Mollusc grazing limits growth and early development of the old forest lichen Lobaria pulmonaria in broadleaved deciduous forests

This study aims: (1) to quantify mollusc grazing on juvenile and mature thalli of the foliose epiphytic lichen Lobaria pulmonaria, and (2) to test the hypothesis inferring a herbivore defensive role of lichen depsidones in forests with indigenous populations of lichen-feeding molluscs. Lichens were transplanted in shaded and less shaded positions in each of two calcareous broadleaved deciduous forests, one poor in lichens, one with a rich Lobarion community. Preventing the access of molluscs significantly reduced the loss of juvenile L. pulmonaria, particularly in the naturally lichen-poor forest. Molluscs also severely grazed mature thalli in the lichen-poor forest, especially thalli placed under the more shading canopies. Furthermore, reducing the natural concentration of depsidones by pre-rinsing with acetone increased subsequent grazing significantly, showing that lichen depsidones function as herbivore defence in natural habitats. Our results suggest that mollusc grazing may play important roles in shaping the epiphytic vegetation in calcareous deciduous forests, and that recently established juvenile L. pulmonaria thalli seem to be particularly vulnerable.     

Fungal depsidones – an inducible or constitutive defence against herbivores in the lichen Lobaria pulmonaria?

Herbivore-deterrent depsidones in the epiphytic lichen Lobaria pulmonaria were quantified after a 104-day exposure to indigenous lichen-feeding mollusc communities in broadleaved deciduous forests in southeastern Norway. Controls and acetone-rinsed living thalli were transplanted under open and shaded tree canopies. Rinsed thalli had their depsidone concentration reduced to 36% of the pre-rinsing level, which is below the level needed to deter grazing molluscs. Grazing did not raise the concentration of depsidones beyond the level occurring in control to which molluscs had no access. Inducible responses were not detected in controls nor in acetone-rinsed thalli. Depsidone resynthesis was negligible in acetone-rinsed thalli regardless of grazing and/or light regimes. Our results suggest that C-based depsidones represent a constitutive type of herbivore defence in L. pulmonaria. A constitutive defence is probably an advantage for stress-tolerant slow-growing lichens inhabiting habitats with a constant presence of generalist invertebrate herbivores.     

Fungal melanins as a sun screen for symbiotic green algae in the lichen<Emphasis Type="Italic"> Lobaria pulmonaria</Emphasis>

The mycobiont of the high-light-susceptible forest lichen Lobaria pulmonaria was shown to deposit brown, melanic compounds in the outer layer of the upper cortex, depending on the long-term level of solar radiation in its natural habitat. Furthermore, pale thalli from a shady habitat produced melanic compounds after transplantation to a sunny habitat. This browning of the cortex appeared to be a physiologically active process, taking place only during periods with frequent hydration. Melanin production was slow. After transplantation, more than 1 year was needed for a shade-adapted thallus to reduce the cortical transmittance (230-1000 nm) to a similar level to that of naturally sun-exposed specimens. Melanic compounds acted as a sun screen, especially reducing UVB and UVA wavelengths, but also visible wavelengths, at the photobiont level. In the near infrared range, there was only a small difference in transmittance between shade- and sun-adapted cortices. A negative correlation was found between the natural light level and the cortical transmittance of wavelengths below 700 nm. However, previous studies have shown that even photobionts of melanic L. pulmonaria thalli are relatively susceptible to high-light exposure. Since melanins also increase the absorbance of solar energy for the whole thallus, it appears that what is gained in terms of UV- and light protection in melanic L. pulmonaria specimens may be offset by increased exposure to excess temperatures for this highly heat-susceptible lichen.     

Light might regulate divergently depside and depsidone accumulation in the lichen Parmotrema hypotropum by affecting thallus temperature and water potential

