class=3DSpellE>Palas</span> and Ferl 1995 Pla=
nt
Cell; Paul and Ferl 1998, Dev Genet; Ferl and Laughner, 1989; Lu et al 1996=
).</span><span
style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt;font-family:Tahoma'><o:=
p></o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><v:shape
 id=3D"_x0000_i1027" type=3D"#_x0000_t75" style=3D'width:425.25pt;height:20=
4pt'
 o:ole=3D"">
 <v:imagedata src=3D"adhchromatinsummaryforwebpage_files/image008.emz" o:ti=
tle=3D""
  croptop=3D"18447f" cropbottom=3D"4369f" cropleft=3D"2751f" cropright=3D"3=
398f"/>
</v:shape><!--[if gte mso 9]><xml>
 <o:OLEObject Type=3D"Embed" ProgID=3D"PowerPoint.Slide.8" ShapeID=3D"_x000=
0_i1027"
  DrawAspect=3D"Content" ObjectID=3D"_1083019425">
 </o:OLEObject>
</xml><![endif]--><o:p></o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt;
font-family:Tahoma'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt;
font-family:Tahoma'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt;
font-family:Tahoma'>The region around the transcription stop site in
Arabidopsis <span class=3DSpellE>Adh</span> is also hypersensitive to <span
class=3DSpellE><span class=3DGramE>DNase</span></span><span class=3DGramE> =
<span
style=3D'mso-spacerun:yes'> </span>(</span></span><span style=3D'font-size:=
11.0pt;
mso-bidi-font-size:10.0pt;font-family:Arial'>Paul and Ferl 1998, Dev Genet)=
.</span><span
style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt;font-family:Tahoma'><o:=
p></o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><o:p>&nbsp;</o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><v:shape
 id=3D"_x0000_i1028" type=3D"#_x0000_t75" style=3D'width:422.25pt;height:18=
2.25pt'
 o:ole=3D"">
 <v:imagedata src=3D"adhchromatinsummaryforwebpage_files/image009.emz" o:ti=
tle=3D""
  croptop=3D"22088f" cropbottom=3D"4612f" cropleft=3D"2104f" cropright=3D"3=
398f"/>
</v:shape><!--[if gte mso 9]><xml>
 <o:OLEObject Type=3D"Embed" ProgID=3D"PowerPoint.Slide.8" ShapeID=3D"_x000=
0_i1028"
  DrawAspect=3D"Content" ObjectID=3D"_1083019426">
 </o:OLEObject>
</xml><![endif]--><o:p></o:p></span></p>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt'><o:p>&nbsp;</o:p></span></p>

<span style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt;font-family:"Time=
s New Roman";
mso-fareast-font-family:"Times New Roman";mso-ansi-language:EN-US;mso-farea=
st-language:
EN-US;mso-bidi-language:AR-SA'><br clear=3Dall style=3D'page-break-before:a=
lways'>
</span>

<div style=3D'border:none;border-bottom:solid windowtext 1.0pt;mso-border-b=
ottom-alt:
solid windowtext .5pt;padding:0in 0in 1.0pt 0in'>

<p class=3DMsoNormal align=3Dcenter style=3D'text-align:center;border:none;
mso-border-bottom-alt:solid windowtext .5pt;padding:0in;mso-padding-alt:0in=
 0in 1.0pt 0in'><b
style=3D'mso-bidi-font-weight:normal'><span style=3D'font-size:11.0pt;mso-b=
idi-font-size:
10.0pt;font-family:Tahoma'>Higher Order Chromatin Structures – Ideas and Mo=
dels<o:p></o:p></span></b></p>

</div>

<p class=3DMsoNormal><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.=
0pt;
font-family:Tahoma'>Paul and Ferl, 1999, PMB<o:p></o:p></span></p>

<p class=3DMsoNormal align=3Dcenter style=3D'text-align:center'><b style=3D=
'mso-bidi-font-weight:
normal'><span style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt;font-fami=
ly:
Tahoma'><o:p>&nbsp;</o:p></span></b></p>

<p class=3DMsoNormal><b style=3D'mso-bidi-font-weight:normal'><span
style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt;font-family:Arial'>At l=
east
two types of matrix attachments may contribute to the organization of <br>
genes and the genome - <span class=3DSpellE><span style=3D'mso-bidi-font-we=
ight:
bold'>MAR</span>s</span> and <span class=3DSpellE><span style=3D'mso-bidi-f=
ont-weight:
bold'>LBAR</span>s</span><o:p></o:p></span></b></p>

<p class=3DMsoNormal><b style=3D'mso-bidi-font-weight:normal'><span
style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt;font-family:Arial'><o:p=
>&nbsp;</o:p></span></b></p>

<p class=3DMsoNormal><b style=3D'mso-bidi-font-weight:normal'><span
style=3D'font-size:11.0pt;mso-bidi-font-size:10.0pt;font-family:Arial'><o:p=
>&nbsp;</o:p></span></b></p>

<p class=3DMsoNormal align=3Dcenter style=3D'text-align:center'><!--[if mso=
 & !supportInlineShapes & supportFields]><b
style=3D'mso-bidi-font-weight:normal'><span style=3D'font-size:11.0pt;mso-b=
idi-font-size:
10.0pt;font-family:Tahoma'><span style=3D'mso-element:field-begin;mso-field=
-lock:
yes'></span><span style=3D'mso-spacerun:yes'> </span>SHAPE<span
style=3D'mso-spacerun:yes'>  </span>\* MERGEFORMAT <span style=3D'mso-eleme=
nt:field-separator'></span></span></b><![endif]--><b
style=3D'mso-bidi-font-weight:normal'><span style=3D'font-size:11.0pt;mso-b=
idi-font-size:
10.0pt;font-family:Tahoma'><v:group id=3D"_x0000_s1079" editas=3D"canvas" s=
tyle=3D'width:6in;
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=3D"10448,6558">
 <o:lock v:ext=3D"edit" aspectratio=3D"t"/>
 <v:shape id=3D"_x0000_s1078" type=3D"#_x0000_t75" style=3D'position:absolu=
te;left:2520;
  top:2620;width:10448;height:6558' o:preferrelative=3D"f">
  <v:fill o:detectmouseclick=3D"t"/>
  <v:path o:extrusionok=3D"t" o:connecttype=3D"none"/>
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ion:absolute;
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9;top:5618;
  width:10056;height:2346;mso-position-horizontal:absolute;
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