Geodermatophilus poikilotrophi sp. nov.: A Multitolerant Actinomycete Isolated from Dolomitic Marble
Identifieur interne : 000403 ( Pmc/Corpus ); précédent : 000402; suivant : 000404Geodermatophilus poikilotrophi sp. nov.: A Multitolerant Actinomycete Isolated from Dolomitic Marble
Auteurs : Maria Del Carmen Montero-Calasanz ; Benjamin Hofner ; Markus Göker ; Manfred Rohde ; Cathrin Spröer ; Karima Hezbri ; Maher Gtari ; Peter Schumann ; Hans-Peter KlenkSource :
- BioMed Research International [ 2314-6133 ] ; 2014.
Abstract
A novel Gram-reaction-positive, aerobic actinobacterium, tolerant to mitomycin C, heavy metals, metalloids, hydrogen peroxide, desiccation, and ionizing- and UV-radiation, designated G18T, was isolated from dolomitic marble collected from outcrops in Samara (Namibia). The growth range was 15–35°C, at pH 5.5–9.5 and in presence of 1% NaCl, forming greenish-black coloured colonies on GYM
Url:
DOI: 10.1155/2014/914767
PubMed: 25114928
PubMed Central: 4119925
Links to Exploration step
PMC:4119925Le document en format XML
<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en"><italic>Geodermatophilus poikilotrophi</italic> sp. nov.: A Multitolerant Actinomycete Isolated from Dolomitic Marble</title><author><name sortKey="Montero Calasanz, Maria Del Carmen" sort="Montero Calasanz, Maria Del Carmen" uniqKey="Montero Calasanz M" first="Maria Del Carmen" last="Montero-Calasanz">Maria Del Carmen Montero-Calasanz</name><affiliation><nlm:aff id="I1">Leibniz Institute DSMZ-German Collection of Microorganisms and Cell Cultures, Inhoffenstraße 7B, 38124 Braunschweig, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="I2">Instituto de Investigacióon y Formacióon Agraria y Pesquera (IFAPA), Centro Las Torres-Tomejil, Carretera Sevilla-Cazalla de la Sierra, Km 12.2, 41200 Alcalá del Río, Sevilla, Spain</nlm:aff></affiliation></author><author><name sortKey="Hofner, Benjamin" sort="Hofner, Benjamin" uniqKey="Hofner B" first="Benjamin" last="Hofner">Benjamin Hofner</name><affiliation><nlm:aff id="I3">Institut für Medizininformatik, Biometrie und Epidemiologie, Friedrich-Alexander-Universität Erlangen-Nürnberg, Waldstraße 6, 91054 Erlangen, Germany</nlm:aff></affiliation></author><author><name sortKey="Goker, Markus" sort="Goker, Markus" uniqKey="Goker M" first="Markus" last="Göker">Markus Göker</name><affiliation><nlm:aff id="I1">Leibniz Institute DSMZ-German Collection of Microorganisms and Cell Cultures, Inhoffenstraße 7B, 38124 Braunschweig, Germany</nlm:aff></affiliation></author><author><name sortKey="Rohde, Manfred" sort="Rohde, Manfred" uniqKey="Rohde M" first="Manfred" last="Rohde">Manfred Rohde</name><affiliation><nlm:aff id="I4">Helmholtz Centre for Infection Research (HZI), Inhoffenstraße 7, 38124 Braunschweig, Germany</nlm:aff></affiliation></author><author><name sortKey="Sproer, Cathrin" sort="Sproer, Cathrin" uniqKey="Sproer C" first="Cathrin" last="Spröer">Cathrin Spröer</name><affiliation><nlm:aff id="I1">Leibniz Institute DSMZ-German Collection of Microorganisms and Cell Cultures, Inhoffenstraße 7B, 38124 Braunschweig, Germany</nlm:aff></affiliation></author><author><name sortKey="Hezbri, Karima" sort="Hezbri, Karima" uniqKey="Hezbri K" first="Karima" last="Hezbri">Karima Hezbri</name><affiliation><nlm:aff id="I5">Laboratoire Microorganismes et Biomolécules Actives, Université de Tunis Elmanar (FST) et Université de Carthage (INSAT), 2092 Tunis, Tunisia</nlm:aff></affiliation></author><author><name sortKey="Gtari, Maher" sort="Gtari, Maher" uniqKey="Gtari M" first="Maher" last="Gtari">Maher Gtari</name><affiliation><nlm:aff id="I5">Laboratoire Microorganismes et Biomolécules Actives, Université de Tunis Elmanar (FST) et Université de Carthage (INSAT), 2092 Tunis, Tunisia</nlm:aff></affiliation></author><author><name sortKey="Schumann, Peter" sort="Schumann, Peter" uniqKey="Schumann P" first="Peter" last="Schumann">Peter Schumann</name><affiliation><nlm:aff id="I1">Leibniz Institute DSMZ-German Collection of Microorganisms and Cell Cultures, Inhoffenstraße 7B, 38124 Braunschweig, Germany</nlm:aff></affiliation></author><author><name sortKey="Klenk, Hans Peter" sort="Klenk, Hans Peter" uniqKey="Klenk H" first="Hans-Peter" last="Klenk">Hans-Peter Klenk</name><affiliation><nlm:aff id="I1">Leibniz Institute DSMZ-German Collection of Microorganisms and Cell Cultures, Inhoffenstraße 7B, 38124 Braunschweig, Germany</nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">25114928</idno><idno type="pmc">4119925</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4119925</idno><idno type="RBID">PMC:4119925</idno><idno type="doi">10.1155/2014/914767</idno><date when="2014">2014</date><idno type="wicri:Area/Pmc/Corpus">000403</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000403</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main"><italic>Geodermatophilus poikilotrophi</italic> sp. nov.: A Multitolerant Actinomycete Isolated from Dolomitic Marble</title><author><name sortKey="Montero Calasanz, Maria Del Carmen" sort="Montero Calasanz, Maria Del Carmen" uniqKey="Montero Calasanz M" first="Maria Del Carmen" last="Montero-Calasanz">Maria Del Carmen Montero-Calasanz</name><affiliation><nlm:aff id="I1">Leibniz Institute DSMZ-German Collection of Microorganisms and Cell Cultures, Inhoffenstraße 7B, 38124 Braunschweig, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="I2">Instituto de Investigacióon y Formacióon Agraria y Pesquera (IFAPA), Centro Las Torres-Tomejil, Carretera Sevilla-Cazalla de la Sierra, Km 12.2, 41200 Alcalá del Río, Sevilla, Spain</nlm:aff></affiliation></author><author><name sortKey="Hofner, Benjamin" sort="Hofner, Benjamin" uniqKey="Hofner B" first="Benjamin" last="Hofner">Benjamin Hofner</name><affiliation><nlm:aff id="I3">Institut für Medizininformatik, Biometrie und Epidemiologie, Friedrich-Alexander-Universität Erlangen-Nürnberg, Waldstraße 6, 91054 Erlangen, Germany</nlm:aff></affiliation></author><author><name sortKey="Goker, Markus" sort="Goker, Markus" uniqKey="Goker M" first="Markus" last="Göker">Markus Göker</name><affiliation><nlm:aff id="I1">Leibniz Institute DSMZ-German Collection of Microorganisms and Cell Cultures, Inhoffenstraße 7B, 38124 Braunschweig, Germany</nlm:aff></affiliation></author><author><name sortKey="Rohde, Manfred" sort="Rohde, Manfred" uniqKey="Rohde M" first="Manfred" last="Rohde">Manfred Rohde</name><affiliation><nlm:aff id="I4">Helmholtz Centre for Infection Research (HZI), Inhoffenstraße 7, 38124 Braunschweig, Germany</nlm:aff></affiliation></author><author><name sortKey="Sproer, Cathrin" sort="Sproer, Cathrin" uniqKey="Sproer C" first="Cathrin" last="Spröer">Cathrin Spröer</name><affiliation><nlm:aff id="I1">Leibniz Institute DSMZ-German Collection of Microorganisms and Cell Cultures, Inhoffenstraße 7B, 38124 Braunschweig, Germany</nlm:aff></affiliation></author><author><name sortKey="Hezbri, Karima" sort="Hezbri, Karima" uniqKey="Hezbri K" first="Karima" last="Hezbri">Karima Hezbri</name><affiliation><nlm:aff id="I5">Laboratoire Microorganismes et Biomolécules Actives, Université de Tunis Elmanar (FST) et Université de Carthage (INSAT), 2092 Tunis, Tunisia</nlm:aff></affiliation></author><author><name sortKey="Gtari, Maher" sort="Gtari, Maher" uniqKey="Gtari M" first="Maher" last="Gtari">Maher Gtari</name><affiliation><nlm:aff id="I5">Laboratoire Microorganismes et Biomolécules Actives, Université de Tunis Elmanar (FST) et Université de Carthage (INSAT), 2092 Tunis, Tunisia</nlm:aff></affiliation></author><author><name sortKey="Schumann, Peter" sort="Schumann, Peter" uniqKey="Schumann P" first="Peter" last="Schumann">Peter Schumann</name><affiliation><nlm:aff id="I1">Leibniz Institute DSMZ-German Collection of Microorganisms and Cell Cultures, Inhoffenstraße 7B, 38124 Braunschweig, Germany</nlm:aff></affiliation></author><author><name sortKey="Klenk, Hans Peter" sort="Klenk, Hans Peter" uniqKey="Klenk H" first="Hans-Peter" last="Klenk">Hans-Peter Klenk</name><affiliation><nlm:aff id="I1">Leibniz Institute DSMZ-German Collection of Microorganisms and Cell Cultures, Inhoffenstraße 7B, 38124 Braunschweig, Germany</nlm:aff></affiliation></author></analytic><series><title level="j">BioMed Research International</title><idno type="ISSN">2314-6133</idno><idno type="eISSN">2314-6141</idno><imprint><date when="2014">2014</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>A novel Gram-reaction-positive, aerobic actinobacterium, tolerant to mitomycin C, heavy metals, metalloids, hydrogen peroxide, desiccation, and ionizing- and UV-radiation, designated G18<sup>T</sup>, was isolated from dolomitic marble collected from outcrops in Samara (Namibia). The growth range was 15–35°C, at pH 5.5–9.5 and in presence of 1% NaCl, forming greenish-black coloured colonies on GYM <italic>Streptomyces</italic> agar. Chemotaxonomic and molecular characteristics of the isolate matched those described for other representatives of the genus <italic>Geodermatophilus</italic>. The peptidoglycan contained <italic>meso</italic>-diaminopimelic acid as diagnostic diaminoacid. The main phospholipids were phosphatidylethanolamine, phosphatidylcholine, phosphatidylinositol, and small amount of diphosphatidylglycerol. MK-9(H<sub>4</sub>) was the dominant menaquinone and galactose was detected as diagnostic sugar. The major cellular fatty acids were branched-chain saturated acids iso-C<sub>16:0</sub> and iso-C<sub>15:0</sub> and the unsaturated C<sub>17:1</sub><italic>ω</italic>8c and C<sub>16:1</sub><italic>ω</italic>7c. The 16S rRNA gene showed 97.4–99.1% sequence identity with the other representatives of genus<italic> Geodermatophilus</italic>. Based on phenotypic results and 16S rRNA gene sequence analysis, strain G18<sup>T</sup> is proposed to represent a novel species, <italic>Geodermatophilus poikilotrophi</italic>. Type strain is G18<sup>T</sup> (= DSM 44209<sup>T</sup> = CCUG 63018<sup>T</sup>). The INSDC accession number is HF970583. The novel R software package <monospace>lethal</monospace> was used to compute the lethal doses with confidence intervals resulting from tolerance experiments.