Seasonal Trends in Nitrogen Status of Antarctic Lichens

The thallus nitrogen (N) concentration of two dominant macrolichensof continental Antarctica (Usnea sphacelata and Umbilicariadecussata) was estimated each month for 1 year on a low roundedknoll on Clark Peninsula, Windmill Islands, Wilkes Land, Antarctica.Thallus N was significantly higher in Umbilicaria decussatathan in Usnea sphacelata and varied according to site. Duringthe winter months, when the lichens were metabolically inactive,thalli gradually accumulated N. At the onset of warmer conditions,thallus N content fell dramatically in both species, with thetiming of the decline being related to microclimatic conditions.The strongly seasonal pattern of metabolic activity in thesespecies is reflected in their nitrogen relations. Copyright2000 Annals of Botany Company Lichen, nitrogen, Antarctica, season, Usnea sphacelata, Umbilicaria decussata     

Vegetation and ecology of ice-free areas of Northern Victoria Land,Antarctica

Summary At Birthday Ridge, a small ice free area in northern Victoria Land (70°48S, 167°00E), cryptogamic vegetation is mostly confined to gaps between granitc rocks. The sheltering effect on lichens and mosses was analyzed by continuous measurements of the microclimate at various levels between the rocks. Although warming by solar radiation was favourable for the existence of cryptogams, rocks strongly insolated were mostly devoid of lichens and mosses. Lichens in the soaked active state were heated to above air temperature but did not reach more than 10°C. The presence of lichens was dependent on the moisture conditions of the habitat. It was observed that snow, the only source of moisture, accumulated in summer only in deeper levels between rocks, and that the snow rapidly melted on contact with the lichens. After a snow shower,Usnea sulphurea gained 67% andUmbilicaria decussata 94% of their maximum water capacity. During one quarter of the time period of 7 days the lichens were soaked and therefore capable of carrying out photosynthesis. The lichens in soaked state had always less than optimum temperatures for net photosynthesis. The rock gaps at Birthday Ridge form oases, the only localities where moisture is provided, and temperature is high enough to enable growth of lichens and mosses.Bryum is also able to exist in the upper 3 cm of soil.     

五种南极地衣的Co、Cr、Pb和Cu元素富集能力的差异

随着人类活动对全球环境的影响,环境重金属污染负荷日益增加,重金属元素向南极地区的迁移和富集已成为该地区面临的严重问题之一。地衣是研究重金属元素大气沉降特征的良好材料,但对南极地衣的重金属元素大气沉降富集能力的比较研究甚少。作者以网袋法实验比较采自西南极乔治王岛菲尔德斯半岛的5种南极优势地衣,即:王橙衣Caloplaca regalis、夹心果衣Himantormia lugubris、孔树花Ramalina terebrata、球粉衣Sphaerophorus globosus和簇花石萝Usnea aurantiacoatra对钴(Co)、铬(Cr)、铅(Pb)和铜(Cu)的富集能力。2个月的网袋法实验结果表明:1)5种地衣对Pb元素均表现出富集能力,其中夹心果衣富集能力最强,簇花石萝和孔树花次之;2)簇花石萝和夹心果衣对Cu元素的富集能力相近,球粉衣对Cu元素的富集能力较低,其他2种未表现出对Cu的富集作用;3)Co和Cr元素的富集仅分别在夹心果衣和簇花石萝中检测出来。因此,在综合检测Co、Cr、Cu和Pb的大气沉降时,可考虑簇花石萝和夹心果衣的组合;在单独检测Co、Pb和Cu的大气沉降时,夹心果衣均为最佳的物种。     

Summer variability, winter dormancy: lichen activity over 3?years at Botany Bay, 77��S latitude, continental Antarctica

