Rainer KurmayerRainer Kurmayer

 

Present position:

Head of Research Institute, Assistant Professor
Head of working group

Research area:

Molecular ecology and ecophysiology of algae (cyanobacteria)

Phone:
Fax:
e-mail:

+43 512 507-50242
+43 512 507-50299
rainer.kurmayer@uibk.ac.at

 


Employment History · Research · Staff · Projects · Teaching · Publications · Memberships


Employment History

 

  • June 2013 – present: Assistant Professor at the Research Institute for Limnology, Mondsee, University of Innsbruck; University Docent at the University of Vienna, Faculty of Life Sciences (since 2007)

  • Sept 1, 2012 - May, 2013: Senior scientist at the Research Institute for Limnology, Mondsee, University of Innsbruck

  • 2007-Aug 31, 2012: Senior scientist at the Institute for Limnology in Mondsee, Austrian Academy of Sciences
  • 2001-2007: Junior scientist at the Institute for Limnology in Mondsee, Austrian Academy of Sciences

  • 1999-2001: Post doc position at the Federal Environmental Agency and Humboldt University, Berlin, Germany

  • 1997-1999: PhD fellow at the University of Vienna, Doc fellowship by the Austrian Academy of Sciences


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Research

Molecular ecology and evolution of toxin-producing cyanobacteria

Harmful algal blooms formed by cyanobacteria (cyanoHABs) are a global phenomenon observed both in fresh and brackish water. The human society is facing a couple of toxic compounds (“cyanotoxins”; e.g. the microcystins) of different chemical structure that are produced by several genera (Microcystis, Planktothrix, Anabaena), with some of them leading to outbreaks and diseases in livestock and humans.

netsample

Sampling of Planktothrix blooms in lakes in the Alps, Photo: R. Kurmayer

 

It is a challenge to identify the factors leading to toxic outbreaks which is due to the development of toxic ecotypes carrying the genes for the production of a certain toxin. In both theoretical and experimental ecology a certain toxin-producing phenotype is considered relatively static, i.e. costs and benefits of toxin synthesis are related to its ecological fitness.

I argue that this approach represents a single snapshot sampling event and cannot address the diversification of toxic and nontoxic ecotypes driven by selective factors also not directly linked to toxin production. Further, as lakes and reservoirs are ephemeral in terms of geological time scales, the current success of some toxic bloom-forming species may at least partly also result from opportunistic abilities than from an adaptation to a specific freshwater habitat.

The elucidation of the genes involved in the synthesis of these toxins paved the way to

(i) Explore time scales and mechanisms in the evolution of responsible genes involved in cyanoHAB formation.

(ii) Postulate hypotheses about factors driving the innovation of toxin synthesis

(i)  Forecast the ecological fitness of toxic/non-toxic genotypes during cyanoHAB development

boats

Microcystis bloom formation in Nyanza Gulf (L. Victoria), Photo: R. Kurmayer

 

Answering these questions would help to understand whether the  biodiversity and evolution of cyanotoxins is favored by the  global increase of cyanoHAB formation. These questions are  addressed by integrating of the ecology and the genetic analysis of ecotypes through a comprehensive field and laboratory research program.

Project:

MOBILOMICS, “Mobilomics of toxin production in cyanobacteria”, Austrian Science Fund, FWF P24070, (2012-2015), http://www.uibk.ac.at/limno/research/projects/mobilomics/



Consequences of toxins produced by algal blooms

Microcystis

Microcystis with epibiontic protozoa and algae and bacteria, Photo: R. Kurmayer

 

The macroscopic colonies of cyanoHAB organisms provide a  habitat to heterotrophic bacteria, protozoans, and epibiontic  algae. Through the close physical coupling the cyanotoxins can have significant effects, though these include probably both  beneficial effects (mutualism) as well as deleterious effects  (parasitism). Thus the effects of bioactive peptides on planktonic diversity are not straightforward. One question is  whether the production of cyanotoxins contributes to the stability/duration of bloom events, for example due to  co-evolutionary adaptation between producers and potential target organisms.

Due to massive accumulation of cyanoHAB organisms at the surface or along the shore, the toxins that are produced can occur in high concentrations. This is of relevance when contaminated raw water is used for drinking as well as for recreation and tourism. Various aquatic biota like fish, crustaceans and mussels that are used for nutrition have been shown to contain cyanotoxins.

