Influence of sun irradiance and water availability on lichen photosynthetic pigments during a Mediterranean summer

This study was carried out to investigate changes in lichen photosynthetic pigments induced by different combinations of light irradiance and water availability during a Mediterranean summer. To this purpose, thalli of three epiphytic lichens with a markedly different ecology concerning photo-hygrophytism, namely Evernia prunastri (hygro-mesophytic), Flavoparmelia caperata (mesophytic) and Xanthoria parietina (xero-mesophytic), were transplanted for 30 days to N- and S-facing cardinal exposures in central Italy. To investigate the effect of thallus hydration, at each cardinal exposure 50% of thalli were hydrated daily with deionised water. The results showed that changes in the concentrations of photosynthetic pigments are species-specific and consist in a general depression of photosynthetic pigments only in the hygro-mesophytic species E. prunastri. The pattern of photosynthetic pigments was also investigated in spontaneous samples along contrasting aspects. In this case, X. parietina from S-facing slopes, adapted to direct solar influx, showed higher pigment content than N-facing thalli; F. caperata and E. prunastri avoid direct extreme solar radiation and assimilation pigments were influenced more by the shadowing of tree canopies than by the cardinal exposure. The influence of drought on lichen photosynthetic pigments in the Mediterranean area is discussed.     

Temperate rainforest lichens in New Zealand: high thallus water content can severely limit photosynthetic CO2 exchange

CO₂ exchange rate in relation to thallus water content (WC, % of dry weight) was determined for 22 species of lichens, mainly members of the genera Pseudocyphellaria and Sticta, from a temperate rainforest, Urewere National Park, New Zealand. All data were obtained in the field, either using a standard technique in which the lichens were initially wetted (soaked or sprayed, then shaken) and allowed to slowly dry, or from periodic measurements on samples that were continuously exposed in their natural habitat. A wide range of WC was found, with species varying from 357 to 3360% for maximal WC in the field, and from 86 to 1300% for optimal WC for photosynthesis. Maximal WC for lichens, wetted by the standard technique, were almost always much less than the field maxima, due to the presence of water on the thalli. The relationships between CO₂ exchange rate and WC could be divided into four response types based on the presence, and degree, of depression of photosynthesis at high WC. Type A lichens showed no depression, and Type B only a little at maximal WC. Type C had a very large depression and, at the highest WC, CO₂ release could occur even in the light. Photosynthetic depression commenced soon after optimal WC was reached. Type D lichens showed a similar depression but the response curve had an inflection so that net photosynthesis was low but almost constant, and never negative, at higher WC. There was little apparent relationship between lichen genus or photobiont type and the response type. It was shown that high WC does limit photosynthetic CO₂ uptake under natural conditions. Lichens, taken directly from the field and allowed to dry under controlled conditions, had net photosynthesis rates that were initially strongly inhibited but rose to an optimum, before declining at low WC. The limiting effects of high WC were clearly shown when, under similar light conditions, severe photosynthetic depression followed a brief, midday, rain storm. Over the whole measuring period the lichens were rarely at their optimal WC for photosynthesis, being mostly too wet or, occasionally, too dry. Photosynthetic performance by the lichens exposed in the field was similar to that expected from the relationship between the photosynthetic rate and WC established by the standard procedure.     

Dirunal leaf movement, chlorophyll fluorescence and carbon assimilation in soybean grown under different nitrogen and water availabilities

Diurnal heliotropic leaf movements, photosynthetic gas exchange, and the ratio of variable fluorescence to maximum fluorescence (Fv/Fm) of unrestrained and horizontally restrained leaves from soybean (Glycine max cv. Cumberland) plants grown in two different water and two different nitrogen treatments were measured. Leaves of plants grown in low water or low nitrogen availability treatments displayed more pronounced diaheliotropism (solar tracking) in the afternoon and a longer period of paraheliotropism (light avoiding) at midday relative to those of well-watered, high-nitrogen-grown plants. Photosaturated photosynthetic rates and the photon flux required to saturate photosynthesis were reduced by water stress and nitrogen deficiency. Compared to horizontal leaves, irradiance on orienting leaves was nearer to the breakpoint of the photosynthetic light response curve, where photosynthesis is co-limited by ribulose biphosphate regeneration and carboxylation. This would increase the carbon return on investments of nitrogen into photosynthesis. A positive linear relationship between Fv/Fm and quantum yield of photosynthesis was measured. Leaves of low-nitrogen-grown plants had earlier and more prolonged reductions in Fv/Fm at midday compared to leaves of high nitrogen grown plants of the same water treatment. Within the same water and nitrogen treatment, horizontally restrained leaves had lower midday Fv/Fm in relation to orienting leaves. Nitrogen deficiency and water stress enhanced this difference such that horizontally restrained leaves of low water and low nitrogen grown plants had earlier and longer midday depressions in Fv/Fm.     

