And Basalt-Hosted Hydrothermal Systems

Home , Deinococcus geothermalis

Comparison of Microbial Communities Inhabiting Ultramafic- and Basalt-Hosted Hydrothermal Systems:

Genetic Characterization of 16S rRNA Genes, Hydrogen

Oxidation and CO 2 Fixation Pathways

DISSERTATION

Dipl.-Biol. Mirjam Perner

Kiel, December 2006

Perner, M., J. Kuever, R. Seifert, T. Pape, A. Koschinsky, K. Schmidt, H. Strauss & J.F. Imhoff.

Perner, M., R. Seifert, S. Weber, A. Koschinsky, K. Schmidt, H. Strauss, M. Peters, K. Haase & J.F. Imhoff.

Perner, M., J. Kuever & J.F. Imhoff.

For Margot

GLOSSARY

ABSTRACT ...... 1

ZUSAMMENFASSUNG ...... 2

1 INTRODUCTION ...... 3

2 MATERIAL AND METHODS ...... 11 in Situ ...... ...... ......

3 RESULTS ...... 18 ...... ...... in Situ ...... cbbL cbbM aclb GLOSSARY

hynL

4DISCUSSION ...... 39 ...... ......

5 ULTRAMAFIC - AND BASALT -HOSTED HYDROTHERMAL SYSTEMS – A COMPARISON - .... 56

REFERENCES ...... 59

ACKNOWLEDGEMENTS ...... 72

TABLES

TABLES cbbL cbbM aclb hynL

FIGURES

Epsilonproteobacteria Deltaproteobacteria Archaea cbbL cbbM aclb hynL cbbL cbbM aclb hynL

ABSTRACT

in situ

ZUSAMMENFASSUNG

in situ

1 INTRODUCTION

1.1 Deep-Sea Hydrothermal Vent Fields

et al. et al. et al. et al. et al.

et al. et al.

1.2 Ultramafic- and Basalt-Hosted Hydrothermal Systems

et al.

1 INTRODUCTION

1.2.1 Ultramafic-Hosted: Logatchev Hydrothermal Field (LHF)

et al. et al. et al. et al. et al. et al. et al. et al. et al. 1 INTRODUCTION

A B

Fig. 1: C

et al. ©

1.2.2 Basalt-Hosted: Lilliput Hydrothermal Field

et al. et al. et al. 1 INTRODUCTION

et al.

Fig. 2:

Bathymodiolus et al. et al.

1.3 Microorganisms Inhabiting Hydrothermal Environments

et al. et al. 1 INTRODUCTION

Methanococcales Aquificales Epsilonproteobacteria Methanococcales Epsilonproteobacteria Aquificales et al. et al. et al. et al. et al.

Methanococcales et al. et al. et al.

et al. et al.

Epsilonproteobacteria et al. et al. et al. et al. et al. et al. et al. et al. et al. Proteobacteria et al. et al. Epsilonproteobacteria et al. Epsilonproteobacteria et al. et al. et al. et al. et al. Epsilonproteobacteria et al. et al. et al. et al.

1 INTRODUCTION

Persephonella et al. Desulfurobacterium et al. Thiomicrospira et al. et al.

et al. et al. et al. et al. et al. et al. et al. et al. et al. Gamma Epsilon Deltaproteobacteria Bacteroidetes Firmicutes Aquificales et al. et al. et al. et al. et al. et al. et al. et al. et al. et al.

1.3.1 Energy Sources

et al.

1 INTRODUCTION

Epsilonproteobacteria et al. et al. et al. hynSL Epsilonproteobacteria et al.

1.3.2 CO 2 Fixation Pathways

et al.

et al. cbbL cbbM

et al. et al. et al.

1 INTRODUCTION

aclBA Deltaproteobacteria et al. Aquificales Chlorobium limicola Epsilonproteobacteria Desulfobacter Crenarchaeota Thermoproteus et al. et al. et al. et al. et al. et al.

Epsilonproteobacteria et al. et al. et al.

et al. et al. et al.

et al.

2 MATERIAL AND METHODS

2.1 Sampling Sites and Fluid Characteristics

2.1.1 Logatchev Hydrothermal Field (LHF)

´´ et al. et al.

Logatchev hydrothermal field (14°45’ N)

Lilliput hydrothermal field (9°33’ S)

Fig. 3:

2 MATERIAL AND METHODS

Fig. 4: et al.

et al. et al.

et al. 2 MATERIAL AND METHODS

2.1.2 Lilliput Hydrothermal Field

2.2 Sample Collection

et al.

2.3 Fluorescence in Situ Hybridization (FISH)

in situ

2 MATERIAL AND METHODS

Table 1:

Position in Probe sequence Fa a Probe Specificity Reference E. Coli (5´→ 3´) (%)

et al. Planctomycetales et al. Verrucomicrobiales et al. et al. Alphaproteobacteria Betaproteobacteria et al. et al. Gammaproteobacteria et al. et al. Epsilonproteobacteria et al. Deltaproteobacteria et al. Gemmimonas et al. Cytophaga Flavobacterium et al. et al.

a b et al.

2.4 DNA Extraction and PCR Amplification

Thiomicrospira

2.4.1 Archaeal and Bacterial 16S rRNA Genes

2 MATERIAL AND METHODS

et al.

2.4.2 Genes Encoding Key Enzymes of the CBB and rTCA cycles

cbbL Thiomicrospira cbbL cbbM aclb cbbL cbbL cbbM cbbM aclb et al. et al.

2.4.3 Genes Encoding NiFe-Uptake Hydrogenase

hynL Thiomicrospira hynL hynL

2.5 Cloning and Sequencing

2 MATERIAL AND METHODS

cbbL hynL Thiomicrospira cbbM cbbL aclb hynL cbbM, cbbL, aclb hynL et al. et al.

2.6 Diversity Estimation of Clone Libraries

b a

2.7 Phylogenetic Analysis

2 MATERIAL AND METHODS

et al. et al. et al. cbbL cbbM aclb hynL

2.8 Nucleotide Sequence Accession Numbers

cbbL cbbM cbbL cbbM aclb hynL cbbL cbbL cbbL Thiomicrospira

3 RESULTS

3.1 Logatchev Hydrothermal Field (LHF)

3.1.1 Diversity Estimation of Clone Libraries

Table 2: Hydrothermal field sites 16S rRNA gene bacteria archaea

Logatchev

(northern MAR)

Fig. 5: 3 RESULTS

3.1.2 Phylogenetic Analysis

et al. . Gammaproteobacteria Vibrionales, Alteromonadales Oceanospirillales Alphaproteobacteria Rhodobacterales Bacteroidetes Betaproteobacteria Burkholderiales Planctomycetales Clostridia Actinobacteria

et al. Epsilonproteobacteria et al.

