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Representative microprofile curves are illustrated. Full size image Long Desulfobulbaceae filaments in natural sediments At all sites with geochemical evidence of electrogenic sulphide oxidation, microscopic examination of the sediments revealed long, unbranched, filamentous bacteria Figure 2a , which were subsequently identified by CARD—FISH as Desulfobulbaceae Figure 2b ; Supplementary Figures S1 and S2.

Enumeration using FISH revealed densities of No filaments were detected at the OSF reference site, where there was no geochemical evidence of electrogenic sulphide oxidation.

SEM images revealed a pattern of evenly spaced external ridges running in parallel along the length of the filaments isolated from all three sites Figure 2c ; Supplementary Figures S1 and S2. Cell diameters were 1. Filaments with larger cell diameters had more ridges and we estimated across all samples between 16 and 58 ridges per filament.

Figure 2 Identification of filamentous Desulfobulbaceae bacteria present in intact sediment from sites described in Figure 1 and Table 1. All images shown here are from MLG.

Closely related sequences of filamentous Desulfobulbaceae from Aarhus Bay and other isolated bacteria are also included.

A summary of the full clone library composition is given in Supplementary Figure 4. A clone library was not constructed from OSF sediments, where the geochemical signature of electrogenic sulphur oxidation was not observed. However, a previous study of similar bioturbated sediments in the Oosterschelde found no Desulfobulbaceae sequences in either the surface-oxidized sediment or the deeper sulphate-reducing sediment Miyatake et al.

Effects of bioturbation on geochemical signal The geochemical fingerprint of electrogenic sulphur oxidation was not present in intact sediment cores collected from the highly bioturbated sandy sediment at OSF collected in May and August However, sediments from OSF that were defaunated by sieving and incubated under darkened laboratory conditions in aerated seawater developed the pH profile indicative of electrogenic sulphur oxidation Figure 3.

The characteristic pH profile indicative of cathodic oxygen consumption was first evident on Day 5 after homogenization, whereby porewater pH was just beginning to increase at the base of the oxic zone Figure 3. This pH peak progressively increased over the days of incubation, with a maximum subsurface pH observed on Day At the start of the incubation, before the onset of cathodic oxygen consumption, the DOU declined from 6.

After Day 5, in concert with the development of a pH maximum in the oxic zone, there was an increase in DOU. Over this same period Days 5—20 , the estimated electrical current density increased from 0. Although CARD—FISH was not performed during this particular experiment, subsequent analysis of sediment collected from OSF and sieved and incubated following the same procedures, confirmed the presence of Desulfobulbaceae filaments whenever the geochemical signature of electrogenic sulphur oxidation was present Supplementary Figure 3.

Figure 3 a A subset of microsensor profiles from sediment collected from a heavily bioturbated sediment from an OSF that has been artificially defaunated and incubated with well-oxygenated overlying water. Despite a low concentration of free sulphide, development of a geochemical signature of long-range electron transport is evident from Day 5 onwards. Full size image To test if infauna specifically can inhibit the geochemical signature of electrogenic sulphide oxidation, we performed a second manipulation experiment, where fauna were first removed by asphyxiation and subsequently reintroduced.

When intact sediment cores were retrieved from the OSF field site and immediately profiled, these sediments did not exhibit evidence of electrogenic sulphur oxidation Figure 4a. In defaunated sediments, the geochemical signature of electrogenic sulphur oxidation again developed within 8 days of exposure to aerated seawater Figure 4b.

The deep burrowing lugworm A. In sediment patches that were not mechanically disrupted by the burrowing lugworms, the electrogenic signature persisted, though with attenuated pH extremes Figure 4c. In sediment patches that were directly overturned by lugworm bioturbation, the electrogenic signature disappeared entirely Figure 4d. Figure 4 Effect of sediment disturbance by fauna on electrogenic sulphur oxidation.

Full size image Discussion Our results provide evidence that electrogenic sulphur oxidation, previously reported from laboratory incubations Nielsen et al. We observed evidence of electrogenic sulphur oxidation, together with Desulfobulbaceae filaments, in a variety of coastal marine sediments that is, a salt marsh drainage channel, a subtidal site recovering from seasonal hypoxia, a coastal area of high mud deposition.

These sediments were characterized by high DOU and high organic matter content. In the narrow oxic zone 0. Below this depth, there was a suboxic zone where oxygen and sulphide were not detectable, which was typically 5—15 mm deep although sometimes extending deeper.

Below the suboxic zone, these sediments exhibited a steep sulphide gradient, indicating high sulphide production. Repeated visits to these three study sites identified that electrogenic sulphur oxidation is a regularly occurring, although not necessarily permanent, feature at these field sites.

Desulfobulbaceae filaments from the suboxic zone of our study sites revealed densities of These densities are similar to those reported in Pfeffer et al. The filamentous bacteria retrieved from the three electrogenic sulphur oxidation study sites RSM, BCZ and MLG were long and unbranched, and retrieved fragments could exceed eight millimetres in length. The unusual morphological feature of raised ridges running in parallel along the length of the filaments, observed previously in laboratory incubations, were also prominent in the Desulfobulbaceae filaments isolated in this study.