In Austria cyanoHABs occur in ponds and gravel pit lakes frequently after sunny periods during summer and autumn. In deep physically stratified lakes in the Alps, high abundances of the cyanobacterium Planktothrix can occur and under bloom conditions microcystin has been shown to be transferred up to the highest trophic levels.

fish

Small fish (mostly Rastineobola argentea) occurring in Lake Victoria contain the hepatotoxin microcystin as revealed by ELISA, Photo: R. Kurmayer

 

As the occurrence of cyanoHABs is of public interest, water samples sent by technical offices or authorities are analyzed for the two most frequent cyanotoxins in freshwater, microcystin and anatoxin-a. Currently, in Austria a national guideline on the estimation of health risks associated with cyanoHABs occurrence is not available. Thus public requests are referred to the recommendation published by the Federal Environmental Agency in Germany.

Projects:

CYANOCOST, “Cyanobacteria booms and toxins in water resources : Occurrence, impacts and management”, EU COST Action ES1105, (2012-2016), http://cyanocost.com/

IPGL, International Training Programs in Limnology, (http://www.wau.boku.ac.at/22151.html ).


Alpine lakes as in situ observatories for climate change effects

Twenger Almsee

Ice break up in Twenger Alm See (2118 m aSL) in the Niedere Tauern Region during June 2012, Photo: V. Schaidreiter

 

Both direct impacts (i.e. increase in water temperature) and indirect impacts (i.e. increase in physical lake stratification, longer vegetation period, eutrophication due to increased precipitation) are predicted to favor the occurrence of nuisance algae such as cyanoHAB organisms. It is possible that the effects of climatic change are likely to run counter to the reduction in nutrient loading which is aimed for by the water management.

We aim to understand the factors that regulate the growth of algae and cyanobacteria both through direct measurements in the field and also through modelling of cyanoHAB growth in the water column. In a recent project we use high-throughput sequencing techniques to monitor plankton community composition as well as the immigration of certain nuisance species as a consequence of climatic change.

For deep, thermally stratified lakes, we apply an established algal growth model developed by Prof. A. E. Walsby (University of Bristol, UK), which is based on an algorithm that describes the change in growth in relation to irradiance and temperature, based on measurement in laboratory cultures. The algorithm is used to calculate the quantity of cells from measurements of irradiance and temperature at hourly intervals and every depth meter down the water column. This model enables the quantification of effects on growth parameters based on gradual changes in water column physical conditions.

Asterionella

Asterionella formosa blooming in Lake Moaralmsee (1825 m aSL) in the Niedere Tauern Region during summer 2011, Photo: R. Kurmayer

 


Projects:

DETECTIVE, “Decadal Detection of Biodiversity in Alpine Lakes” 2010-2013, Alpine Research Program of the ÖAW, http://www.uibk.ac.at/limno/research/projects/detective/ ;

RADICAL, “Risk Analysis of Direct and Indirect Climate effects on deep Austrian Lake Ecosystems (Austrian Climate Research Program (ACRP), 2010-2013, http://www.uibk.ac.at/limno/research/projects/radical/

The ultimate aim is to use these results for a protection, monitoring and restoration of aquatic ecosystems.

Projects:

DETECTIVE, “Decadal Detection of Biodiversity in Alpine Lakes” 2010-2013, Alpine Research Program of the ÖAW, http://www.uibk.ac.at/limno/research/projects/detective/ ;

RADICAL, “Risk Analysis of Direct and Indirect Climate effects on deep Austrian Lake Ecosystems (Austrian Climate Research Program (ACRP), 2010-2013, http://www.uibk.ac.at/limno/research/projects/radical/


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Staff

Former members

PhD students

Post-Doc

  • Guntram Christiansen (FWF projects P18185, P20231, Cyano-Biofuels, DETECTIVE), 2005-2011 

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Projects

  • Cyanobacterial blooms and toxins in water resources: Occurrence, impacts and management. ESSEM COST Action ES1105 (Apr 2012 - Mar 2016, http://www.cost.eu/domains_actions/essem/Actions/ES1105 ). This new COST Action will establish a European network in order to address the issues of cyanobacterial blooms, their toxicity, monitoring, risk assessment, treatment and management and has been approved for involving >30 partners from 24 countries. http://cyanocost.com/

