{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T13:59:58Z","timestamp":1790171998910,"version":"4.1.0"},"reference-count":84,"publisher":"American Society for Microbiology","issue":"9","license":[{"start":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T00:00:00Z","timestamp":1790121600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T00:00:00Z","timestamp":1790121600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/journals.asm.org\/non-commercial-tdm-license"}],"funder":[{"DOI":"10.13039\/100009553","name":"Bundesanstalt f\u00fcr Materialforschung und -Pr\u00fcfung","doi-asserted-by":"publisher","award":["Adolf Martens Fellowship"],"award-info":[{"award-number":["Adolf Martens Fellowship"]}],"id":[{"id":"10.13039\/100009553","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100009553","name":"Bundesanstalt f\u00fcr Materialforschung und -Pr\u00fcfung","doi-asserted-by":"publisher","award":["Ideas: Develop Program (Call 2021,IE2112"],"award-info":[{"award-number":["Ideas: Develop Program (Call 2021,IE2112"]}],"id":[{"id":"10.13039\/100009553","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["journals.asm.org"],"crossmark-restriction":true},"short-container-title":["Appl Environ Microbiol"],"published-print":{"date-parts":[[2026,9,23]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:sec>\n                    <jats:title\/>\n                    <jats:p>\n                      Rock-inhabiting fungi thrive in subaerial oligotrophic environments, such as desert rocks, solar panels, and marble monuments, where organic carbon and nitrogen are scarce. We tested whether the rock-inhabiting fungus\n                      <jats:italic toggle=\"yes\">Knufia petricola<\/jats:italic>\n                      showed a preference regarding nitrogen (\n                      <jats:inline-formula>\n                        <jats:alternatives>\n                          <jats:inline-graphic xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" xlink:href=\"aem.00656-26.im001.tif\"\/>\n                          <mml:math xmlns:mml=\"http:\/\/www.w3.org\/1998\/Math\/MathML\" display=\"inline\" overflow=\"scroll\">\n                            <mml:mi mathvariant=\"normal\">N<\/mml:mi>\n                            <mml:msubsup>\n                              <mml:mrow>\n                                <mml:mi mathvariant=\"normal\">O<\/mml:mi>\n                              <\/mml:mrow>\n                              <mml:mrow>\n                                <mml:mn>3<\/mml:mn>\n                              <\/mml:mrow>\n                              <mml:mrow>\n                                <mml:mo>-<\/mml:mo>\n                              <\/mml:mrow>\n                            <\/mml:msubsup>\n                          <\/mml:math>\n                        <\/jats:alternatives>\n                      <\/jats:inline-formula>\n                      or\n                      <jats:inline-formula>\n                        <jats:alternatives>\n                          <jats:inline-graphic xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" xlink:href=\"aem.00656-26.im002.tif\"\/>\n                          <mml:math xmlns:mml=\"http:\/\/www.w3.org\/1998\/Math\/MathML\" display=\"inline\" overflow=\"scroll\">\n                            <mml:mi mathvariant=\"normal\">N<\/mml:mi>\n                            <mml:msubsup>\n                              <mml:mrow>\n                                <mml:mi mathvariant=\"normal\">H<\/mml:mi>\n                              <\/mml:mrow>\n                              <mml:mrow>\n                                <mml:mn>4<\/mml:mn>\n                              <\/mml:mrow>\n                              <mml:mrow>\n                                <mml:mo>+<\/mml:mo>\n                              <\/mml:mrow>\n                            <\/mml:msubsup>\n                          <\/mml:math>\n                        <\/jats:alternatives>\n                      <\/jats:inline-formula>\n                      ) and carbon (glucose or sucrose) sources, and whether it was sensitive toward carbon and nitrogen limitation. As this fungus produces the carbon-rich, nitrogen-free 1,8-dihydroxynaphthalene (DHN) melanin, we tested whether a melanin-deficient mutant would be less sensitive to carbon limitation. The carbon and nitrogen concentrations were the primary predictors of growth, with a broad optimum partially explained by an optimal fungal C:N ratio. Limiting carbon or nitrogen supply decreased biomass formation,\n                      <jats:inline-formula>\n                        <jats:alternatives>\n                          <jats:inline-graphic xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" xlink:href=\"aem.00656-26.im003.tif\"\/>\n                          <mml:math xmlns:mml=\"http:\/\/www.w3.org\/1998\/Math\/MathML\" display=\"inline\" overflow=\"scroll\">\n                            <mml:mi mathvariant=\"normal\">C<\/mml:mi>\n                            <mml:msub>\n                              <mml:mrow>\n                                <mml:mi mathvariant=\"normal\">O<\/mml:mi>\n                              <\/mml:mrow>\n                              <mml:mrow>\n                                <mml:mn>2<\/mml:mn>\n                              <\/mml:mrow>\n                            <\/mml:msub>\n                          <\/mml:math>\n                        <\/jats:alternatives>\n                      <\/jats:inline-formula>\n                      production, and biofilm thickness, but promoted substratum penetration through filamentous growth. The nitrogen content of the biomass was flexible within limits, increasing with increasing nitrogen supply or decreasing carbon supply. The carbon use efficiency was fairly constant, whereas melanization correlated with a higher nitrogen content of the biomass, despite melanin being nitrogen-free. In conclusion,\n                      <jats:italic toggle=\"yes\">in vitro<\/jats:italic>\n                      ,\n                      <jats:italic toggle=\"yes\">K. petricola<\/jats:italic>\n                      switches to explorative growth under nutrient limitations, like fast-growing fungi, revealing universal fungal resource-acquisition patterns.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>IMPORTANCE<\/jats:title>\n                    <jats:p>\n                      Since their discovery in the 1980s, extremotolerant fungi have been shown to inhabit desert rocks and coastal salterns, deteriorate marble monuments, weather minerals, colonize solar panels, and positively interact with cyanobacteria, algae, and plant microbiomes. These fungi endure sudden shifts in temperature and water availability, as well as intense sunlight, while persisting on very low nutrient levels. Despite their ecological relevance, their biology remains underexplored. The black fungus\n                      <jats:italic toggle=\"yes\">Knufia petricola<\/jats:italic>\n                      offers a rare opportunity to study these organisms in depth because it can be genetically engineered and represents the wider group of resilient surface colonizers. Here, we compared a wild-type strain with a melanin-deficient mutant and generated quantitative data suitable for mathematical modeling of growth. This work provides a foundation for predicting how\n                      <jats:italic toggle=\"yes\">K. petricola<\/jats:italic>\n                      behaves on natural and human-made materials, informing future efforts to harness extremotolerant fungi in sustainable technologies, agriculture, and environmental resilience.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1128\/aem.00656-26","type":"journal-article","created":{"date-parts":[[2026,8,28]],"date-time":"2026-08-28T13:00:38Z","timestamp":1787922038000},"update-policy":"https:\/\/doi.org\/10.1128\/asmj-crossmark-policy-page","source":"Crossref","is-referenced-by-count":1,"title":["Carbon and nitrogen availability affects biofilm growth and morphology of the extremotolerant fungus\n                    <i>Knufia petricola<\/i>"],"prefix":"10.1128","volume":"92","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-7721-3819","authenticated-orcid":true,"given":"Abolfazl","family":"Dehkohneh","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/03x516a66","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Materials and the Environment, Bundesanstalt f\u00fcr Materialforschung und -pr\u00fcfung (BAM)","place":["Berlin, Germany"]},{"id":[{"id":"https:\/\/ror.org\/046ak2485","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Biology, Chemistry and Pharmacy, Freie Universit\u00e4t Berlin","place":["Berlin, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9120-9675","authenticated-orcid":true,"given":"Julia","family":"Schumacher","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/03x516a66","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Materials and the Environment, Bundesanstalt f\u00fcr Materialforschung und -pr\u00fcfung (BAM)","place":["Berlin, Germany"]},{"id":[{"id":"https:\/\/ror.org\/046ak2485","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Biology, Chemistry and Pharmacy, Freie Universit\u00e4t Berlin","place":["Berlin, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3751-0771","authenticated-orcid":true,"given":"Bastiaan J. 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