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SETTING SCIENCE AGENDAS FOR EUROPE
Investigating Life in ExtremeEnvironmentsA European Perspective
InvestigatingLife
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Environm
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www.esf.org
European Science Foundation (ESF)The European Science Foundation (ESF) wasestablished in 1974 to create a common Europeanplatform for cross-border cooperation in all aspectsof scientific research. With its emphasis on a multidisciplinary and pan-European approach, the Foundation provides theleadership necessary to open new frontiers inEuropean science.Its activities include providing science policy advice(Science Strategy); stimulating co-operationbetween researchers and organisations to explorenew directions (Science Synergy); and theadministration of externally funded programmes(Science Management). These take place in thefollowing areas: Physical and engineering sciences;Medical sciences; Life, earth and environmentalsciences; Humanities; Social sciences; Polar;Marine; Space; Radio astronomy frequencies;Nuclear physics.Headquartered in Strasbourg with offices inBrussels, the ESF’s membership comprises 75national funding agencies, research performingagencies and academies from 30 Europeannations.The Foundation’s independence allows the ESF toobjectively represent the priorities of all thesemembers. For more information see:http://www.esf.org
ESF Standing Committee for Life, Earth andEnvironmental Sciences (LESC)LESC encompasses a number of disciplines suchas biology, biotechnology, agriculture, Earthsciences, glaciology, oceanography, meteorology,and other life and environmental sciences. Thecommittee is composed of leading scientistsmandated to represent the ESF MemberOrganisations. Observers from other ESFcommittees/expert groups or externalorganisations are also invited to attend committeemeetings, as are guests from the COST TechnicalCommittees. For more information see:http://www.esf.org/lesc
European Medical Research Councils (EMRC)Set up in 1971, the EMRC evolved into one of thefive Standing Committees of the ESF in 1975. TheEMRC membership is composed of delegates witha high scientific profile who are nominated by theirESF Member Organisations involved in biomedicalsciences, together with observers from Canada,Israel, New Zealand, USA,, WHO-Europe, theEuropean Commission and the ESF StandingCommittee for Life, Earth and EnvironmentalSciences (LESC). Thirty-five representatives of 39Member Organisations participate in the annualplenary meetings of the EMRC. An ExecutiveGroup composed of eight members is responsiblefor the follow-up of EMRC policies between annualmeetings. For more information see:http://www.esf.org/emrc
ESF Standing Committee for the Humanities(SCH)Established in 1978, SCH consists ofrepresentatives of ESF member research councilsand academies, with subject specialists tocomplement ordinary membership. Observersattend from the ESF Standing Committee for theSocial Sciences (SCSS), the COST TechnicalCommittee (TC) in Social Sciences and Humanities,the European Commission, the US NationalEndowment for the Humanities, the CanadianSocial Sciences and Humanities Research Council,and the Israel Academy of Sciences andHumanities. SCH has a policy of encouraginginterdisciplinary work and sees its main task as theindependent evaluation of collaborative researchproposals emanating from the academiccommunity. SCH also has a proactive function inthe identification of priority research areas. Formore information see: http://www.esf.org/sch
2 Investigating Life in Extreme Environments
Cover Page Credits :King penguins – © CNRS Photothèque/Pierre KatellWater ice on Mars – ESA/DLR/FU Berlin (G. Neukum)Physonect siphonophore from the Arctic Ocean – © Kevin RaskoffSand dune in Marocco – © CNRS Photothèque/Raguet HubertDeep sea shrimp – © Ifremer/Campagne Biozaire 2 – 2001Hydrothermal vent sampling – © Ifremer/ Campagne Phare 2002Jupiter’s icy moon Europa – NASAAcidic water, Rio Tinto – J. Segura
3Investigating Life in Extreme Environments
Marine Board-ESF (MB-ESF)The Marine Board-ESF is one of ESF’s expertboards focus on science policy and strategy. It wasestablished in 1995 to facilitate enhancedcoordination between European marine scienceorganisations (both research institutes and fundingagencies) and the development of strategies formarine science in Europe. The Marine Board has23 Member Organisations from 16 countries andoperates by: i) creating a forum for its memberorganisations; ii) identifying strategic scientificissues; iii) providing a voice for European marinescience; and iv) promoting synergy in themanagement of both national programmes andresearch infrastructure facilities and investments.For more information see:http://www.esf.org/marineboard
European Polar Board (EPB)EPB is one of ESF’s expert boards focus onscience policy and strategy. It is Europe’s advisorybody on science policy in the polar regions. It isconcerned with major strategic priorities in theArctic and Antarctic. EPB has 29 members from 19countries. It acts to ensure high-level collaborationand coordination between European nationalfunding agencies, national polar institutes, researchorganisations and the European Commission. Formore information see: http://www.esf.org/epb
European Space Sciences Committee (ESSC)Created in 1975, ESSC is one of ESF’s expertboards focus on science policy and strategy. Itdeals with space research and covers all relatedaspects; i.e. space physical science, Earthobservation, and life and physical sciences inspace. ESSC investigates and presents the view ofthe scientific community in Europe on spaceresearch issues and provides an independent voiceon European space science policy. For moreinformation see: http://www.esf.org/essc
Context and background information on theactivityIn February 2004, in the context of promotinginterdisciplinary research on topics of commoninterest, the European Science Foundation’s (ESF)Standing Committee for Life, Earth andEnvironmental Sciences (LESC) and three of ESF’sExpert Committees, the Marine Board-ESF (MB-ESF), the European Space Science Committee(ESSC) and the European Polar Board (EPB),issued a common call for Expressions of Interest(EoI) on the topic ‘Investigating Life in ExtremeEnvironments’. ESF Standing Committee for theHumanities (SCH) and the European MedicalResearch Councils (EMRC) joined the initiative.
Subsequent to this call, 282 Expressions of Interestfrom 27 countries were received. The maincontributors were from: Germany (18%), UK (16%),Italy (14%) and France (9%). There was alsosignificant participation from the Russia (12 EoIs)and from eastern European countries (25 EoIs).Concerning the topics covered, 33% of the EoIsdealt with the marine environment, 22% with thepolar environment and 16% with the spaceenvironment.
Given the EoIs received, and with the support ofESF and LESC, the Steering Committee of theinitiative recommended that a large-scaleworkshop dealing with the topic Investigating Life inExtreme Environments (ILEE) be organised in orderto debate and put forward research priorities andexplore the recommendations of the communityinvolved. The coordination was tasked to theMarine Board unit of ESF. This workshop, involving128 participants, was held in November 2005 inSant Feliu de Guixols (Spain). The workshop waschaired by Daniel Prieur (IUEM University of Brest,France) with Ricardo Amils (CBMSO UniversidadAutónoma de Madrid and Centro de Astrobiología,Madrid, Spain) as rapporteur.
As a broad spectrum of the European scientificcommunity expressed interest in the initiative, theworkshop was organised around seven sessions(detailed below), three of which dealt withinterdisciplinary issues:
1. Defining and characterising the boundaryconditions of extreme environments - (cross-disciplinary)
2. Molecular adaptation and stability in extremeenvironments - (cross-disciplinary)
3. Microbial life in extreme environments
4. Life strategies of plants in extremeenvironments
5. Life strategies of animals in extremeenvironments
6. Human adaptation to extreme environments
7. Enabling technologies and applications -(cross-disciplinary)
In addition to the presentation sessions, workinggroups were established around five topics:
1. Microbial life in extreme environments
2. Life strategies of plants in extremeenvironments
3. Life strategies of animals in extremeenvironments
4. Human adaptation to extreme environments
5. Enabling technologies and applications
This report results from this workshop and a furtherone held in March 2006 in Strasbourg, the lattergathering the Steering Committee set up for thisinitiative as well as the sessions’ chairpersons andrapporteurs. This report synthesises points of viewregarding priorities and recommendations asexpressed by the scientific community involved inthis ESF initiative.
4 Investigating Life in Extreme Environments
Investigating Life in Extreme Environments - A European PerspectiveEditor: Nicolas Walter • Contributors: Ricardo Amils, Arnoldus Blix, Michael Danson, Christine Ebel, Cynan Ellis-Evans, Françoise Gaill, Helmut Hinghofer-Szalkay, Kai-Uwe Hinrichs,
Francesco Loreto, Daniel Prieur, Farzam Ranjbaran, Klaus Valentin, Elisabeth Vestergaard, Nicolas Walter • Acknowledgement: The European Science Foundation would like to thankthe following scientists who provided valuable comments and inputs on the present report: Stéphane Blanc, Charles Cockell,
Terry Callaghan, Horst Felbeck, Claude Gharib, Yvon le Maho, Steven Scott.
5Investigating Life in Extreme Environments
Foreword
Executive summary and summary of recommendations 8
1. General considerations 13
1.1. Why investigating life in extreme environments is important 131.2. Defining the issue 151.3. The Earth and beyond 171.4. Questions arising 22
2. Scientific priorities 24
2.1. Priorities of cross-cutting relevance 242.1.1. Focus on model organisms and model extreme environments 252.1.2. An ecosystem-based approach 252.1.3. Molecular structural biology and genomics 26
2.2. Issues of specific interest 272.2.1. Microbial life in extreme environments 272.2.2. Life strategies of plants in extreme environments 282.2.3. Life strategies of animals in extreme environments 292.2.4. Human adaptation in extreme environments 30
3. Technological issues 31
3.1. Streamlining the issue 313.2. Crossing the borders 333.3. Collecting information – priorities 34
4. Economic and societal applications 35
5. Policy aspects – the European arena 37
5.1. European scientific landscape 375.2. The need for further interdisciplinarity 385.3. Networking and coordination 395.4. Programme issues 405.5. Public outreach and education 41
References 42
Appendices 45
I Contributors to the report 46
II Steering Committee of the ILEE initiative 47
III ILEE Workshop programme (6-8 November 2005) 48
Contents
6 Investigating Life in Extreme Environments
From the deepest seafloor to the highest mountain, from the hottest region to the cold Antarctic plateau,environments labelled as extreme are numerous on Earth and they present a wide variety of features andcharacteristics. The life processes occurring within these environments are equally diverse, not only depending onstress factors (e.g. temperature, pressure, pH and chemicals), but also on the type of life forms, ranging frommicrobes to higher species.
How is life limited by and adapted to extreme external biotic and abiotic factors? This key questionsummarises the deliberations raised by this exciting and fascinating research area. Addressing the challenge ofanswering this question would help to reveal new insights and refine theories concerning the origin and evolution oflife on our planet, as well as life beyond Earth. Investigating life processes under extreme conditions can also providenew clues towards understanding and predicting ecosystems’ responses to global change. Furthermore, this area ofresearch has a wide application potential in the fields of (bio)technology, chemical and pharmaceutical industry,biomedicine and cosmetics.
Depending on the type of organism studied, addressing extreme environments presents different situations andissues to be considered. However, some transversal priorities stand out and have been identified in this report. Thescientific community is calling for:
• More consideration to be given to the ecosystem-based approach when studying life in extremeenvironments;
• The identification of model organisms in different phyla and for different extreme environments to focusthe research effort and provide a common framework for research activities;
• The use of molecular structural biology and genomics tools.
Looking at the European scientific landscape, the research community is aware of the benefits that would be derivedfrom enhanced information flow between communities with different interests, as well as with the technological sideof their research. It is apparent that an interdisciplinary approach should be extensively promoted andsupported when it comes to examining life processes in extreme environments. It is also important to acknowledgethat scientific outcomes could benefit considerably from better networking and coordination amongscientific communities involved in these areas of research.
The European Science Foundation’s (ESF) report Investigating Life in Extreme Environments is the result of a processthat began in February 2004 with a call for Expressions of Interest addressed to the European scientific community.The high level of response convinced the ESF that the subject was an emerging interdisciplinary research domain thatdeserves attention.
In November 2005 a large-scale multidisciplinary workshop gathering 128 participants was organised in Sant Feliude Guixols (Spain) to debate and put forward the scientific case and needs when considering Life in ExtremeEnvironments in the broadest sense: from microbial life to human adaptation, from the depth of the oceans to outerspace and planetary bodies. Since the marine and polar science communities expressed a strong interest from theearly stages of the initiative, they were largely represented at the workshop and they were very active in providing theirinput and perspectives.
This report synthesises the findings expressed during the Sant Feliu de Guixols workshop. The content andrecommendations presented here are not intended to be exhaustive, but are rather a first expression of integratedissues which can be considered among other priorities to be identified in future initiatives.
While the recommendations are developed in the context of extreme environments, many may be of relevance toother, more conventional research domains. Their application would provide a greater leverage effect on the scientificand technical achievements when considering specifically research on life processes in extreme environments.
These outcomes and findings advocate for the development of further knowledge in this area, and also for structuringthe scientific landscape. This would catalyse scientific and technical achievements for the benefit of science andsociety. It is clear that additional steps and initiatives should be taken in the field of investigating life in extremeenvironments in Europe, furthering the efforts of this ESF initiative and broadening it to include areas yet to beaddressed.
Bertil Andersson Daniel PrieurCEO Chairman of the ESFEuropean Science Foundation Investigating Life in Extreme Environments
workshop
Foreword
7Investigating Life in Extreme Environments
Executive summary and summary of recommendations
General considerations
1. Coping with extreme conditions, microbes,plants and animals have demonstrated theability to adapt and metabolise under highenvironmental stress. Relying on specific andcomplicated mechanisms, some organismseven require such conditions to live and evolve.
2. For most environmental parameters constraininglife, specific limits have been established (but notdefinitely set) beyond which life, in its wholecycle, has never been described to date. Agiven environment, where one or moreparameters show values permanently closeto lower or upper limits known for life, maybe considered as an ‘extreme environment’.
3. When considering research activities andinvestigations on life in extreme environments,the overarching question is ‘How is life limitedby and has adapted to extreme externalbiotic and abiotic factors?’
4. Certain physical and chemical conditions ofsome Earth-based extreme environments couldbe considered analogous to our planet’senvironment in its early stages of developmentand to other planetary bodies. Investigating lifeprocesses in these environments helps toconsider questions of the origin of life on ourplanet and the presence of life beyond the Earth.
5. Earth is becoming increasingly exposed tosevere environmental modifications because ofanthropogenic activity. Changes on a global orregional scale are resulting in someenvironments becoming ‘extreme’ for thefunctioning of their existing ecosystems. It is alsoimportant to note that global change leads someextreme environments to become benign, withthe risk of losing biodiversity of those speciesused to extreme conditions. Investigation of themetabolism and adaptation processes of somespecies accustomed to extreme and highlyvariable environments are relevant in trying tounderstand the foreseeable impacts of globalchange on ecosystems.
6. The study of humans evolving in extremeenvironments provides new keys andunderstanding of psychological, physiological,social and societal functioning of individuals,groups and populations.
Priorities of cross-cuttingrelevance
7. When considering research activities onmicrobes, plants, animals or humans evolving inextreme environments, several transversalscientific priorities arise for the scientificcommunity. These priorities include: i) theidentification of model organisms evolving inextreme environments, and the identification ofmodel extreme environments; ii) the applicationof ecosystem-based approaches to scientificactivities in extreme environments; and iii)developing investigation of life processes inextreme environments through molecularstructural biology and genomics.