  • "Mobilomics of toxin production in cyanobacteria" funded by the Austrian Science Fund P24070,
    www.fwf.ac.at | Summary

    Finished projects
  • Risk Analysis of Direct and Indirect Climate effects on deep Austrian Lake Ecosystems (RADICAL) Climate and Energy Fund of the Federal State, ACRP (2009-2012)
  • “DEcadal deTECTion of biodIVErsity in alpine lakes, DETECTIVE - Global warming threatens biodiversity in (ultra)-sensitive Alpine lakes: an assessment of past, present and future scenarios" funded by the Alpine Research Programme of the Austrian Academy of Sciences (Dec 2009 - Dec 2012), National and International Research Programme.
  • “Natural products from cyanobacteria”, funded by Cyano Biotech GmbH, Chausseestr. 117, D-10115 Berlin, www.cyano-biotech.com
  • “Genotype determined toxin content in cyanobacteria”, funded by the Austrian Science Fund, P20231, www.fwf.at, July 2008 – June 2011 Dez 2011, finished. As a deliverable to disseminate the projects results an international mutagenisation workshop has been held in July 2011 (Link to: announcement; report)
  • “Linking Cyanotoxin production to genetic diversity” funded by the Austrian Science Fund P15709, www.fwf.ac.at , finished
  • "Microevolution of toxin synthesis in cyanobacteria" funded by the Austrian Science Fund P18185, www.fwf.ac.at , finished
  • “Use of cyanobacteria for the production of ethanol”, funded by Cyano BioFuels, Magnusstrasse 11, D-12489 Berlin, www.cyano-biofuels.com, finished
  • “Toxic and other bioactive PEPtides in Cyanobacteria” (PEPCY), funded by the European Community, QLK4-CT-2002-02634, finished, www.pepcy.de , Link to Final report. PDF (FINAL report PEPCY)
    As a deliverable for the PEPCY project “A manual on the use of genetic techniques to quantify genotypes of cyanobacteria in freshwater” has been compiled. Link to manual. PDF

fwf.jpg

  Logo "Klima Energie Fonds"OEAW-logo


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Teaching

Lectures and practica

717031 VU/2 Evolution and Phylogeny of Algae (in German), Winter semester 2013/14, (every second year), University of Innsbruck, Institute for Botany, Sternwartestraße 15, A-6020 Innsbruck

717039 SE/1 Biomonitoring of aquatic environments, part of Elective Module 4A: Special Hydrobotany, Winter semester 2013/14 (every second year), University of Innsbruck, Institute for Botany, Sternwartestraße 15, A-6020 Innsbruck

718041 UE/3 Basic Laboratory Course, Winter semester 2013/14 (yearly), University of Innsbruck, Institute for Microbiology, Labor Biologie 0/406, Technik, A-6020 Innsbruck

300552 VO Physiology and Ecology of Cyanobacteria (in German), Winter Semester (yearly), University of Vienna, UZAI, Althanstrasse 14, Konferenzraum Ökologie

Skriptum: Ökologie und Physiologie von Cyanobakterien (Teile 1-7), (PDF dazu).

Vorbesprechung, Skriptum: “Cyanobakterien-Wen kümmerts” PDF

Diploma theses

  1. Quantification of toxin genes in surface waters by real-time PCR (Thomas Kutzenberger, Univ. of Vienna) 107pp.
  2. Assessment of toxin producing cyanobacteria in Mwanza Gulf - Lake Victoria (Baraka Sekadende, IHE Delft, The Netherlands) 47pp.
  3. Phenotypic and genotypic characterisation of isolates of the toxic cyanobacterium Planktothrix (Marlies Gumpenberger, Univ. of Salzburg) 122pp.
  4. Characterising the content of two cyclic peptides and phytoplankton pigments in isolates of the toxic cyanobacterium Planktothrix spp. under maximum growth rate conditions (Simone Kosol, Univ. of Salzburg) 87pp.
  5. Toxic cyanobacteria in Ugandan freshwater habitats (William Okello, IHE Delft, The Netherlands) 70pp.
  6. Abundance of specific microcystin-deficient genotypes in Planktothrix rubescens populations of the Salzkammergut Area, Upper Austria (Qian Liu, IHE Delft, The Netherlands) 61pp.
  7. Phylogenetic characterisation of mutations inactivating the biosynthesis of the toxic heptapeptide microcystin (Carole Molitor, Univ. of Innsbruck) 150pp.
  8. Seasonal abundance of mutation and recombination in microcystin genes in populations of cyanobacteria (Veronika Ostermaier, Univ. of Munich) 104pp.
  9. Analysis of phytoplankton composition during stratification of Lake Victoria (Mwanza Gulf, Tanzania), Athanasio Mbonde (IHE Delft, The Netherlands) 71pp.
  10. The concentration of microcystins in Rastineobola argentea in Nyanza Gulf, Lake Victoria, Kenya (Benard Simiyu, IHE Delft, the Netherlands) 69pp.
  11. Genetic variation and recombination processes affecting the synthesis of bioactive peptides in the cyanobacterium Planktothrix (Mark Frei, Univ. of Vienna) 121pp.

