On the competitive balance of macroalgae at Potter Cove (King George Island,South Shetlands)

The sublittoral zonation of macroalgae and abundant animals in Potter Cove, King George Island, is described in relation to substrate, exposure to turbulence and impact of grounding icebergs. Implications on the ecological niches of the most prominent phaeophytes are discussed. It is concluded that Desmarestia anceps and D. menziesii exclude Himantothallus grandifolius under favourable conditions. However, Himantothallus, by its potential to inhabit unstable substrates, may be better adapted to withstand the ice impact. The replacement of Desmarestia by Himantothallus at greater depth can be explained only partially at present.     

Photosynthetic characteristics and UV stress tolerance of Antarctic seaweeds along the depth gradient

The photosynthetic characteristics through P-E curves and the effect of UV radiation on photosynthesis (measured as rapid adjustment of photochemistry, F _v/F _m) and DNA damage (as formation of CPDs) were studied in field specimens of green, red and brown algae collected from the eulittoral and sublittoral zone of Fildes Peninsula (King George Island, Antarctic). The content of phenolic compounds (phlorotannins) and the antioxidant activity were also studied in seven brown algae from 0 to 40 m depth. The results indicated that photosynthetic efficiency (α) was high and did not vary between different species and depths, while irradiances for saturation (E _k) averaged 55 μmol m^?2 s^?1 in subtidal and 120 μmol m^?2 s^?1 in eulittoral species. The studied species exhibited notable short-term UV tolerance along the vertical zonation. In intertidal and shallow water species, decreases in F _v/F _m by UV radiation were between 0 and 18 %, while in sublittoral algae, decreases in F _v/F _m varied between 3 and 35 % relative to PAR treatment. In all species, recovery was high averaging 84–100 %. The formation of CPDs increased (15–150 %) under UV exposure, with the highest DNA damage found in some subtidal species. Phlorotannin content varied between 29 mg g^?1 DW in Ascoseira mirabilis from 8 m depth and 156 mg g^?1 DW in Desmarestia menziesii from 17 m depth. In general, phlorotannin concentrations were constitutively high in deeper sublittoral brown algae, which were correlated with higher antioxidant activities of algal extracts and low decreases in photosynthesis. UV radiation caused a strong decrease in phlorotannin content in the deep-water Himantothallus grandifolius, whereas in D. menziesii and Desmarestia anceps, induction of the synthesis of phlorotannins by UV radiation was observed. The antioxidant activity was in general less affected by UV radiation.     

Summer macroalgal biomass in Potter Cove,South Shetland Islands,Antarctica: its production and flux to the ecosystem

Summer macroalgal biomass and production were analyzed at Potter Cove, King George Island, Antarctica and the potential carbon transfer of macroalgal production to the coastal ecosystem of the cove was estimated. A total of 38 algal species were found, with Desmarestia anceps, D. menziesii and Himantothallus grandifolius accounting for almost 80% of the biomass. Biomass data and published growth rates were combined to calculate the production of the five most abundant species. The standing stock for each summer month was estimated as the product of the average biomass and the area of the macroalgal stands. The monthly biomass production was calculated for each species by difference between the expected biomass and the observed biomass at the previous month. The macroalgal production showed a decreasing trend during the summer months. The average standing stock in the whole cove was 792.84 MT and the production was 1,401.33 MT during the summer 1994–1995. The flux of biomass to the ecosystem during the summer period was 1,370.61 MT, which is almost as much as the total summer production. The study demonstrates that macroalgae are one of the main energy sources in Potter Cove, and probably support a large fraction of the secondary production of the benthos.     

Extraction of sterols in brown macroalgae from Antarctica and their identification by liquid chromatography coupled with tandem mass spectrometry

This study reports for the first time the extraction and quantification of sterols in six species of brown macroalgae from Antarctica: Adenocystis utricularis, Ascoseira mirabilis, Cystosphaera jacquinotii, Desmarestia anceps, Desmarestia antarctica, and Himantothallus grandifolius. Ultrasound irradiation was used as a promotor to extract sterols from algal biomass. The extracts were analyzed by high-performance liquid chromatography coupled with tandem mass spectrometry (LC-MS/MS) for sterol quantization (ergosterol, brassicasterol, fucosterol, β-sitosterol, campesterol, cholesterol, and stigmasterol). In general, fucosterol was the most abundant (6.60 to 48.13 mg kg^?1), followed by β-sitosterol (5.29 to 16.49 mg kg^?1), stigmasterol (2.69 to 14.84 mg kg^?1) in five of the six examined algae. The sterol campesterol was found in lower concentrations (0.07 to 0.15 mg kg^?1) in all studied samples.     

