Bernard Henrissat

King Abdulaziz University, Djidda, Makkah, Saudi Arabia

Are you Bernard Henrissat?

Claim your profile

Publications (422)2837.85 Total impact

  • [Show abstract] [Hide abstract]
    ABSTRACT: A bacterial polysaccharide utilization locus (PUL) is a set of physically-linked genes that orchestrate the breakdown of a specific glycan. PULs are prevalent in the Bacteroidetes phylum and are key to the digestion of complex carbohydrates, notably by the human gut microbiota. A given Bacteroidetes genome can encode dozens of different PULs whose boundaries and precise gene content are difficult to predict. Results: Here, we present a fully-automated approach for PUL prediction using genomic-context and domain annotation alone. By combining the detection of a pair of marker genes with operon prediction using intergenic distances, and queries to the carbohydrate-active enzymes database (, our predictor achieved above 86% accuracy in two Bacteroides species with extensive experimental PUL characterization. Availability: PUL predictions in 67 Bacteroidetes genomes from the human gut microbiota and two additional species, from the canine oral sphere and from the environment, are presented in our database accessible at
    Bioinformatics 10/2014; 31(5). DOI:10.1093/bioinformatics/btu716 · 4.62 Impact Factor
  • [Show abstract] [Hide abstract]
    ABSTRACT: Crucial virulence determinants of disease causing Neisseria meningitidis (Nm) species are their extracellular polysaccharide capsules (CPSs). In the serogroups W and Y these are heteropolymers of the repeating units [→6)-α-D-Gal-(1→4)-α-Neu5Ac-(2→]n in NmW and [→6)-α-D-Glc-(1→4)-α-Neu5Ac-(2→]n in NmY. The capsule polymerases, SiaDW and SiaDY, which synthesise these highly unusual polymers, are composed of two predicted GT-B fold domains separated by a large stretch of amino acids (aa399-762). We recently showed that residues critical to the hexosyl- and sialyltransferase activity are found in the predicted N-terminal (aa1-398) and C-terminal (aa763-1037) GT-B fold domains, respectively. Here we use a mutational approach and synthetic fluorescent substrates to define the boundaries of the hexosyl- and sialyltransferase domains. Our results reveal that the active sialyltransferase domain extends well beyond the predicted C-terminal GT-B domain and defines a new glycosyltransferase family, GT97, in the Carbohydrate Active Enzyme database (CAZy).
    Journal of Biological Chemistry 10/2014; 289(49). DOI:10.1074/jbc.M114.597773 · 4.57 Impact Factor
  • Source
    [Show abstract] [Hide abstract]
    ABSTRACT: Background Enzymatic breakdown of lignocellulosic biomass is a known bottleneck for the production of high-value molecules and biofuels from renewable sources. Filamentous fungi are the predominant natural source of enzymes acting on lignocellulose. We describe the extraordinary cellulose-deconstructing capacity of the basidiomycete Laetisaria arvalis, a soil-inhabiting fungus. Results The L. arvalis strain displayed the capacity to grow on wheat straw as the sole carbon source and to fully digest cellulose filter paper. The cellulolytic activity exhibited in the secretomes of L. arvalis was up to 7.5 times higher than that of a reference Trichoderma reesei industrial strain, resulting in a significant improvement of the glucose release from steam-exploded wheat straw. Global transcriptome and secretome analyses revealed that L. arvalis produces a unique repertoire of carbohydrate-active enzymes in the fungal taxa, including a complete set of enzymes acting on cellulose. Temporal analyses of secretomes indicated that the unusual degradation efficiency of L. arvalis relies on its early response to the carbon source, and on the finely tuned sequential secretion of several lytic polysaccharide monooxygenases and hydrolytic enzymes targeting cellulose. Conclusions The present study illustrates the adaptation of a litter-rot fungus to the rapid breakdown of recalcitrant plant biomass. The cellulolytic capabilities of this basidiomycete fungus result from the rapid, selective and successive secretion of oxidative and hydrolytic enzymes. These enzymes expressed at critical times during biomass degradation may inspire the design of improved enzyme cocktails for the conversion of plant cell wall resources into fermentable sugars. Electronic supplementary material The online version of this article (doi:10.1186/s13068-014-0143-5) contains supplementary material, which is available to authorized users.
