Members of Methanocellales are widespread in paddy field soils and play the key role in methane production. These methanogens feature largely in these organisms' adaptation to low H 2 and syntrophic growth with anaerobic fatty acid oxidizers. The adaptive mechanisms, however, remain unknown. In the present study, we determined the transcripts of 21 genes involved in the key steps of methanogenesis and acetate assimilation of Methanocella conradii HZ254, a strain recently isolated from paddy field soil. M. conradii was grown in monoculture and syntrophically with Pelotomaculum thermopropionicum (a propionate syntroph) or Syntrophothermus lipocalidus (a butyrate syntroph). Comparison of the relative transcript abundances showed that three hydrogenase-encoding genes and all methanogenesis-related genes tested were upregulated in cocultures relative to monoculture. The genes encoding formylmethanofuran dehydrogenase (Fwd), heterodisulfide reductase (Hdr), and the membrane-bound energy-converting hydrogenase (Ech) were the most upregulated among the evaluated genes. The expression of the formate dehydrogenase (Fdh)-encoding gene also was significantly upregulated. In contrast, an acetate assimilation gene was downregulated in cocultures. The genes coding for Fwd, Hdr, and the D subunit of F 420 -nonreducing hydrogenase (Mvh) form a large predicted transcription unit; therefore, the Mvh/Hdr/Fwd complex, capable of mediating the electron bifurcation and connecting the first and last steps of methanogenesis, was predicted to be formed in M. conradii. We propose that Methanocella methanogens cope with low H 2 and syntrophic growth by (i) stabilizing the Mvh/Hdr/Fwd complex and (ii) activating formatedependent methanogenesis.T he 16S rRNA gene of a new type of methanogen, Methanocellales, was discovered in 1998 from the surface of rice root (1). The 16S rRNA gene sequences were found phylogenetically branching off between Methanosarcinaceae and Methanomicrobiales. An enrichment culture at 50°C revealed that these organisms contained the methyl-coenzyme M (CoM) reductase-encoding genes, confirming their nature as methanogenic archaea (2, 3). They were found to be widespread in rice field soils and the environment, including desert soil (4-7). Due to the difficulty of cultivation, many molecular studies were conducted to characterize their ecological functions prior to isolation into pure culture. The application of stable isotope probing technology showed that these organisms outcompeted other methanogens under low-H 2 conditions (8). This result suggested, for the first time, that these methanogens are intrinsically adaptive to low H 2 . Moreover, they were found to play the key role in CH 4 production from rootderived material in the rice rhizosphere in situ, illustrating their ecological significance and niche specificity (9).More evidence for the adaptation of Methanocellales to low H 2 came from the investigations of syntrophic oxidation of shortchain fatty acids (10-14). In the investigations of syntrophic oxidations...