<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD Journal Publishing DTD v3.0 20080202//EN" "http://dtd.nlm.nih.gov/publishing/3.0/journalpublishing3.dtd">
<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" article-type="research-article" dtd-version="3.0" xml:lang="en">
<front>
<journal-meta>
<journal-id journal-id-type="publisher">BG</journal-id>
<journal-title-group>
<journal-title>Biogeosciences</journal-title>
<abbrev-journal-title abbrev-type="publisher">BG</abbrev-journal-title>
</journal-title-group>
<issn pub-type="epub">1726-4189</issn>
<publisher><publisher-name>Copernicus GmbH</publisher-name>
<publisher-loc>Göttingen, Germany</publisher-loc>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.5194/bg-8-1465-2011</article-id>
<title-group>
<article-title>Experimental fossilisation of viruses from extremophilic Archaea</article-title>
</title-group>
<contrib-group><contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Orange</surname>
<given-names>F.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Chabin</surname>
<given-names>A.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Gorlas</surname>
<given-names>A.</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Lucas-Staat</surname>
<given-names>S.</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Geslin</surname>
<given-names>C.</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Le Romancer</surname>
<given-names>M.</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Prangishvili</surname>
<given-names>D.</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Forterre</surname>
<given-names>P.</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Westall</surname>
<given-names>F.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
</contrib>
</contrib-group><aff id="aff1">
<label>1</label>
<addr-line>Centre de Biophysique Moléculaire,  UPR4301, CNRS, Rue Charles Sadron, 45071 Orléans Cedex 2, France</addr-line>
</aff>
<aff id="aff2">
<label>2</label>
<addr-line>Observatoire des Sciences de l&apos;Univers en région Centre, UMS3116, 1A Rue de la Férollerie, 45071 Orléans Cedex 2, France</addr-line>
</aff>
<aff id="aff3">
<label>3</label>
<addr-line>Université de Bretagne Occidentale, UMR 6539, CNRS –  Institut Universitaire Européen de la Mer, Technopôle Brest-Iroise, Rue Dumont d&apos;Urville, 29280 Plouzané, France</addr-line>
</aff>
<aff id="aff4">
<label>4</label>
<addr-line>Molecular Biology of the Gene in Extremophiles Unit, Institut Pasteur, 25 rue du Docteur Roux, 75724 Paris Cedex 15, France</addr-line>
</aff>
<pub-date pub-type="epub">
<day>09</day>
<month>06</month>
<year>2011</year>
</pub-date>
<volume>8</volume>
<issue>6</issue>
<fpage>1465</fpage>
<lpage>1475</lpage>
<permissions>
<license xlink:type="simple">
<license-p>This is an open-access article ditributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
</license>
</permissions>
<self-uri xlink:href="http://www.biogeosciences.net/8/1465/2011/bg-8-1465-2011.html">This article is available from http://www.biogeosciences.net/8/1465/2011/bg-8-1465-2011.html</self-uri>
<self-uri xlink:href="http://www.biogeosciences.net/8/1465/2011/bg-8-1465-2011.pdf">The full text article is available as a PDF file from http://www.biogeosciences.net/8/1465/2011/bg-8-1465-2011.pdf</self-uri>
<abstract>
<p>The role of viruses at different stages of the origin of life has recently
been reconsidered. It appears that viruses may have accompanied the earliest
forms of life, allowing the transition from an RNA to a DNA world and
possibly being involved in the shaping of tree of life in the three domains
that we know presently. In addition, a large variety of viruses has been
recently identified in extreme environments, hosted by extremophilic
