@article{BickelTangGrossart2014, author = {Bickel, Samantha L. and Tang, Kam W. and Grossart, Hans-Peter}, title = {Structure and function of zooplankton-associated bacterial communities in a temperate estuary change more with time than with zooplankton species}, series = {Aquatic microbial ecology : international journal}, volume = {72}, journal = {Aquatic microbial ecology : international journal}, number = {1}, publisher = {Institute of Mathematical Statistics}, address = {Oldendorf Luhe}, issn = {0948-3055}, doi = {10.3354/ame01676}, pages = {1 -- 15}, year = {2014}, abstract = {Zooplankton support distinct bacterial communities in high concentrations relative to the surrounding water, but little is known about how the compositions and functionalities of these bacterial communities change through time in relation to environmental conditions. We conducted a year-long field study of bacterial communities associated with common zooplankton groups as well as free-living bacterial communities in the York River, a tributary of Chesapeake Bay. Bacterial community genetic fingerprints and their carbon substrate usage were examined by denaturing gradient gel electrophoresis (DGGE) of amplified 16S rDNA and by Biolog EcoPlates, respectively. Zooplankton-associated communities were genetically distinct from free-living bacterial communities but utilized a similar array of carbon substrates. On average, bacteria associated with different zooplankton groups were genetically more similar to each other within each month (65.4\% similarity) than to bacterial communities of the same zooplankton group from different months (28 to 30\% similarity), which suggests the importance of ambient environmental conditions in shaping resident zooplankton-associated bacterial communities. Monthly changes in carbon substrate utilization were less variable for zooplankton-associated bacteria than for free-living bacteria, suggesting that the zooplankton microhabitat is more stable than the surrounding water and supports specific bacterial groups in the otherwise unfavorable conditions in the water column.}, language = {en} } @article{BickelTangGrossart2012, author = {Bickel, Samantha L. and Tang, Kam W. and Grossart, Hans-Peter}, title = {Ciliate epibionts associated with crustacean zooplankton in German lakes - distribution, motility, and bacterivory}, series = {Frontiers in microbiology}, volume = {3}, journal = {Frontiers in microbiology}, publisher = {Frontiers Research Foundation}, address = {Lausanne}, issn = {1664-302X}, doi = {10.3389/fmicb.2012.00243}, pages = {11}, year = {2012}, abstract = {Ciliate epibionts associated with crustacean zooplankton are widespread in aquatic systems, but their ecological roles are little known. We studied the occurrence of ciliate epibionts on crustacean zooplankton in nine German lakes with different limnological features during the summer of 2011. We also measured the detachment and re-attachment rates of the ciliates, changes in their motility, and the feeding rates of attached vs. detached ciliate epibionts. Epibionts were found in all lakes sampled except an acidic lake with large humic inputs. Epibiont prevalence was as high as 80.96\% on the cladoceran Daphnia cucullata, 67.17\% on the cladoceran Diaphanosoma brachyurum, and 46.67\% on the calanoid copepod Eudiaptomus gracilis. Both cladoceran groups typically had less than 10 epibionts per individual, while the epibiont load on E. gracilis ranged from 1 to >30 epibionts per individual. After the death of the zooplankton host, the peritrich ciliate epibiont Epistylis sp. detached in an exponential fashion with a half-life of 5 min, and 98\% detached within 30 min, leaving behind the stalks used for attachment. Immediately after detachment, the ciliates were immotile, but 62\% became motile within 60 min. When a new host was present, only 27\% reattached after 120 min. The average measured ingestion rate and clearance rate of Epistylis were 11,745 bacteria ciliate(-1) h(-1) and 24.33 mu l ciliate(-1) h(-1), respectively. Despite their high feeding rates, relatively low epibiont abundances were observed in the field, which suggests either diversion of energy to stalk formation, high metabolic loss by the epibionts, or high mortality among the epibiont populations.