Rozprawy doktorskie na temat „Algal Grazers”

Phytoplankton succession and abundance in estuaries is known to be influenced by the relative strengths of various seasonally changing physical and chemical factors. Previous studies of Swan River Estuary phytoplankton biomass and composition have identified salinity, temperature, rainfall and nutrients as the most important controlling factors. These conclusions are generally based on analysis of data from river length transects and depth integrated day-time sampling. They describe influences ,affecting whole system phytoplankton abundance and succession. Many of the typical seasonal bloom that develop are ephemeral and only extend over relatively small areas. The focus of this study is a single site, Ron Courtney Island, considered typical of the upper estuary region. This region of the estuary was chosen as representative of the section of river most influenced by allochthonous nutrient input. It has been the region of most frequent and intense algal blooms over the past decade. The factors, physical, biological or physiological, that have the greatest influence on controlling phytoplankton biomass under various ambient conditions for this system are determined. While previous studies have recognised the importance of nitrogen to phytoplankton growth in the Swan River Estuary, they have focused on NO;, with only anecdotal reference to the importance of the alternative nitrogen source, NH4+. This is the first study to explore the influence of different nitrogen source fluxes on phytoplankton biomass in the upper Swan River Estuary. The roles of physiological adaptation to, and preferences for, 'new' (NO,), recycled (NH4+) and organic (urea) nitrogen sources in relation to ambient nutrient levels are explored.Specific uptake rates (v), normalised to chlorophyll a, for NO;, NH4+ and urea were 0.2 ± 0.04 - 1831.1 ± 779.19, 0.5 ± 0.26 - 1731.6 ± 346.67 and 3.0 ± 0.60 - 2241.2 ± 252.56 ng N μg Chla-1 respectively. Urea concentration (14.8 - 117.7 μg urea-N 1-1) remained relatively constant over the 12 month study period. Measured ambient specific uptake rates for urea represent between 27.5% and 40.4% of total N uptake over the annual period February 1998 -January 1999. Seasonal nitrate uptake over the same period constituted only 11.3% (±10.77%, n=12) to 24.4% (± 13.02%, n=12) with the highest percentage during winter, when nitrate levels are elevated. It is suggested that urea provides a nutrient intermediary over the spring - summer period during transition from autotrophic to heterotrophic dominated communities. Grazing ,and nitrogen recycling are intricately connected by simultaneously providing top-down biomass control and bottom-up nutrient supply. Zooplankton (> 44 μm) grazing has been shown to reduce up to 40% of phytoplankton standing stock at times. Microheterotrophs (<300 pm) can reduce phytoplankton biomass production by up to 100% (potential production grazed, 11.1% day' - 99.6 % day-1) over an annual cycle. This correlated to mean seasonal day-time grazing loss of 80.47 ± 3.5 ngN μg Chla-1 in surface waters and 20.17 ± 9.7 ngN μg Chla-1 at depth (4.5m). Night time grazing for surface and bottom depths resulted in similar nitrogen loss rates (13.03 ± 4.84 ngN μg Chla-1).Uptake rates for nitrate (r2 0.501) and urea (r2 0.512), doing with temperature (r2 0.605) were shown to have the greatest influence on phytoplankton distribution over depth and time. This research emphasises the need for more detailed investigations into the physiology of nutrient uptake and the effects of nutrient fluxes on phytoplankton biomass and distribution. Further research into the roles of organic nitrogen and pico and nanoplankton in this system is recommended.

Phytoplankton succession and abundance in estuaries is known to be influenced by the relative strengths of various seasonally changing physical and chemical factors. Previous studies of Swan River Estuary phytoplankton biomass and composition have identified salinity, temperature, rainfall and nutrients as the most important controlling factors. These conclusions are generally based on analysis of data from river length transects and depth integrated day-time sampling. They describe influences ,affecting whole system phytoplankton abundance and succession. Many of the typical seasonal bloom that develop are ephemeral and only extend over relatively small areas. The focus of this study is a single site, Ron Courtney Island, considered typical of the upper estuary region. This region of the estuary was chosen as representative of the section of river most influenced by allochthonous nutrient input. It has been the region of most frequent and intense algal blooms over the past decade. The factors, physical, biological or physiological, that have the greatest influence on controlling phytoplankton biomass under various ambient conditions for this system are determined. While previous studies have recognised the importance of nitrogen to phytoplankton growth in the Swan River Estuary, they have focused on NO;, with only anecdotal reference to the importance of the alternative nitrogen source, NH4+. This is the first study to explore the influence of different nitrogen source fluxes on phytoplankton biomass in the upper Swan River Estuary. The roles of physiological adaptation to, and preferences for, 'new' (NO,), recycled (NH4+) and organic (urea) nitrogen sources in relation to ambient nutrient levels are explored.
