Water Status of Green and Blue-green Phycobionts in Lichen Thalli after Hydration by Water Vapor Uptake: Do They Become Turgid?

It is known from previous investigations that dry lichens with green algae are able to recover net photosynthesis through rehydration with water vapor, whereas all blue-green lichens tested so far lack this ability. The REM micrographs of the present study show that the green phycobionts (Trebouxia spec.) of Ramalina maciformis become turgid only after water vapor uptake. In contrast, the blue-green phycobionts (Nostoc spec.) of Peltigera rufescens do not differ in appearance from the dry state, even when the thallus has reached equilibrium with the water vapor-saturated air; they require liquid water for turgidity. It is hypothesized that, after humidity hydration, water content is not sufficient for reestablishment of a functioning osmotic cell system in the blue-green phycobiont.     

Chlorophyll Fluorescence of Lichens Containing Green and Blue-Green Algae During Hydration by Water Vapor Uptake and by Addition of Liquid Water*

The fluorescence yield at room temperature of the lichens Ramalina maciformis and Peltigera rufescens, containing either green or blue-green algae (Cyanobacteria) as phycobionts, has been investigated during rehydration of the dry lichens by water vapor uptake or by wetting with liquid water. In the dry state the fluorescence yield with all reaction centers open, F_o, was low and no variable fluorescence could be induced with both species. Whereas R. maciformis, containing green algae, regained normal fluorescence behavior during water vapor uptake, the photosynthetic apparatus of the blue-green algae-containing P. rufescens stayed inhibited and could be reactivated only by addition of liquid water. During stepwise rehydration at increasing air humidities, a pattern was established for the recovery of the different fluorescence parameters in R. maciformis. At a dry-weight related water content between 30 and 40%, F_o rose sharply. Maximal variable fluorescence yield expressed as (F_v)_m/F_o, strongly increased in the same range of water content and remained constant above a water content of 50%. Non-photochemical fluorescence quenching, q_NP, determined at the end of a period of actinic illumination, decreased with increasing water vapor uptake. While spraying the lichen with liquid water did not induce a further decrease of q_NP, slow dehydration at lowered air humidity led to a minimal value of q_NP at a water content of 65 % indicating optimal photosynthetic rate under these conditions. These results extend the conclusions drawn from earlier gas exchange experiments that blue-green algae-containing lichens are unable to reactivate photosynthesis by water vapor uptake. During a re- and de-hydration cycle, no hysteresis in the hydration dependence of the fluorescence parameters was found. From this and the presence of a stable and low F_o value at prolonged incubation in nearly water vapor saturated air, we conclude that the reactivation of photosynthesis in blue-green algae-containing lichens is not prevented through high diffusion resistances for water.     

Further evidence that activation of net photosynthesis by dry cyanobacterial lichens requires liquid water

Abstract: In contrast to green algal lichens, cyanobacterial species of different families, growth forms and habitats proved to be unable to attain positive net CO₂ assimilation when the dry thalli were treated with air of high relative humidity; they needed liquid water for the reactivation of their photosynthetic apparatus. Identical behaviour is shown by all of the 47 lichen species with cyanobacterial photobionts, from six different genera, studied so far. This suggests a widely distributed, if not general, characteristic of cyanobacterial lichens. The difference in performance between both groups of photobionts was maintained when the lichen thallus was macerated. Furthermore, cultures of Chroococcidiopsis were unable to make use of water vapour hydration for positive net photosynthesis, and were similar in this respect to some free-living aerophilic cyanohacteria tested earlier. Possible physiological implications as well as ecological consequences for water-relation-dependent habitat selection of green-algal and cyanobacterial lichens are discussed.     