Depsides and depsidones are the most common secondary products uniquely produced in lichens by the fungal symbiont, and they accumulate on the outer surface of its hyphae. Their biological roles are subject to debate. Quantitatively the compounds typical of a given lichen can vary dramatically from thallus to thallus. Several studies have addressed whether this variability is correlated with the light reaching different thalli, but the conclusions are contradictory. We addressed the question with the lichen Parmotrema hypotropum growing on unshaded, vertical tree trunks, a controlled natural environment where the light absorbed by each thallus over its lifetime is the only major position-dependent variable. The exact north-east-south-west orientation of each thallus was used to calculate its yearly light exposure based on astronomical and meteorological considerations. The calculated irradiation around the trunk, distributed over a continuous 40-fold intensity range, then was compared with the amount of compound per unit thallus weight, determined by quantitative thin layer chromatography. P. hypotropum accumulates the depside atranorin in the cortex and the depsidone norstictic acid in the medulla and around the algae. A direct correlation was observed between the yearly amount of light reaching the lichen and the amount of atranorin. In contrast, the amount of norstictic acid decreased with increasing light. Although we did not measure thallus temperature and water potential, a unifying interpretation of these and other published data is that depside/depsidone accumulation in lichens is mediated by localized changes in temperature and water potential produced by light absorption within each thallus. This suggests water relations-based functions for depsides and depsidones.     

Extremely low lichen growth rates in Taylor Valley,Dry Valleys,continental Antarctica

Estimates of lichen growth rates based on the measurements of several thalli at any site do not exist for continental Antarctica. However, the very limited existing data suggest that lichen growth rate may be a good indicator of climate change in Antarctica. We present measurements made on thalli of the lichen Buellia frigida Darb. growing in the Dry Valleys, Southern Victoria Land, continental Antarctica, which appear to have some of the slowest radial growth rates yet measured. Photographs of thalli at three different sites were analysed for growth over a 25-year period using nano-GIS techniques. At one site, Mt. Falconer Summit, the lichens had a mean growth rate of 0.0052 mm year⁻¹ with one individual as low as 0.0036 mm year⁻¹. Thalli at the other two sites had significantly higher mean growth rates, 0.0136 mm year⁻¹ at Mt. Falconer Ridge and 0.0118 mm year⁻¹ at Rhone Bench. Assuming a constant growth rate, thalli at Mt. Falconer Summit had a mean age of 5,367 years, whilst the thalli at the other two sites were much younger, 840–1,026 years. We suggest that the different ages represent the appearance of new substrate for colonisation following climate changes in the Dry Valleys that altered the amount and duration of snow. The results confirm that lichen growth rate differs by almost two orders of magnitude over a latitudinal range of 15 degrees from south to north across Antarctica.     

In situ development of the foliicolous lichen Phyllophiale (Trichotheliaceae) from propagule germination to propagule production

The vegetative cycle of the foliicolous lichen Phyllophiale, from propagule germination to propagule production, was studied by light microscope observation of thalli colonizing plastic cover slips placed within a lowland tropical forest. Discoid propagules germinated by growth of radially arranged fungal cells and developed directly into lichen thalli. The young lichen comprised a single disc of closely branched, radiating filaments of the algal symbiont Phycopeltis, covered by a network of fungal hyphae extending onto the substrate as a prothallus. The prothallic hyphae incorporated additional Phycopeltis thalli encountered on the substrate. The phycobiont formed a single layer, with individual algal thalli clearly distinguishable within the lichen. Radial growth ceased at points of contact between adjacent phycobiont thalli. The visible shape of the crustose lichen thallus corresponded to the perimeter of the phycobiont thalli within. Propagules were initiated at points corresponding to the margins of the phycobiont thalli, by vertical reorientation of horizontal algal filaments surrounded by fungal hyphae. The lichenized alga produced intercalary gametangia. Degeneration of propagules unsuccessful in lichen establishment sometimes resulted in free growth of the phycobiont. The alga generally maintained its shape, growth pattern, and reproductive independence within the lichen, while also participating in the formation of unique symbiotic propagules.     

Growth and ecophysiological acclimation of the foliose lichen Lobaria pulmonaria in forests with contrasting light climates

This study aims to assess biomass and area growth of 600 thalli of the old forest lichen, Lobaria pulmonaria, transplanted to three successional boreal forest stands with (1) natural rainfall regime, (2) additional moistening during dry days, and (3) additional moistening with added nutrients. Mean biomass growth during 100 days varied from 8.3% in the dark young spruce forest to 23.1% in the clear-cut area, with the old forest in between (16.0%). Additional moistening did not enhance lichen growth, probably because the transplantation period was wet. Nutrient additions slightly increased area growth compared to artificial water additions only. Growth was determined by a combination of external (forest stand, site factors) and internal factors (chlorophyll content, biomass per area). Transplants acclimated to high light by increasing thickness and chlorophyll a/b-ratio. Some visible bleaching and a strong positive correlation between chlorophyll content per area and lichen growth in clear-cuts suggest some high light-induced chlorophyll degradation. We believe that biomass growth and natural occurrence of L. pulmonaria is controlled by a delicate balance between light availability and desiccation risk, and that the species is confined to old forests due to a physiological trade-off between growth potential and fatal desiccation damage, both of which increase with increasing light. The discrepancy between potential and realized ecological niches is probably caused by a long-term risk to be killed in open habitats by high light during long periods with no rain.     