Specific uptake rates (v), normalised to chlorophyll a, for NO;, NH4+ and urea were 0.2 ± 0.04 - 1831.1 ± 779.19, 0.5 ± 0.26 - 1731.6 ± 346.67 and 3.0 ± 0.60 - 2241.2 ± 252.56 ng N μg Chla-1 respectively. Urea concentration (14.8 - 117.7 μg urea-N 1-1) remained relatively constant over the 12 month study period. Measured ambient specific uptake rates for urea represent between 27.5% and 40.4% of total N uptake over the annual period February 1998 -January 1999. Seasonal nitrate uptake over the same period constituted only 11.3% (±10.77%, n=12) to 24.4% (± 13.02%, n=12) with the highest percentage during winter, when nitrate levels are elevated. It is suggested that urea provides a nutrient intermediary over the spring - summer period during transition from autotrophic to heterotrophic dominated communities. Grazing ,and nitrogen recycling are intricately connected by simultaneously providing top-down biomass control and bottom-up nutrient supply. Zooplankton (> 44 μm) grazing has been shown to reduce up to 40% of phytoplankton standing stock at times. Microheterotrophs (<300 pm) can reduce phytoplankton biomass production by up to 100% (potential production grazed, 11.1% day' - 99.6 % day-1) over an annual cycle. This correlated to mean seasonal day-time grazing loss of 80.47 ± 3.5 ngN μg Chla-1 in surface waters and 20.17 ± 9.7 ngN μg Chla-1 at depth (4.5m). Night time grazing for surface and bottom depths resulted in similar nitrogen loss rates (13.03 ± 4.84 ngN μg Chla-1).
Uptake rates for nitrate (r2 0.501) and urea (r2 0.512), doing with temperature (r2 0.605) were shown to have the greatest influence on phytoplankton distribution over depth and time. This research emphasises the need for more detailed investigations into the physiology of nutrient uptake and the effects of nutrient fluxes on phytoplankton biomass and distribution. Further research into the roles of organic nitrogen and pico and nanoplankton in this system is recommended.

The initiation, bloom, and bust of harmful Cyanobacteria and algae blooms (HAB) in lakes are controlled by top-down and bottom-up ecological controls. Excess phosphorous and nitrogen inputs from anthropogenic sources are primary to blame, but eukaryotic grazers may also promote or curb Cyanobacteria dominance. We tracked shifts in bacterial composition, lake chemistry, and eukaryotic grazing community weekly or bi-weekly through spring and summer and modeled the causes of specific Cyanobacterial species blooms and busts across three lakes in Utah, USA, with differing lake trophic states. Regardless of trophic status, all three lakes experienced blooms of varying composition and duration. Aphanizomenon strain MDT14a was the most dominant species in every bloom on Utah Lake, comprising up to 44.16% of the bacterial community. Utah Lake experienced a total of 18 blooms across all sites ranging in duration from one to six weeks. Phormidiaceae sp. (8.5  6.1%) and Microcystis sp. (9.7  4.7%) were the most abundant species in the Deer Creek bloom. Deer creek experienced one bloom at the beginning of fall. Nodularia sp. (9.7  2.1) dominated Great Salt Lake bloom. The Great Salt Lake experienced four separate blooms during the summer months that lasted one to three weeks. Phosphorous concentrations on Utah Lake varied across site and season. Nitrate concentrations on Deer Creek increased over season with a ten-fold increase in concentration. We characterized Cyanobacteria blooms as either bloom communities (growing populations of Cyanobacteria) or as bust communities (declining populations of Cyanobacteria). Using these designations, we modeled the growth and decline of the Cyanobacteria populations across season with top-down and bottom up-controls. Based on generalized least-squared modeling, eukaryotic grazing does not affect relative Cyanobacteria abundances as much as nutrient limitations. Aphanizomenom strain MDT14a was positively correlated with temperature (P < 0.028) and the concentration of K (P = 0.007) and negatively correlated with increases in conductivity (P = 0.0088). Microcystis was positively correlated with increasing levels of SRP (P < 0.001) and negatively correlated with higher Ca concentrations (P = 0.008) and PP (P = 0.008). Busts of Microcystis were related to decreases in nitrate (P = 0.06) and lower total lake depths (P = 0.03). Phormidiaceae sp. relative abundance was negatively correlated with higher levels of TDN (P = 0.01-0.001) and Mg (P = 0.01) and positively correlated with higher S concentrations (P = 0.007). Our findings suggest that micronutrients and more bioavailable forms of P may potentially allow Cyanobacteria to break dormancy and proliferate HAB communities.

Algal morphology is considered to be influenced by physiological and environmental factors such as stress and disturbance; one key disturbance exerted on algae is grazing pressure from herbivores. The factors upon which herbivores base their food preferences include algal attractiveness and edibility. Steneck and Watling's (1982) functional group model advocates the combination of algal species into seven groups based upon morphology and ecological function; the basic premise being that algal attractiveness and edibility will decrease hierarchically from functional group one to functional group seven, and that species within a group will be similarly attractive and edible. Two grazers commonly found in the intertidal area are the gastropod Littorina littorea and the isopod !dotea granulosa. Attractiveness and edibility experiments were conducted in which both grazers were presented with algae representing functional groups two to seven in single- and two-way choice experiments, in order to determine whether their food preferences could be predicted using a functional group approach. Although L. littorea and I. granulosa exhibited preference for algae between functional groups with regards to attractiveness and edibility, preference for both functional group and algal species differed between grazers. When the effect of algal morphology was investigated by presenting grazers with algal homogenates in agar, both grazers displayed a preference for algal extracts in agar over whole plant material, and exhibited preference for algae both between- and within-functional groups. The fact that not all species within a functional group were consumed in similar amounts contradicts the functional group model. Further investigation of the effect of algal availability on the feeding preferences of L. Iittorea showed their food choices could not be predicted based upon the algal species dominant in their habitat of origin. Results from all experimental manipulations suggest that both the habitat and food requirements of a herbivore, along with algal characteristics such as morphological, structural and chemical defences, algal availability and nutritional composition all influence herbivore food choice. As such, the functional group model proposed by Steneck and Wading (1982) cannot be accurately used to predict the feeding preferences of L. littorea or I. granulosa. It is suggested that unless models are modified to meet the requirements of a specific question, current functional group approaches are not a useful tool for predicting algal-grazer interactions.