Water status related photosynthesis and carbon isotope discrimination in species of the lichen genusPseudocyphellaria with green or blue-green photobionts and in photosymbiodemes

Summary Green lichens have been shown to attain positive net photosynthesis in the presence of water vapour while blue-green lichens require liquid water (Lange et al. 1986). This behaviour is confirmed not only for species with differing photobionts in the genusPseudocyphellaria but for green and blue-green photobionts in a single joined thallus (photosymbiodeme), with a single mycobiont, and also when adjacent as co-primary photobionts. The different response is therefore a property of the photobiont. The results are consistent with published photosynthesis/water content response curves. The minimum thallus water content for positive net photosynthesis appears to be much lower in green lichens (15% to 30%, related to dry weight) compared to blue-greens (85% to 100%). Since both types of lichen rehydrate to about 50% water content by water vapour uptake only green lichens will show positive net photosynthesis. It is proposed that the presence of sugar alcohols in green algae allow them to retain a liquid pool (concentrated solution) in their chloroplasts at low water potentials and even to reform it by water vapour uptake after being dried. The previously shown difference in δ¹³C values between blue-green and green lichens is also retained in a photosymbiodeme and must be photobiont determined. The wide range of δ¹³C values in lichens can be explained by a C₃ carboxylation system and the various effects of different limiting processes for photosynthetic CO₂ fixation. If carboxylation is rate limiting, there will be a strong discrimination of¹³CO₂, at high internal CO₂ partial pressure. The resulting very low δ¹³C values (-31 to-35‰) have been found only in green lichens which are able to photosynthesize at low thallus water content by equilibraiton with water vapour. When the liquid phase diffusion of CO₂ becomes more and more rate limiting and the internal CO₂ pressure decreases, the¹³C content of the photosynthates increases and less negative δ¹³C values results, as are found for blue-green lichens.     

Moisture content and CO2 exchange of lichens. II. Depression of net photosynthesis in Ramalina maciformis at high water content is caused by increased thallus carbon dioxide diffusion resistance

Summary Thalli of Ramalina maciformis were moistened to their maximal water holding capacity, thus, simulating actual conditions following a heavy rainfall. Time courses of net photosynthesis at 17° C and 750 E m^-2 s^-1 light intensity (PAR) were obtained during drying of the thalli. At ambient CO₂ concentrations from 200 to 1,000 ppm, CO₂ uptake of the moist lichens was depressed at high water content. After a certain water loss, net photosynthesis increased to a maximal value and decreased again with further drying of the thalli. The degree of initial depression of photosynthesis decreased with increasing ambient CO₂ concentration, and it was fully absent at 1,600 ppm ambient CO₂. Under these conditions of CO₂ saturation, net photosynthesis remained constant at maximum for many hours and decreased only when substantial amounts of water had been lost. We conclude that the carboxylation capacity of the lichen is not affected by high contents of liquid water. Therefore, the depression of CO₂ uptake of the water saturated lichen at lower (e.g. natural) ambient CO₂ must be due exclusively to increased resistance to CO₂ diffusion from the external air to the sites of carboxylation.     

Effect of low water potential on photosynthesis in intact lichens and their liberated algal components

Earlier experiments (T.D. Brock 1975, Planta124, 13–23) addressed the question whether the fungus of the lichen thallus might enable the algal component to function when moisture stress is such that the algal component would be unable to function under free-living conditions. It was concluded that the liberated phycobiont in ground lichen thalli could not photosynthesize at water potentials as low as those at which the same alga could when it was present within the thallus. However, our experience with lichen photosynthesis has not substantiated this finding. Using instrumentation developed since the mid-1970's to measure photosynthesis and control humidity, we repeated Brock's experiments. When applying “matric” water stress (equilibrium with air of constant relative humidity) we were unable to confirm the earlier results for three lichen species including one of the species,Letharia vulpina, had also been used by Brock. We found no difference between the effects of low water potential on intact lichens and their liberated algal components (ground thallus material and isolated algae) and no indication that the fungal component of the lichen symbiosis protects the phycobiont from the adverse effects of desiccation once equilibrium conditions are reached. The photosynthetic apparatus of the phycobiont alone proved to be highly adapted to water stress as it possesses not only the capability of functioning under extremely low degrees of hydration but also of becoming reactivated solely by water vapor uptake.     