    Biotechnology for Biofuels 10/2014; 7(1):143. DOI:10.1186/s13068-014-0143-5 · 6.22 Impact Factor
  • [Show abstract] [Hide abstract]
    ABSTRACT: To study how microbes establish themselves in a mammalian gut environment, we colonized germ-free mice with microbial communities from human, zebrafish, and termite guts, human skin and tongue, soil, and estuarine microbial mats. Bacteria from these foreign environments colonized and persisted in the mouse gut; their capacity to metabolize dietary and host carbohydrates and bile acids correlated with colonization success. Cohousing mice harboring these xenomicrobiota or a mouse cecal microbiota, along with germ-free "bystanders," revealed the success of particular bacterial taxa in invading guts with established communities and empty gut habitats. Unanticipated patterns of ecological succession were observed; for example, a soil-derived bacterium dominated even in the presence of bacteria from other gut communities (zebrafish and termite), and human-derived bacteria colonized germ-free bystander mice before mouse-derived organisms. This approach can be generalized to address a variety of mechanistic questions about succession, including succession in the context of microbiota-directed therapeutics.
    Cell 10/2014; DOI:10.1016/j.cell.2014.09.008 · 33.12 Impact Factor
  • [Show abstract] [Hide abstract]
    ABSTRACT: Glycosylation of proteins and lipids involves over 200 known glycosyltransferases, and deleterious defects in many of the genes encoding these enzymes cause disorders collectively classified as Congenital Disorders of Glycosylation (CDGs). Most known CDGs are caused by defects in glycogenes that effects glycosylation globally. Many glycosyltransferases are members of homologous isoenzyme families and deficiencies in individual isoenzymes may not affect glycosylation globally. In line with this there appears to be an underrepresentation of disease-causing glycogenes among these larger isoenzyme homologous families. However, Genome-Wide-Association Studies (GWAS) have identified such isoenzyme genes as candidates for different diseases, but validation is not straightforward without biomarkers. Large-scale whole exome sequencing (WES) provides access to mutations in e.g. glycosyltransferase genes in populations, which can be used to predict and/or analyze functional deleterious mutations. Here, we constructed a draft of a Functional Mutational Map of glycogenes, GlyMAP, from WES of a rather homogenous population of 2,000 Danes. We catalogued all missense mutations and used prediction algorithms, manual inspection, and in case of CAZy family GT27 experimental analysis of mutations to map deleterious mutations. GlyMAP provides a first global view of the genetic stability of the glycogenome and should serve as a tool for discovery of novel CDGs.
    Glycobiology 09/2014; 25(2). DOI:10.1093/glycob/cwu104 · 3.14 Impact Factor
  • Source
    [Show abstract] [Hide abstract]
    ABSTRACT: Here, we report the draft genome sequence of two ulvan-degrading Alteromonas spp. isolated from the feces of the sea slug, Aplysia. These sequenced genomes display a unique ulvan degradation machinery compared with ulvanolytic enzymes previously identified in Nonlabens ulvanivorans.
    Genome Announcements 09/2014; 2(5). DOI:10.1128/genomeA.01081-14
  • Source
    [Show abstract] [Hide abstract]
    ABSTRACT: Ectomycorrhizal fungi, living in soil forests, are required microorganisms to sustain tree growth and productivity. The establishment of mutualistic interaction with roots to form ectomycorrhiza (ECM) is not well known at the molecular level. In particular, how fungal and plant cell walls are rearranged to establish a fully functional ectomycorrhiza is poorly understood. Nevertheless, it is likely that Carbohydrate Active enZymes (CAZyme) produced by the fungus participate in this process. Genome-wide transcriptome profiling during ECM development was used to examine how the CAZome of L. bicolor is regulated during symbiosis establishment. CAZymes active on fungal cell wall were upregulated during ECM development in particular after 4 weeks of contact when the hyphae are surrounding the root cells and start to colonize the apoplast. We demonstrated that one expansin-like protein, whose expression is specific to symbiotic tissues, localizes within fungal cell wall. Whereas L. bicolor genome contained a constricted repertoire of CAZymes active on cellulose and hemicellulose, these CAZymes were expressed during the first steps of root cells colonization. L. bicolor retained the ability to use homogalacturonan, a pectin-derived substrate, as carbon source. CAZymes likely involved in pectin hydrolysis were mainly expressed at the stage of a fully mature ECM. All together, our data suggest an active remodelling of fungal cell wall with a possible involvement of expansin during ECM development. By contrast, a soft remodelling of the plant cell wall likely occurs through the loosening of the cellulose microfbrils by AA9 or GH12 CAZymes and middle lamella smooth remodelling through pectin (homogalacturonan) hydrolysis likely by GH28, GH12 CAZymes.