The aim of the present study was to determine the potential impact of intertidal grazers on the distribution of macro algae on the south coast of South Africa. Particular attention was paid to the large patellid limpet, Patella oculus, which is found thoughout the intertidal zone. Studies of gut contents revealed that Patella oculus was capable of ingesting not only the thallus of foliose (eg. Ulva spp.) and encrusting coralline macroalgae, but also spores and diatoms. The inclusion of these relatively small particles in the diet was surprising, as electron micrographs of the radula of P.oculus revealed that it is typically docoglossan in structure. Such radulae are thought to be poorly suited for collecting small food particles. Sand made up a significantly higher proportion of the gut contents than other particles at all shore heights, which suggests that P.oculus might be capable of excavating the rocky substratum, or of sweeping up sand, while searching for food. Analysis of the gut contents of other local herbivorous molluscs, was also carried out. These species included the winkles, Oxystele variegata and O.sinensis, and the small pulmonate limpets, Siphonaria concinna, S.capensis, and S.serrata. The guts of all species contained mainly spores and diatoms, although small fragments of Ulva sp. were found. The population structure of Patella oculus was investigated at two sites, Cannon Rocks and Old Woman's River. At Cannon Rocks, mean shell length of low-shore animals was significantly lower than that of both mid- and high-shore animals, while at Old Woman's River, no significant difference was found among shore heights. A regression equation for In (shell length) vs In (dry weight) was calculated, and based on length data, the biomass density (g dry mass.m⁻²) of P.oculus at Old Woman's River was estimated. Values ranged from 2.8 on the low- and midshore to 0.37 on the high-shore. A manipulative field experiment was used to determine the impact of mesograzers and macrograzers (such as Patella oculus) on the distribution of intertidal macro algae on the mid- and low-shore at Old Woman's River. Grazers were excluded using mesh cages (mesh size = 3mm), in two separate experiments, one in winter and the other in spring. Percentage cover of macroalgal species and sessile invertebrates was estimated at approximately 6 week intervals for up to 3 months. MANOV A showed that treatments did not significantly affect cover of macroalgae or barnacles during winter. However, towards the end of the spring experiment (midshore only) cover of barnacles and green foliose turfs did increase in those plots from which mesograzers and/or macro grazers were excluded. The failure of the statistical tests to detect significant differences at some time intervals may have been caused by high levels of variation among replicates. This suggests that factors other than grazing are of overriding importance in determining the distribution of local macroalgae. The existence of a possible symbiotic relationship between Patella oculus and the red foliose alga, Gelidium pristoides, was investigated. The availability of various substratum types, including rock, limpet shells, barnacles etc., and the proportion of the total cover of G.pristoides on each, was calculated. It was shown that a significantly higher proportion of the alga grew on limpet shells, although the availability of this substratum type was low. It is thought that the aggressive behaviour of P.oculus prevents all but juvenile Patella longicosta from grazing on its shell, thus providing a refuge from grazing for G.pristoides.

There has been much interest in the last decade concerning the factors that influence diversity, especially how diversity and ecosystem processes may be linked. This study was based in small, cobble streams in South East Queensland. Its aim was to determine how the diversity and composition of consumers (the grazer guild) is influenced by both the production and composition of benthic algae, at different spatial scales. It also aimed to ascertain whether this response differs among grazer sub guilds with different dispersal capabilities. Ten sites in the Upper Brisbane and Mary catchments were sampled. The sites were selected to provide a range of productivity and composition. Grazers from these sites included snails and elmids, and larval mayflies, moths, and caddisflies. Grazer diversity and composition appeared to be structured by catchment scale influences, but environmental variables also affected which animals colonised patches and microhabitats (cobbles) within catchments. Primary productivity and algal composition could not be separated, with highly productive reaches also having a high cover of filamentous algal taxa. Grazer diversity displayed strongly positive, linear relationships with algal variables at the reach scale. It had a negative relationship with filamentous algae at the cobble scale, and a non-significant hump-shaped relationship with primary productivity. Survey data alone could not separate whether grazers were responding to habitat or food-related drivers, or to variations in productivity. Experimental manipulation of algal variables at the patch scale, using light and nutrients, also could not clearly uncouple the relationship between primary productivity and filamentous algal cover. Once reach scale variation was removed, grazer diversity displayed hump-shaped relationships with algal variables, including algal diversity. Much of this variation was due to patterns in mobile grazers, as sedentary grazers did not respond to algal variation at this scale. The density of the more mobile taxa showed similar patterns to those at the cobble scale (hump-shaped). A second field experiment was carried out in order to further investigate the responses of invertebrates to algal community composition at the cobble scale. Data from all three chapters suggested that as sites shifted to a dominance of filamentous algae, often with an associated increase in GPP, there was also a shift in the grazer community towards more sedentary grazers and away from the more mobile taxa. This also occurred at the cobble scale in the second experiment. The gut analysis and diet studies in the third chapter indicated that while many grazers consumed filamentous algae, it was not assimilated. This suggests that the preferences for sedentary taxa for cobbles and reaches dominated by filamentous algae are likely to be due to some other, possibly habitat-related, factor such as flow or predation refuge. The study provides a rare examination of relationships between primary productivity and consumer diversity in freshwater streams, and finds support for the pattern found in other systems of monotonic relationships of these two variables at large scales and hump-shaped relationships at smaller scales. It emphasises the importance of understanding other, potentially confounding, aspects of communities of producers, and investigates the possible roles of the most important of these (community composition) in structuring consumer communities in the small cobble streams of South-East Queensland.

