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At least 19 recordsLinked to original sources

PubMed2006

Contrasting phylogeographic patterns in Sphagnum fimbriatum and Sphagnum squarrosum (Bryophyta, Sphagnopsida) in Europe.

The chloroplast phylogeography of two peat mosses (Sphagnum fimbriatum and Sphagnum squarrosum) with similar distributions but different life history characteristics was investigated in Europe. Our main aim was to test whether similar distributions reflect similar phylogeographic and phylodemographic processes. Accessions covering the European distributions of the species were collected and approx. 2000 bp of the chloroplast genome of each species was sequenced. Maximum parsimony, statistical parsimony and phylodemographic analyses were used to address the question of whether these species with similar distributions show evidence of similar phylogeographic and phylodemographic processes. The chloroplast haplotypes of the currently spreading species S. fimbriatum showed strong geographic structure, whereas those of S. squarrosum, which has stable historical population sizes, showed only very weak geographic affinity and were widely distributed. We hypothesize that S. fimbriatum survived the last glaciations along the Atlantic coast of Europe, whereas S. squarrosum had numerous, scattered refugia in Europe. The dominance of one haplotype of S. fimbriatum across almost all of Europe suggests rapid colonization after the last glacial maximum. We hypothesize that high colonizing ability is an inherent characteristic of the species and its recent expansion in Europe is a response to climate change.

Biological Evolution↗

PubMed2001

Population structure and genetic diversity of the amphiatlantic haploid peatmoss Sphagnum affine (Sphagnopsida).

Nineteen populations of Sphagnum affine were included in a study of genetic diversity and structure in fragmented and less fragmented landscapes, and differentiation at intercontinental and three regional levels. Isozyme electrophoresis of eight enzyme systems revealed 12 variable loci, which could be used for haplotype identification. A hierachical analysis of variance (AMOVA) revealed no significant intercontinental differentiation, and very limited differentiation among European regions. A trend of decreasing diversity with increasing latitude was apparent. Gametic phase disequilibria was high, suggesting nonrandom mating and regionally high incidences of inbreeding. The partitioning of genetic variation within and among populations in each region varied among regions, the northernmost populations having 86% of the total variation among populations, the southernmost in Scandinavia having 25% of the variation among populations, whereas the American populations displayed 89% of the variation within populations. Fifteen alleles at eight loci occurred in the U.S.A. which were not encountered in Europe, whereas only three European alleles at one locus in three populations were not encountered in U.S.A. The differences in diversity between North America and Europe may result from loss of genetic diversity caused by founder effects during postglacial recolonization of northern Europe. In Europe, the main mountain ranges extend E-W, posing severe barriers to northwards migration of lowland species, compared to the N-S trend of mountain ranges in North America. The decline in genetic diversity and increase in population differentiation and gametic phase disequilibria towards the north in Scandinavia may be caused by a series of founder effects during postglacial migration. These may have corresponded to minor climatic oscillations that influenced the migration front/leading edge in the suboceanic lowlands of Norway. According to this model random genetic drift will be an increasingly important structuring factor with latitude.

Analysis of Variance↗

PubMed2026

The genome sequence of Sphagnum contortum Schultz, 1819 (Sphagnales: Sphagnaceae).

We present a genome assembly of Sphagnum contortum (twisted bog-moss; Streptophyta; Sphagnopsida; Sphagnales; Sphagnaceae). The genome sequence has a total length of 370.55 megabases. Most of the assembly (99.46%) is scaffolded into 21 chromosomal pseudomolecules. The mitochondrial sequence has a length of 141.7 kilobases and the plastid genome assembly has a length of 140.11 kilobases. This assembly was generated as part of the Darwin Tree of Life project, which produces reference genomes for eukaryotic species found in Britain and Ireland.

Sphagnales↗

PubMed2026

The genome sequence of Sphagnum tenellum (Brid.) Bory, 1819 (Sphagnales: Sphagnaceae).

We present a genome assembly of Sphagnum tenellum (soft bog-moss; Streptophyta; Sphagnopsida; Sphagnales; Sphagnaceae). The genome sequence has a total length of 386.16 megabases. Most of the assembly (98.04%) is scaffolded into 21 chromosomal pseudomolecules. The mitochondrial sequence has a length of 141.31 kilobases and the plastid genome assembly has a length of 140.16 kilobases. This assembly was generated as part of the Darwin Tree of Life project, which produces reference genomes for eukaryotic species found in Britain and Ireland.