Lichens make up a major component of Antarctic vegetation; they are also poikilohydric and are metabolically active only when hydrated. Logistic constraints have meant that we have little idea of the length, timing or environmental conditions of activity periods of lichens. We present the results of a three-year monitoring of the activity of the lichen Umbilicaria aprina at Botany Bay (77°S latitude) in the Ross Sea region, continental Antarctica. Chlorophyll fluorescence parameters that allowed hydrated metabolic activity to be detected were recorded with a special fluorometer at 2- or 3-h intervals. Air and thallus temperatures and incident PPFD (photosynthetic photon flux density, μmol photon m⁻² s⁻¹) were also recorded at hourly intervals. Activity was extremely variable between months and years and, overall, lichen was active for 7% of the 28-month period. Spring snow cover often delayed the onset of activity. Whereas the period immediately after snow melt was often very productive, the later months, January to March, often showed low or no activity. Mean thallus temperature when active was just above zero degrees and much higher than the annual mean air temperature of −15 to −19°C. Because major snow melts occurred when incident radiation was high, the lichen was also subjected to very high PPFD when active, often more than 2,500 μmol photon m⁻² s⁻¹. The major environmental stress appeared to be high light rather than low temperatures, and the variability of early season snow fall means that prediction of activity will be very difficult.     

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 degrees C through absorption of water from high air humidity. Physcia dubia showed some activation at temperatures around -5 degrees C but only became fully activated at thallus temperatures of 0 degrees 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.     

Morphology influences water storage in hair lichens from boreal forest canopies

Hair lichens (Alectoria, Bryoria, Usnea) with high surface-area-to-mass ratios rapidly trap moisture. By photography and scanning we examined how internal water storage depended on morphological traits in five species. Specific thallus mass (STM, mg DM cm⁻²) and water holding capacity (WHC, mg H₂O cm⁻²) after shaking and blotting a fully hydrated thallus increased with thallus area. STM was ≈50% higher in Alectoria and Usnea thalli than in Bryoria. WHC was highest in Alectoria while percent water content of freshly blotted thalli was lowest in Usnea. Thallus area overlap ratio (TAO), assessing branch density of the thallus, was highest in the two thinnest Bryoria; lower in the thicker Usnea. Within species, hair lichens increased their water storage by increasing branch density rather than branch diameter. The taxonomically related genera Alectoria and Bryoria shared water storage characteristics, and differed from Usnea. Hair lichens in lower canopies have among the lowest water storage capacity reported in lichens.     

Some Aspects of the Water Relations of Lichens from Habitats of Contrasting Water Status studied using Thermocouple Psychrometry

Thermocouple psychrometry was used to investigate the waterrelations of eight species of lichens from sites of contrastingwater status. Lichens from exposed, dry habitats had lower thallusK concentrations but lower osmotic potentials than plants frommesic sites. As a result, K contributed a much smaller proportionto osmotic potential in plants from dry sites, suggesting thatlow cytoplasmic K concentrations are important for desiccationtolerance. While containing less water at saturation, plantsfrom dry habitats had lower thallus elastic moduli and lowerapoplastic water fractions. They therefore lost turgor at lowerrelative water contents than plants from wetter sites. The significanceof these results for the water relations of lichens is discussed.Copyright1995, 1999 Academic Press Lichen, desiccation, stress, thermocouple psychrometry     

Symbiosis as a successful strategy in continental Antarctica: performance and protection of Trebouxia photosystem II in relation to lichen pigmentation

Lichens as symbiotic associations consisting of a fungus (the mycobiont) and a photosynthetic partner (the photobiont) dominate the terrestrial vegetation of continental Antarctica. The photobiont provides carbon nutrition for the fungus. Therefore, performance and protection of photosystem II is a key factor of lichen survival. Potentials and limitations of photobiont physiology require intense investigation to extend the knowledge on adaptation mechanisms in the lichen symbiosis and to clarify to which extent photobionts benefit from symbiosis. Isolated photobionts and entire lichen thalli have been examined. The contribution of the photobiont concerning adaptation mechanisms to the light regime and temperature conditions was examined by chlorophyll a fluorescence and pigment analysis focusing on the foliose lichen Umbilicaria decussata from North Victoria Land, continental Antarctica. No photoinhibition has been observed in the entire lichen thallus. In the isolated photobionts, photoinhibition was clearly temperature dependent. For the first time, melanin in U. decussata thalli has been proved. Though the isolated photobiont is capable of excess light protection, the results clearly show that photoprotection is significantly increased in the symbiotic state. The closely related photobiont of Pleopsidium chlorophanum, a lichen lacking melanin, showed a higher potential of carotenoid-based excess light tolerance. This fact discriminates the two photobionts of the same Trebouxia clade. Based on the results, it can be concluded that the successful adaptation of lichens to continental Antarctic conditions is in part based on the physiological potential of the photobionts. The findings provide information on the success of symbiotic life in extreme environments.     