microorganisms, in ecosystems considered as analogues to those of the early
Earth. Traces of life on the early Earth were preserved by the precipitation
of silica on the organic structures. We present the results of the first
experimental fossilisation by silica of viruses from extremophilic Archaea
(SIRV2 – &lt;i&gt;Sulfolobus islandicus&lt;/i&gt; rod-shaped virus 2, TPV1 – &lt;i&gt;Thermococcus prieurii&lt;/i&gt; virus 1,
and PAV1 – &lt;i&gt;Pyrococcus abyssi&lt;/i&gt; virus 1). Our results
confirm that viruses can be fossilised, with silica precipitating on the
different viral structures (proteins, envelope) over several months in a
manner similar to that of other experimentally and naturally fossilised
microorganisms. This study thus suggests that viral remains or traces could
be preserved in the rock record although their identification may be
challenging due to the small size of the viral particles.</p>
</abstract>
<counts><page-count count="11"/></counts>
</article-meta>
</front>
<body/>
<back>
<ref-list>
<title>References</title>
<ref id="ref1">
<label>1</label><mixed-citation publication-type="other" xlink:type="simple"> Bamford, D. H.: Do viruses form lineages across different domains of life?, Res. Microbiol. 154, 231–236, http://dx.doi.org/10.1016/S0923-2508(03)00065-2doi:10.1016/S0923-2508(03)00065-2, 2003. </mixed-citation>
</ref>
<ref id="ref2">
<label>2</label><mixed-citation publication-type="other" xlink:type="simple"> Bamford, D. H., Grimes, J. M., and Stuart, D. I.: What does structure tell us about virus evolution?, Curr. Opin. Struc. Biol., 15, 655–663, http://dx.doi.org/10.1016/j.sbi.2005.10.012doi:10.1016/j.sbi.2005.10.012, 2005. </mixed-citation>
</ref>
<ref id="ref3">
<label>3</label><mixed-citation publication-type="other" xlink:type="simple"> Bettstetter, M., Peng, X., Garrett, R. A., and Prangishvili, D.: AFV1, a novel virus infecting hyperthermophilic archaea of the genus \textitAcidianus, Virology, 315, 68–79, 2003. </mixed-citation>
</ref>
<ref id="ref4">
<label>4</label><mixed-citation publication-type="other" xlink:type="simple"> Beveridge, T. J. and Koval, S. F.: Binding metals to cell envelopes of \textitEscherichia coli, Appl. Environ. Microb., 42, 325–335, 1981. </mixed-citation>
</ref>
<ref id="ref5">
<label>5</label><mixed-citation publication-type="other" xlink:type="simple"> Beveridge, T. J. and Murray, R. G. E.: Uptake and retention of metals by cell walls of \textitBacillus subtilis, J. Bacteriol., 127, 1502–1518, 1976. </mixed-citation>
</ref>
<ref id="ref6">
<label>6</label><mixed-citation publication-type="other" xlink:type="simple"> Bird, D. F., Juniper, S. K., Ricciardi-Rigault, M., Martineu, P., Prairie, Y. T., and Calvert, S. E.: Subsurface viruses and bacteria in Holocene/Late Pleistocene sediments of Saanich Inlet, BC: ODP Holes 1033B and 1034B, Leg 169S, Mar. Geol., 174, 227–239, http://dx.doi.org/10.1016/S0025-3227(00)00152-3doi:10.1016/S0025-3227(00)00152-3, 2001. </mixed-citation>
</ref>
<ref id="ref7">
<label>7</label><mixed-citation publication-type="other" xlink:type="simple"> Birnbaum, S. J., Wireman, J. W., and Borowski, R.: Silica precipitation by the anaerobic sulphate reducing bacterium \textitDesulfovibrio desulfuricans: effects upon cellmorphology and implications for preservation, in: Origin, Evolution, and Modern Aspects of Biomineralization in Plants and Animals, Crick, R.E., Plenum Press, New York, USA, 507–516, 1989. </mixed-citation>
</ref>
<ref id="ref8">
<label>8</label><mixed-citation publication-type="other" xlink:type="simple"> Bize, A., Karlsson, E. A., Ekefjärd, K., Quax, T. E. F., Pina, M., Prevost, M. C., Forterre, P., Tenaillon, O., Bernander, R., and Prangishvili, D.: A unique virus release in the Archaea, P. Natl. Acad. Sci. USA, 106, 11306–11311, http://dx.doi.org/10.1073/pnas.0901238106doi:10.1073/pnas.0901238106, 2009. </mixed-citation>