Sphagnales↗

PubMed2003

Structure of microbial communities in Sphagnum peatlands and effect of atmospheric carbon dioxide enrichment.

Little is known about the structure of microbial communities in Sphagnum peatlands, and the potential effects of the increasing atmospheric CO2 concentration on these communities are not known. We analyzed the structure of microbial communities in five Sphagnum-dominated peatlands across Europe and their response to CO2 enrichment using miniFACE systems. After three growing seasons, Sphagnum samples were analyzed for heterotrophic bacteria, cyanobacteria, microalgae, heterotrophic flagellates, ciliates, testate amoebae, fungi, nematodes, and rotifers. Heterotrophic organisms dominated the microbial communities and together represented 78% to 97% of the total microbial biomass. Testate amoebae dominated the protozoan biomass. A canonical correspondence analysis revealed a significant correlation between the microbial community data and four environmental variables (Na+, DOC, water table depth, and DIN), reflecting continentality, hydrology, and nitrogen deposition gradients. Carbon dioxide enrichment modified the structure of microbial communities, but total microbial biomass was unaffected. The biomass of heterotrophic bacteria increased by 48%, and the biomass of testate amoebae decreased by 13%. These results contrast with the absence of overall effect on methane production or on the vegetation, but are in line with an increased below-ground vascular plant biomass at the same sites. We interpret the increase in bacterial biomass as a response to a CO2-induced enhancement of Sphagnum exudation. The causes for the decrease of testate amoebae are unclear but could indicate a top-down rather than a bottom-up control on their density.

Amoeba↗

PubMed2005

Using transgenic Caenorhabditis elegans in soil toxicity testing.

Soil bioassays are important tools for evaluating toxicological effects within the terrestrial environment. The American Society for Testing and Materials E2172-01 Standard Guide outlines a method for conducting laboratory soil toxicity tests using the nematode Caenorhabditis elegans. This method is an efficient tool for extracting C. elegans from soil samples and can be carried out after a 24-h exposure period using relatively small amounts of soil. Drawbacks of this method include problems with (1) recovery of nematodes from soils containing a high percentage of organic matter, and (2) distinguishing indigenous nematode species from nematodes added for the laboratory test. Due in part to these issues, C. elegans has not been extensively accepted for use in soil testing. To address these concerns and improve upon the American Society for Testing and Materials method, this project focused on using transgenic strains of C. elegans carrying a GFP-expressing element. Lethality and behavior tests revealed that the transgenic nematodes respond similarly to the wild-type N2 strain, indicating that they can be used in the same manner in soil testing. The GFP marker is easily identifiable not only within soils containing a large amount of organic matter, but also in field-collected soils containing indigenous nematodes. These results support the use of transgenic GFP C. elegans in soil bioassays as a tool to further the reliability of laboratory toxicity tests.

Aluminum Silicates↗

PubMed2005

The ecology of testate amoebae (Protists) in sphagnum in North-western Poland in relation to peatland ecology.

We studied the relationship between testate amoebae (Protozoa) communities and the depth to the water table (DWT), pH, conductivity, and microhabitat type in Sphagnum dominated peatlands of north-western Poland and built predictive (transfer function) models for inferring DWT and pH based on the testate amoebae community structure. Such models can be used for peatland monitoring and paleoecology. A total of 52 testate amoebae taxa were recorded. In a redundancy analysis, DWT and pH explained 20.1% of the variation in the species data and allowed us to identify three groups of taxa: species that are associated with (1) high DWT and low pH, (2) low DWT and low pH, and (3) high pH and mid-range DWT. Our transfer function models allow DWT and pH to be estimated with mean errors of 9.89 cm and 0.71 pH units. The prediction error of the DWT model and the tolerance of the species both increase with increasing dryness. This pattern mirrors the ecology of Sphagnum mosses: Species growing in wet habitats are more sensitive to change in water table depth than the species growing in drier microhabitats. Our results are consistent with studies of testate amoeba ecology in other regions, and they provide additional support for the use of these organisms in paleoecological and biomonitoring contexts.

Adaptation, Physiological↗

PubMed2006

Comparison of bacterial communities in New England Sphagnum bogs using terminal restriction fragment length polymorphism (T-RFLP).