Photobiont selectivity for lichens and evidence for a possible glacial refugium in the Ross Sea Region, Antarctica

Lichens are a symbiosis consisting of heterotrophic, fungal (mycobiont) and photosynthetic algal or cyanobacterial (photobiont) components. We examined photobiont sequences from lichens in the Ross Sea Region of Antarctica using the internal transcribed spacer region of ribosomal DNA and tested the hypothesis that lichens from this extreme environment would demonstrate low selectivity in their choice of photobionts. Sequence data from three targeted lichen species (Buellia frigida, Umbilicaria aprina and Umbilicaria decussata) showed that all three were associated with a common algal haplotype (an unnamed Trebouxia species) which was present in all taxa and at all sites, suggesting lower selectivity. However, there was also association with unique, local photobionts as well as evidence for species-specific selection. For example, the cosmopolitan U. decussata was associated with two photobiont species, Trebouxia jamesii and an unnamed species. The most commonly collected lichen (B. frigida) had its highest photobiont haplotype diversity in the Dry Valley region, which may have served as a refugium during glacial periods. We conclude that even in these extreme environments, photobiont selectivity still has an influence on the successful colonisation of lichens. However, the level of selectivity is variable among species and may be related to the ability of some (e.g. B. frigida) to colonise a wider range of habitats.     

Carbon dioxide exchange in Cladina lichens from subarctic and temperate habitats

Summary The survival potential of lichens in a given habitat is determined by the response of CO₂ exchange to photosynthetically active radiation (PhAR), thallus temperature, and thallus relative water content (RWC). Therefore morphologically similar lichens from contrasting climatic environments 1) should differ in their CO₂ exchange responses, and 2) these differences should reflect adaptations to their climatic regimes. The CO₂ exchange responses of a subarctic (55°N, 67°W) Cladina stellaris (Opiz) Brodo population and a temperate (29°N, 82°W) Cladina evansii (Abb.) Hale and W. Culb, population were used to test these two related hypotheses.Infrared gas analysis with lichens collected in September–October 1975 established that the two populations differed in their responses to incident PhAR, thallus temperature, and thallus RWC. Net photosynthesis in C. stellaris had an optimum at a lower temperature and a greater relative photosynthetic capacity at low temperatures than did C. evansii. Cladina evansii maintained net photosynthesis above 35°C thallus temperature; C. stellaris did not. In both species the optimum temperature for net photosynthesis increased with increasing irradiance. The C. stellaris light saturation point was consistently lower than that of C. evansii. Both species had maximal rates of net photosynthesis at 70–80% relative water content. In C. evansii the CO₂ exchange rates, expressed as percentages of the maximum rate, declined more rapidly under suboptimal conditions. The absolute CO₂ exchange rates of C. evansii were greater than those of C. stellaris. At 20°C and 90–95% RWC, resaturation respiration occurred in both species and continued until 6–7 h after wetting.Contrasts in the temporal patterns of thallus condition at each collection site suggest that not all differences in the two response surfaces reflect climatic adaptation. The two populations appear well adapted to incident PhAR and thallus temperature regimes but the 70–80% RWC optimum for net photosynthesis common to both species is puzzling since their water regimes differ markedly. The overall adaptedness of the CO₂ exchange responses in the two species cannot be judged without a comprehensive quantitative analysis of carbon balance under differing climatic regimes.     