3.1.2.1 Bacteria

Epsilonproteobacteria. Epsilonproteobacteria Epsilonproteobacteria

3 RESULTS

Escherichia coli K12, AE000406 Lebetimonas acidiphila, AB167820 91 100 Guaymas basin hydrothermal vent clone 182, AF357196 Irina I OTU IB445 (1.3%), AM268870 99 Nautilia lithotrophica, AJ404370 Alvinella pompejana episymbiont Am-H, AF357197 100 Nautilia sp., AJ575809 Group D Caminibacter profundus, AJ535664 100 Irina I OTU IB370 (2.5%), AM268875 Caminibacter mediatlanticus, AY691430 91 Caminibacter hydrogeniphilus, AJ309655 uncultured epsilon proteobacterium, AB113174 Irina II OTU IIB63 (1.5%), AM268709 diffuse outflow Juan de Fuca, AF468762 Iheya North Mid-Okinawa Trough E1825-1, AB175512 Group G Thioreductor micantisoli, AB175498 100 Thioreductor sp. Shim25-G, AB197162 Nitratiruptor tergarcus, AB175499 Iheya North Mid-Okinawa Trough strain SB137-2, AB175507 Iheya North Mid-Okinawa Trough strain NS55-2, AB189149 Nitratiruptor sp. strain 50cm55-A, AB197156 Iheya North Mid-Okinawa Trough strain CBC55-2, AB175505 CIR hydrothermal system clone pCIRB-G, AB113167 Group A 100 deep-sea hydrothermal field strain 365-55-1%, AB091303 Irina II OTU IIB37 (7.7%), AM268716 Hydrogenimonas thermophila, AB105048 Hydrogenimonas thermophila, AB113193 Irina II OTU IIB17 (1.5%), AM268720 68 Snake Pit clone, AF367484 100 mineral surface clone, DQ228643 Irina I OTU IB472 (5.0%), AM268878 Nitratifractor salsuginis, AB175500 92 fluid samples Mid-Okinawa Trough strain 50cm25-F2, AB197161 Irina II OTU IIB194 (1.5%), AM268728 Iheya North Mid-Okinawa Trough E9S37-1, AB175511 98 Sulfurovum sp. 50cm25-F1, AB197160 Iheya North Mid-Okinawa Trough NM25-1, AB175510 95 Sulfurovum lithotrophicum str. 42BKT, AB091292 Rainbow hydrothermal fluids, AJ969500 Irina II OTU IIB6 (5.0%), AM268718 uncultured epsilon proteobacterium, AJ575963 Hydrothermal vent clone, AY075127 Site B OTU BB26 (1.1%), AM268567 Irina II OTU IIB106 (1.5%), AM268727 Rimicaris exoculata ectosymbiont, U29081 Rainbow vent field sediment sample, AY225623 100 uncultured bacterium, DQ228581 Irina II OTU IIB112 (15.4%), AM268719 filamentous epibiont of Vulcanolepas osheai, AB239758 Group F vent snail epibiont clone SF_C23-G9, AY531572 91 Irina II OTU IIB162 (1.5%), AM268729 uncultured epsilon proteobacterium, AF449240 uncultured epsilon proteobacterium, AJ575971 Site B OTU B21C6 (7.5%), AM268592 fluid samples Mid-Okinawa Trough, AB197179 Site B OTU BB18 (1.1%), AM268590 93 deep sea hydrothermal vent clone, AJ575948 Site B OTU B18G2 (10.8%), AM268581 Irina II OTU IIB166 (3.1%), AM268731 Rainbow hydrothermal fluids, AJ969488 Rimicaris exoculata gut clone, AJ515716 uncultured epsilon proteobacterium, AJ515717 67 Site B OTU B18C6 (6.5%), AM268582 Irina II OTU IIB74 (1.5%), AM268711 deep sea sediment clone BD2-1, AB015531 Nankai Trough sediment clone, AB013260 100 deep sea microbial mat clone Milano, AY592854 Site B OTU B22C10 (20.4%), AM268587 sulfide and methane rich cold seep clone, AY768981 fluid samples Mid-Okinawa Trough, AB197180 Irina II OTU IIB64 (1.5%), AM268710 Irina I OTU IB362 (1.3%), AM268881 100 Arcobacter nitrofigilis (T), L14627 58 Arcobacter skirrowii (T), L14625 Campylobacter jejuni (T), L14630 Rainbow hydrothermal fluids, AJ969453 100 Geospirillium sp KoFum, Y18254 Dehalospirillum multivorans, X82931 98 94 Sulfurospirillum barnesii (T), AF038843 Rainbow hydrothermal fluids, AJ969490 Irina I OTU IB394 (1.3%), AM268882 Sulfurospirillum Group Rainbow hydrothermal fluids, AJ969489 bone-eating symbiosis clone C3F4, AY548994 Irina I OTU IB416 (1.3%), AM268868 100 93 Sulfurospirillum arcachonense (T), Y11561 Sulfurospirillum carboxydovorans, AY740528 Sulfuricurvum kujiense, AB080643 87 Site B OTU B16H6 (1.1%), AM279649 Rimicaris exoculata gut clone 22, AJ515713 100 Iheya North Mid-Okinawa Trough, AB175538 diffuse outflow Juan de Fuca, AF468757 endosymbiont of Alviniconcha sp. type2, AB235231 Rainbow hydrothermal fluids, AJ969498 100 Irina II OTU IIB18 (3.1%), AM268714 cold seep sediments, AB189375 100 Rainbow hydrothermal fluids, AJ969482 Sulfurimonas sp.P6.5m25B, AB197157 Group B Sulfurimonas sp. NS25-1, AB175508 Sulfurimonas autotrophica, AB088431 Rainbow hydrothermal fluids, AJ969504 Rainbow vent field sediment sample, AY225613 Irina I OTU IB477 (2.5%), AM268879 Irina I OTU IB440 (3.8%), AM268873 Snake Pit clone, AF367490 Rainbow hydrothermal fluids, AJ969480 Rainbow hydrothermal fluids, AJ969483 10% 91 Sulfurimonas sp. Go25-1, AB197158 Thiomicrospira denitrificans (T), L40808

Fig. 6: Epsilonproteobacteria 3 RESULTS

Sulfurospirillum

Sulfurovum lithotrophicum Sulfurospirillum Hydrogenimonas thermophila

Deeply-Rooted Lineages. Desulfurobacterium Persephonella Oceanithermus profundus

Epsilonproteobacteria et al. et al. et al. et al.