</ref>
<ref id="ref9">
<label>9</label><mixed-citation publication-type="other" xlink:type="simple"> Borriss, M., Helmke, E., Hanschke, R., and Schweder, T.: Isolation and characterization of marine psychrophilic phage–host systems from Arctic sea ice, Extremophiles, 7, 377–384, http://dx.doi.org/10.1007/s00792-003-0334-7doi:10.1007/s00792-003-0334-7, 2003. </mixed-citation>
</ref>
<ref id="ref10">
<label>10</label><mixed-citation publication-type="other" xlink:type="simple"> Daughney, C. J., Châtellier, X., Chan, A., Kenward, P., Fortin, D., Suttle, C. A., and Fowle, D. A.: Adsorption and precipitation of iron from seawater on a marine bacteriophage (PWH3A-P1), Mar. Chem., 91, 101–115, http://dx.doi.org/10.1016/j.marchem.2004.06.003doi:10.1016/j.marchem.2004.06.003, 2004. </mixed-citation>
</ref>
<ref id="ref11">
<label>11</label><mixed-citation publication-type="other" xlink:type="simple"> Dyall-Smith, M., Tang., S. L., and Bath, C.: Haloarchaeal viruses: how diverse are they?, Res. Microbiol., 154, 309–313, http://dx.doi.org/10.1016/S0923-2508(03)00076-7doi:10.1016/S0923-2508(03)00076-7, 2003. </mixed-citation>
</ref>
<ref id="ref12">
<label>12</label><mixed-citation publication-type="other" xlink:type="simple"> Erauso, G., Reysenbach, A. L., Godfroy, A., Meunier, J. R., Crump, B., Partensky, F., Baross, J. A., Marteinsson, V., Barbier, G., Pace, N. R., and Prieur, D.: \textitPyrococcus abyssi sp. nov., a new hyperthermophilic archaeon isolated from a deep-sea hydrothermal vent, Arch. Microbiol., 160, 338–349, 1993. </mixed-citation>
</ref>
<ref id="ref13">
<label>13</label><mixed-citation publication-type="other" xlink:type="simple"> Forterre, P.: A hot story from comparative genomics: reverse gyrase is the only hyperthermophile-specific protein, Trends Genet., 18, 236–238, http://dx.doi.org/10.1016/S0168-9525(02)02650-1doi:10.1016/S0168-9525(02)02650-1, 2002. </mixed-citation>
</ref>
<ref id="ref14">
<label>14</label><mixed-citation publication-type="other" xlink:type="simple"> Forterre, P.: Three RNA cells for ribosomal lineages and three DNA viruses to replicate their genomes : A hypothesis for the origin of cellular domain, P. Natl. Acad. Sci. USA, 103, 3669–3674, http://dx.doi.org/10.1073/pnas.0510333103doi:10.1073/pnas.0510333103, 2006. </mixed-citation>
</ref>
<ref id="ref15">
<label>15</label><mixed-citation publication-type="other" xlink:type="simple"> Forterre, P.: Manipulation of cellular syntheses and the nature of viruses: The virocell concept, C.R. Chim., 14, 392–399, http://dx.doi.org/10.1016/j.crci.2010.06.007doi:10.1016/j.crci.2010.06.007, 2011. </mixed-citation>
</ref>
<ref id="ref16">
<label>16</label><mixed-citation publication-type="other" xlink:type="simple"> Forterre, P. and Prangishvili, D.: The great billion-year war between ribosome- and capsid-encoding organisms (cells and viruses) as the major source of evolutionary novelties, Ann. NY. Acad. Sci., 1178, 65–77, http://dx.doi.org/10.1111/j.1749-6632.2009.04993.xdoi:10.1111/j.1749-6632.2009.04993.x, 2009a. </mixed-citation>
</ref>
<ref id="ref17">
<label>17</label><mixed-citation publication-type="other" xlink:type="simple"> Forterre, P. and Prangishvili, D.: The origin of viruses, Res. Microbiol., 160, 466–472, http://dx.doi.org/10.1016/j.resmic.2009.07.008doi:10.1016/j.resmic.2009.07.008, 2009b. </mixed-citation>
</ref>
<ref id="ref18">
<label>18</label><mixed-citation publication-type="other" xlink:type="simple"> Geslin, C., Le Romancer, M., Erauso, G., Gaillard, M., Perrot, G., and Prieur, D.: PAV1, the first virus-like particle isolated from a hyperthermophilic Euryarchaeote, \textitPyrococcus abyssi, J. Bacteriol., 185, 3888–3894, http://dx.doi.org/10.1128/JB.185.13.3888-3894.2003doi:10.1128/JB.185.13.3888-3894.2003, 2003a. </mixed-citation>
