TY - JOUR A1 - Brothers, Soren M. A1 - Hilt, Sabine A1 - Meyer, Stephanie A1 - Köhler, Jan T1 - Plant community structure determines primary productivity in shallow, eutrophic lakes JF - Freshwater biology N2 - Regime shifts are commonly associated with the loss of submerged macrophytes in shallow lakes; yet, the effects of this on whole-lake primary productivity remain poorly understood. This study compares the annual gross primary production (GPP) of two shallow, eutrophic lakes with different plant community structures but similar nutrient concentrations. Daily GPP rates were substantially higher in the lake containing submerged macrophytes (58623gCm(-2)year(-1)) than in the lake featuring only phytoplankton and periphyton (40823gCm(-2)year(-1); P<0.0001). Comparing lake-centre diel oxygen curves to compartmental estimates of GPP confirmed that single-site oxygen curves may provide unreliable estimates of whole-lake GPP. The discrepancy between approaches was greatest in the macrophyte-dominated lake during the summer, with a high proportion of GPP occurring in the littoral zone. Our empirical results were used to construct a simple conceptual model relating GPP to nutrient availability for these alternative ecological regimes. This model predicted that lakes featuring submerged macrophytes may commonly support higher rates of GPP than phytoplankton-dominated lakes, but only within a moderate range of nutrient availability (total phosphorus ranging from 30 to 100gL(-1)) and with mean lake depths shallower than 3 or 4m. We conclude that shallow lakes with a submerged macrophyte-epiphyton complex may frequently support a higher annual primary production than comparable lakes that contain only phytoplankton and periphyton. We thus suggest that a regime shift involving the loss of submerged macrophytes may decrease the primary productivity of many lakes, with potential consequences for the entire food webs of these ecosystems. KW - macrophytes KW - oxygen curves KW - periphyton KW - regime shift KW - trophic status Y1 - 2013 U6 - https://doi.org/10.1111/fwb.12207 SN - 0046-5070 VL - 58 IS - 11 SP - 2264 EP - 2276 PB - Wiley-Blackwell CY - Hoboken ER - TY - THES A1 - Brothers, Soren M. T1 - Carbon gains, losses, and feedbacks in shallow, eutrophic lakes of phytoplankton and macrophyte dominance T1 - Kohlenstoff-Gewinne, Verluste und Rückkopplungen in flachen, nährstoffreichen Seen mit Dominanz von Phytoplankton und Makrophyten N2 - Lakes are increasingly being recognized as an important component of the global carbon cycle, yet anthropogenic activities that alter their community structure may change the way they transport and process carbon. This research focuses on the relationship between carbon cycling and community structure of primary producers in small, shallow lakes, which are the most abundant lake type in the world, and furthermore subject to intense terrestrial-aquatic coupling due to their high perimeter:area ratio. Shifts between macrophyte and phytoplankton dominance are widespread and common in shallow lakes, with potentially large consequences to regional carbon cycling. I thus compared a lake with clear-water conditions and a submerged macrophyte community to a turbid, phytoplankton-dominated lake, describing differences in the availability, processing, and export of organic and inorganic carbon. I furthermore examined the effects of increasing terrestrial carbon inputs on internal carbon cycling processes. Pelagic diel (24-hour) oxygen curves and independent fluorometric approaches of individual primary producers together indicated that the presence of a submerged macrophyte community facilitated higher annual rates of gross primary production than could be supported in a phytoplankton-dominated lake at similar nutrient concentrations. A simple model constructed from the empirical data suggested that this difference between regime types could be common in moderately eutrophic lakes with mean depths under three to four meters, where benthic primary production is a potentially major contributor to the whole-lake primary production. It thus appears likely that a regime shift from macrophyte to phytoplankton dominance in shallow lakes would typically decrease the quantity of autochthonous organic carbon available to lake food webs. Sediment core analyses indicated that a regime shift from macrophyte to phytoplankton dominance was associated with a four-fold increase in carbon burial rates, signalling a major change in lake carbon cycling dynamics. Carbon mass balances suggested that increasing carbon burial rates were