Photosynthesis,dark respiration and light independent carbon fixation of endemic Antarctic macroalgae

Summary The light saturated photosynthesis, dark respiration and light independent carbon fixation of macroalgal species endemic to the Antarctic were measured. Five brown algae. Ascoseira mirabilis, Desmarestia anceps, D. antarctica, Phaeurus antarcticus, Himantothallus grandifolius and the red alga Palmaria decipiens were included. Rates of these three parameters at 0°C were very similar to those measured in other studies on temperate algae at higher temperature. This indicates a high degree of physiological adaptation to the Antarctic environment within these species. A comparison was made of polarographic and chemical means of measuring oxygen flux during photosynthesis and dark respiration at low temperature. There was a good correlation between measurements of oxygen evolution and carbon fixation, although apparent photosynthetic quotient values were in most cases high.Abbreviations HEPES N-(Hydroxyethyl) piperazine-N-(2 ethane-sulphonic acid) - RuBP D-ribulose 1,5-bisphosphate Contribution No. 415 from the Alfred-Wegener-Institut für Polaru. Meeresforschung     

Animal associations with the dominant species of shallow water macrophytes along the coastline of the Vestfold Hills,Antarctica

Benthic grab samples, taken through core holes in sea-ice 1983, returned eight species of marine macrophytes which had associated fauna on at least one sample. A total of 24 animal taxa were found on these eight macrophytes. Phyllophora antarctica had a significantly greater biomass of attached fauna per unit of macrophyte biomass than any of the other seven species. It also had the greatest variety of epiphytic fauna (21 taxa). Phyllophora antarctica was the only macrophyte collected from comparatively protected Ellis Fjord, apart from filamentous algae, including Cladophora subsimplex and Geminocarpus austrogeorgiae, which were found in the most landward basin. It therefore played an important role in the ecology of the fjord. Palmaria decipiens was commonly collected at more exposed coastal sites and was dominant in Long Fjord, characterized here as intermediate in terms of water movement. Sixteen animal taxa were collected from this macrophyte with the greatest numbers being found in July and August. Only four animal taxa were collected from Iridaea cordata. The other five macrophyte species from which epiphytic animals were collected (Porphyra endiviifolium, Plocamium cartilagineum, Desmarestia menziesii, Himantothallus grandifolius and Cladophora subsimplex had animals associated with them on less than half the occasions when they were found.     

Lack of Physiological Depth Patterns in Conspecifics of Endemic Antarctic Brown Algae: A Trade-Off between UV Stress Tolerance and Shade Adaptation?

A striking characteristic of endemic Antarctic brown algae is their broad vertical distribution. This feature is largely determined by the shade adaptation in order to cope with the seasonal variation in light availability. However, during spring-summer months, when light penetrates deep in the water column these organisms have to withstand high levels of solar radiation, including UV. In the present study we examine the light use characteristics in parallel to a potential for UV tolerance (measured as content of phenolic compounds, antioxidant activity and maximum quantum yield of fluorescence) in conspecific populations of four Antarctic brown algae (Ascoseira mirabilis, Desmarestia menziesii, D. anceps and Himantothallus grandifolius) distributed over a depth gradient between 5 and 30 m. The main results indicated that a) photosynthetic efficiency was uniform along the depth gradient in all the studied species, and b) short-term (6 h) exposure to UV radiation revealed a high tolerance measured as chlorophyll fluorescence, phlorotannin content and antioxidant capacity. Multivariate analysis of similarity indicated that light requirements for photosynthesis, soluble phlorotannins and antioxidant capacity are the variables determining the responses along the depth gradient in all the studied species. The suite of physiological responses of algae with a shallower distribution (A. mirabilis and D. menziesii) differed from those with deeper vertical range (D. anceps and H. grandifolius). These patterns are consistent with the underwater light penetration that defines two zones: 0–15 m, with influence of UV radiation (1% of UV-B and UV-A at 9 m and 15 m respectively) and a zone below 15 m marked by PAR incidence (1% up to 30 m). These results support the prediction that algae show a UV stress tolerance capacity along a broad depth range according to their marked shade adaptation. The high contents of phlorotannins and antioxidant potential appear to be strongly responsible for the lack of clear depth patterns in light demand characteristics and UV tolerance.     

