Do secondary substances in the thallus of a lichen promote CO2 diffusion and prevent depression of net photosynthesis at high water content?

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. Received: 5 April 1997 / Accepted: 26 April 1997     

Light use efficiency of dry matter gain in five macro-lichens: relative impact of microclimate conditions and species-specific traits

Relations between irradiance (I) and lichen growth were investigated for five macro‐lichens growing at two sites in Sweden. The lichens represented different mycobiont–photobiont associations, two morphologies (foliose, fruticose) and two life forms (epiphytic, terricolous). The lichens were transplanted at two geographically distant sites in Sweden (1000 km apart) from Sept 1995 to Sept 1996 in their typical microhabitats, where microclimate and growth were followed. Between April/May and Sept 96, the terricolous species had a dry matter gain of 0·2 to 0·4 g (g DW)^–1 and the epiphytes 0·01 to 0·02 g (g DW)^–1. When related to area, growth amounted to 30 to 70 g m^?2 for the terricolous species and to 1 to 4 g m^?2 for the epiphytes. There was a strong correlation between growth and intercepted irradiance when the lichens were wet (I_wet), with 0·2 to 1·1 g lichen dry matter being produced per MJ solar energy. Across the 10 sets of transplants, light use efficiencies of dry matter yield (e) ranged between 0·5 and 2%, using an energy equivalent of 17·5 kJ g^?1 of lichen dry matter. The higher productivity of the terricolous species was due to longer periods with thallus water contents sufficient for metabolic activity and because of the higher mean photon flux densities of their microhabitat. A four‐fold difference in photosynthetic capacity among the species was also important. It is concluded that lichen dry matter gain was primarily related to net carbon gain during metabolically active periods, which was determined by light duration, photon flux density and photosynthetic capacity.     

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.     

In situ studies of water relations and CO2 exchange of the tropical macrolichen, Sticta tomentosa

Diel (24-h) time courses of CO₂ exchange, water relations, and microclimate of the foliose lichen, Sticta tomentosa (Swartz) Ach., and responses to experimentally manipulated conditions were measured at a forest edge in a lower montane rainforest in Panama.
Similar to earlier observations on two other rain forest lichens, daily desiccation suppressed net photosynthesis (NP) during the period when irradiation was highest. Not surprisingly, the light response curves of NP showed saturation at rather low light levels. Rehydration was associated with an initial resaturation burst of short duration, which could be demonstrated both under natural conditions and experimentally. This additional loss of CO₂ seems too low to be ecologically relevant. Moreover, high thallus hydration was also detrimental to NP: at maximum water content net CO₂ uptake was depressed by >50%. Although NP was well adapted to the prevailing high temperatures, the latter also stimulated dark respiration. On average, almost 60% of the diurnal carbon gain was lost during the night.
In spite of these limitations, the integrated 24-h C gain was quite high, on average 0·5% of the thallus C content. Whilst these figures were determined for horizontally exposed samples, we also assessed the role of different exposures on photosynthetic performance. Diel C gain was highest under conditions of semi-shade (westerly exposure), which allowed long periods of activity, whilst much higher irradiance at other exposures could not be utilized for photosynthetic production: easterly exposed thalli dried out even faster than horizontally exposed samples.     

POROSIMETRIC STUDY OF THE LICHEN FAMILY UMBILICARIACEAE: ANATOMICAL INTERPRETATION AND IMPLICATIONS FOR WATER STORAGE CAPACITY OF THE THALLUS

The pore system within the thalli of 13 lichen taxa belonging to the family Umbilicariaceae has been studied by means of mercury intrusion porosimetry. A characteristic bimodal pore size distribution with a central depression around 0.05 μm of equivalent pore radius was obtained in all lichen samples. However, clear differences were found among the pore size distributions of each lichen taxa. The total thallus porosity was undoubtedly related to the anatomy of the medulla. In general, a radial plectenchymatic medulla conferred larger porosity to the thallus than an arachnoidal one. Maximum thallus water content closely depended on the total thallus porosity in the five lichens possessing rhizinomorphs. The species with a similar type of medulla could be grouped together in a multivariate analysis that considered three porosimetric parameters and the maximum thallus water content. Umbilicaria cinereorufescens was the most distinct species, with the lowest values of total porosity and water storage capacity and the largest value of thallus density, apparently due to its scleroplectenchymatic medulla. The pore size distribution existing inside the thallus of the species studied is discussed in relation to the often opposing problems of CO₂ exchange and optimal water relations. Some results pointed to a large influence of the micropores (<0.05 μm) on the water storage capacity of the thallus, while the macropores would have a more important role in gas exchange.     