Arbuscular mycorrhizae improves low temperature stress in maize via alterations in host water status and photosynthesis

The effect of arbuscular mycorrhizal (AM) fungus, Glomus etunicatum, on growth, water status, chlorophyll concentration and photosynthesis in maize (Zea mays L.) plants was investigated in pot culture under low temperature stress. The maize plants were placed in a sand and soil mixture at 25°C for 7 weeks, and then subjected to 5°C, 15°C and 25°C for 1 week. Low temperature stress decreased AM root colonization. AM symbiosis stimulated plant growth and had higher root dry weight at all temperature treatments. Mycorrhizal plants had better water status than corresponding non-mycorrhizal plants, and significant differences were found in water conservation (WC) and water use efficiency (WUE) regardless of temperature treatments. AM colonization increased the concentrations of chlorophyll a, chlorophyll b and chlorophyll a + b. The maximal fluorescence (Fm), maximum quantum efficiency of PSII primary photochemistry (Fv/Fm) and potential photochemical efficiency (Fv/Fo) were higher, but primary fluorescence (Fo) was lower in AM plants compared with non-AM plants. AM inoculation notably increased net photosynthetic rate (Pn) and transpiration rate (E) of maize plants. Mycorrhizal plants had higher stomatal conductance (g_s) than non-mycorrhizal plants with significant difference only at 5°C. Intercellular CO₂ concentration (Ci) was lower in mycorrhizal than that in non-mycorrhizal plants, especially under low temperature stress. The results indicated that AM symbiosis protect maize plants against low temperature stress through improving the water status and photosynthetic capacity.     

Photosynthetic response of <Emphasis Type="Italic">Bangia fuscopurpurea</Emphasis> (Bangiales,Rhodophyta) towards dehydration and hyposalinity

Bangia fuscopurpurea, an important farmed species in China, inhabits upper intertidal zones where it suffers periodical desiccation and salinity stress. However, the physiological response and acclimation mechanism of Bangia to abiotic stress is unknown. Here, the photosynthetic response of B. fuscopurpurea to desiccation and hyposalinity was investigated by using chlorophyll fluorescence measurement. The optimum photosynthetic efficiency of photosystem II (Fv/Fm), photochemical quenching (qP) and the non-photochemical quenching (NPQ) of B. fuscopurpurea thalli maintained at basal level when the absolute water content (AWC) was 32%. As AWC decreased from 32% to 9%, Fv/Fm dropped from 0.62 to 0.1 and NPQ increased from 0.2 to 1.2. No significant change occurred in the mean qP but great standard deviation was present as AWC was 9%. Fv/Fm, qP and NPQ of the thalli with 9% AWC fully recovered after rehydration. That B. fuscopurpurea kept high photosystem II photochemical reactions even when AWC was mere 32% enabled this species to survive extreme air drying at low tide. Fv/Fm and qP dropped while NPQ increased with 1 h of varying hyposaline treatment and they regained the basal levels after 6–24 h treatment. Nine days later, Fv/Fm, qP and NPQ levels of the thalli in 100% freshwater was equal to the control level (0.62, 0.9, 0.1, respectively). The present finding suggested that this alga has high photosynthetic capacity to survive during low tide, even during heavy rainfall. We hope this study would facilitate further investigation on the stress acclimation mechanism of B. fuscopurpurea.     

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.     

