Knut Asbjørn Solhaug

Parietin, a photoprotective secondary product of the lichen Xanthoria parietina

Secondary lichen products can be extracted from air-dry thalli of Xanthoria parietina, Xanthoparmelia conspersa and Parmelina tiliacea with 100% acetone without affecting either short-or long-term viability. In Xanthoria parientina damage by acetone started to occur as water content reached the critical lower limit for photosystem II (PSII) activity. Extraction of the blue-light absorbing cortical pigment parietin increased the susceptibility of both air-dry and hydrated thalli to high light. Damage by high light levels caused a permanent reduction in Fv/Fm, quantum yield for photosynthetic O2 production and photosynthetic capacity measured after a 2-day recovery period at low light levels (20 μmol photons m-2 s-1). Parietin therefore protects the photosynthetic apparatus of Xanthoria parietina against damage by high light levels. Extraction of UV-absorbing pigments from Xanthoparmelia conspersa and Parmelina tiliacea did not increase photoinhibition after 24 h exposure to high light.

Growth of epiphytic old forest lichens across climatic and successional gradients

This paper aims to assess the influence of canopy cover on lichen growth in boreal forests along a regional forest gradient. Biomass and area gain, and some acclimation traits, were assessed in the ...

The significance of thallus size for the water economy of the cyanobacterial old-forest lichen Degelia plumbea

Abstract Rosette-formed, circular thalli of Degelia plumbea were studied in the laboratory. Regardless of thallus size, the optimal quantum yield of photosystem II (FV/FM) remained at a high, constant level during a drying cycle starting with fully hydrated thalli until the thallus water content reached about 200%. Net photosynthesis reached a maximum level at this hydration level. Thereafter, both FV/FM and net photosynthesis fell rapidly to zero at a water content of somewhat less than 100%. There was a highly significant, positive relationship between thallus size and the water-holding capacity, as well as a strong, negative correlation between size and water loss per thallus area. Consequently, an increase in thallus size from 1 to 36 cm2 lead to a tenfold prolongation of the photosynthetically active period during a drying cycle at a low radiation regime. The improved water-holding capacity in larger thalli is mainly a result of a thicker hypothallus. The fast desiccation of small thalli suggests that the regeneration of D. plumbea could be severely hampered by nearby logging that raises the evaporative demand by increasing radiation loads and wind exposure at remaining lichen sites.

UV triggers the synthesis of the widely distributed secondary lichen compound usnic acid

Synthesis of the cortical dibenzofuran derivative usnic acid and the medullary depsidone salazinic acid was studied in Xanthoparmelia stenophylla thalli from which the compounds had been removed by acetone rinsing prior to a 21-day field experiment with UV absorbing and transmitting screens. Natural levels of ultraviolet radiation clearly induced the re-synthesis of usnic acid. The re-synthesis was boosted by the addition of ribitol, the carbohydrate delivered from the Trebouxia photobiont to the mycobiont. Salazinic acid was also weakly induced by UV. Re-synthesis was relatively low, up to 2.5 and 3.1% of start values for usnic and salazinic acid, respectively. However, given that the natural content of both compounds was high, constituting 12% of thallus dry weight, the absolute amounts of lichen compounds re-synthesised were not so small. We also studied the extractability of nine extracellular lichen compounds in three species X. stenophylla, Hypogymnia tubulosa, and Vulpicida pinastri, and found two distinct fractions of cortical compounds, one major that was completely extractable from living lichens and one minor that was only extractable with grinding. Medullary compounds were completely extracted without grinding. These findings did not influence the relative differences between treatments in our experiment, but may be of importance for future assessments of, e.g., quantitative studies of extracellular lichen compounds.