not due to an increase in primary production or allochthonous loading, but instead were due to a higher carbon burial efficiency (carbon burial / carbon deposition). This, in turn, was associated with diminished benthic mineralization rates and an increase in calcite precipitation, together resulting in lower surface carbon dioxide emissions. Finally, a period of unusually high precipitation led to rising water levels, resulting in a feedback loop linking increasing concentrations of dissolved organic carbon (DOC) to severely anoxic conditions in the phytoplankton-dominated system. High water levels and DOC concentrations diminished benthic primary production (via shading) and boosted pelagic respiration rates, diminishing the hypolimnetic oxygen supply. The resulting anoxia created redox conditions which led to a major release of nutrients, DOC, and iron from the sediments. This further transformed the lake metabolism, providing a prolonged summertime anoxia below a water depth of 1 m, and leading to the near-complete loss of fish and macroinvertebrates. Pelagic pH levels also decreased significantly, increasing surface carbon dioxide emissions by an order of magnitude compared to previous years. Altogether, this thesis adds an important body of knowledge to our understanding of the significance of the benthic zone to carbon cycling in shallow lakes. The contribution of the benthic zone towards whole-lake primary production was quantified, and was identified as an important but vulnerable site for primary production. Benthic mineralization rates were furthermore found to influence carbon burial and surface emission rates, and benthic primary productivity played an important role in determining hypolimnetic oxygen availability, thus controlling the internal sediment loading of nutrients and carbon. This thesis also uniquely demonstrates that the ecological community structure (i.e. stable regime) of a eutrophic, shallow lake can significantly influence carbon availability and processing. By changing carbon cycling pathways, regime shifts in shallow lakes may significantly alter the role of these ecosystems with respect to the global carbon cycle. N2 - Seen werden zunehmend als wichtige Komponente im globalen Kohlenstoffkreislauf anerkannt. Natürliche Veränderungen und anthropogene Aktivitäten beeinflussen die Struktur der Artengemeinschaft von Seen, was Auswirkungen auf den Transport und Umsatz von Kohlenstoff hat. Diese Arbeit konzentriert sich auf die Beziehung zwischen Kohlenstoffkreislauf und der Gemeinschaftsstruktur der Primärproduzenten in kleinen Flachseen. Diese sind der weltweit häufigste Seentyp und weisen durch ihren im Vergleich zur Fläche großen Umfang eine intensive aquatisch-terrestrische Kopplung auf. In Flachseen treten oft Regimewechsel zwischen Makrophyten- und Phytoplankton-Dominanz auf. Diese können potenziell große Konsequenzen für den regionalen Kohlenstoffkreislauf haben. In dieser Dissertation vergleiche ich einen Klarwassersee mit submersen Makrophyten und einen trüben, Phytoplankton-dominierten See hinsichtlich Verfügbarkeit, Umsatz und Export von organischem und anorganischem Kohlenstoff. Des Weiteren habe ich den Effekt der erhöhten Zufuhr von terrestrischem Kohlenstoff auf den internen Kohlenstoffumsatz untersucht. Sowohl die Tagesgänge der pelagischen Sauerstoff-Konzentrationen als auch Fluoreszenz-basierte Messungen der Primärproduktion bewiesen, dass die Präsenz von submersen Makrophyten eine höhere jährliche Brutto-Primärproduktion im Vergleich zu einem Phytoplankton-dominierten See mit ähnlichen Nährstoffkonzentrationen ermöglicht. Ein einfaches, auf den empirischen Daten basierendes Model zeigt, dass diese Unterschiede in der Brutto-Primärproduktion typisch sind für moderat eutrophe Seen mit einer mittleren Tiefe von unter 3 bis vier Metern. In diesen Seen leistet die benthische Primärproduktion den Hauptbeitrag zur Primärproduktion des ganzen Sees. Daraus wird ersichtlich, dass Regimewechsel von Makrophyten- zur Phytoplankton-Dominanz in Flachseen die Verfügbarkeit von autochthonem organischem Kohlenstoff für das Nahrungsnetz reduzieren. Paläolimnologische Analysen in Sedimentkernen beider Seen wiesen darauf hin, dass der Verlust der Makrophyten mit einer vierfachen Zunahme der Kohlenstoff-Speicherraten einhergeht, und somit zu einer großen Veränderung der Dynamik des Kohlenstoffkreislaufs im See führt. Unsere Kohlenstoff-Massenbilanzen zeigen, dass die Erhöhung der Kohlenstoff-Speicherung im Sediment nicht durch die Erhöhung der Primärproduktion oder durch externe Quellen, sondern durch erhöhte