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Normand, P" uniqKey="Normand P">P Normand</name></author><author><name sortKey="Orso, S" uniqKey="Orso S">S Orso</name></author><author><name sortKey="Cournoyer, B" uniqKey="Cournoyer B">B Cournoyer</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Normand, P" uniqKey="Normand P">P Normand</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Luedemann, Gm" uniqKey="Luedemann G">GM Luedemann</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Skerman, Vbd" uniqKey="Skerman V">VBD Skerman</name></author><author><name sortKey="Mcgowan, V" uniqKey="Mcgowan V">V McGowan</name></author><author><name sortKey="Sneath, Pha" uniqKey="Sneath 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<front><journal-meta><journal-id journal-id-type="nlm-ta">Biomed Res Int</journal-id><journal-id journal-id-type="iso-abbrev">Biomed Res Int</journal-id><journal-id journal-id-type="publisher-id">BMRI</journal-id><journal-title-group><journal-title>BioMed Research International</journal-title></journal-title-group><issn pub-type="ppub">2314-6133</issn><issn pub-type="epub">2314-6141</issn><publisher><publisher-name>Hindawi Publishing Corporation</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">25114928</article-id><article-id pub-id-type="pmc">4119925</article-id><article-id pub-id-type="doi">10.1155/2014/914767</article-id><article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group></article-categories><title-group><article-title><italic>Geodermatophilus poikilotrophi</italic> sp. nov.: A Multitolerant Actinomycete Isolated from Dolomitic Marble</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Montero-Calasanz</surname><given-names>Maria del Carmen</given-names></name><xref ref-type="aff" rid="I1"><sup>1</sup></xref><xref ref-type="aff" rid="I2"><sup>2</sup></xref><xref ref-type="corresp" rid="cor1">*</xref></contrib><contrib contrib-type="author"><name><surname>Hofner</surname><given-names>Benjamin</given-names></name><xref ref-type="aff" rid="I3"><sup>3</sup></xref></contrib><contrib contrib-type="author"><name><surname>Göker</surname><given-names>Markus</given-names></name><xref ref-type="aff" rid="I1"><sup>1</sup></xref></contrib><contrib contrib-type="author"><name><surname>Rohde</surname><given-names>Manfred</given-names></name><xref ref-type="aff" rid="I4"><sup>4</sup></xref></contrib><contrib contrib-type="author"><name><surname>Spröer</surname><given-names>Cathrin</given-names></name><xref ref-type="aff" rid="I1"><sup>1</sup></xref></contrib><contrib contrib-type="author"><name><surname>Hezbri</surname><given-names>Karima</given-names></name><xref ref-type="aff" rid="I5"><sup>5</sup></xref></contrib><contrib contrib-type="author"><name><surname>Gtari</surname><given-names>Maher</given-names></name><xref ref-type="aff" rid="I5"><sup>5</sup></xref></contrib><contrib contrib-type="author"><name><surname>Schumann</surname><given-names>Peter</given-names></name><xref ref-type="aff" rid="I1"><sup>1</sup></xref></contrib><contrib contrib-type="author"><name><surname>Klenk</surname><given-names>Hans-Peter</given-names></name><xref ref-type="aff" rid="I1"><sup>1</sup></xref><xref ref-type="corresp" rid="cor2">*</xref></contrib></contrib-group><aff id="I1"><sup>1</sup>Leibniz Institute DSMZ-German Collection of Microorganisms and Cell Cultures, Inhoffenstraße 7B, 38124 Braunschweig, Germany</aff><aff id="I2"><sup>2</sup>Instituto de Investigacióon y Formacióon Agraria y Pesquera (IFAPA), Centro Las Torres-Tomejil, Carretera Sevilla-Cazalla de la Sierra, Km 12.2, 41200 Alcalá del Río, Sevilla, Spain</aff><aff id="I3"><sup>3</sup>Institut für Medizininformatik, Biometrie und Epidemiologie, Friedrich-Alexander-Universität Erlangen-Nürnberg, Waldstraße 6, 91054 Erlangen, Germany</aff><aff id="I4"><sup>4</sup>Helmholtz Centre for Infection Research (HZI), Inhoffenstraße 7, 38124 Braunschweig, Germany</aff><aff id="I5"><sup>5</sup>Laboratoire Microorganismes et Biomolécules Actives, Université de Tunis Elmanar (FST) et Université de Carthage (INSAT), 2092 Tunis, Tunisia</aff><author-notes><corresp id="cor1">*Maria del Carmen Montero-Calasanz: <email>maria.montero-calasanz@dsmz.de</email> and </corresp><corresp id="cor2">*Hans-Peter Klenk: <email>hpk@dsmz.de</email></corresp><fn fn-type="other"><p>Academic Editor: Sara Borin</p></fn></author-notes><pub-date pub-type="ppub"><year>2014</year></pub-date><pub-date pub-type="epub"><day>9</day><month>7</month><year>2014</year></pub-date><volume>2014</volume><elocation-id>914767</elocation-id><history><date date-type="received"><day>1</day><month>4</month><year>2014</year></date><date date-type="rev-recd"><day>3</day><month>6</month><year>2014</year></date><date date-type="accepted"><day>9</day><month>6</month><year>2014</year></date></history><permissions><copyright-statement>Copyright © 2014 Maria del Carmen Montero-Calasanz et al.</copyright-statement><copyright-year>2014</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>A novel Gram-reaction-positive, aerobic actinobacterium, tolerant to mitomycin C, heavy metals, metalloids, hydrogen peroxide, desiccation, and ionizing- and UV-radiation, designated G18<sup>T</sup>, was isolated from dolomitic marble collected from outcrops in Samara (Namibia). The growth range was 15–35°C, at pH 5.5–9.5 and in presence of 1% NaCl, forming greenish-black coloured colonies on GYM <italic>Streptomyces</italic> agar. Chemotaxonomic and molecular characteristics of the isolate matched those described for other representatives of the genus <italic>Geodermatophilus</italic>. The peptidoglycan contained <italic>meso</italic>-diaminopimelic acid as diagnostic diaminoacid. The main phospholipids were phosphatidylethanolamine, phosphatidylcholine, phosphatidylinositol, and small amount of diphosphatidylglycerol. MK-9(H<sub>4</sub>) was the dominant menaquinone and galactose was detected as diagnostic sugar. The major cellular fatty acids were branched-chain saturated acids iso-C<sub>16:0</sub> and iso-C<sub>15:0</sub> and the unsaturated C<sub>17:1</sub><italic>ω</italic>8c and C<sub>16:1</sub><italic>ω</italic>7c. The 16S rRNA gene showed 97.4–99.1% sequence identity with the other representatives of genus<italic> Geodermatophilus</italic>. Based on phenotypic results and 16S rRNA gene sequence analysis, strain G18<sup>T</sup> is proposed to represent a novel species, <italic>Geodermatophilus poikilotrophi</italic>. Type strain is G18<sup>T</sup> (= DSM 44209<sup>T</sup> = CCUG 63018<sup>T</sup>). The INSDC accession number is HF970583. The novel R software package <monospace>lethal</monospace> was used to compute the lethal doses with confidence intervals resulting from tolerance experiments.</p></abstract></article-meta></front><body><sec id="sec1"><title>1. Introduction</title><p>The family cursive was originally proposed by Normand et al. [<xref rid="B1" ref-type="bibr">1</xref>], but a formal description of the family name was only published a decade later [<xref rid="B2" ref-type="bibr">2</xref>]. At the time of writing, the family comprises the genera<italic> Blastococcus, Modestobacter,</italic> and<italic> Geodermatophilus</italic> (as the type genus).