Can mosses serve as model organisms for forest research? Stefanie J. Müller, Desirée D. Gütle, Jean-Pierre Jacquot, Ralf Reski To cite this version: Stefanie J. Müller, Desirée D. Gütle, Jean-Pierre Jacquot, Ralf Reski. Can mosses serve as model organisms for forest research?. Annals of Forest Science, Springer Nature (since 2011)/EDP Science (until 2010), 2016, 73 (1), pp.135-146. 10.1007/s13595-015-0468-7. hal-01557204 HAL Id: hal-01557204 https://hal.archives-ouvertes.fr/hal-01557204 Submitted on 5 Jul 2017 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Copyright Annals of Forest Science (2016) 73:135–146 DOI 10.1007/s13595-015-0468-7 REVIEW PAPER Can mosses serve as model organisms for forest research? Stefanie J. Müller & Desirée D. Gütle & Jean-Pierre Jacquot & Ralf Reski Received: 30 October 2014 /Accepted: 23 February 2015 /Published online: 17 March 2015 # INRA and Springer-Verlag France 2015 Abstract biology techniques and was designated as a JGI plant flagship & Key message Based on their impact on many ecosystems, genome. In this review, we will provide an introduction to we review the relevance of mosses in research regarding moss research and highlight the characteristics of P. patens stress tolerance, metabolism, and cell biology. We intro- and other mosses as a potential complementary model system ducethepotentialuseofmossesascomplementarymodel for forest research. systems in molecular forest research, with an emphasis on & Methods Starting with an introduction into general moss the most developed model moss Physcomitrella patens. biology, we summarize the knowledge about moss physiology & Context and aims Mosses are important components of sev- and differences to seed plants. We provide an overview of the eral ecosystems. The moss P. patens is a well-established non- current research areas utilizing mosses, pinpointing potential vascular model plant with a high amenability to molecular links to tree biology. To complement literature review, we discuss moss advantages and available resources regarding Handling Editor: Jean-Michel Leban molecular biology techniques. & Contribution of the co-authors S.J.M. and D.D.G. wrote the Results and conclusion During the last decade, many fun- manuscript under the supervision of J.P.J. and R.R. damental processes and cell mechanisms have been studied in S. J. Müller : D. D. Gütle : R. Reski (*) mosses and seed plants, increasing our knowledge of plant Plant Biotechnology, Faculty of Biology, University of Freiburg, evolution. Additionally, moss-specific mechanisms of stress Schaenzlestr.1, 79104 Freiburg, Germany tolerance are under investigation to understand their resilience e-mail: [email protected] in ecosystems. Thus, using the advantages of model mosses D. D. Gütle : R. Reski such as P. patens is of high interest for various research ap- Spemann Graduate School of Biology and Medicine (SGBM), proaches, including stress tolerance, organelle biology, cell University of Freiburg, Albertstr. 19A, 79104 Freiburg, Germany polarity, and secondary metabolism. D. D. Gütle : J.<P. Jacquot Interactions Arbres—Microorganismes, UMR1136, Université de Lorraine, 54500 Vandoeuvre-lès-Nancy, France Keywords Physcomitrella patens . Model plant . Gene targeting . Abiotic stress . Evo-devo . Cell biology J.<P. Jacquot Interactions Arbres—Microorganismes, UMR1136, INRA, 54280 Champenoux, France 1 Introduction R. Reski BIOSS Centre for Biological Signalling Studies, Schaenzlestr. 18, Often, the ability to disproof or to support a scientific hypoth- 79104 Freiburg, Germany esis in an experiment depends on the choice of the biological R. Reski model system. In life sciences, several model species have FRIAS Freiburg Institute for Advanced Studies, Albertstr. 19, been used in relation with the different advantages they offer 79104 Freiburg, Germany in terms of generation time, growth, evolutionary position, or R. Reski amenability to molecular biology techniques (Jansson and USIAS Strasbourg Institute for Advanced Study, Strasbourg, France Douglas 2007; Müller and Grossniklaus 2010; Reski 1998). 