Pfeffer et al. It remains to be seen if these morphological differences represent genotypic or phenotypic variation.

A cosmopolitan distribution of electrogenic sulphur oxidation is supported by a number of published datasets in which the geochemical fingerprint of this process is evident, but has not been formally recognized Figure 1d. Shallow subsurface pH maxima, characteristic of cathodic oxygen consumption, have been reported from in situ microsensor profiling in the Santa Barbara Cai et al.

These basins are permanently hypoxic, support high rates of sulphate reduction, and are devoid of bioturbating fauna. Similar pH profiles were also reported from a site situated within an oyster aquaculture park in the Thau lagoon Mediterranean Sea, France; Dedieu, and from sediments near a fish farm in Pillan fjord Chile; Mulsow et al.

Seasonally hypoxic sediments from Tokyo Bay have also demonstrated microprofile evidence of cathodic oxygen consumption Sayama, , similar to those reported here from MLG. At a site characterized by high sulphide fluxes and devoid of large bioturbating infauna on the Mid-Atlantic ridge, high rates of sulphide oxidation were detected which could not be attributed to any other known sulphide removal process, such as sulphur oxidation by large nitrate-accumulating bacteria Schauer et al.

One explanation is that electrogenic sulphur oxidations is responsible for the observed sulphide removal, however, direct evidence is needed to confirm this possibility. Gene sequence archives also support a cosmopolitan distribution of the conductive filamentous bacteria in locations with high rates of sulphide generation, via high organic matter loading or sulphide seepage, and a paucity of bioturbating animals. Previous laboratory experiments have shown that sulphur oxidation by long-range electron transport can play a dominant role in oxygen consumption and can exert a strong imprint on the overall sulphur cycling and pH dynamics in sediments Nielsen et al.

Our observations show that this process likely has a similarly profound effect on mineral cycling in some natural coastal sediments. We observed pH values as high as 8. Filaments with larger cell diameters had more ridges and we estimated across all samples between 16 and 58 ridges per filament. Figure 2 Identification of filamentous Desulfobulbaceae bacteria present in intact sediment from sites described in Figure 1 and Table 1.

All images shown here are from MLG. Closely related sequences of filamentous Desulfobulbaceae from Aarhus Bay and other isolated bacteria are also included. A summary of the full clone library composition is given in Supplementary Figure 4. A clone library was not constructed from OSF sediments, where the geochemical signature of electrogenic sulphur oxidation was not observed.

However, a previous study of similar bioturbated sediments in the Oosterschelde found no Desulfobulbaceae sequences in either the surface-oxidized sediment or the deeper sulphate-reducing sediment Miyatake et al. Effects of bioturbation on geochemical signal The geochemical fingerprint of electrogenic sulphur oxidation was not present in intact sediment cores collected from the highly bioturbated sandy sediment at OSF collected in May and August However, sediments from OSF that were defaunated by sieving and incubated under darkened laboratory conditions in aerated seawater developed the pH profile indicative of electrogenic sulphur oxidation Figure 3.

The characteristic pH profile indicative of cathodic oxygen consumption was first evident on Day 5 after homogenization, whereby porewater pH was just beginning to increase at the base of the oxic zone Figure 3.

This pH peak progressively increased over the days of incubation, with a maximum subsurface pH observed on Day At the start of the incubation, before the onset of cathodic oxygen consumption, the DOU declined from 6. After Day 5, in concert with the development of a pH maximum in the oxic zone, there was an increase in DOU.

Over this same period Days 5—20 , the estimated electrical current density increased from 0. Although CARD—FISH was not performed during this particular experiment, subsequent analysis of sediment collected from OSF and sieved and incubated following the same procedures, confirmed the presence of Desulfobulbaceae filaments whenever the geochemical signature of electrogenic sulphur oxidation was present Supplementary Figure 3.

Figure 3 a A subset of microsensor profiles from sediment collected from a heavily bioturbated sediment from an OSF that has been artificially defaunated and incubated with well-oxygenated overlying water. Despite a low concentration of free sulphide, development of a geochemical signature of long-range electron transport is evident from Day 5 onwards.

Full size image To test if infauna specifically can inhibit the geochemical signature of electrogenic sulphide oxidation, we performed a second manipulation experiment, where fauna were first removed by asphyxiation and subsequently reintroduced. When intact sediment cores were retrieved from the OSF field site and immediately profiled, these sediments did not exhibit evidence of electrogenic sulphur oxidation Figure 4a.

In defaunated sediments, the geochemical signature of electrogenic sulphur oxidation again developed within 8 days of exposure to aerated seawater Figure 4b. The deep burrowing lugworm A. In sediment patches that were not mechanically disrupted by the burrowing lugworms, the electrogenic signature persisted, though with attenuated pH extremes Figure 4c.

In sediment patches that were directly overturned by lugworm bioturbation, the electrogenic signature disappeared entirely Figure 4d. Figure 4 Effect of sediment disturbance by fauna on electrogenic sulphur oxidation. Full size image Discussion Our results provide evidence that electrogenic sulphur oxidation, previously reported from laboratory incubations Nielsen et al.