</ref>
<ref id="ref19">
<label>19</label><mixed-citation publication-type="other" xlink:type="simple"> Geslin, C., Le Romancer, M., Gaillard, M., Erauso, G., and Prieur, P.: Observation of virus-like particles in high temperature enrichment cultures from deep-sea hydrothermal vents, Res Microbiol., 154, 303–307, http://dx.doi.org/10.1016/S0923-2508(03)00075-5doi:10.1016/S0923-2508(03)00075-5, 2003b. </mixed-citation>
</ref>
<ref id="ref20">
<label>20</label><mixed-citation publication-type="other" xlink:type="simple"> Geslin, C., Le Romancer, M., Gaillard, M., and Prieur, D.: Diversité virale associée aux écosystèmes hydrothermaux océaniques profonds et aux sources chaudes terrestres, Virologie, 9, 357–366, 2005. </mixed-citation>
</ref>
<ref id="ref21">
<label>21</label><mixed-citation publication-type="other" xlink:type="simple"> Gibson Jr., E. K., McKay, D. S., Thomas-Keprta, K. L., Wentworth, S. J., Westall, F., Steele, A., Romanek, C. S., Bell, M. S., and Toporski, J.: Life on Mars: evaluation of the evidence within Martian meteorites ALH 84001, Nakhla and Shergotty, Precambrian Res., 106, 15–34, http://dx.doi.org/10.1016/S0301-9268(00)00122-4doi:10.1016/S0301-9268(00)00122-4, 2001. </mixed-citation>
</ref>
<ref id="ref22">
<label>22</label><mixed-citation publication-type="other" xlink:type="simple"> Gorlas, A., Geslin, C., and Prieur, D.: TV1, the first virus-like particle of \textitThermococcus, a hyperthermophilic Archaea genus, Origins Life Evol. B., 39, p 62, 2009. </mixed-citation>
</ref>
<ref id="ref23">
<label>23</label><mixed-citation publication-type="other" xlink:type="simple"> Gorlas, A., Alain, K., Bienvenu, N., Isaac, S., and Geslin, C.: \textitThermococcus prieurii sp. nov., a novel hyperthermophilic archaeon isolated from a deep-sea hydrothermal vent at the East Pacific Rise, Int. J. Syst. Evol. Micr., submitted, 2011. </mixed-citation>
</ref>
<ref id="ref24">
<label>24</label><mixed-citation publication-type="other" xlink:type="simple"> Gowing, M. M.: Large viruses and infected microeukaryotes in Ross Sea summer pack ice habitats, Mar. Biol., 142, 1029–1040, http://dx.doi.org/10.1007/s00227-003-1015-xdoi:10.1007/s00227-003-1015-x, 2003. </mixed-citation>
</ref>
<ref id="ref25">
<label>25</label><mixed-citation publication-type="other" xlink:type="simple"> Gunnarsson, I. and Arnórsson, S.: Amorphous silica solubility and the thermodynamic properties of H&lt;sub&gt;4&lt;/sub&gt;SiO°&lt;sub&gt;4&lt;/sub&gt; in the range of 0° to 350 °C at P$_\rm sat$, Geochim. Cosmochim. Ac., 64, 2295–2307, http://dx.doi.org/10.1016/S0016-7037(99)00426-3doi:10.1016/S0016-7037(99)00426-3, 2000. </mixed-citation>
</ref>
<ref id="ref26">
<label>26</label><mixed-citation publication-type="other" xlink:type="simple"> Jiang, S., Steward, G., Jellison, R., Chu, W., and Choi, S.: Abundance, distribution and diversity of viruses in alkaline hypersaline Mono Lake, California, Microb. Ecol., 47, 9–17, http://dx.doi.org/10.1007/s00248-003-1023-xdoi:10.1007/s00248-003-1023-x, 2004. </mixed-citation>
</ref>
<ref id="ref27">
<label>27</label><mixed-citation publication-type="other" xlink:type="simple"> Kepner, R. L., Wharton Jr., R. A., and Suttle, C. A.: Viruses in Antarctic lakes, Limnol. Oceanogr., 43, 1754–1761, 1998. </mixed-citation>
</ref>
<ref id="ref28">
<label>28</label><mixed-citation publication-type="other" xlink:type="simple"> Konhauser, K. O., Jones, B., Reysenbach, A. L., and Renaut, R. W.: Hot spring sinters: Keys to understanding Earth&apos;s earliest life forms, Can. J. Earth Sci., 40, 1713–1724, 2003. </mixed-citation>
</ref>
<ref id="ref29">