There has been much interest in the last decade concerning the factors that influence diversity, especially how diversity and ecosystem processes may be linked. This study was based in small, cobble streams in South East Queensland. Its aim was to determine how the diversity and composition of consumers (the grazer guild) is influenced by both the production and composition of benthic algae, at different spatial scales. It also aimed to ascertain whether this response differs among grazer sub guilds with different dispersal capabilities. Ten sites in the Upper Brisbane and Mary catchments were sampled. The sites were selected to provide a range of productivity and composition. Grazers from these sites included snails and elmids, and larval mayflies, moths, and caddisflies. Grazer diversity and composition appeared to be structured by catchment scale influences, but environmental variables also affected which animals colonised patches and microhabitats (cobbles) within catchments. Primary productivity and algal composition could not be separated, with highly productive reaches also having a high cover of filamentous algal taxa. Grazer diversity displayed strongly positive, linear relationships with algal variables at the reach scale. It had a negative relationship with filamentous algae at the cobble scale, and a non-significant hump-shaped relationship with primary productivity. Survey data alone could not separate whether grazers were responding to habitat or food-related drivers, or to variations in productivity. Experimental manipulation of algal variables at the patch scale, using light and nutrients, also could not clearly uncouple the relationship between primary productivity and filamentous algal cover. Once reach scale variation was removed, grazer diversity displayed hump-shaped relationships with algal variables, including algal diversity. Much of this variation was due to patterns in mobile grazers, as sedentary grazers did not respond to algal variation at this scale. The density of the more mobile taxa showed similar patterns to those at the cobble scale (hump-shaped). A second field experiment was carried out in order to further investigate the responses of invertebrates to algal community composition at the cobble scale. Data from all three chapters suggested that as sites shifted to a dominance of filamentous algae, often with an associated increase in GPP, there was also a shift in the grazer community towards more sedentary grazers and away from the more mobile taxa. This also occurred at the cobble scale in the second experiment. The gut analysis and diet studies in the third chapter indicated that while many grazers consumed filamentous algae, it was not assimilated. This suggests that the preferences for sedentary taxa for cobbles and reaches dominated by filamentous algae are likely to be due to some other, possibly habitat-related, factor such as flow or predation refuge. The study provides a rare examination of relationships between primary productivity and consumer diversity in freshwater streams, and finds support for the pattern found in other systems of monotonic relationships of these two variables at large scales and hump-shaped relationships at smaller scales. It emphasises the importance of understanding other, potentially confounding, aspects of communities of producers, and investigates the possible roles of the most important of these (community composition) in structuring consumer communities in the small cobble streams of South-East Queensland.
Thesis (PhD Doctorate)
Doctor of Philosophy (PhD)
Australian School of Environmental Studies
Full Text

Benthic environments in streams are areas of high biological activity where primary producers and consumers interact with each other and the environment. Various factors such as light, nutrient availability and temperature affect algal biomass accumulation however disturbance factors and grazing lead to biomass losses as well as changes in the species composition and densities of algae. There are little, if any, studies done on the effects of grazers on benthic algae in South African rivers and as such, this experiment was done in the Berg River, in the Western Cape of South Africa. The aim was to ascertain the effects of herbivores on benthic algae within the river and lit was hypothesized that periphyton biomass would be higher where grazer densities were lower and that the differences observed in biomass of periphyton between treatments would increase over time. It was also hypothesized that there would be a lower diversity of algae where there were lower herbivore densities. A randomized block design experiment was set up in the Berg River and treatments of 'excluded herbivores' and 'included herbivores' were set up, with two pairs of each treatment spread across four replicates. Stones from the stream were randomly selected and placed in the boxes. After five days half of the boxes were sampled and after eight days, the remaining half was sampled. Stones were scrubbed to remove periphyton from the stones and a sub-sample of 50ml was removed from each for later algal species identification. Invertebrates were removed from the boxes, preserved in ethanol and later identified to family level. Chl-a and AEDW (mg/m²) were determined from the samples as were invertebrate and algal densities. The results showed that there were no herbivore density effects on biomass most likely due to the season of sampling and the length of the experiment. However, the effect on algal densities was more pronounced as there was a higher variety of algal forms present where herbivore densities were lower. To get a better idea of how herbivores affect periphyton other factors need to be taken into account like experimental time period, season and different environmental conditions.