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.     

Leaf movement of bush bean: a biometeorological perspective

 Leaf movements of bush bean plants were studied at the relatively low photon flux density of 0.2 mmol/m² per s, and air temperatures of 25° and 35° C in a growth chamber. A beta-ray gauge system was used to monitor continuously pulvinus water status and bending. Leaf angles were below the horizontal and were linearly related to the soil water content (R≥−0.91 at 25° C and R≥−0.93 at 35° C). The beta-ray transmission maxima coincided with the stem temperature minima in darkness and vice versa when brightness prevailed as the growth chamber temperature varied with the photoperiod. Leaf angle increased linearly with increased beta-ray transmission. The Q₁₀ temperature coefficient, a measure of the metabolic energy requirement for leaf movement between 25° and 35° C was estimated at 1.8, and the corresponding mean Arrhenius constant at 423 kJ/mol for bush bean. Received: 19 July 1996 / Accepted: 9 September 1996     

不同水分条件下两种叶状地衣的光合活性对冻融交替的响应

冻融交替是我国北方常见气候现象,其对地衣光合作用的影响尚不清楚。研究了采自雾灵山的卷叶点黄梅Flavopunctelia soredica和平盘软地卷Peltigera elisabethae的光合活性(以净光合速率表示,net photosynthetic rate,Pn)对冻融处理的响应及其与地衣体含水量(干冻组:含水量<20%干重;湿冻组:含水量>200%)和物种的关系。结果显示卷叶点黄梅的干冻组Pn经5次冻融后下降至对照的21%,湿冻组经3次冻融后下降至负值,平盘软地卷的干冻组和湿冻组在5次冻融后Pn均为负值;相对净光合速率(relative net photosynthetic rate,Rpn)与冻融次数的线性回归分析表明,卷叶点黄梅湿冻组的斜率绝对值(58.06)>平盘软地卷湿冻组(41.01)>平盘软地卷干冻组(32.27)>卷叶点黄梅干冻组(11.44)。结果表明冻融胁迫可显著抑制两种地衣的光合活性,这种抑制作用具有物种差异且和地衣体内水分含量有关:水分含量的增高将增强冻融胁迫对地衣光合活性的抑制作用;干燥状态下,卷叶点黄梅的低温耐性远高于平盘软地卷,但在湿润状态下则低于后者。两种地衣对冻融循环的光合响应的物种差异与其微生境气候有关:生长于较干燥开阔地带的卷叶点黄梅与生于阴湿生境中的平盘软地卷相比,可能已形成了更强的低温干燥适应能力,其低温湿润适应能力则较弱。全球气候变化可能会通过冻融事件的时空格局的改变而对地衣的光合作用和分布造成负面影响。     

Acidobacteria in microbial communities of the bog and tundra lichens

Bacterial communities of the lichens from a Sphagnum bog (Karelia) and tundra (Vorkuta oblast) were investigated. Members of the phylum Acidobacteria were numerous in the thallus of living and decaying lichens (3.8 × 10⁸ cells/g), constituting 6 to 32% of the total bacterial number. Pure cultures of acidobacteria were isolated from the samples of living and decaying lichen thallus. Ten of them were identified and classified as members of subgroup 1 of the Acidobacteria. The hydrolytic activity of two strains isolated from the living and decomposing zones of the thallus was investigated. They were capable of growth on xylan, starch, pectin, laminarin, and lichen extract. Acidobacteria were shown to be a stable and numerous component of microbial communities of the bog and tundra lichens.     

