At temperate latitudes where seasonal changing environmental conditions strongly affect the magnitude, duration and species composition of pelagic primary production, macrobenthic organisms living below the photic zone rely on the sedimentation of organic matter as their primary energy source. The succession from nutritious spring blooms to summer cyanobacteria is assumed to reduce food quality for benthic primary consumers and their fatty acid (FA) profiles. In contrast, we find low seasonal variability in FA content of five benthic macroinvertebrates spanning two trophic levels in the Baltic Sea, a system with high seasonal variation in phytoplankton species composition. However, levels of the major FA groups vary greatly between benthic species. The results suggest that benthic macroinvertebrates have evolved FA metabolism adapted to degraded sedimenting material. Moreover, our study shows that species composition of benthic macrofauna rather than seasonal changing conditions affect availability of essential nutrients to higher trophic levels.

Marine soft sediments form the second largest habitat on the planet. Organisms residing in this environment represent a vast reservoir of biodiversity, and play key roles in ecosystem processes. Most benthic organisms depend on organic matter (OM) inputs from phytoplankton in the overlying water column as food supply, but human impacts such as eutrophication and climate change are profoundly altering natural ecosystem dynamics. The consequences of changes in benthic-pelagic coupling for the biodiversity and functioning of soft-sediment communities have yet to be resolved. 

The aim of this thesis is to assess the role of OM settling on soft-sediments microeukaryotic (small organisms < 1 mm) and bacterial communities. The intents are two-fold, to investigate impacts on (1) community structure and diversity (chapters I, II and IV); and (2) ecosystem functioning, notably in relation to nitrogen (N) cycling (chapters I and III). 

Our results show that settling OM quantity and quality both had a significant impact on microeukaryotic alpha-diversity. We observed a decrease in alpha-diversity following settling of diatom-derived spring bloom OM, possibly as a result of competitive exclusion, while cyanobacteria-derived summer bloom OM did not affect alpha-diversity (chapters I and IV). We also found that high biomass of diatoms and others fast sinking phytoplankton groups in the water column led to lower microeukaryotic alpha diversity after this material settled on the seafloor (chapter IV). Presumably, following this large sedimentation event, sediment oxygen (O₂) demand was strongly stimulated, excluding O₂-sensitive taxa. Overall, we propose that the assembly of microeukaryotic communities was primarily mediated by OM settling quantity (chapter IV), while differences in OM quality led to significant but more subtle changes, occurring at fine taxonomic level (chapter I). The response of bacterial communities to OM settling was less pronounced, and probably restricted to the uppermost sediment layer (chapters I and IV). We did, however, observe a significant effect of OM quality on bacterial communities assembly at the sediment-water interface, with taxa favored either by diatom- or by cyanobacteria-derived OM (chapter II). This study also showed that feedback mechanisms from nutrient recycling in the sediment could play a role in this response. Finally, our results indicated a substantial influence of OM quality on N cycling at the sediment-water interface. We found that settling of fresh OM (i.e. low C:N ratio) stimulated denitrification activity (chapters I and III), while simultaneously promoting more N recycling to the water column than settling of degraded OM (i.e. high C:N ratio) did (chapter III).  

Altogether, our results indicate that current changes in OM settling dynamics in marine systems will likely impact microeukaryotic and, to some extent, bacterial biodiversity in soft sediments. Alterations in settling OM quality, in particular, may also affect crucial microbial processes involved in N cycling. This thesis highlights the importance of considering benthic-pelagic coupling mechanisms to better understand likely future changes in marine ecosystems.

Pelagic primary production is the main input of organic energy for benthic production below the photic zone. In the Baltic Sea, spring phytoplankton blooms are dominated by diatoms that sink out rapidly and export nutritionally favorable matter to benthic secondary production, while the summer blooms have more variable sedimentation rates and nutritional profile. Changes in phytoplankton species composition and bloom dynamics, as a consequence of climate change and eutrophication are reducing high quality diatoms reaching the benthic fauna, while promoting cyanobacteria. Here, we test uptake and assimilation of changing phyto- plankton composition for three common benthic invertebrates, a clam, an amphipod and a polychaete under varying degrees of spring-bloom associated diatoms (Skeletonema costatum) and summer-bloom associated cyano- bacteria (Nodularia spumigena). The phytoplankton were labeled with stable isotopes (15N and 13C, respectively) in order to trace assimilation in consumers’ tissues. We found that all three macrofauna species fed on both dia- toms and cyanobacteria. A linear pattern was found for all three species in assimilation of carbon and nitrogen from diatoms, with increasing assimilation associated with higher proportion of diatoms. There was no clear pattern found between proportion of cyanobacteria and assimilation of carbon and nitrogen for any of the spe- cies. This study shows that the investigated macrofaunal species display a selective feeding behavior with prefer- ence for spring-bloom associated diatoms. Thus, changes in phytoplankton bloom composition are likely affecting benthic species composition and production. 