3 RESULTS

Escherichia coli K12, AE000406 55/99 microbial mat clone SRB29, AY193172 100/100 microbial mat clone SRB7, AY193170 68/- deep sea sediment clone BD2-14, AB015542 100/- oral cavity clone X112, AF125207 SR1 uncultured candidate, AB231045 Site B OTU BB12 (1.1%), AM268609 91/85 sulfide rich spring clone ZB18, AY327235 87/- Guaymas basin clone B02R018, AY197400 Irina I OTU IB488 (1.3%), AM268841 Termite gut clone M2PT2-27, AB192119 100/100 Termite gut clone MTG-96, DQ307728 13°N colonization device, AJ575962 97/77 Irina II OTU IIB78 (7.7%), AM268746 RE1 100/100 Haakon Mosby Mud Volcano clone, AJ704700 Irina II OTU IIB33 (1.5%), AM268744 100 Snake Pit growth chamber, AF367488 Snake Pit growth chamber, AF367482 51/98 Rainbow hydrothermal fluids, AJ969468 100/100 Snake Pit growth chamber, AF367483 contaminated aquifer clone WCHB1-11, AF050603 99/- hot spring clone OPB92, AF027030 54/73 contaminated aquifer clone WCHB1-26, AF050599 OP11 contaminated aquifer clone WCHB1-64, AF050606 84/86 98/96 uncultured bacterium, AB179673 uncultured bacterium, AB179678 80/- 100/100 uncultured bacterium, AB179663 96/94 uncultured bacterium, AB179680 uncultured bacterium, AB179676 73/75 OD1 100/100 microbial mat clone SRA1, AY193175 uncultured bacterium, AB179664 52/62 uncultured bacterium, AF419683 Site B OTU B6F1 (1.1%), AM268637 uncultured bacterium, DQ189915 Thermotoga maritima (T), M21774 74/93 100/100 Thermodesulfobacterium thermophilum (T), AF334601 70/78 Thermodesulfotobacterium sp SRI-27, AF255595 100/- Balnearium lithotrophicum, AB105049 Thermovibrio ammoniificans, AY263403 50/- Desulfurobacterium thermolithotrophum (T), AJ001049 89/- Desulfurobacterium sp., AY268939 Desulfurobacterium Desulfurobacterium sp., AY268938 100/61 100/91 Irina I OTU IB505 (1.3%), AM268865 Persephonella guaymasensis, AF385630 100/100 Persephonella marina, AF188332 99 Aquificales str. CIR30126, AF393378 /97 Irina I OTU IB379 (1.3%), AM268867 Aquificales 100/100 100/94 Persephonella sp., AF507960 99/66 Persephonella hydrogeniphila, AB086419 Aquifex aeolicus VF5, AE000709 84/89 Thermus thermophilus, AJ251939 92/94 Iheya North Mid-Okinawa Trough, AB175569 100/93 Oceanithermus profundus, AJ430586 Irina I OTU IB390 (2.5%), AM268866 Thermus/Deinococcus 61/- Smoker Spire East Pacific Rise clone, AY672508 99/89 Oceanithermus desulfurans, AB107956 Deinococcus geothermalis, Y13040 10%

Fig. 7:

Gammaproteobacteria. Bathymodiolus Bathymodiolus Escarpia spicata et al. Codakia orbicularis et al.

3 RESULTS

Deltaproteobacteria. Deltaproteobacteria Desulfobulbaceae Desulfocapsa sulfexigens

Desulfosarcina variabilis (T), M34407 Desulfobulbus mediterraneus, AF354663 93 Desulfobulbus rhabdoformis (T), U12253 92 Desulfobulbus elongatus (T), X95180 95 Rainbow hydrothermal fluids, AJ969450 52 Irina II OTU IIB152 (1.5%), AM268766 100 Irina II OTU IIB133 (1.5%), AM268767 Riftia pachyptila`s tube clone R103-B27, AF449228 98 Riftia pachyptila`s tube clone R103-B13, AF449229 Guaymas Basin hydrothermal vent sediment isolate a1b012, AF420336 vent snail foot surface clone SF_C6-D1, AY327886 13°N colonization device, AJ575987 70 Desulfocapsa sulfexigens (T), Y13672 96 vent snail epibiont clone C11-G5, AY355297 Irina II OTU IIB75 (4.6%), AM268749 Site B OTU B21C9 (10.8%), AM268636 90 uncultured eubacterium WCHB1-67, AF050536 70 Desulfocapsa thiozymogenes (T), X95181 100 uncultured sulfate-reducing bacterium 330, AJ389627 67 Desulfofustis glycolicus (T), X99707 Desulfotalea arctica (T), AF099061 100 Desulfotalea psychrophila (T), AF099062 Haakon Mosby Mud Volcano clone, AJ704668 83 Desulfobacterium corrodens, AY274450 68 Haakon Mosby Mud Volcano clone, AJ704669 Haakon Mosby Mud Volcano clone, AJ704695 97 Site B OTU BB27 (2.2%), AM268611 uncultured delta proteobacterium, AB015588 Haakon Mosby Mud Volcano clone, AJ704692 uncultured Desulforhopalus, AY177789 Haakon Mosby Mud Volcano clone, AJ704698 90 uncultured delta proteobacterium Sva1036, AJ240990 uncultured delta proteobacterium Sva1037, AJ240992 Desulfobacterium catecholicum (T), AJ237602 52 Rainbow hydrothermal fluids, AJ969502 Site B OTU B21D8 (2.2%), AM268646 uncultured sulfate-reducing bacterium 141, AJ389624 100 uncultured sulfate-reducing bacterium 22, AJ389622 10% Desulforhopalus singaporensis (T), AF118453

Fig. 8: Deltaproteobacteria

3.1.2.2 Archaea

Methanococcales

3 RESULTS

Methanocaldococcus infernus Methanothermococcus thermolithotrophicus Methanocaldococcus janaschii