Moisture content and CO2 exchange of lichens

Summary Net photosynthesis (10 klx light intensity, 150 E m^-2 s^-1 PAR) and dark respiration of the lichen Ramalina maciformis at different temperatures are measured in relation to thallus water content. Both first increase with increasing hydration. Dark respiration then remains constant with increased water content until thallus saturation. In contrast, a further increase in water content leads to a depression of net photosynthesis, as shown in previous studies, after a maximum of CO₂ uptake has been attained. However, the extent of this depression depends strongly on temperature. In saturated thalli (160% water content in relation to lichen dry weight) the depression amounts to about 15% and 63% of the maximum unsaturated rate at 5°C and 25°C thallus temperature, respectively. The moisture compensation-point of net photosynthesis is also decisively determined by temperature (for 0°C at 20% water content; for 25°C at 15%), and the water content that allows maximum rates of CO₂ uptake (for 0°C at 80%; for 25°C at less than 40% water content). An electrical analogue of CO₂ exchange in a lichen thallus is presented, and it is suggested that the experimental results may be interpreted in terms of temperature-dependent CO₂ diffusion resistances in imbibed lichen thalli.     

Carbon-dioxide exchange in lichens: determination of transport and carboxylation characteristics

Measurements were made of net rates of CO₂ assimilation in lichens at various ambient concentrations of CO₂ in air and in helox (79% He, 21% O₂). Because of the faster rate of CO₂ diffusion in the pores of lichen thalli when filled with helox than when filled with air, a given net rate of assimilation was achieved at a lower ambient concentration of CO₂ in helox. The differences were used to estimate resistances to diffusion through the gas-filled pore systems in lichens. The technique was first tested with five lichen species, and then applied in a detailed study with Ramalina maciformis, in which gas-phase resistances were determined in samples at four different states of hydration and with two irradiances. By assuming, on the basis of previous evidence, that the phycobiont in R. maciformis is fully turgid and photosynthetically competent at the smallest hydration imposed (equilibration with vapour at 97% relative humidity), and that, with this state of hydration, diffusion of CO₂ to the phycobiont takes place through continuously gas-filled pores, it was possible also to determine both the dependence of net rate of assimilation in the phycobiont on local concentration of CO₂ in the algal layer, and, with the wetter samples, the extents to which diffusion of CO₂ to the phycobiont was impeded by water films. In equilibrium with air of 97% relative humidity, the thallus water content being 0.5 g per g dry weight, the resistance to CO₂ diffusion through the thallus was about twice as large as the resistance to CO₂ uptake within the phycobiont. Total resistance to diffusion increased rapidly with increase in hydration. At a water content of 2 g per g it was about 50 times as great as the resistance to uptake within the phycobiont and more than two-thirds of it was attributable to impedance of transfer by water. The influences of water content on rate of assimilation at various irradiances are discussed. The analysis shows that the local CO₂ compensation concentration of the phycobiont in R. maciformis is close to zero, indicating that photorespiratory release of CO₂ does not take place in the alga, Trebouxia sp., under the conditions of these experiments.Symbols and Units rate of CO₂ diffusion in air relative to that in carrier gas (unity if the carrier gas is air and 0.43 if is helox) - A₁ net rate of CO₂ uptake by the lichen - A_p gross rate of carboxylation minus photorespiratory decarboxylation in the phycobiont, i.e. net rate of light-activated CO₂ exchange - A_* maximum, CO₂-saturated magnitude of A_p - c concentration of CO₂ - c_a ambient concentration of CO₂ - c_i c_a minus difference in CO₂ concentration across air-filled pore space in the thallus - c₈ CO₂ concentration equivalent to partial pressure of CO₂ at the surface of the phycobiont - ₁ magnitude of c_a at which A₁ = 0 - _* magnitude of c_* at which A_p = 0 - R rate of dark respiration in the lichen (mycobiont and phycobiont) - R rate of dark respiration in region between the surface of the lichen and an arbitrary distance from the surface within the thallus - r resistance to CO₂ transfer from lichen surface to the surface of the phycobiont - r resistance to CO₂ transfer between effective source of dark respiration in the lichen and the surface of the phycobiont - r_g, r _g components of r and r, respectively, attributable to transfer in air-phase - r_w, r _w components of r and r, respectively, attributable to transfer in water-phase - r component of r between surface of lichen and an arbitrary distance from the surface within the thallus - r_* resistance to CO₂ transfer and carboxylation in the phycobiont - RH relative humidity     

