Burkhard Schroeter

Monitoring photosynthetic activity of crustose lichens using a PAM-2000 fluorescence system

SummaryCO2 exchange and fluorescence yield of the crustose lichen Buellia frigida were measured in situ by means of a CO2 porometer and a PAM-2000, a newly developed portable fluorescence system. The pulse amplitude modulation system of the PAM-2000 allows measurements in the field under ambient light, temperature and moisture conditions without dark adaptation of the sample. CO2 exchange and fluorescence measurements were well correlated when measured under natural conditions in continental Antarctica during a drying cycle of melt-water-soaked lichen thalli. It was shown that the fluorescence parameter ΔF/Fm′ is a measure of the photosynthetic activity of the lichen. It proved possible, using the PAM-2000, to differentiate the physiological performance of the thallus centre and the marginal lobes. The distribution of water in the thallus during a drying cycle was shown to be inhomogeneous. The photosynthetic rates of B. frigida calculated on an area basis are comparatively high and indicate that this lichen is well adapted to its habitat conditions in this part of continental Antarctica.

Are lichens active under snow in continental Antarctica

Photosynthetic activity, detected as chlorophyll a fluorescence, was measured for lichens under undisturbed snow in continental Antarctica using fibre optics. The fibre optics had been buried by winter snowfall after being put in place the previous year under snow-free conditions. The fibre optics were fixed in place using specially designed holding devices so that the fibre ends were in close proximity to selected lichens. Several temperature and PPFD (photosynthetic photon flux density) sensors were also installed in or close to the lichens. By attaching a chlorophyll a fluorometer to the previously placed fibre optics it proved possible to measure in vivo potential photosynthetic activity of continental Antarctic lichens under undisturbed snow. The snow cover proved to be a very good insulator for the mosses and lichens but, in contrast to the situation reported for the maritime Antarctic, it retained the severe cold of the winter and prevented early warming. Therefore, the lichens and mosses under snow were kept inactive at subzero temperatures for a prolonged time, even though the external ambient air temperatures would have allowed metabolic activity. The results suggest that the major activity period of the lichens was at the time of final disappearance of the snow and lasted about 10–14 days. The activation of lichens under snow by high air humidity appeared to be very variable and species specific. Xanthoria mawsonii was activated at temperatures below −10°C through absorption of water from high air humidity. Physcia dubia showed some activation at temperatures around –5°C but only became fully activated at thallus temperatures of 0°C through liquid water. Candelariella flava stayed inactive until thallus temperatures close to zero indicated that liquid water had become available. Although the snow cover represented the major water supply for the lichens, lichens only became active for a brief time at or close to the time the snow disappeared. The snow did not provide a protected environment, as reported for alpine habitats, but appeared to limit lichen activity. This provides at least one explanation for the observed negative effect of extended snow cover on lichen growth.

Structural and functional processes during water vapour uptake and desiccation in selected lichens with green algal photobionts

Structural alterations of the photobiont and mycobiont cells of lichens have been related to CO2-gas exchange during experiments involving water vapour uptake and desiccation of liquid-water-saturated thalli. Increasing water vapour uptake of air dry lichens led to a gradual unfolding of the photobiont cells in Lobaria pulmonaria, Pseudevernia furfuracea, Ramalina maciformis and Teloschistes lacunosus as studied by low-temperature scanning electron microscopy. The data indicated that globular, probably turgid, cells and also slightly infolded or even heavily collapsed cells contributed to positive net photosynthesis, which was reached after water vapour uptake by the four species studied. During desiccation of fully water-saturated thalli of L. pulmonaria, extrathalline water films gradually evaporated before maximum values of CO2-gas exchange were measured and before photobiont cells started to shrivel. In contrast, in P. furfuracea the CO2-gas exchange maximum was reached when a considerable percentage of photobiont cells had already collapsed and while other parts of the thalli were still covered with liquid water. Further desiccation led to cavitation of the cortical cells in both species, this occurring at water contents at which net photosynthesis was still positive.

Chlorophyll a fluorescence and CO2 exchange of Umbilicaria aprina under extreme light stress in the cold

Abstract A lichen growing in a continental Antarctic region with low temperatures and strong irradiance in summer was investigated for evidence of photoinhibition. Field experiments with Umbilicaria aprina from a sheltered site with heavy snowpack showed no effects of photoinhibition when the lichen was exposed to strong sun irradiance for nearly 11u2009h a day. This was evident from CO2 exchange and simultaneous chlorophyll a fluorescence measurements. CO2 exchange was also not affected if quartz glass allowing greater UV penetration, was used as a lid for the cuvette. The dependency of net photosynthesis on photosynthetic photon flux density suggests that the lichen is photophilous.