<italic> Geodermatophilus</italic> was proposed by Luedemann [<xref rid="B3" ref-type="bibr">3</xref>] and was included in the Approved Lists of Bacterial Names [<xref rid="B4" ref-type="bibr">4</xref>]. This genus was poorly studied for a long time due to difficulties in culturing isolates [<xref rid="B5" ref-type="bibr">5</xref>], in spite of the fact that its members are frequently isolated from arid soils [<xref rid="B5" ref-type="bibr">5</xref>] and occasionally from arid and semiarid rock substrates such as rock vanish and marble [<xref rid="B6" ref-type="bibr">6</xref>, <xref rid="B7" ref-type="bibr">7</xref>], where a variety of environmental changing factors influence their settlement, growth, and development [<xref rid="B8" ref-type="bibr">8</xref>]. Some of them were also isolated from rhizosphere soil [<xref rid="B9" ref-type="bibr">9</xref>, <xref rid="B10" ref-type="bibr">10</xref>]. To enable the survival in such extreme ecological niches, where bacterial cells are suppressed to reactive oxygen species (ROS) generating-stresses, those should exhibit a very broad range of tolerance to multiple and fluctuating environmental stresses, such as solar radiation, desiccation and rehydration, temperature fluctuations, salts, and metals [<xref rid="B8" ref-type="bibr">8</xref>, <xref rid="B11" ref-type="bibr">11</xref>], and a probable ionizing-radiation (IR) resistance. The origin of this last capability cannot be explained as adaptation to environment, suggesting an “incidental” result of tolerance to desiccation, whose DNA damage pattern is similar to that generated by ionizing radiation in<italic> Deinococcus </italic>species [<xref rid="B12" ref-type="bibr">12</xref>]. Furthermore, tolerance to hydrogen peroxide and mitomycin C as indicators of the presence of an efficient microbial oxidative stress repair and double-strand break repair system, characteristics also attributed to radiation resistance, have been widely studied [<xref rid="B13" ref-type="bibr">13</xref>, <xref rid="B14" ref-type="bibr">14</xref>]. Multiple-stress tolerance of the type strain<italic> Geodermatophilus obscurus</italic> was already described by Gtari et al. [<xref rid="B11" ref-type="bibr">11</xref>], suggesting a correlation between tolerance profiles to desiccation, mitomycin C, hydrogen peroxide, and ionizing- and UV-radiation. Previous works of Rainey et al. [<xref rid="B15" ref-type="bibr">15</xref>] and Giongo et al. [<xref rid="B16" ref-type="bibr">16</xref>] already revealed the prevalence of IR resistant<italic> Geodermatophilus</italic> isolates from arid soil sample at comparatively the same radiation levels as observed for<italic> Deinococcus</italic> species and the predominance of species belongs to the family<italic> Geodermatophilaceae</italic> detected from intercontinental dust, illustrating, therefore, to resist radiation and desiccation stresses during travel in the high atmosphere.</p><p>Fourteen named species have been classified in the genus<italic> Geodermatophilus </italic>(ordered by the dates of effective publication of the names):<italic> G. obscurus</italic> [<xref rid="B3" ref-type="bibr">3</xref>],<italic> G. ruber</italic> [<xref rid="B9" ref-type="bibr">9</xref>],<italic> G. nigrescens</italic> [<xref rid="B17" ref-type="bibr">17</xref>],<italic> G. arenarius</italic> [<xref rid="B18" ref-type="bibr">18</xref>],<italic> G. siccatus</italic> [<xref rid="B19" ref-type="bibr">19</xref>, <xref rid="B20" ref-type="bibr">20</xref>],<italic> G. saharensis</italic> [<xref rid="B20" ref-type="bibr">20</xref>, <xref rid="B21" ref-type="bibr">21</xref>],<italic> G. tzadiensis </italic>[<xref rid="B22" ref-type="bibr">22</xref>, <xref rid="B23" ref-type="bibr">23</xref>],<italic> G. telluris</italic> [<xref rid="B24" ref-type="bibr">24</xref>],<italic> G. soli</italic> and<italic> G. terrae</italic> [<xref rid="B10" ref-type="bibr">10</xref>],<italic> G. africanus</italic> [<xref rid="B5" ref-type="bibr">5</xref>, <xref rid="B23" ref-type="bibr">23</xref>],<italic> G. normandii</italic> [<xref rid="B25" ref-type="bibr">25</xref>],<italic> G. taihuensis</italic> [<xref rid="B26" ref-type="bibr">26</xref>], and<italic> G. amargosae</italic> [<xref rid="B27" ref-type="bibr">27</xref>, <xref rid="B28" ref-type="bibr">28</xref>]. Until now, only the genome of the type strain of the type species,<italic> G. obscurus </italic>G-20<sup>T</sup>, has been sequenced [<xref rid="B29" ref-type="bibr">29</xref>]. Moreover, three subspecies have been identified and named, but their names were not validly published yet: “<italic>G. obscurus</italic> subsp.<italic> utahensis,</italic>” “<italic>G. obscurus</italic> subsp.<italic> dictyosporus</italic>” [<xref rid="B3" ref-type="bibr">3</xref>], and “<italic>G. obscurus </italic>subsp.<italic> everesti</italic>” [<xref rid="B30" ref-type="bibr">30</xref>, <xref rid="B31" ref-type="bibr">31</xref>]. This study describes the taxonomic position of a novel species into the genus<italic> Geodermatophilus</italic> based on a polyphasic approach and its tolerance to different environmental stresses.</p></sec><sec id="sec2"><title>2. Materials and Methods</title><sec id="sec2.1"><title>2.1. Isolation</title><p>During screening for microorganisms from dolomitic marble outcrops in an agriculture area at 1150 masl in Samara, near to Namib desert (Namibia), a greenish-black strain designated as G18<sup>T</sup> was isolated (in 1993) and purified as described by Eppard et al. [<xref rid="B7" ref-type="bibr">7</xref>].</p></sec><sec id="sec2.2"><title>2.2. Morphological and Biochemical Characterization</title><p>Cultural characteristics were tested on GYM<italic> Streptomyces</italic> medium (DSMZ medium 65), TSB agar (DSMZ medium 535), GPHF medium (DSMZ medium 553), R2A medium (DSMZ medium 830), GEO medium (DSMZ medium 714), PYGV medium (DSMZ medium 621), and Luedemann medium (DSMZ medium 877) for 15 days at 28°C. To determine its morphological characteristics, strain G18<sup>T</sup> was cultivated on GYM<italic> Streptomyces</italic> medium at 28°C. Colony features were observed at 4 and 15 days under a binocular microscope according to Pelczar Jr. [<xref rid="B32" ref-type="bibr">32</xref>]. Exponentially growing bacterial cultures were observed with an optical microscope (Zeiss AxioScope A1) with a 100-fold magnification and phase-contrast illumination. Micrographs of bacterial cells grown on GYM<italic> Streptomyces</italic> broth after 7 days were taken with a field-emission scanning electron microscope (FE-SEM Merlin, Zeiss, Germany). Gram reaction was performed using the KOH test described by Gregersen [<xref rid="B33" ref-type="bibr">33</xref>]. Cell motility was observed on modified ISP2 [<xref rid="B34" ref-type="bibr">34</xref>] swarming agar (0.3%, w/v) at pH 7.2 supplemented with (l<sup>−1</sup>) 4.0 g dextrin, 4.0 g yeast extract, and 10.0 g malt extract. Oxidase activity was analysed using filter-paper disks (Sartorius grade 388) impregnated with 1% solution of<italic> N,N,N</italic>′<italic>,N</italic>′-tetramethyl-<italic>p</italic>-phenylenediamine (Sigma-Aldrich); a positive test was defined by the development of a blue-purple colour after applying biomass to the filter paper. Catalase activity was determined based on formation of bubbles following the addition of 1 drop of 3% H<sub>2</sub>O<sub>2</sub>. Growth rates were determined on plates of GYM<italic> Streptomyces</italic> medium for temperatures from 10 to 50°C at 5°C increments and for pH values from 4.0 to 12.5 (in increments of 0.5 pH units) on modified ISP2 medium by adding NaOH or HCl, respectively, since the use of a buffer system inhibited growth of the strains. The utilization of carbon compounds and acid production were tested at 28°C using API 20 NE strips (bioMérieux) and GEN III Microplates in an Omnilog device (BIOLOG Inc., Hayward, CA, USA) in comparison with the reference strains<italic> G. africanus</italic> DSM 45422<sup>T</sup>,<italic> G. amargosae</italic> DSM 46136<sup>T</sup>,<italic> G. arenarius </italic>DSM 45418<sup>T</sup>,<italic> G. nigrescens </italic>DSM 45408<sup>T</sup>,<italic> G. normandii</italic> DSM 45417<sup>T</sup>,<italic> G. obscurus </italic>DSM 43160<sup>T</sup>,<italic> G. ruber</italic> DSM 45317<sup>T</sup>,<italic> G. saharensis</italic> DSM 45423<sup>T</sup>,<italic> G. siccatus</italic> DSM 45419<sup>T</sup>,<italic> G. soli </italic>DSM 45843<sup>T</sup><italic>, G. taihuensis </italic>DSM 45962<sup>T</sup>,<italic> G. telluris</italic> DSM 45421<sup>T</sup>,<italic> G. terrae</italic> DSM 45844<sup>T</sup>, and<italic> G. tzadiensis</italic> DSM 45416<sup>T</sup> in parallel assays. The GEN III Microplates were inoculated with cells suspended in a viscous inoculating fluid (IF C) provided by the manufacturer at a cell density of 70% transmittance (T) for<italic> G. amargosae </italic>DSM 46136<sup>T</sup>, at 75–79% T for<italic> G. africanus </italic>DSM 45422<sup>T</sup>, at 90% T for<italic> G. arenarius</italic> DSM 45418<sup>T</sup> and<italic> G. taihuensis </italic>DSM 45962<sup>T</sup>, and at 80–83% T for all other reference strains. Respiration rates (and growth) were measured yielding a total running time of 5 or 10 days, depending on the strain, in phenotype microarray mode. Each strain was studied in two independent repetitions. Data were exported and analysed using the <monospace> opm</monospace> package for R [<xref rid="B35" ref-type="bibr">35</xref>, <xref rid="B36" ref-type="bibr">36</xref>] v.1.0.6. Reactions with a distinct behaviour between the two repetitions were regarded as ambiguous. Clustering analyses of the phenotypic microarrays were constructed using the <monospace> pvclust</monospace> package for R v.1.2.2. [<xref rid="B37" ref-type="bibr">37</xref>]. Enzymatic activities were tested using API ZYM galleries according to the instructions of the manufacturer (bioMérieux).