8. Model organisms and extremeenvironments - Recommendation:
• Identify and agree on i) model organisms indifferent phyla and for different extremeenvironments; and ii) model extremeenvironments, on which research activitiesacross Europe would focus. Not only wouldthese systems focus research activities in aconcerted manner but they would bring along-term perspective and an observatorycomponent to the scene. Providing acommon framework, they would also allowthe development of comparative studiesacross domains and disciplines(Recommendation 1).
9. An ecosystem-based approach applied toextreme environments - Recommendations:
• Favour an ecosystem-based multidisciplina-ry approach when considering scientific acti-vities in extreme environments. When dealingwith the complex interplay of biotic and abio-tic processes structuring ecosystems,emphasis should be given to describing theinterface between biology and geochemistry(Recommendation 2).
• All environments have some degree ofseasonal and spatial variability. Spatial andtemporal variability, including long-termmonitoring, need to be considered whenplanning and performing research activities inextreme environments (Recommendation 3).
• Evolution, dynamics and recovery ofecosystems in response to disturbances
8 Investigating Life in Extreme Environments
should be investigated to further understandcurrent and future factors influencingecosystems’ structure and functioning. Thisis of particular relevance when consideringglobal change and anthropogenic pressurescurrently put on the Earth’s environments(Recommendation 4).
10. Molecular structural biology andgenomics - Recommendations:
• Initiate efforts to integrate studiesencompassing genomics, genetics, cellularand molecular biochemistry, biophysics andstructural biology aimed at investigating life inextreme environments (Recommendation 5).
• Compare genomes and proteomes andidentify intracellular chemical and physicalparameters for related organisms or cellcommunities adapted to different extremeenvironments (Recommendation 6).
• Through functional and structural genomics,exploit the potential of the archaea and othermicroorganisms adapted to extremes astractable models for difficult-to-studyhomologous human and pathogenmacromolecular systems (Recommendation 7).
Scientific issues of specific interest
11. In addition to cross-cutting priorities and recom-mendations, several specific recommendationsare expressed concerning: i) microbial life inextreme environments; ii) life strategies of plantsin extreme environments; iii) life strategies of ani-mals in extreme environments; and iv) humanadaptation to extreme environments.
12. Recommendations on microbial life inextreme environments include:
• Investigate further the molecular basis of thephysiological adaptation of microbes toextreme environments, especially with regardto DNA, RNA and protection of cellularmetabolites (Recommendation M1).
• The deep subsurface (oceanic andcontinental) exhibits unique features in termsof temperature, pressure and availability ofenergy. Investigating the subsurface microbialprocesses further, especially with regard tocell division, would deepen the fundamental
understanding of life and concepts of cellmaintenance. Moreover, understanding therole of the subsurface in biogeochemicalcycles is essential for an assessment ofelement budgets on a global scale(Recommendation M2).
• Microbial extremophiles cover all threedomains of microbial life, bacteria, archaeaand eukaryotes. Comparative adaptation todifferent environmental stressors wouldprovide a very useful insight into the natureand evolution of extremophilic organisms(Recommendation M3).
13. Recommendations on life strategies of plantsin extreme environments include:
• Give focus to boosting progress in molecularand biochemical studies and to unravelling themechanistic basis of plant life under extremeconditions (Recommendation P1).
• Study the strategies of adaptation (from themolecular to the ecosystem level) which allowplant life in extreme environments, as well asplants’ strategies of reproduction anddissemination in their long-term efforts toavoid environments becoming hostile(Recommendation P2).
• When considering studies on plant life inextreme environments, a particular attentionshould be given on fragile ecosystems,including ecosystems that are starting to becolonised by plants, or that are becomingunfavourable to plant life, especially as aconsequence of global change and human-driven land use change. Study strategies ofsurvival at this boundary that may be – or maybecome – of utmost importance for plantsurvival, for plant loss of diversity, and for theentire food-chain based on plant life(Recommendation P3).
14. Recommendations on life strategies ofanimals in extreme environments include:
• Comparative studies are needed to determinethe specificities of animals, communities andecosystem responses to different types ofenvironmental stress (Recommendation A1).
• Favour characterisation studies on extremeenvironments and their interactions withanimals; this would help in understandingadaptation processes (RecommendationA2).
9Investigating Life in Extreme Environments
• The analysis of the biodiversity dynamics ofanimal communities and ecosystems found inextreme environments, includingpalaeontological approaches, has to bepromoted to understand the evolution of life inextreme environments (Recommendation A3).
• In vivo approaches simulating the naturalenvironment are needed to help in theknowledge and prediction of the tolerance ofanimals to environmental changes(Recommendation A4).
15. Recommendations on human adaptation inextreme environments include:
• Bring together humanities, social sciences,psychology and medical sciences under thelabel of Human Sciences in studies on humanadaptation to extreme environments(Recommendation H1).
• Issues of environmental monitoring andcontrol have an important role both in artificialand confined environments as well as innatural environments. These aspects shouldbe further investigated when consideringhumans, not only because the environmentinfluences them, but also because there is anadaptive effect that humans impose upon theenvironment (Recommendation H2).
• When considering human species, favourcross-disciplinary dialogue and interactionswhile dealing with different levels of systemshierarchy at the same time.(Recommendation H3).
• Address systematically and carefully ethicalissues when considering human adaptation toextreme environments (Recommendation H4).
Technological issues
16. Technology sees itself as a clear cross-cuttingissue of major relevance and its barriers as oneof the main limiting factors for the developmentof knowledge on life processes in extremeenvironments. Recommendations ontechnological issues include:
• Develop interdisciplinary initiatives to defineenvironment-specific technological andtechnical requirements, their availability anda road-map for their development(Recommendation T1).
• Support the development of initiatives aimingto improve information exchange onavailable technologies and new
developments throughout the scientificcommunity involved in extreme environments(Recommendation T2).
• Develop and support mechanisms to facilitateaccess to and sharing of existing and futuretechnical platforms and infrastructures to beused when accessing extreme environments(Recommendation T3).
• Consider further the enhancement of theEuropean simulation capability in the field ofextreme environments. Relevant simulationfacilities (e.g. microcosms) should bedeveloped (Recommendation T4).
• Develop in situ sampling, measurement andmonitoring technologies (e.g. light-weight,low-power, and robust sensors) and capacity(e.g. in situ observatories and bio-logging).Categorisation and inventory of existingsystems, technologies and techniques arealso recommended. (Recommendation T5).
Economic and societalapplications
17. Extremophilic species may have an importantrole to play in improving and creating someindustrial processes. Furthermore, theypotentially allow the development of newproducts.
18. Development of high-quality and robustfacilities and technological capabilitiesdeveloped for scientific activities in extremeenvironments brings together industries andresearch institutions. Through their involvementin such activities, they develop and gainknowledge, competencies and capabilitiesrelevant to a wide range of industrial andeconomic sectors.
Policy aspects — the Europeanarena
19. At the international level, Europe currently leadsin areas such as polar research, biodiversity,and global change, although without a specificfocus on extreme conditions. Polar and deep-sea research fields appear to be areas ofexcellence in Europe. In considering the type oforganisms, the study of microbes, plants andanimals are reasonably mature areas ofresearch.
10 Investigating Life in Extreme Environments
Executive summary and summary of recommendations
20. A strong benefit can be expected frominternational collaboration. This is of particularrelevance in areas of global significance.
The need for furtherinterdisciplinarity
21. Compartmentalisation between disciplines andtheir associated technological developments isseen as a major hurdle to greater efficiency inscientific activities, even if this statementapplies to a whole range of scientific activities,it is of particular relevance when consideringscientific and technological activities in extremeenvironments. Therefore, interdisciplinarity andmultidisciplinarity between scientific domainsand between the technological and scientificspheres is identified as a major source ofpotential increase in scientific achievements.When relevant, these aspects should beextensively promoted and supported as muchas possible when considering life processes inextreme environments (Recommendation 8).
Networking and coordination
22. While high-level expertise and know-how havebeen developed at the European level, thefragmentation of this knowledge and the lack ofcoordination across initiatives are majorconstraints to the further development of apan-European capacity. Therefore, it isrecommended:
• To create as soon as possible an overarchinginterdisciplinary group of experts to definethe necessary actions to build a criticalEuropean mass in the field of InvestigatingLife in Extreme Environments(Recommendation 9).
• That effort should be made to support andimprove information exchange, coordinationand networking of the European communityinvolved in scientific activities in extremeenvironments. This could be achievedthrough a Coordination Action supported bythe European Commission and include thedevelopment of a Web portal and associateddevelopments (database of experts,dissemination of information). In this context,large-scale multidisciplinary events (e.g.conferences, fora and workshops) should be
organised regularly in order to build andmaintain momentum at the European level.This should be done in the context of thenumerous specialised events that are alreadyorganised (Recommendation 10).
Programme issues
23. Overall, and especially because of theirspecificities and positioning at the frontier ofscience, research activities dealing with lifeprocesses in extreme environments raisesissues such as: i) the long planning horizonsthat are necessary for field expeditions and thedifficulty for the scientific community tobecome part of them; ii) the cost of carrying outscientific activities in such environments; and iii)the difficulty in accessing national researchresources. It is recommended that:
• Extreme environments are integrated intonational science strategy and clearly identifiedin national and European scientific calls andprogrammes (Recommendation 11).
• When considering scientific activitiesinvolving extreme environments, fundingagencies should ensure that theirassessment procedures can adequatelyreview proposals involving a multidisciplinaryapproach as opposed to the more typical,strongly focused proposals(Recommendation 12).
Public outreach and education
24. It is important to inform the public on issuesrelated to extreme environments, highlightingthe exotic nature of these environments, theirbiota and their societal and economic potential.
25. It is fundamental to secure the transmission ofextreme environments scientific knowledge tostudents and young scientists.
11Investigating Life in Extreme Environments
12 Investigating Life in Extreme Environments
1. General considerations
1.1. Why investigating life inextreme environments isimportant
Life and extreme environments
To our present knowledge, the Earth seems to bethe single habitable and inhabited planet within thesolar system. The concept of habitability is oftendefined in terms of the requirements for humanexistence. However, that part of the Earth’sbiosphere permanently inhabited by human beingsis rather small and most of the planet, its deep coreor mantle, will clearly never see a living organism. Inbetween these two zones (inhabited anduninhabited), a variety of environments exist wherehuman beings cannot live permanently, or
physically access, although other forms of life existwithin them.
Coping with these latter environments, microbes,plants and animals have demonstrated the ability toadapt and metabolise under high environmentalstress. Relying on specific and complicatedmechanisms, some organisms even require suchconditions to live and evolve.
From the bottom of the seas to the highestmountains, from the hottest regions to the coldAntarctic plateau, environments labelled as‘extreme’ are numerous on Earth and they presenta wide variety of features and characteristics. The
life processes occurring within these environmentsare equally diverse, not only depending on stressfactors but also on the type organisms, rangingfrom microbes to higher species.
Polar regions, outer space or deep-sea, theseenvironments have always triggered humancuriosity and their exploration has fascinated notonly scientists but also the general public.However, they have been explored or sampled onlyvery recently, mainly because of earlier technicallimitations. This exploration is still in its infancy andnovel organisms (e.g. extremophilic microbes),novel habitats (e.g. the deep subsurface, subglaciallakes or unexplored regions of the seafloor) andnovel technological applications (e.g.bacteriorhodopsine for optical computers andinformation storage, acidophilic chemolithotrophicmicroorganisms for biomining and polymerasechain reaction) continue to be discovered.
Furthermore — and this is a key point aboutextreme environments — because of their particularcharacteristics and features, they provide a widerange of unique natural laboratories for thescientific community. Similarly unique are themultidisciplinary aspects of means andinfrastructures that are utilised to conduct scientificactivities within such environments. For example,polar stations are designed to facilitate a largespectrum of scientific activity, including biology,geology, atmospheric or medical research whilstunderwater Remotely Operated Vehicles (ROVs)can be used for geological as well as biologicalsurveys and sampling.
13Investigating Life in Extreme Environments
Hydothermal vent — 2 ,630m deep
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The relevance of studying life processes inextreme environments
Certain physical and chemical conditions of someEarth-based extreme environments could beconsidered analogous to our planet’s environmentin its early stages of development. Extending ourknowledge of life processes in conditions broadlycomparable to the paleo-environments of Earthwould help reveal new insights and refine theoriesabout the origin of life and how it has evolved overtime on a constantly changing planet. In thiscontext, attention should be given to fossil lifeforms such as microorganisms brought back to lifefrom a state of deep sleep/dormancy for hundredsof thousands of years in ice (ice cores from LakeVostock), a million years in permafrost, 25-40million years in amber and an astounding 250million years in salt crystals [1.1].
Furthermore, consideration should also be given topast extreme environments that resulted in massextinctions and new evolution events (e.g. whenoceans were anoxic, when there was darkness andlow temperatures following the meteorite crash intothe Gulf of Mexico, and when UV-B radiation waslikely to be exceptionally high following massivevolcanic activity of the Siberian plates). Much canbe learned from these episodes.
Not only are extreme environments thought toprovide some analogy with those existing on theearly Earth, but also some can be considered toshow some similarities with other planetary bodies.One salient example of this is the analogy betweenAntarctica’s subglacial lakes and the Jovian icymoon Europa [1.2], where a large amount of liquidwater is thought to be present under a thick layerof ice. Investigating life processes in extremeenvironments may provide potential scenarios forpossible extraterrestrial life, thus helping toconsider that major question: Is there life beyondthe Earth?
At present, planet Earth is increasingly exposed tosevere environmental modifications, because ofanthropogenic activities. Changes on a global orregional scale are resulting in some environmentsbecoming ‘extreme’ for the functioning of theirexisting ecosystems; examples include acidificationof the oceans and more frequent extreme climaticevents such as severe droughts or flooding. It isalso important to note that global change leadssome extreme environments to become benign,with the risk of losing biodiversity of those speciesused to extreme conditions. In this context,investigation of the metabolism and adaptationprocesses of some species accustomed toextreme and highly variable environments arerelevant in trying to understand the foreseeableimpacts of global change on ecosystems.
The high stress level that can be induced in livingorganism by extreme conditions leads to thedevelopment of mechanisms that mitigate thepotential negative effect of such environments.Depending on the stress factor, some mechanismsare fundamental for survival and are extremelydeveloped and active. While in more conventionalconditions they would be hardly noticeable, thesephysiological mechanisms are then much morevisible and identifiable and can be better studied inextreme environments.
14 Investigating Life in Extreme Environments
Spectra from different Earth-like planets
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1. General considerations
The study of humans evolving in extremeenvironments provides new keys to theunderstanding of psychological, physiological,social and societal functioning of individuals andgroups. The isolation and hazardousness ofexternal environments are sources of majorpsychological stress; e.g. microgravity and the levelof physical activities in confined areas havephysiological impacts. Studying these aspects andtheir feedback mechanisms is fully relevant toscientific activities in extreme environments as theynot only provide information about humanfunctioning but also because scientific activities inharsh conditions are partially dependent on thehuman presence and its ability to perform research.