Aquifex aeolicus VF5, AE000751 100 Palaeococcus ferriphilus, AB019239 95 Thermococcus kodakaraensis, D38650 Thermococcus profundus, Z75233 86 Thermococcales 99 Pyrococcus furiosus, U20163 Pyrococcus abyssi, AJ225071 100 Pyrococcus horikoshii, AP000001 Irina II OTU II26B8 (8.7%), AM268692 97 Methanocaldococcus jannaschii, M59126 Methanocaldococcus vulcanius, AF051404 98 78 Rainbow hydrothermal fluids, AJ969473 Rainbow hydrothermal fluids, AJ969471 100 Rainbow hydrothermal fluids, AJ969474 Rainbow hydrothermal fluids, AJ969469 58 Irina II OTU IIA119 (4.4%), AM268675 Irina I OTU IAG2 (14.3%), AM268777 Methanocaldococcus infernus, AF025822 Site B OTU B16E4 (2.9%), AM268544 100 Methanotorris formicicus, AB100884 100 Irina I OTU IF5 (5.7%), AM268779 Irina I OTU IAF8 (11.4%), AM268783 Methanococcales petroleum reservoir clone vp183, AF220345 Methanothermococcus sp., AF220347 Methanothermococcus thermolithotrophicus, M59128 100 Methanococcus vannielii, AY196675 100 Methanococcus maripaludis, U38484 Methanococcus voltae, U38461 85 Methanothermococcus okinawensis, AB057722 91 "Methanococcus aeolicus", U39016 Irina I OTU I25G8 (5.7%), AM268789 Site B OTU B20H4 (47.1%), AM268557 Irina II OTU II26C11 (54.4%), AM268694 Irina I OTU I28A12 (37.2%), AM268786 100 100 Snake Pit vent cap clone, AF068824 100 Ferroglobus placidus, AF220166 Archaeoglobus fulgidus, Y00275 Archaeoglobales 100 Archaeoglobus veneficus, Y10011 uncultured archaeon, AF419627 100 uncultured archaeon, AF419652 100 87 uncultured archaeon, AF419654 ANME-1 63 uncultured archaeon, AF419649 100 uncultured archaeon, AF419626 97 Methanogenium cariaci, M59130 100 Methanomicrobium mobile, M59142 72 Methanocorpusculum parvum, M59147 84 Methanospirillum hungatei, M60880 Methanosaeta thermoacetophila, M59141 100 100 Methanosaeta sp., AF424772 Methanosaeta soehngenii, X16932 X51423 Methano- 54 100 Methanosaeta concilii, X51423 100 Methanococcoides burtonii, X65537 sarcinales 69 Methanococcoides methylutens, M59127 Site B OTU B20G6 (2.9%), AM268564 archaeon 2C83, AF015977 99 uncultured archaeon, AF354130 uncultured archaeon, AF354134 ANME-2 100 uncultured archaeon, AF354138 10% 100 uncultured archaeon, AF354140

Fig. 9:

3 RESULTS

3.2 Lilliput Hydrothermal Field

3.2.1 Diversity Estimation of Clone Libraries

in situ

Table 3:

Hydrothermal field site 16S rRNA gene bacteria archaea Lilliput (southern MAR)

Fig. 10:

3 RESULTS

3.2.2 Fluorescence in Situ Hybridization (FISH)

Cytophaga Flavobacterium Epsilon Gamma Deltaproteobacteria Alpha Betaproteobacteria

Epsilonproteobacteria

Table 4: Probe Specificity Abundance a Alphaproteobacteria Betaproteobacteria Gammaproteobacteria Epsilonproteobacteria Deltaproteobacteria Gemmimonas

Cytophaga Flavobacterium

3 RESULTS

Fig. 11: Epsilonproteobacteria

3.2.3 Phylogenetic Analysis

Beta- Gamma- Epsilon- Deltaproteobacteria Bacteroidetes Firmicutes Acidobacteria Crenarchaeota

3.2.3.1 Bacteria

Epsilonproteobacteria Arcobacter Deltaproteobacteria Desulfobacterales Desulfovibrionales