Differences in the utilization of water vapour and free water in two co‐occurring foliose lichens from semi‐arid southern Australia

Hydration and dehydration kinetics were investigated in two xerophytic lichens with contrasting thallus morphologies, Chondropsis semiviridis (F.Muell. ex Nyl.) Nyl. and Xanthoparmelia convoluta (Krempehl.) Hale. Pulse‐modulated chlorophyll fluorescence was used to measure photosynthetic activity in thalli hydrated with either liquid water or water vapour in the laboratory and in the field. Water content (WC) and photosynthetic activity of thalli in both species increased rapidly on contact with liquid water. When exposed to water vapour, C. semiviridis hydrated more rapidly and achieved higher WC than X. convoluta. Both lichens achieved maximum Fv/Fm at low WC, regardless of hydration source. Rates of water loss were slower, and Fv/Fm remained high for longer, in X. convoluta than in C. semiviridis. Light saturated electron transport rates of both lichens were low compared with a homoiohydric plant from the same environment. Our results suggest that X. convoluta, which has a more complex morphology, retains water and remains photosynthetically active for significantly longer periods than C. semiviridis, providing X. convoluta with a potential advantage in the semi‐arid environment in which both species are found.     

Flechten. Der Erfolg einer Symbiose

The success of a symbiosis: Lichens Lichens are a unique group of organisms composed of one or two alga and a fungus. Together they form species specific thalli. Their common eco‐physiological properties allow colonizing almost all terrestrial habitats, even the most hostile climatic zones on earth. However, as poikilohydrous organisms they also suffer from disadvantages related with their nature. As water content cannot be actively controlled, many lichens experience water‐oversaturation, thus being not able to gain full photosynthetic rates, even though they have otherwise optimal conditions. These eco‐physiological properties set up the frame for which microclimatic situation the realized thallus construction might do best. As all optimizations regarding water uptake also count for water loss, lichens are always at the edge of having either too much or not enough water for optimal carbon gain. So each habitat has its own challenge for the lichen thallus construction and lichens have to fit well into a specific ecological niche.     

Competitive behaviour of umbilicate lichens – an experimental approach

The lichens Lasallia pustulata and Umbilicaria spodochroa grow in dense monospecific or mixed populations on the coastal cliffs of southern Scandinavia. Attached to the substrate by only a thin central holdfast, their shield-shaped thalli compete for light and space for growth by overlapping each other. Matched pair experiments in the laboratory and field observations of interacting pairs show that different behavioural responses to precipitation tend to result in the margins of U. spodochroa overlapping those of L. pustulata within a few minutes. The behaviour is apparently caused by different capacities for water absorption in the upper and lower cortices of the species. An initial period of repeated encounter caused by thallus expansion and contraction during precipitation will be followed by a period in which U. spodochroa grows to overlap L. pustulata more and more. When the overlapping lichens are wet, flexible and photosynthetically active, the thallus above rests directly on the upper surface of the one below. Very little light is transmitted through thalli of U. spodochroa, and the shaded parts of L. pustulata are retarded in their growth and die off. Received: 29 August 1996 / Accepted: 19 April 1997     

New Insights into the Biological Activity of Lichens: Bioavailable Secondary Metabolites of Umbilicaria decussata as Potential Anticoagulants

This study reports the in vitro anticoagulation activity of acetonic extract (AE) of 42 lichen species and the identification of potential bioavailable anticoagulant compounds from Umbilicaria decussata as a competent anticoagulant lichen species. Lichens’ AEs were evaluated for their anticoagulant activity by monitoring activated partial thromboplastin time (APTT) and prothrombin time (PT) assays. A strong, positive correlation was observed between total phenolics concentration (TPC) of species and blood coagulation parameters. U. decussata was the only species with the longest clotting time in both APTT and PT assays. The research was moved forward by performing in vivo assays using rats. The results corroborated the dose-dependent impact of U. decussata’s AE on rats’ clotting time. Major secondary metabolites of U. decussata and their plasma-related bioavailability were also investigated using LC-ESI-MS/MS. Atranol, orsellinic acid, D-mannitol, lecanoric acid, and evernic acid were detected as possible bioavailable anticoagulants of U. decussata. Our findings suggest that U. decussata might be a potential anticoagulant lichen species that can be used for the prevention or treatment of coagulation-related issues such as cardiovascular diseases (CVDs).     