    Fungal Genetics and Biology 08/2014; 72. DOI:10.1016/j.fgb.2014.08.007 · 3.26 Impact Factor
  • Source
    [Show abstract] [Hide abstract]
    ABSTRACT: Clostridium termitidis strain CT1112 is an anaerobic, gram positive, mesophilic, cellulolytic bacillus isolated from the gut of the wood-feeding termite, Nasutitermes lujae. It produces biofuels such as hydrogen and ethanol from cellulose, cellobiose, xylan, xylose, glucose, and other sugars, and therefore could be used for biofuel production from biomass through consolidated bioprocessing. The first step in the production of biofuel from biomass by microorganisms is the hydrolysis of complex carbohydrates present in biomass. This is achieved through the presence of a repertoire of secreted or complexed carbohydrate active enzymes (CAZymes), sometimes organized in an extracellular organelle called cellulosome. To assess the ability and understand the mechanism of polysaccharide hydrolysis in C. termitidis, the recently sequenced strain CT1112 of C. termitidis was analyzed for both CAZymes and cellulosomal components, and compared to other cellulolytic bacteria. A total of 355 CAZyme sequences were identified in C. termitidis, significantly higher than other Clostridial species. Of these, high numbers of glycoside hydrolases (199) and carbohydrate binding modules (95) were identified. The presence of a variety of CAZymes involved with polysaccharide utilization/degradation ability suggests hydrolysis potential for a wide range of polysaccharides. In addition, dockerin-bearing enzymes, cohesion domains and a cellulosomal gene cluster were identified, indicating the presence of potential cellulosome assembly.
    PLoS ONE 08/2014; 9(8):e104260. DOI:10.1371/journal.pone.0104260 · 3.53 Impact Factor
  • Source
    [Show abstract] [Hide abstract]
    ABSTRACT: Background Growing interest in cellulolytic clostridia with potential for consolidated biofuels production is mitigated by low conversion of raw substrates to desired end products. Strategies to improve conversion are likely to benefit from emerging techniques to define molecular systems biology of these organisms. Clostridium stercorarium DSM8532T is an anaerobic thermophile with demonstrated high ethanol production on cellulose and hemicellulose. Although several lignocellulolytic enzymes in this organism have been well-characterized, details concerning carbohydrate transporters and central metabolism have not been described. Therefore, the goal of this study is to define an improved whole genome sequence (WGS) for this organism using in-depth molecular profiling by RNA-seq transcriptomics and tandem mass spectrometry-based proteomics. Results A paired-end Roche/454 WGS assembly was closed through application of an in silico algorithm designed to resolve repetitive sequence regions, resulting in a circular replicon with one gap and a region of 2 kilobases with 10 ambiguous bases. RNA-seq transcriptomics resulted in nearly complete coverage of the genome, identifying errors in homopolymer length attributable to 454 sequencing. Peptide sequences resulting from high-throughput tandem mass spectrometry of trypsin-digested protein extracts were mapped to 1,755 annotated proteins (68% of all protein-coding regions). Proteogenomic analysis confirmed the quality of annotation and improvement pipelines, identifying a missing gene and an alternative reading frame. Peptide coverage of genes hypothetically involved in substrate hydrolysis, transport and utilization confirmed multiple pathways for glycolysis, pyruvate conversion and recycling of intermediates. No sequences homologous to transaldolase, a central enzyme in the pentose phosphate pathway, were observed by any method, despite demonstrated growth of this organism on xylose and xylan hemicellulose. Conclusions Complementary omics techniques confirm the quality of genome sequence assembly, annotation and error-reporting. Nearly complete genome coverage by RNA-seq likely indicates background DNA in RNA extracts, however these preps resulted in WGS enhancement and transcriptome profiling in a single Illumina run. No detection of transaldolase by any method despite xylose utilization by this organism indicates an alternative pathway for sedoheptulose-7-phosphate degradation. This report combines next-generation omics techniques to elucidate previously undefined features of substrate transport and central metabolism for this organism and its potential for consolidated biofuels production from lignocellulose. Electronic supplementary material The online version of this article (doi:10.1186/1471-2164-15-567) contains supplementary material, which is available to authorized users.