Photosynthesis of Marine Macroalgae from Antarctica: Light and Temperature Requirements

The photosynthetic performance of macroalgae isolated in Antarctica was studied in the laboratory. Species investigated were the brown algae Himantothallus grandifolius, Desmarestia anceps, Ascoseira mirabilis, the red algae Palmaria decipiens, Iridaea cordata, Gigartina skottsbergii, and the green algae Enteromorpha bulbosa, Acrosiphonia arcta, Ulothrix subflaccida and U. implexa. Unialgal cultures of the brown and red algae were maintained at 0°C, the green algae were cultivated at 10°C. I_K values were between 18 and 53 μmol m^?2 s^?1 characteristic or low light adapted algae. Only the two Ulothrix species showed higher I_K values between 70 and 74 μmol m^?2 s^?1. Photosynthesis compensated dark respiration at very low photon fluence rates between 1.6 and 10.6 μmol m^?2 s^?1. Values of α were high: between 0.4 and 1.1 μmol O₂ g^?1 FW h^?1 (μmol m^?2 s^?1)^?1 in the brown and red algae and between 2.1 and 4.9 μmol O₂ g^?1 FW h^?1 (μmol m^?2 s^?1)^?1 in the green algal species. At 0°C P_max values of the brown and red algae ranged from 6.8 to 19.1 μmol O₂ g^?1 FW h^?1 and were similarly high or higher than those of comparable Arctic-cold temperate species. Optimum temperatures for photosynthesis were 5 to 10°C in A. mirabilis, 10°C in H. grandifolius, 15°C in G. skottsbergii and 20°C or higher in D. anceps and I. cordata. P: R ratios strongly decreased in most brown and red algae with increasing temperatures due to different Q₁₀ values for photosynthesis (1.4 to 2.5) and dark respiration (2.5 to 4.1). These features indicate considerable physiological adaptation to the prevailing low light conditions and temperatures of Antarctic waters. In this respect the lower depth distribution limits and the northern distribution boundaries of these species partly depend on the physiological properties described here.     

Seaweed biodiversity in the south-western Antarctic Peninsula: surveying macroalgal community composition in the Adelaide Island/Marguerite Bay region over a 35-year time span

The diversity of seaweed species of the south-western Antarctic Peninsula region is poorly studied, contrasting with the substantial knowledge available for the northern parts of the Peninsula. However, this is a key region affected by contemporary climate change. Significant consequences of this change include sea ice recession, increased iceberg scouring and increased inputs of glacial melt water, all of which can have major impacts on benthic communities. We present a baseline seaweed species checklist for the southern Adelaide Island and northern Marguerite Bay region, combining data obtained during a small number of surveys completed in 1973–1975 and a 6-week intensive diving-based field campaign in 2010–2011. Overall, with a total of 41 macroalgal species recorded (7 brown, 27 red, 6 green, 1 chrysophyte), the region is species-poor compared to the north of the Antarctic Peninsula, and even more so in comparison with the sub-Antarctic. The key canopy-forming species is Desmarestia menziesii, which is abundant in Antarctic Peninsula waters, but lacking in the sub-Antarctic. Himantothallus grandifolius, which is a common species further north in the Antarctic phytobenthos, was absent in our recent collections. This paper also reports the first record of Aplanochytrium sp. (Labyrinthulomycetes) from this part of Antarctica and in association with Elachista sp.     

Direct observations on the sublittoral marine algae of Argyll,Scotland

Summary The distribution with depth of sublittoral marine algae has been investigated at 11 sites on the west coast of Scotland. More than half of the species found were confined to the sublittoral zone. The greatest variety of species was found in shallow water at sites sheltered from excessive turbulence. With increasing depth the number of species found steadily decreased. There was no evidence of a specifically distinct algal flora confined to deeper water. No algae were found deeper than 36 m below ELWS at any site, but the lower limit of algal growth was reduced to 9 m in a turbid water loch and to 3 m at a site where the herbivorous echinoderm Ophiocomina nigra was abundant. Fairly distinct algal communities were found on different substrates. The major communities recognised underwater were algae attached to stable substrata, algae attached to unstable substrata and epiphytic on other algae, especially on the stipes of Laminaria hyperborea. On stable rock in areas subject to water movement L. hyperborea forest was the dominant vegetation whereas on unstable substrata and in sheltered localities L. saccharina was dominant.     

