This study addresses organic matter decomposition in permeable sediment of a sloping intertidal sand flat (German Wadden Sea) affected by current-induced pore water exchange and pore fluid drainage. Seasonal and spatial scales of aerobic and anaerobic mineralization were investigated at 2 sites, one near the water line and one on the upper flat. Hydrodynamic forcing during inundation caused deeper oxygen penetration through flushing of the uppermost sediment layer. This flushing resulted in higher areal oxygen consumption rates and lower depth integrated sulfate reduction rates in the submerged flat compared to the rates measured during exposure. Mineralization rates in the top 15 cm of the sediment were similar between both study sites and ranged from 38 (winter) to 280 mmol C m -2 d -1 (summer), with sulfate reduction contributing 3 to 25% to total mineralization, depending on the season. At the upper flat, these seasonal differences were reflected in the pore water concentrations of nutrients, dissolved inorganic carbon (DIC) and dissolved organic carbon (DOC). Near the low water line, however, pore water nutrient and DIC concentrations were independent of the season and up to 15 times higher compared to the values recorded in the upper flat. The differences in concentrations of metabolic products between the 2 sites resulted from a low tide drainage extending deep below the uppermost flushed layer and causing seepage of pore water near the low water line. Mineralization and nutrient release in these permeable intertidal sediments is affected by 2 circulation processes that work on distinctly different temporal and spatial scales: (1) rapid 'skin circulation' through the uppermost sediment layer during inundation that is characterized by short flow paths, low pore water residence time and immediate feedback to the ecosystem, and (2) slow 'body circulation' through deeper sediment layers during low tide that is characterized by long flow paths and pore water residence times, and acts as a buffered nutrient source to the ecosystem.

Nitrogen (N) input to the coastal oceans has increased considerably because of anthropogenic activities, however, concurrent increases have not occurred in open oceans. It has been suggested that benthic denitrification in sandy coastal sediments is a sink for this N. Sandy sediments are dynamic permeable environments, where electron acceptor and donor concentrations fluctuate over short temporal and spatial scales. The response of denitrifiers to these fluctuations are largely unknown, although previous observations suggest they may denitrify under aerobic conditions. We examined the response of benthic denitrification to fluctuating oxygen concentrations, finding that denitrification not only occurred at high O concentrations but was stimulated by frequent switches between oxic and anoxic conditions. Throughout a tidal cycle, in situtranscription of genes for aerobic respiration and denitrification were positively correlated within diverse bacterial classes, regardless of O concentrations, indicating that denitrification gene transcription is not strongly regulated by O in sandy sediments. This allows microbes to respond rapidly to changing environmental conditions, but also means that denitrification is utilized as an auxiliary respiration under aerobic conditions when imbalances occur in electron donor and acceptor supply. Aerobic denitrification therefore contributes significantly to N-loss in permeable sediments making the process an important sink for anthropogenic N-inputs.

In this article, we describe the dynamics of pH, O 2 and H 2 S in the top 5-10 cm of an intertidal flat consisting of permeable sand. These dynamics were measured at the low water line and higher up the flat and during several seasons. Together with pore water nutrient data, the dynamics confirm that two types of transport act as driving forces for the cycling of elements (Billerbeck et al. 2006b): Fast surface dynamics of pore water chemistry occur only during inundation. Thus, they must be driven by hydraulics (tidal and wave action) and are highly dependent on weather conditions. This was demonstrated clearly by quick variation in oxygen penetration depth: Seeps are active at low tide only, indicating that the pore water flow in them is driven by a pressure head developing at low tide. The seeps are fed by slow transport of pore water over long distances in the deeper sediment. In the seeps, high concentrations of degradation products such as nutrients and sulphide were found, showing them to be the outlets of deep-seated degradation processes. The degradation products appear toxic for bioturbating/ bioirrigating organisms, as a consequence of which, these were absent in the wider seep areas. These two mechanisms driving advection determine oxygen dynamics in these flats, whereas bioirrigation plays a minor role. The deep circulation causes a characteristic distribution of strongly reduced pore water near the low water line and rather more oxidised sediments in the centre of the flats. The two combined transport phenomena determine the fluxes of solutes and gases from the sediment to the surface water and in this way create specific niches for various types of microorganisms.