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The eIF4F and eIFiso4F Complexes of Plants: An Evolutionary Perspective

Identifieur interne : 001355 ( Pmc/Curation ); précédent : 001354; suivant : 001356

The eIF4F and eIFiso4F Complexes of Plants: An Evolutionary Perspective

Auteurs : Ryan M. Patrick [États-Unis] ; Karen S. Browning [États-Unis]

Source :

RBID : PMC:3352336

Abstract

Translation initiation in eukaryotes requires a number of initiation factors to recruit the assembled ribosome to mRNA. The eIF4F complex plays a key role in initiation and is a common target point for regulation of protein synthesis. Most work on the translation machinery of plants to date has focused on flowering plants, which have both the eIF4F complex (eIF4E and eIF4G) as well as the plant-specific eIFiso4F complex (eIFiso4E and eIFiso4G). The increasing availability of plant genome sequence data has made it possible to trace the evolutionary history of these two complexes in plants, leading to several interesting discoveries. eIFiso4G is conserved throughout plants, while eIFiso4E only appears with the evolution of flowering plants. The eIF4G N-terminus, which has been difficult to annotate, appears to be well conserved throughout the plant lineage and contains two motifs of unknown function. Comparison of eIFiso4G and eIF4G sequence data suggests conserved features unique to eIFiso4G and eIF4G proteins. These findings have answered some questions about the evolutionary history of the two eIF4F complexes of plants, while raising new ones.