Indeed, other human groups than scientificpersonnel are evolving in environments that can beconsidered as extreme, arctic populations being anevident example. In this perspective, it is importantto note that it is not only physico-chemical factorsthat can lead an environment to be extreme for itspopulation, but also political, social and culturalfactors. When considering populations evolving insuch environments a particular feature is the highdegree of transdisciplinarity and multidimensionalityof the research involved. Understanding themechanisms which make an impact on populationsand individuals, studying their response and tryingto develop mitigation means is extremelychallenging and requires commitment from thesocial and human sciences communities.
On the applications side, industrial processes ofteninduce conditions that are extreme in terms of pH,temperature and pressure. In these conditions, thedevelopment of extremophilic enzymes may havean important role to play in improving the efficiencyof these processes and could even form the basisof new ones (e.g. for the textile or the food-processing industries). Furthermore, studies ofextremophilic species and their properties haverevealed features that potentially allow thedevelopment of new products in various fields suchas cosmetics, food stabilisation, oil spill response,de-pollution or pharmaceutical applications.
Overall, extreme environments have features thatare challenging for any research activity and theirassociated technological development. At theforefront of the technological challenges inducedby extreme environments are issues of automationof sampling, monitoring and analysing equipment,miniaturisation of sensors and robotics. Industriesand research institutions involved in generating
such capabilities are developing competencies andknow-how applicable to a whole range of highadded-value activities.
It is important to foster and catalyse researchactivities on life processes in extremeenvironments, not only because they are applicableto some of the main scientific questions such asunderstanding the basics of life processes, theorigin of life or the potential existence ofextraterrestrial life, but also because the economicpotential behind these activities is important. For allthese reasons, ‘investigating life in extremeenvironments’ represents a major challenge forEarth’s inhabitants at the beginning of the 21stcentury.
1.2 Defining the issue
Life, as we know it, is constrained by a series ofenvironmental parameters (physical and chemical),whose values may vary according to space andtime. Combinations of specific ranges for each ofthese parameters within a particular area definebiotopes, or sets of environmental conditions,suitable for a given species. In its habitat, a givenspecies carries out its complete life cycle.
For most environmental parameters constraininglife, specific limits have been established (but notdefinitely set) beyond which life, in its whole cycle,has never been described to date (Table 1). Forexample, the highest temperature allowing thecomplete life cycle of an organism (the archaeonPyrolobus fumarii) is 113°C, the lowest is -18°C. Agiven environment, where one or moreparameters show values permanently close tothe lower or upper limits known for life may beconsidered as an ‘extreme environment’.
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More generally, environments with parameters thatput some particular species under threat(approaching its own limits), can be considered as‘extreme’. This is of particular relevance whenconsidering global change issues, rapidenvironmental change such as drought or floodingand exceptional conditions; these issues are alsoconsidered in this report.
Extreme conditions are a requirement for someorganisms to carry out their complete life cycle;some organisms have developed mechanisms tosurvive (without cell division or reproduction) forshort or long (even very long) periods in extreme
physico-chemical conditions, that mostly consist ofdormancy stages. It is interesting to note that notonly microorganisms, spores, seeds or cysts areable to go into a dormancy stage but also higherspecies such as tardigrades or nematodes – eventhe African lungfish can survive for years withoutwater in dried up wadis of North Africa.
Overall, it is important to note that whenconsidering extreme environments, Homo sapiens,represents a particular case as it has developed avariety of technologies to adapt more or lesspermanently to extreme conditions.
Environmental Known lower limit Known higher limit Commentsparameters for life for life
Temperature -18°C [1.3] 113°C [1.4] Mean human body temperature 37°C
Pressure 10-14 Pa [1.5] 130 MPa [1.6] Atmospheric pressure: 101 KPa
pH Negative [1.7] 13.2 [1.8] Neutral pH: 7
Salt concentration None 5 M [1.6] Human blood saltiness: 3.5 %
Water activity 0.7 [1.9] 1.0 [1.9]
Light Darkness Variable High light levels can cause photoinhibition and free radical formation
Highly energetic None 1.5 MRad [1.10] 1.5 MRad is 3,000 times radiation the lethal level for humans
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1. General considerations
Table 1: Examples of parameters constraining life processes
1.3. The Earth and beyond
A few typical extreme environments can be identified; the tables below (not exhaustive) describe them along withthe environmental stressors that lead them to be considered as ‘extreme’.
Location: Between 700 and 10,000m deep in seas and oceans
Environmental stressors:• Temperature Low – between 2°C and 4°C
• Pressure Very high – up to 1,000 MPa
• Light Non-existent
Comments:- Broad gradients of parameters on a large spatial scale.- World’s deepest seafloor is at 10,898 m below the ocean surface (Mariana trench).- Bacteria [1.11] and eukaryotes [1.12] have been found in the deepest depths of the oceans.
Marine extreme environments
Deep-sea
Location: Sparsely distributed on geoactive seafloors at different depths (Somevents are found in shallow water)
Environmental stressors:• Temperature Very high at the vent chimney - up to 420°C while low in the
surroundings
• Pressure High to very high - depending on the vent’s depth
• Light Low to non-existent
• Chemical stressors High concentration of chemicals dissolved in the vent fluids, and lowoxygen concentration
• pH Low
Comments: - Steep gradients of parameters on a small spatial scale. - High concentrations of potentially toxic chemicals. - The most thermo-tolerant organism, the archaea Pyrolobus fumarii, was found in such
environments. [1.4]
Hydrothermal vents
Location: On the leading edge of a continental plate where it is colliding withan oceanic plate (e.g. western Pacific subduction zones) or on thetrailing edge where continental crust is rifting (e.g. Atlantic passivemargins).
Environmental stressors:
• Temperature High, because of tectonic activity in active marginsLow in passive margins
• Pressure High to very high - depending on the depth
• Light Non-existent
• Chemical stressors High concentration of chemicals, low oxygen, high salinity brines
Comments:- Hosts various types of geobiological features such as benthic ecosystems fuelled by cold seeps
and/or mud volcanoes. - Dynamic processes impacting the environment and disturbing the animal communities and more
generally the existing ecosystems.
Continental margins (active and passive)
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Location: Beneath the continental surface or seafloor
Environmental stressors:
• Temperature Low to very high
• Pressure High to very high
• Light Non-existent
• Chemical stressors Depending on the geological context
Comments:- Extremely low energy availability and long-term life processes.- Bacteria found 3,200 m beneath surface. [1.13]
Marine and terrestrial extreme environments
Continental and seafloor subsurfaces
Location: Areas of the globe surrounding the poles, north of the Arctic circle, orsouth of the Antarctic circle.
Environmental stressors:
• Temperature Very low – world’s coldest record: -89.2°C in Vostok station inAntarctica
• Dryness Very high in mainland - mean humidity 0.03% in Antarctica dry desert.
• Light Non-existent during the 4-month winter in latitudes higher than thepolar circles
• High energetic High due to the characteristics of the Earth magnetic field at theseradiations latitudes
Comments: - Relevant for human adaptation studies.
Terrestrial extreme environments
Polar regions
Location: On the seashore, between low tide and high tide level, at the junctionof sea, atmosphere and lithosphere
Environmental stressors:
• Temperature High variability of temperature
• Light High variability of exposure
• Dryness High variability of the level of humidity
Comments: - Very dynamic environments, strong water flow.
Intertidal coastal areas
Location: On the ground, in lakes or in the sea
Environmental stressors:
• Chemical stressors High concentration of salt
Comments:- Different ions according to geological context. - For terrestrial hypersaline media, intense evaporation, high UV and high temperatures
because of their location.
Hypersaline medium
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Location: Sparsely distributed on the Earth’s surface, usually in geoactive zone
Environmental stressors:
• Temperature High to very high
• Chemical stressors High concentration of chemicals
Hot springs
Location: In mountains, above 1,500m altitude, up to more that 8,800m
Environmental stressors:
• Temperature Low – decreasing with altitude
• Atmospheric pressure Low atmospheric pressure – decreasing with altitude
• Dryness High – increasing with altitude
• High energetic High – increasing with altituderadiations
Comment: - Relevant for human adaptation studies.
High altitude (and glaciers)
Location: Sparsely distributed on the Earth’s surface. For example the Saharaor Atacama deserts.
Environmental stressors:
• Temperature High and wide daily amplitude – 57.7°C measured at El Azizia in Libya
• Dryness High – constant lack of water
• Light High solar radiation pressure
Comment: - Relevant for human adaptation studies.
Hot arid regions and deserts
Location: Sparsely distributed on the Earth’s surface, often following volcanicactivities and metallic mining activities
Environmental stressors:
• Temperature Moderate to high
• Chemical stressors High concentration of heavy metals
• pH Low (<7) to negative – Hyperacidic media have a ph<<1
Comment: - Some of these extreme environments are the product of microbial activity.
Acidic environments
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Location: Sparsely distributed on the Earth’s surface. Soda lakes and sodadeserts are typical examples
Environmental stressors:
• Temperature Depending on the location
• Chemical stressors Usually high concentration of salt and sodium carbonate (a fewalkaline lakes are non-saline)
• pH High: between 7 and 14 – Hyperalkaline media have a pH>10
Comments: - Microbial life up to pH 13.2, some fish and shrimps evolve at pH 10.
Alkaline environments
Location: Anywhere on Earth where human activity has induced major changes
Environmental stressors:
• Temperature Global warning induces increased temperatures
• Light Enhanced UV-B radiation because of ozone layer depletion
• Chemical stressors High concentration of polluting chemicals (heavy metals,hydrocarbons)
Comments: - Wide variety of disturbances, with local (e.g. mercury concentration in Amazonian rivers or acid mine
drainage) and global (acidification of the oceans or global warming) impact, including over-exploitedecosystems.
Environments strongly shaped by humans
Location: In outer space, large celestial masses of rock, metal, or gas orbitingaround a star.
Environmental stressors:
• Temperature All ranges of temperature from extremely low (less than -200°C) toextremely high (more than 300°C) – High variability because ofrotation
• Pressure From null to very high, depending on the atmosphere
• Light From very low to very high exposure, depending on the position withregards to the star
• High energetic Typically very high, because of cosmic rays and central star’s radiations radiations
Comment: - Wide variety of environments, some include liquid water, as probably under the thick ice layer of the
Jovian Moon Europa.- Different level of gravity, depending on the mass.
Planetary bodies
Outer space
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Location: In outer space, spacecraft, evolving in microgravity.
Environmental stressors:
• Temperature All range of temperatures from extremely low to very high on externalexposed facilities
• Pressure Null on external exposed facilities
• High energetic From high in shielded areas to very high on external exposedradiations facilities.
Comment:- Unique characteristic: Very low level of gravity (impossible to be simulated for a long period on
Earth). - Space vessels are confined and isolated environments – relevant for human adaptation studies. - Bacillus subtilis can survive six years of exposure to the space environment (dormancy stage).
Space vessels/space environment
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1.4. Questions arising
When considering research activities andinvestigations on life in extreme environments, theoverarching question is: ‘How is life limited byand how has it adapted to extreme externalbiotic and abiotic factors?’ Considering thisquestion and investigating this field involvesdeveloping new approaches, methods and tools aswell as fostering those already in use; e.g. inaccessing to extreme environments, sampling, insitu monitoring, laboratory simulation, or highperformance analysis.
At a more detailed level, several fundamentalquestions arise that are relevant to any life formsfrom microbes to vertebrates.
What limits life on Earth in general?
This encompasses an understanding of what limitslife under extreme conditions of e.g. temperature,salinity, pressure, pH, chemical composition of theenvironment or radiation (including light/darkness)and understanding how extremophilic organismsare different from conventional organisms.
What limits life with respect to certain groupsor processes?
Considering this issue requires the use ofdisciplines such as molecular biology, ecology andphysiology in order to understand how speciescolonise new environments or confront newstressors (e.g. high concentration of metals in soilsand water, salinisation of soils and urbanenvironment pollution).
All stages in the life cycle and the total life cycle asa whole need to be investigated to answer thisquestion.
What limits individual species and to whatextent can they usually acclimatise or adapt?
This encompasses the adaptation mechanismsassociated with specific extreme environments:how have life forms been shaped by evolution tocope with hostile conditions? This question is ofparticular relevance when considering that globalchange leads some environments to becomehostile for their living organisms (e.g. oceanacidification, enhanced UV radiation in polarregions, extension of deserts, pollution, reductionin sea ice and warming of the ocean). Investigatingthe capacity of species to face variations in theirenvironmental conditions would allow a definition ofthe limits of these adaptive mechanisms and is akey to predicting future patterns of biodiversity lossor increase or ecosystem change. In this context itis important to consider not only adaptation tosingle stressors but also adaptation tomultistressors because some environmentsprovide a combination of environmental stressors.
What limits communities or ecosystemsevolving in extreme environments; i.e. whichparameters shape an ecosystem so that anychange may drastically alter that ecosystem?
Some characteristic extremophilic species (e.g.resurrection plants of arid environments oracidophilic chemolithotrophs from acidicenvironments) have been investigated in somedetail but little is known about interactions amongspecies; e.g. multi-trophic relationships involved inlife in extreme conditions. A multi-disciplinary effortis needed to characterise ecosystems in extremeenvironments. A particular issue would be toinvestigate how extremophiles shape theirenvironment, another would be to compare thefunctions of ecosystems across a variety ofextreme environments. (See also Section 2.1.2).
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What happens to species facing a newenvironment?
This encompasses the study of the molecular andbiochemical bases of response to conditionsinvolving factors and stresses not yet experienced.A comprehensive understanding of the survivalstrategies of species through comparativegenomics and proteomics would provide biologicalinsights. These insights would not only involve newaspects of physiology and evolution that may beunique to extremophiles, but could also uncoversome unknown adaptations required for life instressful environments that may be shared acrossall domains of life. A special focus should be givento adaptations to absence of gravity, lowtemperature, low pressure, pollution and unusualradiation.
Besides these life process-focused issues, extremeenvironments also raise questions per se:
In what way are certain extreme environmentsanalogues for ancient terrestrial and extra-terrestrial environments?
The study of current life forms adapted to extremeconditions will certainly bring insights to the earlyhistory of life on Earth and its potential analogues inthe universe. First, it is recognised that hostileconditions (temperature, radiations and perhapshigh salinity) prevailed while life emerged on Earth.Second, planetary exploration missions providedevidence for internal water, high salt conditions andlow temperatures in the solar system, analogous toconditions found in certain areas on Earth; e.g.polar regions or the deep hypersaline ecosystemsfrom the seafloor of the Mediterranean Sea. It isalso important to mention that salt-adaptedorganisms display a remarkable ability to withstand
dehydration, multiple stresses and to survive overgeological time when trapped in salt crystals.Understanding the mechanisms responsible for thisrobustness may guide the search for traces of lifeon other planets.
What is the role of a particular extremeenvironment for regional/global ecology andelement cycles?