Os estudos apresentados nesta tese tiveram como objetivo compreender aspectos da organização de assembleias de algas bênticas em riachos. Foi avaliado especificamente o efeito da heterogeneidade de habitat na escala de rugosidade de substratos e sua interação com distúrbios hidrológicos e organismos pastejadores (grazers). Utilizou-se uma abordagem experimental in situ com substratos lisos e rugosos (com fendas) constituindo os tratamentos. Os experimentos foram conduzidos no Rio do Marco, um riacho de quarta ordem na região dos Campos de Cima da Serra, em São José dos Ausentes, Rio Grande do Sul. Após uma introdução geral acerca do assunto são apresentados os capítulos com os estudos. O Capítulo 1 avaliou os efeitos da rugosidade de substratos sobre a riqueza de espécies, densidade, composição e padrões de distribuição de assembleias de algas bênticas. Assembleias em substratos rugosos foram mais ricas que assembleias em substratos lisos, porém esse resultado diferiu entre grupos com diferentes formas de vida, assim como as diferenças na composição de espécies, indicando que somente alguns grupos foram beneficiados pela ocorrência de refúgios em substratos rugosos. Observou-se também que diferenças na composição da assembleia entre substratos lisos e rugosos resultaram de aninhamento e substituição de espécies. No Capítulo 2 foi testada a hipótese de que substratos rugosos abrigam assembleias mais persistentes que substratos lisos. Os resultados encontrados confirmaram esta hipótese e foram atribuídos ao efeito da rugosidade per se após a exclusão de alguns fatores de confundimento, como estabilidade física dos substratos e diferenças na riqueza de espécies. Concluiu-se que o mecanismo mais plausível para explicar a maior persistência em substratos rugosos é a maior disponibilidade de refúgios em relação a substratos lisos. O objetivo do Capítulo 3 foi avaliar o efeito da rugosidade de substratos sobre assembleias de algas frente a um distúrbio hidrológico experimental, testando especificamente se a resistência e a resiliência de algas é maior em substratos rugosos que em substratos lisos e se a resposta de diferentes formas de vida é distinta. Os resultados indicaram que a rugosidade de substratos não apresenta efeitos pronunciados sobre a resistência e a resiliência de algas bênticas. Porém, grupos de algas com distintas formas de vida diferiram quanto à resistência e resiliência, de forma que o padrão observado está relacionado a atributos biológicos que conferem habilidades para suportar distúrbios. Os resultados deste estudo em conjunto com os resultados obtidos no Capítulo 2 e resultados obtidos por outros autores permitiram que fosse proposto que a importância da heterogeneidade de habitas e de refúgios é mediada pela intensidade de distúrbios. No Capítulo 4 foram avaliados os efeitos independentes e as interações de rugosidade de substratos, peixes pastejadores e tipos de mesohabitats (corredeiras e remansos) sobre a biomassa algal, matéria orgânica e peso seco total. O efeito de peixes pastejadores não foi mediado pelos mesohabitats, sendo que a exclusão de peixes causou aumento nas três variáveis resposta, independente do tipo de mesohabitat. Por outro lado, houve uma interação entre mesohabitats e substratos para determinar o acúmulo de peso seco total, sendo que substratos rugosos acumularam maior quantidade de material em remansos que em corredeiras, enquanto substratos lisos acumularam quantidades similares entre os dois tipos de mesohabitats. Já a biomassa algal e matéria orgânica apresentaram maior acúmulo nos substratos rugosos em relação aos lisos, independente do mesohabitat. Os resultados encontrados indicam que a perda de organismos pastejadores pode afetar o acúmulo de biomassa algal e material orgânico, importantes recursos alimentares em riachos. Finalmente, na última seção são apresentadas as considerações finais.
The studies presented in this thesis aimed at understanding some aspects of the organization of benthic algal assemblages in streams. Specifically, the studies evaluated the effect of habitat heterogeneity on the scale of substrate roughness and its interaction with hydrological disturbances and grazing organisms. An experimental in situ approach was used, in which the treatments were constituted by smooth and rough (with crevices) substrates. The experiments were conducted in the Marco River, a fourth-order stream at the region of Campos de Cima da Serra, São José dos Ausentes, Rio Grande do Sul, Brazil. After a general introduction on the subjects of this thesis, the chapters with the studies are presented. The Chapter 1 describes a study to evaluate the effects of substrate roughness on species richness, density, composition and distribution patterns of benthic algal assemblages. Algal assemblages on rough substrates were richer than on smooth substrates, but this result differed among algal life forms. Similarly, results on species composition also differed among life forms, indicating that only some groups were benefited by the occurrence of refuges on rough substrates. It was also observed that differences in the composition of assemblages between rough and smooth substrates resulted from nestedness and turnover. The Chapter 2 describes a study in which was tested the hypothesis that rough substrates harbor more persistent assemblages than smooth substrates. The results confirmed this hypothesis and were attributed to the effect of roughness per se after the exclusion of some confounding factors, such as the physical stability of substrates and differences in species richness. It was concluded that the most plausible mechanism that could explain the higher persistence in rough substrates is the greater availability of refuges in relation to smooth substrates. The main aim of the study described in Chapter 3 was to evaluate the effect of substrate roughness on algal assemblages in the face of an experimental hydrological disturbance. Specifically, it was tested if the resistance and resilience of algae is higher on rough than on smooth substrates and if algal life forms differ in their response to disturbance. The results indicated that substrate roughness had no pronounced effects on benthic algal resistance and resilience. However, algal life forms differ in their resistance and resilience, and the observed pattern was mostly related to biological traits that confer abilities to support disturbances. The results of this study, in addition to the results obtained in Chapters 1 and 2, allowed us to suggest that the importance of habitat heterogeneity and refuges is mediated by the intensity of disturbances. The Chapter 4 describes a study that evaluated the independent and interacting effects of substrate roughness, grazing fish and mesohabitats (pools and riffles) on algal biomass, organic matter and total dry mass accrual. The effect of grazers was not mediated by mesohabitats, and their exclusion caused an increase in the three response variables, regardless of mesohabitat. On the other hand, mesohabitats and substrates interacted to determine the accrual of total dry mass, since rough substrates accumulated greater amounts of dry mass in pools than in riffles, while smooth substrates accumulated similar amounts in both mesohabitats. Algal biomass and organic matter showed greater accrual on rough than on smooth substrates, independently of mesohabitat. These results indicate that the loss of grazers may affect the accrual of algal biomass and organic matter, which are important food resources in streams. Finally, in the last section are presented the concluding remarks.