An assessment of the relationship between chlorophyll a fluorescence and CO2 gas exchange from field measurements on a moss and lichen

Abstract. The relationship between CO2 exchange and relative electron-transport rate through photosystem II (ETR, measured using chlorophyll a fluorescence) was determined for a moss and a green algal lichen, photobiont probably Trebouxia sp., in the field in Antarctica. Net photosynthesis (NP) and dark respiration (DR) were measured over temperatures from zero to 25u2009°C and gross photosynthesis (GP) calculated (GPu2009=u2009NP + DR). The strong response of DR to temperature in these organisms resulted in substantial changes in CO2 exchange rates. The moss Bryum argenteum Hedw. showed a strong, linear relationship between GP and ETR. This was an unexpected result since mosses are C3 plants and, in higher plants, this group normally has a curvilinear GP versus ETR relationship. It is suggested that suppression of DR in the light might be involved. The lichen, Umbilicaria aprina Nyl., had nonlinear relationships between ETR and GP that were different at each measurement temperature. In some cases the lowest ETR was at the higher CO2 exchange rates. It is suggested that these relationships are the result of strong quenching mechanisms that are inversely proportional to GP. The results support a growing impression that the relationships between ETR and CO2 exchange are complex in these organisms and different from those found for higher plants.

rDNA ITS and β-tubulin gene sequence analyses reveal two monophyletic groups within the cosmopolitan lichen Parmelia saxatilis

A considerable number of species of lichen-forming fungi have wide geographical distributions, but studies of their genetic variability are minimal. ITS rDNA sequences of 32 populations of Parmelia saxatilis from five continents revealed two monophyletic groups. β-tublin gene sequences from a subset of nine collections supported these conclusions. While the number of collections sequenced is limited, one monophyletic group (the Atlantic Population. AtP) was recognized as occurring in Arctic and Antarctic regions and also included collections from more atlantic sites. Samples from more mesic environments in the Mediterranean region belonged to a second monophyletic group (the Mediterranean Population, MeP). In addition, four subgroups were distinguishable within the Atlantic Population. Norstictic and protocetraric acids are reported from the species for the first time, the norstictic acid only being found in the Atlantic Population. Living thalli from the Atlantic Population were provenance-tested; specimens transported from the UK to central Spain where the Mediterranean Population occurs showed adverse symptoms after six months. These results demonstrate that there can be substantial large-scale genotypic variability within widespread lichen phenospecies, something which has implications for comparative ecological, physiological, and air pollution sensitivity studies as well as for lichen conservation.

Ecophysiological investigations of Usnea antarctica in the maritime Antarctic I. Annual microclimatic conditions and potential primary production

Photosynthetic photon flux density (PPFD), air relative humidity and thallus temperature (TT) were measured in Usnea antarctica over a period of 12 months at a coastal rock on King George Island, South Shetland Islands. TT had an annual amplitude of c. 54 K with the maximum of +27.4°C recorded in December and minimum of −27.3°C in July. Daily maximum TT exceeded 0°C every month except in June and July when they were below −0.4°C and −1.6°C, respectively. Daily minimum temperatures were always below zero. Diel courses of PPFD showed pronounced seasonal differences between summer (more than 2000 μmol m −2 s −1 ) and winter (less than 50 μmol m −2 s −1 in July). Daily sum of PPFD was highest (more than 30 mol m −2 d −1 ) in December and lowest (0.1 mol m −2 d −1 ) in July. A photosynthesis model was used to estimate the potential annual primary production from habitat PPFD and TT. The estimated potential annual carbon balance of U. antarctica was 323 mg CO 2 g −1 dry weight y −1 assuming that the lichen was always at optimal moisture. The potential carbon balance indicates that primary production is severely limited by low PPFD and subzero TT in the austral winter. Nevertheless, the PPFD and TT would allow metabolic activity during winter but with a negative carbon balance during May–August. Spring and autumnal months are revealed as probably the most important periods for lichen primary production because PPFD exceeds photosynthetic light compensation point during the daytime and ambient moisture conditions frequently favour a positive carbon balance.

Improved appreciation of the functioning and importance of biological soil crusts in Europe: the Soil Crust International Project (SCIN)

Here we report details of the European research initiative “Soil Crust International” (SCIN) focusing on the biodiversity of biological soil crusts (BSC, composed of bacteria, algae, lichens, and bryophytes) and on functional aspects in their specific environment. Known as the so-called “colored soil lichen community” (Bunte Erdflechtengesellschaft), these BSCs occur all over Europe, extending into subtropical and arid regions. Our goal is to study the uniqueness of these BSCs on the regional scale and investigate how this community can cope with large macroclimatic differences. One of the major aims of this project is to develop biodiversity conservation and sustainable management strategies for European BSCs. To achieve this, we established a latitudinal transect from the Great Alvar of Öland, Sweden in the north over Gössenheim, Central Germany and Hochtor in the Hohe Tauern National Park, Austria down to the badlands of Tabernas, Spain in the south. The transect stretches over 20° latitude and 2,300xa0m in altitude, including natural (Hochtor, Tabernas) and semi-natural sites that require maintenance such as by grazing activities (Öland, Gössenheim). At all four sites BSC coverage exceeded 30xa0% of the referring landscape, with the alpine site (Hochtor) reaching the highest cyanobacterial cover and the two semi-natural sites (Öland, Gössenheim) the highest bryophyte cover. Although BSCs of the four European sites share a common set of bacteria, algae (including cyanobacteria) lichens and bryophytes, first results indicate not only climate specific additions of species, but also genetic/phenotypic uniqueness of species between the four sites. While macroclimatic conditions are rather different, microclimatic conditions and partly soil properties seem fairly homogeneous between the four sites, with the exception of water availability. Continuous activity monitoring of photosystem II revealed the BSCs of the Spanish site as the least active in terms of photosynthetic active periods.