Url:
DOI: 10.1155/2012/287814
PubMed: 22611336
PubMed Central: 3352336

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<journal-meta>
<journal-id journal-id-type="nlm-ta">Comp Funct Genomics</journal-id>
<journal-id journal-id-type="iso-abbrev">Comp. Funct. Genomics</journal-id>
<journal-id journal-id-type="publisher-id">CFG</journal-id>
<journal-title-group>
<journal-title>Comparative and Functional Genomics</journal-title>
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<issn pub-type="ppub">1531-6912</issn>
<issn pub-type="epub">1532-6268</issn>
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<publisher-name>Hindawi Publishing Corporation</publisher-name>
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<article-id pub-id-type="pmid">22611336</article-id>
<article-id pub-id-type="pmc">3352336</article-id>
<article-id pub-id-type="doi">10.1155/2012/287814</article-id>
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<subj-group subj-group-type="heading">
<subject>Review Article</subject>
</subj-group>
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<title-group>
<article-title>The eIF4F and eIFiso4F Complexes of Plants: An Evolutionary Perspective</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Patrick</surname>
<given-names>Ryan M.</given-names>
</name>
<xref ref-type="aff" rid="I1"></xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Browning</surname>
<given-names>Karen S.</given-names>
</name>
<xref ref-type="aff" rid="I1"></xref>
<xref ref-type="corresp" rid="cor1">*</xref>
</contrib>
</contrib-group>
<aff id="I1">Department of Chemistry and Biochemistry and the Institute for Cell and Molecular Biology, The University of Texas at Austin, Austin, TX 78712, USA</aff>
<author-notes>
<corresp id="cor1">*Karen S. Browning:
<email>kbrowning@mail.utexas.edu</email>
</corresp>
<fn fn-type="other">
<p>Academic Editor: Thomas Preiss</p>
</fn>
</author-notes>
<pub-date pub-type="ppub">
<year>2012</year>
</pub-date>
<pub-date pub-type="epub">
<day>7</day>
<month>5</month>
<year>2012</year>
</pub-date>
<volume>2012</volume>
<elocation-id>287814</elocation-id>
<history>
<date date-type="received">
<day>12</day>
<month>1</month>
<year>2012</year>
</date>
<date date-type="accepted">
<day>16</day>
<month>2</month>
<year>2012</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright © 2012 R. M. Patrick and K. S. Browning.</copyright-statement>
<copyright-year>2012</copyright-year>
<license license-type="open-access">
<license-p>This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.</license-p>
</license>
</permissions>
<abstract>
<p>Translation initiation in eukaryotes requires a number of initiation factors to recruit the assembled ribosome to mRNA. The eIF4F complex plays a key role in initiation and is a common target point for regulation of protein synthesis. Most work on the translation machinery of plants to date has focused on flowering plants, which have both the eIF4F complex (eIF4E and eIF4G) as well as the plant-specific eIFiso4F complex (eIFiso4E and eIFiso4G). The increasing availability of plant genome sequence data has made it possible to trace the evolutionary history of these two complexes in plants, leading to several interesting discoveries. eIFiso4G is conserved throughout plants, while eIFiso4E only appears with the evolution of flowering plants. The eIF4G N-terminus, which has been difficult to annotate, appears to be well conserved throughout the plant lineage and contains two motifs of unknown function. Comparison of eIFiso4G and eIF4G sequence data suggests conserved features unique to eIFiso4G and eIF4G proteins. These findings have answered some questions about the evolutionary history of the two eIF4F complexes of plants, while raising new ones.</p>
</abstract>
</article-meta>
</front>
<floats-group>
<fig id="fig1" position="float">
<label>Figure 1</label>
<caption>
<p>Phylogeny of Viridiplantae eIF4E and eIFiso4E. eIF4E
<sub>gs</sub>
genes of gymnosperms are labeled eIF4EG. Phylogeny generated by alignment of eIF4E, eIFiso4E, and eIF4E
<sub>gs</sub>
genes using MAFFT version 6 [
<xref ref-type="bibr" rid="B21">21</xref>
].</p>
</caption>
<graphic xlink:href="CFG2012-287814.001"></graphic>
</fig>
<fig id="fig2" position="float">
<label>Figure 2</label>
<caption>
<p>Domain organization of eIF4G and eIFiso4G from mammals, angiosperms, and the green algae
<italic>Chlamydomonas reinhardtii</italic>
. Plant eIF4G and eIFiso4G share the core organization of the eIF4E-binding site, HEAT-1/MIF4G domain, H1-CT motif, and HEAT-2/MA3 domain with mammals, but do not have the C-terminal HEAT-3 domain. The eIF3- and eIF4A-binding regions are thought to be maintained between all shown isoforms. Plant eIF4G has a longer N-terminus than mammals and contains the plant-specific 4G-PN1 and 4G-PN2 motifs as shown.
<italic>Chlamydomonas </italic>
eIF4G has a 4G-PN1-like sequence but no 4G-PN2 motif, while other green algae may have a 4G-PN2 motif but no 4G-PN1 motif. eIFiso4G is remarkably well conserved across plants, with the N-terminal XSLRPGG motif maintained from green algae to angiosperms.</p>
</caption>
<graphic xlink:href="CFG2012-287814.002"></graphic>
</fig>
<fig id="fig3" position="float">
<label>Figure 3</label>
<caption>
<p>The N-terminal motifs of eIF4G. Residues highlighted in green have identity to the consensus sequence, and residues highlighted in blue have similarity. Genes of nonflowering plants and green algae are shaded grey. (a) The PG-N1 motif with consensus sequence PARTSAPPNxDEQKRxQ. (b) The PGN-2 motif with consensus sequence VKITxPxTHEELxLD.</p>
</caption>
<graphic xlink:href="CFG2012-287814.003"></graphic>
</fig>
<fig id="fig4" position="float">
<label>Figure 4</label>
<caption>
<p>The H1-CT motif of eIF4G and eIFiso4G. Residues highlighted in green have identity to the shared core sequence RRx
<sub>5</sub>
KxIxExHxxA. The arrow identifies the site of the
<italic>cum2</italic>
mutation in eIF4G. (a) The H1-CT motif of eIF4G. Residues highlighted in purple have identity to the unique residues of the eIF4G H1-CT motif RRVEGPKKI(D/E)EVHRDA. Genes of nonflowering plants and green algae are shaded grey. (b) The H1-CT motif of eIFiso4G. Residues highlighted in yellow have identity to the unique residues of the eIFiso4G H1-CT motif PRREExKAKTIxEHxEAExxLG. Genes of nonflowering plants and green algae are shaded grey. (c) The H1-CT motif of eIF4G genes of heterokonts, animals, and fungi. Residues are highlighted according to their identity to the shared core motif (green), the motif of plant eIF4G (purple), or the motif of eIFiso4G (yellow).</p>
</caption>
<graphic xlink:href="CFG2012-287814.004"></graphic>
</fig>
<fig id="fig5" position="float">
<label>Figure 5</label>