    BMC Genomics 07/2014; 15(1):567. DOI:10.1186/1471-2164-15-567 · 4.04 Impact Factor
  • Source
    [Show abstract] [Hide abstract]
    ABSTRACT: Background A complex community of microorganisms is responsible for efficient plant cell wall digestion by many herbivores, notably the ruminants. Understanding the different fibrolytic mechanisms utilized by these bacteria has been of great interest in agricultural and technological fields, reinforced more recently by current efforts to convert cellulosic biomass to biofuels. Methodology/Principal Findings Here, we have used a bioinformatics-based approach to explore the cellulosome-related components of six genomes from two of the primary fiber-degrading bacteria in the rumen: Ruminococcus flavefaciens (strains FD-1, 007c and 17) and Ruminococcus albus (strains 7, 8 and SY3). The genomes of two of these strains are reported for the first time herein. The data reveal that the three R. flavefaciens strains encode for an elaborate reservoir of cohesin- and dockerin-containing proteins, whereas the three R. albus strains are cohesin-deficient and encode mainly dockerins and a unique family of cell-anchoring carbohydrate-binding modules (family 37). Conclusions/Significance Our comparative genome-wide analysis pinpoints rare and novel strain-specific protein architectures and provides an exhaustive profile of their numerous lignocellulose-degrading enzymes. This work provides blueprints of the divergent cellulolytic systems in these two prominent fibrolytic rumen bacterial species, each of which reflects a distinct mechanistic model for efficient degradation of cellulosic biomass.
    PLoS ONE 07/2014; 9(7):e99221. DOI:10.1371/journal.pone.0099221 · 3.53 Impact Factor
  • Source
    [Show abstract] [Hide abstract]
    ABSTRACT: Here we report the draft genome sequence of the bacterium Nonlabens ulvanivorans, which was recently isolated. To our knowledge, this is the first published genome of a characterized ulvan-degrading bacterium. Revealing the ulvan utilization pathways may provide access to a vast marine biomass source that has yet to be exploited.
    Genome Announcements 07/2014; 2(4). DOI:10.1128/genomeA.00793-14
  • Source
    [Show abstract] [Hide abstract]
    ABSTRACT: Brown macroalgae are considered an attractive option as sustainable feedstock for the production of biofuels and commodity chemicals due to their high carbohydrate content (such as alginates), and the absence of hemicellulose and lignin. Microbial communities from cold coastal environments represent promising sources of novel alginate-depolymerizing enzymes, as brown algae constitute a large primary biomass in these environments. We used a nested sampling strategy to obtain coastal sediment samples from four high-latitude regions (Svalbard Archipelago, Norway; Baltic Sea, Sweden; Ushuaia Bay, Argentina and Potter Cove, Antarctic Peninsula). Metagenomes were sequenced using 23 lanes of Illumina HiSeq™ 1500, and were annotated at the IMG/M pipeline. The complete assembled metagenome dataset contains 5.6 Gb and 1.4 x 107 protein coding genes. We mined this dataset with the goal of identifying alginate lyase homologs using both blastp searches and assigned product names. We retrieved 2,705 sequences ≥ 100 amino acids in length, 75% of them belonging to the samples from polar regions. Most sequences were classified as belonging to PL17 (30.4%), PL7 (28%) or PL6 (22%) families (CAZy database), and 13% of them were too distant to reference sequences and could not be classified. Phylogenetic analysis of full length sequences and gene order conservation for long scaffolds show moderate relatedness to sequences from isolated bacteria. This study revealed a large diversity of alginate lyase sequences from marine yet-to-be cultured microorganisms, which could aid in the engineering of microorganisms for biorefineries.
    New Biotechnology; 07/2014
  • Source
    [Show abstract] [Hide abstract]
    ABSTRACT: Basidiomycota (basidiomycetes) make up 32% of the described fungi and include most wood-decaying species, as well as pathogens and mutualistic symbionts. Wood-decaying basidiomycetes have typically been classified as either white rot or brown rot, based on the ability (in white rot only) to degrade lignin along with cellulose and hemicellulose. Prior genomic comparisons suggested that the two decay modes can be distinguished based on the presence or absence of ligninolytic class II peroxidases (PODs), as well as the abundance of enzymes acting directly on crystalline cellulose (reduced in brown rot). To assess the generality of the white-rot/brown-rot classification paradigm, we compared the genomes of 33 basidiomycetes, including four newly sequenced wood decayers, and performed phylogenetically informed principal-components analysis (PCA) of a broad range of gene families encoding plant biomass-degrading enzymes. The newly sequenced Botryobasidium botryosum and Jaapia argillacea genomes lack PODs but possess diverse enzymes acting on crystalline cellulose, and they group close to the model white-rot species Phanerochaete chrysosporium in the PCA. Furthermore, laboratory assays showed that both B. botryosum and J. argillacea can degrade all polymeric components of woody plant cell walls, a characteristic of white rot. We also found expansions in reducing polyketide synthase genes specific to the brown-rot fungi. Our results suggest a continuum rather than a dichotomy between the white-rot and brown-rot modes of wood decay. A more nuanced categorization of rot types is needed, based on an improved understanding of the genomics and biochemistry of wood decay.