Defensive compound concentration in boreal lichens in response to simulated nitrogen deposition

Nitrogen (N) deposition is expected to increase in northwestern Europe the next 50–100 years. The effects of higher N availability on lichens will presumably depend on their capacity to acquire carbon (C), that is, of the timing and duration of the wet and active state. If lichens respond like plants, their C and N status may affect their concentration of carbon-based secondary compounds (CBSCs), and thus their defence against herbivores, detrimental radiation, pathogens and parasites. In the present study we have manipulated N availability and timing and duration of the metabolically active state by spraying lichen transplants in an old spruce forest with rainwater or rainwater with added N corresponding to 50 kg N ha⁻¹ yr⁻¹. The spraying was applied either at night, in the morning or at noon to also investigate the effect of timing and duration of the active state. Concentrations of N, chlorophyll a (Chl a ) and CBSCs were measured before and after one summer's spraying of 10 thalli in each of four different lichen species; Alectoria sarmentosa, Lobaria scrobiculata, Platismatia glauca , and Xanthoria aureola . The added N was readily taken up by all the lichen species. A. sarmentosa, P. glauca , and X. aureola increased their Chl a concentration in response to increased N, while L. scrobiculata increased Chl a in response to increased active time. None of the studied species reduced their concentration of secondary compounds during the experimental period, but in P. glauca the concentration of all compounds were significantly lower in N-treated thalli compared with those that got only rainwater. The results are consistent with a high degree of constitutive defence in three of four species, and we conclude that all the investigated lichens seem to have rather robust chemical defence systems despite considerable manipulation of the environmental conditions.     

Competitive trade-off strategies in ArcticDaphnia linked to melanism and UV-B stress

 Different high-Arctic, freshwater localities at Ny-A˚lesund, Svalbard (79°N) were examined for their UV-absorbing properties, depth and presence of melanic or non-melanic morphs of the planktonic crustacean Daphnia pulex. Light regimes in two localities with each of these morphs were measured by using underwater spectroradiometer. Most localities have low absorbance of short-waved light, but no clearcut relationship between UV transparency and occurrence of melanic morphs was detected. Yet, in the laboratory, the melanic morph showed far lower growth rates, thus being competitively inferior to the non-melanic. Conversely, the melanic morph was more resistant to UV light, suggesting a trade-off between the metabolic tax paid for the melanin synthesis and its UV-protecting abilities, or a staggered growth capacity possibly owing to polyploidy. Frequency of melanic or non-melanic clones could thus be directly linked to ambient UV-B stress and serve as an indicator thereof, but the apparently extensive need for UV protection under the Arctic light regimes is still puzzling, and the role of melanism and polyploidy in these organisms cannot be considered finally settled. Received: 29 September 1995/Accepted: 29 January 1996     

Use of 14C/C ratios to provide insights into the magnitude of carbon turnover in the crustose saxicolous lichen Caloplaca trachyphylla

Abstract:¹⁴C/C ratios in samples from radial transects across individual thalli ofCaloplaca trachyphylla collected at two sites were measured and the results used to investigate whether ¹⁴C/C data might provide some insight into the magnitude of carbon turnover in this lichen species. The ¹⁴C/C data suggest that significant internal recycling/translocation of carbon is unlikely in the sampled thalli. However, converting the¹⁴ C/C data for the larger intact thalli sampled at each site to calendar years, using the atmospheric ¹⁴C record, does not yield constant or even monotonically varying growth rates. Since crustose lichen growth rates are constant or decrease with thallus size, and since the ¹⁴C/C data from these larger thalli show a relatively small spread in¹⁴ C/C data values compared to the Northern Hemisphere atmospheric ¹⁴C record over the past 50 years, the¹⁴ C/C data suggest that carbon turnover may be occurring. Carbon turnover was modelled starting with the atmospheric¹⁴ C record. Turnover was incorporated so that for each year in the record a constant percentage of the total carbon was lost annually and replaced by new photosynthetically fixed carbon with a ¹⁴C/C ratio equal to that of the contemporary atmosphere. The ¹⁴C/C data from the radial samples were then converted to a calendar year using the model record. Constant annual carbon turnover values of 0, 0·5, 1, 1·5, 2, 2·5, 3, 3·5, 4, 4·5, 5, 5·5, 6, 7, 8, 9, 10, 15, 20, 25 and 50% were modelled. Carbon turnover values between 3 and 6% created ¹⁴C model records that when applied to ¹⁴C/C data from the thalli produced constant radial growth rates that were: (1) identical for all lichens at a given site, and (2) independent of lichen size at a given site. The ¹⁴C/C data further indicate that annual carbon turnover in this species of lichen is <10%, independent of the nature of thallus radial growth. The data and modelling suggest that carbon turnover might provide a simple explanation for the ¹⁴C/C data from the thalli and might explain the discrepancies between the standard atmospheric ¹⁴C record and the ¹⁴C/C ratios observed in C. trachyphylla.     