Photosynthetic electron transport at low temperatures in the green algal foliose lichens <Emphasis Type="Italic">Lasallia pustulata</Emphasis> and <Emphasis Type="Italic">Umbilicaria hirsuta</Emphasis> affected by manipulated levels of ribitol

In lichens, ribitol is known as a carbon storage compound, an osmotic agens involved effectively in cell compartments protection during dehydration of lichen thalli and as a cryoprotective compound. In our study, we investigated the effect of ribitol on photochemical processes of photosynthesis in foliose lichens [Lasallia pustulata (L.) Mérat., Umbilicaria hirsuta (Sw. ex Westr.) Hoffm.] at low temperature. The effects of three concentrations of ribitol, added externally to thalli segments on several chlorophyll (Chl) fluorescence parameters, were evaluated. The 72 h exposition to 8, 16, and 26 mM ribitol led to a concentration-dependent increase in F_V/F_M, decrease in non-photochemical quenching (NPQ) but no change in quantum yield of photosystem II photochemistry (Φ_PSII) values at −5 °C). At higher temperature (0, +5 °C), no effect of ribitol addition on the photosynthetic parameters was apparent.     

Morphology influences water storage in hair lichens from boreal forest canopies

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

Will climate warming exceed lethal photosynthetic temperature thresholds of lichens in a southern African arid region?

Predicted elevated temperatures and a shift from a winter to summer rainfall pattern associated with global warming could result in the exposure of hydrated lichens during summer to more numerous temperature extremes that exceed their thermal thresholds. This hypothesis was tested by measuring lethal temperature thresholds under laboratory and natural conditions for four epilithic lichen species (Xanthoparmelia austro‐africana, X. hyporhytida, Xanthoparmelia sp., Xanthomaculina hottentotta) occurring on quartz gravel substrates at a hot arid inland site two epigeous lichen species (Teloschistes capensis, Ramalina sp.) occurring on gypsum‐rich topsoil at a warm humid coastal site. Extrapolated lethal temperatures for photosynthetic quantum yield under laboratory conditions were up to 4°C higher for lichens from a dry inland site than those from a humid coastal site. Lethal temperatures extrapolated for photosynthetic quantum yield at a saturating photosynthetic photon flux density of ≥11,000 μmol photons m^?2 s^?1 under natural conditions were up to 6°C higher for lichens from the dry inland site than the more humid coastal site. It is concluded that only under atypical conditions of lichen exposure in a hydrated state to temperature extremes at high midday solar irradiances during summer could lethal photosynthetic thresholds in sensitive lichen species be potentially exceeded, but whether the increased frequency of such conditions with climate warming would lead to increased likelihood of lichen mortality is debatable.     

Effects of ocean acidification on growth and physiology of Ulva lactuca (Chlorophyta) in a rockpool‐scenario

Rising atmospheric CO₂‐concentrations will have severe consequences for a variety of biological processes. We investigated the responses of the green alga Ulva lactuca (Linnaeus) to rising CO₂‐concentrations in a rockpool scenario. U. lactuca was cultured under aeration with air containing either preindustrial pCO₂ (280 μatm) or the pCO₂ predicted by the end of the 21st century (700 μatm) for 31 days. We addressed the following question: Will elevated CO₂‐concentrations affect photosynthesis (net photosynthesis, maximum relative electron transport rate (rETR(max)), maximum quantum yield (Fv/Fm), pigment composition) and growth of U. lactuca in rockpools with limited water exchange? Two phases of the experiment were distinguished: In the initial phase (day 1–4) the Seawater Carbonate System (SWCS) of the culture medium could be adjusted to the selected atmospheric pCO₂ condition by continuous aeration with target pCO₂ values. In the second phase (day 4–31) the SWCS was largely determined by the metabolism of the growing U. lactuca biomass. In the initial phase, Fv/Fm and rETR(max) were only slightly elevated at high CO₂‐concentrations, whereas growth was significantly enhanced. After 31 days the Chl a content of the thalli was significantly lower under future conditions and the photosynthesis of thalli grown under preindustrial conditions was not dependent on external carbonic anhydrase. Biomass increased significantly at high CO₂‐concentrations. At low CO₂‐concentrations most adult thalli disintegrated between day 14 and 21, whereas at high CO₂‐concentrations most thalli remained integer until day 31. Thallus disintegration at low CO₂‐concentrations was mirrored by a drastic decline in seawater dissolved inorganic carbon and HCO₃^?. Accordingly, the SWCS differed significantly between the treatments. Our results indicated a slight enhancement of photosynthetic performance and significantly elevated growth of U. lactuca at future CO₂‐concentrations. The accelerated thallus disintegration at high CO₂‐concentrations under conditions of limited water exchange indicates additional CO₂ effects on the life cycle of U. lactuca when living in rockpools.     

