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Many lake water bodies not attained the goal of the European Water Framework Directive (WFD) to achieve a good ecological state by 2015. This is, among other things, because the assessment and improvement of hydromorphological conditions of lakeshores has been neglected as an important component ensuring the ecological integrity of lake ecosystems. In recent years, macroinvertebrates were emphasized to be useful indicators for the assessment of lakeshore hydromorphology. Hence, in Europe macroinvertebrate-based assessment methods were developed to evaluate the hydromorphological conditions of lakeshores.
In this thesis, some of the uncertainties and missing aspects of existing macroinvertebrate-based assessment methods were addressed. The results were obtained by sampling macroinvertebrates and macrophytes at natural shores and at shores modified by marinas and beaches in three depth zones between April and November 2011 in a large lowland lake (Lake Scharmützelsee, Germany).
Firstly, I clarified that upper littoral macroinvertebrates should be used for assessing the hydromorphology of lakeshores. It was shown that the effect of lakeshore modification on macroinvertebrate diversity and community composition was most pronounced in the upper littoral and decreased to the profundal zone. Secondly, I demonstrated that a single seasonal sampling is sufficient to capture the compositional differences of macroinvertebrate communities associated with human lakeshore modification. Seasonal effects on upper littoral macroinvertebrate diversity and composition were less important than shore type in comparison with the middle littoral and profundal zone. Thirdly, upper littoral macrophyte communities were also affected by lakeshore modification and at the same time the most important variable structuring macroinvertebrate communities. Hence, the effects of different shore types on macrophytes were transferred to macroinvertebrates, but artificial substrates were also able to partly substitute macrophyte habitats as it was shown for the studied marinas. Since lakeshore modification affected macrophytes slightly differently than macroinvertebrates, macrophytes should be considered as an additional component in lakeshore assessment. Finally, secondary production as proxy to determine the effect of lakeshore modification on the functioning of macroinvertebrates was estimated. Estimation of secondary production requires the determination of biomass. Biomass was indirectly determined by using length-mass regressions established for macroinvertebrates from temperate lakes of the central European lowland. The result showed that total secondary production and secondary production of native taxa in the upper littoral was substantially lower at the studied beaches compared to natural sites. In contrast, upper littoral secondary production at marinas did not differ to natural sites, but secondary production of non-native taxa was significantly higher at marinas. No effects of lakeshore modification on secondary production were found with increasing depth. Different scenarios based on upscaling of site-specific production to whole lake ecosystem level gave evidence that the observed local impacts of lakeshore modification can translate into alterations of the functioning of macroinvertebrates at whole lake ecosystem level. In addition, it was emphasized that secondary production as a functional measure is more sensitive in detecting hydromorphological alterations than the structural measures diversity or biomass. Secondary production should therefore be included in existing lakeshore assessment methods. In order to obtain a comprehensive overview about the changes in the functioning of macroinvertebrates following lakeshore modification, it is recommended to consider not only total secondary production but also secondary production of functional groups. With these results, this thesis contributes to the mechanistic understanding of the effect of lakeshore modification on the functioning of macroinvertebrates and the consequences for the functioning of the whole lake ecosystem. The newly generated knowledge helps to optimize the development of successful lakeshore assessment tools and identification of management measures.
The main goal of this dissertation was to explore the interactions between the hydrogeomorphology of the streambed in sandy lowland low-order streams and the microbial community inhabiting it. In particular, (i) the influence of the vertical water exchange across the streambed and (ii) of the sediment transport on the function and structure of the streambed microbial community, (iii) and the potential of the microbial community to influence these physical factors were explored. The influences were studied with a model system approach (micro- and mesocosms).
Firstly, I examined the significance of vertical water exchange across the streambed for the microbial community. I determined the differences in the microbial community structure and function associated with sediments of differing grain sizes. The grain sizes differed in surface-to-volume ratio and hydraulic conductivity. The results revealed vertical water exchange as the major factor for the structure and function of the microbial community.
