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Background and Aims: Cyanolichens are usually stated to be bipartite (mycobiont plus cyanobacterial photobiont). Analyses revealed green algal carbohydrates in supposedly cyanobacterial lichens (in the genera Pseudocyphellaria, Sticta and Peltigera). Investigations were carried out to determine if both cyanobacteria and green algae were present in these lichens and, if so, what were their roles., Methods: The types of photobiont present were determined by light and fluorescence microscopy. Small carbohydrates were analysed to detect the presence of green algal metabolites. Thalli were treated with selected strengths of Zn(2+) solutions that stop cyanobacterial but not green algal photosynthesis. CO(2) exchange was measured before and after treatment to determine the contribution of each photobiont to total thallus photosynthesis. Heterocyst frequencies were determined to clarify whether the cyanobacteria were modified for increased nitrogen fixation (high heterocyst frequencies) or were normal, vegetative cells., Key Results: Several cyanobacterial lichens had green algae present in the photosynthetic layer of the thallus. The presence of the green algal transfer carbohydrate (ribitol) and the incomplete inhibition of thallus photosynthesis upon treatment with Zn(2+) solutions showed that both photobionts contributed to the photosynthesis of the lichen thallus. Low heterocyst frequencies showed that, despite the presence of adjacent green algae, the cyanobacteria were not altered to increase nitrogen fixation., Conclusions: These cyanobacterial lichens are a tripartite lichen symbiont combination in which the mycobiont has two primarily photosynthetic photobionts, 'co-primary photobionts', a cyanobacterium (dominant) and a green alga. This demonstrates high flexibility in photobiont choice by the mycobiont in the Peltigerales. Overall thallus appearance does not change whether one or two photobionts are present in the cyanobacterial thallus. This suggests that, if there is a photobiont effect on thallus structure, it is not specific to one or the other photobiont.

Facilitative effects and plant-plant interactions are well known for higher plants, but there is a lack of information about their relevance in cryptogams. Additional information about facilitative effects between bryophytes and lichens would be an important contribution to recent research on positive plant-plant interactions, as these can have striking influences not only on the organisation of early successional terrestrial communities but also on succession dynamics by kick-starting ecosystem development through the import of key nutrients. We investigated and quantified these mechanisms between Peltigera rufescens and its associated mosses. Moss-associated thalli had a different morphology that led to several benefits from the association. They had 66% higher net photosynthetic rate and, because the majority of the gas exchange of lichen thalli took place through the lower surface, there was a further increase as the CO(2) concentration was >25% higher beneath moss-associated thalli. Microclimatic measurements showed that mean light levels were substantially lower and temperature extremes slightly ameliorated for moss-associated thalli. As a consequence, desiccation was slower which is, together with an increase in thallus thickness and water storage, the reason for extended periods of optimal net photosynthesis for the moss-associated thalli. All these benefits combined to produce a growth rate of the moss-associated thalli which was significantly higher, twice that of non-associated thalli [0.75 ± 0.4 vs. 0.30 ± 0.1 mm/month (mean ± SD)]. This appears to be the first demonstration of a strong mechanistic basis for facilitative effects between lichens and bryophytes.

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.

Extensive investigations made in the past two decades on lichen photosynthesis in relation to water content have shown two features of particular interest: first, the depression of net photosynthesis at high water contents, suprasaturation (i.e. the lichen contains more water than necessary to saturate photosynthesis), and, second, the ability of green algal lichens to regain photosynthetic activity by uptake of water from humid air. Evidence from several investigators is presented to confirm that both phenomena are now well substantiated through laboratory investigations. It has been questioned whether these features do actually occur in nature and, if they do, to what extent. Recent work is summarized that demonstrates that for many of the lichens studied suprasaturation is of major importance and can result in depressed photosynthesis for around a third of the time that the lichens are photosynthetically active. Reactivation of photosynthesis of green algal lichens by high humidity is also, apparently, very common in some environments, for example, humid temperate rainforests, occurring almost every night. It is possible that the dominance of green algal lichens, rather than cyanobacterial species, in these habitats is a result of their ability to utilize water vapour. If so, then the phenomenon must have major ecological importance for lichen productivity. In general, laboratory studies seem to be able to predict extremely well the behaviour of lichens in their natural habitat.