Changes of endogenous ABA and ACC,and their correlations to photosynthesis and water relations in mungbean (Vigna radiata (L.) Wilczak cv. KPS1) during waterlogging

The effects of waterlogging on the dynamics of leaf abscisic acid (ABA) and root 1-aminocyclopropane-1-carboxilic acid (ACC, a precursor of ethylene) contents together with those on photosynthetic rate, leaf water potential and chlorophyll fluorescence were studied in mungbean (Vigna radiata (L.) Wilczak cv. KPS1) plants under greenhouse conditions. Waterlogging reduced the photosynthetic rate and water use efficiency rapidly without any changes of stomatal conductance, transpiration rate and ABA concentrations. Rapid reduction of photosynthetic rate and Fv/Fm ratio of chlorophyll fluorescence without increase of ABA indicates that early reduction of photosynthetic rate may not be related to ABA. In addition, the slower recovery of P, P/T_r and Fv/Fm values than ABA implies that ABA is not completely involved in photosynthetic reduction. Increased concentration of ACC during the waterlogging period and after the end of waterlogging may indicate the involvement of ethylene in photosynthetic reduction through the reduction of PSII activities, although early reduction of photosynthesis could not be explained by ethylene. After 2 days of waterlogging, ABA was increased concomitantly with the rapid reduction of P, T_r and g_s. It may suggest that ABA reduces photosynthesis through some ABA-related reactions, such as stomatal closure.     

Photosynthetic Depression at High Thallus Water Contents in Lichens: Concurrent Use of Gas Exchange and Fluorescence Techniques with a Cyanobacterial and a Green Algal Peltigera Species

Abstract: Lichens, being poikilohydric, have varying thallus water contents (WC) and show a complex interaction between net photosynthesis (NP) and WC. NP can be depressed at low WC (desiccation effects) and, in some species, also at high WC. In the latter case the depression is normally ascribed to increased CO₂ diffusion resistances through water blockage. Recently, an earlier explanation, that the depression at high WC is due to recycling of CO₂ from increased dark respiration processes (DR), has been given renewed prominence. The two explanations were distinguished by the concurrent use of gas exchange and chlorophyll fluorescence techniques to investigate NP: WC relationships in the lichens Peltigera leucophlebia (green algal) and P. neckeri (cyanobacterial). Both species had a distinct optimal WC for NP with depressed values at low and high WC. The maximal quantum yield for both CO₂ fixation (initial slope of light response curves of NP) and photosystem II (fluorescence signals of dark-adapted thalli) was depressed only at low WC and remained high at optimal and greater WC. In contrast, the relative electron transport rate (ETR, derived from fluorescence signals of thalli in the light) tracked NP and was depressed at low and high WC. The depression of both NP and ETR at high WC (not that at low WC) could be prevented by using elevated external CO₂ concentrations. A single, linear relationship was found between all values of gross photosynthesis (NP + DR) and ETR regardless of external CO₂ concentration or WC. Our results show that, for these lichens, the depression in NP at high WC is a real fall in photosynthetic rate of the photobionts and is not due to recycling of CO₂. The removal of the depression in NP and ETR at high WC by using elevated external CO₂ levels allows us to conclude that an additional CO₂ diffusion resistance is present.     

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.     

干旱胁迫对不同葡萄砧木光合特性和荧光参数的影响

干旱胁迫导致葡萄砧木实生苗叶片光合能力下降.在正常供水和轻度干旱下,砧木的P_n 以3309C最高,其次是1103P,420A较低,各砧木的G_s和T_r差异不显著;中度干旱下,则以1103P的P_n最高,3309C最低;而严重干旱胁迫下,1103P的P_n比3309C高出124%,水分利用效率是3309C的1.95倍.干旱胁迫下,3种砧木的共同趋势是可变荧光 (F_o) 升高,最大荧光 (F_m)、实际光能转化效率 (ФPSⅡ）和可变荧光与最大荧光比 (F_v/F_m) 降低,但品种变幅不同.中度干旱使3309C的F _o升高17.1%,F_v/F_m降低了8.5%,而1103P的F_o升高6.8%,F_v/F_m降低了5.8%;严重干旱则使3309C的F_o升高36.2%,F_v/F_m降低了20.1%,而1103P的F_o升高9.9%,F_v/F_m降低了10.2%.干旱胁迫对不同葡萄砧木光合和荧光参数的影响与其抗旱性密切相关,其中F_v/F_m和P_n的相关系数最大（r=0.9883）.     