Secondly, I studied the ability of the microbial community to influence the vertical water exchange across two sandy streambeds: leveled and rippled. My results showed that the microbial community can reduce and even block the vertical water exchange by reducing pore space with gas bubbles formed due to high primary production.
Thirdly, I determined the effect of short-term sediment transport events on the function of the microbial community and on the influence of the microbial community on vertical water exchange. The results show that the mechanical stress associated with short-term sediment transport events does not influence the microbial community function. However, a single short-term sediment transport event increased vertical water exchange by (i) releasing the gas bubbles produced by the microbial community and (ii) creating irregularities in the flume bed.
Lastly, I ascertained the potential of benthic algal mats to transport sediment by means of buoyancy-mediated detachment from the bed. The results revealed the detachment of algal mats as a novel mechanism of sediment transport during low-flow periods. Overall, the interactions studied show that in sandy streambeds (i) the pattern of vertical water exchange is the primary physical template for the microbial community, and (ii) the activity of the microbial community and sediment transport are stochastic sources of spatiotemporal heterogeneity in vertical water exchange. These results contribute to the understanding and prediction of stream ecosystem functions in sandy streams, which is of special significance in light of the increase in fine sediment load in streams worldwide.
Nostocalean cyanobacteria differentiate heterocysts to fix dissolved nitrogen (N2) and dormant cells (akinetes) to survive harsh environmental conditions. The distribution and proliferation of Nostocales and their hibernation strategies in deep stratified lakes of different trophic states and climate zones were investigated. Studies were carried out in the meso-eutrophic Lake Kinneret (Israel) and in the mesotrophic Lake Scharmützelsee and the oligo-mesotrophic Lake Stechlinsee (Germany).
The spatio-temporal distribution of akinetes in sediments was analysed. The abundance in the sediment surface increased with water depth and differed due to basin morphometry. Light and temperature distribution revealed shallow areas of sedimentary akinete pool as a potential inoculum to contribute to the formation of a pelagic population. Hence, a small viable akinete pool was deposited in shallow sediments.
A CARD-FISH protocol was implemented to identify and quantify the akinetes in sediments. Tested Nostocales strains and akinetes in sediment samples were successfully labelled with 16S rRNA targeted probes and microscopically detected by their fluorescence signal. Akinete enumeration in field samples confirmed the suitability of the CARD-FISH approach.
Species- or genus-specific differences in the overwintering strategies of Nostocales were observed. Akinetes were the only overwintering form of Anabaena spp. On the other hand, Aphanizomenon flos-aquae overwintered as vegetative filaments in the pelagial of Lake Stechlinsee. In the other lakes studied, Aphanizomenon spp. performed an intermediate life cycle, with akinetes and a small pelagic population in winter. This suggested that Aphanizomenon spp. had another overwintering strategy contrary to the exclusively dormant strategy of Anabaena spp.
Putative strains of cylindrospermopsin (CYN)-producing Aphanizomenon ovalisporum were isolated from Lake Kinneret. Their morphologic and phylogenetic diversity, as well as the presence of CYN-encoding gene cluster and CYN production, were analysed. Four of the six strains were assigned to Anabaena bergii. Although all the isolated strains possess at least fragments of CYN gene cluster, no CYN production was detected. Non-CYN-producing Aphanizomenon ovalisporum strains and the presence of non-toxic Anabaena bergii in Lake Kinneret have not been reported before. This suggested that toxic and non-toxic Nostocales strains can coexist in lakes.