<label>29</label><mixed-citation publication-type="other" xlink:type="simple"> Konhauser, K. O., Jones, B., Phoenix, V. R., Ferris, G., and Renaut, R. W.: The microbial role in hot spring silicification, Ambio, 33, 552–558, 2004. </mixed-citation>
</ref>
<ref id="ref30">
<label>30</label><mixed-citation publication-type="other" xlink:type="simple"> Koonin, E. V. and Dolja, V. V.: Evolution of complexity in the viral world: the dawn of a new vision. Virus Res., 117, 1–4, http://dx.doi.org/10.1016/j.virusres.2006.01.018doi:10.1016/j.virusres.2006.01.018, 2006. </mixed-citation>
</ref>
<ref id="ref31">
<label>31</label><mixed-citation publication-type="other" xlink:type="simple"> Koonin, E. V., Senkevich, T. G., and Dolja, V. V.: The ancient virus world and evolution of cells, Biol. Direct, 1, 29, http://dx.doi.org/10.1186/1745-6150-1-29doi:10.1186/1745-6150-1-29, 2006. </mixed-citation>
</ref>
<ref id="ref32">
<label>32</label><mixed-citation publication-type="other" xlink:type="simple"> Krupovic, M. and Bamford, D. H.: Virus evolution: how far does the double beta-barrel viral lineage extend?, Nat. Rev. Microbiol., 6, 941–9488, http://dx.doi.org/10.1038/nrmicro2033doi:10.1038/nrmicro2033, 2008. </mixed-citation>
</ref>
<ref id="ref33">
<label>33</label><mixed-citation publication-type="other" xlink:type="simple"> Kyle, J. E., Eydal, H. S. C., Ferris, F. G., and Pedersen, K.: Viruses in granitic groundwater from 69 to 450 m depth of the Äspö hard rock laboratory, Sweden, ISME J., 2, 571–574, http://dx.doi.org/10.1038/ismej.2008.18doi:10.1038/ismej.2008.18, 2008a. </mixed-citation>
</ref>
<ref id="ref34">
<label>34</label><mixed-citation publication-type="other" xlink:type="simple"> Kyle, J. E., Pedersen, K., and Ferris, F. G.: Virus mineralization at low pH in the Rio Tinto, Spain, Geomicrobiol. J., 25, 338–345, http://dx.doi.org/10.1080/01490450802402703doi:10.1080/01490450802402703, 2008b. </mixed-citation>
</ref>
<ref id="ref35">
<label>35</label><mixed-citation publication-type="other" xlink:type="simple"> Laidler, J. R. and Stedman, K. M.: Virus silicification under simulated hot spring conditions, Astrobiology, 10, 569–576, http://dx.doi.org/10.1089/ast.2010.0463doi:10.1089/ast.2010.0463, 2010. </mixed-citation>
</ref>
<ref id="ref36">
<label>36</label><mixed-citation publication-type="other" xlink:type="simple"> Lalonde, S. V., Konhauser, K. O., Reysenbach, A. L., and Ferris, F. G.: The experimental silicification of Aquificales and their role in hot spring formation, Geobiology, 3, 41–52, http://dx.doi.org/10.1111/j.1472-4669.2005.00042.xdoi:10.1111/j.1472-4669.2005.00042.x, 2005. </mixed-citation>
</ref>
<ref id="ref37">
<label>37</label><mixed-citation publication-type="other" xlink:type="simple"> La Scola, B., Audic, S., Robert, C., Jungang, L., de Lamballerie, X., Drancourt, M., Birtles, R., Claverie, J. M., and Raoult, D.: A giant virus in Amoebae, Science, 299, p. 2033, http://dx.doi.org/10.1126/science.1081867doi:10.1126/science.1081867, 2003. </mixed-citation>
</ref>
<ref id="ref38">
<label>38</label><mixed-citation publication-type="other" xlink:type="simple"> Le Romancer, M., Gaillard, M., Geslin, C., and Prieur, D.: Viruses in extremes environments, Reviews in Environmental Science and Biotechnology, 6, 17–31, http://dx.doi.org/10.1007/s11157-006-0011-2doi:10.1007/s11157-006-0011-2, 2007. </mixed-citation>
</ref>
<ref id="ref39">
<label>39</label><mixed-citation publication-type="other" xlink:type="simple"> Maranger, R., Bird, D. F., and Juniper, S. K.: Viral and bacterial dynamics in Arctic sea ice during the spring algal bloom near Resolute, N.W.T., Canada, Mar. Ecol.-Prog. Ser., 111, 121–127, 1994. </mixed-citation>
</ref>
<ref id="ref40">
