Leopoldo G. Sancho

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.

Photosynthesis and water relations and the role of anatomy in Umbilicariaceae (lichenes) from Central Spain

SummaryThe response of net photosynthesis and dark respiration in eight species of Umbilicariaceae (lichenes) to temperature (-5, 0, 5, 10, 15, 20, 25, 30°C) and irradiance (55, 110, 220, 400, 620 μmol photons m-2 s-1 PAR) was studied. The samples were collected in montane and alpine localities of the Spanish Sistema Central. The species differed widely in their net photosynthetic rates. The optimal temperature for net photosynthesis in alpine species was significantly lower than in montane species. Montane species were more photophytic than alpine ones. Water saturation and water loss rate were dependent on morphology and particularly anatomy of the thallus. The physiological and structural data are useful in the interpretation of the ecology and altitudinal distribution of the Umbilicariaceae. No adaptation could be linked to particularities of the mediterranean climate.

Exploring the physiological state of continental Antarctic endolithic microorganisms by microscopy

In this microscopy study, we show that microorganisms comprising many endolithic communities of the McMurdo Dry Valleys of Antarctica appear in different physiological states. Live/dead microbial fluorescence stains were used to identify the state of microorganisms in the biofilms. The ultrastructures of these microorganisms were then characterized by transmission electron microscopy. Cyanobacteria were associated with heterotrophic bacterial cells, while fungal cells were free-living or formed partners with green alga as lichens. The extracellular polymeric substances, in which the endolithic microorganisms were embedded, formed an integral part of the biofilms observed. Extracellular polymeric substances probably play a significant role in nutrient interactions and protection of microorganisms from the environmental conditions outside the film. Living, moribund, dormant and dead microorganisms shared this microhabitat. The ecological impacts of the observed physiological dynamics are discussed.

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.

Ecology of endolithic lichens colonizing granite in continental Antarctica

In this study, the symbiont cells of several endolithic lichens colonizing granite in continental Antarctica and the relationships they have with the abiotic environment were analyzed in situ, in order to characterize the microecosystems integrating these lichens, from a microecological perspective. Mycobiont and photobiont cells, the majority classified as living by fluoresecent vitality testing, were observed distributed through the fissures of the granite. The fact that extracellular polymeric substances were commonly observed close to these cells and the features of these compounds, suggest a certain protective role for these substances against the harsh conditions of the environment. Different chemical, physical and biological relationships take place within the endo- lithic biofilms where the lichens are found, possibly affecting also the survival and distribution of these organisms. The alteration of bedrock minerals and synthesis of biominerals in the proximity of these lichens gives rise to different chemical microenvironments and suggests their participation in mineral nutrient cycling.

Ecophysiology of Desiccation/Rehydration Cycles in Mosses and Lichens

Although both lichens and bryophytes are all poikilohydric the groups seem to behave very differently. Bryophytes also show a clear preference for wetter areas and this seems to be a result of the different structures of the organisms. A lichen is algae (or cyanobacteria) suspended in a mycobiont with excess water often having a negative effect on photosynthesis. Bryophytes, in contrast, are true multicellular plants and can construct photosynthetic tissues that can effectively separate their photosynthetic and water storage functions. Under dry atmospheric conditions lichens and bryophytes will desiccate to low water contents and they become dormant. Ability to tolerate desiccation varies considerably both between and within the groups. Somewhat surprisingly, lichens appear to show less ability to tolerate long periods of desiccation than bryophytes, and even some vascular plants. Actual mechanisms of desiccation have been best studied in bryophytes and appear to be constitutive, no protein synthesis is required on rehydration to enable the commencement of metabolism and the necessary protection appears to be always present. Consistently high sucrose levels, for instance are reported from bryophytes. Cellular structure is often maintained when desiccated. Recovery from dryness also differs between the groups with bryophytes generally hydrating more slowly but there are large species differences. In general, rate of recovery may be related to the length of the hydrated activity period, species that hydrate and then dry rapidly, as on rock surfaces, recover rapidly. Species in habitats that remain wet for long periods once hydrated appear to recover more slowly from dryness. In addition to a photosynthetic response to light and temperature, the poikilohydric lichens and bryophytes also have a photosynthetic response to thallus water content. Starting with a dry thallus, addition of water will both increase the thallus water content and also allow photosynthesis and respiration to commence. Both processes increase almost linearly with further hydration at low water contents. Photosynthesis reaches a maximum at an optimal thallus water content (WCopt) that is strongly species dependant. In both groups this photosynthetic optimum represents full cellular turgor. At water contents above this optimum surface or external water can interfere with carbon dioxide uptake and can severely limit photosynthetic rates, especially in lichens. When thallus water contents are normalised to WCopt = 1, then the net photosynthesis (NP) response curves at water contents below WCopt are very similar for liverworts, mosses and higher plants, suggesting a common mechanism in controlling NP. It is suggested that this might be an inhibitor acting on Rubisco activity. In contrast to vascular plants both groups can carry out photosynthesis at lower, suboptimal thallus water contents and very low water potentials but the contribution that this makes to total carbon budget appears to be a major difference between the groups. Bryophytes seem to pass rapidly through this water content range when both drying and hydrating for tens of minutes are often enough. In contrast, it is now apparent that lichens are often active at low thallus water contents. They can not only hydrate from humid air alone, or from dew and fog, but can use these water sources very effectively, often achieving a major part of their annual carbon gain. Information on when the lichens and bryophytes are actually active is only recently starting to appear but, again, the groups seem to differ. Bryophytes strongly prefer wetter habitats and can be active and fully hydrated for long periods and seem to have excellent capacity to tolerate high light and UV radiation when wet. In contrast many lichens, in particular those with green algal symbionts, rarely seem to be hydrated for long periods, especially in high light conditions, and rapidly dry out. Lichens seem to be active mainly under suboptimal conditions one of which is suboptimal water content.