<italic> Chemotaxonomic procedures</italic>. Whole-cell sugars were prepared according to Lechevalier and Lechevalier [<xref rid="B38" ref-type="bibr">38</xref>], followed by thin layer chromatography (TLC) analysis [<xref rid="B39" ref-type="bibr">39</xref>]. Polar lipids were extracted, separated by two-dimensional TLC, and identified according to procedures outlined by Minnikin et al. [<xref rid="B40" ref-type="bibr">40</xref>] with modifications proposed by Kroppenstedt and Goodfellow [<xref rid="B41" ref-type="bibr">41</xref>]. Additionally, choline-containing lipids were detected by spraying with Dragendorff's reagent (Merck) [<xref rid="B42" ref-type="bibr">42</xref>]. Menaquinones (MK) were extracted from freeze-dried cell material using methanol as described by Collins et al. [<xref rid="B43" ref-type="bibr">43</xref>] and analysed by high-performance liquid chromatography (HPLC) [<xref rid="B44" ref-type="bibr">44</xref>]. The extraction and analysis of cellular fatty acids were carried out in two independent repetitions from biomass grown on GYM agar plates held at 28°C for 4 days and harvested always from the same sector (the last quadrant streak). Analysis was conducted using the microbial identification system (MIDI) Sherlock Version 4.5 (method TSBA40, ACTIN6 database) as described by Sasser [<xref rid="B45" ref-type="bibr">45</xref>]. The annotation of the fatty acids in the ACTIN6 peak naming table is consistent with IUPAC nomenclature (i.e., double bond positions identified with reference to the carboxyl group of the fatty acid), but for consistency with other publications this has been altered to numbering from the aliphatic end of the molecule (i.e., 16 : 1 CIS 9 become 16 : 1<italic>ω</italic>7c, etc.). The composition of peptidoglycan hydrolysates (6 N HCl, 100°C for 16 h) was examined by TLC as described by Schleifer and Kandler [<xref rid="B46" ref-type="bibr">46</xref>]. All chemotaxonomical analyses were conducted under standardized conditions with strain G18<sup>T</sup> and cultures of the same set of reference strains as listed above for morphological and biochemical characterisations.</p></sec><sec id="sec2.3"><title>2.3. Genetic and Phylogenetic Analysis</title><p>G + C content of chromosomal DNA of strain G18<sup>T</sup> was determined by HPLC according to Mesbah et al. [<xref rid="B47" ref-type="bibr">47</xref>]. Genomic DNA extraction, PCR-mediated amplification of the 16S rRNA gene, and purification of the PCR product were carried out as described by Rainey et al. [<xref rid="B48" ref-type="bibr">48</xref>]. Phylogenetic analysis was based on an alignment of 16S rRNA gene sequences from type strains of all species with effectively published names in the<italic> Geodermatophilaceae</italic> inferred as described by Montero-Calasanz et al. [<xref rid="B5" ref-type="bibr">5</xref>]. Pairwise similarities were calculated as recommended by Meier-Kolthoff et al. [<xref rid="B49" ref-type="bibr">49</xref>]. For DNA-DNA hybridization tests, cells were disrupted by using a Constant Systems TS 0.75 KW (IUL Instruments, Germany). DNA in the crude lysate was purified by chromatography on hydroxyapatite as described by Cashion et al. [<xref rid="B50" ref-type="bibr">50</xref>]. DNA-DNA hybridization was carried out as described by De Ley et al. [<xref rid="B51" ref-type="bibr">51</xref>] under consideration of the modifications described by Huss et al. [<xref rid="B52" ref-type="bibr">52</xref>] using a model Cary 100 Bio UV/VIS-spectrophotometer equipped with a Peltier-thermostatted 6 × 6 multicell changer and a temperature controller with<italic> in situ </italic>temperature probe (Varian).</p></sec><sec id="sec2.4"><title>2.4. Tolerance Experiments</title><p>The tolerance of strain G18<sup>T</sup> and<italic> G. obscurus</italic> G-20<sup>T</sup> (DSM 43160), as a positive control [<xref rid="B11" ref-type="bibr">11</xref>], to ionizing- and UV-radiation, mitomycin C, hydrogen peroxide, desiccation, and heavy metals/metalloids, was assayed using nonsporulating cultures obtained by growth in TYB medium [<xref rid="B53" ref-type="bibr">53</xref>] at 28°C for 5 days, washed twice with 0.9% NaCl, homogenized, and subsequently resuspended in saline solution. Ionizing-radiation experiments were carried out according to a protocol outlined by Gtari et al. [<xref rid="B11" ref-type="bibr">11</xref>]. To test the resistance to UV-radiation, 0.5 mL aliquots of culture suspensions was spread onto GYM<italic> Streptomyces</italic> agar plates in duplicate in two independent experiments and then exposed to a dose of 5–10 J<italic>·</italic>s<sup>−1</sup> m<sup>−2</sup> in a laminar flow hood equipped with crossbeam 254 nm UV sources in both side walls (Safe 2020, Thermo Scientific) for 1, 10, 30, 60, 120, 240, and 600 min. After 2 weeks at 28°C, the survival fractions were calculated based on the c.f.u. mL<sup>−1</sup>. The UV shadow zone was avoided. The tolerance to DNA damaging agent mitomycin C was tested in two independent experiments by incubation of cell suspension at room temperature with the antibiotic at a final concentration of 5 <italic>μ</italic>g<italic>·</italic>mL<sup>−1</sup>. After 1, 5, 10, 20, 40, 60, and 120 min, samples were centrifuged at 3500 rpm for 4 min, washed twice in 0.9% NaCl, and, subsequently, serially diluted. Aliquots were spread on GYM<italic> Streptomyces</italic> agar in duplicate. After incubation, the survival fractions were calculated based on the c.f.u. mL<sup>−1</sup>. To test the resistance to hydrogen peroxide, equal volumes of cell suspensions and 0.5% hydrogen peroxide were incubated at room temperature in two independent experiments. After 1, 5, 10, 20, 40, 60, and 120 min, samples were handled as was previously described in mitomycin experiments to calculate the survival fractions. For desiccation tolerance, 25 <italic>μ</italic>L of cell suspension were transferred to individual wells of microtiter plates in triplicate. Unsealed microtiter plates were placed in a desiccator (23.5% relative humidity) containing silica gel rubin (Fluka) at room temperature. After 20, 40, 60, 80, and 100 days, 250 <italic>μ</italic>L of sterile water was added to individual wells to rehydrate the desiccated cells and then incubated at room temperature for 1 hour and plated on GYM<italic> Streptomyces</italic> agar. The determination of survival fractions was conducted as described above. The sensitivity of strain G18<sup>T</sup> to heavy metals and metalloids was determined by a growth inhibition plate assay as described by Richards et al. [<xref rid="B54" ref-type="bibr">54</xref>]. AgNO<sub>3</sub>, CuCl<sub>2</sub>, CoCl<sub>2</sub>, NiCl<sub>2</sub>, K<sub>2</sub>CrO<sub>4</sub>, Pb(NO<sub>3</sub>)<sub>2</sub>, and Na<sub>2</sub>HAsO<sub>4</sub> were added to GYM<italic> Streptomyces</italic> medium at 0.1, 0.3, 0.5, 1.0, 2.0, 4.0, 8.0, 10.0, 30.0, and 50.0 mM. Growth was evaluated after 1 month at 28°C, determining minimum inhibitory concentration (MIC).</p></sec><sec id="sec2.5"><title>2.5. Statistical Analysis of Tolerance Experiments</title><p>To evaluate the tolerance of strain G18<sup>T</sup> and<italic> G. obscurus</italic> DSM 43160<sup>T</sup> with respect to the various physiological challenges, the median lethal dose (LD50) and the lethal dose 10 (LD10) values were computed for both strains. As the number of bacteria initially used in each experiment cannot directly be obtained and consequently, death rates or survival rates cannot be directly computed; standard models based on logistic regression models to obtain LD values are thus not available. A negative binomial model for count data [<xref rid="B55" ref-type="bibr">55</xref>] was used to estimate of number of survivors dependent on dose, strain, and experiment. Penalized splines [<xref rid="B56" ref-type="bibr">56</xref>], one for each strain, were used to allow the dose to have a nonlinear influence on survival fractions. The estimation process was stabilised by using of a square root transformation on dose. LD50 and LD10 values were subsequently estimated from the model and 95% confidence intervals were obtained using a parametric bootstrap approach [<xref rid="B57" ref-type="bibr">57</xref>, Chapter 5.4]. Details on model fitting and the estimation of the confidence intervals as well as code to derive LD values from survival count data with one or two strains can be found in the supplementary material (see Figure S4 in Supplementary Material available online at <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1155/2014/914767">http://dx.doi.org/10.1155/2014/914767</ext-link>). All computations were done with R [<xref rid="B58" ref-type="bibr">58</xref>] using the R software packages <monospace> mgcv</monospace> [<xref rid="B57" ref-type="bibr">57</xref>] and <monospace> lethal</monospace> [<xref rid="B59" ref-type="bibr">59</xref>].