3 RESULTS

Archaeoglobus fulgidus , AE000965 Lilliput diffuse fluid clone LDDD11 (2.5%), AM295212 100 Ralstonia pickettii , AB004790 Rhodoferax antarcticus (T), AF084947 Betaproteobacteria 100 Lilliput diffuse fluid clone LDGF10 (1.3%), AM295238 54 Monochlorobenzene Contaminated Groundwater clone, AY050604 Lilliput diffuse fluid clone LDGD7 (1.3%), AM295225 Pseudomonas viridiflava , AF094751 100 Oceanobacter kriegii (T), AB006767 Bacterioplankton ZA3706c, AF382127 100 Lilliput diffuse fluid clone LDGB5 (1.3%), AM295237 Lilliput diffuse fluid clone GG6 (1.3%), AM295236 Gammaproteobacteria Pseudoalteromonas sp KT0812A, AF239705 62 Lilliput diffuse fluid clone LDDA12 (1.3%), AM295203 100 Thiomicrospira crunogena , AF069959 Lilliput diffuse fluid clone LDGD10 (1.3%), AM295235 71 Thiomicrospira thermophila , AB166731 100 Sulfurimonas autotrophica , AB088431 Lilliput diffuse fluid clone LDGC7 (1.3%), AM295226 Group B 78 Mucous secretion Paralvinella palmiformis AJ441199 Sulfurovum lithotrophicum , AB091292 98 Lilliput diffuse fluid clone LDGE7 (1.3%), AM295218 99 Rainbow vent field sediment sample, AY225617 Lilliput diffuse fluid clone LDB01_H (1.3%), AM295247 Group F vent snail epibiont clone SF_C23-H11, AY531570 Lilliput diffuse fluid clone LDDF12 (1.3%), AM295216 100 Lilliput diffuse fluid clone LDGG8 (3.8%), AM295219 56 Sulfurospirillum carboxydovorans , AY740528 100 Bacteroides ureolyticus (T), L04321 71 Campylobacter jejuni (T), L04315 Lilliput diffuse fluid clone LDDD9 (1.3%), AM295210 Mucous secretion Paralvinella sulfincola AY280411 Epsilonproteobacteria 100 Lilliput diffuse fluid clone LDGE8 (1.3%), AM295227 13°N colonization device, AJ575943 72 vent snail foot surface clone SF_C11-F3, AY327877 Lilliput diffuse fluid clone LDGB7 (1.3%), AM295228 Diffuse outflow Juan de Fuca, AF468752 Lilliput diffuse fluid clone LDGG7 (1.3%), AM295224 100 Arcobacter halophilus , AF513455 Candidatus Arcobacter sulfidicus , AY035822 100 Diffuse outflow Juan de Fuca, AF468779 vestimentiferan tubeworm clone, D83061 100 Lilliput diffuse fluid clone LDGG9 (2.5%), AM295221 Arcobacter Rainbow vent field sediment sample, AY225610 100 Lilliput diffuse fluid clone LDGD9 (1.3%), AM295220 relatives deep sea hydrothermal vent chminey CHA3-437, AJ132728 Lilliput diffuse fluid clone LDGA7 (3.8%)), AM295222 Diffuse outflow Juan de Fuca, AF468711 55 Lilliput diffuse fluid clone LDGG4 (2.5%), AM295223 Desulfovibrio caledoniensis , U53465 Lilliput diffuse fluid clone LDDE11 (1.3%), AM295207 Lilliput diffuse fluid clone LDGC5 (1.3%), AM295243 54 marine sediment clone MBMS30, AY193150 Desulfobacterium corrodens , AY274450 Lilliput diffuse fluid clone LDDH10 (11.3%), AM295208 Site B OTU B21D8, AM268646 Deltaproteobacteria 100 Desulfofaba gelida (T), AF099063 100 Lilliput diffuse fluid clone LDDB11 (1.3%), AM295206 100 Desulfobacula phenolica (T), AJ237606 Lilliput diffuse fluid clone LDGA10 (1.3%), AM295245 62 Antarctic sediment clone SB1-49 , AY177795 batch reactor clone SBRH58, AF268992 100 Lilliput diffuse fluid clone LDGD4 (1.3%), AM295229 Candidate Division TM6 harbor sediment clone VHS-B3-74, DQ394961 Lilliput diffuse fluid clone LDFC11 (1.3%), AM295200 100 Guaymas basin hydrothermal sediment clone C1_B017, AF419691 Candidate Division OP8 97 Riftia pachyptila‘s tube clone R76-B102, AF449263 uncultured Holophaga/Acidobacterium Sva0450, AJ240998 Lilliput diffuse fluid clone LDGD6 (1.3%), AM295244 Acidobacteria Salt marsh clone SIMO-2332, AY711698 71 Clostridium aminobutyricum , X76161 Lilliput diffuse fluid clone LDGH9 (2.5%), AM295242 100 uncultured Low G+C Gram-positive bacterium Sva1064, AJ241007 Fusibacter paucivorans (T), AF050099 Firmicutes 79 Lilliput diffuse fluid clone LDGD8 (1.3%), AM295232 100 Mucous secretion Paralvinella sulfincola AY280415 100 Flexibacter aggregans, M58791 Lilliput diffuse fluid clone LDDH9 (1.3%), AM295213 100 Guaymas Basin bottom water, AF419687 sulfidic cave microbial mat clone LKC, AY510257 Site B OTU B21C5, AM268643 Lilliput diffuse fluid clone LDDE12 (1.3%), AM295214 Lilliput diffuse fluid clone GB10 (2.5%), AM295234 100 Cytophaga fermentans (T), M58766 Bacteroidetes Lilliput diffuse fluid clone LDDC9 (10.0%), AM295204 54 Mucous secretion Paralvinella palmiformis AJ441219 Lilliput diffuse fluid clone LDDF11 (1.3%), AM295205 vent snail epibiont clone SF_C9-B4, AY531558 Lilliput diffuse fluid clone LDGH4 (2.5%), AM295231 57 sulfidic cave microbial mat clone LKC, AY510256 100 deep sea sediment clone BD2-14, AB015542 microbial mat clone SRB1, AY193174 Lilliput diffuse fluid clone LDDD10 (1.3%), AM295211 Candidate Division SR1 sulfide rich spring clone ZB18, AY327235 91 Lilliput diffuse fluid clone LDGF9 (1.3%), AM295233 enrichment clone SHA-95, AJ306788 Lilliput diffuse fluid clone LDDF10 (2.5%), AM295217 groundwater clone KNA6-NB10, AB179666 Candidate Division WS5 100 trickling water clone koll6, AJ224539 Lake Gossenkoellesee clone GKS2-30, AJ290044 Lilliput diffuse fluid clone LDFG11 (1.3%), AM295248 Lilliput diffuse fluid clone LDDE9 (3.8%), AM295249 57 Guaymas Basin hydrothermal sediment clone C1_B032, AF419694 100 Lilliput diffuse fluid clone LDGC6 (1.3%), AM295240 99 mangrove bacterioplankton clone DS020, DQ234104 Lilliput diffuse fluid clone LDGH7 (1.3%), AM295241 Candidate Division OP11 marine sediment clone Bol26, AY193135 82 Guaymas basin clone B04R033, AY197423 Lilliput diffuse fluid clone LDGG5 (1.3%), AM295246 marine sediment clone BMS_29, AF172926 Candidate Division WS6 51 100 uncultured bacterium, AY532576 deep sea sediment clone BD7-1, AB015577 Lilliput diffuse fluid clone LDDC10 (1.3%), AM295209 100 Lilliput diffuse fluid clone LDGA6 (1.3%), AM295230 100 subsurface clone uEV818BHEB5102702SAS79, DQ256360 63 deep sea sediment clone BD5-13, AB015569 Lilliput diffuse fluid clone LDDA11 (1.3%), AM295201 Candidate Division OD1 100 Suiyo Seamount clone Sc-NB03, AB193914 Lilliput diffuse fluid clone LDDH12 (1.3%), AM295215 98 Anaerobic swine lagoons, DQ018805 Lilliput diffuse fluid clone LDGB9 (1.3%), AM295239 10% Lilliput diffuse fluid clone LDDA9 (3.8%), AM295202 microbial mat Guerrero Negro, DQ330686 Fig. 12:

3 RESULTS

Thiomicrospira thermophila Thiomicrospira crunogena Riftia pachyptila et al. Acidobacterium

Bacteroidetes Paralvinella palmiformis et al. et al. et al. et al. Firmicutes et al. Paralvinella sulfincola et al.

et al. et al. et al. et al. et al.

3.2.3.2 Archaea

3 RESULTS

Nitrosopumilus maritimus et al. et al. Aciduliprofundum boonei et al. et al. et al.