Coastal sediments are major contributors to global carbon (C) mineralization and nutrient cycling and are tightly linked to processes in the pelagic environment. In this study, we aimed to investigate the regulating potential of quantity and quality of planktonic organic matter (OM) deposition on benthic metabolism, with a particular focus on nitrogen (N) cycling processes. We simulated inputs of spring (C : N 10.9) and summer (C : N 5.6) plankton communities in high and low quantities to sediment cores, and followed oxygen consumption, nutrient fluxes as well as nitrate reduction rates, that is, denitrification and dissimilatory nitrate reduction to ammonium for 10 d. Our results demonstrate the primary importance of OM quality in determining the fate of organic N once it settles to the sediment surface. Settling of N-rich summer plankton material resulted in a similar to twofold lower denitrification efficiency (40-56%) compared to N-poor spring plankton (88-115%). This indicates that N-rich plankton deposition favors recycling of inorganic nutrients to the water column over N-loss via denitrification. OM quantity was positively related to mineralization activity, but this neither directly affected N fluxes nor denitrification activity, highlighting the complex interplay between the OM quantity and quality in regulating N cycling. In light of these new findings, we support the use of simple qualitative indicators such as C : N ratio of OM to investigate how future changes in benthic-pelagic coupling might influence N budgets at the sediment-water interface.

In coastal aphotic sediments, organic matter (OM) input from phytoplankton is the primary food resource for benthic organisms. Current observations from temperate ecosystems like the Baltic Sea report a decline in spring bloom diatoms, while summer cyanobacteria blooms are becoming more frequent and intense. These climate-driven changes in phytoplankton communities may in turn have important consequences for benthic biodiversity and ecosystem functions, but such questions are not yet sufficiently explored experimentally. Here, in a 4-week experiment, we investigated the response of microeukaryotic and bacterial communities to different types of OM inputs comprising five ratios of two common phytoplankton species in the Baltic Sea, the diatom Skeletonema marinoi and filamentous cyanobacterium Nodularia spumigena. Metabarcoding analyses on 16S and 18S ribosomal RNA (rRNA) at the experiment termination revealed subtle but significant changes in diversity and community composition of microeukaryotes in response to settling OM quality. Sediment bacteria were less affected, although we observed a clear effect on denitrification gene expression (nirS and nosZ), which was positively correlated with increasing proportions of cyanobacteria. Altogether, these results suggest that future changes in OM input to the seafloor may have important effects on both the composition and function of microbenthic communities.

Phytoplankton comprises a large fraction of the vertical carbon flux to deep water via the sinking of particulate organic matter (POM). However, despite the importance of phytoplankton in the coupling of benthic-pelagic productivity, the extent to which its deposition in the sediment affects bacterial dynamics at the water-sediment interface is poorly understood. Here, we conducted a microcosm experiment in which varying mixtures of diatom and cyanobacteria, representing phytoplankton-derived POM of differing quality, served as inputs to sediment cores. Characterization of 16S rRNA gene of the bacterial communities at the water-sediment interface showed that bacterial α-diversity was not affected by POM addition, while bacterial β-diversity changed significantly along the POM quality gradient, with the variation driven by changes in relative abundance rather than in taxon replacement. Analysing individual taxa abundances across the POM gradient revealed two distinct bacterial responses, in which taxa within either diatom- or cyanobacteria-favoured groups were more phylogenetically closely related to one another than other taxa found in the water. Moreover, there was little overlap in taxon identity between sediment and water communities, suggesting the minor role played by sediment bacteria in influencing the observed changes in bacterial communities in the overlying water. Together, these results showed that variability in phytoplankton-originated POM can impact bacterial dynamics at the water-sediment interface. Our findings highlight the importance of considering the potential interactions between phytoplankton and bacteria in benthic-pelagic coupling in efforts to understand the structure and function of bacterial communities under a changing climate.