</p></sec></sec><sec id="sec3"><title>3. Results and Discussion</title><sec id="sec3.1"><title>3.1. Morphological and Biochemical Characteristics</title><p>Cells of strain G18<sup>T</sup> were pleiomorphic and Gram-reaction-positive. Individual cells and aggregates were observed, confirming reports by Ishiguro and Wolfe [<xref rid="B53" ref-type="bibr">53</xref>] of synchronous morphogenesis on unspecific media and previous observations on other representatives of the genus<italic> Geodermatophilus</italic> [<xref rid="B27" ref-type="bibr">27</xref>]. In line with the original description by Luedemann [<xref rid="B3" ref-type="bibr">3</xref>], circular or elliptical motile zoospores and septated filaments from zoospore germination were observed (<xref ref-type="fig" rid="fig1">Figure 1</xref>). Young colonies were light-red in colour and turned greenish-black at maturity. Similar colours conversions were already observed by Nie et al. [<xref rid="B17" ref-type="bibr">17</xref>] and Montero-Calasanz et al. [<xref rid="B18" ref-type="bibr">18</xref>, <xref rid="B19" ref-type="bibr">19</xref>, <xref rid="B21" ref-type="bibr">21</xref>, <xref rid="B22" ref-type="bibr">22</xref>, <xref rid="B25" ref-type="bibr">25</xref>] for type strains of other representatives of the genus, such as<italic> G. nigrescens</italic>,<italic> G. arenarius, G. siccatus, G. saharensis, G. tzadiensis, </italic>and<italic> G. normandii</italic>, when cultivated under the same growth conditions (<xref ref-type="table" rid="tab1">Table 1</xref>). Colonies were convex, nearly circular and opaque with a moist surface and an entire margin. Strain G18<sup>T</sup> grew well on GYM<italic> Streptomyces</italic> and GEO media but did not grow on TSA, R2A, GPHF, PYGV, and Luedemann media. It grew best at 25–30°C but did not grow below 15°C or above 35°C. Growth was observed in presence of 1% NaCl and between pH 5.5–9.5 (optimal range pH 7.0–9.5). Results from phenotype microarray analysis are shown as a heatmap in the supplementary material (Figure S1) in comparison to the reference type strains of the genus<italic> Geodermatophilus. </italic>A summary of selected differential phenotypic characteristics is presented in <xref ref-type="table" rid="tab1">Table 1</xref>. In the phenotypic clustering significant support (>95%) is obtained for<italic> G. poikilotrophi</italic> DSM 44209<sup>T</sup>,<italic> G. nigrescens</italic> DSM 45408<sup>T</sup> and<italic> G. normandii</italic> DSM 45417<sup>T</sup> being most similar to each other regarding the characters present in GEN III Microplates (Suppl. Figure S2).</p></sec><sec id="sec3.2"><title>3.2. Chemotaxonomic Characteristics</title><p>Analysis of cell wall components revealed the presence of<italic> DL</italic>-diaminopimelic acid (cell wall type III), which is consistent with other species of the genus<italic> Geodermatophilus</italic> [<xref rid="B27" ref-type="bibr">27</xref>, <xref rid="B38" ref-type="bibr">38</xref>]. Strain G18<sup>T</sup> displayed primarily menaquinone MK-9(H<sub>4</sub>) (82.5%), in agreement with values reported for the family<italic> Geodermatophilaceae</italic> [<xref rid="B2" ref-type="bibr">2</xref>], but also MK-9(H<sub>0</sub>) (8.8%) and MK-9(H<sub>2</sub>) (4.8%). Major fatty acids were iso-C<sub>16:0</sub> (24.5 ± 0.2%), iso-C<sub>15:0</sub> (16.6 ± 1.3%), C<sub>17:1</sub><italic>ω</italic>8c (13.9 ± 0.1%) and C<sub>16:1</sub><italic>ω</italic>7c (8.3 ± 0.1%), complemented by iso-C<sub>16:1</sub> H (5.6 ± 0.9%), anteiso-C<sub>15:0</sub> (4.1 ± 0.4%), anteiso-C<sub>17:0</sub> (4.4 ± 0.2%), C<sub>18:1</sub><italic>ω</italic>9c (3.6 ± 0.1%) and C<sub>16:0</sub> (2.4 ± 0.9%). The phospholipids pattern consisted of phosphatidylethanolamine (PE), phosphatidylcholine (PC), phosphatidylinositol (PI), and small amount of diphosphatidylglycerol (DPG) in accordance with profiles obtained for representatives of other<italic> Geodermatophilus</italic> species investigated in this study (<xref ref-type="table" rid="tab1">Table 1</xref>). Phosphatidylglycerol was not detectable (see Supplementary Figure S3). This fact was already predictable based on phospholipids profiles displayed for other representatives of the genus such as<italic> G. arenarius, G. siccatus, G. tzadiensis, G. normandii, </italic>or<italic> G. amargosae</italic>, whose phosphatidylglycerol amounts were nearly imperceptible. Whole-cell sugar analysis revealed galactose as the diagnostic sugar [<xref rid="B38" ref-type="bibr">38</xref>] but also glucose and ribose. Genomic G + C content was 74.4 mol%.</p></sec><sec id="sec3.3"><title>3.3. Molecular Analysis</title><p>The almost complete (1514 bp) 16S rRNA gene sequence of strain G18<sup>T</sup> was determined. The 16S rRNA sequence showed the highest degree of similarity with the type strains of<italic> G. siccatus</italic> (99.1%),<italic> G. africanus </italic>(99.0%)<italic>, G. amargosae</italic> (98.5%),<italic> G. normandii</italic> (98.4%),<italic> G. obscurus </italic>(98.3%),<italic> G. tzadiensis</italic> (98.2%),<italic> G. nigrescens </italic>(98.1%),<italic> G. ruber</italic> (98.0%), and<italic> G. arenarius</italic> (98.0%). All listed closely related type strains were placed within the same phylogenetic group by both, maximum likelihood and maximum-parsimony estimations (<xref ref-type="fig" rid="fig2">Figure 2</xref>). The 16S rRNA gene sequences analysis thus strongly supports the assignment of strain G18<sup>T</sup> to the genus<italic> Geodermatophilus. </italic>However, similarities in 16S rRNA gene sequence between G18<sup>T</sup> and some closely related type strains indicated the need to prove the genomic distinctness of the type strain representing the novel species by DNA-DNA hybridization. Strain G18<sup>T</sup> displayed a DNA-DNA relatedness of 35.3 ± 1.0% with the type strain of<italic> G. siccatus </italic>and 28.1 ± 2.1% with<italic> G. africanus.</italic> DNA-DNA hybridizations of strain G18<sup>T</sup> with the type strains of<italic> G. amargosae</italic>,<italic> G. normandii</italic>,<italic> G. obscurus</italic>,<italic> G. tzadiensis</italic>,<italic> G. nigrescens</italic>,<italic> G. ruber,</italic> and<italic> G. arenarius</italic> were not conducted, according to Meier-Kolthoff et al. [<xref rid="B49" ref-type="bibr">49</xref>] that statistically confirmed that the threshold value previously established at 97% 16S rRNA gene sequence similarity was too conservative in microbial species discrimination and determined a<italic> Actinobacteria</italic>-specific 16S rRNA threshold at 99.0% with a maximun probability of error of 1.00% to get DNA-DNA hybridization values above the 70% threshold recommended by Wayne et al. [<xref rid="B60" ref-type="bibr">60</xref>] to confirm the species status of novel strains.</p></sec><sec id="sec3.4"><title>3.4. Tolerance</title><p>Gamma-radiation survival of strain G18<sup>T</sup> (<xref ref-type="fig" rid="fig3">Figure 3(a)</xref>) showed not significantly different inactivation kinetic as for<italic> G. obscurus</italic> DSM 43160<sup>T</sup>, which is considered as highly resistant, according to data reported by Gtari et al. [<xref rid="B11" ref-type="bibr">11</xref>]. Strain G18<sup>T</sup> strains exhibited a shoulder of resistance similar to<italic> D. radiodurans</italic> R1 to approximately 5 KGy [<xref rid="B61" ref-type="bibr">61</xref>], but comparatively lower than the observed one by<italic> G. obscurus</italic> DSM 43160<sup>T</sup>. Nevertheless, LD10 of both, G18<sup>T</sup> and<italic> G. obscurus</italic> DSM 43160<sup>T</sup>, was around 9 KGy, a dose comparatively higher than the displayed one for the high radiation resistant strain<italic> D. radiodurans</italic> R1 [<xref rid="B61" ref-type="bibr">61</xref>], although other authors reported a LD10 around 10 KGy by using the same strain [<xref rid="B62" ref-type="bibr">62</xref>]. UV-radiation survival curves revealed a similar progressive loss of viability in both strains during the first 10 min of exposure until levels below 50%. However, the differences between the two resistant phenotypes increased along the curve, observing a significant variation on viability at 10% survival (<xref ref-type="fig" rid="fig3">Figure 3(b)</xref>). According to radiated doses, strain G18<sup>T</sup> and<italic> G. obscurus</italic> DSM 43160<sup>T</sup> were capable to support the lethal effects of 6300–12600 J<italic>·</italic>s<sup>−1</sup> m<sup>2</sup> and 63600–31800 J<italic>·</italic>s<sup>−1</sup> m<sup>2</sup>, respectively, sustaining a survival rate higher than 10%. Battista [<xref rid="B63" ref-type="bibr">63</xref>] and Shukla et al. [<xref rid="B62" ref-type="bibr">62</xref>] reported LD10 values of 700–1000 J<italic>·</italic>s<sup>−1</sup> m<sup>2</sup> for the highly resistant<italic> D. radiodurans </italic>R1. The tolerance to UV-radiation in the genus<italic> Geodermatophilus</italic> was already observed, in addition to<italic> G. obscurus </italic>DSM 43160<sup>T</sup>, in<italic> G. tzadiensis</italic> DSM 45416<sup>T</sup> by Montero-Calasanz et al. [<xref rid="B22" ref-type="bibr">22</xref>]. Cultures of strain G18<sup>T</sup> tolerated an exposure to mitomycin of nearly 120 min showing a viability rate of 10%, a value significantly higher than the one observed for the positive control (LD10 = 71 min) (<xref ref-type="fig" rid="fig3">Figure 3(c)</xref>). Tolerance of strain G18<sup>T</sup> (LD10 = 7 min) in comparison with the positive control<italic> G. obscurus </italic>DSM 43160<sup>T</sup> (LD10 = 8 min) to 0.5% hydrogen peroxide along the curves did not show any significant differences (<xref ref-type="fig" rid="fig3">Figure 