<label>40</label><mixed-citation publication-type="other" xlink:type="simple"> Marteinsson, V. T., Watrin, L., Prieur, D., Caprais, J. C., Raguénès, G., and Erauso G.: Phenotypic characterization, DNA similarities, and protein profiles of twenty sulfur-metabolizing hyperthermophilic anaerobic Archaea isolated from hydrothermal vents in the southwestern Pacific Ocean, Int. J. Syst. Bacteriol., 45, 623–632, http://dx.doi.org/10.1099/00207713-45-4-623doi:10.1099/00207713-45-4-623, 1995. </mixed-citation>
</ref>
<ref id="ref41">
<label>41</label><mixed-citation publication-type="other" xlink:type="simple"> Martin, A., Yeats, S., Janekovic, D., Reiter, W. D., Aicher, W., and Zillig, W.: SAV1, a temperate u.v.-inducible DNA virus-like particle from the archarbacterium \textitSulfolobus acidocaldarius isolate B12, EMBO J., 3, 2165–2168, 1984. </mixed-citation>
</ref>
<ref id="ref42">
<label>42</label><mixed-citation publication-type="other" xlink:type="simple"> McKay, D. S., Gibson Jr., E. K., Thomas-Keprta, K. L., Vali, H., Romanek, C. S., Clemett, S. J., Chillier, X. D. F., Maechling, C. R., and Zare, R. N.: Search for Past Life on Mars: Possible Relic Biogenic Activity in Martian Meteorite ALH84001, Science, 273, 924–930, http://dx.doi.org/10.1126/science.273.5277.924doi:10.1126/science.273.5277.924, 1996. </mixed-citation>
</ref>
<ref id="ref43">
<label>43</label><mixed-citation publication-type="other" xlink:type="simple"> Moreira, D. and López-García, P.: Ten reasons to exclude viruses from the tree of life, Nat. Rev. Microbiol., 7, 306–311, http://dx.doi.org/10.1038/nrmicro2108doi:10.1038/nrmicro2108, 2009. </mixed-citation>
</ref>
<ref id="ref44">
<label>44</label><mixed-citation publication-type="other" xlink:type="simple"> Nisbet, E. G. and Sleep, N. H.: The habitat and nature of early life, Nature, 409, 1083–1091, http://dx.doi.org/10.1038/35059210doi:10.1038/35059210, 2001. </mixed-citation>
</ref>
<ref id="ref45">
<label>45</label><mixed-citation publication-type="other" xlink:type="simple"> Orange, F., Westall, F., Disnar, J. R., Prieur, D., Bienvenu, N., Le Romancer, M., and Défarge, C.: Experimental silicification of the extremophilic Archaea \textitPyrococcus abyssi and \textitMethanocaldococcus jannaschii. Applications in the search for evidence of life in early Earth and extraterrestrial rocks, Geobiology, 7, 403–418, http://dx.doi.org/10.1111/j.1472-4669.2009.00212.xdoi:10.1111/j.1472-4669.2009.00212.x, 2009. </mixed-citation>
</ref>
<ref id="ref46">
<label>46</label><mixed-citation publication-type="other" xlink:type="simple"> Orange, F., Disnar, J. R., Westall, F., Prieur, D., and Baillif, P.: Metal cation binding by the hyperthermophilic microorganism, Archaea \textitMethanocaldococcus jannaschii, and its effects on silicification, Palaeontology, accepted, 2011. </mixed-citation>
</ref>
<ref id="ref47">
<label>47</label><mixed-citation publication-type="other" xlink:type="simple"> Oren, A., Bratbak, G., and Heldal, M.: Occurrence of virus like particles in the Dead Sea, Extremophiles, 1, 143–149, http://dx.doi.org/10.1007/s007920050027doi:10.1007/s007920050027, 1997. </mixed-citation>
</ref>
<ref id="ref48">
<label>48</label><mixed-citation publication-type="other" xlink:type="simple"> Ortmann, A. C. and Suttle, C. A.: High abundances of viruses in deep-sea hydrothermal vent system indicate viral mediated microbial mortality, Deep-Sea Res. Pt I, 52, 1515–1527, http://dx.doi.org/10.1016/j.dsr.2005.04.002doi:10.1016/j.dsr.2005.04.002, 2005. </mixed-citation>
</ref>
<ref id="ref49">
<label>49</label><mixed-citation publication-type="other" xlink:type="simple"> Ortmann, A. C., Wiedenheft, B., Douglas, T., and Young, M.: Hot crenarchaeal viruses reveal deep evolutionary connections, Nat. Rev. Microbiol., 4, 520–528, http://dx.doi.org/10.1038/nrmicro1444doi:10.1038/nrmicro1444, 2006. </mixed-citation>