der Effizienz der Speicherung begründet war. Dies geht mit einer reduzierten benthischen Mineralisierungsrate und einer erhöhten Calcitfällung einher und führt zu reduzierten Kohlendioxid-Emissionen. Eine Periode ungewöhnlich hoher Niederschläge mit erhöhten Wasserständen führte im Phytoplankton-dominierten See zu zu einem starken Anstieg der Konzentrationen an gelöstem organischem Kohlenstoff (DOC) und zu anoxischen Bedingungen. Es wurde postuliert, dass zwischen diesen Prozessen eine positive Rückkopplung besteht. Die hohen Wasserstände und DOC-Konzentrationen reduzierten die Lichtversorgung und damit die Primärproduktion im Benthal und erhöhten die pelagischen Respirationsraten. Dadurch verringerte sich die Sauerstoffverfügbarkeit im Hypolimnion. Die dadurch erzeugten Redox-Verhältnisse führten zu einer Freisetzung großer Mengen an Nährstoffen, DOC und Eisen aus dem Sediment. Die während des gesamten Sommers andauernden anoxischen Verhältnisse in Wassertiefen unter 1 m führten zu einem fast vollständigen Verlust von Fischen und Makroinvertebraten. Zusätzlich wurde der pH-Wert im Pelagial signifikant erniedrigt und die Kohlenstoffdioxid-Emissionen im Vergleich zu früheren Jahren verzehnfacht. Insgesamt trägt diese Dissertation wesentliche Aspekte zum besseren Verständnis der Bedeutung des Benthals für den Kohlenstoffkreislauf in Flachseen bei. Der Anteil der benthischen Zone an der Primärproduktion in kleinen Flachseen wurde in Relation zur Gesamtproduktion des Systems quantifiziert. Letztlich zeigt diese Arbeit, dass die Gemeinschaftsstruktur der Primärproduzenten eines eutrophen Flachsees die Verfügbarkeit und den Umsatz von Kohlenstoff signifikant beeinflusst. Regimewechsel in Flachseen können durch Änderungen im internen Kohlenstoffkreislauf deren Rolle im globalen Kohlenstoffkreislauf verändern. KW - Carbon Cycling KW - Primärproduktion KW - Anoxie KW - Brownification KW - Carbon cycling KW - primary production KW - anoxia KW - brownification Y1 - 2013 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus-68200 ER - TY - JOUR A1 - Brothers, Soren M. A1 - Hilt, Sabine A1 - Attermeyer, Katrin A1 - Grossart, Hans-Peter A1 - Kosten, Sarian A1 - Lischke, Betty A1 - Mehner, Thomas A1 - Meyer, Nils A1 - Scharnweber, Inga Kristin A1 - Köhler, Jan T1 - A regime shift from macrophyte to phytoplankton dominance enhances carbon burial in a shallow, eutrophic lake JF - Ecosphere : the magazine of the International Ecology University N2 - Ecological regime shifts and carbon cycling in aquatic systems have both been subject to increasing attention in recent years, yet the direct connection between these topics has remained poorly understood. A four-fold increase in sedimentation rates was observed within the past 50 years in a shallow eutrophic lake with no surface in-or outflows. This change coincided with an ecological regime shift involving the complete loss of submerged macrophytes, leading to a more turbid, phytoplankton-dominated state. To determine whether the increase in carbon (C) burial resulted from a comprehensive transformation of C cycling pathways in parallel to this regime shift, we compared the annual C balances (mass balance and ecosystem budget) of this turbid lake to a similar nearby lake with submerged macrophytes, a higher transparency, and similar nutrient concentrations. C balances indicated that roughly 80% of the C input was permanently buried in the turbid lake sediments, compared to 40% in the clearer macrophyte-dominated lake. This was due to a higher measured C burial efficiency in the turbid lake, which could be explained by lower benthic C mineralization rates. These lower mineralization rates were associated with a decrease in benthic oxygen availability coinciding with the loss of submerged macrophytes. In contrast to previous assumptions that a regime shift to phytoplankton dominance decreases lake heterotrophy by boosting whole-lake primary production, our results suggest that an equivalent net metabolic shift may also result from lower C mineralization rates in a shallow, turbid lake. The widespread occurrence of such shifts may thus fundamentally alter the role of shallow lakes in the global C cycle, away from channeling terrestrial C to the atmosphere and towards burying an increasing amount of C. KW - calcite precipitation KW - CO2 emissions KW - global carbon cycle KW - metabolism KW - regime shift KW - sedimentation KW - submerged macrophytes KW - temperate zone KW - trophic status Y1 - 2013 U6 - https://doi.org/10.1890/ES13-00247.1 SN - 2150-8925 VL - 4 IS - 11 PB - Wiley CY - Washington ER -