Edge and land-use effects on epiphytic lichen diversity in the forest-steppe ecotone of the Mongolian Altai

Epiphytic lichen diversity was studied in forests of Siberian larch (Larix sibirica) in the forest-steppe ecotone of the Mongolian Altai. These forests are utilized for livestock grazing, fuelwood collection and occasional logging by pastoral nomads. The density of nomad households in the proximity of the forests influences epiphytic lichen diversity more strongly than the position of sample trees in the forest interior or at the forest line to the steppe. This suggests that land use exerts a stronger effect on lichen diversity than the distinct gradient in microclimate between the forest interior and the forest edge. The co-occurrence of nitrophytes with anitrophytic acidophytes on a small spatial scale as well as higher N and Ca concentrations and pH values in the bark of larch trees at the forest edge than in the forest interior indicate that moderate livestock grazing increases the epiphytic lichen diversity due to an increase of the diversity of chemically different microhabitats. Preference of many lichen species (and of rare species in particular) for overmature and decaying trees suggests that logging and fuelwood collection has adverse effects on epiphytic lichen diversity. This adverse effect is likely to be more crucial for lichen diversity than the putatively positive effect of livestock grazing, since more lichens with a preference for old and decaying trees than for nitrogen-enriched bark were found. The present study is the first one investigating the impact of pastoral nomadism in Central Asia on epiphyte diversity.     

亚热带山地森林附生地衣的移植生长及其对环境变化的响应

本研究选取云南哀牢山亚热带森林系统6种常见的附生大型地衣进行为期2年的移植实验,分析其在原生林林外、林缘以及林内3种生境下的生物量增长速率和健康率等的差异,并探讨其生长对环境因子变化的响应。结果显示,网肺衣Lobaria retigera和平滑牛皮叶Sticta nylanderiana在林缘处生物量增长最快;而槽枝Sulcaria sulcata、多花松萝Usnea florida、皮革肾岛衣Nephromopsis pallescens、针芽肺衣Lobaria isidiophora 4种地衣则都是在林外光照更强、湿度更低的条件下生长得最好,林内最差。除网肺衣外,所有地衣的健康率均在林外较好,林缘次之,林内最差。非参数相关分析表明,6种地衣的生长速率和光照以及温度呈正相关,与大气湿度呈负相关;温度和光照是影响地衣生长最主要的生境因子。     

Field photosynthetic activity of lichens in the Windmill Islands oasis, Wilkes Land, continental Antarctica

In order to ascertain whether the major species of continental antarctic macrolichens are photosynthetically active during summer conditions, the chlorophyll fluorescence of three lichen species [Umbilicarin decussata (Vill. Zahlbr., Pseudephebe minuscula (Ny-l. ex Arnold) Brodo and Hawksw. and Usnea sphacetala R. Br.] was monitored in the vicinity of Casey Station. Wilkes Land, continental Antarctica using a PAM-2000 modulated fluorescence system. Lichens were studied when in equilibrium with the atmosphere as well as when moistened by snow showers. Photochemical quantum yield was estimated as ΔF/F′_m and related to thallus water content as well as microclimatic conditions. Lichens were photosynthetically active only when moistened by snow fall or by run-off from snow melt. The levels of photosynthetic activity in the field for all species were influenced by microenvironmental conditions and patterns in response were site and species specific. Highest levels of photosynthetic efficiency occurred when thalli were at intermediate water contents. Photosynthetic activity was reduced by cold as well as warm, bright conditions. Highest thallus water contents occurred during the middle of the day after substantial “falls of snow. P. minuscula maintained highest thallus water contents at all sites and appears to have a high water compensation point which is related to its observed distribution patterns. Umbilicaria decussata studied in the laboratory did not become photosynthetically active even when exposed to 95% relative humidity (RH) for 51 h and. when dehydrating after artificial wetting, showed an optimum thallus water content for photosynthesis of ca 90% dry weight and a thallus water compensation point of about 35% dry weighl. In the field U. decussata did not become pholosynthetically active except when moistened by snow. Usnea sphacelata exposed to the atmosphere had low thallus water contents (ca 30%) which was not related to RH. The results indicate that the lichens are photosynthetically inactive for most of the summer period and are totally reliant on snow as a water supply. This i.s important when modelling carbon gain and growth rates of continental antarctic lichens.     

