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Subject: Nature Reviews Genetics - Highlights
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    <TD>
      <P><BR><FONT face=3Dtimes size=3D2><I>Nature Reviews Genetics</I> =
<B>4</B>,=20
      934 (2003); doi:10.1038/nrg1242</FONT> <!-- pdf link --><BR><IMG =
height=3D5=20
      alt=3D"" src=3D"http://www.nature.com/nrg/images/spacer.gif" =
width=3D1=20
      border=3D0><BR><A=20
      =
href=3D"http://www.nature.com/cgi-taf/DynaPage.taf?file=3D/nrg/journal/v4=
/n12/full/nrg1242_fs.html&amp;filetype=3Dpdf"=20
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      src=3D"http://www.nature.com/nrg/images/printable_pdf.gif" =
width=3D94=20
      align=3Dbottom border=3D0> </A><BR><FONT =
class=3Dpdfsize>[722K]</FONT> <!-- link to a correction --><!-- article =
title -->
      <P><FONT face=3DVerdana,Geneva,Arial,Helvetica,sans-serif><FONT=20
      size=3D2><B>EVOLUTION</B></FONT><BR><FONT class=3Dtitle=20
      face=3D"Verdana, Geneva, Arial, Helvetica, sans-serif" =
color=3D#cc0033=20
      size=3D4><SPAN class=3Dtitle><B>More variation than meets the=20
      eye</B></SPAN></FONT>=20
      <TABLE cellSpacing=3D0 cellPadding=3D0 width=3D"100%" =
bgColor=3D#000000=20
        border=3D0><TBODY>
        <TR>
          <TD><IMG height=3D1 alt=3D""=20
            src=3D"http://www.nature.com/nrg/images/spacer.gif" =
width=3D1=20
          border=3D0></TD></TR></TBODY></TABLE><!--authors --><IMG =
height=3D3 alt=3D""=20
      src=3D"http://www.nature.com/nrg/images/spacer.gif" width=3D1 =
border=3D0><BR><!--authors --><SPAN=20
      class=3Dauthor><B>Tanita&nbsp;Casci</B></SPAN>=20
      <P>Genetic variation is so pervasive that it would seem impossible =
to=20
      ignore. Yet ignore it we invariably do, because much of it is =
hidden in=20
      our genomes and surfaces only under certain genetic or =
environmental=20
      conditions. The precise molecular basis of such 'cryptic' =
variation has=20
      not been investigated, but Ian Dworkin and co-workers have now =
pinpointed=20
      the underlying nucleotides in one <I>Drosophila</I> gene =97 the=20
      <I>Epidermal growth factor receptor</I> (<A=20
      =
href=3D"http://www.ncbi.nlm.nih.gov/LocusLink/LocRpt.cgi?l=3D37455"=20
      target=3D_new><I>Egfr</I></A>) =97 by exploiting the =
characteristic ease with=20
      which the phenotype of the fly eye can be perturbed.
      <P>
      <TABLE cellSpacing=3D0 cellPadding=3D0 width=3D"5%" align=3Dright =
border=3D0>
        <TBODY>
        <TR>
          <TD><IMG height=3D1 alt=3D""=20
            src=3D"http://www.nature.com/nrg/images/spacer.gif" =
width=3D1 align=3Dleft=20
            border=3D0></TD>
          <TD vAlign=3Dtop align=3Dright><IMG alt=3D""=20
            =
src=3D"http://www.nature.com/nrg/journal/v4/n12/images/nrg1242-i1.jpg"=20
            border=3D0></TD></TR>
        <TR>
          <TD></TD>
          <TD>&nbsp;</TD></TR></TBODY></TABLE></P>
      <P>Flies that have a dominant gain-of-function allele of =
<I>Egfr</I>,=20
      called <I>Ellipse<SUP>1</SUP></I> (<I>Egfr<SUP>E1</SUP></I>), have =
a rough=20
      eye surface that is caused by the presence of too many =
photoreceptors,=20
      which reflects the normal role of <I>Egfr</I> in recruiting cells =
to the=20
      photoreceptor fate. The <I>Egfr<SUP>E1</SUP></I> phenotype is =
reasonably=20
      stable within a line; however, when <I>Egfr<SUP>E1</SUP></I> was =
crossed=20
      into 210 wild-type strains, the eye roughness of the progeny =
varied widely=20
      in severity from severely blistered to almost normal =97 an effect =
that had=20
      a strong genetic component.</P>
      <P>Could this effect be the result of cryptic variation in the =
wild-type=20
      <I>Egfr</I> gene, which interacts with <I>Egfr<SUP>E1</SUP></I> to =
modify=20
      its phenotype but is otherwise invisible? This theory is supported =
by the=20
      strong and independent association between half a dozen of the 267 =

      bi-allelic SNPs along the <I>Egfr</I> gene and the ability to =
modulate the=20
      rough-eye phenotype.</P>