Intraspecific variation in photosynthetic responses of trebouxioid lichens with reference to the activity of a carbon-concentrating mechanism

The photosynthetic responses of a range of trebouxioid lichens were investigated to determine whether variations in net assimilation rates shown by populations of the same species collected from different habitats could be correlated with adjustments in carbon-concentrating mechanism (CCM) activity. The activity of a CCM was inferred from the high affinity for CO₂ [i.e. low CO₂ compensation point (Γ); low external CO₂ concentration at which half-maximal assimilation rates are reached (K _{0.5 CO2})], the release of a pool of accumulated dissolved inorganic carbon (C_i) during light/dark transient measurements of CO₂ exchange and values for carbon isotope discrimination intermediate between those characteristic of C₃ and C₄ terrestrial plants. Higher net and gross assimilation rates were expressed by lichens collected from shaded woodland habitats. The higher rates were not accounted for by variations in chlorophyll content. Lichens with high assimilation rates also showed an increased affinity for CO₂ as demonstrated by low CO₂ compensation points and K _0.5 values and the magnitude of the C_i pool accumulated upon illumination and released after darkening of the thalli. However, there was no correlation between assimilation rates and organic matter or instantaneous carbon isotope discrimination measurements, with the latter remaining roughly consistent whatever the provenance or species of the lichen material. The data are discussed with reference to significant environmental factors which are likely to control photosynthesis in the habitats studied. Received: 5 April 1997 / Accepted: 9 September 1997     

Measurement of photorespiration in algae

The rates of true and apparent photosynthesis of two unicellular green algae, one diatom and four blue-green algae were measured in buffer at pH 8.0 at subsaturating concentrations of dissolved inorganic carbon (13-27 micromolar). Initial rates of depletion from the medium of inorganic carbon and ¹⁴C activity caused by the algae in a closed system were measured by gas chromatography and by liquid scintillation counting, respectively. The rate of photorespiration was calculated as the difference between the rates of apparent and true photosynthesis. The three eucaryotic algae and two blue-green algae had photorespiratory rates of 10 to 28% that of true photosynthesis at air levels of O₂. Reduction of the O₂ level to 2% caused a 52 to 91% reduction in photorespiratory rate. Two other blue-green algae displayed low photorespiratory rates, 2.4 to 6.2% that of true photosynthesis at air levels of O₂, and reduction of the O₂ concentration had no effect on these rates.     

The complexity of symbiotic interactions influences the ecological amplitude of the host: A case study in Stereocaulon (lichenized Ascomycota)

Symbiosis plays a fundamental role in nature. Lichens are among the best known, globally distributed symbiotic systems whose ecology is shaped by the requirements of all symbionts forming the holobiont. The widespread lichen‐forming fungal genus Stereocaulon provides a suitable model to study the ecology of microscopic green algal symbionts (i.e., phycobionts) within the lichen symbiosis. We analysed 282 Stereocaulon specimens, collected in diverse habitats worldwide, using the algal ITS rDNA and actin gene sequences and fungal ITS rDNA sequences. Phylogenetic analyses revealed a great diversity among the predominant phycobionts. The algal genus Asterochloris (Trebouxiophyceae) was recovered in most sampled thalli, but two additional genera, Vulcanochloris and Chloroidium, were also found. We used variation‐partitioning analyses to investigate the effects of climatic conditions, substrate/habitat characteristic, spatial distribution and mycobionts on phycobiont distribution. Based on an analogy, we examined the effects of climate, substrate/habitat, spatial distribution and phycobionts on mycobiont distribution. According to our analyses, the distribution of phycobionts is primarily driven by mycobionts and vice versa. Specificity and selectivity of both partners, as well as their ecological requirements and the width of their niches, vary significantly among the species‐level lineages. We demonstrated that species‐level lineages, which accept more symbiotic partners, have wider climatic niches, overlapping with the niches of their partners. Furthermore, the survival of lichens on substrates with high concentrations of heavy metals appears to be supported by their association with toxicity‐tolerant phycobionts. In general, low specificity towards phycobionts allows the host to associate with ecologically diversified algae, thereby broadening its ecological amplitude.     