Summary: Chronosequences at the forefront of retreating glaciers provide information about colonization rates of bare surfaces. In the northern hemisphere, forest development can take centuries, with rates often limited by low nutrient availability. By contrast, in front of the retreating Pia Glacier (Tierra del Fuego, Chile), a Nothofagus forest is in place after only 34 yr of development, while total soil nitrogen (N) increased from near zero to 1.5%, suggesting a strong input of this nutrient.We measured N‐fixation rates, carbon fluxes, leaf N and phosphorus contents and leaf δ15N in the dominant plants, including the herb Gunnera magellanica, which is endosymbiotically associated with a cyanobacterium, in order to investigate the role of N‐fixing and mycorrhizal symbionts in N‐budgets during successional transition.G. magellanica presented some of the highest nitrogenase activities yet reported (potential maximal contribution of 300 kg N ha−1 yr−1). Foliar δ15N results support the framework of a highly efficient N‐uptake and transfer system based on mycorrhizas, with c. 80% of N taken up by the mycorrhizas potentially transferred to the host plant.Our results suggest the symbiosis of G. magellanica with cyanobacteria, and trees and shrubs with mycorrhizas, to be the key processes driving this rapid succession. [ABSTRACT FROM AUTHOR]

Lichens 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. [ABSTRACT FROM AUTHOR]

Within Antarctica there are large gradients both in climate and in vegetation which offer opportunities to investigate links between the two. The activity (% total time active) of lichens and bryophytes in hydric and xeric environments was monitored at Livingston Island (62°39'S). This adds a northern site with a maritime, cloudy climate to previous studies in the southern Antarctic Peninsula and the Dry Valleys (78°S). Annual activity increases northwards from less than 1% to nearly 100%. There is a major and consistent difference between hydric sites which, with snow melt, can be 100% active in summer months even in the Dry Valleys, and xeric sites which, depending on precipitation, rarely exceed 80% activity even at Livingston Island. Mosses dominate hydric sites and lichens the xeric sites all along the gradient. Mean temperatures when active are 2-4°C at all sites, as liquid water is required. Light is a potential major stress reaching 880 µmol m-2 s-1 when active in continental sites. The lack of extremes in temperatures and light combined with high activity levels means that summer at Livingston Island is one of the better sites for lichen and bryophyte growth in the world. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Biological soil crusts (BSC) are the dominant functional vegetation unit in some of the harshest habitats in the world. We assessed BSC response to stress through changes in biotic composition, CO2 gas exchange and carbon allocation in three lichen-dominated BSC from habitats with different stress levels, two more extreme sites in Antarctica and one moderate site in Germany. Maximal net photosynthesis (NP) was identical, whereas the water content to achieve maximal NP was substantially lower in the Antarctic sites, this apparently being achieved by changes in biomass allocation. Optimal NP temperatures reflected local climate. The Antarctic BSC allocated fixed carbon (tracked using 14CO2) mostly to the alcohol soluble pool (low-molecular weight sugars, sugar alcohols), which has an important role in desiccation and freezing resistance and antioxidant protection. In contrast, BSC at the moderate site showed greater carbon allocation into the polysaccharide pool, indicating a tendency towards growth. The results indicate that the BSC of the more stressed Antarctic sites emphasise survival rather than growth. Changes in BSC are adaptive and at multiple levels and we identify benefits and risks attached to changing life traits, as well as describing the ecophysiological mechanisms that underlie them. [ABSTRACT FROM AUTHOR]

Antarctica, with its almost pristine conditions and relatively simple vegetation, offers excellent opportunities to investigate the influence of environmental factors on species performance, such information being crucial if the effects of possible climate change are to be understood. Antarctic vegetation is mainly cryptogamic. Cryptogams are poikilohydric and are only metabolically and photosynthetically active when hydrated. Activity patterns of the main life forms present, bryophytes (10 species, ecto- and endohydric), lichens (5 species) and phanerogams (2 species), were monitored for 21 days using chlorophyll a fluorescence as an indicator of metabolic activity and, therefore, of water regime at a mesic (hydration by meltwater) and a xeric (hydration by precipitation) site on Léonie Island/West Antarctic Peninsula (67°36′S). Length of activity depended mainly on site and form of hydration. Plants at the mesic site that were hydrated by meltwater were active for long periods, up to 100 % of the measurement period, whilst activity was much shorter at the xeric site where hydration was entirely by precipitation. There were also differences due to life form, with phanerogams and mesic bryophytes being most active and lichens generally much less so. The length of the active period for lichens was longer than in continental Antarctica but shorter than in the more northern Antarctic Peninsula. Light intensity when hydrated was positively related to the length of the active period. High activity species were strongly coupled to the incident light whilst low activity species were active under lower light levels and essentially uncoupled from incident light. Temperatures were little different between sites and also almost identical to temperatures, when active, for lichens in continental and peninsular Antarctica. Gradients in vegetation cover and growth rates across Antarctica are, therefore, not likely to be due to differences in temperature but more likely to the length of the hydrated (active) period. The strong effect on activity of the mode of hydration and the life form, plus the uncoupling from incident light for less active species, all make modelling of vegetation change with climate a more difficult exercise. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The activity pattern of the moss Hennediella heimii (Hedw.) Zand. was monitored over a period of 18 days during the austral summer season 2000/2001 at the Canada Flush in Taylor Valley, continental Antarctica. Provided with melt water from the massive Canada Glacier, the moss showed a constant potential photosynthetic activity during the entire measurement period. Permanently hydrated, the moss faced high light levels at surprisingly low moss temperatures, which is commonly supposed a deleterious situation for plants. The electron transport rate response of the moss to photosynthetic photon flux densities was linear at all temperatures and did not show a sign of saturation or photoinhibition. H. heimii seems to be well adapted to its environment and tolerates the ambient conditions without apparent harm. This might be due to the fact that mosses can acclimatise to high light conditions by building up highly effective non-photochemical quenching systems. [ABSTRACT FROM AUTHOR]

