Impacts of long-term enhanced UV-B radiation on bryophytes in two sub-Arctic heathland sites of contrasting water availability

Abstract

Background and aims: 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.

Fig. 1.

(A) Percentage cover, (B) new growth, (C) sclerophylly index (SI; dry matter per surface area), (D) maximum quantum yield of PSII (F_v/F_m), (E) maximal non-photochemical quenching (NPQ_max) and (F) methanol-extractable UV-absorbing compounds (MEUVACs) of the three bryophytes (the mosses Hylocomium splendens and Polytrichum commune, and the foliose liverwort Barbilophozia lycopodioides), studied in plots exposed to two radiation regimes (control or enhanced UV-B, as indicated) under two different water regimes (M, mesic plots; D, drier plots). Means ± s.e. are shown (n = 4). The main factor effects (two-way ANOVA) of water regime (WR) and UV-B radiation (UV), and the interactions between both (WR×UV), are indicated for each species when significant (***P < 0·001; **P < 0·01; *P < 0·05). Differences between control and +UV-B plots for each species and water regime separately are shown over the corresponding bars (Student́s t-test adjusted with Bonferroni; °P <0·025; °°P < 0·005).

Fig. 2.

Curves of the electron transport rate through PSII (ETR) against photosynthetic photon flux density (PPFD); species and treatment details as for Fig. 1. The s.e. are not shown for clarity. The overall effect (two-way ANOVA) of the water regime (WR) is indicated when significant (*** P <0·001; **P < 0·01; *P < 0·05). The effect of the radiation regime (UV, UV-B effect) was not significant in any species.