3(d)</xref>). Based on desiccation survival curves given in <xref ref-type="fig" rid="fig3">Figure 3(e)</xref>, both strains initially exhibited a similar resistance (LD50). At the first sample point (20 days), strain G18<sup>T</sup> showed a survival of less than 10%, a value comparatively different to the results observed by<italic> G. obscurus </italic>DSM 43160<sup>T</sup>, whose LD10 is reached after 38 days. However, it is worth mentioning that after 110 days a remaining bacterial population of strain G18<sup>T</sup> was still observed. Strain G18<sup>T</sup> demonstrated thus a high tolerance to ROS-generating stresses gamma- and UV-radiation, mitomycin C, hydrogen peroxide, and desiccation comparable to the positive control<italic> G. obscurus</italic> DSM 43160<sup>T</sup> and, in general terms, to DNA damaging-resistant<italic> D. radiodurans</italic> R1. This correlative tolerance between ROS-generating stresses was already widely described [<xref rid="B11" ref-type="bibr">11</xref>, <xref rid="B62" ref-type="bibr">62</xref>] and support the hypothesis of efficient and common cellular DNA repair mechanisms. Strain G18<sup>T</sup> showed the highest tolerance to AsO<sub>4</sub><sup>3−</sup> (MIC = 8.0 mM) followed by Pb<sup>2+</sup> (MIC = 4.0 mM), CrO<sub>4</sub><sup>2−</sup> (MIC = 4.0 mM) and Ag<sup>1+</sup> (MIC = 1.0 mM). Whereas the growth of<italic> G. obscurus</italic> DSM 43160<sup>T</sup> was mainly inhibited by concentrations below 1.0 mM, except AsO<sub>4</sub><sup>3−</sup> whose sensitivity was 10 times higher (MIC = 80.0 mM) than the one observed for strain G18<sup>T</sup> (<xref ref-type="table" rid="tab2">Table 2</xref>). It has been widely described that the heavy metals/metalloids exposure also produces ROS generation [<xref rid="B64" ref-type="bibr">64</xref>]. In this study, a correspondence with other ROS-generating stresses was not observed, in agreement with data reported by Gtari et al. [<xref rid="B11" ref-type="bibr">11</xref>] for<italic> G. obscurus </italic>DSM 43160<sup>T</sup>, but also for<italic> Modestobacter multiseptatus </italic>BC501 and<italic> Blastococcus saxobsidens </italic>DD2, suggesting the presence of alternative mechanisms to counteract the heavy metals/metalloids stress, such as transport outside the cells [<xref rid="B65" ref-type="bibr">65</xref>], adsorption on exocellular structures such as melanin [<xref rid="B66" ref-type="bibr">66</xref>], or enzymatic reduction to less toxic forms [<xref rid="B67" ref-type="bibr">67</xref>, <xref rid="B68" ref-type="bibr">68</xref>]. Although it is noteworthy that toxicity levels of lead and copper in<italic> G. obscurus</italic> DSM 43160<sup>T</sup> by comparison with the results displayed by Gtari et al. [<xref rid="B11" ref-type="bibr">11</xref>] were much different from each other. These divergences in the levels of tolerance might be due to the differences in the media compositions [<xref rid="B69" ref-type="bibr">69</xref>]. In addition, it was confirmed that neither phosphate buffer nor carbon source concentration present in GYM<italic> Streptomyces</italic> medium caused an overestimated metals tolerance of strains, justified by the different tolerance range found in both strains and its mostly correlation with the results described by Gtari et al. [<xref rid="B11" ref-type="bibr">11</xref>].</p><p>Apart from the phylogenetic analysis based on 16S rRNA gene sequences, several phenotypic features support the distinctiveness of strain G18<sup>T</sup> from representatives of all other<italic> Geodermatophilus</italic> species (<xref ref-type="table" rid="tab1">Table 1</xref>). Based on the phenotypic and genotypic data presented, we propose that strain G18<sup>T</sup> represents a novel species within the genus<italic> Geodermatophilus</italic>, with the name<italic> Geodermatophilus poikilotrophi </italic>sp. nov.</p><p><italic>Description of Geodermatophilus poikilotrophi sp. nov</italic>..<italic> Geodermatophilus poikilotrophi</italic> (poi.kil.o.troph'i N. L. fem. gen. n.<italic> poikilotrophi</italic> referring to a bacterium that can tolerate diverse environmental stresses). </p><p>Colonies are greenish-black-coloured, circular, and convex with a moist surface. Cells are Gram-reaction-positive, catalase positive, and oxidase negative. No diffusible pigments are produced on any of the tested media. Utilizes dextrin, D-maltose, D-trehalose, D-cellobiose, sucrose, stachyose, D-glucose, D-mannose, D-fructose, D-galactose, L-rhamnose, D-sorbitol, D-mannnitol, myo-inositol, glycerol, L-arginine, pectin, D-gluconic acid, quinic acid, methyl pyruvate, D-lactic acid methyl ester, <italic>α</italic>-ketoglutaric acid, D-malic acid, bromosuccinic acid, potassium tellurite, <italic>ϒ</italic>-amino-N-butyric acid, acetoacetic acid, propionic acid, acetic acid, as sole carbon source for energy and growth, but not turanose, D-raffinose, D-melibiose, <italic>β</italic>-methyl-D-glucoside, D-salicin,<italic> N</italic>-acetyl-D-glucosamine,<italic> N</italic>-acetyl-D-galactosamine,<italic> N</italic>-acetylneuraminic acid, 3-<italic>O</italic>-methyl-D-glucose, D-fucose, inosine, sodium lactate, D- and L-serine, D-arabitol, D-glucose-6-phosphate, D-aspartic acid, glycyl-L-proline, L-alanine, L-glutamic acid, L-histidine, L-pyroglutamic acid, L-galactonic acid-<italic>γ</italic>-lactone, glucuronamide, mucic acid, D-saccharic acid,<italic> p</italic>-hydroxyphenylacetic acid, citric acid, <italic>γ</italic>-amino-n-butyric acid, and butyric acid. Acid is produced from L-arginine and <italic>ϒ</italic>-amino-N-butyric acid and can be used as a sole nitrogen source but not<italic> N</italic>-acetyl-D-glucosamine,<italic> N</italic>-acetyl-D-galactosamine,<italic> N</italic>-acetyl-neuraminic acid, D- and L-serine, D-aspartic acid, glycyl-L-proline, L-alanine, L-histidine, L-glutamic acid, L-histidine, L-pyroglutamic acid, glucuronamide, and <italic>γ</italic>-amino-n-butyric acid. Positive for aesculin degradation. Negative for reduction of nitrate, denitrification, indole production and gelatin degradation. Tests for alkaline phosphatase, esterase lipase (C8), esterase (C4), leucine arylamidase and <italic>α</italic>-glucosidase are positive, but those for urease, <italic>β</italic>-glucosidase, acid phosphatase, valine arylamidase, Naphthol-AS-BI-phosphohydrolase, lipase (C14), cystine arylamidase, trypsin, <italic>α</italic>-chymotrypsin, <italic>α</italic>- and <italic>β</italic>-galactosidase, <italic>β</italic>-glucuronidase, N-acetyl-<italic>β</italic>-glucosamidase, <italic>α</italic>-mannosidase and <italic>α</italic>-fucosidase are negative. Cell growth ranges from 15 to 35°C and from pH 5.5 to 9.5. It is tolerant to gamma- and UV-radiation, mitomycin C, hydrogen peroxide, desiccation and heavy metals/metalloids AsO<sub>4</sub><sup>3−</sup>, Pb<sup>2+</sup>, CrO<sub>4</sub><sup>2−</sup> and Ag<sup>1+</sup>. The peptidoglycan in the cell wall contains<italic> meso</italic>-diaminopimelic acid as diamino acid, with galactose as the diagnostic sugar. The predominant menaquinone is MK-9(H<sub>4</sub>). The main polar lipids are phosphatidylethanolamine, phosphatidylcholine, phosphatidylinositol, and small amount of diphosphatidylglycerol. Cellular fatty acids consist mainly of iso-C<sub>16:0</sub>, iso-C<sub>15:0</sub>, C<sub>17:1</sub><italic>ω</italic>8c, and C<sub>16:1</sub><italic>ω</italic>7c. The type strain has a genomic DNA G + C content of 74.4 mol %. The INSDC accession number for the 16S rRNA gene sequences of the type strain G18<sup>T</sup> (= DSM 44209<sup>T</sup> = CCUG 63018<sup>T</sup>) is HF970583.</p></sec></sec><sec sec-type="supplementary-material" id="supplementary-material-sec"><title>Supplementary Material</title><supplementary-material content-type="local-data" id="f1"><caption><p>Supplementary Fig. S1: The parameter “Maximum Height” estimated from the respiration curves as measured with the OmniLog phenotyping device and discretized and visualized as heatmap using the opm package. Plates and substrates are rearranged according to their overall similarity (as depicted using the row and column dendrograms). Ochre colour indicates positive reaction; blue colour indicates negative reaction; white colour indicates ambiguous reaction. Letters (A/B) indicate each replicate of experiment.</p><p>Supplementary Fig. S2: Phenotypic dendrogram based on the parameter “Maximun Height” estimated from the respiration curves as measured with the OmniLog phenotyping device using Ward algorithm for agglomerative hierarchical clustering and correlation coefficient as a distance metric using the pvclust package. Support values approximately unbiased (AU, left) and bootstrapping (BP, right) are shown above the branches. Well supported clusters are defined by red squares. </p><p>Supplementary Fig. S3: Polar lipids profile of Geodermatophilus poikilotrophi sp. nov. G18T, after separation by two-dimensional TLC. Plate was sprayed with molydatophosphoric acid for detection of total polar lipid. DPG, diphosphadidylglycerol; PE, phosphatidylethanolamine; PC, phosphatidylcholine; PI, phosphatidylinositol; GL, unknown glycolipid.