PhD theses

  1. Diversity in abundance of toxic genotypes in populations of the cyanobacterium Planktothrix spp. (Eva Schober, Univ. of Vienna) 129pp.
  2. Molecular and chemical characterisation of toxic cyanobacteria in Ugandan freshwater (William Okello, Univ. of Vienna) 109pp.
  3. Effects of cage fish farming systems on phytoplankton and water quality in small ponds of East Africa (Nadja Straubinger, Univ. of Vienna), EU project BOMOSA, http://bomosa.oeaw.ac.at ongoing
  4. Time in microevolution of microcystin synthesis (Veronika Ostermaier, Univ. of Vienna), DOC-FORTE fellowship 2009-2011. 161pp.
  5. Assessment of planktonic diversity in alpine lakes by means of ultra-deep sequencing (Stephan Blank, Univ. of Vienna), project DETECTIVE (Link Project website), ongoing.
  6. Molecular Ecology and Physiology of Cyanobacteria in the Ecosystems of Freshwater and Sediments (Qin Chen, Northwest A&F University, Yangling, Shaanxi Province), ongoing.



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Publications

2014

  • Niedermeyer T. J., Schmieder P., Kurmayer R. (2014) Isolation of Microcystins from the Cyanobacterium Planktothrix rubescens Strain No80. Natural Products and Bioprospecting: 1-9. DOI 10.1007/s13659-013-0001-3

2013

  • Ostermaier, V., Christiansen G., Schanz, F., Kurmayer, R. (2013) Genetic variability of microcystin biosynthesis genes in Planktothrix as elucidated from samples preserved by heat desiccation during three decades. PLOS One. DOI: 10.1371/journal.pone.0080177 PDF

  • Nevalainen, L., Ketola, M., Korosi, J.B., Manca, M., Kurmayer, R., Koinig, K., Psenner, R., Luoto, TP. (2013) Zooplankton (Cladocera) species turnover and long-term decline of Daphnia in two high mountain lakes in the Austrian Alps. Hydrobiologia, DOI 10.1007/s10750-013-1676-5.
  • Rohrlack, T., Christiansen, G., Kurmayer, R. (2013). A putative anti-parasite defensive system in the cyanobacterium Planktothrix involving ribosomal and non-ribosomal oligopeptides. Applied and Environmental Microbiology 79, 2642-2647 PDF

2012

  • Ostermaier V., Schanz F., Köster  O., and Kurmayer R. (2012) Stability of toxin gene proportion in red-pigmented populations of the cyanobacterium Planktothrix during 29 years of re-oligotrophication of Lake Zürich. BMC Biology 10:100, doi:10.1186/1741-7007-10-100.

  • Sitoki,·L.,·Kurmayer,·R.,·Rott,·E.·(2012)· Spatial·variation·of·phytoplankton·composition,·biovolume,·and· resulting·microcystin·concentrations·in·the·Nyanza·Gulf·(Lake· Victoria,·Kenya).·Hydrobiologia 691:109-122. DOI·10.1007/s10750-012-1062-8 PDF

2011

  • Savichtcheva, O., Debroas, D., Kurmayer, R., Villar, C., Jenny J.P., Fabien A., Perga M.E., Domaizon I. (2011) Quantitative PCR enumeration of total and toxic Planktothrix rubescens/agardhii and other cyanobacteria in preserved DNA isolated from lake sediments. Applied and Environmental Microbiology 77:8744-8753. doi: 10.1128/AEM.06106-11 PDF
  • Christiansen, G., Philmus, B., Hemscheidt, T., and Kurmayer, R. (2011) Genetic variation of adenylation domains of the anabaenopeptin synthesis operon and the evolution of substrate promiscuity. Journal of Bacteriology 193, 3822-3831 PDF

  • Okello, W., Kurmayer, R. (2011) Seasonal development of cyanobacteria and microcystin production in Ugandan freshwater lakes. Lakes & Reservoirs: Research & Management 16:123-135.