Carbon Dioxide Exchange in Lichens: Apparent Photorespiration and Possible Role of CO2 Refixation in Some Members of the Stictaceae (Lichenes)

At low O₂ concentration (1% O₂, v/v) the true photosynthetic(TP) rates of Pseudocyphellaria homoeophylla and Sticta latifronsare stimulated by up to 46% and 55% respectively. This stimulation,which is considered to be due to the repression of photorespiration(PR), varied with the thallus water content of the lichens.At intermediate and low water contents PR rates approximatedthe 43% of TP theoretically expected for C3 plants, but at veryhigh water contents PR rates were near zero. Results for P.billardierii show that in this species PR rates are low (upto 11% TP) at all water contents. Comparison of light/dark CO₂ evolution ratios under low O₂ conditionsindicated that up to 70% of respired CO₂ could be refixed withinthe lichen thallus. The lack of stimulation at 1% O₂ and highwater content is suggested to be a result of a high surfaceCO₂ diffusion resistance encouraging internal CO₂ refixation.Lichens thus join the group of plants with low CO₂ compensationvalues and photorespiration.     

Comparative study of the effects of temperature on net photosynthesis and respiration in lichens from the antarctic and subalpine zones in Japan

Photosynthetic activity and dark respiration were measured in some species of lichens,Umbilicaria, Cladonia, Stereocaulon, Usnea, etc. sampled in the Antarctic and subalpine zones of central and northern Japan. On the basis of the responses of their activities to thallus temperature, the response patterns were divived into three groups, and further, each group was subdvided into some adaptation types for net photosynthesis and repiration. Lichens collected in the Antarctic were adapted to cool condition and some species collected in subalpine zones were adapted to warm condition. For exmaple, the optimal temperatures for net photosynthetic rates in Antarctic lichens were lower than 5 C and those in lichens which lived on rock surface at a southern slope in mountains of Japan were higher than 20 C.     

Field Measurements of Photosynthesis of Umbilicarious Lichens in Winter

The photosynthetic performance of Lasallia pustulata and Umbilicaria spodochroa was measured both in the field on granitic rock in southern Norway and in the laboratory under controlled conditions of light and temperature. In the field thallus temperatures varied between ? 2 and + 5 °C during the daylight period in January 1994. In situ water contents were between 50 and 400% d.wt. in L. pustulata and between 100 and 500% d.wt. in U. spodochroa. The lichens were active during the whole period of investigation. Photosynthetic rates reached 13.03 μmol CO₂ g Chl^?1 s^?1 in L. pustulata and 5.56 μmol CO₂ g Chl^?1 s^?1 in U. spodochroa. Ice formation on the thallus surface did not impede CO₂ exchange. Light was mainly the limiting factor, as could be confirmed by the laboratory experiments. In general, habitat conditions never provided optimum photosynthetic rates but photosynthetic carbon balance was positive during 4 of the 5 days investigated. The coldest day was photosynthetically almost as efficient as the warmest day during this period.     

Response of the lichen-eating moth <Emphasis Type="Italic">Cleorodes lichenaria</Emphasis> larvae to varying amounts of usnic acid in the lichens

Lichens are characterized by a great variety of secondary metabolites. The function of these substances remains partly unknown. In this study, we propose that some of these metabolites may expel insect herbivores. To test this hypothesis, we reared larvae of the lichenivorous moth Cleorodes lichenaria on three selected lichens, Cladonia arbuscula subsp. mitis, Usnea hirta, and Usnea dasypoga. In experimental setup, the secondary metabolite usnic acid was removed from the lichens with acetone prior to feeding, whereas a control was left untreated. On all three lichens, removal of usnic acid from the lichens using acetone significantly prolonged survival of larvae and increased their viability. Larvae reared on control lichens contained significantly more usnic acid than those reared on treated lichens, both in their biomass and their faeces. These results support the hypothesis that usnic acid serves as a repellent against insect feeding, besides its well established functions of UV protection and antimicrobial properties.     