In this study, the effects of biological and physical disturbance on algal assemblages on cobbles in rainforest streams were investigated. Study sites in two rainforest streams, Stony Creek and Booloumba Creek, located in south- east Queensland, were used for both a regular sampling program and an extensive experimental schedule. Previous research has shown that physical disturbance due to spates and, in the absence of high discharge events, biological disturbance due to invertebrate grazing exert significant structuring and controlling forces on periphyton assemblages in streams. The principal objectives in this study were to deternine the relative importance of physical disturbance in the form of spates and desiccation, to outline the significance of biological effects, in particular as a result of grazing pressure, and to describe algal succession on substrates cleared after disturbance events. Periphyton chlorophyll a, biomass values and grazer numbers were monitored over a period of two years in relation to rainfall and stream discharge (to pinpoint the incidence of spates). Periphyton assemblages were composed mainly of diatoms (Cocconeis and Navicula being the dominant genera). Physical disturbance by spates was clearly an important factor structuring stream algal assemblages. Periphyton chlorophyll a and biomass values were reduced following a high discharge event, but then recovered relatively fast in the absence of any other disturbance. Furthermore, chlorophyll a values were correlated with grazer numbers. Differences were observed in the epilithon of Stony Creek and Booloumba Creek: while cobbles in the former supported more chlorophyll a containing material of algal origin and less non-algal biomass, the reverse was true for latter stream. This was most probably due to the physical characteristics of the two stream channels, primarily their slope, which was considerably steeper in Stony Creek, thus requiring less depth to facilitate a major disturbance event. This means that the physical characteristics of streams are most important determining factors regarding the effects of high discharge events, with the disturbance regime playing a significant role in the composition of the stream benthos. However, some major changes in chlorophyll a and biomass occurred in the absence of big disturbances. Periphyton on cobbles collected in the streams was subjected to various time periods out of the water to determine the effects of desiccation. Diatom assemblages on cobbles were not resistant to desiccation and did not recover following re-immersion in the streams. Thus, periods of low flow are capable of reducing periphyton assemblages, however, this type of disturbance would take place over a longer period compared to spates. A colonization experiment where polyethylene foils were introduced into the streams was used to monitor algal succession on bare substrates. Diatoms were the only algae which colonized experimental foils, exhibiting distinct successional patterns in both streams. A series of grazer manipulation experiments were employed to test for any effects of grazing; this involved the inclusion / exclusion of caddisfly larvae from cobbles enclosed in screened perspex cylinders. Signficantly, the effects of grazing invertebrates resulted in an increase in chlorophyll a and biomass values, contrary to expectations and the majority of past studies. This could mean that grazers at a certain density may actually stimulate algal growth and there exists a fine balance between when grazer numbers enhance periphyton productivity or periphyton is overgrazed. Overall, results obtained indicated that variations in stream algal assemblages were a consequence of the combined effects of spates, low flows and, to a lesser extent, grazing.
Thesis (PhD Doctorate)
Doctor of Philosophy (PhD)
Graduate School of Environmental Science and Engineering
Science, Environment, Engineering and Technology
Full Text

Univ., Diss. u.d.T.: Kim, Dohong: Seasonal dynamics of an Antarctic rocky intertidal, with emphasis on the role of the limpet Nacella concinna Strebel (Gastropoda: Patellidae)--Bremen, 2000.