Possible functional roles of cortical depsides and medullary depsidones in the foliose lichen Hypogymnia physodes

Secondary compounds were quantified in 75 thalli of the foliose lichen Hypogymnia physodes collected in habitats along a natural shade-sun gradient from dark spruce forests to sun-exposed sea cliffs. The irradiance in all the 75 lichen sites was estimated from hemispherical photographs. The content of lichen compounds per thallus area correlated positively with irradiance level (r²=0.73), and the mean concentration increased from 6.7% in the spruce forest to 14.4% on sea cliffs. The medullary depsidones, physodic, physodalic and protocetraric acids, constituted >95% of the total pool of extractable secondary compounds, the cortical depsides, atranorin and chloratranorin, represented <5%. Both cortical compounds correlated well with direct and with diffuse radiation, whereas the three medullary compounds correlated better with diffuse than with direct radiation. Mentioned trends are consistent with a solar radiation screening hypothesis of both groups of these colourless compounds occuring as tiny crystals outside fungal hyphae. However, the UV-B protective hypothesis of the compounds was further tested in a lab experiment. Unnaturally high UV-B doses were required to significantly reduce the PSII efficiency (F_V/F_M) of symbiotic algae. Removal of the major pool of secondary compounds with acetone did not increase photobiont susceptibility to UV-B. Therefore, the main functional role of the UV-B absorbing secondary compounds in H. physodes is hardly UV-B screening. Other roles such as PAR-screening and defence against herbivores and pathogenic organisms are discussed.     

地衣次生代谢产物及其生物活性研究进展

地衣是一类独特的生物有机体,其体内合成了多种特殊的次生代谢产物,主要为缩酚酸、缩酚酸环醚、缩酚酮、二苯并呋喃等类化合物。这些次生代谢产物具有多样的生物活性,包括抗氧化、抗辐射、抗菌、抗病毒、抗肿瘤、植物生长抑制和昆虫拒食等活性。本文对近40年来有关地衣化学及生物活性方面的研究进行综述,为地衣资源的进一步研究和开发提供参考。     

Photosynthetic acclimation to different light levels in the brown marine macroalga, <Emphasis Type="Italic">Hizikia fusiformis</Emphasis> (Sargassaceae,Phaeophyta)

Hizikia fusiformis thalli experience dynamic incident light conditions during the period of growth. The present study was designed to examine how changing photon irradiance affects the photosynthesis both in the short and long terms by culturing H. fusiformis under three different light levels: 35 μmol photons m^-2 s^-1 (low light, LL), 85 μmol photons m^-2 s^-1 (intermediate light, IL), and 165 μmol photons m^-2 s^-1 (high light, HL). A similar relative growth rate was observed between IL- and HL-grown algae, but the growth rate was significantly reduced in LL-grown algae. The photosynthetic rates (P _n) measured at their respective growth light levels were found to be lowest in the thalli grown at LL and highest at HL. However, LL-grown algae exhibited much higher P _n in comparison with IL- and the HL-grown thalli at the same measuring photosynthetic photon flux density, indicating the photosynthetic acclimation to low growth light in H. fusiformis. The photosynthesis–light curves showed that LL-grown algae had a highest light-saturating maximum P _n (P _max) in comparison with IL- or HL-grown algae when the photosynthetic rates were expressed on the biomass basis. However, P _max was highest in HL-grown algae compared to IL- or LL-grown algae when the rates were normalized to chlorophyll a. The photosynthesis–inorganic carbon (Ci) response curves were also significantly affected by the growth light conditions. The highest value of apparent photosynthetic conductance occurred in LL-grown algae while the lowest value in HL-grown algae. Additionally, the activity of external carbonic anhydrase (CA) tended to increase while the total CA activity inclined to decrease in H. fusiformis thalli when the growth light level altered from 35 to 165 μmol photons per square meter per second. The external CA inhibitors showed a higher inhibition in HL-grown algae compared with LL-grown algae. It was proposed that photosynthetic acclimation to low light condition in H. fusiformis was achieved through an increase in the number of reaction centers and increased capacities of electron transport and of Ci transport within cells. The ability of photosynthetic acclimation to low light confers H. fusiformis thalli to overcome the environmental low light condition as a result of the attenuation of seawater or self-shading through enhancing its photosynthetic performance and carbon assimilation necessary for growth.     