Aquifex aeolicus VF5, AE000751 100 Methanothermococcus thermolithotrophicus, M59128 Methanococcales 81 Methanocaldococcus jannaschii , M59126 100 Pyrococcus abyssi , AJ225071 Thermococcales 85 Thermococcus celer , M21529 Euryarchaeota 100 Archaeoglobus veneficus , Y10011 79 Archaeoglobus profundus , AF297529 Archaeoglobales Ferroglobus placidus , AF220166 76 Methanobacterium formicicum , AF028689 68 Halobacterium salinarium NCIMB 786, AJ002947 Methanomicrobium mobile , M59142 Lilliput diffuse fluid clone LDG12C (4.8%), AM295196 DHVE 3 74 uncultured euryarchaeote, AY396622 Aciduliprofundum boonei , DQ451875 100 DHVE 2 100 Picrophilus oshimae , X84901 Thermoplasma acidophilum , M38637 100 100 hydrothermal field Japan area, AB019738 hydrothermal field Japan area, AB019737 Diffuse outflow Juan de Fuca, AF355815 DHVE 1 Lilliput diffuse fluid clone LDG12E (4.8%), AM295197 96 hydrothermal field Japan area, AB019731 94 Santa Barbara Channel clone SB95-57, U78199 Lilliput diffuse fluid clone LD37B4 (4.8%), AM295191 hydrothermal field Japan area, AB019728 Lilliput diffuse fluid clone LDG12H (19.1%), AM295193 67 hydrothermal field Japan area, AB019730 Lilliput diffuse fluid clone LDG12D (28.6%), AM295195 100 deep sea hydrothermal vent water, AB193984 99 hydrothermal field Japan area, AB019729 Crenarchaeota hydrothermal field Japan area, AB019727 67 Lilliput diffuse fluid clone LD37B5 (9.5%), AM295190 Marine Cenarchaeum symbiosum, U51469 Group I Lost City carbonate chimney, AY505049 82 70 Candidatus Nitrosopumilus maritimus , DQ085097 88 North Atlantic clone C46, U71117 100 ShabanDeep Brine-Seawater Interface clone ST-3K4A, AJ347774 Lilliput diffuse fluid clone LDH02_H (9.5%), AM295198 Lilliput diffuse fluid clone LDG11D (9.5%), AM295194 100 Lost City carbonate chimney, AY505047 Lilliput diffuse fluid clone LDC03_H (4.8%), AM295199 Lost City carbonate chimney, AY505051 87 100 uncultured marine crenarchaeote, DQ085102 Lilliput diffuse fluid clone LD22A2 (4.8%), AM295192 Guaymas Basin sediment clone AT_R021, AF419653 99 Haakon Mosby Mud Volcano clone, AJ579323 100 100 wastewater clone, AF424775 100 Lake Griffy sediment clone, U59986 100 Desulfurococcus mobilis, M36474 Sulfolobus solfataricus, X90478 10%

Fig. 13:

3 RESULTS

3.3 Functional Genes at the Logatchev and Lilliput hydrothermal fields

3.3.1 Diversity Estimation of the Clone Libraries

cbbL cbbM aclb hynL cbbL cbbM aclb hynL

Table 5: cbbL cbbM aclb hynL Hydrothermal sites PCR detection of functional genes field CBB rTCA H2 cycle cycle oxidation

cbbL cbbM aclb hynL

Logatchev (northern MAR) Lilliput (southern MAR)

CbbL cbbM aclb hynL

cbbM hynL

3 RESULTS

cbbL cbbM cbbL cbbM aclb hynL

Fig. 14: cbbL cbbM aclb hynL

3 RESULTS

aclb

3.3.2 The CBB (cbbL and cbbM Genes) and the rTCA (aclb Gene) Cycles

Thiomicrospira Thiomicrospira thermophila Thiomicrospira cbbL cbbL cbbL Thiomicrospira

cbbL Solemya Bathymodiolus Thiomicrospira thermophila Thiomicrospira cbbL cbbL

Thiomicrospira thermophila Thiomicrospira crunogena

3 RESULTS

Methanocaldococcus janaschii DSM 2661, L77117 98 LHF Site B B05 (2.6%), AM403045

LHF Irina II F10 (14.3%), AM403034 ( RubisCOI form LHF Site B C06 (47.4%), AM403044 LHF Irina II H07 (10.7%), AM403035 LHF Site B D07 (7.9%), AM403043 Lilliput H06 (6.7%), AM403039 LHF Irina II G08 (10.7%), AM403037 94 LHF Site B C07 (5.3%), AM403041 LHF Site B F10 (36.8%), AM403042 94 Lilliput G08 (53.3%), AM403040 Endosymbiont of Bathymodiolus azoricus , AY945760 92 LHF Irina II A02 (3.6%), AM403036 85 Solemya velum gill symbiont, AY531637 Acidithiobacillus ferrooxidans ATCC 23270, AF307091 Thiomicrospira crunogena cbbL1, DQ272534 Thiomicrospira kuenenii cbbL1, DQ272531 cbbL 99 Thiomicrospira sp. strain Lilliput-1 cbbL-1, AM404076 LHF Irina II F12 (53.6%), AM403032 100 Lilliput G06 (40.0%), AM403038 93 Thiomicrospira crunogena cbbL2, DQ272533 ) Thiomicrospira kuenenii cbbL2, DQ272532 LHF Irina II G05 (7.1%), AM403033 96 Thiomicrospira sp. strain Lilliput-1 cbbL-2, AM404077 100 Riftia pachyptila endosymbiont, AF047688 100 LHF Irina II B07 (3.7%), AM40362 Lilliput B9 (5.9%), AM403011 100 hydrothermal plume clone Suiyo (II)-5, AB174751 LHF Irina II H2 (3.7%), AM403012 LHF Site B C04 (16.7%), AM403018 Rhodospirillum rubrum , X00286 ( RubisCOII form 100 Hydrogenovibrio marinus cbbM, D28135 100 Thiomicrospira kuenenii cbbM, DQ272537 Lilliput A08 (58.8%), AM403019 100 Thiomicrospira thermophila cbbM, AY958067 hydrothermal plume clone Fryer (II)-13, AB206056 LHF Irina II E01 (3.7%), AM403061 magnetite-containing magnetic Vibrio, AF442518 Thiomicrospira pelophila cbbM, DQ272536 LHF Site B B09 (8.3%), AM403015 LHF Irina II A07 (3.7%), AM403013 96 Lilliput A3 (5.9%), AM403010 Lilliput E04 (5.9%), AM403008 groundwater and aquifer clone, AY099396

