Table Of Content~ SpringerWienNewYork
Helga Stan-Lotter
Sergiu Fendrihan
Editors
Adaption of Microbial Life
to Environmental Extremes
Novel Research Results and Application
SpringerWienNewYork
Prof. Dr. Helga Stan-Lotter
Department of Microbiology, University of Salzburg, Salzburg, Austria
Dr. Sergiu Fendrihan
Romanian Bioresource Center and Advanced Research Association,
Bucharest, Romania
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Foreword
Lifeontheedge,lifeatthephysico-chemicallimits,weirdandeccentriclife–these
are the descriptive terms assigned to extremophiles. The first European workshop
entitled“Microbialadaptationtoextremeenvironments”washeldattheUniversity
of Nijmegenin theNetherlandsin 1973 (Heinenetal. 1974).The first useof the
term “extremophiles” is credited to Robert MacElroy, a NASA exobiologist, who
participated in this workshop (MacElroy 1974).
Manymoreconferencesonextremophilesfollowed,thejournal“Extremophiles”
was launched and the CAREX initiative (Coordination Action for Research
Activities on Life in Extreme Environments) was started (www.carex-eu.org),
providing an extensive data base and recently a new roadmap.
The often surprising ranges of physico-chemical factors, within which life is
possible,stimulatedmanyscientiststoexploreastrobiologicalperspectives,withthe
reasoning that findings on the evolution and mechanisms of adaptation of life at
extremes would help to understand the environments of other planets or moons.
Theseastrobiologicalaspectsaredealtwithinseveralchaptersofthisvolume(Billi;
Mapelli et al.; Gomez and Parro; Moissl-Eichinger; Stan-Lotter).
Thebiodiversityofextremeenvironmentsappearsofunexpectedandenormous
size;itsmagnitudeisstilllargelyunknown.Classicalandmolecularapproachesare
used for its investigation and described by Enache et al., Fendrihan and Negoiţă,
Mapelli et al., Heulin et al., Hreggvidsson et al. and Pearce.
Several updated lists of taxonomic descriptions of extremophilic species and
strains are provided: halophilic archaea and bacteria (Enache et al.), desiccation-
resistant and thermotolerant bacteria (Heulin et al.), thermophilic bacteria
(Hreggvidsson et al.), psychrophilic bacteria, archaea, algae and fungi (Fendrihan
and Negoiţă; Pearce).
Adaptations to extreme environmental conditions, especially on the molecular
level,areatopicofuninterruptedfascinationsincethefirstinvestigationsweremade,
and are described by Billi, Heulin et al., Fendrihan and Negoiţă, and Pearce.
Numerous applications of extremophiles for biotechnological purposes have
already been implemented, although the whole potential is certainly not realized
v
Foreword
yet – treatments of this subject are included in the chapters by Billi, Enache et al.,
Fendrihan and Negoiţă, and Pearce.
Sadly,ourdearcolleagueTeodorNegoiţăpassedawayduringthecompletionof
thisvolume(onMarch23,2011attheageof64).Dr.Negoiţăwasthefounderof
the Romanian Institute of Polar Research. He had organized several international
expeditionsintotheArcticandintheyear2005managedtoestablishtheRomanian
Law-Racoviţă Research Station in the Antarctic.
Our thanks go to the initiators and organizers of the CAREX project – Cynan
Ellis-Evans,NicolasWalter,PetraRettberg–andfortheopportunityofparticipating
in great meetings at wonderful places – Sant Feliu de Guixols, Sasbachwalden, –
which facilitated the interactions between people who would otherwise not have
met, and which was also the partial foundation for the contributions in this book.
We are also very grateful to Amrei Strehl and Eva-Maria Oberhauser from the
Springer-Verlag for helpful suggestions and excellent cooperation.