So far, we do not understand the role of extremeenvironments in global biogeochemical cycles. Forexample, the contribution of the extreme marineenvironments to budgets of climatically relevantgasses (e.g. methane hydrates in continentalmargins) is largely unknown. Extreme environmentscan be sources of elements potentially hazardousto the hydrosphere and atmosphere, on theregional to global scale. Strong interactionsbetween living organisms and inanimate matter,such as those that have shaped the Earth systemover geological time, can play a fundamental role inbuffering these emissions. These interactions alsodrive the functioning of unique ecosystems,sustaining energy transfer from the geosphere tothe biosphere.
An additional consideration is:
What is the potential societal and economicvalue of extreme adaptation?
Organisms living in extreme environments mustcope with higher levels of stressors and mutagenicagents than normal. Consequently, they arethought to have developed mechanisms thatconfer a high degree of resistance to damagingagents. Investigating these properties brings newpotentialities of relevance to industrial processesincluding biotechnology, pharmaceuticalapplications or cosmetics that could be used forthe benefit of human health, economic growth andsustainability. (See also Section 4)
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Life under extreme conditions is a reality for awhole range of life forms. When considering futureresearch activities on microbes, plants, animals orhumans evolving in extreme environments, severalscientific priorities arise for the Europeancommunity involved. Among the priorities identifiedby the participants to the ESF workshop, three arecross-cutting to the study of microbes, plants andanimals (and to a lesser extent, humans). Inaddition, scientific priorities specific to each ofthem have also been identified. From priorities,recommendations have been developed.
2.1 Priorities of cross-cuttingrelevance
2.1.1. Focus on model organisms and modelextreme environments
Since the beginning of the 20th century, theDrosophila fly has been workhorse of genetics anddevelopmental biology research. This modelorganism is practical to study and work with andhas allowed major scientific progress. A similar,while more detailed, approach should be adoptedwhen considering life processes in extremeenvironments. Some species (microorganisms,animals or plants) are well adapted to theirenvironment and may be considered as accuraterepresentatives for understanding adaptationmechanisms and survival strategies in a givenextreme environment.
Such species may encompass a wide range ofcharacteristics, including the originality of molecularmechanisms, the specificity of some cellmetabolism, the characteristics of variousphysiological responses, or the links betweendifferent stress responses and the regulation of thegenomic activity. Focusing and convergingresearch on such model species at different stagesof their life cycle would reveal a large amount ofknowledge on the mechanisms and strategiesdeveloped to cope with extreme environments.
Depending on the parameter considered, a numberof model organisms are available. Considering hightemperatures, examples could include thepolychaete worm Alvinella pompejana fromhydrothermal vents which is considered as the
most thermotolerant metazoan known to scienceand which can survive in highly-pressurisedaquariums [2.1], or the highly thermotolerant desertant Cataglyphis bicolour (that is easier to collect).Other examples could include the archaeonHaloferax volcanii, which is adapted to high-saltconditions, or the extreme acidophilic archaea,Ferroplasma acidophilum, capable of growing atnegative pH and that has been recently isolatedfrom an acidic mine site. Even if this would raiseinherent difficulties and require mobilisation ofresources and expertise, there is a need for thecommunity to agree on a set of model organismsto be further studied.
A similar assessment scheme can be establishedfor specific types of extreme environments. Thiswould allow some questions to be considered for abroad range of species; e.g. is the physiology ofspecies inhabiting the polar regions similar indifferent geographical areas? What is the range ofthe metabolic response of polar organismsregarding a specific function (e.g. reproduction)? Isthere a large diversity of responses depending onthe species considered?
2. Scientific priorities
Recommendation 1: Identify and agree on i) model organisms in different phyla and for different extreme environments; andii) model extreme environments, on which research activities across Europe would focus. Not only wouldthese systems focus research activities in a concerted manner but they would bring a long-termperspective and an observatory component to the scene. Providing a common framework, they wouldalso allow the development of comparative studies across domains and disciplines.
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2.1.2. An ecosystem-based approach appliedto extreme environments
Single-species studies have considerable merit butit is increasingly recognised that greater value willbe obtained from research that acknowledges theecosystems within extreme environments and theinterdependence of the various ecosystemcomponents. This is referred to as ‘ecosystem-based research’ applied to extreme environments.Though identified by the scientific communityinvolved in the ESF initiative profiled here, therehave been relatively few examples to-date of anecosystem-based approach being applied toextreme environments [2.2]. This is in part becauseof the often considerable difficulties in accessingand researching these environments. Also, extremeenvironments have been subjected to moreextensive studies only in relatively recent times.
In many cases, there is no detailed inventory of thebiodiversity of extreme environments or fulldescription of the temporal and spatial variability oftheir abiotic parameters. Nevertheless, it isrecognised that understanding how various lifeforms survive at limits and interact with otherorganisms and their environment requiresrecognition of the ecosystem itself and where thetarget organisms fit into this system.
It is also pertinent to mention that there is limitedexperience within funding agencies to effectivelyassess approach. Ecosystem-based research is bydefinition multidisciplinary and may also involveconsiderable technical and logistical resources, sofunding bodies need to clearly identify theseelements in programme calls for proposals so thatreviewers are properly briefed (see also Section 5.2).
Recommendation 2: Favour an ecosystem-based multidisciplinary approach when considering scientific activities in extremeenvironments. When dealing with the complex interplay of biotic and abiotic processes structuringecosystems, emphasis should be placed on describing the interface between biology and geochemistry.
Recommendation 3: All environments have some degree of seasonal and spatial variability. Spatial and temporal variability,including long-term monitoring, need to be considered when planning and performing research activitiesin extreme environments.
Recommendation 4:Evolution, dynamics and recovery of ecosystems in response to disturbances should be investigated tofurther understand current and future factors influencing ecosystem structure and functioning. This is ofparticular relevance when considering global change and anthropogenic pressures currently put on theEarth’s environments.
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2.1.3. Molecular structural biology andgenomics
The investigation of species and processes throughmolecular genetics, biochemistry, physiology,biophysics and structural biology has the potentialto enlarge and deepen our knowledge of cellularand molecular adaptation processes. In thisrespect, the extremophile research can be dividedinto three broad areas: i) cellular adaptation toextreme conditions; ii) molecular adaptation toextreme conditions; and iii) extremophilic molecularsystems as tractable models for complexhomologous human and pathogen systems.
Another important aspect is tolerance. Forexample, to cope with low pH some cells haveadapted to pump out H+ from the cell. However,some cellular structures can be just tolerant of, forexample, high salt. Understanding the boundarybetween tolerance and adaptation in differentorganisms would be useful.
Mechanisms of cellular adaptation to extremeconditions
All cells protect themselves against stressors byinducing systems to rescue or eliminate damagedproteins (chaperones and proteases), to protect thegenetic material (DNA repair systems), to eliminateunwanted molecules (transporters, detoxificationsystems) or to stabilise molecules and metabolites(compatible solutes etc.). Extremophiles areexcellent models to identify novel protectionsystems and non-conventional metabolicpathways. For this purpose, genomes, proteomes,interactomes and metabolomes from organismsadapted to a wide range of environments shouldbe compared at an integrated level. Expectedoutcomes are the characterisation of novelmolecules and metabolic pathways that stabilisecellular macromolecules.
Understanding molecular adaptation toextreme conditions
Life under extreme conditions is limited primarily bythe stability of macromolecules and their ability toexert their biochemical functions under non-conventional conditions that normally lead to thecollapse of structures or to functional inactivation.Very little is known about the mechanisms involvedin these processes. Interdisciplinary studiescombining comparative genomics, enzymology,structural biology and biophysics should reveal themolecular strategy and the evolutionary processesthat shaped macromolecules from the origin of lifeto present environmental constraints.
The expected outcomes include: i) anunderstanding of the structural and dynamicrequirements of macromolecules for function; ii) adefinition of the physicochemical limits for life; iii) aguide for research in astrobiology and origin-of-lifestudies; and iv) strategies for rational modificationsof enzyme or pharmaceutical compounds inindustrial processes.
Structural and functional genomics onextremophilic systems
Extremophilic organisms represent a uniqueresource characterising novel molecular systems.Those originating from extremophilic archaea are ofspecial interest since they represent ancestral,simpler versions that can be models forhomologous systems in eukaryotes (human) orbacterial pathogens. Archaeal genetic tools haverecently been developed and we can expect thediscovery of unknown interacting molecules andcellular processes for biotechnological and medicalapplications [2.3 and 2.4].
Recommendation 5: Initiate efforts to integrate studies encompassing genomics, genetics, cellular and molecularbiochemistry, biophysics and structural biology aimed at investigating life in extreme environments.
Recommendation 6:Compare genomes and proteomes and identify intracellular chemical and physical parameters forrelated organisms or cell communities adapted to different extreme environments.
Recommendation 7: Through functional and structural genomics, exploit the potential of the archaea and othermicroorganisms adapted to extremes as tractable models for difficult-to-study homologous human andpathogen macromolecular systems.
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2. Scientific priorities
2.2. Issues of specific interest
2.2.1. Microbial life in extreme environments
Microbes living in extreme environments have anumber of special features that provide fascinatinginsights into the nature of cellular and biochemicaladaptation to extreme conditions, whilst beingsome of the most exploitable of extremophilicorganisms for biotechnological processes. Inaddition, contemporary extreme environments areuseful analogues of environments existing on earlyEarth and on other planetary bodies. Thephylogenetically most deeply branching microbeson Earth proliferate in extreme environments.Studying these organisms and their habitats maythus provide a window of information on extinctorganisms and early life on Earth.
Microbial extremophiles are the most extreme of allorganisms: some can grow at temperatures ashigh as 113ºC [1.4], others at temperatures as lowas -18 ºC [1.3], at negative pH [1.7] or at pH 13.2[1.8], under extreme pressures of up to 130 MPa[1.6], in complete absence of oxygen [2.5], in saltbrines [2.6], or even in conditions of nearly absoluteenergy starvation [2.7]. Particularly with respect tohigh temperatures, their cellular components haveto be resistant to such conditions.
Whilst we have a partial understanding of proteinsand enzymes with regards to extreme conditions,this is far less well developed for DNA and RNA,and we have little knowledge of how cellularmetabolites are protected. Defining the molecularbasis of these properties will make invaluablecontributions to our understanding of the molecularand physiological adaptation to extremeenvironments. It will also allow us to define thelimits of life on Earth, and hence of terrestrial-like lifeon other planetary bodies in our universe.
The vast, largely untapped subsurface of thecontinents and seafloor is one of the greatestrealms of microbial life on Earth, both in terms ofbiomass and volume. Recently an internationalteam of ocean scientists discovered that ethaneand propane are being produced bymicroorganisms in deeply buried sediments [2.8].The findings suggest that microbes below theseafloor carry out processes which are highlyrelevant to both the understanding of global cyclesand the metabolic abilities of microbes.Investigating microbial processes in theseenvironments needs to be explored further.
Extremes of temperature and pressure are met insuch environments and in the subsurface, lifeunder extremely low energy availability is not anexception but appears to be the rule [2.9].Extremely slow growth rates suggested bygeochemical turnover rates with estimated celldivision times of the order of hundreds tothousands of years challenge our fundamentalunderstanding of life and concepts of cellmaintenance.
Finally, because microbial extremophiles cover allthree domains of microbial life, bacteria, archaeaand eukaryotes, comparative adaptation studieswould provide a very useful insight into the natureand evolution of extremophilic organisms.
Recommendation M1: Investigate further the molecular basis of the physiological adaptation of microbes to extremeenvironments, especially with regard to DNA, RNA and protection of cellular metabolites.
Recommendation M2: The deep subsurface (oceanic and continental) exhibits unique features in terms of temperature,pressure and availability of energy. Investigating the subsurface microbial processes further, especiallywith regard to cell division, would deepen the fundamental understanding of life and concepts of cellmaintenance. Moreover, understanding the role of the subsurface in biogeochemical cycles is essentialfor an assessment of element budgets on a global scale.
Recommendation M3: Microbial extremophiles cover all three domains of microbial life, bacteria, archaea and eukaryotes.Comparative adaptation to different environmental stressors and the results of gene arrays andproteomic studies would provide a very useful insight into the nature and evolution of extremophilicorganisms (Recommendation M3).
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2.2.2. Life strategies of plants in extremeenvironments
The most obvious difference between plants andother organisms is their ability to perform oxygenicphotosynthesis using light and carbon dioxide(CO2) in their chloroplasts. Photosystems makeplants more susceptible to radiation andtemperature when compared for example tobacteria. Because they are designed to acceptlight, they are inherently sensitive to radiationquality (especially UV) and quantity. As theirthylakoid membrane system requires a certaindegree of fluidity, they can tolerate only a limitedtemperature range and are restricted to areas thatreceive sunlight. Plants are less dependent onexternal resources and mainly need water, CO2 andlight. Investigating mechanisms of autotrophy havestarted at the molecular level and this transfer ofknowledge is strongly needed for studies aboutplant life in extreme environments.
Plants survive in extreme environments such as inanoxic conditions (e.g. the CO2 springs),hypersaline soils and water, extreme temperatures,and also in extreme conditions caused by bioticstresses such as anthropic pollution and animalgrazing. Many strategies allow autothropic carbon
fixation and plant survival under extremeconditions, based on biochemical adaptation,molecular adaptation and acclimationmechanisms.
A second common characteristic for plants is that,when grown, their sessile life style does not allowactive avoidance mechanisms as in motile (and/ormobile) organisms. Consequently, plants possessmore protective mechanisms than mobileorganisms. This applies in particular to juvenile orlong-term survival stages; e.g. spores or seeds,which consequently deserve special attention. Thepeculiar strategies of adaptation developed bysessile plants need special studies, ranging fromthe mechanism level to the ecosystem level (e.g. itsconsequences for the plant-based life-chainpyramid). Exceptions to sessile life are found in theplant kingdom (e.g. clonal plants) and should bealso investigated.
Finally, plants are extremely important elements inthe Earth life-chain, because of their position asprimary drivers, in many situations, of the trophicchain. They can also play a role as naturalengineers producing environmental conditionssuitable to life and as they can be extremelysensitive to environmental changes, they are usefulas bio-indicators.
Recommendation P1: Put the emphasis on boosting progress in molecular and biochemical studies and on unravelling themechanistic basis of plant life under extreme conditions.
Recommendation P2: Study the strategies of adaptation (from the molecular to the ecosystem level) which allow plant life inextreme environments, as well as plants’ strategies of reproduction and dissemination in their long-termefforts to avoid environments becoming hostile (e.g. because of global change factors).
Recommendation P3: When considering studies on plant life in extreme environments, a particular attention should be givenon fragile ecosystems, including ecosystems that start to be colonised by plants, or that are becomingunfavourable to plant life, especially as a consequence of global change and human-driven land usechange. Strategies of survival at this boundary that may be or may become of utmost importance forplant survival, for plant loss of diversity, and for the entire food-chain based on plant life.