LHF Site B D05 (16.7%), AM403016 cbbM 96 Hydrothermal Plume clone Fryer(II)-15, AB206058 Lilliput B12 (5.9%), AM403009 83 LHF Site B B05 (16.7%), AM403014 Hydrothermal Plume clone Suiyo (II)-4, AB174750

86 ) LHF Irina II D01 (3.7%), AM403060 LHF Irina II D02 (81.5%), AM403059 98 hypersaline anoxic basin clone DB-cbbm 2, DQ149110 LHF Site B C07 (41.6%), AM403017 94 Lilliput A12 (17.6%), AM403020 10% Thiomicrospira crunogena XCL-2, CP000109

Fig. 15: cbbL cbbM

aclb aclb aclb Arcobacter 3 RESULTS

Epsilonproteobacteria Arcobacter Epsilonproteobacteria

Chlorobium limicola , AB054669 LHF Irina II H04 (15.4%),AM403113 Microbial mat EPR, AM181243 Nautilia sp. Am-H, AY430879 Group D Hydrothermal vent EPR, AY430967 100 Lebetimonas acidiphila strain Pd55, AY929803 Group D Thioreductor micantisoli strain BKB25Ts-Y, AY929800 Group G Hydrothermal vent Guaymas Basin, AY430989 97 LHF Irina II F4b (35.9%), AM403114 LHF Irina II G1b (2.6%), AM403115 LHF Site B A12 (93.1%), AM403119 85 Sulfurovum lithotrophicum strain BKB 42, AY929802 Group F Hydrothermal vent EPR, AY430944 Nitratifractor salsuginis strain E9I37-1, AY929801 Group F 85 Hydrogenimonas thermophila strain EP1-55-1%, AY929804 Group A LHF Irina II F6b (5.1%), AM403116 85 Sulfurimonas autotrophica strain OK10, AY929806 Group B Sulfurimonas paralvinella strain GO25-1, AY929805 LHF Irina II F9b (5.1%), AM403117 Hydrothermal vent EPR, AY430941 85 Thiomicrospira denitrificans strain DSM1251, AY885677 Group B LHF Irina II G5b (7.7%), AM404112 LHF Irina II G8 (12.8%), AM404109 Lilliput E11 (3.0%), AM403108 sulfur oxidizing enrichment EPR, AY553060 Lilliput B10 (21.2%), AM403103 Lilliput B12 (3.0%), AM403104 Lilliput A12 (3.0%), AM403102 uncultured episymbiont of Alvinella pompejana , AY308600 LHF Irina II H1 (5.1%), AM403118 LHF Site B E07 (6.9%), AM403120 Hydrothermal vent Guaymas Basin, AY430938 Hydrothermal vent EPR, AY430977 LHF Irina II F1b (5.1%), AM403111 Lilliput C02 (39.4%), AM403105 Lilliput D05 (21.2%), AM403106 Candidatus Arcobacter sulfidicus, AY308640 Arcobacter Hydrothermal vent Guaymas Basin, AY430984 LHF Irina II H6b (5.1%), AM403110 Hydrothermal vent Guaymas Basin, AY430939 Lilliput B09 (9.1%), AM403107 uncultured episymbiont of Alvinella pompejana , AY308630 10%

Fig. 16: aclb

3 RESULTS

Table 6: cbbL Solemya velum Solemya velum Solemya velum Solemya velum Bathymodiolus azoricus Solemya velum Solemya velum Bathymodiolus azoricus Bathymodiolus azoricus Thiomicrospira cbbl Thiomicrospira cbbL Solemya velum Bathymodiolus azoricus Thiomicrospira cbbl cbbM Rhodospirillum rubrum Riftia pachyptila Vibrio Thiomicrospira crunogena Riftia pachyptila Thiomicrospira thermophila Thiomicrospira crunogena aclb Sulfurovum lithotrophicum Nautilia Sulfurovum lithotrophicum Hydrogenimonas thermophila Thiomicrospira denitrificans hynL Flavobacterium johnsoniae Flavobacterium johnsoniae Rhodobacter sphaeroides Hydrogenimonas thermophila Thiomicrospira crunogena Thiomicrospira crunogena Thiomicrospira denitrificans Thiomicrospira denitrificans Flavobacterium johnsoniae Oligotropha carboxydoverans Hydrogenimonas thermophila Hydrogenimonas thermophila Thiomicrospira denitrificans Thiomicrospira denitrificans Azotobacter vinelandii Oligotropha carboxydoverans Thiomicrospira crunogena Lebetimonas acidiphila

3 RESULTS

3.3.3 Hydrogen Oxidation (NiFe-Uptake Hydrogenase gene: hynL )

hynL Epsilonproteobacteria Campylobacter jejuni Thioreductor micantsoli Lebetimonas acidiphila Epsilonproteobacteria

Aquifex aeolicus strain VF5, AE000657 Thiocapsa roseopersicina , AF002817 LHF Site B 33F01 (2.6%), AM403046 Flavobacterium johnsoniae UW101, AAPM01000004 92 LHF Irina II 33F07 (6.7%), AM403047 100 LHF Site B 33E01 (18.4%), AM403048 LHF Site B 33H03 (5.3%), AM403049 95 Rhodobacter sphaeroides ATCC 17025, NZ_AAME01000007 97 100 Paracoccus denitrificans PD1222, NZ_AAIT01000002 95 Azotobacter vinelandii , L23970 Lilliput 33B05 (10.5%), AM403050 100 Alcaligens hydrogenophilus , S56898 100 Pseudomonas hydrogenovora , D86986 86 Ralstonia eutropha , U82564 Oligotropha carboxydovorans , X82447 LHF Irina II 33A08 (6.7%), AM403051 Lilliput 33D05 (10.5%), AM403052 97 Hydrogenimonas thermophila strain EP1-55-1%, AB206664 100 LHF Site B 33D01 (5.3%), AM403053 87 LHF Irina II 33E08 (13.3%), AM403054 88 LHF Irina II 33C09 (6.7%), AM403056 97 Nitratifractor salsuginis strain E9I37-1, AB206665 "Sulfurimonas paralvinella" strain GO25, AB206666 100 100 LHF Irina II 33F09 (30.0%), AM403027 LHF Irina II 33A09 (6.7%),AM403028 LHF Site B 33F03 (10.5%), AM403026 100 LHF Irina II 33C08 (3.3%), AM403023 LHF Site B 33E02 (5.3%), AM403025 100 LHF Site B 33D02 (31.6%), AM403022 100 LHF Site B 33C01 (5.3%), AM403030 100 Thiomicrospira crunogena XCL-2, CP000109 Lilliput 33A06 (73.7%), AM403057 LHF Site B 33G03 (5.3%), AM403055 LHF Irina II 33F08 (13.3%), AM403021 Thiomicrospira denitrificans ATCC 33889, NC_007575 LHF Irina II 33H10 (13.3%), AM403058 100 99 LHF Site B 33D03 (7.9%), AM403024 100 LHF Site B 33H01 (2.6%), AM403029 Helicobacter pylori , CP000241 Wollinella succinogenes strain DSM1740, BX571661 Campylobacter jejuni strain NCTC11168, AL139077 100 86 Lilliput 33B04 (5.3%), AM403031 Lebetimonas acidiphila strain Pd55, AB206667 10% Thioreductor micantsoli strain BKB25Ts-Y, AB206668

