Thallus Water Content Research Articles

The Hydration-Dependent Dynamics of Greenhouse Gas Fluxes of Epiphytic Lichens in the Permafrost-Affected Region

Recent studies actively debate oxic methane (CH4) production processes in water and terrestrial ecosystems. This previously unknown source of CH4 on a regional and global scale has the potential to alter our understanding of climate-driving processes in vulnerable ecosystems, particularly high-latitude ecosystems. Thus, the main objective of this study is to use the incubation approach to explore possible greenhouse gas (GHG) fluxes by the most widely distributed species of epiphytic lichens (ELs; Evernia mesomorpha Nyl. and Bryoria simplicior (Vain.) Brodo et D. Hawksw.) in the permafrost zone of Central Siberia. We observed CH4 production by hydrated (50%–400% of thallus water content) ELs during 2 h incubation under illumination. Moreover, in agreement with other studies, we found evidence that oxic CH4 production by Els is linked to the CO2 photoassimilation process, and the EL thallus water content regulates that relationship. Although the GHG fluxes presented here were obtained under a controlled environment and are probably not representative of actual emissions in the field, more research is needed to fully comprehend ELs’ function in the C cycle. This particular research provides a solid foundation for future studies into the role of ELs in the C cycle of permafrost forest ecosystems under ongoing climate change (as non-methanogenesis processes in oxic environments).

Modelling the carbon balance in bryophytes and lichens: Presentation of PoiCarb 1.0, a new model for explaining distribution patterns and predicting climate-change effects.

Bryophytes and lichens have important functional roles in many ecosystems. Insight into their CO2 -exchange responses to climatic conditions is essential for understanding current and predicting future productivity and biomass patterns, but responses are hard to quantify at time scales beyond instantaneous measurements. We present PoiCarb 1.0, a model to study how CO2 -exchange rates of these poikilohydric organisms change through time as a function of weather conditions. PoiCarb simulates diel fluctuations of CO2 exchange and estimates long-term carbon balances, identifying optimal and limiting climatic patterns. Modelled processes were net photosynthesis, dark respiration, evaporation and water uptake. Measured CO2 -exchange responses to light, temperature, atmospheric CO2 concentration, and thallus water content (calculated in a separate module) were used to parameterize the model's carbon module. We validated the model by comparing modelled diel courses of net CO2 exchange to such courses from field measurements on the tropical lichen Crocodia aurata. To demonstrate the model's usefulness, we simulated potential climate-change effects. Diel patterns were reproduced well, and the modelled and observed diel carbon balances were strongly positively correlated. Simulated warming effects via changes in metabolic rates were consistently negative, while effects via faster drying were variable, depending on the timing of hydration. Reproducing weather-dependent variation in diel carbon balances is a clear improvement compared to simply extrapolating short-term measurements or potential photosynthetic rates. Apart from predicting climate-change effects, future uses of PoiCarb include testing hypotheses about distribution patterns of poikilohydric organisms and guiding conservation strategies for species.

How dry is dry? Molecular mobility in relation to thallus water content in a lichen.

Lichens can withstand extreme desiccation to water contents of ≤ 0.1 g H2O g-1 DW, and in the desiccated state are among the most extremotolerant organisms known. Desiccation-tolerant life-forms such as seeds, mosses and lichens survive 'vitrification', that is the transition of their cytoplasm to a 'glassy' state, which causes metabolism to cease. However, our understanding of the mechanisms of desiccation tolerance is hindered by poor knowledge of what reactions occur in the desiccated state. Using Flavoparmelia caperata as a model lichen, we determined at what water contents vitrification occurred upon desiccation. Molecular mobility was assessed by dynamic mechanical thermal analysis, and the de- and re-epoxidation of the xanthophyll cycle pigments (measured by HPLC) was used as a proxy to assess enzyme activity. At 20 °C vitrification occurred between 0.12-0.08 g H2O g-1 DW and enzymes were active in a 'rubbery' state (0.17 g H2O g-1 DW) but not in a glassy state (0.03 g H2O g-1 DW). Therefore, desiccated tissues may appear to be 'dry' in the conventional sense, but subtle differences in water content will have substantial consequences on the types of (bio)chemical reactions that can occur, with downstream effects on longevity in the desiccated state.