28 Investigating Life in Extreme Environments
Espeletia timotensis on a semi-desert plateau at an altitude of 4,200m
Phytoremediation of a former gas-station in Rønnede (Denmark)using different species of Salix.
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2.2.3. Life strategies of animals in extremeenvironments
The wide scope of this subject shows thecomplexity of the scientific community investigatinglife strategies of animals in extreme environments.This results from the diversity of animals andenvironments studied and, to a lesser extent, fromthe questions raised (ecology, evolution,physiology). Each subgroup is well organised andexhibits a high degree of maturity, but it seemsclear that so far there are no links among thesesubgroups. There are several questions which cutacross these subgroups (e.g. communityresponses, ecosystem functioning, hypoxia andthermal behaviour) and which require morecoordination. The scientists concerned with thequestions of life strategies of animals in extremeenvironments are numerous and highly diverse;e.g. groups working on polar animals orhydrothermal vents fauna.
Apart from the two environments generally studied(polar and deep-sea environments), more specificones have also been studied, such as those thatare under increased anthropogenic pressures; e.g.on coastal zones or polluted areas.
Within the research community a strong interestexists in the question of genomics and stress
response. Another major point of interest is theneed to develop comparative studies to identify thecommon characteristics and to determine thespecificities of cells, animals, communities andecosystem responses to environmental stress ofvarious types including climate change.
There is also a consensus about the importance ofcharacterising environments and their interactionswith animals from the microscale (cellular) to theglobal scale (Earth), this is of particular relevancewhen considering animals evolving in extremeenvironments. Such approaches may help, forexample, in understanding the establishment ofsymbiotic interactions as well as in theirdisappearance. Questions about variability ofbiodiversity are stressed by different subgroups ofscientists. Approaches involving palaeontologymay help the understanding of the evolution of lifein extreme environments. In vivo approachesmimicking the natural environment would help inthe understanding and prediction of the toleranceof animals to environmental changes.
Recommendation A1: Comparative studies are needed to determine the specificities of animals, communities and ecosystemresponses to different types of environmental stress.
Recommendation A2: Favour characterisation studies on extreme environments and their interactions with animals; this wouldhelp in understanding adaptation processes.
Recommendation A3: The analysis of the biodiversity dynamics of animal communities and ecosystems found in extremeenvironments, including palaeontological approaches, has to be promoted to understand the evolutionof life in extreme environments.
Recommendation A4: In vivo approaches simulating the natural environment are needed to help in the knowledge andprediction of the tolerance of animals to environmental changes.
29Investigating Life in Extreme Environments
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Onymacris unguicularis, fog-basking beetle of the Namib desert
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2.2.4. Human adaptation in extremeenvironments
Human adaptations to extreme environments are ofdifferent types: they can be biomedical,physiological, psychological but also cultural,societal and technological. With this diversity ofadaptation aspects, it seems important thatscientists should collaborate in studying humanadaptation to extreme environments under thelabel Human Sciences.
Humans adapt to extreme environments but thereare also adaptive effects that they impose upon agiven environment. Environmental monitoring andcontrol have an important role both in artificial andconfined environments (i.e. parts of thetechnosphere, ranging from spaceships to cities)as well as in natural environments.
In the context of scientific research conducted inextreme environments, studying human adaptationhas a twofold aspect. First, it allows a betterunderstanding of the mechanisms of humanresponse to physiological and psychologicalstressors, paving the way for new therapies andapplications in daily life. Second, with thedevelopment of countermeasures, it acts as anenabler aiming at optimising the well-being andsecurity of the personnel involved in extremeenvironment research. This is of particularrelevance when considering polar station crew, andastronauts performing research in microgravity. Inthese contexts, one can identify three major factorsthat would impact on human activity:
Isolation and confinement: crew in spacevessels or polar stations are disconnected fromtheir usual social environment and thehazardousness of the surroundings is a majorsource of stress for them. This psychologicalpressure impacts on the coherence of the groupand its ability to perform research.
Recommendation H1: Bring together humanities, social sciences, psychology and medical sciences under the label of HumanSciences in studies on human adaptation to extreme environments.
Recommendation H2: Issues of environmental monitoring and control have an important role both in artificial and confinedenvironments as well as in natural environments. These aspects should be further investigated whenconsidering humans, not only because the environment influences them, but also because there is anadaptive effect that humans impose upon the environment.
Recommendation H3: When considering human species, favour cross-disciplinary dialogue and interactions while dealing withdifferent levels of systems hierarchy at the same time.
Recommendation H4: Address ethical issues systematically and carefully when considering human adaptation to extremeenvironments.
30 Investigating Life in Extreme Environments
Level of activity: the low level of physical activityarising from confinement in space and polarstations may lead to hypokinesia (diminishedmovement of body musculature) with its associatedimpact on the human body. Understanding theimpacts and associated mechanisms may help usto understand the role of physical inactivity on theaetiology of obesity and diabetes, boneosteoporosis and also muscle atrophy observed inthe elderly.
Reduced level of gravity (only for spacemissions): without gravity the complicated bonearchitecture or muscle organisation would not benecessary. The intricacies of the cardiovascularsystem are mainly organised to counteractcontinuous fluid transfer induced by gravity. Thesame evidence emerged from the organisation ofthe central nervous system given all themechanisms developed to counteract gravity (e.g.the vestibular system). Studying human adaptationto microgravity therefore provides potentialities in awide range of areas in fundamental sciences andalso in pathology (i.e. ageing, cardiovasculardysfunctions, osteoporosis and muscle atrophy).
Finally, it can be observed that extremeenvironments are not necessarily disadvantageous,but may sometimes be favourable: there might bepositive and negative impacts at the same time.This is one example where ethical issues enter thepicture.
ESA astronaut Ulf Merbold soon after leaving the Soyuz-TM 19descent capsule
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3.1. Streamlining the issue
When considering scientific activities in extremeenvironments, technology sets itself as a clear andvital cross-cutting resource of major relevance.However, technological barriers are the mainlimiting factors for the development of newknowledge on life processes in extremeenvironments. Furthermore, technologicaldevelopments are crucial elements that allowhumans to cope with extreme environments.Without these, human beings would not be able toexplore and evolve in some harsh conditions.Evident examples are space suits for planetaryexploration and polar stations for the exploration ofthe polar regions.
Thus, it is important to develop a comprehensivecharacterisation of the barriers, technologicalneeds and capabilities for investigation of life inextreme environments. It is also important toconsider a common technology development planto which different scientific communities (e.g. thosestudying microbes, plants, animals, humans) cancontribute with their specific technologyrequirements, strengths and capabilities. Table 2proposes a common functional grouping undervarious technology thrusts of relevance whenconsidering investigation of life in extremeenvironments.
Furthermore, through the development of a road-mapping exercise on technology development,added benefit can be given to each of the diversebut connected research communities involved bybringing to them the existing tools and capabilitiesin use or under development by their neighbouringcommunities.
Harsh environments impose major constraints onthe capabilities and sustainability of technologiesthat are required to undertake high qualityresearch. Examination of the extremeness withrespect to a chosen frame of reference requiresmeasurement of the environmental parameters(physical, chemical and biological). Conductingscientific experiments accurately and robustlyunder extreme conditions requires technologicaltools with extreme capabilities needed duringdifferent phases of the scientific investigations suchas discovery, access, measurement, analysis,acquisition and transmission of results or samples,life support systems, computer software etc. It ismainly through the enhancement andadvancement of the technological capabilities thatscientific experiments under extreme conditionscan be further envisaged and enabled. In thiscontext, selecting analogous sites to testtechnologies would be useful.
Recommendation T1: Develop interdisciplinary initiatives to define environment-specific technological and technicalrequirements, their availability and a road-map for their development.
3. Technological issues
31Investigating Life in Extreme Environments
Victor Remotely Operated Vehicle
European ExoMars rover — artist's view
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Technology thrust Specialised systems, Important issues subsystems and tools at the design stage
Exploration Where to go?When to go?How to get there?How to navigate?How to communicate?What to look for?What to bring back?Environmental impactsEnergy and life support sustainability
Identification Sensing Measurement of predefined physical,biological and environmental parameters. Recognition of targets
Recognition and samples of interest
Monitoring Sensing Continuous and/or intermittent measurement of various parameters of interest on microbial, plants, animalsand human subjects. Size, energy supply, communications
Acquisition Containment Isolation and insulations. Environmental control. Specialised sampling, capturing,
Samples and/or data retrieval and containment systems
Retrieval
Analysis In situ Autonomous or semi-autonomousmobile or stationary laboratories. Interfaces, communications,
Lab processing environmental control, energy, mobility, life support systems
Data storage and communication
Sample/species Samples/species monitoring Autonomous, remotely operated orreturn Acquisition manned vehicles.
Contamination control, isolation Life support systems, communications and containmentLife support systemsRetrievalTransportation
Data acquisition, Data acquisition and storage Power, data rate, volume, bandwidthstorage and Data miningtransmission Data reduction and compression
Communication
32 Investigating Life in Extreme Environments
3. Technological issues
Table 2: Functional grouping of technologies under various technology thrusts
Planetary and interplanetary access
Terrestrial remote access
Marine remote and/or deep access
Polar remote, deep and subglacial access
Local deployment, command and control
Specialised deep drilling
3.2. Crossing the borders
While some specific research and technologyprogrammes have allowed significant progress, thefragmentation of scientific activities and strengthsprevents spin-offs across disciplines. It is importantthat technology-related information exchange isfacilitated between communities involved inresearch activities dealing with extremeenvironments. This is of particular relevance whenconsidering, for instance, the effort put into thedevelopment of space technologies. The highquality, robustness and capabilities of light-weightand small-size sensors and instruments developedfor space missions would surely have the potentialto benefit terrestrial research in extremeenvironments.
Interactions between communities already exist;e.g. the Antarctica (subglacial) Lake Ellsworthexploration team (from the UK) involves scientistsfrom the Beagle 2 Mars lander mission for thedevelopment of a robot exploration probe [3.1].This use across domains would in turn benefitfuture planetary exploration systems throughfurther advancements in the capabilities andoperational knowledge and experiences gained. Asanother example, the European Space Agency(ESA) developed, together with the Italian nationalAntarctic research programme (PNRA) and Frenchpolar institute (IPEV), the water recycling system ofthe French-Italian Concordia Antarctic station [3.2].These types of interactions and spin-offs should beencouraged.
Regarding infrastructures, the ones that are used toreach and undertake scientific activities in extremeenvironments (e.g. icebreakers, polar stations orunderwater vehicles), have very peculiarcharacteristics; they are typically rare (sometimesunique) complex systems, expensive to developand operate. In order to maximise the scientificreturn on such investments at the European level, itis important to develop the mechanisms and toolsnecessary to facilitate access to and sharing oftechnical platforms and infrastructures (e.g. barteragreements, common programmes andinternational consortia).
Recommendation T2: Support the development of initiatives aiming to improve information exchange on available technologiesand new developments throughout the scientific community involved in extreme environments.
Recommendation T3: Develop and support mechanisms to facilitate access to and sharing of existing and future technicalplatforms and infrastructures to be used when accessing extreme environments.
33Investigating Life in Extreme Environments
The AWI-operated German icebreaker Polarstern docked on the sea ice for unloading.
Melting Probe technology of potential use for penetratinglayers of ice.
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3.3. Collecting information –priorities
As already stated, access to extreme environmentsis time and resource limited. Laboratory simulationtechniques and capabilities permit lowering thecost of investigations while allowing real-time long-term observation. An example of such a facility isthe LabHorta in the Department of Oceanographyand Fisheries of the University of the Azores(Portugal). LabHorta simulates the physical andchemical conditions found around deep-seahydrothermal vents. This facility was developedand built to support and expand the capacity ofresearch cruises together with experimental studiesof the biology, physiology and behaviour of deep-sea hydrothermal vent fauna and taxa from otherdeep-sea environments [3.3]. Development oflaboratory simulation techniques and facilitieswould stimulate and enhance the volume ofresearch focused on extreme environments; theyshould be developed in Europe for relevantenvironments and made available to the Europeanscientific community involved.
Besides the simulation issue, development of insitu measuring and monitoring capacity in harshconditions is required. For example, this includesdeep-sea observatories, polar observatories andRemotely Operated Vehicles (ROVs). Developingthis capacity (quantitatively and qualitatively) meansobtaining the highest quality information on thephysical and chemical conditions as well as
imaging. This also requires continuousdevelopment of, and access to, sensor, robotic andtelemetry technology.
In this context, at the convergence of space-basedinfrastructure with global capacity, (GPS,telecommunication satellites), and newdevelopments in microelectronics, on sensors,data storage and data transmitter technologies,bio-logging is a new field of activity that allows alarge amount of information based on continuousmeasurements to be gathered remotely. Thisinformation can concern animal physiology (e.g.temperature or heart rate), behaviour (e.g. itinerary,speed, depth) but also the environment where theanimal is located (e.g. ambient temperature, salinityor light). Any decrease in size, weight and cost ofelectronic components together with an increase intheir capability enable bio-logging techniques to bevery promising in scientific terms.
In addition to the development and use of in situcapabilities, there is a need to develop the use ofremote measurement capability such as satelliteremote sensing (e.g. for plants and algaemonitoring on Earth-based environments ordetections of gas resulting from biological activityon planetary bodies).
Recommendation T4: Consider further the enhancement of the European simulation capability in the field of extremeenvironments. Relevant simulation facilities (e.g. microcosms) should be developed.
Recommendation T5: Develop in situ sampling, measurement and monitoring technologies (e.g. light-weight, low-power, androbust sensors) and capacity (e.g. in situ observatories and bio-logging). Categorisation and inventoryof existing systems, technologies and techniques are also recommended.
34 Investigating Life in Extreme Environments
The LabHorta facility designed for the simulation of deep seahydrothermal vents’ conditions
The experimental Life on Ice: Robotic Antarctic Explorer(LORAX) autonomous rover
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Benefiting from extremophilic organisms
Organisms that live in extreme physico-chemicalconditions, in high concentrations of deleterioussubstances or heavy metals, represent one of themost important frontiers for the development ofnew biotechnological applications. Actually, thebiotechnological applications of extremophiles andtheir components (e.g. extremozymes) have beenthe main driving force for the research in this area[4.1].
The most direct application of extremophiles inbiotechnological processes involves the organismsthemselves. Among the most established we canfind biomining, in which microbial consortia thatoperate at acidic pH are used to extract metalsfrom minerals [4.2]. Most applications involvingextremophiles are based on their biomolecules(primarily enzymes, but also other componentssuch as proteins, lipids or small molecules). Thebest-known example of a successful application ofan extremophile product is Taq DNA polymerasefrom the bacteria Thermus aquaticus [1.6] whichfacilitated a revolution in molecular biologymethodology, but also other commercialisedproducts (e.g. ligases, proteases, phosphatases,cellulases or bacteriorhodopsin) have resulted frominvestigation of extremophiles isolated fromdifferent environments.
