Yngvar Gauslaa

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.

Lichen palatability depends on investments in herbivore defence

Lichens are well-suited organisms for experimental herbivory studies because their secondary compounds, assumed to deter grazing, can be non-destructively extracted. Thalli of 17 lichen species from various habitats were cut in two equal parts; compounds were extracted from one part by acetone, the other served as a control. These two pieces were offered as a paired choice to the generalist herbivore snail Cepaea hortensis. Control thalli of all lichens were consumed at a low rate regardless of their investments in acetone-extractable lichen compounds; naturally compound-deficient lichen species were not preferred compared to those with high contents. However, for extracted thalli, there was a highly significant positive correlation between rate of consumption and the extracted compound contents. These data imply that herbivore defence has evolved in different directions in different lichens. Studied members of Parmeliaceae, common in oligotrophic habitats, have high contents of carbon-rich acetone-soluble compounds; these lichens became highly palatable to snails subsequent to acetone rinsing. Extracted lichen compounds were applied to pieces of filter paper and fed to snails. Extracts from members of the Parmeliaceae significantly deterred feeding on paper. Such data suggest that generalist herbivores may have shaped evolution in the widespread and highly diverse Parmeliaceae towards high investments in lichen compounds. On the other hand, lichens belonging to the Physciaceae and Teloschistales, common in nutrient-enriched habitats, are deficient in, or have low concentrations of, lichen compounds. Such lichens did not become more palatable after acetone rinsing. The orange anthraquinone compound parietin, restricted to the Teloschistales, and which has previously been found to protect against excess light, did not deter grazing.

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.

Acidity of boreal Picea abies-canopy lichens and their substratum, modified by local soils and airborne acidic depositions

Summary Lichens of the rare and predominantly epiphytic old forest community Lobarion were attached to Picea abies twigs with significantly higher bark pH (KCl) than members of the common and ubiquitous Pseudevernion community. The Lobarion species of spruce twigs, mainly members of the order Peltigerales, had distincly higher thallus pH than substratum pH, while the greenalgal Pseudevernion species, mainly belonging to the Lecanorales, were consistently more acidic than their substratum. pH of lichens and bark of P. abies twigs responded to a forest vegetation gradient reflecting the soil nutrient condition at the forest floor. However, the two groups of epiphytes seem to modify the bark pH in a way that enlarges a difference originally determined by soils in the root zone of the phorophyte. The lack of the Lobarion on spruce twigs in eastern Norway is probably a result of acid rain, as the bark and thallus pH in spruce canopies of eastern Norway appeared too low to support the Lobarion, even in stands with a species-rich epiphytic assemblage of alectorioid species.

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 .

Mollusc grazing limits growth and early development of the old forest lichen Lobaria pulmonaria in broadleaved deciduous forests

This study aims: (1) to quantify mollusc grazing on juvenile and mature thalli of the foliose epiphytic lichen Lobaria pulmonaria, and (2) to test the hypothesis inferring a herbivore defensive role of lichen depsidones in forests with indigenous populations of lichen-feeding molluscs. Lichens were transplanted in shaded and less shaded positions in each of two calcareous broadleaved deciduous forests, one poor in lichens, one with a rich Lobarion community. Preventing the access of molluscs significantly reduced the loss of juvenile L. pulmonaria, particularly in the naturally lichen-poor forest. Molluscs also severely grazed mature thalli in the lichen-poor forest, especially thalli placed under the more shading canopies. Furthermore, reducing the natural concentration of depsidones by pre-rinsing with acetone increased subsequent grazing significantly, showing that lichen depsidones function as herbivore defence in natural habitats. Our results suggest that mollusc grazing may play important roles in shaping the epiphytic vegetation in calcareous deciduous forests, and that recently established juvenile L. pulmonaria thalli seem to be particularly vulnerable.

Rain, dew, and humid air as drivers of morphology, function and spatial distribution in epiphytic lichens

This review is a first attempt to combine and compare spatial distribution of the three main water sources, rain, dew and humid air, with water-related traits of mainly epiphytic macrolichens in a conceptual and functional model. By comparing climatic and lichenological knowledge, various effects of dewfall, rainfall and humid air on epiphytic lichen morphology and function are analyzed to search for traits and patterns. Although dew, rain and humid air cause lichen hydration and activate photosynthesis, these atmospheric hydration sources influence and shape lichens differently. In order to visualize hydration patterns, dew, rain and humid air are shown as corners in a triangle exhibiting the various combinations of these hydration sources. The sources of hydration vary on temporal scales, and on the spatial scales: regional, landscape, stand and tree. Lichen growth form, photobiont type, water-holding capacity (WHC) and suprasaturation depression show clear patterns within the hydration triangle. For most lichen species, one average pre-dawn dewfall approximately fills their average internal WHC. This suggests that lichens are optimally designed to utilize dew rather than rain, consistent with literature emphasizing dew as a driver for annual C-assimilation in chlorolichens. However, rain is needed to fill their external WHC and to fully hydrate most cyanolichens. Including the sources of hydration and internal lichen variables, such as water-holding capacity, will improve modelling of local and global future scenarios on lichen distribution and biomass production.

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.