title: Impacts of long-term enhanced UV-B radiation on bryophytes in two sub-Arctic heathland sites of contrasting water availability.
abstract: OBJECTIVE
  Anthropogenic depletion of stratospheric ozone in Arctic latitudes has resulted in an increase of ultraviolet-B radiation (UV-B) reaching the biosphere. UV-B exposure is known to reduce above-ground biomass and plant height, to increase DNA damage and cause accumulation of UV-absorbing compounds in polar plants. However, many studies on Arctic mosses tended to be inconclusive. The importance of different water availability in influencing UV-B impacts on lower plants in the Arctic has been poorly explored and might partially explain the observed wide variation of responses, given the importance of water in controlling bryophyte physiology. This study aimed to assess the long-term responses of three common sub-Arctic bryophytes to enhanced UV-B radiation (+UV-B) and to elucidate the influence of water supply on those responses.
  
  
  METHODS
  Responses of three sub-Arctic bryophytes (the mosses Hylocomium splendens and Polytrichum commune and the liverwort Barbilophozia lycopodioides) to +UV-B for 15 and 13 years were studied in two field experiments using lamps for UV-B enhancement with identical design and located in neighbouring areas with contrasting water availability (naturally mesic and drier sites). Responses evaluated included bryophyte abundance, growth, sporophyte production and sclerophylly; cellular protection by accumulation of UV-absorbing compounds, β-carotene, xanthophylls and development of non-photochemical quenching (NPQ); and impacts on photosynthesis performance by maximum quantum yield (F(v) /F(m)) and electron transport rate (ETR) through photosystem II (PSII) and chlorophyll concentrations.
  
  
  RESULTS
  Responses were species specific: H. splendens responded most to +UV-B, with reduction in both annual growth (-22 %) and sporophyte production (-44 %), together with increased β-carotene, violaxanthin, total chlorophyll and NPQ, and decreased zeaxanthin and de-epoxidation of the xanthophyll cycle pool (DES). Barbilophozia lycopodioides responded less to +UV-B, showing increased β-carotene and sclerophylly and decreased UV-absorbing compounds. Polytrichum commune only showed small morphogenetic changes. No effect of UV-B on bryophyte cover was observed. Water availability had profound effects on bryophyte ecophysiology, and plants showed, in general, lower growth and ETR, together with a higher photoprotection in the drier site. Water availability also influenced bryophyte responses to +UV-B and, in particular, responses were less detectable in the drier site.
  
  
  CONCLUSIONS
  Impacts of UV-B exposure on Arctic bryophytes were significant, in contrast to modest or absent UV-B effects measured in previous studies. The impacts were more easily detectable in species with high plasticity such as H. splendens and less obvious, or more subtle, under drier conditions. Species biology and water supply greatly influences the impact of UV-B on at least some Arctic bryophytes and could contribute to the wide variation of responses observed previously.
authors.affiliation: Department of Animal and Plant Sciences, University of Sheffield, Western Bank, Sheffield S10 2TN, UK. maria.arroniz@gmail.com
authors.forename: M
authors.initials: M
authors.lastname: Arróniz-Crespo
authors.forename: D
authors.initials: D
authors.lastname: Gwynn-Jones
authors.forename: T V
authors.initials: TV
authors.lastname: Callaghan
authors.forename: E
authors.initials: E
authors.lastname: Núñez-Olivera
authors.forename: J
authors.initials: J
authors.lastname: Martínez-Abaigar
authors.forename: P
authors.initials: P
authors.lastname: Horton
authors.forename: G K
authors.initials: GK
authors.lastname: Phoenix
mesh_terms: D044002:Bryophyta / Q000737:chemistry / Q000502:physiology / Q000528:radiation effects*; D002338:Carotenoids / Q000378:metabolism; D002734:Chlorophyll / Q000378:metabolism; D004579:Electron Transport / Q000528:radiation effects; D010788:Photosynthesis / Q000528:radiation effects; D045332:Photosystem II Protein Complex / Q000528:radiation effects; D012098:Reproduction / Q000528:radiation effects; D014466:Ultraviolet Rays; D014867:Water / Q000502:physiology*
chemical_list: D045332:Photosystem II Protein Complex; D014867:Water; D002734:Chlorophyll; D002338:Carotenoids