The lifestyle of lichens in soil crusts

AbstractLichens are one of the common dominant biota in biological soil crusts (biocrusts), a community that is one of the largest in extent in the world. Here we present a summary of the main features of the lifestyle of soil crust lichens, emphasizing their habitat, ecophysiology and versatility. The soil crust is exposed to full light, often to high temperatures and has an additional water source, the soil beneath the lichens. However, despite the open nature of the habitat the lichens are active under shady and cooler conditions and avoid climate extremes of high temperature and light. In temperate and alpine habitats they can also be active for long periods, several months in some cases. They show a mixture of physiological constancy (e.g. similar activity periods and net photosynthetic rates) but also adaptations to the habitat (e.g. the response of net photosynthesis to thallus water content can differ for the same lichen species in Europe and the USA and some species show extensive rhizomorph development). Despite recent increased research, aspects of soil crust ecology, for example under snow, remain little understood.

Dry-hot stress significantly reduced the nitrogenase activity of epiphytic cyanolichen

Nitrogen (N) fixed by epiphytic cyanolichens (i.e. lichens that contain cyanobacterial symbionts) is thought to be the most important resource of this nutrient in some natural forest ecosystems. Although a great deal of work has been carried out to evaluate the biomass of this group as well as its contribution to ecosystem N budgets, empirical studies are needed to confirm the N input responses by cyanolichens under climate change conditions (dry-hot stress) as well as to determine the factors that control this process. We simulated climate change conditions by transplanting Lobaria retigera, a common cyanolichen in the area, to lower elevations, and measured nitrogenase activity in response to warmer and drier conditions. In addition, we conducted a series of laboratory and greenhouse experiments to determine the dominant factors influencing nitrogenase activity in this species. The results of this study show that mean annual nitrogenase activity at the higher site was 1.5 and 2.4 times that at the simulated warmer and drier (middle and lower) sites, respectively. Combining laboratory experimental conclusions, we show that thallus water content is a key factor determining the nitrogenase activity of L. retigera in early transplantation while insufficient carbon storage resulting from a combination of warming and desiccation was likely responsible for reducing nitrogenase activity in later months of the transplant experiment. The results of this study imply that the negative impact of climate change (dry-hot stress) on ecosystems not only impacts the distribution and growth of species, but also nutrient circles and budgets.

Distributional and ecophysiological study on the Antarctic lichens species pair Usnea antarctica/Usnea aurantiaco-atra

The factors that control lichen distribution in Antarctica are still not well understood, and in this investigation we focused on the distribution, local and continental, and gas exchange of a species pair, closely related lichens with differing reproductive strategies, Usnea aurantiaco-atra (fertile) and Usnea antarctica (sterile, sorediate). The local distributions of these species were recorded along an altitudinal gradient of nearly 300 m at South Bay, Livingston Island, and microclimate was also recorded over 1 year. The photosynthetic responses to temperature, light and thallus water content were determined under controlled conditions in the laboratory. The species were almost identical in their photosynthetic profiles. Locally, on Livingston Island, U. antarctica was confined to low altitude sites which were warmer and drier, whilst U. aurantiaco-atra was present at all altitudes. This contrasts with its distribution across Antarctica where U. antarctica grows 9° latitude further south than U. aurantiaco-atra. Temperature appears not to be the main controller of distribution in these species, but dryness of habitat, which will influence length of activity periods, may be important.

TWO SPECIFIC CAUSES OF CELL MORTALITY IN FREEZE-THAW CYCLE OF YOUNG THALLI OF PORPHYRA YEZOENSIS (BANGIALES, RHODOPHYTA)1

Porphyra yezoensis Ueda is an important marine aquaculture crop with single-layered gametophytic thalli. In this work, the influences of thallus dehydration level, cold-preservation (freezing) time, and thawing temperature on the photosynthetic recovery of young P. yezoensis thalli were investigated employing an imaging pulse-amplitude-modulation (PAM) fluorometer. The results showed that after 40 d of frozen storage when performing thallus thawing under 10°C, the water content of the thalli showed obvious effects on the photosynthetic recovery of the frozen thalli. The thalli with absolute water content (AWC) of 10%–40% manifested obvious superiority compared to the thalli with other AWCs, while the thalli thawed at 20°C showed very high survival rate (93.10%) and no obvious correlation between thallus AWCs and thallus viabilities. These results indicated that inappropriate thallus water content contributed to the cell damage during the freeze-thaw cycle and that proper thawing temperature is very crucial. Therefore, AWC between 10% and 40% is the suitable thallus water content range for frozen storage, and the thawing process should be as short as possible. However, it is also shown that for short-term cold storage the Porphyra thallus water content also showed no obvious effect on the photosynthetic recovery of the thalli, and the survival rate was extremely high (100%). These results indicated that freezing time is also a paramount contributor of the cell damage during the freeze-thaw cycle. Therefore, the frozen nets should be used as soon as time permits.