Seasonal acclimation in the epiphytic lichen Parmelia sulcata is influenced by change in photobiont population density

CO₂ gas exchange, radial growth, chlorophyll (Chl) content and photobiont density of an epiphytic population of Parmelia sulcata were monitored every 2 months during 1 year in a temperate deciduous forest of Central Italy, to verify possible seasonal variations. Light response curves of south-exposed thalli, built up in the laboratory at 6 and 27°C at optimal thallus hydration, showed that CO₂ gas exchange changed significantly during the year, with a maximum for gross photosynthesis in December at both temperatures. Photoinhibition phenomena occurred in early spring, immediately before tree leaves sprouted. The principal component analysis of CO₂ gas exchange parameters clearly separated the months with from the months without tree canopy cover. Radial growth, measured on marginal lobes of north- and south-exposed thalli, was the highest in December, and the lowest in April. Photobiont density, measured in lobes of south- and north-exposed thalli with a sedimentation chamber, also changed during the year: the number of photobionts was highest in June and December, and lowest in April, although no significant change in cell size and Chl content per cell was evident throughout the year. South-exposed thalli had slightly, but constantly higher photobiont density both on a weight and an area basis. The acclimation of lichen photosynthesis and Chl content to seasonal temperature and light changes should partially be re-visited on the basis of the significant variation in photobiont population density. This phenomenon still awaits, however, a satisfactory explanation, although it is probably related to the seasonal change in nutrient availability.     

Genetic structure in a fragmented Northern Hemisphere rainforest: large effective sizes and high connectivity among populations of the epiphytic lichen Lobaria pulmonaria

An extraordinary diversity of epiphytic lichens is found in the boreal rainforest of central Norway, the highest-latitude rainforest in the world. These rainforest relicts are located in ravine systems, and clear cutting has increased the distance between remaining patches. We hypothesized that the relatively small lichen populations in the remaining forest stands have suffered a depletion of genetic diversity through bottlenecks and founder events. To test this hypothesis, we assessed genetic diversity and structure in the populations of the tripartite lichen Lobaria pulmonaria using eight SSR loci. We sampled thalli growing on Picea abies branches and propagules deposited in snow at three localities. Contrary to expectations, we found high genetic diversity in lichen and snow samples, and high effective sizes of the studied populations. Also, limited genetic differentiation between populations, high historical migration rates, and a high proportion of first generation immigrants were estimated, implying high connectivity across distances <30km. Almost all genetic variation was attributed to variation within sites; spatial genetic structures within populations were absent or appeared on small scales (5-10m). The high genetic diversity in the remaining old boreal rainforests shows that even relict forest patches might be suitable for conservation of genetic diversity.     

Sapling growth and survivorship as a function of light in a mesic forest of southeast Texas, USA

For seven species in a mature mesic forest in southeast Texas, we estimated species-specific parameters representing radial growth in high light and low light for tree saplings. Shade-intolerant species had higher asymptotic growth rates and lower low-light growth than tolerant species. Inspection of species positions on graphs of low-light growth versus high-light growth suggested that there was a trade-off between these two processes across species. By linking functions of growth versus light and mortality versus growth, we also found that shade-intolerant species had higher mortality risk at low light and stronger sensitivity of mortality to light than shade-tolerant species. Moreover, we found that low-light survival and high-light growth were negatively correlated across species. In contrast to northern hardwood forests, where sapling survival in low light may be achieved at the expense of growth, our results suggested that shade-tolerant species in this southern mixed forest can grow faster as well as survive better than shade-intolerant species in low light. We conclude that both sapling growth and survival are important components of shade tolerance and their relationships may be system-specific.     