Ecological research on acidic lakes has historically focused on soft-waters, whereas biota in natural and man-made acidic hard-water lakes remains poorly studied. My thesis deals with the effects of multiple abiotic stressors on species and genetic biodiversity of littoral Cladocera in two types of acidic waters in Germany: hard-water mining and soft-water bog lakes. I found that these two types of acidic lakes supported highly distinct cladoceran communities, both with respect to their species diversity and composition, what led to the rejection of my null hypothesis that pH was the only abiotic factor, determining species diversity patterns. Further analyses revealed that, apart from pH, TDS-related parameters and trophy significantly affected species composition of littoral Cladocera in the studied lakes. Particularly the role of TDS and its constituents for colonization of acidic lakes was virtually neglected by previous studies, based on soft-water lakes’ data sets. My study on the genetic diversity of a generalist cladoceran Chydorus sphaericus provided evidence for genetic erosion, i.e. negative effects of abiotic stress in the acidic hard-water mining lakes, but not in the acidic soft-water bog lakes. I conclude that the observed genetic erosion in acidic mining lakes may have negative long-term consequences for population persistence even in broadly tolerant species. In general, biodiversity at both species and genetic levels was much more impaired in the acidic hard-water mining lakes, compared to the naturally acidic soft-water bog lakes. This was possibly due to both ecological (higher abiotic stress) and evolutionary (insufficient time for long-term adaptations to evolve) mechanisms. My results suggest that colonization of acidic hard- and soft-water lakes in each case should require specific physiological adaptations at both species and population levels. The practical relevance of my studies is that they provide the scientific basis for monitoring of acidic hard-water mining lakes based on littoral Cladocera. Especially genetic diversity proved a valuable indicator of water quality in acidic mining lakes, as it provided unique information on population-level responses, being complementary to species diversity assessment. Overall, my results demonstrate that inclusion of acidic hard-water lakes into the research framework as well as comparisons among communities in chemically diverse acidic lakes can result in important new insights, concerning the community organization and adaptations in acidic environments.
Phytoplankton succession and diversity-productivity relation in German and Brazilian shallow lakes
(2012)
Phytoplankton succession linked to productivity-diversity relationship at steady-state/non steady state conditions, mixing events as predominant disturbance in sense of Connell (1978) and influence of conditional differentiation (temperature) on phytoplankton diversity were investigated. Water samples were taken two times a week in 4 German shallow lakes in Brandenbug region (Melangsee, Petersdorfer See, Wolziger See, and Langer See) from 2007 to 2010 and in 2009/2010 during cold and rainy season in a Brazilian coastal lagoon Peri. A comparison between literature data from 4 Brazilian lakes, in order to investigate the influence of temperature on diversity and dynamic of phytoplankton species was also done. The cyanobacterial species dominated all studied lakes and the mixing had a positive effect on the supremacy of this group in tropical lakes. The high biovolume of cyanobacteria contributed to the achievement of steady state conditions in sense of Sommer et al. (1993) during consecutive periods in Petersdorfer See and in Peri lagoon practically during all phases. The monodominance of Cylindrospermopsis raciborskii was observed in warm water, under no stratification regime. Oscillatoriales of group S1 (Reynolds 2002), represented by Limnothrix redekei and Pseudanabaena limnetica, typical in turbid mixed layers and with high nitrogen affinity, were typical in Petersdorfer See. In Melangsee, steady state was observed during colder periods under dominance of Bacillariophyceae and Chrysophyceae. Lower diversity coincided with steady state conditions and higher values of resource use efficiency (RUE), while higher diversity occurred mainly at transitional state during the spring in temperate lakes, supporting the non-equilibrium concept (Hutchinson 1961) at lower RUE values. Higher biovolume occurred mainly during the late summer in temperate lakes under dominance of cyanobacteria, and/or at steady state. Concerning the temporal variation of phytoplankton, the highest peak of species richness preceded the resource use efficiency, suggesting the productivity as consequence of diversity. The richness was much higher in temperate shallow lakes, which were more similar to each other in terms of seasonality, species number and biovolume. A positive correlation between richness and biovolume of temperate lakes was found and the opposite was observed in tropical lakes. The biovolume and richness in tropical lakes were more variable and in temperate lakes presented higher richness at intermediate level of biovolume (annual mean from 10 to 18 mm3/l). Low level of biodiversity allows an ecosystem to function under constant conditions, but a greater biodiversity occurred in fluctuating environments.