Photosynthesis in chlorolichens: the influence of the habitat light regime

The hypothesis that CO(2) gas exchange and chlorophyll a fluorescence (ChlaF) of lichens vary according to the light regimes of their original habitat, as observed in vascular plants, was tested by analysing the photosynthetic performance of 12 populations of seven dorsoventral, foliose lichens collected from open, south-exposed rocks to densely shaded forests. Light response curves were induced at optimum thallus water content and ChlaF emission curves at the species-specific photon flux at which the quantum yield of CO(2) assimilation is the highest and is saturating the photosynthetic process. Photosynthetic pigments were quantified in crude extracts. The results confirm that the maximum rate of gross photosynthesis is correlated with the chlorophyll content of lichens, which is influenced by light as well as by nitrogen availability. Like leaves, shade tolerant lichens emit more ChlaF than sun-loving ones, whereas the photosynthetic quantum conversion is higher in the latter.

Diel and seasonal courses of ambient carbon dioxide concentration and their effect on productivity of the epilithic lichen Lecanora muralis in a temperate, suburban habitat

Abstract:Ambient CO2 concentration (together with CO2 exchange and microclimate) was recorded every 30 min for 15 months for Lecanora muralis growing in the Botanical Garden Würzburg (Germany, northern Bavaria), a habitat on the outskirts of the city. Annual mean CO2 was around 17 ppm higher than the global average reported for the time of measurement (361 ppm; 1995/96), and daily values ranged from 317 to 490 ppm. Diel courses of CO2 could be classified into three different types. Type A, when CO2 levels rose overnight and then fell strongly to below global levels during the day, which predominated in the summer (about 75 of days); Type B, irregular diel courses occurred during all seasons with often very rapid changes apparently due to advective CO2 transport; Type C, CO2 concentration was typically almost stable at generally between c. 330 and 430 ppm which predominated in the winter (63 of days).Under controlled conditions, CO2 saturation of net photosynthesis (NP) of L. muralis at optimal hydration and light occurred at around 1000 ppm. NP was also affected by low CO2 at limiting light and thallus water contents. Based upon these data, we estimated the improvement of NP of L. muralis due to transient increase of ambient CO2 (as compared with the global average) for one selected combination of environmental factors (nocturnal dew or frost). This combination is an important source of water for the lichen, resulting in 40 of its annual production and, especially in these situations, photosynthesis was increased by high ambient CO2 in the early morning under prevailing Type A conditions. After dew activation, light compensation point of NP occurred at an average concentration of 413 ppm and diel maxima of NP at 402 ppm. This allows a rough estimate that the transiently elevated CO2 increased the photosynthetic gain of the lichen after dew of 7, or an improvement to its annual carbon balance of about 3. Conditions, especially interrelationships between lichen hydration, light and CO2 are so complex that we are not yet able to extend our estimates to other environmental situations of photosynthetic activity of L. muralis.

Nocturnal respiration of lichens in their natural habitat is not affected by preceding diurnal net photosynthesis

Dark respiration (DR) of lichens is reported to be higher in species with a high photosynthetic potential (suggesting a metabolic maintenance cost effect) and also, often in laboratory studies, transiently after photosynthesis (suggesting a substrate-driven effect). We investigated the occurrence of the latter, the effect of diurnal net photosynthesis (NP) on subsequent nocturnal DR, under natural temperate climate conditions in the chlorolichens Lecanora muralis and Cladonia convoluta and the cyanolichen Collema cristatum. Data sets totalling 15 months, 106 and 113 days, respectively, were obtained from automatic cuvettes that continually measured CO2 exchange and ambient conditions at 30 min intervals. For each 24 h period (sunrise to following sunrise), several measures of NP and DR were extracted, including maximal and mean rates and daily sums. No statistically significant correlations between the various measures of DR and preceding NP were found for L. muralis, only one weak correlation for Co. cristatum (the means of DR and NP) and three for Cl. convoluta (sums and means of DR and NP). It is proposed that even these significant correlations are actually a result of embedded codependencies between NP, DR and thallus water content. Overall it is concluded that no substrate-driven dependency of DR on preceding NP under natural conditions could be recognised. The periods of desiccation that often occur between the NP and following DR as well as the wide range of combinations of conditions would certainly contribute to this lack of relationship.