The weathering action of saxicolous lichens in maritime Antarctica

SummaryXanthoria elegans (Link) Th Fr. and Lecidea lapicida (Ach.) were studied on volcanic andesite, and Rhizocarpon geographicum (L.) DC. and Bacidia stipata Lamb on a volcanigenic sediment, using light microscopy, infrared spectroscopy, X-ray diffraction and transmission electron microscopy. Feldspars were present in the rocklichen interface to a lesser extent than in the underlying rock. R. geographicum was found to alter the minerals in the rock on which it grew without producing any new minerals in the rock/lichen interface, in contrast to the observations for this species on granite in temperate regions. Beneath of the thallus of L. lapicida there was calcium oxalate and some micas of the illite type, which may have been degradation products of various phyllosilicates in the rock. B. stipata, an endemic Antarctic lichen, had the greatest capacity to weather the rock and had weddellite (dihydrate calcium oxalate) and calcite in the contact area as well as many bacteria. The presence of crystalline oxalate, imogolite, allophane, carbonates (calcite) and amorphous material not found in the parent rock indicates biomineralization processes attributable to the lichens.

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.

Iron-Rich Diagenetic Minerals are Biomarkers of Microbial Activity in Antarctic Rocks

The cold, dry ecosystems of Antarctica have been shown to harbor traces left behind by microbial activity within certain types of rocks, but only two indirect biomarkers of cryptoendolithic activity in the Antarctic cold desert zone have been described to date. These are the geophysical and geochemical bioweathering patterns macroscopically observed in sandstone rock. Here we show that in this extreme environment, minerals are biologically transformed, and as a result, Fe-rich diagenetic minerals in the form of iron hydroxide nanocrystals and biogenic clays are deposited around chasmoendolithic hyphae and bacterial cells. Thus, when microbial life decays, these characteristic neocrystalized minerals act as distinct biomarkers of previous endolithic activity. The ability to recognize these traces may have potential astrobiological implications because the Antarctic Ross Desert is considered a terrestrial analogue of a possible ecosystem on early Mars.

Functional and spatial pressures on terrestrial vegetation in Antarctica forced by global warming

There is growing interest in what controls the present distribution of terrestrial vegetation in Antarctica because of the potential use of biodiversity as an indicator or predictor of the effects of climate change. Recent advances in knowledge of distribution and ecophysiological performance of terrestrial vegetation means that an initial analysis of the potential influence of temperature is now possible. Regressions of species numbers of lichens, mosses and hepatics on latitude and mean annual temperature (standard macroclimatic data) were carried out, and the terrestrial vegetation in Antarctica could be divided into two zones. The microenvironmental zone lies south of around 72°S, and biodiversity (richness and location) is uncoupled from the macroenvironment and is, instead, determined by the occasional coincidences of warmth, water, light and shelter. The macroenvironmental zone lies north of about 72°S, and biodiversity (richness, cover and growth) is strongly positively linked to mean annual temperature; species numbers increase at about 9–10% per K (24.0, 9.3 and 1.8 species for lichens, mosses and hepatics, respectively) probably due to improved water availability through increased precipitation and longer active period (monthly degree-days also reach zero at about 72°S) allowing greater productivity, completion of metabolic processes and a switch from survival to growth strategies. Cyanobacterial lichens appear to be a special case and may be expanding after being forced into northerly refugia. Warming will cause a southward movement of the boundary between the two zones but distribution in the microenvironmental zone will remain determined by local coincidences of environment and resources.