Wetlands are major sources of carbon dioxide, methane, and other greenhouse gases released during microbial degradation. Despite the fact that decomposition is mainly driven by bacteria and fungi, little is known about the taxonomic diversity of bacterial communities in wetlands, particularly Sphagnum bogs. To explore bacterial community composition, 24 bogs in Vermont and Massachusetts were censused for bacterial diversity at the surface (oxic) and 1 m (anoxic) regions. Bacterial diversity was characterized by a terminal restriction fragment length (T-RFLP) fingerprinting technique and a cloning strategy that targeted the 16S rRNA gene. T-RFLP analysis revealed a high level of diversity, and a canonical correspondence analysis demonstrated marked similarity among bogs, but consistent differences between surface and subsurface assemblages. 16S rDNA sequences derived from one of the sites showed high numbers of clones belonging to the Deltaproteobacteria group. Several other phyla were represented, as well as two Candidate Division-level taxonomic groups. These data suggest that bog microbial communities are complex, possibly stratified, and similar among multiple sites.

Bacteria↗

PubMed2005

Possible input of nitrogen of visitors' origin on a protected peatland.

The Ozegahara peatland, in the Nikko-Oze National Park in Japan, is ecologically significant because of its oligotrophic environment; it is one of the most strictly preserved areas in the country. The isotope ratio of nitrogen (15N/14N) and carbon (13C/12C) and C/N ratio of peat moss (Sphagnum spp.) and sundew (Drosera rotundifolia) in the peatland were analyzed. The correlation of the isotope ratio with some parameters (sundew population density, number of trapped insects, water level, bog myrtle coverage, and visitor density) was investigated. The nitrogen isotope ratio of sundew showed the most significant covariation with visitor density, where sundew from lunch areas or along busy walkways showed a higher nitrogen isotope ratio. The nitrogen isotope ratio of peat moss covaried, not only with route traffic frequency but also with water level, bog myrtle coverage, and number of trapped insects by sundew, indicating that factors other than the visitor level influence the local nitrogen cycle. This study suggests that the nitrogen imported into the peatland by visitors is a principal factor to be monitored for the maintenance of the natural environment.

Animals↗

PubMed2004

Distribution of cell wall components in Sphagnum hyaline cells and in liverwort and hornwort elaters.

Spiral secondary walls are found in hyaline cells of Sphagnum, in the elaters of most liverworts, and in elaters of the hornwort Megaceros. Recent studies on these cells suggest that cytoskeletal and ultrastructural processes involved in cell differentiation and secondary wall formation are similar in bryophytes and vascular plant tracheary elements. To examine differences in wall structure, primary and secondary wall constituents of the hyaline cells of Sphagnum novo-zelandicum and elaters of the liverwort Radula buccinifera and the hornwort Megaceros gracilis were analyzed by immunohistochemical and chemical methods. Anti-arabinogalactan-protein antibodies, JIM8 and JIM13, labeled the central fibrillar secondary wall layer of Megaceros elaters and the walls of Sphagnum leaf cells, but did not label the walls of Radula elaters. The CCRC-M7 antibody, which detects an arabinosylated (1-->6)-linked beta-galactan epitope, exclusively labeled hyaline cells in Sphagnum leaves and the secondary walls of Radula elaters. Anti-pectin antibodies, LM5 and JIM5, labeled the primary wall in Megaceros elaters. LM5 also labeled the central layer of the secondary wall but only during formation. In Radula elaters, JIM5 and another anti-pectin antibody, JIM7, labeled the primary wall. The distribution of arabinogalactan-proteins and pectic polysaccharides restricted to specific wall types and stages of development provides evidence for the developmental and functional regulation of cell wall composition in bryophytes. Monosaccharide-linkage analysis of Sphagnum leaf cell walls suggests they contain polysaccharides similar to those of higher plants. The most abundant linkage was 4-Glc, typical of cellulose, but there was also evidence for xyloglucans, 4-linked mannans, 4-linked xylans and rhamnogalacturonan-type polysaccharides.

Cell Wall↗

PubMed2004

Systemic induction of traumatic resin ducts and resin flow in Austrian pine by wounding and inoculation with Sphaeropsis sapinea and Diplodia scrobiculata.