Variation in phlorotannin content within two species of brown macroalgae (Desmarestia anceps and D. menziesii) from the Western Antarctic Peninsula

Phlorotannins are metabolites found only in the Phaeophyceae (brown algae) and have a variety of metabolic roles, including both primary (e.g. cell wall construction) and secondary (e.g. herbivore defence and UV protection). This study aimed to establish the level of variation of phlorotannins in two species of dominant Antarctic macroalgae (Desmarestia anceps and D. menziesii). Thirteen samples were taken from specific locations throughout the thallus of multiple individuals at two depths at three locations near Anvers Island, Antarctica. Overall average concentrations were 0.117±0.003 g g^–1dwt in D. anceps and 0.052±0.002 g g^–1dwt in D. menziesii. Concentrations varied greatly at the 13 sampling locations in each individual, but not in any consistent manner. Significant variation occurred between locations in both species and between depths in D. anceps.     

<Superscript>137</Superscript>Cs, <Superscript>40</Superscript>K, <Superscript>238</Superscript>Pu, <Superscript>239+240</Superscript>Pu and <Superscript>90</Superscript>Sr in biological samples from King George Island (Southern Shetlands) in Antarctica

There are few data reported on radionuclide contamination in Antarctica. The aim of this paper is to report ¹³⁷Cs, ⁹⁰Sr and ^238,239+240Pu and ⁴⁰K activity concentrations measured in biological samples collected from King George Island (Southern Shetlands, Antarctica), mostly during 2001–2002. The samples included: bones, eggshells and feathers of penguin Pygoscelis papua, bones and feathers of petrel Daption capense, bones and fur of seal Mirounga leonina, algae Himantothallus grandifolius, Desmarestia anceps and Cystosphaera jacquinotii, fish Notothenia corriceps, sea invertebrates Amphipoda, shells of limpet Nacella concina, lichen Usnea aurantiaco-atra, vascular plants Deschampsia antarctica and Colobanthus quitensis, fungi Omphalina pyxidata, moss Sanionia uncinata and soil. The results show a large variation in some activity concentrations. Samples from the marine environment had lower contamination levels than those from terrestrial ecosystems. The highest activity concentrations for all radionuclides were found in lichen and, to a lesser extent, in mosses, probably because lichens take up atmospheric pollutants and retain them. The only significant correlation (except for that expected between ²³⁸Pu and ^239+240Pu) was noted for moss and lichen samples between plutonium and ⁹⁰Sr. A tendency to a slow decrease with time seems to be occurring. Analyses of the activity ratios show varying fractionation between various radionuclides in different organisms. Algae were relatively more highly contaminated with plutonium and radiostrontium, and depleted with radiocesium. Feathers had the lowest plutonium concentrations. Radiostrontium and, to a lesser extent, Pu accumulated in bones. The present low intensity of fallout in Antarctic has a lower ²³⁸Pu/^239+240Pu activity ratio than that expected for global fallout.     

Desmarestia antarctica (Desmarestiales,Phaeophyceae), a new ligulate Antarctic species with an endophytic gametophyte

Desmarestia antarctica Moe & Silva, a new species from the Antarctic Peninsula and the Scotia Arc, is distinguished from all other ligulate members of the genus by the arrangement of zoosporangia, which are produced together with paraphyses in a raised sorus. Among all species in the genus, onlyD. anceps Montagne shares this feature.Desmarestia antarctica is the first species of the genus for which an endophytic gametophyte is demonstrated, the usual host beingCurdiea racovitzae De Wildeman, a red alga of the familyGracilariaceae. The sporophyte contains only a moderate concentration of acid compared to Northern Hemisphere ligulate species, while physodes that probably contain polyphenolic compounds form noticeable speckles and dark margins in preserved plants. Often more than one axis arises from a single holdfast, probably as the result of compounding rather than proliferation.Dedicated to Prof. DrLothar Geitler on the occasion of the 90th anniversary of his birthday.     