Supplementary R-code File. S4. Exemplary code to fit the model and compute the confidence intervals using the R package lethal.
</p></caption><media xlink:href="914767.f1.pdf" mimetype="application" mime-subtype="pdf" orientation="portrait" id="d35e1635" position="anchor"></media></supplementary-material></sec></body><back><ack><title>Acknowledgments</title><p>The authors would like to acknowledge the help of Bettina Sträubler and Birgit Grün for DNA-DNA hybridization analyses, Gabi Pötter for assistance with chemotaxonomy, Brian J. Tindall (all at DSMZ, Germany) for his guidance in the chemotaxonomic analyses, and Haitham Sghaier (CNSTN, Tunisia) for providing access to the gamma irradiation facility. 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person-group-type="author"><name><surname>Rathnayake</surname><given-names>IVN</given-names></name><name><surname>Megharaj</surname><given-names>M</given-names></name><name><surname>Krishnamurti</surname><given-names>GSR</given-names></name><name><surname>Bolan</surname><given-names>NS</given-names></name><name><surname>Naidu</surname><given-names>R</given-names></name></person-group><article-title>Heavy metal toxicity to bacteria: are the existing growth media accurate enough to determine heavy metal toxicity?</article-title><source><italic>Chemosphere</italic></source><year>2013</year><volume>90</volume><issue>3</issue><fpage>1195</fpage><lpage>1200</lpage><pub-id pub-id-type="other">2-s2.0-84869875791</pub-id><pub-id pub-id-type="pmid">23040649</pub-id></element-citation></ref></ref-list></back><floats-group><fig id="fig1" orientation="portrait" position="float"><label>Figure 1</label><caption><p>Scanning electron micrograph of strain G18<sup>T</sup> grown on GYM<italic> Streptomyces</italic> medium for 7 days at 28°C.</p></caption><graphic xlink:href="BMRI2014-914767.001"></graphic></fig><fig id="fig2" orientation="portrait" position="float"><label>Figure 2</label><caption><p>Maximum likelihood phylogenetic tree inferred from 16S rRNA gene sequences, showing the phylogenetic position of strain G18<sup>T</sup> relative to the type strains within the family cursive. The branches are scaled in terms of the expected number of substitutions per site (see size bar). Support values from maximum-likelihood (left) and maximum-parsimony (right) bootstrapping are shown above the branches if equal to or larger than 60%.</p></caption><graphic xlink:href="BMRI2014-914767.002"></graphic></fig><fig id="fig3" orientation="portrait" position="float"><label>Figure 3</label><caption><p>Estimation of survival following exposure to gamma-radiation (a), UV-radiation (b), mitomycin C (c), hydrogen peroxide (d), and desiccation (e) for strain G18<sup>T</sup> and<italic> G. obscurus</italic> DSM 43160<sup>T</sup> as positive control. The mean c.f.u.mL<sup>−1</sup> per strain is given together with the LD50 and LD10 values in the upper panel of each figure; <italic>y</italic>-axis is on a logarithmic scale ((a)–(c), (e)), or on a square root scale (d). The lower panel depicts LD10 and LD50 values per strain and the differences between strains together with confidence intervals. Confidence intervals that do not contain zero (dashed vertical line) indicate significant differences to zero; in case of strain differences this indicates significant differences between strains.</p></caption><graphic xlink:href="BMRI2014-914767.003"></graphic></fig><table-wrap id="tab1" orientation="portrait" position="float"><label>Table 1</label><caption><p>Differential phenotypic characteristics of strain G18<sup>T</sup> and the type strains of other <italic>Geodermatophilus</italic> species. Strains: 1, <italic>G. poikilotrophi</italic> sp. nov. G18<sup>T</sup>; 2, <italic>G. obscurus</italic> DSM 43160<sup>T</sup>; 3, <italic>G. ruber</italic> DSM 45317<sup>T</sup>; 4, <italic>G. nigrescens</italic> DSM 45408<sup>T</sup>; 5, <italic>G. arenarius</italic> DSM 45418<sup>T</sup>; 6, <italic>G. siccatus</italic> DSM 45419<sup>T</sup>; 7, <italic>G. saharensis </italic>DSM 45423<sup>T</sup>; 8, <italic>G. tzadiensis</italic> DSM 45416<sup>T</sup>; 9, <italic>G. telluris</italic> DSM 45421<sup>T</sup>; 10, <italic>G. soli</italic> DSM 45843<sup>T</sup>; 11, <italic>G. terrae </italic>DSM 45844<sup>T</sup>; 12, <italic>G. africanus</italic> DSM 45422<sup>T</sup>; 13, <italic>G. normandii</italic> DSM 45417<sup>T</sup>; 14, <italic>G. taihuensis</italic> DSM 45962<sup>T</sup>; 15, <italic>G. amargosae</italic> DSM 46136<sup>T</sup>. All physiological data are from this study.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" rowspan="1" colspan="1">Characteristics</th><th align="center" rowspan="1" colspan="1">1</th><th align="center" rowspan="1" colspan="1">2</th><th align="center" rowspan="1" colspan="1">3</th><th align="center" rowspan="1" colspan="1">4</th><th align="center" rowspan="1" colspan="1">5</th><th align="center" rowspan="1" colspan="1">6</th><th align="center" rowspan="1" colspan="1">7</th><th align="center" rowspan="1" colspan="1">8</th><th align="center" rowspan="1" colspan="1">9</th><th align="center" rowspan="1" colspan="1">10</th><th align="center" rowspan="1" colspan="1">11</th><th align="center" rowspan="1" colspan="1">12</th><th align="center" rowspan="1" colspan="1">13</th><th align="center" rowspan="1" colspan="1">14</th><th align="center" rowspan="1" colspan="1">15</th></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1">Colony colour on GYM</td><td align="center" rowspan="1" colspan="1">Light-red, greenish-black</td><td align="center" rowspan="1" colspan="1">Black</td><td align="center" rowspan="1" colspan="1">Light-red, red</td><td align="center" rowspan="1" colspan="1">Light-red, black</td><td align="center" rowspan="1" colspan="1">Light-red, brown</td><td align="center" rowspan="1" colspan="1">Light-red, black</td><td align="center" rowspan="1" colspan="1">Light-red, black</td><td align="center" rowspan="1" colspan="1">Light-red, greenish-black</td><td align="center" rowspan="1" colspan="1">Black</td><td align="center" rowspan="1" colspan="1">Light red</td><td align="center" rowspan="1" colspan="1">Light red</td><td align="center" rowspan="1" colspan="1">Black</td><td align="center" rowspan="1" colspan="1">Light-red, greenish-black</td><td align="center" rowspan="1" colspan="1">Coral pink</td><td align="center" rowspan="1" colspan="1">Black</td></tr><tr><td align="left" rowspan="1" colspan="1">Colony surface on GYM</td><td align="center" rowspan="1" colspan="1">Moist</td><td align="center" rowspan="1" colspan="1">Dry</td><td align="center" rowspan="1" colspan="1">Moist</td><td align="center" rowspan="1" colspan="1">Moist</td><td align="center" rowspan="1" colspan="1">Moist</td><td align="center" rowspan="1" colspan="1">Moist</td><td align="center" rowspan="1" colspan="1">Moist</td><td align="center" rowspan="1" colspan="1">Moist</td><td align="center" rowspan="1" colspan="1">Dry</td><td align="center" rowspan="1" colspan="1">Moist</td><td align="center" rowspan="1" colspan="1">Moist</td><td align="center" rowspan="1" colspan="1">Dry</td><td align="center" rowspan="1" colspan="1">Moist</td><td align="center" rowspan="1" colspan="1">Moist</td><td align="center" rowspan="1" colspan="1">Dry</td></tr><tr><td align="left" rowspan="1" colspan="1">Utilization of</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" rowspan="1" colspan="1"> Turanose</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td></tr><tr><td align="left" rowspan="1" colspan="1"> Stachyose</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">−</td></tr><tr><td align="left" rowspan="1" colspan="1"> D-Melibiose</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td></tr><tr><td align="left" rowspan="1" colspan="1"> D-Salicin</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td></tr><tr><td align="left" rowspan="1" colspan="1">NaCl range (w/v)</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" rowspan="1" colspan="1"> 1%</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+/−</td></tr><tr><td align="left" rowspan="1" colspan="1"> 4% </td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+/−</td></tr><tr><td align="left" rowspan="1" colspan="1"> D-Mannose</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+/−</td></tr><tr><td align="left" rowspan="1" colspan="1"> L-Rhamnose</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td></tr><tr><td align="left" rowspan="1" colspan="1"> Inosine</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td></tr><tr><td align="left" rowspan="1" colspan="1"> D-Sorbitol</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">+/−</td></tr><tr><td