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Methanothermobacter – Biokatalysator Für Die Energiewende

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Methanothermobacter – Biokatalysator Für Die Energiewende 14 WISSENSCHAFT Mikrobe des Jahres 2021 Methanothermobacter – Biokatalysator für die Energiewende autotrophen, wie Knallgasbakterien und nitri- SEIGO SHIMA, RUDOLF K. THAUER fi zierende Bakterien, benötigen Sauerstoff. MAX-PLANCK-INSTITUT FÜR TERRESTRISCHE MIKROBIOLOGIE, MARBURG Heute gibt es acht beschriebene Methano- thermobacter-Spezies. Sie wurden aus anaero- Methanothermobacter is a thermophilic genus within the kingdom of ben Klärschlämmen, industriellen thermo- philen Biogasanlagen, heißen Quellen oder Euryarchaeota. Chemolithoautotrophic growth on H2 and CO2 at 65 °C is rapid and to high cell concentrations. Champions in this respect are Quellwasser von Gasfeldern isoliert und wachsen alle auf H und CO mit einem Tem- the species M. thermautotrophicus and M. marburgensis, which were 2 2 peraturoptimum um 65 °C. Ihre Genome sind used to elucidate the unique biochemistry of methane formation from ringförmig, 1,6–1,8 ∗ 106 Basenpaare groß H2 and CO2. These two species are presently also being explored as und haben einem DNA-G+C-Gehalt um die biocatalysts in the industrial conversion of electrolytically produced 50 Prozent. Wichtigste Spezies sind M. therma uto trophicus und M. marburgensis H2 to “green” methane. (M. marburgensis wurde zunächst Methano- DOI: 10.1007/s12268-021-1530-8 bacterium thermoautotrophicum, Stamm Mar- © Die Autoren 2021 burg, genannt [2]). ó In einer Kläranlage in Urbana, Illinois, handelt [1, 2], die wie ein Bakterium aussieht Beide Spezies vermehren sich exponentiell USA beginnt diese Geschichte. Aus dem (Abb. 1, [3]). mit Verdoppelungszeiten von rund zwei anaeroben Schlamm isolierten Gregory Zei- Die Entdeckung von M. thermautotrophicus Stunden zu höheren Zelldichten als alle kus und Ralph Wolfe 1972 eine hitzeliebende 1972 war ein Meilenstein, weil sie erstmals anderen bisher bekannten Methanbildner. Mikrobe, die in Abwesenheit von Sauerstoff zeigte, dass auch strikte Anaerobier bei Tem- Deshalb gelten sie als Modellorganismen für aus Wasserstoff (H2) und CO2 Methan (CH4, peraturen von 65 °C optimal wachsen kön- Untersuchungen der Biochemie der Metha- Biogas oder Erdgas) bildet [1, 2]. Sie wächst nen; alle damals bekannten extrem Thermo- nogenese aus H2 und CO2 [4, 5]. Sie dienen bei 65 °C optimal. Das Isolat wurde zunächst philen – wie Bacillus acidocaldarius und die als Biokatalysatoren in Pilotanlagen zur Methanobacterium thermoautotrophicum und Gattungen Thermus, Thermoplasma und Sul- Umsetzung von H2 und CO2 zu Methan für später Methanothermobacter thermautotrophi- folobus – benötigen Sauerstoff zum Leben. energetische Zwecke. cus benannt, da es sich um eine wärmelie- Sie zeigte ferner, dass chemotrophe Mikro- Methanothermobacter sind Archaea bende (thermophile), nur von CO2 als Kohlen- ben auch ohne Sauerstoff autotroph leben stoffquelle lebende (autotrophe) Mikrobe können; alle damals beschriebenen Chemo- Ende der 1970er-Jahre erkannte Karl Woese, dass es zwei Arten von Prokaryoten gibt: Bacte ria und Archaea, die nur sehr entfernt AB miteinander verwandt sind. Methanothermo- bacter