Studies toward the comprehension of fungal-macroalgae interaction in cold marine regions from a biotechnological perspective

In marine ecosystems, macroalgae are the habitat for several microorganisms, fungi being among them. In the Antarctic benthic coastal ecosystem, macroalgae play a key role in organic matter cycling. In this study, 13 different macroalgae from Potter Cove and surrounding areas were sampled and 48 fungal isolates were obtained from six species, four Rhodophyta Ballia callitricha, Gigartina skottsbergii, Neuroglossum delesseriae and Palmaria decipiens, and two Phaeophyceae: Adenocystis utricularis and Ascoseira mirabilis. Fungal isolates mostly belonged to the Ascomycota phylum (Antarctomyces, Cadophora, Cladosporium, Penicillium, Phialocephala, and Pseudogymnoascus) and only one to the phylum Mucoromycota. Two of the isolates could not be identified to genus level, implying that Antarctica is a source of probable novel fungal taxa with enormous bioprospecting and biotechnological potential. 73% of the fungal isolates were moderate eurypsychrophilic (they grew at 5–25 °C), 12.5% were eurypsychrophilic and grew in the whole range, 12.5% of the isolates were narrow eurypsychrophilic (growth at 15–25 °C), and Mucoromycota AUe4 was classified as stenopsychrophilic as it grew at 5–15 °C. Organic extracts of seven macroalgae from which no fungal growth was obtained (three red algae Georgiella confluens, Gymnogongrus turquetii, Plocamium cartlagineum, and four brown algae Desmarestia anceps, D. Antarctica, Desmarestia menziesii, Himantothallus grandifolius) were tested against representative fungi of the genera isolated in this work. All extracts presented fungal inhibition, those from Plocamium cartilagineum and G. turquetii showed the best results, and for most of these macroalgae, this represents the first report of antifungal activity and constitute a promising source of compounds for future evaluation.     

Algal canopy as a proxy for the disturbance history of understorey communities in East Antarctica

Much of the macroalgal zonation on Antarctic coasts is thought to be maintained by ice scour. The frequency and severity of ice scour typically decrease with depth, which is hypothesized to drive the zonation of two canopy-forming macroalgae, Desmarestia menziesii and Himantothallus grandifolius. If true, understorey communities should share the same history of ice scour as their respective canopies, and their composition should vary accordingly. To evaluate this prediction we collected boulders from under each canopy species at two depths, 6 and 12 m, at two sites on the coast of East Antarctica. We examined the hard-substrate communities growing on boulders and tested for differences in community composition with respect to canopy species, surface orientation, and depth. Communities under the different canopies showed some variation consistent with the hypothesized difference in disturbance history. Those under H. grandifolius accommodated a greater abundance and diversity of sponges, which is usually characteristic of older, later successional communities. Differences were subtle, however, suggesting that canopies might be maintained by ice disturbance over large temporal scale relative to those at which understorey communities develop, and/or that canopies themselves influence understorey composition. This study describes patterns associated with one of the most prominent examples of bathymetric zonation in shallow Antarctic benthos, and experimental work is now needed to partition the processes at work.     

Antarctic patterns of shallow subtidal habitat and inhabitants in Wilke’s Land

Studies of east Antarctic marine assemblages on hard substrata are rare. In relation to sea-ice breakout, we assessed benthic patterns of habitat and inhabitants between islands and bays at each of two depths (6 and 12 m) across the Windmill Islands coast. Island sites experience sea-ice breakout in the austral spring, while bay sites typically retain sea-ice cover into the summer and in some places the cover is virtually permanent. Composition of assemblages differed between sheltered bays and exposed islands. Islands were dominated by macroalgae, which also varied with depth. Immediately below the ice–foot zone at 6 m, substratum space were monopolised by foliose red (Palmaria decipiens) and foliose brown (Desmarestia sp.) algae, whereas at 12 m large canopies of Himantothallus grandifolius was abundant. The understorey consisted of a mixture of turfs and encrusting red algae at 6 m, and coralline algae at 12 m. Sheltered bays had large areas of sediment/algal complex and no canopy-forming macroalgae. We found more sponges and hydroids in bays, and more brittle stars around islands. Experiments testing factors that covary with exposure and depth in Antarctica, such as light, sedimentation and ice scour are necessary to determine processes that maintain these striking patterns.     