Water relations and CO 2 exchange of the terrestrial lichen Teloschistes capensis in the Namib fog desert: Measurements during two seasons in the field and under controlled conditions

Although the coastal zone of the Central Namib Desert (Namibia) has negligible rainfall, frequent fog, dew and high air humidity support a luxurious lichen flora. Large areas of soil crust communities are dominated by the multibranched, fruticose Teloschistes capensis interspersed by a (still indeterminable) Ramalina species. In earlier communications, based on field measurements in autumn, we began the analysis of functional mechanisms that allow these lichens to exist under the special conditions of a fog desert. We have extended this work by monitoring lichen CO 2 exchange and water relations in spring and by experiments under controlled conditions. In both seasons, nocturnal hydration, by fog and/or dew, activated dark respiration of the lichens which was followed, after sunrise, by a short period of positive net photosynthesis (NP) that continued until metabolic inactivation occurred from desiccation. Dry thalli of T. capensis were able to reactivate NP through water vapour uptake alone, beginning at an air relative humidity of 82%, i.e. at a water potential of −26.3 MPa; the moisture compensation point during desiccation was at 13% thallus water content (WC, dry weight related). Optimal WC for photosynthesis was around 100%, and both species showed a large and extended suprasaturation depression of CO 2 assimilation. Light response showed “sun-plant” characteristics with saturation >1000 μmol m −2 s −1 photosynthetically active photon flux density (PPFD). However, due to rapid desiccation, the combination of light saturation with optimal WC very rarely occurred under field conditions. Light compensation point after sunrise was highly dependent on actual WC: at low hydration, it amounted to only ca. 10 μmol m −2 s −1 PPFD so that even the smallest levels of hydration could be used for carbon gain before desiccation took place again. This phenomenon was probably due to a hydration gradient in the thallus branches during transient moistening so that the outer photobiont layer was favoured in contrast to the internal mycobiont which remained dry longer and did not contribute respiratory CO 2 loss. Fully hydrated thalli had light compensation points around 50 μmol m −2 s −1 PPFD. Extended desiccation of 1–3 days had no impact on the magnitude and recovery of photosynthesis but, imposed desiccation of 10 days reduced NP in lab and field experiments and caused an extended period of recovery. “Resaturation respiration” was not detected in the field data, although it was present after experimental moistening of dry thalli. In spring, the higher fog frequency and intensity increased maximal nocturnal WC, maximal attained NP as well as integrated daily carbon income (ΣNP) compared to the autumn measurements. NP max and ΣNP depended on maximal nocturnal WC with a saturation-type response. In terms of carbon gain both species seem to be optimally adapted to nocturnal moistening up to 160% WC and were not able to make use of higher degrees of hydration, a feature that might well influence their habitat selection. Maximal daily carbon-related ΣNP for T. capensis was 4.6 mg C (g C) −1 day −1. A rough estimate of the annual (projected) area-related carbon balance (photosynthetic income minus respiratory losses) based on published fog and dew frequencies and personal observations was 15–34 mg C m −2 yr −1.

Estimation of the annual primary production of the lichen Cetrariella delisei in a glacier foreland in the High Arctic, Ny-�lesund, Svalbard

The fruticose lichen Cetrariella delisei is among the dominant lichen species in the deglaciated High Arctic areas of Svalbard. As part of a study of carbon cycling in the High Arctic, we aimed to estimate the primary production of lichen in a deglaciated area in Ny-Alesund, Svalbard (79° N), by examining the effects of abiotic factors on the net photosynthesis (Pn) and dark respiration (R) rates of C. delisei. Experiments were conducted in the snow-free season of 2000 using an open-fl ow gas exchange system with an infrared gas analyser. Positive photosynthetic activities were observed on rainy days or soon after rainfall when the thallus water content was high, whereas photosynthetic activities dropped below the detectable limit on clear days because of the low thallus water content. Under a suffi ciently high thallus water content and close to light saturation, Pn was nearly constant over a wide temperature range of 4 - 20 °C, while R increased with increasing temperature. We constructed a model for estimating the net primary production (NPP) of lichen based on the relationships between abiotic factors and the CO2 exchange rate. The mean, minimum and maximum NPP values in the snow-free season, estimated using meteorological data obtained from 1995–2003, were 5.1, 1.0 and 8.4 g dry wt. m–2 snow-free season–1, respectively. These results suggest that NPP is highly variable and the contribution of lichen to carbon input is small compared with that of vascular plants and mosses in the study site.

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.