align="left" rowspan="1" colspan="1"> D-Mannitol</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+/−</td></tr><tr><td align="left" rowspan="1" colspan="1"> D-Arabitol</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td></tr><tr><td align="left" rowspan="1" colspan="1"> Glycerol</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td></tr><tr><td align="left" rowspan="1" colspan="1"> L-Alanine</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+/−</td><td align="center" rowspan="1" colspan="1">+</td></tr><tr><td align="left" rowspan="1" colspan="1"> L-Arginine</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td></tr><tr><td align="left" rowspan="1" colspan="1"> L-Histidine</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+/−</td></tr><tr><td align="left" rowspan="1" colspan="1"> Pectin</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td></tr><tr><td align="left" rowspan="1" colspan="1"> D-Gluconic acid</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td></tr><tr><td align="left" rowspan="1" colspan="1"> Quinic acid</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">−</td><td align="center" rowspan="1" colspan="1">+</td><td align="center" rowspan="1" colspan="1">−</td></tr><tr><td align="left" rowspan="1" colspan="1">Predominant menaquinone(s)<sup>a</sup></td><td align="center" rowspan="1" colspan="1">MK-9(H<sub>4</sub>)</td><td align="center" rowspan="1" colspan="1">MK-9(H<sub>4</sub>), MK-9(H<sub>2</sub>), MK-8(H<sub>4</sub>)</td><td align="center" rowspan="1" colspan="1">MK-9(H<sub>4</sub>)</td><td align="center" rowspan="1" colspan="1">MK-9(H<sub>4</sub>)</td><td align="center" rowspan="1" colspan="1">MK-9(H<sub>4</sub>), MK-8(H<sub>4</sub>), MK-9 (H<sub>0</sub>)</td><td align="center" rowspan="1" colspan="1">MK-9(H<sub>4</sub>), MK-8(H<sub>4</sub>), MK-9(H<sub>0</sub>)</td><td align="center" rowspan="1" colspan="1">MK-9(H<sub>4</sub>), MK-8(H<sub>4</sub>)</td><td align="center" rowspan="1" colspan="1">MK-9(H<sub>4</sub>), MK-9(H<sub>0</sub>)</td><td align="center" rowspan="1" colspan="1">MK-9(H<sub>4</sub>)</td><td align="center" rowspan="1" colspan="1">MK-9(H<sub>4</sub>), MK-9(H<sub>0</sub>), MK-9(H<sub>2</sub>)</td><td align="center" rowspan="1" colspan="1">MK-9(H<sub>4</sub>), MK-9(H<sub>0</sub>)</td><td align="center" rowspan="1" colspan="1">MK-9(H<sub>4</sub>)</td><td align="center" rowspan="1" colspan="1">MK-9(H<sub>4</sub>)</td><td align="center" rowspan="1" colspan="1">MK-9 (H<sub>4</sub>), MK-9(H<sub>0</sub>)<sup>#</sup></td><td align="center" rowspan="1" colspan="1">MK-9(H<sub>4</sub>)</td></tr><tr><td align="left" rowspan="1" colspan="1">Phospholipids∗</td><td align="center" rowspan="1" colspan="1">PE, PC, PI, DPG</td><td align="center" rowspan="1" colspan="1">DPG, PC, PE, PI, PG</td><td align="center" rowspan="1" colspan="1">DPG, PE, PC, PI, 2PL, PG</td><td align="center" rowspan="1" colspan="1">DPG, PE, PC, PI, PG</td><td align="center" rowspan="1" colspan="1">PE, PC, DPG, PI, PG</td><td align="center" rowspan="1" colspan="1">PE, PC, PI, DPG, PG</td><td align="center" rowspan="1" colspan="1">DPG, PC, PI, PE, PG</td><td align="center" rowspan="1" colspan="1">DPG, PC, PE, PI, PG</td><td align="center" rowspan="1" colspan="1">DPG, PC, PE, PI, APL, PG</td><td align="center" rowspan="1" colspan="1">DPG, PME, PE, PI, 3PL<sup>†</sup></td><td align="center" rowspan="1" colspan="1">DPG, PME, PE, PI, 5PL<sup>†</sup></td><td align="center" rowspan="1" colspan="1">DPG, PC, PE, PI, PG</td><td align="center" rowspan="1" colspan="1">DPG, PC, PE, PI, PG</td><td align="center" rowspan="1" colspan="1">DPG, PE, PI, PIM<sup>#</sup></td><td align="center" rowspan="1" colspan="1">DPG, PC, PE, PI, PG</td></tr><tr><td align="left" rowspan="1" colspan="1">Major fatty acids<sup>b</sup></td><td align="center" rowspan="1" colspan="1">i-C<sub>15:0</sub>, <break></break>i-C<sub>16:0</sub>, <break></break>C<sub>17:1</sub><italic>ω</italic>8c</td><td align="center" rowspan="1" colspan="1">i-C<sub>15:0</sub>, <break></break>i-C<sub>16:0</sub>, <break></break>C<sub>17:1</sub><italic>ω</italic>8c</td><td align="center" rowspan="1" colspan="1">i-C<sub>15:0</sub>, <break></break>ai-C<sub>15:0</sub>, <break></break>i-C<sub>16:0</sub>, <break></break>ai-C<sub>17:0</sub>, <break></break>C<sub>17:1</sub><italic>ω</italic>8c</td><td align="center" rowspan="1" colspan="1">i-C<sub>15:0</sub>, <break></break>i-C<sub>16:0</sub></td><td align="center" rowspan="1" colspan="1">i-C<sub>15:0</sub>, <break></break>i-C<sub>16:0</sub></td><td align="center" rowspan="1" colspan="1">i-C<sub>15:0</sub>, <break></break>i-C<sub>16:0</sub>, <break></break>C<sub>17:1</sub><italic>ω</italic>8c</td><td align="center" rowspan="1" colspan="1">i-C<sub>15:0</sub>, <break></break>i-C<sub>16:0</sub>, <break></break>i-H-C<sub>16:1</sub></td><td align="center" rowspan="1" colspan="1">i-C<sub>15:0</sub>, <break></break>i-C<sub>16:0</sub></td><td align="center" rowspan="1" colspan="1">i-C<sub>15:0</sub>, <break></break>i-C<sub>16:0</sub></td><td align="center" rowspan="1" colspan="1">i-C<sub>15:0</sub>, <break></break>i-C<sub>16:0</sub>, <break></break>C<sub>17:0</sub></td><td align="center" rowspan="1" colspan="1">i-C<sub>15:0</sub>, <break></break>i-C<sub>16:0</sub>, <break></break>C<sub>18:1</sub><italic>ω</italic>9c</td><td align="center" rowspan="1" colspan="1">i-C<sub>16:0</sub></td><td align="center" rowspan="1" colspan="1">i-C<sub>15:0</sub>, <break></break>i-C<sub>16:0</sub></td><td align="center" rowspan="1" colspan="1">i-C<sub>15:0</sub>, <break></break>i-C<sub>16:0</sub>, <break></break>C<sub>17:1</sub><italic>ω</italic>8c</td><td align="center" rowspan="1" colspan="1">i-C<sub>15:0</sub>, <break></break>i-C<sub>16:0</sub></td></tr></tbody></table><table-wrap-foot><fn><p>+, positive reaction; −, negative reaction; +/−, ambiguous; MK, menaquinones; DPG, diphosphatidylglycerol; PE, phosphatidylethanolamine; PME, phosphatidyl-N-methylethanolamine; PE-OH, hydroxyphosphatidylethanolamine; PG, phosphatidylglycerol; PC, phosphatidylcholine; PI, phosphatidylinositol; PIM, phosphatidylinositol mannoside; PL, unknown phospholipid; APL, unknown amino-phospholipid; i-, iso-branched; ai-, anteiso-branched.</p></fn><fn><p><sup>a</sup>Only components making up ≥ 5% peak area ratio are shown; <sup>b</sup>only components making up ≥ 10% peak area ratio are shown; ∗the components are listed in decreasing order of quantity.</p></fn><fn><p><sup>†</sup>Data taken from Jin et al. [<xref rid="B10" ref-type="bibr">10</xref>]. <sup>#</sup>Data taken from Qu et al. [<xref rid="B26" ref-type="bibr">26</xref>].</p></fn></table-wrap-foot></table-wrap><table-wrap id="tab2" orientation="portrait" position="float"><label>Table 2</label><caption><p>Minimum inhibitory concentration of seven heavy metals and metalloids for strain G18<sup>T</sup> and <italic>G. obscurus</italic> DSM 43160<sup>T</sup>. </p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" rowspan="2" colspan="1">Strain</th><th align="center" colspan="7" rowspan="1">MIC (mM) of</th></tr><tr><th align="center" rowspan="1" colspan="1">AgNO<sub>3</sub></th><th align="center" rowspan="1" colspan="1">CuCl<sub>2</sub></th><th align="center" rowspan="1" colspan="1">CoCl<sub>2</sub></th><th align="center" rowspan="1" colspan="1">NiCl<sub>2</sub></th><th align="center" rowspan="1" colspan="1">K<sub>2</sub>CrO<sub>4</sub></th><th align="center" rowspan="1" colspan="1">Pb(NO<sub>3</sub>)<sub>2</sub></th><th align="center" rowspan="1" colspan="1">NaHAsO<sub>4</sub></th></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1">G18<sup>T</sup></td><td align="center" rowspan="1" colspan="1">1.0</td><td align="center" rowspan="1" colspan="1">0.1</td><td align="center" rowspan="1" colspan="1">0.3</td><td align="center" rowspan="1" colspan="1">0.5</td><td align="center" rowspan="1" colspan="1">4.0</td><td align="center" rowspan="1" colspan="1">4.0</td><td align="center" rowspan="1" colspan="1">8.0</td></tr><tr><td align="left" rowspan="1" colspan="1">DSM 43160<sup>T</sup></td><td align="center" rowspan="1" colspan="1">0.3</td><td align="center" rowspan="1" colspan="1">0.1</td><td align="center" rowspan="1" colspan="1">0.3</td><td align="center" rowspan="1" colspan="1">0.3</td><td align="center" rowspan="1" colspan="1">1.0</td><td align="center" rowspan="1" colspan="1">1.0</td><td align="center" rowspan="1" colspan="1">80.0</td></tr></tbody></table></table-wrap></floats-group></pmc></record>Pour manipuler ce document sous Unix (Dilib)
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