We observed evidence of electrogenic sulphur oxidation, together with Desulfobulbaceae filaments, in a variety of coastal marine sediments that is, a salt marsh drainage channel, a subtidal site recovering from seasonal hypoxia, a coastal area of high mud deposition.

These sediments were characterized by high DOU and high organic matter content. In the narrow oxic zone 0.

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Below this depth, there was a suboxic zone where oxygen and sulphide were not detectable, which was typically 5—15 mm deep although sometimes extending deeper. Below the suboxic zone, these sediments exhibited a steep sulphide gradient, indicating high sulphide production.

Repeated visits to these three study sites identified that electrogenic sulphur oxidation is a regularly occurring, although not necessarily permanent, feature at these field sites. Desulfobulbaceae filaments from the suboxic zone of our study sites revealed densities of These densities are similar to those reported in Pfeffer et al. The filamentous bacteria retrieved from the three electrogenic sulphur oxidation study sites RSM, BCZ and MLG were long and unbranched, and retrieved fragments could exceed eight millimetres in length.

The unusual morphological feature of raised ridges running in parallel along the length of the filaments, observed previously in laboratory incubations, were also prominent in the Desulfobulbaceae filaments isolated in this study. Pfeffer et al. It remains to be seen if these morphological differences represent genotypic or phenotypic variation.

A cosmopolitan distribution of electrogenic sulphur oxidation is supported by a number of published datasets in which the geochemical fingerprint of this process is evident, but has not been formally recognized Figure 1d. Shallow subsurface pH maxima, characteristic of cathodic oxygen consumption, have been reported from in situ microsensor profiling in the Santa Barbara Cai et al. These basins are permanently hypoxic, support high rates of sulphate reduction, and are devoid of bioturbating fauna.

Similar pH profiles were also reported from a site situated within an oyster aquaculture park in the Thau lagoon Mediterranean Sea, France; Dedieu, and from sediments near a fish farm in Pillan fjord Chile; Mulsow et al. Seasonally hypoxic sediments from Tokyo Bay have also demonstrated microprofile evidence of cathodic oxygen consumption Sayama, , similar to those reported here from MLG.

At a site characterized by high sulphide fluxes and devoid of large bioturbating infauna on the Mid-Atlantic ridge, high rates of sulphide oxidation were detected which could not be attributed to any other known sulphide removal process, such as sulphur oxidation by large nitrate-accumulating bacteria Schauer et al.

One explanation is that electrogenic sulphur oxidations is responsible for the observed sulphide removal, however, direct evidence is needed to confirm this possibility. Gene sequence archives also support a cosmopolitan distribution of the conductive filamentous bacteria in locations with high rates of sulphide generation, via high organic matter loading or sulphide seepage, and a paucity of bioturbating animals.

Previous laboratory experiments have shown that sulphur oxidation by long-range electron transport can play a dominant role in oxygen consumption and can exert a strong imprint on the overall sulphur cycling and pH dynamics in sediments Nielsen et al. Our observations show that this process likely has a similarly profound effect on mineral cycling in some natural coastal sediments.

We observed pH values as high as 8. These pH extremes lead to dissolution of metal sulphides and calcium carbonate within the deep suboxic zone, followed by re-precipitation of iron hydr oxides and carbonate near the sediment—water interface Risgaard-Petersen et al.

High alkalinity effluxes have been reported from muddy sites on the BCZ, including Station Braeckman et al. We estimated that a minimum of 5. Our data further suggest that bioturbation may exert a major control on the natural distribution of electrogenic sulphur oxidation in coastal marine sediments. In heavily bioturbated sediment from OSF, the geochemical signature of electrogenic sulphide oxidation was not detected under field conditions, but could be induced in laboratory experiments, as long as bioturbation was excluded.

In these sediments, a seed population of Desulfobulbaceae bacteria must have existed, from which these filaments were able to proliferate when burrowing animals are removed.

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This website uses cookies to provide you with the best possible experience and to optimize the website to best fit the needs of our visitors. By using this website, you automatically agree to the use of cookies. For detailed information on the use of cookies on this website, please see our Privacy Policy. OK Close. First Name:These basins are permanently hypoxic, support high rates of sulphate reduction, and are devoid of bioturbating fauna.

Thus, the constant voyage enables one to voyage the true essence. In laboratory experiments, the development of this characteristic geochemical fingerprint has been used as a sign of electrogenic sulphur oxidation activity Nielsen et al. When intact sediment cores were retrieved from the OSF field site and immediately profiled, these sediments did not exhibit evidence of electrogenic sulphur oxidation Figure 4a.

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In sediment patches that were directly overturned by lugworm bioturbation, the electrogenic signature disappeared entirely Figure 4d. Compelling evidence indicates that electrogenic sulphur oxidation is likely carried out by long filamentous bacteria. Repeated visits to these three study sites identified that electrogenic sulphur oxidation is a regularly occurring, although not necessarily permanent, feature at these field sites.