    Proceedings of the National Academy of Sciences 06/2014; 111(41). DOI:10.1073/pnas.1400592111 · 9.81 Impact Factor
  • Source
    [Show abstract] [Hide abstract]
    ABSTRACT: Saprophytic filamentous fungi are ubiquitous micro-organisms that play an essential role in photosynthetic carbon recycling. The wood-decayer Pycnoporus cinnabarinus is a model fungus for the study of plant cell wall decomposition and is used for a number of applications in green and white biotechnology.
  • Source
  • Didier Raoult, Bernard Henrissat
    European Journal of Epidemiology 05/2014; 29(5). DOI:10.1007/s10654-014-9905-4 · 5.15 Impact Factor
  • Source
    [Show abstract] [Hide abstract]
    ABSTRACT: The draft genome sequence of the smut fungus Tilletiaria anomala UBC 951 (Basidiomycota, Ustilaginomycotina) is presented. The sequenced genome size is 18.7 Mb, consisting of 289 scaffolds and a total of 6,810 predicted genes. This is the first genome sequence published for a fungus in the order Georgefisheriales (Exobasidiomycetes).
    Genome Announcements 05/2014; 2(3). DOI:10.1128/genomeA.00539-14
  • Source
    [Show abstract] [Hide abstract]
    ABSTRACT: The zygomycete fungi like Rhizomucor miehei have been extensively exploited for the production of various enzymes. As a thermophilic fungus, R. miehei is capable of growing at temperatures that approach the upper limits for all eukaryotes. To date, over hundreds of fungal genomes are publicly available. However, Zygomycetes have been rarely investigated both genetically and genomically. Here, we report the genome of R. miehei CAU432 to explore the thermostable enzymatic repertoire of this fungus. The assembled genome size is 27.6-million-base (Mb) with 10,345 predicted protein-coding genes. Even being thermophilic, the G + C contents of fungal whole genome (43.8%) and coding genes (47.4%) are less than 50%. Phylogenetically, R. miehei is more closerly related to Phycomyces blakesleeanus than to Mucor circinelloides and Rhizopus oryzae. The genome of R. miehei harbors a large number of genes encoding secreted proteases, which is consistent with the characteristics of R. miehei being a rich producer of proteases. The transcriptome profile of R. miehei showed that the genes responsible for degrading starch, glucan, protein and lipid were highly expressed. The genome information of R. miehei will facilitate future studies to better understand the mechanisms of fungal thermophilic adaptation and the exploring of the potential of R. miehei in industrial-scale production of thermostable enzymes. Based on the existence of a large repertoire of amylolytic, proteolytic and lipolytic genes in the genome, R. miehei has potential in the production of a variety of such enzymes.
    BMC Genomics 04/2014; 15(1):294. DOI:10.1186/1471-2164-15-294 · 4.04 Impact Factor
  • Source
    [Show abstract] [Hide abstract]
    ABSTRACT: The genome of the coprophilous fungus Podospora anserina harbors a large and highly diverse set of putative lignocellulose-acting enzymes. In this study, we investigated the enzymatic diversity of a broad range of P. anserina secretomes induced by various carbon sources (dextrin, glucose, xylose, arabinose, lactose, cellobiose, saccharose, Avicel, Solka-floc, birchwood xylan, wheat straw, maize bran, and sugar beet pulp (SBP)). Compared with the Trichoderma reesei enzymatic cocktail, P. anserina secretomes displayed similar cellulase, xylanase, and pectinase activities and greater arabinofuranosidase, arabinanase, and galactanase activities. The secretomes were further tested for their capacity to supplement a T. reesei cocktail. Four of them improved significantly the saccharification yield of steam-exploded wheat straw up to 48 %. Fine analysis of the P. anserina secretomes produced with Avicel and SBP using proteomics revealed a large array of CAZymes with a high number of GH6 and GH7 cellulases, CE1 esterases, GH43 arabinofuranosidases, and AA1 laccase-like multicopper oxidases. Moreover, a preponderance of AA9 (formerly GH61) was exclusively produced in the SBP condition. This study brings additional insights into the P. anserina enzymatic machinery and will facilitate the selection of promising targets for the development of future biorefineries.