}, language = {en} } @article{DubovskayaTangGladyshevetal.2015, author = {Dubovskaya, Olga P. and Tang, Kam W. and Gladyshev, Michail I. and Kirillin, Georgiy and Buseva, Zhanna and Kasprzak, Peter and Tolomeev, Aleksandr P. and Grossart, Hans-Peter}, title = {Estimating In Situ Zooplankton Non-Predation Mortality in an Oligo-Mesotrophic Lake from Sediment Trap Data: Caveats and Reality Check}, series = {PLoS one}, volume = {10}, journal = {PLoS one}, number = {7}, publisher = {PLoS}, address = {San Fransisco}, issn = {1932-6203}, doi = {10.1371/journal.pone.0131431}, pages = {17}, year = {2015}, abstract = {Background Mortality is a main driver in zooplankton population biology but it is poorly constrained in models that describe zooplankton population dynamics, food web interactions and nutrient dynamics. Mortality due to non-predation factors is often ignored even though anecdotal evidence of non-predation mass mortality of zooplankton has been reported repeatedly. One way to estimate non-predation mortality rate is to measure the removal rate of carcasses, for which sinking is the primary removal mechanism especially in quiescent shallow water bodies. Objectives and Results We used sediment traps to quantify in situ carcass sinking velocity and non-predation mortality rate on eight consecutive days in 2013 for the cladoceran Bosmina longirostris in the oligo-mesotrophic Lake Stechlin; the outcomes were compared against estimates derived from in vitro carcass sinking velocity measurements and an empirical model correcting in vitro sinking velocity for turbulence resuspension and microbial decomposition of carcasses. Our results show that the latter two approaches produced unrealistically high mortality rates of 0.58-1.04 d(-1), whereas the sediment trap approach, when used properly, yielded a mortality rate estimate of 0.015 d(-1), which is more consistent with concurrent population abundance data and comparable to physiological death rate from the literature. Ecological implications Zooplankton carcasses may be exposed to water column microbes for days before entering the benthos; therefore, non-predation mortality affects not only zooplankton population dynamics but also microbial and benthic food webs. This would be particularly important for carbon and nitrogen cycles in systems where recurring mid-summer decline of zooplankton population due to non-predation mortality is observed.}, language = {en} } @article{DziallasGrossartTangetal.2013, author = {Dziallas, Claudia and Grossart, Hans-Peter and Tang, Kam W. and Nielsen, Torkel Gissel}, title = {Distinct Communities of Free-Living and Copepod-Associated Microorganisms along a Salinity Gradient in Godthabsfjord, West Greenland}, series = {ARCTIC ANTARCTIC AND ALPINE RESEARCH}, volume = {45}, journal = {ARCTIC ANTARCTIC AND ALPINE RESEARCH}, number = {4}, publisher = {INST ARCTIC ALPINE RES}, address = {BOULDER}, issn = {1523-0430}, doi = {10.1657/1938-4246.45.4.471}, pages = {471 -- 480}, year = {2013}, abstract = {Microorganisms such as Bacteria and Archaea play important roles in the Arctic food web and biogeochemical cycles. Nevertheless, knowledge of microbial community composition in Greenland waters is scarce, and information on microorganisms associated with Arctic zooplankton species is virtually non-existent. We compared free-living microbial communities with those associated with two key copepod species (Calanus finmarchicus and Metridia longa) along a salinity gradient from the deep waters beyond Fyllas Banke to the inner part of Godthabsfjord, West Greenland, in summer 2008. Using genetic fingerprinting we found that free-living Bacteria (in particular Alphaproteobacteria) and Archaea varied with environmental factors and formed different communities along the fjord. Microbial communities associated with the two copepod species were clearly different from those in the ambient water. Surprisingly, Archaea could not be detected on the copepods. Our results show that zooplankton form "microbial islands" in the Arctic pelagic realm with a distinctive community composition and presumably functionality different from the free-living Bacteria. Changes in intensity and timing of meltwater runoff due to global warming are expected to affect these microbial assemblages differently, with potentially significant ramifications for Arctic food webs and biogeochemistry.}, language = {en} } @article{GludGrossartLarsenetal.2015, author = {Glud, Ronnie N. and Grossart, Hans-Peter and Larsen, Morten and Tang, Kam W. and Arendt, Kristine E. and Rysgaard, Soren and Thamdrup, Bo and Gissel Nielsen, Torkel}, title = {Copepod carcasses as microbial hot spots for pelagic denitrification}, series = {Limnology and oceanography}, volume = {60}, journal = {Limnology and oceanography}, number = {6}, publisher = {Wiley-Blackwell}, address = {Hoboken}, issn = {0024-3590}, doi = {10.1002/lno.10149}, pages = {2026 -- 2036}, year = {2015}, abstract = {Copepods are exposed to a high non-predatory mortality and their decomposing carcasses act as microniches with intensified microbial activity. Sinking carcasses could thereby represent anoxic microenvironment sustaining anaerobic microbial pathways in otherwise oxic water columns. Using non-invasive O-2 imaging, we document that carcasses of Calanus finmarchicus had an anoxic interior even at fully air-saturated ambient O-2 level. The extent of anoxia gradually expanded with decreasing ambient O-2 levels. Concurrent microbial sampling showed the expression of nitrite reductase genes (nirS) in all investigated carcass samples and thereby documented the potential for microbial denitrification in carcasses. The nirS gene was