</ref>
<ref id="ref50">
<label>50</label><mixed-citation publication-type="other" xlink:type="simple"> Pagaling, E., Haigh, R. D., Grant, W. D., Cowan, D. A., Jones, B. E., Ma, Y., Ventosa, A., and Heaphy, S.: Sequence analysis of an Archaeal virus isolated from a hypersaline lake in Inner Mongolia, China, BMC Genomics, 8, 410, http://dx.doi.org/10.1186/1471-2164-8-410doi:10.1186/1471-2164-8-410, 2007. </mixed-citation>
</ref>
<ref id="ref51">
<label>51</label><mixed-citation publication-type="other" xlink:type="simple"> Poinar, G. P. and Poinar, R.: Fossil evidence of insect pathogens, J. Invertebr. Pathol., 89, 243–250, http://dx.doi.org/10.1016/j.jip.2005.05.007doi:10.1016/j.jip.2005.05.007, 2005. </mixed-citation>
</ref>
<ref id="ref52">
<label>52</label><mixed-citation publication-type="other" xlink:type="simple"> Prangishvili, D., Arnold, H. P., Götz, D., Ziese, U., Holz, I., Kristjansson, J. K., and Zillig, W.: A novel virus family, the \textitRudiviridae: Structure, virus-host like interactions and genome variability of the Sulfolobus viruses SIRV1 and SIRV2, Genetics, 152, 1387–1396, 1999. </mixed-citation>
</ref>
<ref id="ref53">
<label>53</label><mixed-citation publication-type="other" xlink:type="simple"> Prangishvili, D., Forterre, P., and Garrett, R. A.: Viruses of the Archaea: a unifying view, Nat. Rev. Microbiol., 4, 837–848, http://dx.doi.org/10.1038/nrmicro1527doi:10.1038/nrmicro1527, 2006a. </mixed-citation>
</ref>
<ref id="ref54">
<label>54</label><mixed-citation publication-type="other" xlink:type="simple"> Prangishvili, D., Garrett, R. A., and Koonin, E. V.: Evolutionary genomics of archaeal viruses: unique viral genomes in the third domain of life, Virus Res., 117, 52–67, http://dx.doi.org/10.1016/j.virusres.2006.01.007doi:10.1016/j.virusres.2006.01.007, 2006b. </mixed-citation>
</ref>
<ref id="ref55">
<label>55</label><mixed-citation publication-type="other" xlink:type="simple"> Prigent, M., Leroy, M., Confalonieri, F., Dutertre, M., and DuBow M. S.: A diversity of bacteriophage forms and genomes can be isolated from the surface sands of the Sahara Desert, Extremophiles, 9, 289–296, http://dx.doi.org/10.1007/s00792-005-0444-5doi:10.1007/s00792-005-0444-5, 2005. </mixed-citation>
</ref>
<ref id="ref56">
<label>56</label><mixed-citation publication-type="other" xlink:type="simple"> Rachel, R., Bettstetter, M., Hedlund, B. P., Häring, M., Kessler, A., Stetter, K. O., and Prangishvili, D.: Remarkable morphological diversity of viruses and virus-like particles in hot terrestrial environments, Arch. Virol., 147, 2419–2429, http://dx.doi.org/10.1007/s00705-002-0895-2doi:10.1007/s00705-002-0895-2, 2002. </mixed-citation>
</ref>
<ref id="ref57">
<label>57</label><mixed-citation publication-type="other" xlink:type="simple"> Raoult, D., Audic, S., Robert, C., Abergel, C., Renesto, P., Ogata H., La Scola, B., Suzan, M., and Claverie, J. M.: The 1.2-megabase genome sequence of Mimivirus, Science, 306, 1344–1350, http://dx.doi.org/10.1126/science.1101485doi:10.1126/science.1101485, 2004. </mixed-citation>
</ref>
<ref id="ref58">
<label>58</label><mixed-citation publication-type="other" xlink:type="simple"> Rice, G., Stedman, K., Snyder, J., Wiedenheft, B., Willits, D., Brumfield, S., and McDermott, T.: Viruses from extreme thermal environments, P. Natl. Acad. Sci. USA, 98, 13341–13345, http://dx.doi.org/10.1073/pnas.231170198doi:10.1073/pnas.231170198, 2001. </mixed-citation>
</ref>
<ref id="ref59">
<label>59</label><mixed-citation publication-type="other" xlink:type="simple"> Schelble, R. T., Hall, J. A., Nealson, K. H., and Steele, A.: DNA perseverance of microorganisms exposed to silica: an experimental study, Geobiology, 6, 503–511, http://dx.doi.org/10.1111/j.1472-4669.2008.00177.xdoi:10.1111/j.1472-4669.2008.00177.x, 2008. </mixed-citation>