Preliminary comparison of atmospheric CO2 enhancement to photosynthesis of Pyropia yezoensis (Bangiales,Rhodophyta) leafy thalli and filamentous thalli

Intertidal macroalgae are submerged in seawater at high tide and exposed to air at low tide. When they are exposed to the air, CO₂ is the main inorganic carbon source. In this study, the photosynthetic performances of PSI and PSII were measured in different generations of Pyropia yezoensis (leafy thalli and filamentous thalli) that had been exposed to air containing different CO₂ concentrations. Changes in photosynthesis during dehydration and salt treatment under the different CO₂ concentrations were also analyzed. The results showed that in leafy thalli, the effective photochemical quantum yield of PSII (YII) was enhanced as CO₂ increased, which suggested that CO₂ assimilation was enhanced and that they can utilize CO₂ in the air directly, even when they are subjected to moderate stress. These findings could explain why, in P. yezoensis aquaculture, moderate exposure to air does not lead to a decrease in crop yield. However, in filamentous thalli, there were no significant differences in YII at the CO₂ concentrations tested. The expression of genes involved in the Calvin cycle in leafy thalli was higher than that in filamentous thalli. CO₂ uptake and biomass of P. yezoensis leafy thalli is larger than filamentous thalli, which may be due to its different carbon utilization mechanism and the adaptation of intertidal environment in the evolutionary process.     

Photosynthetic activity and proteomic analysis highlights the utilization of atmospheric CO2 by Ulva prolifera (Chlorophyta) for rapid growth

Free‐floating Ulva prolifera is one of the causative species of green tides. When green tides occur, massive mats of floating U. prolifera thalli accumulate rapidly in surface waters with daily growth rates as high as 56%. The upper thalli of the mats experience environmental changes such as the change in carbon source, high salinity, and desiccation. In this study, the photosynthetic performances of PSI and PSII in U. prolifera thalli exposed to different atmospheric carbon dioxide (CO₂) levels were measured. Changes in photosynthesis within salinity treatments and dehydration under different CO₂ concentrations were also analyzed. The results showed that PSII activity was enhanced as CO₂ increased, suggesting that CO₂ assimilation was enhanced and U. prolifera thalli can utilize CO₂ in the atmosphere directly, even when under moderate stress. In addition, changes in the proteome of U. prolifera in response to salt stress were investigated. Stress‐tolerance proteins appeared to have an important role in the response to salinity stress, whereas the abundance of proteins related to metabolism showed no significant change under low salinity treatments. These findings may be one of the main reasons for the extremely high growth rate of free‐floating U. prolifera when green tides occur.     

Effect of high light on the efficiency of photochemical energy conversion in a variety of lichen species with green and blue-green phycobionts