Processes of microbial carbon transformation and accumulation during initial stream succession were investigated. Studies were carried out in the experimental watershed Chicken Creek, constructed to investigate ecosystem succession, and additionally in experimental flumes simulating sand-bed streams. In a one year investigation, microbial respiration in soils and sediments along the hydrologic flow path of three stream corridors in the Chicken Creek watershed was measured. Contrary to expectation, respiration rates of rewetted soil and sediment from dry stream channels were similar to rates measured with sediments collected in the perennial channel sections. This suggested that permanent water availability was not a main factor determining metabolic potential in this early successional watershed. In an outdoor flume experiment it was determined whether shallow (1cm) and deep (4cm) sediment disturbances in small sand-bed streams have similar effects on whole-stream metabolism, and whether autotrophic and heterotrophic processes and organisms respond in similar ways. Results suggested that disturbing sediments during early successional stages had no effect on whole-stream metabolism, whereas in advanced stages, deep but not shallow sediment disturbance could lead to a transitory shift towards heterotrophy. Changes in riparian and in-stream vegetation during stream succession come along with different amounts and types of organic matter input in stream ecosystems. It was tested to determine if increasing quality and quantity of litter input changes whole-stream metabolism and activity and structure of microbial communities associated with sediments and leaves. Whole-stream metabolism was found to be similar in all treatments because sediments and leaves were constrained by oxygen and nutrient availability. There seemed to be compensation between the effect of algae fueling microbial activity in open-land treatments and microbial use of allochthonous carbon sources in the litter treatments. Fungal and bacterial activity associated with leaves was unaffected by the background litter standing stock, but the structure of communities was affected. However, leaf quality had a clear effect on microbial activity and community structure with higher activity on tree compared to grass leaves.
Viele biotische und abiotische Faktoren, wie die Nahrungsqualität und diverse chemische Stressoren, bestimmen die Lebensparameter von Cladoceren. Diese Arbeit konzentriert sich auf die Effekte der unterschiedlichen Futteralgen (Pseudokirchneriella subcapitata, Monoraphidium minutum und Desmodesmus armatus) auf die Lebensparameter von vier Cladoceren-Arten (Daphnia magna, D. galeata, Moina macrocopa und M. micrura) in An- und Abwesenheit von chemischem Stress, ausgelöst durch den Huminstoff HuminFeed® (HF). Um die Effekte von HF auf D. magna zu ermitteln, wurden Lebenszyklusexperimente durchgeführt. In umweltrelevanten Konzentrationen wirkte HF lebensverlängernd, jedoch zeigten sich Verminderung in der Reproduktion. Die Ergebnisse wurden im Licht der Disposable Soma Theorie diskutiert. Es konnte keine Steuerung der Energieallokation durch HF und durch die Futteralgenqualität festgestellt werden. In einem Multigenerationsexperiment wurde der Einfluss von HF auf D. magna über drei Generationen hinweg betrachtet. Dabei bewirkte HF stets eine Verminderung der Reproduktion und eine Erhöhung der Lebensspanne. Die Ergebnisse des ersten Filialgenerationsversuchs F1 haben gezeigt, dass ein signifikanter Unterschied in der Fruchtbarkeit von Versuchstieren besteht, die in derselben Umwelt aufgewachsen sind (ohne HF), aber deren Mütter (P) unterschiedlichen Behandlungen ausgesetzt waren. Die Ergebnisse des zweiten Filialgenerationsversuchs F2 haben aber aufgezeigt, dass die Umwelt, in der die Parentalgeneration aufgewachsen ist, die Fruchtbarkeit der F2 Generation nicht nachhaltig beeinflusst. Es handelt sich vermutlich um Transgenerationseffekte (epigenetische Programmierung). In einem Langzeitexperiment wurden die Auswirkungen vom HF auf die Reproduktion und die Lebensspanne von D. galeata untersucht. Es zeigte sich, dass HF erneut eine Steigerung der Lebensdauer und eine Verringerung der Reproduktion bewirkt. Die Ergebnisse wurden im Licht der Disposable Soma Theorie diskutiert. Des Weiteren