Growth in relation to microclimatic conditions and physiological characteristics of four Lobaria pulmonaria populations in two contrasting habitats

The aim of the present study was to compare the physiological characteristics of various populations of the lichen Lobaria pulmonaria in Portugal and Sweden. For this, indirect markers of algal (photobiont) and fungal (mycobiont) activity were measured, as well as their CO₂ gasexchange characteristics. Microclimatic conditions and the lichens growth performance in the two countries were compared using reciprocal transplantation. Two populations of L. pulmonaria represented each country: one collected from forest interior conditions and one from forest edge habitats. A non-transplanted "wild" population was also studied in each country, in order to evaluate any transplantation effects per se. The main hypothesis were that; 1) growth should be faster in Portugal due to higher light availability; 2) the energy use efficiency of lichen biomass gain should be similar for the native populations in their respective native habitat; 3) if the lichens were able to adapt to the environmental conditions in the foreign habitat this should be revealed as similar growth rates among all thalli transplanted at the same site, regardless of their origin. Physiologically, the Portuguese and Swedish populations were very similar, both concerning their CO₂ gas exchange characteristics and distribution of resources between photo- and mycobiont tissue. Environmental conditions were more advantageous for L. pulmonaria growth in Portugal, i.e. higher photon flux densities and ambient temperatures when the lichens were wet and active, and a lower fraction of the active time occurring in darkness. However, despite similar physiological characteristics of all the studied populations, the Swedish lichens were not able to grow as well in Portugal as the native, while all populations had similarly low growth rates in Sweden.     

The lichens <Emphasis Type="Italic">Xanthoria elegans</Emphasis> and <Emphasis Type="Italic">Cetraria islandica</Emphasis> maintain a high protection against UV-B radiation in Arctic habitats

This study reports UV screening pigments in the upper cortices of two widespread lichens collected in three sun-exposed locations along a latitudinal gradient from the Arctic lowland to alpine sites of the Central European Alps. Populations from the Alps receive 3–5 times higher UV-B irradiance than their Arctic counterparts from Svalbard because of latitudinal and altitudinal gradients in UV-B irradiance. In Cetraria islandica, the screening capacity of melanin in the upper cortices was assessed by direct measurements of cortical transmittance (250–1,000 nm). A comparison of cortical transmittances in brown sun-exposed and pale shade-adapted forest C. islandica thalli showed that fungal melanins strongly absorb both UV-B and photosynthetically active radiation (PAR). For Xanthoria elegans cortical UV-B absorbing pigments, mainly the orange parietin, were extracted and quantified. Field experiments with extracted, parietin-deficient X. elegans thalli cultivated under various filters showed that UV-B was essential for the induction of parietin synthesis. The parietin resynthesis in these parietin-deficient samples increased with decreasing latitude of their location in which they had been sampled, which may imply that the synthesis of pigments is habitat specific. However, no latitudinal gradient in cortical screening capacity was detected for any of the two species investigated in the field. This implies that Arctic populations maintain a high level of screening pigments in spite of low ambient UV-B, and that the studied lichen species presumably may tolerate an increase in UV-B radiation due to the predicted thinning of the ozone layer over polar areas     

QUANTITATIVE VARIATION IN VULPINIC AND PINASTRIC ACIDS PRODUCED BY TUCKERMANNOPSIS PINASTRI (LICHEN-FORMING ASCOMYCOTINA,PARMELIACEAE)

Tuckermannopsis pinastri is a lichen species found commonly on rocks and tree branches in boreal and alpine habitats in the northern United States. Members of this species produce three yellow-pigmented phenolic compounds: usnic, pinastric, and vulpinic acids. The objective of our study was to quantify the variation in concentration of the latter two of these compounds in relation to substrate factors, chlorophyll content, and thallus size. Using high-performance liquid chromatography, we analyzed 120 thalli of T. pinastri collected randomly from a single large population located at Spruce Knob, West Virginia. Although individuals were sampled from both tree and rock substrates that differed markedly in light intensity, these environmental factors were not correlated with observed variations in vulpinic or pinastric acid concentrations. Instead, compound concentrations were correlated most closely with thallus size, with small rather than large thalli having the highest concentrations of the two compounds. Small thalli did not have higher concentrations of chlorophylls than large thalli, however, which suggests that the rate of production of secondary compounds by the fungus in T. pinastri is independent of algal biomass. Inasmuch as lichen secondary compounds serve a defensive role against microorganisms and herbivores, our results suggest that small, juvenile thalli are better defended than more mature thalli.     

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。