The potential role of the resin system in the response of Austrian pine (Pinus nigra Arn.) seedlings to mechanical injury and fungal infection was studied in greenhouse experiments. Anatomical observations were performed on 2-year-old plants wounded at collar level and inoculated with Sphaeropsis sapinea (Fr.: Fr.) Dyko & Sutton in Sutton or Diplodia scrobiculata (J. de Wet, B. Slippers & M. J. Wingfield, sp. nov.; sensu de Wet et al. 2003), two fungal pathogens that cause shoot blight and canker on conifers, and that are characterized by different levels of aggressiveness. Histological examination of host tissue taken from the stem at 0, 8, and 12 cm above the treatment site revealed significant treatment- and time-dependent effects on the course of locally and systemically induced traumatic resin duct (TRD) development. Occurrence of TRDs was observed after 4 days only in seedlings inoculated with D. scrobiculata. At 12 days, TRDs were present also in mock-inoculated controls. No TRDs appeared in seedlings inoculated with S. sapinea. However, S. sapinea caused loss of vacuolar phenolics, severe disruption of cambial tissue and invaded the host xylem quickly and apparently unimpeded, whereas D. scrobiculata was never detected in the host xylem. Five-year-old Austrian pines subjected to the same stem base treatments were used to determine the resin mass flowing from the stem 30 cm above the treatment sites. Wounding and/or inoculation induced a significant, 8.3-fold average increase in systemic resin flow over the untreated trees 3 weeks after basal treatment, suggesting that wounding is the sole prerequisite for systemic induction of resin flow. The results are discussed in the context of current disease resistance models.

Ascomycota↗

PubMed2003

Does initial litter chemistry explain litter mixture effects on decomposition?

We tested the hypothesis that interactions in litter mixtures (expressed as the difference between observed and expected decomposition rates) are greater when the component species differ more in their initial litter chemistry. Thereto, we collected freshly senesced leaf litter from a wide range of species from an old field and woodland vegetation, and a fen ecosystem in The Netherlands. Litterbags with either mono-specific litter (20 and 15 species), or litter mixtures (50 and 42 species pairs) of randomly drawn combinations of two representatives from different plant functional types were incubated for 20, 35 and 54 weeks in a purpose-built decomposition bed (woodland/old field) or in the field (fen). Species showed a wide range of decomposition rates. For the woodland/old field species, initial litter C and P concentrations were significantly correlated with litter decomposition rate. For the fen species, litter phenolics concentration was correlated with decomposition rate. If the Sphagnum species were left out of the analyses, initial litter P and phenolics concentration both showed a significant relationship, albeit only with the remaining mass after 1 year. Differences between observed and expected decomposition were often considerable in individual litter mixtures. Regression analysis showed that such differences were not related to the differences in litter chemistry of the component species. Furthermore, litter mixtures containing species with very different initial litter chemistry did not show stronger interaction when tested against litter mixtures containing chemically similar litter types. From these observations we conclude that the difference in initial single litter chemistry parameters of the component is not a useful concept to explain interactions in litter mixtures.

Biodegradation, Environmental↗

PubMed2003

Carbon dioxide and methane fluxes in boreal peatland microcosms with different vegetation cover--effects of ozone or ultraviolet-B exposure.

O(3) concentrations in the troposphere are rising and those in the stratosphere decreasing, the latter resulting in higher fluxes of solar ultraviolet-B (UV-B) radiation to the earth's surface. We assessed whether the fluxes of CO(2) and CH(4) are altered by enhanced UV-B radiation or elevated tropospheric O(3) concentrations in boreal peatland microcosms (core depth 40 cm, diameter 10.5 cm) with different vegetation cover. At the end of the UV-B experiment which lasted for a growing season, net CO(2) exchange (NEE) and dark ecosystem respiration ( R(TOT)) were sevenfold higher, and CH(4) efflux 12-fold higher, in microcosms with intact vegetation dominated by Eriophorum vaginatum L. and Sphagnum spp., compared to microcosms from which we removed E. vaginatum. Vegetation treatment had minor effects on CH(4) production and consumption potentials in the peat, suggesting that the large difference in CH(4) efflux is mainly due to efficient CH(4) transport via the aerenchyma of E. vaginatum. Ambient UV-B supplemented with 30% and elevated O(3) concentrations (100 and 200 ppb, for 7 weeks) significantly increased R(TOT) in both vegetation treatments. Elevated O(3) concentrations reduced NEE over time, while UV-B had no clear effects on the fluxes of CO(2) or CH(4) in the cloudy summer of the study. Field experiments are needed to assess the significance of increasing UV-B radiation and elevated tropospheric O(3) concentration on peatland gas exchange in the long-term.