Exposure to high light induced a quantitatively similar decrease in the rate of photosynthesis at limiting photon flux density (PFD) and of photosystem II (PSII) photochemical efficiency, F_V/F_M, in both green and blue-green algal lichens which were fully hydrated. Such depressions in the efficiency of photochemical energy conversion were generally reversible in green algal lichens but rather sustained in blue-green algal lichens. This greater susceptibility of blue-green algal lichens to sustained photoinhibition was not related to differences in the capacity to utilize light in photosynthesis, since the light-and CO₂-saturated rates of photosynthetic O₂ evolution were similar in the two groups. These reductions of PSII photochemical efficiency were, however, largely prevented in lichen thalli which were fully desiccated prior to exposure to high PFD. Thalli of green algal lichens which were allowed to desiccate during the exposure to high light exhibited similar recovery kinetics to those which were kept fully hydrated, whereas bluegreen algal lichens which became desiccated during a similar exposure exhibited greatly accelerated recovery compared to those which were kept fully hydrated. Thus, green algal lichens were able to recover from exposure to excessive PFDs when thalli were in either the hydrated or desiccated state during such an exposure, whereas in blue-green algal lichens the decrease in photochemical efficiency was reversible in thalli illuminated in the desiccated state but rather sustained subsequent to illumination of thalli in the hydrated state.Abbreviations and Symbols F_o yield of instantaneous fluorescence - F_M maximum yield of fluorescence induced by pulses of saturating light - F_V variable yield of fluorescence - PFD photon flux density (400–700 nm) - PSII photosystem II This work was supported by the Deutsche Forschungsgeneinschaft (Forscherguppe Ökophysiologic and Sonderforschungsbereich 251 of the University of Würzburg) and the Fonds der Chemischen Industrie. W.W.A. gratefully acknowledges the support of a fellowship from the Alexander von Humboldt Foundation. We thank Professor T.G.A. Green for identifying and supplying all of the New Zealand lichen material and Professor F.-C. Czygan for advice concerning the chlorophyll analyses which were performed by Johanna Leisner.     

Changes in gas exchange characteristics and water use efficiency of mangroves in response to salinity and vapour pressure deficit

Summary Measurements were made of the photosynthetic gas exchange properties and water use efficiency of 19 species of mangrove in 9 estuaries with different salinity and climatic regimes in north eastern Australia and Papua New Guinea. Stomatal conductance and CO₂ assimilation rates differed significantly between species at the same locality, with the salt-secreting species, Avicennia marina, consistently having the highest CO₂ assimilation rates and stomatal conductances. Proportional changes in stomatal conductance and CO₂ assimilation rate resulted in constant and similar intercellular CO₂ concentrations for leaves exposed to photon flux densities above 800 mol·m^-2·s^-1 in all species at a particular locality. In consequence, all species at the same locality had similar water use efficiencies. There were, however, significant differences in gas exchange properties between different localities. Stomatal conductance and CO₂ assimilation rate both decreased with increasing salinity and with increasing leaf to air vapour pressure deficit (VPD). Furthermore, the slope of the relationship between assimilation rate and stomatal conductance increased, while intercellular CO₂ concentration decreased, with increasing salinity and with decreasing ambient relative humidity. It is concluded from these results that the water use efficiency of mangroves increases with increasing environmental stress, in this case aridity, thereby maximising photosynthetic carbon fixation while minimising water loss.Contribution No. 459 from the Australian Institute of Marine Science     

Über Cyanotrophie bei Flechten

In contrast to the well-known blue-green algal containing lichens several green algal containing lichens, belonging to very different genera, show regular connections to free-living or ± lichenized blue-green algae, mainlyStigonema. Most of these lichens have squamulose thalli. This lichen-algal relationship, regarded as cyanotrophy, may be either facultative or obligate. Some of the species occur only on very poor, acidic rocks onStigonema, while they occur independent ofStigonema in high nutrient biotops. Obligate species cover the blue-green algae with hyphae and some of these species cover the algae so extensively that one can call these connections paracephalodia. — Two species and one variety are new to science from the mainly Himalayan genusBryonora. They occur in high elevations in Nepal and are cyanotrophic.Bryonora selenospora has thick, halfmoon-shaped to slightly twisted ascospores. The other two new taxa areB. reducta andB. rhypariza var.cyanotropha. There are several other cyanotrophic lichen taxa besides the ones described here. They will be introduced at a later occasion.

Frau Prof. Dr.Elisabeth Tschermak-Woess zu ihrem 70. Geburtstag gewidmet.     