In order to develop biotechnological processes, it isimportant to isolate the organism. In addition, theuse of new methodologies, such as comparativegenomics is helping to sort out the genetic andmolecular bases of adaptation to extremeconditions, facilitating the design of improvedproducts by the introduction of appropriatemodifications by protein engineering. Thisapproach has been used not only for improving thethermostability of enzymes but to design cryo-enzymes for the food industry to operate at lowtemperatures.
Most microbial communities are complex andcurrently only a few components can be cultured.Sequence-based approaches to study themetabolism of microbial communities are beingused to retrieve genomic information from thecommunity of potential use in biotechnology(metagenomics). Strategies based on thegeneration of environmental genomic libraries havebeen developed to directly identify enzymes fromthe environment with the required specificity or theappropriate operational conditions.
Plants adapted to extreme environments alsoprovide potential economic and societalapplications. Extremophilic plants can surviveunder conditions toxic or harmful to crop plants.Therefore there is the potential to transfer, e.g. bymolecular cloning, some of these abilities fromextremophiles to crop plants with the aim ofproducing frost, salt, heavy metal or droughttolerance or enhanced UV stability.
Plants can also be useful to remediate pollutedareas where life is made difficult or impossible.Phytoremediation is an innovative technology thatuses the natural properties of plants in engineeredsystems to remediate hazardous waste sites.Within the phytoremediation technologies,phytoextraction (uptake and concentration ofsubstances from the environment into plantbiomass) and phytotransformation (chemicalmodification of environmental substances as adirect result of plant metabolism) are of applicativeinterest. Phytoremediation has been effectivelyused for the decontamination of soils and waterspolluted by high concentrations of hazardousorganic (e.g. pesticides) or inorganic (e.g. arsenicand mercury) substances. It is also a promisingtechnology for the remediation of atmosphericpollutants (hydrocarbons, ozone) [4.3].
Biomedical applications of adaptive mechanisms ofanimals should also be thoroughly investigated.Such potentialities are real as illustrated by thesubantarctic King penguin that has developed theability to preserve fish in its stomach for threeweeks at a temperature of 38°C [4.4]. With furtherdevelopments, the antimicrobial and antifungalpeptide involved in this conservation process mightbe used, for example, to fight some nosocomialinfections.
4. Economic and societal applications
35Investigating Life in Extreme Environments
Bacteria Thermus aquaticus
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It is obvious that the combination of newmethodologies to study extreme environments willprovide important biotechnological breakthroughs.
Thus it is extremely important to support thedevelopment of this knowledge in Europe in order tobe at the forefront of the new biotechnologicaladvancements.
Benefiting from research activities
Conducting scientific activities in extremeenvironments requires high-quality and highlyrobust facilities and technological capabilities. Inthis context, efforts in sensor development andoptimisation, miniaturisation of scientific devices,automation of processes, robotics, softwaredevelopment or material engineering areundertaken. This allows industries and researchinstitutions involved in such activities to gainknowledge, competencies and capabilities relevantto a wide range of industrial and economic sectors.
36 Investigating Life in Extreme Environments
4. Economic and societal applications
5. Policy aspects – the European arena
5.1. European scientific landscape
The scope of research activities in extremeenvironments is very broad and the state ofdevelopment of this area in Europe is very diverse,depending on: i) which extreme environment isstudied and ii) which type of organism isconsidered.
Maturity
Historically in Europe, the consideration ofextremophiles within funding programmes of theEuropean Union Fourth (FP4) and Fifth (FP5)Framework Programmes has greatly contributed toestablishing and structuring the European scientificcommunity dealing with these topics. Currently, atthe international level, Europe leads in areas suchas polar research, biodiversity, and global change,although without a specific focus on extremeconditions.
Polar and deep-sea research fields, includingassociated technological developments, appear tobe areas of excellence in Europe. In considering thetype of organisms, the study of microbes, plantsand animals are reasonably mature areas ofresearch. However, gaps exist within each of thedifferent fields (regarding species andenvironments) and there is a significant potential toinvestigate common questions (especiallyphysiological and ecological ones) across variousextreme environments. There is then a substantialpotential to expand our functional knowledge of asyet relatively unexplored fields, exploiting recentadvances in molecular and structural biology anddeveloping new technologies and modelorganisms for extreme environments. These shouldinclude ecosystem approaches and in situmeasurements.
A concerted European initiative could helpconsiderably to achieve a visibility similar to currentor past large-scale initiatives in the United States(e.g. the National Science Foundation/NASA Life inExtreme Environments Programme - LExEn) and torealise the field’s full growth potential. Such aninitiative should not only focus on organisms butalso on the environments themselves, providingknowledge on interactions within ecosystems andan original integrated approach full of potentialities.
Existing initiatives
There is a general challenge across the topic areasof very different paradigms, methods and
philosophies among the scientific disciplinesinvolved. However, a patchwork of interdisciplinaryprojects and collaborative networks of interestalready exists and is indicative of the dynamicnature of the research fields covered. Theseinitiatives are spread across the European scientificlandscape but are either dedicated to specificenvironments (e.g. European participation in theInternational Polar Year (2007-08), the EuropeanPolar Consortium, HERMES, ESO-NET and MarsExpress), to specific geological contexts (e.g.European participation in IODP, InterRidge andEuroMARC), or focused on a thematic topic suchas biodiversity (e.g. EuroCoML and MarBef).Connections with generalist networks are alsoimportant considering their advances inmethodologies relevant to the field of investigatinglife in extreme environments (e.g. FP6 Network ofExcellence Marine Genomics Europe).
International collaboration
A strong benefit can be expected from internationalcollaboration especially with Canada, China, India,Japan, Russia and the United States. This is ofparticular relevance in areas of global significance,e.g. desertification, pollution, and the effects ofglobal change in polar regions and other fragileareas. International collaboration is indeed crucialboth for large-scale initiatives such as theestablishment of seafloor observatories, the use oflarge-scale (expensive) infrastructures, the planningand execution of space missions, and coordinationof expeditions. Also, developing countries, wheremany of the as yet unexplored extremeenvironments exist, should be partners incooperative ventures. Special attention should bepaid to the education of local scientists andstudents on these topics; e.g. through the use offellowships or organisation of joint investigation.
37Investigating Life in Extreme Environments
French-Italian Concordia Antarctic station: a platform for inter-national cooperation
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5.2. The need for furtherinterdisciplinarity
It is important to emphasise the unique opportunitythat the interdisciplinary approach currently offersto investigations of life in extreme environmentsand that such an approach should be furtherdeveloped. In order to deal with broader aspects oflife in extreme environments in a scientific way andto secure access to sufficient future funding,appropriate mechanisms of peer review, adapted ifnecessary, and significantly increased opennesswould be needed.
Existing compartmentalisation between disciplinesand their associated technological developments isseen as a major barrier to greater efficiency inscientific activities. Even if this statement applies toa whole range of scientific activities, it is ofparticular relevance when considering extremeenvironments because science and technologyactivities dealing with or relevant to extremeenvironments are widespread across the scientificlandscape. Cross-over exists between disciplines,therefore others should be further studied oridentified; e.g., using microbial and plant lifeprocesses in some particular conditions could lead
to the development of better life-support systemsin Antarctic or space stations. Interdisciplinaryaspects should be considered as much as possiblein any relevant scientific activity involving extremeenvironments, and especially when considering thestudy of ecosystems (see also Section 2.1.2).Larger and/or more frequent interdisciplinaryinitiatives should be developed to facilitate greaterunderstanding of specific problems, ethical issuesand even to develop a common scientific language.It is obvious that sharing common questions andtechniques, developing the study of interfaces,undertaking comparative projects and integratingnew tools and knowledge would definitely catalysethe development of scientific excellence for eacharea involved. Given Europe's academic history,tradition and capability, there is clearly potential toachieve truly remarkable results in this area.
Alongside the knowledge-based approach, on amore practical side, interdisciplinary andmultidisciplinary approaches are also necessary inremote research areas requiring expensiveexpeditions or space technology. A multitude ofmethods needs to be applied to maximise thescientific outcome samples that are expensive tocollect. This also applies to the field of genomics ofextreme model organisms where the generation ofthe data is fast but still expensive. Furthermore, theanalysis of results would benefit tremendously fromthe synergistic effect of multidisciplinary expertise,ranging across ecophysiology to DNAbioinformatics. It is evident that, whenever suchtransdisciplinary research approaches aresuccessful, the benefits are remarkable.
Palmatogecko rangel in the Nabib desert
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The archea Methanococcus jannaschiia evolves in the area ofhydrothermal vents
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Recommendation 8: Because of the transversal nature of science and technology activities in extreme environments,interdisciplinarity and multidisciplinarity between scientific domains and between the technological andscientific spheres is identified as a major source of potential increase in scientific achievement. Whenrelevant, these aspects should be extensively promoted and supported as much as possible whenconsidering life processes in extreme environments.
5. Policy aspects – the European arena
38 Investigating Life in Extreme Environments
5.3. Networking and coordination
While high-level expertise and know-how havebeen developed at the European level, thefragmentation of this knowledge and the lack ofcoordination across initiatives are major constraintsto the further development of a pan-Europeancapacity.
Thus, there is a clear need to develop an efficientcapability in networking and information exchangefor the European scientific community involved inresearch on extreme environments. Thisnetworking should concern not only individualscientists but also existing and future collaborativeinitiatives. As stated above, it is important that sucha capability is multidisciplinary and designedaccording to the requirements and expectations ofthe scientific community. This could include anInternet portal gathering and disseminatinginformation on relevant programmes and projects,publications, funding opportunities and the use ofinfrastructures. To facilitate the networking of thecommunity, it should also include a dynamicdatabase of experts and specific scientific andtechnical expertises. An associated newslettercould also be considered in order to inform thescientific and technological communities and topromote interdisciplinary approaches in the field ofextreme environments.
In addition, large-scale multidisciplinary events(conferences, workshops, fora) such as the ESF’sInvestigating Life in Extreme Environments (ILEE)workshop held in November 2005, should beorganised regularly in order to build and maintainmomentum at the European level and to shapeinitiatives and possibly develop relevant projectswith a bottom-up approach. In this context, thecompetitive ESF Research Conferences schemecould be a very useful tool for the scientificcommunity.
There is also a need to identify potentialcoordination (and possibly clustering) mechanismsto be initiated among existing networks andprogrammes as well as to investigate mechanismsfor sharing costs associated with infrastructures
(e.g. ROVs and ships) or national large-scaleresources.
The use of instruments such as the EuropeanCommission’s Framework Programmes’Coordination Actions (CA) would be fully relevant todevelop networking and coordination within theEuropean research community involved in extremeenvironments.
These initiatives and undertakings need to beconsidered in detail and an overarchinginterdisciplinary group of experts should beestablished. This focal point, possibly set up underthe auspices of ESF, would consider and steer theissues of coordination, networking anddissemination of activities. Furthermore, it couldidentify priorities and investigate new opportunitiesfor Europe’s scientific community. Theoretical andmethodological approaches should also bediscussed and defined to coordinate the variousviews of the community involved. It is important tonote that acquiring a common language among thevarious fields considered is a prerequisite for thispurpose.
Finally, reaching a critical mass, getting to knoweach other and identifying common researchpriorities through networking and cooperationwould be the first steps towards the developmentof a European Research Area in the field of extremeenvironments and potential European-wideresearch projects and programmes.
The ESF Investigating Life in Extreme Environments workshop,November 2005
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Recommendation 9: Create an overarching interdisciplinary group of experts to define the necessary actions to build a criticalEuropean mass in the field of Investigating Life in Extreme Environments.
Recommendation 10: Efforts should be made to support and improve the information exchange, coordination and networkingof the European community involved in scientific activities in extreme environments. This could beachieved through a Coordination Action in the European Union Seventh Framework Programme (FP7)and include the development of a Web portal and associated developments (database of experts,dissemination of information). In this context, large-scale multidisciplinary events (e.g. conferences, foraand workshops) should be organised regularly in order to build and maintain the momentum at theEuropean level. This should be done in the context of the numerous specialised events that are alreadyorganised).
39Investigating Life in Extreme Environments
5.4. Programme issues
Overall, and especially because of their specificitiesand positioning at the frontier of science, researchactivities dealing with life processes in extremeenvironments should adopt a bottom-up approachto be open and flexible in order to avoid any rigidityand possible obstruction of creativity, to take fulladvantage of current trends and possibilities, andto tap into the entire spectrum of Europeanpotential. In this context, the scientific communityneeds to continually generate new ideas andconcepts.
A general difficulty from an operational point of viewinvolves the long planning horizons of fieldexpeditions to sample and monitor extremeenvironments that are often remote and hard toreach (e.g. oceanic sites, polar regions or planetarybodies). Because these expeditions (or missions)have a long planning interval relative to fundingcycles, it is often difficult for the scientificcommunity to take part in them.
The current level of coordination of Europeanfunding for investigating life in extremeenvironments is certainly viewed as a constraintthat needs to be overcome. Carrying out scientificactivities in such environments is often veryexpensive, because of the cost of facilities such aspolar stations, research vessels, RemotelyOperated Vehicles (ROVs) or interplanetary probes
and landers. This represents a major constraint onscientific progress in these fields.
National research funding may be very difficult toaccess (deep-sea and high altitude research beingparticular examples). A strong European networkcould create stimuli for national research councilsand funding organisations to commit themselvesand include extreme environments as such in theirnational science strategy. A consideration at theEuropean level of the funding needed for suchinitiatives, the tools and schemes to beimplemented as well as the identification ofpotential synergies and scale effects would fallunder the remit of the overarching interdisciplinarygroup of experts mentioned above. Furthermore, itwould seem pertinent that extreme environmentsshould be clearly identified in scientific calls andprogrammes. It is preferable to fund leadingscientists in the field defining research goals ratherthan urging scientists to mould their research intoexisting funding schemes.
Finally, a new field needs new peer groups in orderto avoid tunnel effects in scientific evaluation.Review and evaluation processes need to include amultidisciplinary aspect to be more open toexternal influences, and to embrace a broader view(systems biology being an example). Decisionguidelines need to be appropriately flexible.
Recommendation 11: Extreme environments should be integrated into national science strategy and clearly identified innational and European scientific calls and programmes.
Recommendation 12: When considering scientific activities involving extreme environments, funding agencies need to ensurethat their assessment procedures can adequately review proposals involving a multidisciplinaryapproach as opposed to the more typical, strongly focused proposals.
40 Investigating Life in Extreme Environments
Merged ice pancakes in Antarctica
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41Investigating Life in Extreme Environments
5.5. Public outreach and education
Securing public awareness that science is excitingis crucial, especially as the general public isinterested in seeing the outcome of investments inscience. It is important to inform the public onissues related to extreme environments,highlighting the exotic nature of these environmentsand their biota. Extreme environments such aspolar regions, deserts or outer space are excitingand easily communicated to the public becauseeven the non-scientific audience intuitively seesthat these environments impose a challenge for life.Moreover, observing and understanding life inextreme environments brings the public to questionitself on very important issues such as the origin oflife or potential life on other planetary bodies. Otherextreme environments, such as intertidal sites, salt-water marshes or heavy metal polluted sites, mayinitially appear less attractive and exotic but are inpractice more accessible to the general public andprobably of greater relevance to daily life.