Are tropical lowlands a marginal habitat for macrolichens? Evidence from a field study with Parmotrema endosulphureum in Panama

Summary There is a striking increase in the abundance of non-vascular organisms (mosses, liverworts, and macrolichens) with altitude in the tropics. The proposed mechanism behind this altitudinal gradient still awaits experimental verification. This study addresses this question using diel (= 24-h) time courses of CO 2 exchange, thallus water relations, and microclimate of the foliose lichen, Parmotrema endosulphureum , under natural tropical lowland conditions. In addition, we studied the responses of lichen gas exchange to experimental manipulations of incident light, temperature and thallus water content. Similar to the results of the only other field study on CO 2 exchange in lowland lichens, 24-h carbon gain was strongly limited by nocturnal CO 2 loss: the cumulative dark respiration reached almost 90% of diurnal carbon gain. Our estimates of long-term carbon gain are consistent with direct measurements of growth, and are substantially lower than those for lichens from montane sites in the tropics.

SEASONAL VARIATION OF P CONTENT AND MAJOR N POOLS IN PALMARIA PALMATA (RHODOPHYTA)1

The annual variation of major nitrogen pools, phosphorus, carbon, ash, and thallus water content in relation to seasonal environmental changes was studied in two northern Spanish populations of the edible seaweed Palmaria palmata (Linnaeus) Kuntze. Observed patterns were investigated using Spearman rank order correlation coefficients. There were significant relationships between thallus nutrient content and nitrate and orthophosphate seawater concentration, irradiance, temperature, and wave force. The highest levels of total N and P and nitrogenous compounds were observed during autumn and winter because the thallus stored N‐ and P‐rich compounds in response to high nutrient seawater concentration when growth was limited by low light and temperature. Phycoerythrin and other proteins were the main N reserves. Thallus P content was higher in algae from the eutrophic site. During spring, reduced N and P thallus content and increased ash, water, and C content were observed in the growing fronds. N and P seawater concentrations were undetectable during summer when nutrient reserves were low and growth was reduced and eventually suppressed, suggesting nutrient limiting conditions. Palmaria palmata clearly could take advantage of elevated N and P concentrations to create storage reserves in winter to support early summer growth. This storage response reduced the dependence of algal nutrition on the external nutrient supply and supports the use of pulse fertilization to diminish summer nutrient limitation of cultured algae.

Ecological significance of net photosynthesis activation by water vapour uptake in Ramalina capitata from rain-protected habitats in central Spain

The reactivation of net photosynthesis and dark respiration from water vapour and from liquid water in the lichen Ramalina capitata var. protecta (H. Magn.) Nimis from sheltered habitats in Central Spain was investigated. CO 2 gas-exchange response and thallus water content were measured in the laboratory as dry thalli were allowed to equilibrate by water vapour uptake under different relative humidities, and as saturated thalli dried. Water content of thalli from different habitats was also measured in the field during rainfall. The results show that the lichen is very efficient in using water vapour from the air for photosynthesis. Net photosynthetic rates were measured at very low water potentials (up to −26·9 Megapascal) with water contents of the thalli below 30% dry weight. Maximum net photosynthetic rates measured after water vapour uptake were in some cases similar to those obtained after hydration by liquid water, while respiration rates were always lower after water vapour uptake, indicating a relatively larger activation of the photobiont. Moreover, photosynthetic rates at thallus water contents below 60% dry weight were higher when hydration was achieved through water vapour uptake than with liquid water. Field measurements had shown that this lichen reached water contents through water vapour absorption of 40% dry weight that would allow a positive balance of CO 2 uptake. Possible ecological implications of differences in photosynthetic activation depending on the water source are discussed in terms of the carbon gain under natural conditions of this ombrophobous lichen.

Photosynthetic productivity of the epilithic lichen Lecanora muralis: Long-term field monitoring of CO2 exchange and its physiological interpretation. I. Dependence of photosynthesis on water content, light, temperature, and CO2 concentration from laboratory measurements