Spatial variation in carbon isotope discrimination across the thalli of several lichen species

Variations in stable carbon isotope discrimination (δ) were investigated across the thalli of several lichen species possessing different photobiont associations. Lichens containing (i) green algae (phycobiont), (ii) green algae in association with cyanobacteria confined in cephalodia, or (iii) cyanobacteria (cyanobiont) as the photobiont partner were studied. Carbon isotope discrimination was analysed in different thallus sections, which varied in distance from the margin and in age. The marginal thallus region is considered to be youngest, while the central region is thought to be oldest. This analysis showed a clear variation in δ across the thallus related to distance from the growing margin. In most of the species examined, the highest δ values were found in marginal regions (younger), while the central and basal regions (older) showed significantly lower δ. To investigate the effects of the historical increase in atmospheric CO₂ concentration and the concurrent decrease in the ¹³C content of atmospheric CO₂ on the δ of lichens, experiments were carried out on herbarium samples of Lobaria pulmonaria collected from the mid 19th Century to 1953. The results obtained showed a pattern of variation of δ consistent with that of freshly collected samples; δ decreased substantially with increasing distance from the thallus margin, irrespective of the collection date. Moreover, no consistent variation of discrimination was found among different collection dates. These results demonstrate that the observed variation in δ is caused by age-related changes in the physiological behaviour of different thallus sections, and that the past 150 years of increasing CO₂ concentration have not had significant effects on A in L. pulmonaria. Photosynthesis measurements, chlorophyll analysis and observations using optical microscopy, performed on freshly collected lichens, showed significant changes in morphological and physiological characteristics across the thallus. Particularly, remarkable variations in thickness were found across the thallus. These anatomical changes may be responsible for the variation in δ, through variations in CO2 transfer resistance and, consequently, in CO₂ availability across the thallus. The lack of age-dependent variation in δ in cyanobiont lichens is possibly attributable to the operation of a CO₂-coneentrating mechanism and, therefore, to a more constant CO₂ environment across the thallus in this lichen group.     

Photoprotection in the lichen Parmelia sulcata: the origins of desiccation-induced fluorescence quenching

Lichens, a symbiotic relationship between a fungus (mycobiont) and a photosynthetic green algae or cyanobacteria (photobiont), belong to an elite group of survivalist organisms termed resurrection species. When lichens are desiccated, they are photosynthetically inactive, but upon rehydration they can perform photosynthesis within seconds. Desiccation is correlated with both a loss of variable chlorophyll a fluorescence and a decrease in overall fluorescence yield. The fluorescence quenching likely reflects photoprotection mechanisms that may be based on desiccation-induced changes in lichen structure that limit light exposure to the photobiont (sunshade effect) and/or active quenching of excitation energy absorbed by the photosynthetic apparatus. To separate and quantify these possible mechanisms, we have investigated the origins of fluorescence quenching in desiccated lichens with steady-state, low temperature, and time-resolved chlorophyll fluorescence spectroscopy. We found the most dramatic target of quenching to be photosystem II (PSII), which produces negligible levels of fluorescence in desiccated lichens. We show that fluorescence decay in desiccated lichens was dominated by a short lifetime, long-wavelength component energetically coupled to PSII. Remaining fluorescence was primarily from PSI and although diminished in amplitude, PSI decay kinetics were unaffected by desiccation. The long-wavelength-quenching species was responsible for most (about 80%) of the fluorescence quenching observed in desiccated lichens; the rest of the quenching was attributed to the sunshade effect induced by structural changes in the lichen thallus.