<caption>
<p>The N-terminal XSLRPGG motif of eIFiso4G. Residues highlighted in green have identity to the consensus sequence, and the variable hydrophobic residue is highlighted in blue. Genes of nonflowering plants and green algae are shaded grey. Genes of the heterokont eIF4G sequences containing this motif are shaded in brown.</p>
</caption>
<graphic xlink:href="CFG2012-287814.005"></graphic>
</fig>
<fig id="fig6" position="float">
<label>Figure 6</label>
<caption>
<p>The 4E-binding site of plant eIF4G. Residues highlighted in green have identity to the consensus sequence KKYSRDFLLx
<sub>8</sub>
LPxxF, and residues highlighted blue have similarity. Genes of nonflowering plants and green algae are shaded grey.</p>
</caption>
<graphic xlink:href="CFG2012-287814.006"></graphic>
</fig>
<fig id="fig7" position="float">
<label>Figure 7</label>
<caption>
<p>The 4E-binding site of plant eIFiso4G. Residues highlighted in green have identity to the consensus sequence ERVRYTR(D/E)QLLZLRE, and residues highlighted blue have similarity. Genes of nonflowering plants and green algae are shaded grey. Plants generally have one copy of eIFiso4G that closely resembles the consensus sequence; this primary copy is highlighted in yellow. Secondary copies, which are unhighlighted, may diverge from this sequence.</p>
</caption>
<graphic xlink:href="CFG2012-287814.007"></graphic>
</fig>
<table-wrap id="tab1" position="float">
<label>Table 1</label>
<caption>
<p>Distribution of eIF4F subunit genes in Viridiplantae. Nonflowering plants and green algae are bold.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" rowspan="1" colspan="1"></th>
<th align="center" rowspan="1" colspan="1">eIF4G</th>
<th align="center" rowspan="1" colspan="1">eIFiso4G</th>
<th align="center" rowspan="1" colspan="1">eIF4E</th>
<th align="center" rowspan="1" colspan="1">eIFiso4E</th>
<th align="center" rowspan="1" colspan="1">4EHP</th>
<th align="center" rowspan="1" colspan="1">eIF4E1b</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Arabidopsis thaliana</italic>
</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">2</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Arabidopsis lyrata</italic>
</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">3</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">2</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Thellungiella halophila</italic>
</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Carica papaya</italic>
</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Theobroma cacao</italic>
</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Citrus clementina</italic>
</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Citrus sinensis</italic>
</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Eucalyptus grandis</italic>
</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Solanum tuberosum</italic>
</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Prunus persica</italic>
</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Fragaria vesca</italic>
</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Cucumis sativus</italic>
</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Glycine max</italic>
</td>
<td align="center" rowspan="1" colspan="1">4</td>
<td align="center" rowspan="1" colspan="1">4</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Medicago truncatula</italic>
</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Populus trichocarpa</italic>
</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">4</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Ricinus communis</italic>
</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Manihot esculenta</italic>
</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Vitis vinifera</italic>
</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Mimulus guttatus</italic>
</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Aquilegia coerulea</italic>
</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Sorghum bicolor</italic>
</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Zea mays</italic>
</td>
<td align="center" rowspan="1" colspan="1">3</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Setaria italica</italic>
</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Oryza sativa</italic>
</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<italic>Brachypodium distachyon</italic>
</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="center" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<bold>Selaginella moellendorffii</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>2</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>2</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>4</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>0</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>1</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>0</bold>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<bold>Physcomitrella patens</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>2</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>5</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>4</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>0</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>1</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>0</bold>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<bold>Chlamydomonas reinhardtii</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>1</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>1</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>1</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>0</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>0</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>0</bold>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<bold>Volvox carteri</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>1</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>1</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>1</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>0</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>0</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>0</bold>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">
<bold>Micromonas pusilla</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>1</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>1</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>1</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>0</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>0</bold>
</td>
<td align="center" rowspan="1" colspan="1">
<bold>0</bold>
</td>
</tr>
</tbody>
</table>
</table-wrap>
</floats-group>
</pmc>
</record>

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