Summary CO 2 exchange of the epilithic, green algal lichen Lecanora muralis was monitored continuously for 15 months by means of an automatic cuvette in its natural habitat in the Botanical Garden, Wurzburg (Bavaria, Germany). As a first stage in the interpretation of the results, identically treated parallel samples were used, under controlled laboratory conditions, to determine the dependence of net photosynthesis (NP) and dark respiration (DR) on light, CO 2 concentration, thallus hydration and temperature. Results of these measurements are communicated in the present, first part of a series of publications. The lichen showed an extremely large and extended suprasaturation depression of its CO 2 assimilation. NP at high hydration was reduced almost to the compensation point. Maximal NP was only possible in a very small window of thallus water content (WC). Thallus water relations dominated the lichen’s responses to the other environmental factors and the combination of ‘optimal’ moisture and light conditions was a rare event and lasted only for short times under natural conditions. Light response curves showed typical saturation kinetics with ‘sun-plant’ characteristics. Maximal NP, light saturation, maximal quantum use efficiency, and light compensation point were highly dependent on WC. Temperature optimum of NP was about 22°C. But, at 2°C, NP was still 60% of maximal NP. Under optimal WC, NP became saturated at c 1000 ppm external CO 2 and, at natural, ambient CO 2 , NP was 62% of maximal NP with a CO 2 compensation point of 16 ppm. Suprasaturation depression of NP was not completely removed by 5000 ppm external CO 2 . Seasonal changes in photosynthetic capacity and other photosynthetic parameters were extremely small. However, dark respiratory capacity showed a clear adaptation to low winter temperatures. This resulted in a higher light compensation point for CO 2 exchange in winter than in summer. The daily and seasonal courses of CO 2 exchange of L. muralis under natural conditions will be reported in further communications. The field performance of the lichen will be interpreted and analysed by means of the data presented here.

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 35days. 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 30min 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 3days. Activation level depended on the overnight relative humidity. P. knightii was activated and had positive ETR on only 13days 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 80µmol 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.

Environmental variables controlling the uptake of carbonyl sulfide by lichens

The uptake of atmospheric carbonyl sulfide (COS) by the lichen species Ramalina menziesii, representative for the open oak woodland in central California, was studied under laboratory conditions. By use of a dynamic cuvette system, the controlling parameters for the COS uptake were investigated under climate chamber conditions. The thallus water content, essential for the overall physiology of lichens, was found to be of basic importance for the trace gas exchange. A water content of 30% was the approximate minimum for COS uptake, with increasing activity up to a water content of 200%. Additionally, actual atmospheric mixing ratios have a significant influence on the exchange. The COS uptake was found to be a linear function of the ambient COS mixing ratio resulting in a compensation point as low as 37 ppt. A temperature optimum of 25°C was indicative of a physiological basis of the COS uptake. The inhibition of the COS consumption in the presence of a specific inhibitor for the enzyme carbonic anhydrase proved this enzyme to be of key relevance for the uptake. All these variables controlling the COS deposition were integrated into an uptake algorithm to model the exchange behavior of this lichen. The applicability of the model to field data is demonstrated. Uptake rates on a dry weight basis normalized to optimized conditions (25°C; 450 ppt COS) reached 0.17±0.09 pmol g−1 s−1 (i.e. 4.2±2.2 pmol m−2 s−1 thallus surface area, respectively). The contribution of lichens to the global COS sink strength is assigned to be about 0.3 Tg a−1, representing not a major but a significant sink.

Water relations and photosynthetic performance of fruticose lichens from the semiarid southeast of spain

The lichen species Teloschistes lacunosus (RUPR.) SAV. and Ramalina bourgeana NYL. are characteristic of the semiarid southeast Iberian Peninsula. These species showing different distribution patterns and ecological requirements, were collected at the coast ( R. bourgeana ) and in the inland ( T. lacunosus ). Laboratory experiments revealed that net photosynthesis (NP) and dark respiration (DR) responses to temperature differed between both species. Optimal temperatures of NP were higher in T. lacunosus than in R. bourgeana at optimal thallus water content at the various photosynthetic photon flux density (PPFD) levels. In the upper temperature range studied (20–25°C), R. bourgeana proved to be less efficient than T. lacunosus . In both species, lower and upper temperature compensation points were below 0°C and above 30°C, for moderate and high PPFD levels, respectively. At high temperatures and high thallus water content, T. lacunosus showed lower DR rates than R. bourgeana . NP response to PPFD also showed differences between both species. The light compensation points were high in both species for temperatures above 20°C (at 20°C around 50 μmol m −2 s −1 in T. lacunosus , and around 60 µmol m −2 s −1 in R. bourgeana ). Light saturation of NP was higher in T. lacunasus at all temperatures in the experiments. In T. lacunasus , the value was below 1200 µmol m −2 s −1 at 0-15°C, but at temperatures above 15°C, it was above 1200 µmol m −2 s −1 . In R. bourgeana the values were always below 1200 µmol m −2 s −1 . In both species, NP was strongly depressed at supraoptimal water content. The physiological differences observed between R. bourgeana and T. lacunasus , and their relation to the distribution patterns of these lichens are discussed.