  • Babanazarova, O. V., Kurmayer R., Sidelev S. I., Aleksandrina E. M., and Sakharova E .G. (2011) Phytoplankton structure and microcystine concentration in the highly eutrophic Nero Lake . Water Resources 39(2):229-236. PDF

  • Kurmayer, R., Schober E., Tonk L., Visser P., and Christiansen G. (2011) Spatial divergence in the proportions of genes encoding toxic peptide synthesis among populations of the cyanobacterium Planktothrix in European lakes. FEMS Microbiol Letters 317:127-137. PDF

  • Kurmayer, R. (2011) The toxic cyanobacterium Nostoc strain 152 produces highest amounts of microcystin and nostophycin under stress conditions. Journal of Phycology 47:200-207. PDF

2010

  • Ostermaier, V., and Kurmayer, R. (2010) Application of real-time PCR to estimate toxin production by the cyanobacterium Planktothrix sp. Applied and Environmental Microbiology 76:3495-3502. PDF

  • Okello, W; Ostermaier, V; Portmann, C; Gademann, K, and Kurmayer, R. (2010) Spatial isolation favours the divergence in microcystin net production by Microcystis in Ugandan freshwater lakes. Water Research 44: 2803-2814. PDF

  • Okello, W., Portmann, C., Erhard, M., Gademann, K., Kurmayer, R. (2010) Occurrence of microcystin-producing cyanobacteria in Ugandan freshwater habitats. Environ. Tox. 25: 367-380. PDF

2009

  • Kosol, S., Schmidt, J., and Kurmayer, R. (2009): Variation in peptide net production and growth among strains of the toxic cyanobacterium Planktothrix spp. Eur. J. Phycol. 44:49-62. PDF

  • Kurmayer, R. and Christiansen, G. (2009): The genetic basis of toxin production in Cyanobacteria. Freshwater Reviews 2:31-50 PDF

  • Ostermaier, V. and Kurmayer, R. (2009): Distribution and abundance of nontoxic mutants of cyanobacteria in lakes of the Alps. Microbial Ecology 58:323-333. PDF

2008

  • Christiansen, G., Yoshida W.Y., Blom, J., Portmann, C., Gademann, KG., Hemscheidt, T., and Kurmayer, R. (2008): Isolation and structure determination of two microcystins and sequence comparisons of McyABC adenylation domains in Planktothrix species. Journal of Natural Products 71:1881-1886. PDF Supplement

  • Christiansen G., Molitor C., Philmus B., and Kurmayer R. (2008): Non-toxic strains of cyanobacteria are the result of major gene deletion events induced by a transposable element. Molecular Biology and Evolution 25:1695-1704 Link

2007

  • Naselli-Flores, L., Barone, R., Chorus, I., and Kurmayer, R. (2007): Toxic cyanobacterial blooms under a semiarid mediterranean climate: The magnification of a problem. Environmental Toxicology 22:399-404. PDF

  • Ishida K., Christiansen G., Yoshida WY., Kurmayer R., Welker , Bonjoch J., Hertweck C., Börner T., Hemscheidt T., and Dittmann E. (2007): Biosynthetic pathway and structure analysis of aeruginoside 126A and B, cyanobacterial peptide glycosides bearing an unusual 2-carboxy-6-hydroxyoctahydroindole moiety. Chemistry and Biology 14: 565-576. Links to PDF's (Ishida et al. 2007), (preface Ishida et al. 2007)

  • Schober E., Werndl M., Laakso K., Korschineck I., Sivonen K., and Kurmayer R. (2007): Interlaboratory comparison of Taq Nuclease Assays for the quantification of the toxic cyanobacteria Microcystis sp. Journal of Microbiological Methods 69: 122-128. PDF

