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By modifying the physical environment, ecosystem engineers can have inordinately large effects on surrounding communities and ecosystem functioning. However, the significance of engineering in ecosystems greatly depends on the physical characteristics of the engineered habitats. Mechanisms underlying such context-dependent impact of engineers remain poorly understood even though they are crucial to establish general predictions concerning the contribution of engineers to ecosystem structure and function. The present study aimed to decrypt such mechanisms by determining how the environmental context modulates the effects of ecosystem engineers (bioturbators) on microorganisms in river sediments. To test the effects of environmental context on the role of bioturbators in sediments, we used mesocosms and recreated two sedimentary contexts in the laboratory by adding a layer of either fine or coarse sand at the top of a gravel-sand matrix. For each sediment context, we examined how the sediment reworking activity of a bioturbating tubificid worm ( Tubifex tubifex) generated changes in the physical (sediment structure and permeability) and abiotic environments (hydraulic discharge, water chemistry) of microorganisms. Microbial characteristics (abundances, activities) and leaf litter decomposition - a major microbially-mediated ecological process - were measured to evaluate the impact of bioturbation on biotic compartment. Our results showed that the permeability, the availability of oxygen and the activities of microorganisms were reduced in sediments covered with fine sand, in comparison with sediments covered with coarse sand. Tubifex tubifex significantly increased permeability (by about six-fold), restored aerobic conditions and ultimately stimulated microbial communities (resulting in a 30% increase in leaf litter breakdown rate) in sediments covered with fine sand. In contrast T. tubifex had low effects in sediments topped by coarse sand, where O2 was already available for hyporheic microorganisms. Our study supports the idea that context dependency mainly modulates the effects of engineering by controlling the ability of engineers to create changes on abiotic (O2 in the present study) factors that are limiting for surrounding communities. [ABSTRACT FROM AUTHOR]

1. A substantial portion of particulate organic matter (POM) is stored in the sediment of rivers and streams. Leaf litter breakdown as an ecosystem process mediated by microorganisms and invertebrates is well documented in surface waters. In contrast, this process and especially the implication for invertebrates in subsurface environments remain poorly studied. 2. In the hyporheic zone, sediment grain size distribution exerts a strong influence on hydrodynamics and habitability for invertebrates. We expected that the influence of shredders on organic matter breakdown in river sediments would be influenced strongly by the physical structure of the interstitial habitat. 3. To test this hypothesis, the influence of gammarids (shredders commonly encountered in the hyporheos) on degradation of buried leaf litter was measured in experimental systems (slow filtration columns). We manipulated the structure of the sedimentary habitat by addition of sand to a gravel-based sediment column to reproduce three conditions of accessible pore volume. Ten gammarids were introduced in columns together with litter bags containing alder leaves at a depth of 8 cm in sediment. Leaves were collected after 28 days to determine leaf mass loss and associated microbial activity (fungal biomass, bacterial abundance and glucosidase, xylosidase and aminopeptidase activities). 4. As predicted, the consumption of buried leaf litter by shredders was strongly influenced by the sediment structure. Effective porosity of 35% and 25% allowed the access to buried leaf litter for gammarids, whereas a lower porosity (12%) did not. As a consequence, leaf litter breakdown rates in columns with 35% and 25% effective porosity were twice as high as in the 12% condition. Microbial activity was poorly stimulated by gammarids, suggesting a low microbial contribution to leaf mass loss and a direct effect of gammarids through feeding activity. 5. Our results show that breakdown of POM in subsurface waters depends on the accessibility of food patches to shredders. [ABSTRACT FROM AUTHOR]