wurde der Einfluss von HF auf die Lebensparameter von M. macrocopa untersucht. Durch die Zugabe von HF bei Fütterung mit D. armatus wurde teilweise eine Verkürzung der Lebensspanne und eine Erhöhung der Nachkommenzahl hervorgerufen. Bei Fütterung mit P. subcapitata zeigte sich, dass HF keine signifikanten Effekte erzielte. Bei Fütterung mit M. minutum konnte in exponierten Ansätzen eine Verlängerung der Lebensspanne und Erhöhung der durchschnittlichen kumulativen Nachkommenzahl festgestellt werden. Hormonähnliche Effekte der Huminstoffe wurden in Betracht gezogen. Die unterschiedliche Antwort von M. macrocopa auf Huminstoffe bei verschiedenen Futteralgen kann darauf hindeuten, dass die Energieallokation in M. macrocopa bei HF-Exposition durch die Futteralgenqualität gesteuert wird. Ebenfalls wurde bei M. micrura festgestellt, dass die Energieallokation unter dem chemischen Stressor HF durch die Futteralgenqualität gesteuert wurde. So wurde gezeigt, dass HF einen (teilweise signifikant) negativen Einfluss sowohl auf die Lebensspanne als auch auf die Fruchtbarkeit hat, wenn die Versuchstiere mit P. subcapitata gefüttert wurden. Werden M. micrura hingegen mit M. minutum gefüttert, wirkte HF teilweise lebensverkürzend und ein Trend zur Erhöhung der Nachkommenzahl war sichtbar. Diese Arbeit gibt Hinweise darauf, dass zwischen den Zooplanktonarten Unterschiede hinsichtlich der Antwort auf chemischen Stress und des Bedarfs an einzelnen Futterqualitätsfaktoren bestehen.
Rapidly developing countries are facing severe environmental problems at an extent that industrialized countries did not have to solve in comparable periods. Freshwaters belong to the ecosystems with the most severe impacts despite they are one of the most vital resources. The Lerma River is one example of a heavily degraded freshwater ecosystem in the rapidly developing country of Mexico. A number of NGOs and governmental institutions have made great efforts to elaborate strategies for the restoration and rehabilitation of the river system during the last years. Scientists from different disciplines and institutions met for a workshop at ININ in February 2009 to discuss the actual degradation of the upper Lerma River and argue about solutions for its sustainable restoration. This also included the presentation and discussion of new technologies that are currently in an experimental stage, but have the potential to solve different river pollution problems in the near future.
Die EU-Wasserrahmenrichtlinie (EU-WRRL 2000) beschäftigt Wasserforscher und Behörden in einer erfreulichen Weise: Erstmalig werden ökologische und gewässertypbezogene Bewertungsverfahren im nationalen Rahmen entwickelt und auf europäischer Ebene angepasst. Seit 999 sind die Autoren dieses Gewässerreportes in Projekten zur Erstellung eines Bewertungsverfahrens für das Phytoplankton in natürlichen Seen eingebunden. Die Verfahrensentwicklung begann mit einer umfassenden Literaturstudie zu Klassifizierungsverfahren von Seen und Flüssen anhand des Phytoplanktons. Diese internationale Literaturrecherche ergab, dass zwar gute Ansätze aus anderen Ökoregionen Europas vorhanden sind, diese sich jedoch nicht einfach auf die naturräumlichen Verhältnisse in Deutschland übertragen lassen oder nicht den vielfältigen Anforderungen der EU-WRRL genügen. Folglich beauftragte die Länderarbeitsgemeinschaft Wasser die Autoren, ein WRRL konformes Bewertungssystem zu entwickeln, das aus vorhandenen Überwachungsdaten gewonnen werden sollte. Allerdings machte die Zusammenführung dieser heterogenen Datenbestände deutlich, wie dringend eine Vereinheitlichung der Probenahmen und der mikroskopischen Erfassung erfolgen muss, da die einzelnen Bundesländer hierfür bisher sehr unterschiedliche Strategien praktizierten. Im Abschlussbericht wurden im Ergebnis die ersten Vorschläge für ein multimetrisches Bewertungssystem mittels Phytoplankton sowie zu seiner einheitlichen methodischen Erfassung beschrieben. In dem darauf folgenden 2-jährigen Praxistest wurden diese Empfehlungen einer bundesweiten Prüfung insbesondere auf Praktikabilität und Plausibilität unterzogen. Vielfältige Modifikationen des ersten Bewertungsentwurfes wurden notwendig infolge der Abstimmungsprozesse mit den Praxispartnern in den Bundesländern, mit den Anwendern in speziell zu diesem Thema