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

Structural alterations of the photobiont and mycobiont cells of lichens have been related to CO₂-gas exchange during experiments involving water vapour uptake and desiccation of liquid-water-saturated thalli. Increasing water vapour uptake of air dry lichens led to a gradual unfolding of the photobiont cells in Lobaria pulmonaria, Pseudevernia furfuracea, Ramalina maciformis and Teloschistes lacunosus as studied by low-temperature scanning electron microscopy. The data indicated that globular, probably turgid, cells and also slightly infolded or even heavily collapsed cells contributed to positive net photosynthesis, which was reached after water vapour uptake by the four species studied. During desiccation of fully water-saturated thalli of L. pulmonaria, extrathalline water films gradually evaporated before maximum values of CO₂-gas exchange were measured and before photobiont cells started to shrivel. In contrast, in P. furfuracea the CO₂-gas exchange maximum was reached when a considerable percentage of photobiont cells had already collapsed and while other parts of the thalli were still covered with liquid water. Further desiccation led to cavitation of the cortical cells in both species, this occurring at water contents at which net photosynthesis was still positive.Abbreviations EF exoplasmic fracture face - LTSEM low-temperature scanning electron microscopy - NP net photosynthesis - PAR photosynthetic active radiation (400–700 nm) - PF plasmic fracture face We thank D. Pichier, P. Hatvani, H. Müller, Birmensdorf, and J.B. Winkler, Kiel, for technical assistance, and J. Innes, Birmensdorf, for correcting the English text. Stimulating discussion with R. Honegger (Institut für Pflanzenbiologie, Universität Zürich, Switzerland), L. Kappen (Botanisches Institut, Universität Kiel, Germany), T.G.A. Green (Department of Biological Sciences, Hamilton, New Zealand), and O.L. Lange (Julius-von-Sachs-Institut für Biowissenschaften, Universität Würzburg, Germany) are gratefully acknowledged.     

STEM AND LEAF GAS EXCHANGE OF TWO ARID LAND SHRUBS

Gas exchange studies were conducted on two shrub species found in cool shrub-steppe communities of the American West, big sagebrush (Artemisia tridentata subsp. tridentata Nutt.) and broom snakeweed (Gutierrezia sarothrae [Pursh] Britt. and Rusby), with a goal of evaluating characteristics and relative contributions of green stem and leaf material to total shoot CO₂ exchange at different temperatures. Variations in tissue temperature exerted a pronounced effect on CO₂ exchange—net photosynthesis and dark respiration—of green stems and leaves of both species. Definite temperature optima of net photosynthesis were noted, and dark respiration rates consistently increased with increases in temperature. Green stems of both species exhibited sizable dark respiration rates, although stem rates at all temperatures were lower than corresponding leaf rates. Artemisia tridentata did not exhibit sizeable green stem net photosynthesis even under conditions of optimal temperature and water availability, and leaf net photosynthesis rates were much lower than those of G. sarothrae. However, A. tridentata in general possessed a greater leaf biomass than G. sarothrae. Green stems of G. sarothrae exhibited considerable rates of net photosynthesis under both optimal and sub-optimal temperature and water availability conditions. A higher optimum temperature of net photosynthesis was noted for stems than for leaves of G. sarothrae. The adaptive significance of these interspecific differences in CO₂ exchange characteristics is discussed.     

Water relations and CO2 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 CO₂ 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 CO₂ 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 CO₂ 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.     

Rates of photosynthesis of two lichens and of their isolated algal components

Abstract

The rate of photosynthesis of two lichen species (Peltigera leucophlebia and Ramalina farinacea) was found to be 30 to 40% that of spinach leaf dises and 20% that of the free-living alga Chlorella when the results were expressed on a per mg chlorophyll basis. When the algae were isolated from the thalli, the rate of photosynthesis per mg chlorophyll increased for Ramalina farinacea and decreased for Peltigera leucophlebia. Product analysis indicated that the products of photosynthesis depended on the association of the alga with the fungus: algae isolated from the thalli showed a «shift» in products from sugars and sugar alcohols. to compounds such as organic acids. The results suggest that a symbiotic relationship with a fungus alters both the rate and products of algal photosynthesis.