The photobiont determines the pattern of photosynthetic activity within a single lichen thallus containing cyanobacterial and green algal sectors (photosymbiodeme)

Photosystem activity status of the green algal (Pseudocyphellaria lividofusca) and cyanobacterial (P. knightii) components of a photosymbiodeme were continuously monitored in the field over a period of 35xa0days. The photosymbiodeme grew on a Nothofagus menziesii tree at Lake Waikaremoana, Urewera National Park, North Island, New Zealand. Two Mini-PAM fluorometers were placed so that the chlorophyll a fluorescence, temperature and PPFD (photosynthetically active photon flux density) could be recorded every 30xa0min for green algal and cyanobacterial parts of the thallus. Microclimate conditions were also recorded with a datalogger. The study confirmed the already known ability of green algal lichens to reactivate from high humidity alone whilst cyanobacterial species need liquid water, here obtained from rainfall. The photosystems of P. lividofusca were activated on every day and positive ETR (relative electron transport rate) occurred on all but 3xa0days. Activation level depended on the overnight relative humidity. P. knightii was activated and had positive ETR on only 13xa0days when rainfall had occurred. Both species were mostly inactive above 12°C but differed at low temperatures. P. knightii showed no activation at very low temperatures, –2 to 0°C, since these only occurred on clear, rain-free nights. PPFD was always very low, mostly below 80xa0µmol m–2 s–1, and both species were inactive at higher PPFD. The three-dimensional structure of the thallus seemed to contribute to the hydration. The cyanobacterial sectors were more appressed to the trunk and needed substantial rainfall to rewet whereas the green algal lobes were more distant from the trunk and this probably caused more rapid desiccation as well as lower temperatures. It is suggested that the longer active periods for photosynthesis by P. lividofusca are balanced by several factors: first, depressed net photosynthesis at high thallus water contents after rainfall, a feature not shown by P. knightii; second, possible lower maximal net photosynthetic rates; and third, the possibility of greater respiratory rates when thalli have been hydrated by high relative humidity. There is little evidence for high PPFD differently affecting the photosymbiodeme components since sustained, high PPFD did not occur. It has been reported that the photosystems of cyanobacterial species from photosymbiodemes can reactivate at high relative humidity but the results obtained here suggest that it is not ecologically significant.

Photosynthetic responses of three common mosses from continental Antarctica

Predicting the effects of climate change on Antarctic terrestrial vegetation requires a better knowledge of the ecophysiology of common moss species. In this paper we provide a comprehensive matrix for photosynthesis and major environmental parameters for three dominant Antarctic moss species (Bryum subrotundifolium, B. pseudotriquetrum and Ceratodon purpureus). Using locations in southern Victoria Land, (Granite Harbour, 77°S) and northern Victoria Land (Cape Hallett, 72°S) we determined the responses of net photosynthesis and dark respiration to thallus water content, thallus temperature, photosynthetic photon flux densities and CO2 concentration over several summer seasons. The studies also included microclimate recordings at all sites where the research was carried out in field laboratories. Plant temperature was influenced predominantly by the water regime at the site with dry mosses being warmer. Optimal temperatures for net photosynthesis were 13.7°C, 12.0°C and 6.6°C for B. subrotundifolium, B. pseudotriquetrum and C. purpureus, respectively and fall within the known range for Antarctic mosses. Maximal net photosynthesis at 10°C ranked as B. subrotundifolium > B. pseudotriquetrum > C. purpureus. Net photosynthesis was strongly depressed at subzero temperatures but was substantial at 0°C. Net photosynthesis of the mosses was not saturated by light at optimal water content and thallus temperature. Response of net photosynthesis to increase in water content was as expected for mosses although B. subrotundifolium showed a large depression (60%) at the highest hydrations. Net photosynthesis of both B. subrotundifolium and B. pseudotriquetrum showed a large response to increase in CO2 concentration and this rose with increase in temperature; saturation was not reached for B. pseudotriquetrum at 20°C. There was a high level of variability for species at the same sites in different years and between different locations. This was substantial enough to make prediction of the effects of climate change very difficult at the moment.