Photoinhibition in lichens depends on cortical characteristics and hydration

Two shade-adapted ( Lobaria pulmonaria and Cetraria islandica ) and two sun-adapted lichen populations ( Xanthoria parietina and Cetraria nivalis ) were exposed to three irradiance regimes (1: photosynthetic radiation—PAR, 2: PAR+UV-A, 3: PAR+UV-A+UV-B) and two hydration regimes (1: no hydration, 2: daily hydration) in a growth cabinet for three weeks. Shade-adapted thalli had transparent upper cortices without coloured pigments, whereas sun-adapted thalli had coloured UV-B absorbing cortical pigments masking the photobiont. Manipulation of pigment concentration was the third factor used in the factorial design (1: pigments intact, 2: pigments non-destructively extracted from air-dry living thalli by acetone). Inhibition of the photobiont due to PAR alone was severe in the two shade-adapted populations, but no applied UV wavelength bands caused additional aggravation of photoinhibition. Shade-adapted thalli of the ubiquitous C. islandica were more PAR-susceptible than of the rare old forest lichen L. pulmonaria , suggesting that screening by the mycobiont rather than photobiont characteristics, account for their different success in sun-exposed localities. Hydration of shade-adapted species during exposure reduced their photoinhibition substantially, probably because of moisture-activated repair mechanisms. On the contrary, the sun-adapted X. parietina was most phototolerant in the desiccated state, whereas hydration caused increased photoinhibition. When removing the orange cortical pigment parietin, the photoinhibition in moist thalli was aggravated, confirming a PAR-protective function of parietin. No effects of irradiance treatment, pigment extraction (usnic acid), or hydration level were observed in C. nivalis .

The paradox of higher light tolerance during desiccation in rare old forest cyanolichens than in more widespread co‐occurring chloro‐ and cephalolichens

Desiccation tolerance was quantified in four cyanolichens (Lobaria hallii, Lobaria retigera, Lobaria scrobiculata, Pseudocyphellaria anomala), one cephalolichen (Lobaria pulmonaria) and one chlorolichen (Platismatia glauca) from xeric and mesic, open and closed North American boreal forests. These sympatric epiphytes were exposed to 0%, 33%, 55% and 75% relative humidity with or without medium light (200 μmol m−2 s−1) for 7 d. Permanent and temporary photoinhibitory damage was recorded as viability measures. All species tolerated well the drying in darkness, but L. hallii and L. retigera, associated with a very humid climate, showed minor damage at the hardest drying (silica gel). Simultaneous exposure to medium light severely aggravated the drying damage at all relative humidity levels. Combined drying–light exposure was particularly devastating for the widespread chloro- and cephalolichens, whereas cyanolichens, including rare old forest species, were fairly resistant. The ability to recover after combined drying–light stress (this study) correlated positively with increasing species-specific water holding capacities (from the literature). Cyanolichens, depending on liquid water and large internal water storage, probably require strong drying–light resistance to handle long periods between hydration events, whereas chlorolichens can regularly maintain their photosynthetic apparatus during frequent and rapid activation by humid air on clear mornings.

Forest successional stage affects the cortical secondary chemistry of three old forest lichens.

Three epiphytic old forest lichens (Usnea longissima, Pseudocyphellaria crocata, and Lobaria pulmonaria) were transplanted along a natural shade–sun gradient comprising three successional stages in boreal spruce forests (dense young forest, open old forest, and clear-cut) for one summer. After harvest, extractable secondary compounds were analyzed by high-performance liquid chromatography, and the brown pigmentation in melanic species was quantified by reflectance measurements. Cortical compounds in all species increased from shady young forests to exposed clear-cuts. Usnic acid, the major cortical, secondary compound in U. longissima, showed consistently higher concentration in the clear-cut than in the two forested stands. Pseudocyphellaria crocata and L. pulmonaria, lacking extractable secondary compounds in the cortex, significantly increased their amounts of cortical melanins in well-lit stands. The medullary compounds showed more complex responses. Many were not influenced by environmental conditions during the transplantation, whereas the majority of those that responded showed the lowest concentration in clear-cut transplants. Only a few medullary compounds showed the highest concentration in the clear-cut, and at a low level of significance. The synthesis of UV-B-absorbing usnic acid and melanins seems to be part of an acclimation to increased light exposure. The medullary compounds in studied species barely function as solar screens despite their strong UV-B absorbance.