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The factors that determine the efficiency of energy transfer in aquatic food webs have been investigated for many decades. The plant-animal interface is the most variable and least predictable of all levels in the food web. In order to study determinants of food quality in a large lake and to test the recently proposed central importance of the long-chained eicosapentaenoic acid (EPA) at the pelagic producer-grazer interface, we tested the importance of polyunsaturated fatty acids (PUFAs) at the pelagic producerconsumer interface by correlating sestonic food parameters with somatic growth rates of a clone of Daphnia galeata. Daphnia growth rates were obtained from standardized laboratory experiments spanning one season with Daphnia feeding on natural seston from Lake Constance, a large pre-alpine lake. Somatic growth rates were fitted to sestonic parameters by using a saturation function. A moderate amount of variation was explained when the model included the elemental parameters carbon (r2 = 0.6) and nitrogen (r2 = 0.71). A tighter fit was obtained when sestonic phosphorus was incorporated (r2 = 0.86). The nonlinear regression with EPA was relatively weak (r2 = 0.77), whereas the highest degree of variance was explained by three C18-PUFAs. The best (r2 = 0.95), and only significant, correlation of Daphnia's growth was found with the C18-PUFA α-linolenic acid (α-LA; C18:3n-3). This correlation was weakest in late August when C:P values increased to 300, suggesting that mineral and PUFA-limitation of Daphnia's growth changed seasonally. Sestonic phosphorus and some PUFAs showed not only tight correlations with growth, but also with sestonic α-LA content. We computed Monte Carlo simulations to test whether the observed effects of α-LA on growth could be accounted for by EPA, phosphorus, or one of the two C18-PUFAs, stearidonic acid (C18:4n-3) and linoleic acid (C18:2n-6). With >99 % probability, the correlation of growth with α-LA could not be explained by any of these parameters. In order to test for EPA limitation of Daphnia's growth, in parallel with experiments on pure seston, growth was determined on seston supplemented with chemostat-grown, P-limited Stephanodiscus hantzschii, which is rich in EPA. Although supplementation increased the EPA content 80-800x, no significant changes in the nonlinear regression of the growth rates with α-LA were found, indicating that growth of Daphnia on pure seston was not EPA limited. This indicates that the two fatty acids, EPA and α-LA, were not mutually substitutable biochemical resources and points to different physiological functions of these two PUFAs. These results support the PUFA-limitation hypothesis for sestonic C:P < 300 but are contrary to the hypothesis of a general importance of EPA, since no evidence for EPA limitation was found. It is suggested that the resource ratios of EPA and α-LA rather than the absolute concentrations determine which of the two resources is limiting growth.

En raison d’une utilisation croissante et massive, les nanoparticules manufacturées apparaissentcomme de potentiels contaminants émergents pour l’environnement, incluant notammentles écosystèmes aquatiques. Alors que le transfert trophique semble constituer unevoie d’exposition majeure pour les organismes, une connaissance lacunaire dans la littératurescientifique persiste, résultant pour partie des difficultés expérimentales inhérentes àce type d’exposition. Pour ce travail en conditions contrôlées de laboratoire, les nanoparticulesd’or (sphériques, 10 nm, fonctionnalisées aux PEG-amines), stables en solution, ontété choisies pour l’étude du transfert trophique et des impacts toxiques sur des organismesaquatiques. Ce continuum trophique considère la base des réseaux trophiques (biofilms naturels,algues), des niveaux intermédiaires (poissons brouteurs, bivalves suspensivores), jusqu’auxorganismes de haut de chaînes trophiques, avec l’anguille européenne. Avec des expositionsréalisées à de relatives faibles doses, ce travail tend à la représentativité environnementale.Des approches méthodologiques intégratives des niveaux subcellulaire à tissulaire(RT-qPCR, séquençage haut-débit, histologie) ont permis d’évaluer les impacts toxiques.Les résultats indiquent une importante capacité de rétention des nanoparticules par les biofilmsnaturels. À la suite d’une exposition de 21 jours, les dosages d’or révèlent un transfertdes biofilms aux poissons brouteurs, avec une distribution de l’or dans tous les organes. Deplus, ce transfert est associé à une réponse inflammatoire au regard des lésions histologiquesobservés dans les foies, rates et muscles des poissons exposés. Une chaîne alimentaire « naturelle» à trois maillons trophiques, impliquant algues - bivalves - anguilles européennes,atteste d’un transfert significatif jusqu’au poisson prédateur. Enfin, l’analyse du transcriptome,par une approche de séquençage haut-débit, des foies et cerveaux d’anguilles exposéesaux nanoparticules par nourriture enrichie, a permis de mettre en évidence une réponseconjointe à ces deux organes dans des processus biologiques associés au système immunitaireet sa régulation, dont des récepteurs NOD-like impliqués dans l’inflammasome.L’ensemble des résultats expérimentaux interpellent quant aux effets délétères à long-termequ’engendreraient les nanoparticules sur les écosystèmes aquatiques, illustrant par ailleursla propension de ces contaminants à être transférés dans les chaînes trophiques
Due to an increasing and massive use, engineered nanoparticles are raising as potentialemerging contaminants in the environment, including aquatic ecosystems. While trophictransfer appears to constitute a major exposure route for organisms, scientific literature hasdifficulties to respond to the questions raised to explore the range of the interactions existingbetween nanoparticles and living organisms at different scales from the trophic interactionsto the cellular impacts. This problem is partly due to experimental difficulties inherent tothis exposure type. For this work performed in controlled laboratory conditions, sphericalgold nanoparticles (10 nm, coated with PEG-amines, positively charged) were chosen tostudy the trophic transfer and toxic effects on aquatic organisms. Trophic chains concernedseveral trophic levels (up to three) with a variety of species considered : the basis of thetrophic web with natural biofilms or microalgae, intermediate levels with grazing fish orsuspensivorous bivalves, and up to top food chain organisms, with the European eel, a carnivorousfish.With relatively low doses for exposures, this work tends to represent environmentalconditions. Integrative methodological approaches from subcellular to tissue levels(RT-qPCR, RNA-sequencing, histology) were performed in order to assess toxic impacts.The results indicate a high retention capacity of nanoparticles by natural biofilms. Followinga 21-day exposure, gold quantifications reveal a transfer from biofilms to grazing fish, witha gold distribution in all organs. Moreover, this transfer is associated with an inflammatoryresponse according to the histological lesions observed in the liver, spleen and muscle ofexposed fish. A longer food chain, with three trophic levels involving microalgae - bivalves- European eels, is set up to give a better representation of the complexity of trophic interactionsin the aquatic environment. It shows a significant transfer to the predatory fish.Transcriptomic analyses, using the RNA-sequencing approach, for the liver and the brain ofexposed eels by nanoparticles’ enriched food, highlight a joint response for these two organsin the biological processes associated with the immune system and its regulation, includingNOD-like receptors involved in inflammasome.All the experimental results suggest long-term harmful effects that nanoparticles would generatein aquatic ecosystems, emphasizing the ability of these contaminants to be transferredthroughout trophic chains