Fig. 17: hynL Epsilonproteobacteria 3 RESULTS

Epsilonproteobacteria Thiomicrospira crunogena Thiomicrospira crunogena hynL hynL Alpha Gammaproteobacteria Bacteroidetes et al.

4 DISCUSSION

et al. et al. et al. et al.

et al. et al. Epsilonproteobacteria et al.

4.1 Logatchev Hydrothermal Field (LHF)

4 DISCUSSION

Methanococcales

4.1.1 The Role of Host Rocks

Methanococcales et al. Methanococcales Methanococcales

Epsilonproteobacteria et al. et al. et al. Aquificales Epsilonproteobacteria Proteobacteria 4 DISCUSSION

Epsilonproteobacteria et al. Epsilonproteobacteria

et al. et al. Aquificales Epsilonproteobacteria et al. Methanococcales et al.

et al. et al. et al.

et al. et al. et al. 4 DISCUSSION

4.1.2 The Importance of Mixing Processes

Epsilonproteobacteria et al. et al. et al. Epsilonproteobacteria Methanococcales Epsilonproteobacteria et al.

4 DISCUSSION

Deltaproteobacteria

Epsilonproteobacteria et al. Desulfobulbaceae

et al. Bathymodiolus Solemya Thiomicrospira Thiomicrospira Bathymodiolus Bathymodiolus et al. et al. Bathymodiolus aclb aclb

4 DISCUSSION

cbbM Epsilonproteobacteria

Methanococcales et al. et al. et al. Methanococcales

4.1.3 Other Environmental Parameters

et al. et al. et al.

Methanosarcinales et al. et al.

4 DISCUSSION

4.2 Lilliput Hydrothermal Field

≤ et al. Thiomicrospira Epsilonproteobacteria

Table 8: Microorganisms Lilliput hydrothermal field

Thiomicrospira γ + Methylococcales - δ +

- + - ε

ACTERIA + Proteobacteria B - Arcobacter + Aquificales - Desulfurobacterium -

Methanococcales - Methanosarcinales - RCHAEA A

4.2.1 Novelty of the Ecosystem

et al. et al.

4 DISCUSSION

Bathymodiolus Bathymodiolus Bathymodiolus Mytilus Bathymodiolus Bathymodiolus

4.2.2 Habitats Characterized by Microorganisms

cbbL cbbM aclb

4 DISCUSSION

et al. et al. aclb

Fig. 18:

4 DISCUSSION

et al. et al. et al.

4.2.2.1 Archaea of an Oxic Environment

Crenarchaeota Euryarchaeota Crenarchaeota 4 DISCUSSION

Crenarchaeota Nitrosopumilus maritimus et al. Nitrosopumilus Aciduliprofundum boonei et al.

Table 9:

Physico-chemical parameters ambient seawater Lilliput hydrothermal field

End-member fluid portion T pH H2 CH 4 13 δ Cmethane ‰ H2S Fe 2+ Fe 3+

et al.

4 DISCUSSION

4.2.2.2 Bacteria of the Oxic/Anoxic Interface

Gorgonaria et al.

Thiomicrospira et al. Thiomicrospira Thiomicrospira Thiomicrospira

Gammaproteobacteria

Thiomicrospira Epsilonproteobacteria et al. Thiomicrospira et al. Epsilonproteobacteria et al. Epsilonproteobacteria Arcobacter 4 DISCUSSION

et al. Aquificales Epsilonproteobacteria Epsilonproteobacteria Epsilonproteobacteria et al. et al. Epsilonproteobacteria et al.

Epsilonproteobacteria et al. et al. et al. et al. et al. Gammaproteobacteria Thiomicrospira Epsilonproteobacteria

et al. et al. aclb

Arcobacter Campylobacter aclb

4 DISCUSSION

aclb aclb

Epsilonproteobacteria et al. Epsilonproteobacteria

et al. et al. Cytophaga Flavobacterium et al. Firmicutes et al.

4.2.2.3 Bacteria of the Anoxic Zone

Epsilonproteobacteria et al. et al. Deltaproteobacteria

4 DISCUSSION

Epsilonproteobacteria et al. Deltaproteobacteria Epsilonproteobacteria et al.

Desulfobacula

Desulfovibrio

Desulfobacterium

et al. et al. et al. et al. et al. et al. et al. et al. et al. et al.

4 DISCUSSION

et al.

et al. et al.

et al.

5 ULTRAMAFIC - AND BASALT -HOSTED HYDROTHERMAL SYSTEMS – A COMPARISON -

5 ULTRAMAFIC - AND BASALT -HOSTED HYDROTHERMAL SYSTEMS

– A COMPARISON -

5.1 Microorganisms and Energy Sources

et al. Aquificales Epsilonproteobacteria Methanococcales et al. et al. et al.

5 ULTRAMAFIC - AND BASALT -HOSTED HYDROTHERMAL SYSTEMS – A COMPARISON -

5.2 CO 2 Fixation Pathways

et al. et al.

5 ULTRAMAFIC - AND BASALT -HOSTED HYDROTHERMAL SYSTEMS – A COMPARISON -

Thiomicrospira thermophila

aclb

et al. et al. aclb

aclb

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ACKNOWLEDGMENTS

ACKNOWLEDGEMENTS

ERKLÄRUNG