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 25 °C and gross photosynthesis (GP) calculated (GP = NP + 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. [ABSTRACT FROM AUTHOR]

Carbon dioxide. (CO[Sub2]) exchange of Sticta latifrons Rich. was studied using an infrared gas analyzer. Slight differences in the relationship between net photosynthetic rate (NPR) and water content were found when thalli were dried by using air directed solely onto the upper or lower surfaces. Further studies using a simple split chamber which allowed separate analysis of the CO[Sub2] exchange of upper and lower thallus surfaces demonstrated that the vast majority of CO[Sub2] exchanged through the lower surface. The results provide strong circumstantial evidence for the suggestion that cyphellae act as air pores and the possible importance of such pores in the water relations of the species is discussed. [ABSTRACT FROM AUTHOR]

The gaseous exchange pathways of Sticta latifrons Rich. and Pseudocyphellaria amphisticta Kremp. were examined using both light and scanning electron microscopes. The size and frequency of the pores in the gas exchange structures (cyphellae and pseudocyphellae) and in the medulla were measured and from these CO2 diffusion resistances were calculated. Pseudocyphellae were found to be smaller and more widely spaced than cyphellae, consequently the resistance of the pseudocyphellae, was much greater than that of the cyphellae. Medulla resistances were low in both lichens and are probably unimportant, even at high water contents. No evidence of hyphal swelling was found. Gas exchange structure resistances were more than five-fold greater than medulla resistances. It is suggested that this arrangement of resistances may simultaneously encourage refixation of respired CO2 and maintain a non desiccating environment for the lichen algae. The internal transport resistances calculated in this work approximate experimentally obtained values. [ABSTRACT FROM AUTHOR]

The total resistances to CO2 uptake by Sticta latifrons Rich, and Pseudocyphellaria amphisticta Kremp. were separated into transport and carboxylation components by calculation after transformation of net photosynthesis rate against CO2 concentration curves into a linear form. The use of this technique circumvented the problem of measuring the internal CO2 concentration of the lichen thalli. Both species exhibited an increase in transport resistance at high thallus water contents and an increase in both transport and carboxylation resistances at low water contents. At low and intermediate water contents internal transport resistances were larger than carboxylation resistances when measured at limiting CO2 concentrations. However, at ambient CO2 concentrations carboxylation processes were the dominant factors limiting photosynthesis at all, except the high, water contents. [ABSTRACT FROM AUTHOR]

Many lichens show seriously depressed net photosynthesis (NP) at high thallus water contents due to increased carbon dioxide diffusion resistance through blockage of diffusion pathways by water. The soil lichen Diploschistes muscorum, however, shows no depression and NP is close to maximal even at the highest thallus water content. We investigated whether lichen substances (lecanoric and diploschistesic acids) in the cortex and medulla contributed to this ability to maintain high NP. Dry thalli were extracted with water-free acetone and, after this treatment, were found to be fully viable to the extent of continued growth after replanting in the field. No differences were found in the response of NP to thallus water content between the normal and extracted thalli, in fact the response curves were often nearly identical. Thus, in this species it seems that lichen substances did not maintain the water-free diffusion pathways and some other explanation, possibly structural, needs to be sought. [ABSTRACT FROM AUTHOR]