Salzburg,Bucharest,May2011 HelgaStan-Lotter,SergiuFendrihan
References
Heinen W, Skinner FA, Tempest WD, van den Ende G, Vogels GD, van den Drift C, Dundas I,
HarderW,WeerkampA,GounotAM,vanBeeumenJ,MacElroyRD,KnoopeP,SchwartzAW,
Schlegel HG (1974) Proceedings of the first European workshop on microbial adaptation to
extremeenvironments.Biosystems6:57–80
MacElroyRD(1974)Somecommentsontheevolutionofextremophiles.Biosystems6:74–75
vi
Contents
Physico-chemical boundaries of life
Helga Stan-Lotter......................................................................... 1
1 Introduction.............................................................................. 1
2 Brief history of lifeon Earth............................................................. 3
3 Prokaryotes, eukaryotes, the tree of life and viruses..................................... 4
4 Extreme environments and their inhabitants............................................. 7
4.1 Temperature ranges of microorganisms......................................... 7
4.2 Low water activity: halophiles, osmophiles, and xerophiles..................... 8
4.3 Extremes of pH and low nutrients.............................................. 9
4.4 Highpressure...................................................................... 10
4.5 Viruses........................................................................... 10
4.6 Extremophilic multicellular organisms........................................... 11
5 Microbial survival of extreme conditions ................................................ 12
6 Concluding remarks...................................................................... 14
Microbial diversity in deep hypersaline anoxic basins
Francesca Mapelli, Sara Borin, Daniele Daffonchio................................... 21
1 Introduction.............................................................................. 21
2 Localization and origin of DHABs....................................................... 21
2.1 The anoxic brine basins in the Red Sea: the Shaban and Kebrit Deeps........ 22
2.2 The Orca basin and other seafloor anoxicbrines in the Gulf of Mexico....... 22
2.3 Eastern Mediterranean Sea: DHABsand the Chefren mud volcano ........... 23
3 Geochemical features of DHABs......................................................... 23
3.1 DHABsof the Red Sea ......................................................... 24
3.2 DHABsof the Gulf of Mexico.................................................. 24
3.3 DHABsof the Eastern Mediterranean Sea...................................... 25
3.4 DHABsoftheMediterraneanSea:theChefrenmudvolcano(NileDeepSeaFan)... 27
4 Microbial life in the DHABs: biodiversity and adaptation............................... 27
4.1 Prokaryotic biodiversity of the Kebrit and Shaban Deeps, Red Sea............ 28
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Contents
4.2 Phylogenetic analyses of the microbial communities colonizing different
seafloor hypersaline brines of the Gulf of Mexico.............................. 29
4.3 Investigation of the microbiota composition inhabiting the Chefren
mud volcano (Nile Deep Sea Fan) ............................................. 30
4.4 Stratified microbial communitiesat the deep seawater-brine interface
of hypersaline anoxic basins of the Eastern Mediterranean Sea................. 31
5 Concluding remarks...................................................................... 33
Microbial speciation in the geothermal ecosystem
Gudmundur Oli Hreggvidsson, Solveig K. Petursdottir, Snaedis H. Bj€ornsdottir,
Olafur H. Fridjonsson.................................................................... 37
1 Introduction.............................................................................. 37
2 Geothermal areas ........................................................................ 39
2.1 High- and low-temperature fields............................................... 41
2.2 Origin of hot spring water...................................................... 42
2.3 Surface characteristics of geothermal areas...................................... 43
3 Ecology of thermophiles................................................................. 48
3.1 Temperature..................................................................... 49
3.2 pH............................................................................... 49
3.3 Energy sources as a selective pressure .......................................... 50
4 Biogeography of thermophiles........................................................... 52
4.1 Dispersal of thermophiles....................................................... 55
4.2 Biogeography of Thermus....................................................... 60
4.3 Ecological adaptations of Thermus.............................................. 61
Bacterial adaptation to hot and dry deserts
Thierry Heulin, Gilles De Luca, Mohamed Barakat, Arjan de Groot,
Laurence Blanchard, Philippe Ortet, Wafa Achouak.................................. 69
1 Introduction.............................................................................. 69
2 Characteristic of hot and dry deserts with emphasis on Sahara......................... 70
3 Mechanisms for desiccation tolerance ................................................... 71
4 Counting and describing bacterial populations in desert environments................. 73
5 Ramlibacter and its life cycle............................................................. 78
6 Deinococcus and DNA repair............................................................. 80
7 Conclusions.............................................................................. 81
Extremophiles in Antarctica: life at low temperatures
David A. Pearce........................................................................... 87
1 Introduction.............................................................................. 88
2 Environmental extremes associated with the Antarctic.................................. 90
viii
Contents
3 Extreme environments that also occur in the Antarctic ................................ 92
4 Extreme environments particular to the Antarctic...................................... 97
5 Key Antarctic extremophiles............................................................ 99
6 Biodiversity.............................................................................. 100
7 Methodology............................................................................ 102
8 Adaptations ............................................................................. 104
8.1 Pseudoalteromonas haloplanktis ................................................. 107
8.2 M. burtonii...................................................................... 108
8.3 M. frigidum..................................................................... 108
9 Discussion and future perspectives...................................................... 109
Anhydrobiotic rock-inhabiting cyanobacteria: potential for astrobiology
and biotechnology
Daniela Billi.............................................................................. 119
1 Introduction............................................................................. 119
2 Cyanobacteria in hot and cold desert rocks............................................ 120
3 Cyanobacterial adaptation to Earth’s deserts............................................ 123