136 S.J. Müller et al. In plant sciences, several seed plant model species are regarding ecology as well as regarding fundamental processes established, whereas lycophytes and ferns are more problem- and evolution of land plants. Moss model species include, atic model organisms due to their low transformability or large e.g., the Funariaceae Funaria hygrometrica and genome size (Plackett et al. 2014). Liverworts, mosses, and Physcomitrella patens, the Ditrichacea Ceratodon purpureus, hornworts constitute the paraphyletic group of bryophytes and and peat moss (Sphagnum) species. Moreover, some model represent early diverging lineages of land plants with an inter- mosses provide extensive advantages for research projects mediate position between vascular plants and algae (Kenrick employing molecular biology. The currently most developed and Crane 1997;Langetal.2010). Many moss features are model moss is P. patens, a moss of the temperate zones which regarded as ancestral, though their special life style has led to canbefoundonsoilexposedbyfallingwaterlevelsoronfields the evolution of many moss-specific morphological and phys- (Cove 2005). It provides a fully sequenced genome (Rensing iological traits, which support alternative survival strategies et al. 2008), many resources for -omics techniques, a high ame- among land plants (Mishler and Oliver 2009). While the sim- nability to microscopy, and a valuable experimental platform for ple morphological structure of mosses is largely conserved comparative studies with other model organisms, as, e.g., trees. since at least 330 million years (Hubers and Kerp 2012), they Therefore, the Physcomitrella genome was designated, like pop- colonize an extensive spectrum of diverse habitats (Turetsky lar, as a “JGI plant flagship genome” (http://jgi.doe.gov/our- 2003; Turetsky et al. 2012). Their occurrence in many extreme science/science-programs/plant-genomics/plant-flagship- habitats such as the Antarctic tundra and deserts implies the genomes/). Potential research areas include general moss existence of many yet unknown moss-specific survival mech- physiology, moss-specific stress resistance mechanisms, puta- anisms (Oliver et al. 2005; Roads et al. 2014). The resilience tively interesting for the transfer to seed plants, as well as research of mosses could recently be demonstrated with the successful on fundamental mechanisms which evolved early and are con- regeneration of subglacial bryophytes following 400 years of served in many land plants. In addition, the technical advantages ice entombment (Farge et al. 2013) and even by the regrowth of the moss system can offer complementing approaches to many of over 1500-year-old moss from the Antarctic permafrost open questions in seed plant research by, e.g., enabling the ma- (Roads et al. 2014). nipulation of the genome down to the single base pair level by However, besides these extreme examples, mosses are also homologous recombination. important components of tropical systems (Gradstein et al. This review gives a basic introduction into moss biology 2001), boreal forests, and woodlands or unshaded habitats in and the current reference model Physcomitrella and its advan- temperate zones (Turetsky 2003). Despite their small size, tages, and provides an overview of the current use of mosses mosses have a huge impact on various ecosystems and are to gain knowledge in different research areas, with an empha- essential contributors to complex biological cycles. Besides sis on potential links with molecular tree research. the prevention of soil erosion, the development of microtopography, and the regulation of soil climate and water availability, main aspects are nitrogen fixation and carbon 2 The moss model: particularities and advantages cycling. For instance, mosses are important contributors to bog for- Plants started to colonize the land mass of Earth about 450– mation and carbon trapping (Jones et al. 1994; Shaw et al. 500 million years ago (Kenrick and Crane 1997;Langetal. 2003) in peat lands, where almost 100 % of the ground layer 2010), shaping the new environment and establishing the ba- is covered by mosses (Vitt and Wieder 2009)suchas sis of land ecosystems, paving the way for the colonization by Sphagnum species. The relevance of biotechnology ap- animals. During land plant evolution, many adaptations oc- proaches regarding the production and cultivation of peat curred, some of them leading to large radiation of new groups, mosses is increasing due to the rising demand of peat as, as, e.g., the flowering plants. Ancestral features of land plants e.g., substrate for horticulture (Beike et al. 2015a). can be either reconstructed from fossils or by the investigation In forest ecosystems such as rain forests, in which mosses of early diverging land plant groups, as mosses. Though for live in immediate vicinity to vascular plants, they can serve as some aspects moss biology is similar, in others