Light screening in lichen cortices can be quantified by chlorophyll fluorescence techniques for both reflecting and absorbing pigments

Lichens, representing mutualistic symbioses between photobionts and mycobionts, often accumulate high concentrations of secondary compounds synthesized by the fungal partner. Light screening is one function for cortical compounds being deposited as crystals outside fungal hyphae. These compounds can non-destructively be extracted by 100% acetone from air-dry living thalli. Extraction of atranorin from Physcia aipolia changed the lichen colour from pale grey to green in the hydrated state, whereas acetone-rinsed and control thalli were all pale grey when dry. Removal of parietin from Xanthoria parietina changed the colour of desiccated thalli from orange to grey. Colour changes were quantified by reflectance measurements. By a new chlorophyll fluorescence method, screening was assessed as the decrease in incident irradiance (PAR) necessary to reach identical effective quantum yields of PSII (ΦPSII) in acetone-rinsed and control thalli. Thereby, we estimated a screening efficiency due to cortical atranorin crystals at 61, 38, and 40% of blue, green and red light, respectively, whereas parietin screened 81, 27 and 1% of these wavelength ranges. Removal of atranorin caused similar levels of increased photoinhibition for P. aipolia in blue, green and red light, whereas parietin-deficient thalli of X. parietina exhibited increased photoinhibition with decreasing wavelengths. Atranorin possibly prevents water from entering the spaces between the hyphae in the cortex. The air-filled cavities with white atranorin crystals reflect excess light, whereas the yellow compound parietin absorbs excess light. Thereby, both atranorin and parietin play significant photoprotective roles for symbiotic green algae, but with compound-specific screening mechanisms.

Secondary Lichen Compounds as Protection Against Excess Solar Radiation and Herbivores

The functional roles of secondary lichen compounds are reviewed with focus on sun-screening and herbivore-deterring functions. Hypotheses on ecological functions can be tested because lichen compounds can nondestructively be extracted from air-dry lichens with 100% acetone. Substantial evidence supports a sun-screening function of cortical compounds. They screen solar radiation by absorptance (parietin, melanins) or by reflectance (atranorin). Their concentration correlates with light exposure and they protect the photobiont against excessive visible light. UV-B induces the formation of parietin, usnic acid, and melanins; the synthesis of the two first compounds has been shown to be boosted by photosynthates. The numerous extractable medullary lichen compounds hardly function as sun-screens. Some of these carbon-based compounds deter generalist herbivores, particularly in lichens in oligotrophic sites. Lichens in nitrogen-rich sites often deter grazing animals as efficient as those from oligotrophic sites despite low contents of lichen compounds. Acetone rinsing of nitrophytic lichens does not lead to increased grazing, meaning that their defense remains to be described. Thanks to grazing experiments using acetone rinsing, there is now solid support for the optimal defense theory in lichen–herbivore interactions. Recent studies have shown that lichen-feeding gastropods can shape epiphytic lichen communities.

Size-dependent growth of two old-growth associated macrolichen species.

Relationships between thallus size and growth variables were analysed for the foliose Lobaria pulmonaria and the pendulous Usnea longissima with the aim of elucidating their morphogenesis and the factors determining thallus area (A) versus biomass (dry weight (DW) gain. Size and growth data originated from a factorial transplantation experiment that included three boreal climate zones (Atlantic, suboceanic and continental), each with three successional forest stands (clear-cut, young and old). When A was replaced by the estimated photobiont layer area in an area-DW scatterplot including all thalli (n = 1080), the two separate species clusters merged into one, suggesting similar allocation patterns between photobionts and mycobionts across growth forms. During transplantation, stand-specific water availability boosted area gain in foliose transplants, consistent with a positive role of water in fungal expansion. In pendulous lichens, A gain greatly exceeded DW gain, particularly in small transplants. The A gain in U. longissima increased with increasing DW:A ratio, consistent with a reallocation of carbon, presumably mobilized from the dense central chord. Pendulous lichens with cylindrical photobiont layers harvest light from all sides. Rapid and flexible three-dimensional A gain allows the colonization of spaces between canopy branches to utilize temporary windows of light in a growing canopy. Foliose lichens with a two-dimensional photobiont layer have more coupled A and DW gains.