The popularity of television channels such as theDiscovery Channel and National GeographicChannel in Europe and the USA suggests that thepublic is interested to learn how certain organismssurvive and even thrive under extreme conditions.Raising public interest can be achieved by reportsin the media detailing new findings such as themysterious animals of the deep-sea, but also inreporting on expeditions to extreme environmentson Earth or missions to outer space.Demonstrating applications of the data, e.g. howresults from studies in extreme environments caninfluence crop production, would also be ofimportance.
It is also fundamental to secure the transmission ofscientific knowledge to students and youngscientists. This could be achieved throughsupporting PhDs and postdoctoral positions aswell as involving students and young scientists inscientific events. The latter could be achieved, forexample, through the organisation ofinterdisciplinary summer schools focused on lifeprocesses in extreme environments.
The Norwegian University Centre in Svalbard (UNIS) located at78° North
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[1.1] Duxbury N.S. et al.Time machine: Ancient life on Earth and in the cosmos EOS, Trans., American Geophysical Union, v. 87, no. 39: 401-406, (2006)
[1.2] Priscu, JC. et alGeomicrobiology of Subglacial Ice Above Lake Vostok, Antarctica Science 286: 2141-2144 (1999).
[1.3] Clarke, A. Evolution on Planet Earth: The Impact of the Physical Environment Editors: Rothschild, L. &Lister, A. - Academic, London (2003).
[1.4] Blochl, E. et al. Pyrolobus fumarii, gen. and sp. nov., represents a novel group of archaea, extending the uppertemperature limit for life to 113°C.Extremophiles 1, 14–21 (1997).
[1.5] Horneck G. Responses of Bacillus subtilis spores to the space environment: results from experiments in space. Origins of Life and Evolution of the Biosphere. 23:37–52 (1993).
[1.6] Rothschild, L. J., and Mancinelli, R. L.Life in extreme environments. Nature 409:1092-101 (2001).
[1.7] Schleper, C et al. Picrophilus gen. nov. , fam. nov.: a nobel aerobic, heterotrophic, thermoacidophilic genus and familycomprising archaea capable of growth around pH 0. J. Bacteriol., 177, 7050-7059 (1995).
[1.8] E Roadcap, G. et al.Extremely Alkaline (pH > 12) Ground Water Hosts Diverse Microbial Community Ground Water 44: 511-517 (2006).
[1.9] Madigan MM, et al.In Brock Biology of Microorganisms 11th Edition Pearson Education, Inc., Upper Saddle River, NJ (2006)
[1.10] Levin-Zaidman, S., et al.Ringlike Structure of the Deinococcus radiodurans Genome: A Key to Radioresistance?Science 299: 254-256 (2003).
[1.11] Kato, C. et al.Extremely Barophilic Bacteria Isolated from the Mariana Trench, Challenger Deep, at a Depth of11,000 MetersApplied and Environmental Microbiology 64(4): 1510–1513 (1998).
[1.12] Blankenship LE, et al.Vertical zonation patterns of scavenging amphipods from the Hadal zone of the Tonga and KermadecTrenches Deep-Sea Research Part I-Oceanographic Research Papers 53 (1): 48-61 (2006)
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42 Investigating Life in Extreme Environments
[1.13] Extreme Life BriefingAstrobiology Magazine http://www.astrobio.net/news/article2107.html
[2.1] Shillito, B et al.First Access to Live AlvinellasHigh Pressure Research, 24, 169- 172 (2004).
[2.2] Allen, E.F., Banfield, J.F.Community genomics in microbial ecology and evolution. Nature Reviews Microbiology, 3: 489-498 (2005).
[2.3] Ferrer, M. et al. Microbial enzymes mined from the Urania deep-sea hypersaline anoxic basin. Chem Biol. 12(8):895-904 (2005)
[2.4] Franzetti, B. et al.Tetrahedral aminopeptidase: a novel large protease complex from archaea. EMBO J. 21, 2132-2138 (2002)
[2.5] Rainey, F.A. and Oren, A. (eds.) Extremophiles, Methods in MicrobiologyVol. 35, Elsevier, London (2006).
[2.6] Michaelis, W et al.Microbial reefs in the Black Sea fuelled by anaerobic oxidation of methane. Science, 297, 1013-1015(2002).
[2.7] Sorensen, K.B. et al.Archaeal phylotypes in a metal-rich and low-activity deep subsurface sediments of the Peru Basin,ODP Leg 201, Site 1231. Geobiology, 2, 151-161 (2004)
[2.8] Hinrichs, K.-U. et al.Biological formation of ethane and propane in the deep marine subsurface Proceedings of the National Academy of Sciences 103 (Oct. 3):14684-14689 (2006).
[2.9] Physiology and Biochemistry of ExtremophilesEditors: C. Gerday and N Glansdorff - ASM Press, Washington, DC (2007).
[3.1] UK races to find life under ice Times Higher Education Supplement, 17 September 2004http://www.geos.ed.ac.uk/research/ellsworth/THES.htm
[3.2] Water recycling systems for the Antarctic Concordia stationESA Publications Division. - (ESA BR; 217) 2004.www.esa.int/esapub/br/br217/br217.pdf
43Investigating Life in Extreme Environments
[3.3] Compendium of Specialist Marine Research Infrastructures - PORTUGAL For the MarinERA project www.marinera.net/about/partners/documents/MarinERA_Portugal_Rev1.pdf
[4.1] Antranikian, G. et al.Extreme environments as a source for microorganisms and novel biocatalists. Advances inBiochemical Engineering/Biotechnology., 96: 219-262 (2005).
[4.2] Rawlings, D.E. Characteristics and adaptability of iron- and sulfur- oxidizing microorganisms used for the recovery ofmetals from minerals and their concentrates. Microbial Cell Factories, 4: 13 (2005).
[4.3] Krämer, U. Phytoremediation: novel approaches to cleaning up polluted soils. Current Opinion in Biotechnology, v.2, p.133-141 (2005).
[4.4] Thouzeau C, et al. Spheniscins:avian β-defensins in preserved stomach contents of the king penguin, Aptenodytespatagonicus.J.Biol.Chem., 278: 51053-51058 (2003).
References
44 Investigating Life in Extreme Environments
Appendices
45Investigating Life in Extreme Environments
Appendix I: Contributors to the report
Name Organisation Town Country
Dr Ricardo Amils Universidad Autonoma de Madrid - Centro de Biología Molecular del CSIC Madrid Spain
Dr Arnoldus Blix University of Tromso Tromso Norway
Dr Michael Danson University of Bath - Centre for Extremophile Research Bath United
Kingdom
Dr Christine Ebel CEA-CNRS-UJF – Institut de Biologie Structurale Grenoble France
Dr Cynan Ellis-Evans British Antarctic Survey Cambridge United Kingdom
Dr Françoise Gaill University Pierre et Marie Curie - Laboratoire ‘Systématique, adaptation, évolution’ Paris France
Dr Helmut Hinghofer- Medical University Graz - InstituteSzalkay of Physiology, Center of Physiological
Medicine; and IAP Institute Graz Graz Austria
Dr Kai-Uwe Hinrichs Universität Bremen - Fachbereich Geowissenschaften Bremen Germany
Dr Francesco Loreto CNR – Istituto di Biologia Agroambientale e Forestale (IBAF) Rome Italy
Dr Daniel Prieur Université de Bretagne Occidentale - Laboratoire de Microbiologie des Environnements Extrêmes Plouzane France
Dr Farzam Ranjbaran European Science Foundation Strasbourg France
Dr Klaus Valentin Alfred-Wegener-Institut für Polar und Meeresforschung Bremerhaven Germany
Dr Elisabeth Vestergaard Danish Institute for Studies in Research and Research Policy Aarhus Denmark
Mr. Nicolas Walter European Science Foundation Strasbourg France
46 Investigating Life in Extreme Environments
Appendix II: Steering Committee of the ILEE initiative
Name Organisation Town Country
Dr Demir Altiner Middle East Technical University Ankara Turkey
Dr Ricardo Amils Universidad Autonoma de Madrid – Centro de Biología Molecular del CSIC Madrid Spain
Dr Fernando Barriga Universidade de Lisboa Lisbon Portugal
Dr Cynan Ellis-Evans British Antarctic Survey Cambridge United Kingdom
Dr Lucien Hoffmann CRP Gabriel Lippmann Belvaux Luxembourg
Dr Jesus Martinez-Frias CSIC/INTA – Centro de Astrobiologia Torrejón de Ardoz Spain
Dr Daniel Prieur Université de Bretagne Occidentale - Laboratoire de Microbiologie des Environnements Extrêmes Plouzane France
Dr Zoltán Varga Debrecen University Debrecen Hungary
47Investigating Life in Extreme Environments
Saturday 5 November 2005
14.30 Registration at the hotel reception and ESF desk18.00 Icebreaker20.00 Dinner
Sunday 6 November 2005
08.30-08.45 Conference OpeningDaniel Prieur – Chair of the Workshop
08.45-09.00 ESF PresentationBertil Andersson (ESF Chief Executive)
09.00-09.20 Keynote AddressInvestigating Life in Extreme Environment – Challenges and Perspectives Jean-François Minster (CNRS, Paris, France – Marine Board Chairperson)
09.20-09.40 Keynote AddressExtreme life at deep-sea hydrothermal vents from a geological perspective Steven Scott (University of Toronto, Canada)
Session 1Defining and characterising the boundary conditions of ‘extreme’ environmentsSession chair: Daniel PrieurRapporteur: Ricardo Amils
09.40-10.00 Biocatalysis under extreme conditions – harvesting the potential ofextremophiles - Garo Antranikian
10.00-10.10 Discussion
10.10-10.30 Human adaptation – from the white to the red planet - Didier Schmitt10.30-10.40 Discussion
10.40-11.10 Coffee break
Session 2Molecular adaptation and stability in extreme environments
11.10-11.30 Peptides under extreme conditions: the foundation for the origin of life?Bernd Rode
11.30-11.40 Discussion
11.40-12.00 The interplay between metabolic programmes and stress-responseprogrammes in hydrocarbon-degrading bacteria - Victor de Lorenzo
12.00-12.10 Discussion
12.30 Lunch
14.00-14.50 Round table discussion
Appendix III: ILEE Workshop Programme (6-8 November 2005)
48 Investigating Life in Extreme Environments
Microbial life in extremeenvironments
Chair: Michael DansonRapporteur: Kaï-Uwe Hinrichs
General principles ofmicrobial ecosystems inextreme environmentsHelga Stan Lotter
Discussion
Microorganisms in snow and iceRoland Psenner
Discussion
Diversity ofhyperthermophilicarchaea and theirviruses in extremethermal environmentsDavid Prangishvili
Discussion
Life strategies of plantsin extremeenvironments
Chair: Francesco LoretoRapporteur:Klaus-Ulrich Valentin
Life strategies andevolution of rock blackfungi from Antarcticcold desertSilvano Onofri
Discussion
Photosynthesis inAntarctic lichens inextreme environments:plant stress physiologyapproachesMilos Bartak
Discussion
Life strategies ofseaweeds and ice-algaein polar watersChristian Wiencke
Discussion
Life strategies ofanimals in extremeenvironments
Chair: Arnoldus BlixRapporteur:Françoise Gail
The influence of highcarbon dioxideatmosphere on marineorganisms on a macroto micro scaleHenry Elderfield
Discussion
Metazoans living arounddeep-sea vents:evidence for avoidancemechanismsDavid Dixon
Discussion
How polar birds copewith extremetemperaturesYvon Le Maho
Discussion
Human adaptation inextreme environments
Co-Rapporteur :Helmut Hinghofer-SzalkayCo-Rapporteur :Elisabeth Vestergaard
Human aspects ofspaceflight andplanetary explorationDidier Schmitt
Discussion
Vascular adaptations inextreme environmentsSergio Pillon
Discussion
Personnel in extremeenvironments:psychosocial challengesGro Sandal
Discussion
Session 3 Session 4 Session 5 Session 6
Splinter Presentation Session15.00-17.30
15.00-15.15
15.15-15.20
15.20-15.35
15.35-15.40
15.40-15.55
15.55-16.00
16.00-16.30 Coffee Break
49Investigating Life in Extreme Environments
Terrestrial ecosystemsin extremeenvironmentsYoseph Steinberger
Discussion
Microbial life and spaceenvironmentGerda Horneck
Discussion
Acidic environmentsunder the control of ironRicardo Amils
Discussion
Poster session
Role of the cell wall inthe ability of theresurrection plantCraterostigmaplantagineum to survive dryingSimon McQueen-Mason
Discussion
Plant growing in spaceFranck Ditengou
Discussion
Adaptation ofphotosyntheticorganisms to extremeultraviolet conditionsImre Vass
Discussion
Ecophysiology ofinvertebrates living inperiglacial habitat in ice-retaining stony debris.Miroslav Zacharda
Discussion
Food web structure and dynamics along a productive/latitudegradient in extremeenvironments at presentand in the pastErik Jeppesen
Discussion
The adaptive evolutionof polar fishes: anexciting challenge forbiologyGuido di Prisco
Discussion
Round table discussionSocial aspects ofhuman adaptation inextreme environments
16.30-16.45
16.45-16.50
16.50-17.05
17.10-17.25
17.25-17.30
17.30-19.30
17.05-17.10
Appendix III: ILEE Workshop Programme
50 Investigating Life in Extreme Environments
Monday 7 November 2005
Session 7Enabling technologies and applicationsModerator: Farzam Ranjbaran
08.30-08.50 Subglacial waters - melting probe technology - Stephan Ulamec
08.50-09.00 Discussion
09.00-09.20 Technological challenges of planetary protection - Gerhard Schwehm
09.20-09.30 Discussion
09.30-09.50 The concept of LabHorta, the first land-based laboratory/experimental platformfor conducting experimental studies on hydrothermal-vent organisms from theMid-Atlantic Ridge - Eniko Kadar
09.50-10.00 Discussion
10.00-10.30 Coffee break
10.30-12.30 Flash presentations
12.30 Lunch
14.00-16.30 Working Group Discussions
16.30-17.00 Coffee break
17.00-18.00 Report from the Working Groups
18.00-19.00 Round table discussion: Life in Extreme Environments at the European Level –ESF Boards and Committees Perspectives Niamh Connolly (Marine Board), Arja Kallio (LESC)
19.00-19.20 Conclusion of the Workshop
20.30 Gala Dinner
Tuesday 8 November 2005
Breakfast and departure
Closed Session
09.00-12.00 Wrap Up, Synthesis, Recommendations, Report Structure
12.00-13.30 Lunch and departure
WG1 WG2 WG3 WG4 WG5Enabling technologies Microbial life Life strategies Life strategies Human adaptationand applications in extreme of plants in of animals in in extreme
environments extreme extreme environmentsenvironments environments
Discussion Discussion Discussion Discussion Discussion
51Investigating Life in Extreme Environments
Monday 7 November 2005
10.30-12.30
1. Proteins from Extremophiles as Probes for the Design of Advanced Nano-Sensors forAnalyses of High Social InterestsSabato D'Auria (Institute of Protein Biochemistry, Naples, Italy)
2. Raman Spectroscopic Analysis of Biologically Modified Geological Substrates : Markers forthe Astrobiological Exploration of MarsHowell Edwards (Chemical and Forensic Sciences, School of Pharmacy, Univ. of Bradford, UK)