Carbon Dioxide↗

PubMed2004

Reduction of solar UV-B mediates changes in the Sphagnum capitulum microenvironment and the peatland microfungal community.

The influence of near-ambient and reduced solar UV-B radiation on a peatland microfungal community was assessed by exposing experimental plots to UV-selective filtration. Replicate plots were covered with special plastic films to effect treatments of near-ambient and attenuated solar UV-B. The microfungal community from the top 1 cm of Sphagnum capitulum in a Tierra del Fuego peatland was censused throughout three growing seasons, between 1999 and 2002. Sphagnum capitula under near-ambient UV-B were more compressed and held more water than capitula under reduced UV-B. This water had a greater conductivity and was more acidic under near-ambient UV-B, as would be expected with increased leaching from the Sphagnum leaves. Nine regularly occurring hyphal fungi from the peatland were identified, at least to genus. Over three field seasons, no treatment effect on total fungal colony abundance was recorded, but individual species abundance was increased (Mortierella alpina), decreased (Penicillium frequentans), or was unaffected (P. thomii, Aureobasidium) by near-ambient UV-B. Species richness was also slightly lower under near-ambient UV-B. These treatment differences were smaller than seasonal or inter-annual fluctuations in abundance and species richness. In a growth chamber experiment, lamp UV-B treatments indicated that realistic fluxes of UV-B can inhibit fungal growth in some species. In addition to this direct UV-B effect, we suggest that changes in the peatland fungal community under near-ambient solar UV-B may also result from increased nutrient and moisture availability in the Sphagnum capitulum. The subtle nature of the responses of peatland fungi to solar UV-B suggests that most fungal species we encountered are well adapted to current solar UV-B fluxes in Tierra del Fuego.

Biomass↗

PubMed2005

Stabilization of metals in acidic mine spoil with amendments and red fescue (Festuca rubra L.) growth.

Stabilization of metals with amendments and red fescue (Festuca rubra, cv. Keszthelyi 2) growth was studied on an acidic and phytotoxic mine spoil (pH(KCl) 3.20-3.26; Cd 7.1 mg kg(-1), Cu 120 mg kg(-1), Pb 2154 mg kg(-1) and Zn 605 mg kg(-1)) from Gyöngyösoroszi, Hungary in a pot experiment. Raising the pH above 5.0 by lime (CaCO(3)), and supplementing with 40 mg kg(-1)nitrogen (NH(4)NO(3)) made this material suitable for plant growth. All cultures were limed with 0.5% (m/m) CaCO(3) (treatment 1), which was combined with 5% (m/m) municipal sewage sludge compost (treatment 2), 5% (m/m) peat (treatment 3), 7.5% (m/m) natural zeolite (clinoptilolite) (treatment 4), and 0.5 (m/m) KH(2)PO(4) (treatment 5). Treatments 1-5 were combined with each other (treatment 6). After 60 days of red fescue growth, pH of the limed mine spoil decreased in all cultures units. Application of peat caused the highest pH decrease (1.15), while decrease of pH was less than 0.23 in treatments 2, 5 or 6. Application of lime significantly reduced concentrations of metals in the 'plant available' fraction of mine spoil compared to non-limed mine spoil. Amendments added to limed mine spoil changed variously the ratio of Cd, Cu, Pb and Zn in exchangeable or 'plant available' fractions, differently influencing the phytoavailability of these metals. Most of the metals were captured in the roots of test plants. Treatment 2 caused the appearance of less Cd in shoots (<0.1 microg g(-1)) or roots (3.11 microg g(-1)), while treatment 5 resulted in the highest Cd concentration (2.13 microg g(-1)) in shoots. Treatments did not influence significantly the Cu accumulation in shoots. The Pb accumulation of roots (44.7 microg g(-1)) was most effectively inhibited by combined treatment, while the highest value (136 microg g(-1)) was found in the culture treated with potassium phosphate. Pb concentration in shoots was below the detection limit, except for treatments 5 and 6. Peat application resulted in higher Zn concentration (448 microg g(-1)) in shoots than other amendments, where these values were around 100 microg g(-1). All amendments influenced positively the dry matter yield of red fescue grown in limed mine spoil, however the application of 0.5 phosphate was less favourable. Liming, application of amendments and growth of red fescue can stabilize metals in acidic and phytotoxic mine spoil, and by phytostabilization they can reduce the risk of metal contamination of the food chain.