The effects of gap size on some microclimate variables during late summer and autumn in a temperate broadleaved deciduous forest

The creation of gaps can strongly influence forest regeneration and habitat diversity within forest ecosystems. However, the precise characteristics of such effects depend, to a large extent, upon the way in which gaps modify microclimate and soil water content. Hence, the aim of this study was to understand the effects of gap creation and variations in gap size on forest microclimate and soil water content. The study site, in North West England, was a mixed temperate broadleaved deciduous forest dominated by mature sessile oak (Quercus petraea), beech (Fagus sylvatica) and ash (Fraxinus excelsior) with some representatives of sycamore (Acer pseudoplatanus). Solar radiation (I), air temperature (T _A), soil temperature (T _S), relative humidity (h), wind speed (v) and soil water content (Ψ) were measured at four natural treefall gaps created after a severe storm in 2006 and adjacent sub-canopy sites. I, T _A, T _S, and Ψ increased significantly with gap size; h was consistently lower in gaps than the sub-canopy but did not vary with gap size, while the variability of v could not be explained by the presence or size of gaps. There were systematic diurnal patterns in all microclimate variables in response to gaps, but no such patterns existed for Ψ. These results further our understanding of the abiotic and consequent biotic responses to gaps in broadleaved deciduous forests created by natural treefalls, and provide a useful basis for evaluating the implications of forest management practices.     

Growth of Corynebacterium glutamicum in ammonium- and potassium-limited continuous cultures under high osmotic pressure

 In order to determine the possible effect of nutrient limitations on the response of Corynebacterium glutamicum to a saline osmotic up-shock, the bacteria were grown in continuous cultures, at osmotic pressures of 0.4 osmol/kg and 1.2 osmol/kg, under ammonia and potassium limitation. At the low osmolality of 0.4 osmol/kg, the glutamate and proline levels of 15 mg/g and 5 mg/g dry weight respectively were lower than previously reported in glucose-limited continuous cultures (50 mg/g and 10 mg/g dry weight respectively). On the other hand, the internal trehalose pool was much higher at 40 mg/g dry weight. When the medium osmolality was increased to 1.2 osmol/kg by NaCl addition, under ammonia limitation, the proline content rose from 5 mg/g to 20 mg/g dry weight and the trehalose content from 40 mg/g to 70 mg/g dry weight, whereas the intracellular pool of glutamate remained essentially constant. An increase in the internal sodium content was also observed. Similar results were found for the internal pool of glutamate, proline and trehalose when C. glutamicum was grown under potassium limitations at an osmolality of 1.2 osmol/kg. There were also higher levels of sodium ions, glutamine and alanine. According to the present results, whereas proline was previously reported to be the dominantly accumulated osmoprotectant in C. glutamicum grown under glucose limitations, under ammonia and potassium limitations trehalose represented the dominantly synthesized metabolite. Received: 19 December 1995/Received revision: 9 April 1996/Accepted: 15 April 1996     

Lichenizing Rhizomorphs and Thallus Development in the Squamulose Lichen Aspicilia crespiana Rico ined. (Lecanorales,Ascomycetes)

This work describes the rhizomorphs and their relation to thallus development in the squamulose lichen Aspicilia crespiana Rico ined. The rhizomorphs capture algal cells, giving rise to new squamules terminally or laterally. The thallus thus consists of a network of lichenized squamules interlinked by mycobiontic rhizomorphs. A previously proposed comparison of lichen rhizomorphs to the prothallus (hypothallus) of crustose lichens is applied and expanded upon. Other less organized prothallic structures sometimes produced by A. crespiana are also described. The importance of lichenizing rhizomorphs as an essential feature of thallus growth in this species is emphasized and a competitive role in substrate occupation and thallus expansion is suggested.     

Growth of foliose lichens: a review

This review considers various aspects of the growth of foliose lichens including early growth and development, variation in radial growth rate (RaGR) of different species, growth to maturity, lobe growth variation, senescence and fragmentation, growth models, the influence of environmental variables, and the maintenance of thallus symmetry. The data suggest that a foliose lichen thallus is essentially a ‘colony’ in which the individual lobes exhibit a considerable degree of autonomy in their growth processes. During development, recognisable juvenile thalli are usually formed by 15 months to 4 years while most mature thalli exhibit RaGR between 1 and 5 mm yr⁻¹. RaGR within a species is highly variable. The growth rate-size curve of a foliose lichen thallus may result from growth processes that take place at the tips of individual lobes together with size-related changes in the intensity of competition for space between the marginal lobes. Radial growth and growth in mass is influenced by climatic and microclimatic factors and also by substratum factors such as rock and bark texture, chemistry, and nutrient enrichment. Possible future research topics include: (1) measuring fast growing foliose species through life, (2) the three dimensional changes that occur during lobe growth, (3) the cellular changes that occur during regeneration, growth, and division of lobes, and (4) the distribution and allocation of the major lichen carbohydrates within lobes.     