</ref>
<ref id="ref60">
<label>60</label><mixed-citation publication-type="other" xlink:type="simple"> Schultze-Lam, S., Ferris, F. G., Kohnauser, K. O., and Wiese, R. G.: In situ silicification of an Icelandic microbial mat: implications for microfossil formation, Can. J. Earth Sci., 32, 2021–2026, 1995. </mixed-citation>
</ref>
<ref id="ref61">
<label>61</label><mixed-citation publication-type="other" xlink:type="simple"> Sime-Ngando, T., Lucas, S., Robin, A., Pause Tucker, K., Colombet, J., Bettarel, Y., Desmond, E., Gribaldo, S., Forterre, P., Breitbart, M., and Prangishvili, D.: Diversity of virus–host systems in hypersaline Lake Retba, Senegal, Environ. Microbiol., http://dx.doi.org/10.1111/j.1462-2920.2010.02323.xdoi:10.1111/j.1462-2920.2010.02323.x, in press, 2010. </mixed-citation>
</ref>
<ref id="ref62">
<label>62</label><mixed-citation publication-type="other" xlink:type="simple"> Templeton, M. R., Andrews, R. C., and Hofmann, R.: Impact of iron particles in groundwater on the UV inactivation of bacteriophages MS2 and T4, J. Appl. Microbiol., 101, 732–741, http://dx.doi.org/10.1111/j.1365-2672.2006.02980.xdoi:10.1111/j.1365-2672.2006.02980.x, 2006. </mixed-citation>
</ref>
<ref id="ref63">
<label>63</label><mixed-citation publication-type="other" xlink:type="simple"> Toporski, J. K. W., Steele, A., Westall, F., Thomas-Keprta, K. L., and McKay, D. S.: The simulated silicification of bacteria – new clues to the modes and timing of bacterial preservation and implications for the search for extraterrestrial microfossils, Astrobiology, 2, 1–26, 2002. </mixed-citation>
</ref>
<ref id="ref64">
<label>64</label><mixed-citation publication-type="other" xlink:type="simple"> Vestergaard, G., Shah, S. A., Bize, A., Reitberger, W., Reuteur, M., Phan, H., Brigel, A., Rachel, R., Garrett, R. A., and Prangishvili, D.: \textitStygiolobus Rod-Shaped Virus and the interplay of Crenarchaeal Rudiviruses with the CRISPR Antiviral System, J. Bacteriol., 190, 6837–6845, http://dx.doi.org/10.1128/JB.00795-08doi:10.1128/JB.00795-08, 2008. </mixed-citation>
</ref>
<ref id="ref65">
<label>65</label><mixed-citation publication-type="other" xlink:type="simple"> Westall, F.: Early life: nature, distribution and evolution, in: Origins and evolution of life, an astrobiological perspective, edited by: Gargaud, M., López-García, P., and Martin, H., Cambridge University Press, Cambridge, United-Kingdom, 391–413, 2010. </mixed-citation>
</ref>
<ref id="ref66">
<label>66</label><mixed-citation publication-type="other" xlink:type="simple"> Westall, F. and Southam, G.: The early record of life, Geoph. Monog. Series, 164, 283–304, 2006. </mixed-citation>
</ref>
<ref id="ref67">
<label>67</label><mixed-citation publication-type="other" xlink:type="simple"> Westall, F., Boni, L., and Guerzoni, E.: The experimental silicification of microorganisms, Palaeontology, 38, 495–528, 1995. </mixed-citation>
</ref>
<ref id="ref68">
<label>68</label><mixed-citation publication-type="other" xlink:type="simple"> You, Y., Han, J., Chiu, P. C., and Jin, Y.: Removal and inactivation of waterborne viruses using zerovalent iron, Environ. Sci. Technol., 39, 9263–9269, http://dx.doi.org/10.1021/es050829jdoi:10.1021/es050829j, 2005. </mixed-citation>
</ref>
<ref id="ref69">
<label>69</label><mixed-citation publication-type="other" xlink:type="simple"> Zillig, W., Kletzin, A., Schleper, C., Holz, I., Janekovic, D., Hain, J., Lanzendörfer, M., and Kristjansoon, J. K.: Screening for Sulfolobales, their plasmids, and their viruses in Icelandic solfataras, Syst. Appl. Microbiol., 16, 609–628, 1994. </mixed-citation>
</ref>
</ref-list>
</back>
</article>