Harmful algal blooms (HABs) are becoming more prevalent throughout the world’s aquatic systems. These blooms have been the subjects of numerous studies because they can cause human health issues and economic impact through fish kills, contaminated shellfish and decreased tourism. Much research has focused on the “bottom-up” aspect of these blooms; namely, the potential role of increased nutrient input into coastal waters from anthropogenic sources causing increased growth in harmful algal species. However, there are also potential “top-down” controls affecting the rate at which harmful algal species are consumed by grazers. The aim of this project was to determine protozoan grazer population fluctuations and their grazing impact on HAB species through field monitoring and laboratory grazing experiments. Protozoan grazers were chosen because their growth rates could potentially keep up with those of HAB species. Declines in grazer populations before the onset of a bloom could be indicative of a release of the HAB from a “top-down” grazing control. Field samples taken during bloom and non-bloom events helped elucidate any microplankton community changes. After establishing that there appear to be changes to the grazer population before and after a bloom, ingestion experiments including direct epifluorescence microscopy and DNA analyses were conducted to determine if it is possible that a chosen protozoan grazer can ingest a HAB species. Finally, experiments were conducted to determine whether the HAB species was a favorable food source for the grazer. Population growth experiments in which grazers are fed a HAB species, 50:50 mixture, or normal culture food source were used to determine the survival and growth rate of the grazer. Although certain ciliates and heterotrophic dinoflagellates were found to feed on HAB species in the lab and in natural bloom samples, the HAB species as a food source produces lower grazer growth rates than on control food. Protozoan grazers may be a more effective control during bloom initiation than after the bloom has been established.
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Algal biofuel has been shown to have great potential to solve the World’s sustainable energy crisis but technologies for large-scale cultivation are still elusive. While photobioreactors meet very high value algal products there is still no technology for producing algae by the millions of tons. Flooded paddy lands of India offer excellent opportunities for co-cultivation of algae with paddy crop provided it meets various sustainability criteria, not to mention very low cost options. This research examines sustainability, technology and climate change challenges to this above concept termed “Algiculture”. The potential of naturally emerging algal consortia to overcome travails of pure-culture, the ability to scavenge ‘lost’ plant nutrients in flooded paddy, overcome threats by grazers and predators, evolving naturally mediated techniques to harvest algae, impact on methane emissions, etc. were examined critically under laboratory and flooded paddy conditions. Experimental results indicate that many of the sustainability criteria can be met by growing algae simultaneously with a paddy crop for the first 60-75d which doubles the overall biomass yield from such lands. Algae raised can scavenge ammoniacal-N that generally occurs as unavoidable losses in flooded paddy system and can thus be raised without additional fertilizer inputs. This simultaneously ameliorates the N-pollution from paddy runoff to water bodies. Algal cultivation with paddy (Algiculture) alters micro-environmental conditions e.g. oxygen supersaturation, to make methane emissions unfavourable and by contrast algae even take up the C hitherto wasted away as methane and thereby converting an environmental liability to conservation. Consortia dominated by Chlorella and Chlorococcum sp. along with a small number of Cyanophycaeans facilitate simple low energy algal harvest techniques employing clumping and floc-formation that enables maintaining appropriate stocking density of algae and allowing continuous operation. The pattern of grazer /predator occurrence in such systems, techniques to minimize their influence by merely altering the cultivation conditions have been worked out and tested successful. The causes of reduction in methane emissions and C-source identification have been assessed with 13C discrimination studies. The research creates a new potential reducing GHG on the one hand for also raising sustainable bioenergy options in India as well as in all flooded paddy lands of the world

Dissertação mest., Estudos Marinhos e Costeiros, Universidade do Algarve, 2007
Serrão, Ester
Herbivores have been repeatedly shown to induce defences in terrestrial plants, but few examples exist from marine macroalgae. In two bio-assayed laboratory experiments we tested the effects of (1) direct amphipod consumption, (2) water borne cues from nearby grazed conspecifics, and (3) water-borne cues from non grazing consumers on the induction of anti-herbivore defences in macroalgae of the southern coast of Portugal. Both the brown alga Fucus vesiculosus and the red alga Sphaerococcus coronopifolius displayed distinct defence regulation. They induced defences when consumers were present and reduced defences again when consumption ceased. A reduction in palatability was considered to be indicative of an alga’s defence mechanism. Defence regulation reacted to consumption stimuli in less than two weeks. For F. vesiculosus we found the defence activity in the lipophilic fraction of the alga’s chemistry. Both algal species induce defences after receiving water-borne signals from nearby grazed conspecifics as well as non-grazing amphipods. These results imply that species of red and brown algae can recognise a suite of cues indicating herbivore pressure, allowing them to adjust their defensive traits in finely tuned short-term responses. Inducible defences may be more common in macroalgae than suspected.