Habitat fragmentation impacts on biodiversity of evergreen broadleaved forests in Jinyun Mountains, China

The plant communities and their microclimates were surveyed and observed, and the soil fertilities were determined in six plots of evergreen broadleaved forests of different sizes and similar slope aspects on Jinyun Mountains of Chongqing in China from April to October, 2003. The relationships of biotic and abiotic factors were analyzed using the Simpson, Shannon—Wiener, and Hill diversity indices, and stepwise multilinear regression analyses techniques. The results showed that compared with continuous evergreen broadleaved forests, five fragmentations had a lower species diversity index, and different life forms showed differences in diversity index. With the decrease in patch areas, the daily differences in air temperature (ΔTa), ground surface temperature (ΔTs), daily differences in relative humidity (ΔRH), maximum wind velocity (Vmax), differences in photosynthetic available radiation (ΔPAR) (at noon) of both edges and interiors, all tended to increase. Maximum wind velocity (Vmax) and photo effective radiation in forest edges were higher than those in interior forest, which presented a stronger temperature-gained edge effect. In all the fragmentations of evergreen broadleaved forests, the depth of the edge effect was the nearest from interior forest in the biggest patch (about 15 meters away from interior forest), while the depth of the edge effect was the farthest from interior forest in the smallest patch (about 25 meters away from interior forest). With regard to the water conservation function, soil water content improved along with increasing species diversity. Some of the nutritional function substances of soil increased with increasing species diversity. The elements of microclimate, such as Ta, ΔTa, ΔTs, ΔRH, Vmax, and PAR, changed along with the extent of fragmented forest. Translated from Acta Ecologica Sinica, 2005, 25(7): 1642–1648 [译自: 生态学报]     

Flechten. Der Erfolg einer Symbiose

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

Respiration‐induced weathering patterns of two endolithically growing lichens

The two endolithic lichen species Hymenelia prevostii and Hymenelia coerulea were investigated with regard to their thallus morphology and their effects on the surrounding substrate. The physiological processes responsible for the observed alterations of the rock were identified. Whereas the thallus surface of H. coerulea was level, H. prevostii formed small depressions that were deepest in the thallus center. In a cross‐section, both species revealed an algal zone consisting of algal cavities parallel to the substrate surface and a fungal zone below. However, H. prevostii revealed significantly larger cavities with more than twice the cell number and a denser pattern of cavities than H. coerulea, resulting in a biomass per surface area being more than twice as large. Below H. prevostii the layer of macroscopically visibly altered rock material was about twice as deep and within this layer, the depletion of calcium and manganese was considerably higher. In simultaneous measurements of the oxygen uptake/oxygen release and pH shift, the isolated algal strains of both lichens revealed respiration‐induced acidification of the medium in the dark. At higher light intensities, H. coerulea and to a lesser extent also H. prevostii alkalized the medium which may lessen the acidification effect somewhat under natural conditions. In a long‐term growth experiment, the isolated algal strains of both lichens revealed acidification of the medium to a similar extent. Neither acidic lichen substances nor oxalic acid was identified. The significant differences between the weathering patterns of both species are based on the same respiration‐induced acidification mechanism, with H. prevostii having a greater effect due to its higher biomass per area.     

CYANOPHYTE CEPHALODIA IN THE LICHEN GENUS NEPHROMA

The lichens, Nephroma expallidum (Nyl.) Nyl. and N. arcticum (L.) Torss., consistently have at least two symbionts in a single thallus: a green alga in the algal layer and a blue-green alga in the internal cephalodia. The cephalodia originate from algal cells in contact with the lower surface of the lichen, in the zone of rhizine formation. The rhizines surround the epiphytic algal colony and form a second cortical layer; following dissociation of the original lower cortex, further growth of the two organisms results in the cyanophyte colony being enveloped by a compact layer of fungal tissue and positioned in the lichen medulla. The colony may eventually assume a superior or inferior position in relation to the lichen thallus, depending in part on the lichen species. Nephroma anticum may have two distinct morphological forms of blue-green algae in the same thallus and occasionally in the same cephalodium. It appears that the relationship that exists between the cephalodial algae and the lichen thallus is antagonistic and results, in some cases, in the exclusion of the green algal layer and death to the cephalodial cyanophytes.