    Applied Microbiology and Biotechnology 04/2014; 98(17). DOI:10.1007/s00253-014-5698-3 · 3.81 Impact Factor
  • Nicolas Terrapon, Bernard Henrissat
    [Show abstract] [Hide abstract]
    ABSTRACT: Trillions of commensal bacteria in our colon thrive on what we do not digest in our small intestine. Many have evolved multiple sophisticated machineries, termed polysaccharide utilization loci or PULs, for carbohydrate breakdown; each PUL may target a particular complex carbohydrate. Until now, studies have focused on the structural and functional characterization of individual PUL constituents. A recent work by Larsbrink et al. moves the scope from single-gene analysis to the entire PUL dissection.
    Trends in Biochemical Sciences 03/2014; 39(4). DOI:10.1016/j.tibs.2014.02.005 · 13.52 Impact Factor

Publication Stats

34k Citations
2,837.85 Total Impact Points


  • 2014–2015
    • King Abdulaziz University
      • • Department of Biological Science
      • • Faculty of Sciences
      Djidda, Makkah, Saudi Arabia
    • Universidade Católica de Brasília
      • Pós-Graduação em Ciências Genômicas e Biotecnologia
      Brasília, Federal, Brazil
  • 2002–2015
    • Aix-Marseille Université
      • Unité de Recherche d'Architecture et Fonction des Macromolécules Biologiques (UMR 7257 AFMB)
      Marsiglia, Provence-Alpes-Côte d'Azur, France
    • CUNY Graduate Center
      New York, New York, United States
    • European Synchrotron Radiation Facility
      Grenoble, Rhône-Alpes, France
  • 1998–2015
    • Architecture et Fonction des Macromolécules Biologiques
      Marsiglia, Provence-Alpes-Côte d'Azur, France
  • 1990–2014
    • French National Centre for Scientific Research
      • • Laboratoire Information Génomique et Structurale (IGS)
      • • Laboratoire de Architecture et Fonction des Macromolécules Biologiques
      • • Centre de Recherches sur les Macromolécules Végétales
      Lutetia Parisorum, Île-de-France, France
    • John Innes Centre
      • The Sainsbury Laboratory
      Norwich, England, United Kingdom
  • 2013
    • Cornell University
      • Department of Plant Pathology and Plant-Microbe Biology
      Итак, New York, United States
  • 1988–2013
    • French National Institute for Agricultural Research
      • Biotechnologie des Champignons Filamenteux
      Lutetia Parisorum, Île-de-France, France
  • 2012
    • Clark University
      • Department of Biology
      Worcester, MA, United States
  • 2011
    • Washington University in St. Louis
      San Luis, Missouri, United States
  • 2008
    • Howard University
      • Department of Biology
      Washington, West Virginia, United States
  • 2007
    • Harvard University
      Cambridge, Massachusetts, United States
    • Lawrence Livermore National Laboratory
      Livermore, California, United States
  • 2004
    • Technion - Israel Institute of Technology
      H̱efa, Haifa, Israel
    • Kyoto University
      • Department of Pathology and Tumor Biology
      Kyoto, Kyoto-fu, Japan
  • 2003
    • Hebrew University of Jerusalem
      • Department of Inorganic Chemistry
      Yerushalayim, Jerusalem, Israel
  • 2001
    • Cea Leti
      Grenoble, Rhône-Alpes, France
    • Virginia Polytechnic Institute and State University
      Blacksburg, Virginia, United States
  • 1995–2001
    • The University of York
      • • York Structural Biology Laboratory
      • • Department of Chemistry
      York, England, United Kingdom
  • 1997
    • Newcastle University
      • School of Chemistry
      Newcastle-on-Tyne, England, United Kingdom
  • 1995–1997
    • University Joseph Fourier - Grenoble 1
      • Centre de Recherche sur les MAcromolécules Végétales
      Grenoble, Rhône-Alpes, France
  • 1996
    • Max-Delbrück-Centrum für Molekulare Medizin
      Berlín, Berlin, Germany
  • 1994
    • Niigata University
      • Faculty of Agriculture
      Niahi-niigata, Niigata, Japan
  • 1993
    • Technical University of Denmark
      København, Capital Region, Denmark
  • 1991
    • University of British Columbia - Vancouver
      • Department of Microbiology and Immunology
      Vancouver, British Columbia, Canada
  • 1987
    • Institute of Food Research
      Norwich, England, United Kingdom
  • 1985
    • University of Grenoble
      Grenoble, Rhône-Alpes, France