occasionally expressed in live copepods, but not as consistently as in carcasses. Incubations of sinking carcasses in (15)NO3-amended seawater demonstrated denitrification, of which on average 34\%+/- 17\% (n=28) was sustained by nitrification. However, the activity was highly variable and was strongly dependent on the ambient O-2 levels. While denitrification was present even at air-saturation (302 mol L-1), the average carcass specific activity increased several orders of magnitude to approximate to 1 nmol d(-1) at 20\% air-saturation (55 mol O-2 L-1) at an ambient temperature of 7 degrees C. Sinking carcasses of C. finmarchicus therefore represent hotspots of pelagic denitrification, but the quantitative importance as a sink for bioavailable nitrogen is strongly dependent on the ambient O-2 level. The importance of carcass associated denitrification could be highly significant in O-2 depleted environments such as Oxygen Minimum Zones (OMZ).}, language = {en} } @article{GrossartFrindteDziallasetal.2011, author = {Grossart, Hans-Peter and Frindte, Katharina and Dziallas, Claudia and Eckert, Werner and Tang, Kam W.}, title = {Microbial methane production in oxygenated water column of an oligotrophic lake}, series = {Proceedings of the National Academy of Sciences of the United States of America}, volume = {108}, journal = {Proceedings of the National Academy of Sciences of the United States of America}, number = {49}, publisher = {National Acad. of Sciences}, address = {Washington}, issn = {0027-8424}, doi = {10.1073/pnas.1110716108}, pages = {19657 -- 19661}, year = {2011}, abstract = {The prevailing paradigm in aquatic science is that microbial methanogenesis happens primarily in anoxic environments. Here, we used multiple complementary approaches to show that microbial methane production could and did occur in the well-oxygenated water column of an oligotrophic lake (Lake Stechlin, Germany). Oversaturation of methane was repeatedly recorded in the well-oxygenated upper 10 m of the water column, and the methane maxima coincided with oxygen oversaturation at 6 m. Laboratory incubations of unamended epilimnetic lake water and inoculations of photoautotrophs with a lake-enrichment culture both led to methane production even in the presence of oxygen, and the production was not affected by the addition of inorganic phosphate or methylated compounds. Methane production was also detected by in-lake incubations of lake water, and the highest production rate was 1.8-2.4 nM.h(-1) at 6 m, which could explain 33-44\% of the observed ambient methane accumulation in the same month. Temporal and spatial uncoupling between methanogenesis and methanotrophy was supported by field and laboratory measurements, which also helped explain the oversaturation of methane in the upper water column. Potentially methanogenic Archaea were detected in situ in the oxygenated, methane-rich epilimnion, and their attachment to photoautotrophs might allow for anaerobic growth and direct transfer of substrates for methane production. Specific PCR on mRNA of the methyl coenzyme M reductase A gene revealed active methanogenesis. Microbial methane production in oxygenated water represents a hitherto overlooked source of methane and can be important for carbon cycling in the aquatic environments and water to air methane flux.}, language = {en} } @article{GuenthelDonisKirillinetal.2019, author = {G{\"u}nthel, Marco and Donis, Daphne and Kirillin, Georgiy and Ionescu, Danny and Bizic, Mina and McGinnis, Daniel F. and Grossart, Hans-Peter and Tang, Kam W.}, title = {Contribution of oxic methane production to surface methane emission in lakes and its global importance}, series = {Nature Communications}, volume = {10}, journal = {Nature Communications}, publisher = {Nature Publishing Group UK}, address = {London}, issn = {2041-1723}, doi = {10.1038/s41467-019-13320-0}, pages = {10}, year = {2019}, abstract = {Recent discovery of oxic methane production in sea and lake waters, as well as wetlands, demands re-thinking of the global methane cycle and re-assessment of the contribution of oxic waters to atmospheric methane emission. Here we analysed system-wide sources and sinks of surface-water methane in a temperate lake. Using a mass balance analysis, we show that internal methane production in well-oxygenated surface water is an important source for surface-water methane during the stratified period. Combining our results and literature reports, oxic methane contribution to emission follows a predictive function of littoral sediment area and surface mixed layer volume. The contribution of oxic methane source(s) is predicted to increase with lake size, accounting for the majority (>50\%) of surface methane emission for lakes with surface areas >1 km(2).}, language = {en} } @article{KirillinGrossartTang2012, author = {Kirillin, Georgiy and Grossart, Hans-Peter and Tang, Kam W.