4 Survival of desert cyanobacteria beyond Earth.......................................... 125
5 Biotechnological exploitation of anhydrobiosis ......................................... 127
Psychrophilic microorganisms as important source for
biotechnological processes
Sergiu Fendrihan, Teodor G. Negoiţă ................................................. 133
1 Introduction............................................................................. 133
2 Diversity of cold-adapted microorganisms.............................................. 134
3 Ecology and biology .................................................................... 135
4 Cold environments...................................................................... 136
5 Adaptation to cold environments....................................................... 140
6 Applications of psychrophilic microorganisms.......................................... 144
7 Conclusions............................................................................. 157
Halophilic microorganisms from man-made and natural hypersaline
environments: physiology, ecology, and biotechnological potential
Madalin Enache, Gabriela Popescu, Takashi Itoh, Masahiro Kamekura.......... 173
1 Introduction............................................................................. 173
2 Halophilic microorganisms.............................................................. 173
2.1 Haloarchaea.................................................................... 174
2.2 Halophilic Bacteria............................................................. 181
ix
Contents
2.3 Romanian hypersaline environments........................................... 182
2.4 Halophiles from salty environments in Romania............................... 185
3 Biotechnological potential............................................................... 186
3.1 Bioremediation................................................................. 186
3.2 Nanobiotechnology............................................................. 187
3.3 S-layers......................................................................... 187
3.4 Extracellular enzymes........................................................... 188
3.5 Halocins........................................................................ 189
3.6 Exopolysaccharides............................................................. 189
3.7 Resistance to heavy metals..................................................... 190
3.8 Therapeutical value............................................................. 191
4 Concluding remarks..................................................................... 192
Applications of extremophiles in astrobiology: habitability
and life detection strategies
Felipe Gómez, Víctor Parro............................................................. 199
1 Introduction............................................................................. 199
2 Extremophiles and astrobiology......................................................... 200
2.1 Extremophiles.................................................................. 200
2.2 Habitability..................................................................... 203
3 Extremophiles as sources of biomarkers for the search for extraterrestrial life......... 204
3.1 Preservation of molecular biomarkers.......................................... 207
3.2 Stability of organic and living matter in planetary bodies:
the case of Mars................................................................ 211
4 Life detection strategies................................................................. 212
4.1 Immunological systems for the detection of signs of life...................... 213
4.2 Detection systems for extant life based on metabolic activity ................. 217
4.3 Other systems for the detection of signs of life................................ 219
4.4 Signs of lifeby remote sensing systems........................................ 221
Extremophiles in spacecraft assembly clean rooms
Christine Moissl-Eichinger............................................................... 231
1 Introduction............................................................................. 231
1.1 Planetary protection............................................................ 231
1.2 Clean rooms.................................................................... 233
1.3 Contamination control and examinations...................................... 233
1.4 Clean roommicrobiology...................................................... 234
1.5 Extremophiles and extremotolerants – definition.............................. 237
2 Spore-forming microorganisms.......................................................... 237
2.1 Background..................................................................... 237
2.2 Results.......................................................................... 239
x
Contents
3 Oligotrophic microorganisms.......................................................... 240
3.1 Background..................................................................... 240
3.2 Results.......................................................................... 241
4 Alkaliphiles and acidophiles............................................................ 242
4.1 Background..................................................................... 242
4.2 Results.......................................................................... 242
5 Autotrophic and nitrogen fixing microorganisms...................................... 246
5.1 Background..................................................................... 246
5.2 Results.......................................................................... 246
6 Anaerobes.............................................................................. 246
6.1 Background..................................................................... 246
6.2 Results.......................................................................... 247
7 Thermophiles and psychrophiles...................................................... 249
7.1 Background..................................................................... 249
7.2 Results.......................................................................... 249
8 Halophiles............................................................................. 250
8.1 Background..................................................................... 250
8.2 Results.......................................................................... 250
9 Archaea................................................................................ 251
9.1 Background..................................................................... 251
9.2 Results.......................................................................... 251
10 Other extremophiles................................................................... 252
11 Lessons learned from the Herschel campaign: extremophiles are everywhere ........ 253
12 The bacterial diversity beyond cultivation, or cultivation vs. molecular analyses...... 255
Subject index .............................................................................. 263
Organism index............................................................................ 271
List of contributors........................................................................ 281
xi