LACK OF DEFENSE OR PHLOROTANNIN INDUCTION BY UV RADIATION OR MESOGRAZERS IN DESMARESTIA ANCEPS AND D. MENZIESII (PHAEOPHYCEAE)1

Phlorotannins are polyphenoloic metabolites occurring only in the Phaeophyceae that have numerous putative primary roles (e.g. cell‐wall construction and storage) as well as secondary metabolic roles, which include herbivore feeding deterrence and protection from UV radiation. The proposed role of phlorotannins in the defense against UV radiation is of particular importance in the Antarctic due to depletion of the stratospheric ozone layer in that area. Several studies of brown algae have found evidence of an induction response (the production of defensive metabolites, including phlorotannins) after grazing by various mesograzers, after simulated grazing/wounding, and after exposure to increases in UV radiation. This study aimed to determine if phlorotannin production or other defenses in two dominant, endemic Antarctic species (Desmarestia menziesii Montagne and Desmarestia anceps J. Agardh) could be induced by an increase in exposure to UV radiation or by natural and artificial grazing. An in situ experiment failed to detect any effect of UV radiation on phlorotannin concentrations in either species or on subsequent palatability in feeding bioassays. A laboratory‐based experiment did not detect any effect of mesoherbivore grazing or simulated grazing (wounding) on palatability or the concentration of phlorotannins in D. menziesii. Instead, phlorotannin concentrations increased in all treatments in both experiments, consistent with an increase in overall resource availability due to an increase in available PAR compared with the in situ irradiance at the algal collection sites.     

The phylogenetic position of the subantarctic marine macroalga Desmarestia chordalis (Phaeophyceae) inferred from nuclear ribosomal ITS sequences

The phylogenetic position of the subantarctic brown macroalga Desmarestia chordalis was inferred from nuclear ribosomal DNA internal transcribed spacer sequences, which were compared with published homologous sequences of putative relatives. According to the molecular data, D. chordalis is closely related to the Antarctic species D. menziesii, and more distantly related to other taxa of similar morphology, i.e. Antarctic D. anceps and northern hemisphere D. aculeata. Accepted: 2 August 1999     

Polychaete fauna associated with holdfasts of the large brown alga Himantothallus grandifolius in Admiralty Bay, King George Island, Antarctic

The polychaete community associated with holdfasts of the brown alga Himantothallus grandifolius in Admiralty Bay has been studied. It is the first study of its kind in this area and only the second in the Antarctic. Samples were collected in the summer season of 1979/1980 from a depth range of 10–75 m. Seventy-eight species were found on 19 holdfasts. The community was dominated by Brania rhopalophora and Neanthes kerguelensis. Analysis of similarity showed that polychaete fauna associated with this habitat did not show any partitioning related to depth. Regression analysis showed that densities of both species and individuals decreased with increased holdfast volume. A positive correlation was found between the number of individuals and holdfast volume. Polychaetes from 10 feeding guilds were found with dominance of macrophagous motile herbivores and sessile filter feeders. The complex habitat provided by holdfasts is a shelter for a rich polychaete fauna and may function as important protection from disturbance in the shallow areas of Admiralty Bay.     

Fungal community associated with marine macroalgae from Antarctica

Filamentous fungi and yeasts associated with the marine algae Adenocystis utricularis, Desmarestia anceps, and Palmaria decipiens from Antarctica were studied. A total of 75 fungal isolates, represented by 27 filamentous fungi and 48 yeasts, were isolated from the three algal species and identified by morphological, physiological, and sequence analyses of the internal transcribed spacer region and D1/D2 variable domains of the large-subunit rRNA gene. The filamentous fungi and yeasts obtained were identified as belonging to the genera Geomyces, Antarctomyces, Oidiodendron, Penicillium, Phaeosphaeria, Aureobasidium, Cryptococcus, Leucosporidium, Metschnikowia, and Rhodotorula. The prevalent species were the filamentous fungus Geomyces pannorum and the yeast Metschnikowia australis. Two fungal species isolated in our study, Antarctomyces psychrotrophicus and M. australis, are endemic to Antarctica. This work is the first study of fungi associated with Antarctic marine macroalgae, and contributes to the taxonomy and ecology of the marine fungi living in polar environments. These fungal species may have an important role in the ecosystem and in organic matter recycling.