Biodegradation, Environmental↗

PubMed2005

Active moss biomonitoring applied to an industrial site in Romania: relative accumulation of 36 elements in moss-bags.

Active moss biomonitoring using the species Sphagnum girgensohnii was tested at a strongly polluted site in Romania (Baia Mare) according to a novel sampling design. Nine moss transplants from each of the two background areas (Dubna, Russia and Vitosha Mountain, Bulgaria) were deployed in parallel on balconies about 24 m above street level for 4 months. The samples were analyzed for 36 elements using instrumental neutron activation analysis (INAA). Based on the results obtained the sampling variability is discussed in relation to the analytical variability, and the relative uptake of the different elements is assessed. The moss-bags using Sphagnum girgensohnii demonstrate a high or a very high relative uptake for a majority of the 36 investigated elements, but the values depend on the initial element concentration in the moss. Moss leaves analyzed separately showed somewhat higher levels than stems for many elements. Practical considerations however still speak in favor of using the whole moss for transplants.

Air Pollutants↗

PubMed2005

Efficacy of a biomonitoring (moss bag) technique for determining element deposition trends on a mid-range (375 km) scale.

National networks detect multi-state trends in element deposition using direct measurement methods. Biomonitoring techniques have been used to examine deposition in local areas and around point sources. We sought to determine the efficacy of a moss bag technique to detect element deposition trends on a mid-range (state) scale, and to compare these results with those of the National Acid Deposition Program/National Trends Network (NADP/NTN, 1999). We sampled heavy metals, sulfur, and nitrogen deposition (21 elements) using mesh bags containing Sphagnum russowii at nine sites, over a 375 km transect crossing southern Wisconsin (upper Midwest, USA). We found statistically significant trends of decreasing deposition in a northwesterly direction for 13 elements: Al, B, Ca, Cd, Co, Cu, Cr, Fe, Mg, Mn, Ni, S, and Zn. Six of these have moderate to large changes in concentration (14-37%). The trends for Ca, Mg, and S are consistent with regional deposition patterns in 1998 isopleth maps from the NADP/NTN (1999) which are derived from a sampling array far less dense than the transect sites. This national network indicates that Ca and Mg increase to the southeast, beyond Wisconsin borders. The fact that the present study demonstrates strong correlations between both of these elements (Ca and Mg) and Al, B, Cr, Cu, Fe, Mn, Ni, and Zn (mean r for all correlations = 0.75, p < 0.02) implies that these correlated elements also increase to the southeast in neighboring states.

Environmental Monitoring↗

PubMed2005

Removal of H(2)S by Thiobacillus denitrificans immobilized on different matrices.

Biological removal of high concentrations of H(2)S was studied using the immobilized Thiobacillus denitrificans with peat moss, wood chip, ceramic and granular activated carbon (GAC) separately. Experiments on the physical adsorption capacity of matrix, retention time and pressure drop were carried out; the ability of bioreactor to buffer shock loading and the removal efficiency with different packing materials were also investigated. Besides, the kinetics of single-stage biodesulfuration was analyzed. The results showed that GAC provided higher bacteria adsorption capacity, showed a more resistance to shock loading and allowed better operational control with respect to pressure drop than other inert carriers. When the retention time was changed from 30 to 100 s at an influent concentration of 100 mg/L of H(2)S, the removal efficiencies were above 98%; when the inlet concentration of H(2)S were changed from 110 to 120 mg/L, an average 96.8% removal efficiency was achieved during the long-term operation for GAC bioreactor. Next to GAC, wood chip was found to be a good packing material; however, peat moss and ceramic had limited effectiveness and their removal efficiencies were less of 90%. The kinetic analysis showed that the maximum removal rate and the half-saturation constant of the GAC bioreactor were 666.7 mg (H(2)S)/(L.d) and 20.8 mg/L, respectively.

Adsorption↗