2006

  • Kurmayer R. and Gumpenberger M. (2006): Diversity of microcystin genotypes among populations of the filamentous cyanobacteria Planktothrix rubescens and Planktothrix agardhii. Molecular Ecology 15: 3849-3861. PDF

  • Schober E. and Kurmayer R. (2006): Evaluation of different DNA sampling techniques for the application of the real-time PCR method for the quantification of cyanobacteria in water. Letters in Applied Microbiology 42: 412-417. PDF

  • Christiansen G., Kurmayer R., Liu Q., and Börner T. (2006): Transposons inactivate the biosynthesis of the nonribosomal peptide microcystin in naturally occurring Planktothrix spp. Applied and Environmental Microbiology 72: 117-123. PDF, Cover

2005

  • Sekadende BC., Lyimo T. J., and Kurmayer R. (2005): Microcystin production by cyanobacteria in the Mwanza Gulf (Lake Victoria, Tanzania). Hydrobiologia 543: 299-304. PDF

  • Kurmayer R., Christiansen G., Gumpenberger M., and Fastner J. (2005): Genetic identification of microcystin ecotypes in toxic cyanobacteria of the genus Planktothrix. Microbiology 151: 1525-1533. PDF

2004

  • Via-Ordorika L., Fastner J., Kurmayer R., Hisbergues M., Dittmann E., Komarek J., Erhard M., and Chorus I. (2004): Distribution of microcystin-producing and non-microcystin-producing Microcystis sp. in European freshwater bodies: detection of microcystins and microcystin genes in individual colonies. Systematic and Applied Microbiology 27: 592-603. PDF

  • Kurmayer R., Christiansen G., Fastner J., and Börner T. (2004): Abundance of active and inactive microcystin genotypes in populations of the toxic cyanobacterium Planktothrix spp. Environmental Microbiology 6: 831-841. PDF, Cover

2003

  • Kurmayer R. and Kutzenberger T. (2003): Application of real-time PCR for the quantification of microcystin genotypes in a population of the toxic cyanobacterium Microcystis sp. Applied and Environmental Microbiology 69:6723-6730. PDF

  • Kurmayer R., Christiansen G., and Chorus I. (2003): The abundance of microcystin-producing genotypes correlates positively with colony size in Microcystis and determines its microcystin net production in Lake Wannsee. Applied and Environmental Microbiology 69/2: 787-795. PDF

2002

  • Kurmayer R., Dittmann E., Fastner J.and Chorus I. (2002): Diversity of microcystin genes within a population of the toxic cyanobacterium Microcystis spp. in Lake Wannsee (Berlin, Germany). Microbial Ecology 43: 107-118. PDF

2001

  • Kurmayer R. (2001): Competitive ability of Daphnia under dominance of non-toxic filamentous cyanobacteria, M. Boersma & K. Wiltshire (eds.): Proceedings of the V symposium on cladocera in Plön (Germany), Hydrobiologia 442: 279-289. PDF

2000

  • Kurmayer R. (2000): Nutritive vs. interfering effects of Anabaena sp. on Daphnia and Ceriodaphnia. Verh. Int. Ver. Limnol. 27: 3001-3005.

1999

  • Kurmayer R., and Jüttner F. (1999): Strategies for the coexistence of zooplankton with the toxic cyanobacterium Planktothrix rubescens in Lake Zürich. J. Plankt. Res. 21: 659-683. PDF

Popular scientific contributions

  • Kurmayer, R. (2006): Bedingungen und Steuerung der Toxinproduktion bei Cyanobakterien – aktueller Stand der Forschung. In: Toxinbildende Cyanobakterien (Blaualgen) in bayrischen Gewässern. Materalienband Nr. 125, Bayrisches Landesamt für Umwelt, pp. 116-143. Link to PDF (Bayr. Landesamt Bericht)

  • Kurmayer R. (2005): Dreckige Gewinner. Team vom Mondsee erforscht die Karriere von Blaualgen. Universum April 2005, No4: 91. PDF

  • ORF Science Channel, Young Science, http://science.orf.at/science/news/45445 - http://science.orf.at/science/news/96757

Editorial Board

Memberships

American Association for the Advancement of Science (AAAS)
 Ecological Society of America (ESA)
Österreichischer Alpenverein (ÖAV)
Societas Internationalis Limnologiae (SIL Austria) 
World Wild Life Fund (WWF)


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