The influence of water on CO2 exchange in the lichen Parmelia praesignis Nyl.

Net photosynthetic rates for the lichen Parmelia praesignis Nyl. were obtained as a function of 5 light levels, 5 temperature levels, and of water content as thalli dried from saturated conditions. Data are described as second order polynomials in the light, and as saturation curves in the dark. Rates in the light were depressed at high water contents reaching maximal rates between 110% and 180% water content and declining as thalli dried. Physiological parameters were derived from the drying curves to investigate temperature and light interactions. Dark respiration parameters are the maximal rate, the water content where the rate is half-maximal, the water content at which respiration is zero, and the maximal water efficiency. In the light, parameters are the maximal net photosynthetic rate, the water content at the maximal rate, the water compensation point, the maximal water efficiency, and the sensitivity of net photosynthesis to change in water content.Values of half-maximal rate water contents for respiration were found to increase as temperatures increased. The greatest maximal net photosynthetic rate occurred at higher temperatures as the light intensity increased. In the light, maximal water efficiency and the sensitivity to changes in water content were generally maximal at temperatures yielding the greatest maximal net photosynthetic rates.     

Effects of winter cold stress on photosynthesis and photochemical efficiency of PSII of the Mediterranean <Emphasis Type="Italic">Cistus albidus</Emphasis> L. and <Emphasis Type="Italic">Quercus ilex</Emphasis> L.

This study examined the photosynthetic and growth performances of potted plants of Cistus albidus L. and Quercus ilex L. submitted either to natural Mediterranean winter conditions or to mild greenhouse conditions. Plants grown outdoors exhibited lower light and CO₂-saturated CO₂ assimilation rates (A_sat) and apparent quantum yield (_i) than those indoors. Until mid-winter, C. albidus had higher A_sat than Q. ilex, but differences disappeared after a period of severe cold. Maximal photochemical efficiency of PSII (Fv/Fm) measured predawn was higher in C. albidus than in Q. ilex, and decreased throughout the season in outdoor plants. Fv/Fm also decreased at light saturation (A_sat) in both species. Fv/Fm was correlated with photosynthetic capacity and efficiency (quantum yield), but the resulting regression slopes were different between the two species. At the physiological level, C. albidus seemed to cope better with cold stress than Q. ilex. However, winter stress induced reduction of leaf absorptance, increased leaf mass per area, extensive leaf damage and high plant mortality in C. albidus. This suggests that the high performance of C. albidus leaves is not likely to be maintained for long periods of cold stress, and may therefore depend on continuous leaf replacement. Quercus ilex showed a conservative behaviour, with low net assimilation rates but greater leaf and plant survival than C. albidus.     

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.     

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.     

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.     

CHANGES IN WATER ECONOMY IN RELATION TO ANATOMICAL AND MORPHOLOGICAL CHARACTERISTICS DURING THALLUS DEVELOPMENT INPARMELIA ACETABULUM

The influence on uptake and water loss of the structural changes experienced by Parmelia acetabulum during thallus development were investigated. Small specimens were characterized by flat lobes and a thin thallus and cortex. Large specimens appeared strongly rugose, imbricate and irregularly folded, and had a significantly thicker cortex and medulla than small thalli. Maximum water storage capacity did not differ between large and small thalli, although water retention was much higher in large thalli, presumably due to the interaction of structural characteristics and a higher boundary layer resistance. This translated into a longer duration of the period of thallus hydration in large thalli compared to small thalli. Incubation of thalli in water-vapour-saturated atmospheres induced full recovery of photosynthetic electron transport to the values before desiccation in small thalli but only induced a partial recovery in large thalli. The close correlation between photosynthetic electron transport and net photosynthesis during desiccation found in this species suggested that carbon-fixation activity could be regained to a larger extent by incubation of thalli in water vapour in small compared to large thalli. The higher ability for water vapour uptake of small thalli might allow them to efficiently use small amounts of intermittently available water or periods of high relative humidity. The significance of this differential ability to utilize water is discussed with regard to the known ecological preferences of the species.