ausgerichteten Workshops, aber auch innerhalb der EU („Interkalibrierung“). Ökologische Bewertung nach WRRL heißt, die Reaktion des Phytoplanktons auf eine anthropogene Belastung auf Ebene seiner Biomasse und seiner taxonomischen Zusammensetzung (Arten / Algenklassen) in Form von biologischen Kenngrößen (Metrics) zu berücksichtigen. Für die vorliegende Bewertungsherleitung für die Biokomponente Phytoplankton wird der Begriff „Belastung“ eingrenzend als eine anthropogen beschleunigte Eutrophierung und als Abweichung von einem Referenzzustand verstanden, wobei dieser Zustand nicht mehr allein biomasseorientiert, sondern auch durch die völlig neu entwickelte Liste an Indikatorarten (PTSI) definiert wird. Die Bewertung des Phytoplanktons erfolgt demnach mit einem multimetrischen Ansatz im Phyto-See-Index (PSI), der durch die Einbeziehung des Arteninventars sowie durch die neu definierten trophischen Referenzzustände in den meisten Fällen strenger ist als der in Deutschland bislang zur trophischen Einstufung von Seen verwendete LAWA-Index (1999). Um eine routinemäßige Bewertung einschließlich der Indikatorarten zu ermöglichen, wurde eine operative Taxaliste des Phytoplanktons mit Hinweisen für die Bestimmungstiefe sowie das Auswertungsprogramm „PhytoSee“ zur automatisierten Berechnung entwickelt, die zusammen eine schnelle Auswertung der standardisierten Messdaten erlauben.
In recent years Cylindrospermopsis raciborskii (Woloszynska) Seenayya et Subba Raju a potentially toxic freshwater cyanobacterium of the order Nostocales initially classified as a tropical species, has invaded the temperate region and has established in lakes in northern Germany. The formation of dormant cells (akinetes) is an important part of the life cycle which enables Nostocales species to survive harsh conditions like the winter in the temperate zone. In this thesis, my aim is to throw more light on the life cycle of the invasive cyanobacterium Cylindrospermopsis raciborskii and also to compare it to native Nostocales species of the genus Aphanizomenon using derived field and laboratory data. This would help in the understanding of the invasion of C. raciborskii and spread and how it can be contained. Abiotic factors influencing the development and growth of photoautotrophic planktonic organisms in nature are examined and their variation with the years is discussed. A comprehensive look at C. raciborskii’s akinetes in the sediment and the filaments and akinetes in the water column is done. The timing and temperature condition of akinete germination, recruitment, formation in the pelagic and importation into the sediment is determined. An assessment of the role of abiotic factors on the dynamics and abundance of filaments is done with a statistical model used to isolate which abiotic factors are most influential on the population growth of C. raciborskii. Akinetes distribution on the bed of the lake and the effect of loss processes on their abundance are looked into. Cylindrospermopsis is compared to native species Aphanizomenon spp in terms of the seasonal dynamics and the germination condition of their akinetes. The main findings in this thesis were that the seasonal dynamics of C. raciborskii had a similar trend but different timing of each event in the four years of the study. The variabilities were due to the different timing of the abiotic conditions. But in all temperature was identified as the most important factor influencing the timing of most events with germination of akinetes occurring at 13°C which is lower than so far reported in literature Subsequent increment in temperature to at least 23°C was found to be essential for the development of high populations of C. raciborskii in the water column. With a drop of temperature in autumn the filaments are stimulated in to producing akinetes which settle out into the sediment and remain there until the next spring when favourable conditions force them to germinate. They might also be consumed before germination by non identified grazer or losses due to microbial infections are discussed. Overall, the native Nostocales species of the genus Aphanizomenon have a selective advantage over the invaded species C. raciborskii under low temperatures and temperature fluctuations in spring as observed from the field and laboratory studies.