3. Information and Communication Technologies for remote support in investigating life in extreme environmentsDiego Liberati (CNR, Dipartimento di Elettronica e Informazione – Politecnico, Milano, Italy)
4. Exploration of subglacial lakesMartin Siegert (University of Bristol, Bristol Glaciology Centre, UK)
5. Molecular Basis of Response Mechanisms to Environmental Stresses in ThermophilesSimonetta Bartolucci (Dipartimento di Biologia Strutturale e Funzionale, Università degli Studi di Napoli Federico II, Italy)
6. On the possible relevance of biotic control on the microbial food web in antarctic lakesAntonio Camacho (Department of Microbiology and Ecology, University of Valencia, Spain)
7. Microbial ecology and biotechnology of hydrothermal and sulphuric marine cavesFrancesco Canganella (Dept. of Agrobiology and Agrochemistry, University of Tuscia, Viterbo, Italy)
8. Death and longevity at the salinity limits of lifeTerence McGenity (University of Essex, Department of Biological Sciences, Colchester, UK)
9. Microbial life in extreme acidic aquatic environmentsChristian Wilhelm (University of Leipzig, Department of Plant Physiology, Germany)
10. Technical challenges for probing subglacial lakes in Antarctica Jean Robert Petit (LGGE/CNRS, St Martin d’Hères, France)
11. Molecular aspects of plant adaptation to continuous heavy metal exposure: Thlaspi caerulescensMark Aarts (Lab of Genetics, Wageningen University, Wageningen, The Netherlands)
12. Effects of global environmental changes on terrestrial vegetation: Implications for extreme environmentsMauro Centritto (CNR - Istituto sull’Inquinamento Atmosferico, Monterotondo Scalo, Monterotondo Scalo, Italy)
ESF Joint InitiativeInvestigation Life in Extreme EnvironmentFlash Presentations*
Appendix III: ILEE Workshop Programme
52 Investigating Life in Extreme Environments
13. Cyanobacteria and algae polar ecologyJosef Elster (Academy of Sciences of the Czech Republic, Institute of Botany)
14. Adaptive thermogenesis in cold acclimation and hibernationBarbara Cannon (The Wenner-Gren Institute, Stockholm University, Sweden)
15. Strategies of autotrophic organisms in extremely acidic waters to overcome limitationsinduced by acidity.Dieter Lessmann (Department of Freshwater Conservation Brandenburg University of Technology, Germany)
16. Evolution of zooplankton communities in extreme acidic environmentsRainer Deneke and Brigitte Nixdorf (Brandenburg University of Technology, Environmental Science and Process Engineering, Bad Saarow, Germany)
17. Life shaped by Antarctic ice shelves: spatially explicit modelling can predict climate induced community shiftsJulian Gutt (Alfred Wegener Institute for Polar and Marine Research, Bremerhaven, Germany)
18. How ‘bioengineer’ animals shape their extreme environment at deep-sea ventsNadine Le Bris (IFREMER, Département Etude des Ecosystèmes Profonds, Plouzané, France)
19. Vertebrates life in oxygen free environmentsGøran Nilsson (University of Oslo, Department of Molecular Biosciences, Norway)
* Each flash-presentation is of 5 minutes (including discussion)
53Investigating Life in Extreme Environments
Poster Session 1: Life Strategies of animals in extreme environments
ANIM-01 Adaptive thermogenesis in cold acclimation and hibernationBarbara Cannon (The Wenner-Gren Institute, Stockholm University, Sweden)
ANIM-02 Evolution of aquatic communities in extremely acidic environments (pH 0 - 4)Rainer Deneke (Brandenburg University of Technology, Environmental Science and ProcessEngineering, Bad Saarow, Germany)
ANIM-03 Anthropogenic Radionuclides in European Arctic Marine HabitatsDirk Dethleff (Institute for Polar Ecology, Kiel University, Germany)
ANIM-04 Life shaped by Antarctic ice shelves: spatially explicit modelling can predictclimate induced community shiftsJulian Gutt (Alfred Wegener Institute for Polar and Marine Research, Bremerhaven,Germany)
ANIM-05 Adaptation of terrestrial invertebrates to extreme cold environmentsIan Hodkinson (Liverpool John Moores University, dpt.of Biological and Earth Sciences, UK)
ANIM-06 Industrial barrens: exploration of the novel extreme environmentMikhail V. Kozlov (University of Turku, Section of Ecology, Finland)
ANIM-07 How ‘bioengineer’ animals shape their extreme environment at deep-sea ventsNadine Le Bris (IFREMER, Département Etude des Ecosystèmes Profonds, Plouzané,France)
ANIM-08 Ecological peculiarities of the symbiosis between the scale-worm Branchipolynoeseepensis and the giant mussels Bathymodiolus spp. rom hydrothermal ventsDaniel Martin
ANIM-09 Vertebrates life in oxygen free environmentsGøran Nilsson (University of Oslo, Department of Molecular Biosciences, Norway)
ANIM-10 Living in a critical salinity zone – the physiology of the marine amphipodGammarus oceanicus from the Baltic Sea Monika Normant (University of Gdansk, Institute of Oceanography, Gdynia, Poland)
ANIM-11 The different sensibility to temperature of fish and mammal metallothioneins islinked to the evolution of protein domainsElio Parisi (CNR Institute of Protein Biochemistry, Napoli, Italy)
ANIM-12 Colonisation of deep extreme environment: how to break the pressure barrier?Florence Pradillon (Max-Planck Institute for Marine Microbiology, Molecular Ecology,Bremen, Germany)
ANIM-13 Diversity, distribution and functioning of deep-water chemosynthetic ecosystems- ChEss, a Census of Marine Life pilot projectEva Ramirez-Llodra (Institut de Ciències del Mar, CMIMA-CSIC, Barcelona, Spain)
ESF Joint InitiativeInvestigation Life in Extreme EnvironmentList of Posters
Appendix III: ILEE Workshop Programme
54 Investigating Life in Extreme Environments
ANIM-14 Extreme Oxidative Threats in Deep-Sea OrganismsJean-François Rees (Université de Louvain, Institut des Sciences de la Vie, Belgium)
ANIM-15 Ecophysiology of invertebrates living in periglacial habitat in ice-retaining stonydebris: Physiological questionsVladimir Sustr (Academy of Sciences of the Czech Republic, Institute of Soil Biology, _eskéBud_jovice, Czech Republic)Miloslav Zacharda (Academy of Sciences of the Czech Republic, Institute of SystemsBiology and Ecology, _eské Bud_jovice, Czech Republic)
Poster Session 2: Microbial life in extreme environments
MICR-01 Molecular Basis of Response Mechanisms to Environmental Stresses inThermophilesSimonetta Bartolucci (Dipartimento di Biologia Strutturale e Funzionale, Università degli Studi di Napoli Federico II, Italy)
MICR-02 On the possible relevance of biotic control on the microbial food web in antarcticlakesAntonio Camacho (Department of Microbiology and Ecology, University of Valencia, Spain
MICR-03 Microbial ecology and biotechnology of hydrothermal and sulphuric marine cavesFrancesco Canganella (Dept. of Agrobiology and Agrochemistry, University of Tuscia, Viterbo, Italy)
MICR-04 Anoxia in sulfide rich aquatic environments: microbial life under extremely lowredox potentialsEmilio Casamayor (Centre d’Estudis Avançats de Blanes (CSIC), Blanes, Spain)
MICR-05 Thioautotrophic symbiosis: towards a new step in eukaryote evolution?Stéphane Hourdez (CNRS/UPMC - Station Biologique, Roscoff, France)
MICR-06 Use of continuous culture to access the cultivable thermophilic microbial diversityfrom deep-sea hydrothermal ventAnne Godfroy (IFREMER, Laboratoire de Microbiologie des Environnements Extrêmes ,Plouzané, France)
MICR-07 Halophilic fungi Nina Gunde-Cimerman (University of Ljubljana, Slovenia)
MICR-08 Life in sea iceJohanna Ikävalko (University of Helsinki, Dept. of Biological and EcologicalSciences/Aquatic Sciences, Finland)
MICR-09 Activity and diversity of methanogenic Archaea under extreme environmentalconditions in Late Pleistocene permafrost sediments of the Lena Delta, SiberiaGerman Jurgens (University of Helsinki, Dept. of Applied Chemistry and Microbiology,Finland)
55Investigating Life in Extreme Environments
MICR-10 Morphological diversity of hyperthermophilic viruses and virus-like particles indeep-sea hydrothermal ventsMarc Le Romancer (Université de Bretagne Occidentale, Institut Universitaire Européen dela Mer (IUEM), Plouzané, France)
MICR-12 Translation in extremely thermophilic archaea: insights on the early evolution ofthe decoding machineryPaola Londei (University of Bari, Dpt. DIBIFIM, Italy)
MICR-13 Death and longevity at the salinity limits of lifeTerence McGenity (University of Essex, Department of Biological Sciences, Colchester, UK)
MICR-17 Search for extraterrestrial hereditable informationWilfred F.M. Röling (Vrije Universiteit, Molecular Cell Physiology, FALW , Amsterdam, TheNetherlands)
MICR-18 Microbial diversity of snow collected from Himalayan peaks: a study system for stratospheric transfer of microorganisms and colonization of extremely coldenvironmentsDavid Pearce (British Antarctic Survey, Cambridge, UK)
MICR-19 Phenotypic and genotypic diversity of photosynthetic protists in microbial mats ofAntarctic lakesKoen Sabbe (Ghent University, Laboratory of Protistology & Aquatic Ecology, Ghent,Belgium)
MICR-20 Microbial life in extreme acidic aquatic environmentsChristian Wilhelm (University of Leipzig, Department of Plant Physiology, Germany)
MICR-21 The Vostock Project (TBC)Jean-Robert Petit (LGGE - Laboratoire de Glaciologie et Géophysique de l'Environnement,St Martin d'Hères, France)
MICRO-22 Extremely acidophilic prokaryotes: new insights into their physiological andphylogenetic biodiversitiesD. Barrie Johnson and Kevin B. Hallberg (School of Biological Sciences, University ofWales, Bangor, UK)
Poster Session 3: Life Strategies of plants in extreme environments
PLAN-01 Molecular aspects of plant adaptation to continuous heavy metal exposure:Thlaspi caerulescensMark Aarts (Lab of Genetics, Wageningen University, Wageningen, The Netherlands)
PLAN-02 Effects of global environmental changes on terrestrial vegetation: Implications forextreme environmentsMauro Centritto (CNR - Istituto sull’Inquinamento Atmosferico, Monterotondo Scalo, Monterotondo Scalo, Italy)
Appendix III: ILEE Workshop Programme
56 Investigating Life in Extreme Environments
PLAN-03 Preservation of extinct and extant forms of life under terrestrial magneticanomaliesCristina Dobrota ( Babes-Bolyai University, Experimental Biology Dept., Cluj-Napoca,Romania)
PLAN-04 Cyanobacteria and algae polar ecologyJosef Elster (Academy of Sciences of the Czech Republic, Institute of Botany)
PLAN-05 Ecophysiological responses of halophytes to saline environements -Sustainableuse, conservation and developmentHans-Werner Koyro (Justus-Liebig-University of Giessen, Institute for Plant Ecology,Germany)Vladimir Sustr (Academy of Sciences of the Czech Republic, Institute of Soil Biology)
PLAN-06 Arabidopsis root growth in simulated microgravity conditionsFernando Migliaccio (CNR, Institute of Agroenvironmental and Forest Biology,Monterotondo, Italy)
PLAN-07 Life expansion in NE Sorkapp Land, Spitsbergen, under the contemporaryclimate warmingWieslaw Ziaja (Jagiellonian University, Institute of Geography and Spatial Management,Krakow, Poland)
PLAN-08 Adaptation of high plants or cyanobacteria to high visible light intensity or UVBLello Zolla (University of Tuscia, Dip. Environmental Science, Viterbo, Italy)
PLAN-09 “United we stand”: The role of positive plant-plant interactions in the extreme environmentsElena Zvereva (University of Turku, Section of Ecology, Turku, Finland)
Poster Session 4: Enabling technologies and applications
TECH-01 Proteins from Extremophiles as Probes for the Design of Advanced Nano-Sensorsfor Analyses of High Social InterestsSabato D'Auria (Institute of Protein Biochemistry, Naples, Italy)
TECH-02 Raman Spectroscopic Analysis of Biologically Modified Geological Substrates :Markers for the Astrobiological Exploration of MarsHowell Edwards (Chemical and Forensic Sciences, School of Pharmacy, Univ. of Bradford, UK)
TECH-03 Information and Communication Technologies for remote support in investigatinglife in extreme environmentsDiego Liberati (CNR, Dipartimento di Elettronica e Informazione – Politecnico, Milano, Italy)
TECH-04 Fluorescence in situ hybridisation coupled to ultra-small immunogold detection toidentify prokaryotic cells on minerals by electron and X-ray microscopyBénédicte Menez (CNRS/Institut de Physique du Globe de Paris, France)
57Investigating Life in Extreme Environments
TECH-05 Multiparametric cytometry and High-speed cell sorting to investigate Mechanismsof cold adaptation of psychrophilic microorganismsSusann Müller (UFZ Centre for Environmental Research Leipzig-Halle, Department ofEnvironmental Microbiology, Leipzig, Germany)
TECH-06 ‘Sensory mechanisms in the deep sea and appropriate methods for observingdeep sea animalsJulian Partridge (University of Bristol, School of Biological Sciences, UK)
TECH-07 The exploration of Subglacial Lake Ellsworth, West AntarcticaMartin J. Siegert (Bristol Glaciology Centre, School of Geographical Sciences, University ofBristol, UK)
Poster Session 5: Human Adaptation to Extreme Environments
HUMA-01 The Post-Tsunami crops through salt tolerant speciesAdriana Galvani (Dipartimento di Scienze Economiche, Università di Bologna, Italy)
HUMA-02 Phenogenotypic factors in the formation of non-specific human resistance andthe level of biological aging under extreme environmentsEugene Kobyliansky (Tel-Aviv University, Israel)
HUMA-03 Research needs related on human adaptation to global change for ahead in thenorthern most EuropeJuhani Hassi (Centre for Arctic Medicine, University of Oulu, Finland)
HUMA-04 DADLE: a Challenge Towards Human “Hibernation”? (TBC)Carlo Zancanaro (Department of Morphological & Biomedical Sciences, University ofVerona, Italy)
HUMA-05 Strategies of human adaptation to high altitude Jean-Paul Richalet (Université Paris 13, France)
Appendix III: ILEE Workshop Programme
58 Investigating Life in Extreme Environments
59Investigating Life in Extreme Environments
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Investigating Life in Extrem
e Environm
ents • Europ
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