      <P>So, the authors showed that a certain <I>Egfr</I> nucleotide =
variant=20
      might be associated with the ability to modify the=20
      <I>Egfr<SUP>E1</SUP></I> eye, but they had yet to confirm a causal =
link=20
      between the two. To do this, they turned to the progeny of 1,000 =
crosses=20
      between the <I>Egfr<SUP>E1</SUP></I> allele and a large outbred=20
      population; indeed, four of the candidate SNPs identified in the =
first=20
      study were associated with the more extreme rough-eye =
modifications in=20
      this cross.</P>
      <P>Although even this result does not constitute formal proof of=20
      causality, it is as close as we can get =97 short of putting the =
nucleotide=20
      variants back in the fly. Such an experiment will no doubt follow, =
along=20
      with others that aim to match the genetic structure observed here =
with=20
      that of hidden variation at other loci, or indeed at <I>Egfr</I> =
in other=20
      tissues in which it acts.</P>
      <P>
      <P>
      <TABLE cellSpacing=3D0 cellPadding=3D0 width=3D"100%" =
bgColor=3D#000000=20
        border=3D0><TBODY>
        <TR>
          <TD><IMG height=3D1 alt=3D""=20
            src=3D"http://www.nature.com/nrg/images/spacer.gif" =
width=3D1=20
          border=3D0></TD></TR></TBODY></TABLE><!-- references -->
      <P><A name=3Drefs></A>
      <H3><FONT color=3D#cc0033>References and links</FONT></H3>
      <TABLE cellSpacing=3D0 cellPadding=3D0 border=3D0>
        <TBODY>
        <TR>
          <TD vAlign=3Dtop><A name=3DB1></A>
            <P>&nbsp;</P></TD>
          <TD vAlign=3Dtop>
            <P><B>ORIGINAL RESEARCH PAPER</B><BR>Dworkin, I. <I>et =
al</I>.=20
            Evidence that <I>Egfr</I> contributes to cryptic genetic =
variation=20
            for photoreceptor determination in natural populations of=20
            <I>Drosophila melanogaster</I>. <I>Curr. Biol.</I> =
<B>13</B>,=20
            1888=961893 (2003)&nbsp;|&nbsp;<A=20
            href=3D"http://dx.doi.org/10.1016/j.cub.2003.10.001"=20
            target=3Dexternal><SPAN=20
            class=3Dintlink>Article</SPAN></A>&nbsp;|&nbsp;<A=20
            =
href=3D"http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?holding=3Dnpg&amp;c=
md=3DRetrieve&amp;db=3DPubMed&amp;list_uids=3D14588245&amp;dopt=3DAbstrac=
t"=20
            target=3Dexternal><SPAN =
class=3Dintlink>PubMed</SPAN></A>&nbsp;|&nbsp;<A=20
            =
href=3D"http://links.isiglobalnet2.com/gateway/Gateway.cgi?&amp;GWVersion=
=3D2&amp;SrcAuth=3DNature&amp;SrcApp=3DNature&amp;DestLinkType=3DFullReco=
rd&amp;KeyUT=3D000186263900021&amp;DestApp=3DWOS_CPL"=20
            target=3Dexternal><SPAN =
class=3Dintlink>ISI</SPAN></A>&nbsp;|&nbsp;<A=20
            =
href=3D"http://chemport.cas.org/cgi-bin/sdcgi?APP=3Dftslink&amp;action=3D=
reflink&amp;origin=3Dnpg&amp;version=3D1.0&amp;coi=3D1:CAS:528:DC%2BD3sXo=
slSjs7c%3D&amp;pissn=3D1471-0056&amp;pyear=3D2003&amp;md5=3D5ac6d08dd4a00=
139d5ca6dd0410160fd"=20
            target=3Dexternal alt=3D"Chemical Abstracts Service"><SPAN=20
            =
class=3Dintlink>ChemPort</SPAN></A>&nbsp;|</P></TD></TR></TBODY></TABLE><=
!-- end references -->
      <TABLE cellSpacing=3D0 cellPadding=3D0 border=3D0>
        <TBODY>
        <TR>
          <TD>
            <P>&nbsp;</P></TD>
          <TD>
            <P><A name=3Dlinks></A><!--links subcategory--><B>WEB =
SITE</B><BR><A=20
            href=3D"http://statgen.ncsu.edu/ggibson" target=3D_new>Greg =
Gibson's=20
            laboratory</A><BR></P></TD></TR></TBODY></TABLE>
      <P>&nbsp; <!-- footer -->
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      <TABLE cellSpacing=3D2 cellPadding=3D2 width=3D460>
        <TBODY>
        <TR>
          <TD><A=20
            =
href=3D"http://www.nature.com/cgi-taf/DynaPage.taf?file=3D/nrg/journal/v4=
/n12/full/nrg1242_r.html&amp;filetype=3D&amp;dynoptions=3D#top">
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            border=3D0><BR>&nbsp;<SPAN class=3Dfooter>back to =
top</SPAN></P></TD>
          <TD class=3Dfooter align=3Dright>NATURE =
REVIEWS&nbsp;|&nbsp;<SPAN=20
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