}, title = {Modeling sinking rate of zooplankton carcasses effects of stratification and mixing}, series = {Limnology and oceanography}, volume = {57}, journal = {Limnology and oceanography}, number = {3}, publisher = {Wiley}, address = {Waco}, issn = {0024-3590}, doi = {10.4319/lo.2012.57.3.0881}, pages = {881 -- 894}, year = {2012}, abstract = {Using the carcass sinking rate and density determined in laboratory for several freshwater zooplankton species, we developed a model of zooplankton carcass sinking as affected by turbulence and stratification. The model was subsequently used to estimate the residence time of zooplankton carcasses in the water column of Lake Stechlin, a typical temperate dimictic lake in northeastern Germany. The residence time varied among the different species and was strongly affected by thermal stratification. At the peak of summer stratification, the carcasses stayed up to 5 d in the 70 m-deep water column before reaching the lake bottom. Residence time was long enough that zooplankton carcasses could serve as an important matter and energy source for bacteria in the lake's pelagic zone and hence have the potential to significantly affect aquatic carbon and nutrient cycling. The proposed model of sinking rates, based on physically sound relationships, can be easily applied to other passively sinking particles, and be integrated into large ecosystem models.}, language = {en} } @article{McGinnisFluryTangetal.2017, author = {McGinnis, Daniel F. and Flury, Sabine and Tang, Kam W. and Grossart, Hans-Peter}, title = {Porewater methane transport within the gas vesicles of diurnally migrating Chaoborus spp.}, series = {Scientific reports}, volume = {7}, journal = {Scientific reports}, publisher = {Nature Publ. Group}, address = {London}, issn = {2045-2322}, doi = {10.1038/srep44478}, pages = {7}, year = {2017}, abstract = {Diurnally-migrating Chaoborus spp. reach populations of up to 130,000 individuals m-2 in lakes up to 70 meters deep on all continents except Antarctica. Linked to eutrophication, migrating Chaoborus spp. dwell in the anoxic sediment during daytime and feed in the oxic surface layer at night. Our experiments show that by burrowing into the sediment, Chaoborus spp. utilize the high dissolved gas partial pressure of sediment methane to inflate their tracheal sacs. This mechanism provides a significant energetic advantage that allows the larvae to migrate via passive buoyancy rather than more energy-costly swimming. The Chaoborus spp. larvae, in addition to potentially releasing sediment methane bubbles twice a day by entering and leaving the sediment, also transport porewater methane within their gas vesicles into the water column, resulting in a flux of 0.01-2 mol m-2 yr-1 depending on population density and water depth. Chaoborus spp. emerging annually as flies also result in 0.1-6 mol m-2 yr-1 of carbon export from the system. Finding the tipping point in lake eutrophication enabling this methane-powered migration mechanism is crucial for ultimately reconstructing the geographical expansion of Chaoborus spp., and the corresponding shifts in the lake's biogeochemistry, carbon cycling and food web structure.}, language = {en} } @article{TangBackhausRiemannetal.2019, author = {Tang, Kam W. and Backhaus, Liv and Riemann, Lasse and Koski, Marja and Grossart, Hans-Peter and Munk, Peter and Nielsen, Torkel Gissel}, title = {Copepod carcasses in the subtropical convergence zone of the Sargasso Sea}, series = {Journal of plankton research}, volume = {41}, journal = {Journal of plankton research}, number = {4}, publisher = {Oxford Univ. Press}, address = {Oxford}, issn = {0142-7873}, doi = {10.1093/plankt/fbz038}, pages = {549 -- 560}, year = {2019}, abstract = {The oligotrophic subtropical gyre covers a vast area of the Atlantic Ocean. Decades of time-series monitoring have generated detailed temporal information about zooplankton species and abundances at fixed locations within the gyre, but their live/dead status is often omitted, especially in the dynamic subtropical convergence zone (STCZ) where the water column stratification pattern can change considerably across the front as warm and cold water masses converge. We conducted a detailed survey in the North Atlantic STCZ and showed that over 85\% of the copepods were typically concentrated in the upper 200 m. Copepod carcasses were present in all samples and their proportional numerical abundances increased with depth, reaching up to 91\% at 300-400 m. Overall, 14-19\% of the copepods within the upper 200 m were carcasses. Shipboard experiments showed that during carcass decomposition, microbial respiration increased, and the bacterial community associated with the carcasses diverged from that in the ambient water. Combining field and experimental data, we estimated that decomposing copepod carcasses constitute a negligible oxygen sink in the STCZ, but sinking carcasses may represent an overlooked portion of the passive carbon sinking flux and should be incorporated in future studies of carbon flux in this area.}, language = {en} }