und alle anderen anaeroben Methan- bildner gehören zu den Archaea. Dazu passt der frühe Befund, dass M. thermautotrophicus eine Gram-positive Zellwand ohne den Wand- baustoff Murein hat und deshalb nicht durch Penicillin im Wachstum gehemmt wird. Anstatt aus Murein besteht die Zellwand von M. thermautotrophicus aus einem mehr- schichtigen Pseudomurein-Sacculus. Auch die Beschaffenheit der Cytoplasmamembran (aus Isoprenoidlipiden) und der Aufbau der ˚ Abb. 1: Methanothermobacter thermautotrophicus sind unbewegliche krumme Stäbchen. RNA-Polymerase deuteten früh darauf hin, A, elektronenmikroskopische Aufnahme des patentierten M. thermautotrophicus-Stamms, den dass M. thermautotrophicus kein Bakterium, die Firma Electrochaea als Biokatalysator verwendet. (Foto (bearbeitet): Andreas Klingl, LMU sondern ein Archaeon ist. München) B, fl uoreszenzmikroskopische Aufnahme einer planktonischen M. thermautotrophicus- Die Gattung Methanothermobacter gehört Zelle nach Anfärben mit AlexaFluor®-488 [3]. Methanothermobacter heftet sich mit feinen Härchen (Fimbriae) an Oberfl ächen an. (mit freundlicher Genehmigung aus [3]) zur Familie der Methanobacteriaceae, die wie- derum zu der Ordnung der Methanobacte- BIOspektrum | 01.21 | 27. Jahrgang 15 riales gehört, eine von inzwischen acht methanogenen Ordnungen im Reich der Euryarchaeota. AB Energie und Kohlenstoff nur aus H2 und CO2 Die farblosen unbeweglichen Archaea leben optimal bei etwa 65 °C, pH 7 und niedrigem Salzgehalt auf H2 und CO2 als einzigen Ener- gie- und Kohlenstoffquellen. Die meisten Spezies wachsen in rein mineralischem Medium ohne Zusätze von Vitaminen, Ami- nosäuren oder Hefeextrakt. Methanothermo- bacter sind bezüglich ihres Energiestoffwech- sels lithotroph (anorganischer Elektronendo- nator) und bezüglich ihres Baustoffwechsels autotroph (Zellkohlenstoff aus CO ). 2 ˚ Die Cytoplasmamembran von Methano- Abb. 2: Schematische Darstellung des Energiestoffwechsels (A) und des autotrophen Baustoff- wechsels (B) von Methanothermobacter bei Wachstum auf H2 und CO2. In der Berechnung der thermobacter ist für H2, CO2 und Methan benötigten Menge ATP für die Synthese von 72 g Zellen aus 3 CO2 wurde angenommen, dass die durchlässig, nicht aber für Zucker, Amino- Zellen zu 50 % aus Kohlenstoff bestehen und dass der theoretische ATP-Wachstumsertrag 5 g säuren, Pyruvat und Acetat, für die es auch Zellen/mol ATP ist. Der Wachstumsertrag pro mol Methan wurde für M. marburgensis zu 2,5 g keine Transportsysteme gibt. Es fehlen Enzy- Zellen/mol Methan bestimmt [5]. MFR: Methanofuran; H4MPT: Tetrahydromethanopterin; HS-CoM: Coenzym M; HS-CoB: Coenzym B; F : Coenzym F ; pFd: Polyferredoxin, das pro me, die diese Verbindungen in der Zelle ver- 420 420 [4Fe4S]-Cluster jeweils ein Elektron überträgt; Acs: Acetyl-CoA-Synthase/Decarbonylase; ATPase: stoffwechseln könnten. Infolgedessen gibt es ATP-Synthetase; Coo: Kohlenmonoxid-Dehydrogenase; Eha und Ehb: energiekonservierende [NiFe]- für Methanothermobacter keine Alternativen Hydrogenasen; Frh: F420-reduzierende [NiFe]-Hydrogenase; Fwd und Fmd: Wolfram- bzw. Molyb- zur Methanbildung aus H2 und CO2 als Ener- dän-abhängige Formylmethanofuran-Dehydrogenase; Hmd: [Fe]-Hydrogenase; Mcr und Mrt: Methyl-Coenzym M-Reduktase; Mtr: Methyltransferase; Mvh-Hdr: Methylviologen-reduzierender giequelle und zur autotrophen CO2-Fixie- rung. Methanothermobacter sind daher obli- [NiFe]-Hydrogenase-Heterodisulfi d-Reduktase-Komplex. gat lithoautotroph, allerdings mit einer Ein- schränkung: Einige Stämme können nämlich Oxidationsstufe +4) zu Methan (–4) erfolgt nosarcinales ist der Kopplungsmechanismus Ameisensäure (Formiat) anstelle von H2 und über Formyl-MFR (+2), Formyl-H4MPT (+2), wesentlich anders. Während im Energiestoff- CO2 zum Wachstum verwenden, wobei aller- Methenyl-H4MPT (+2), Methylen-H4MPT (0), wechsel von Methanothermobacter die ender- dings die Reduktion der Ameisensäure zu Methyl-H4MPT (–2) und Methyl-Coenzym M gone Reduktion von Polyferredoxin (pFd) mit Methan über CO2 läuft. (–2) als Zwischenprodukte. Reduziertes Poly- H2 an die exergone Reduktion von CoM-S-S- 2– Als Stickstoffquelle verwendet die Mikrobe ferredoxin (pFdred ), F420H2 und HS-CoB die- CoB mit H2 durch Flavin-basierte Elektronen- ausschließlich NH3 und als Schwefelquelle nen dabei als Elek tronendonatoren, die ihrer- bifurkation gekoppelt ist (Abb. 2A, [8]), erfolgt ausschließlich H2S. Als Spurenelemente seits durch H2 re-reduziert werden. diese Kopplung in Methanosarcina-Spezies benötigt sie Eisen, Kobalt, Nickel, Molybdän Die bei der Reduktion von CO2 zu Methan chemiosmotisch [5]. und/oder Wolfram, nicht aber Mangan, Kupfer freiwerdende Energie wird zum Aufbau eines und Selen [6]. Wie alle Lebewesen brauchen elektrochemischen Na+-Potenzials verwen- Der Nickelbedarf war eine Zufalls- Methanothermobacter auch Kalium- und det, das seinerseits die endergone Synthese entdeckung Magne siumionen zum Wachstum. Methan- von ATP (Abb. 2A) und die endergone Reduk- Bei Wachstumsversuchen mit M. marburgen- bildung und Wachstum dieser Archaea sind tion von Polyferredoxin (pFd) mit H2 treibt sis beobachteten Peter Schönheit et al. 1979 von Na triumionen abhängig, obwohl Methano- (Abb. 2B, [7]). zufällig, dass Kulturen in Rührfermentern thermobacter keine marinen Organismen sind. Im Falle des Baustoffwechsels wird CO2 zu mit Anteilen aus Edelstahl besser wachsen Kohlenmonoxid (CO) (+2) reduziert, das mit als solche in Fermentern ohne Edelstahlan- Natriumionen für den Stoffwechsel Methyl-H4MPT (–2) (aus dem Energiestoff- teile [6]. Analysen des Edelstahls ergab eine Der Energie- und Baustoffwechsel von Metha- wechsel) und Coenzym A zu Acetyl-CoA Zusammensetzung aus Eisen, Nickel und nothermobacter sind in Abbildung 2 skizziert reagiert, das wiederum reduktiv zu Pyruvat Chrom. Es stellte sich schnell heraus, dass [5, 7]. Am Stoffwechsel sind sechs Co enzyme carboxyliert wird. Von Pyruvat aus werden Nickel das Wachstum beschleunigte, was beteiligt, die erstmals in Methanothermobac- ATP-abhängig fast alle Zellbausteine synthe- eine große Überraschung war. Damals glaub- ter entdeckt wurden. Es handelt sich dabei tisiert (Abb. 2B). te man noch, dass Nickel keine biologische + um die C1-tragenden Coenzyme Methanofu- Der Na -abhängige Energiestoffwechsel von Funktion in Mikroorganismen hat. Dass ran (MFR), Tetrahydromethanopterin CO2 und H2 zu Methan, wie er in Abbildung Methanothermobacter-Spezies zuvor über- (H4MPT) und Coenzym M (HS-CoM) und die 2A aufgezeichnet ist, fi ndet sich mit kleineren
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