In detritus-based ecosystems, autochthonous primary production contributes very little to the detritus pool. Yet primary producers may still influence the functioning of these ecosystems through complex interactions with decomposers and detritivores. Recent studies have suggested that, in aquatic systems, small amounts of labile carbon (C) (e.g., producer exudates), could increase the mineralization of more recalcitrant organic-matter pools (e.g., leaf litter). This process, called priming effect, should be exacerbated under low-nutrient conditions and may alter the nature of interactions among microbial groups, from competition under low-nutrient conditions to indirect mutualism under high-nutrient conditions. Theoretical models further predict that primary producers may be competitively excluded when allochthonous C sources enter an ecosystem. In this study, the effects of a benthic diatom on aquatic hyphomycetes, bacteria, and leaf litter decomposition were investigated under two nutrient levels in a factorial microcosm experiment simulating detritus-based, headwater stream ecosystems. Contrary to theoretical expectations, diatoms and decomposers were able to coexist under both nutrient conditions. Under low-nutrient conditions, diatoms increased leaf litter decomposition rate by 20% compared to treatments where they were absent. No effect was observed under high-nutrient conditions. The increase in leaf litter mineralization rate induced a positive feedback on diatom densities. We attribute these results to the priming effect of labile C exudates from primary producers. The presence of diatoms in combination with fungal decomposers also promoted decomposer diversity and, under low-nutrient conditions, led to a significant decrease in leaf litter C:P ratio that could improve secondary production. Results from our microcosm experiment suggest new mechanisms by which primary producers may influence organic matter dynamics even in ecosystems where autochthonous primary production is low.

1. Leaf litter constitutes the major source of organic matter and energy in woodland stream ecosystems. A substantial part of leaf litter entering running waters may be buried in the streambed as a consequence of flooding and sediment movement. While decomposition of leaf litter in surface waters is relatively well understood, its fate when incorporated into river sediments, as well as the involvement of invertebrate and fungal decomposers in such conditions, remain poorly documented. 2. We tested experimentally the hypotheses that the small interstices of the sediment restrict the access of the largest shredders to buried organic matter without compromising that of aquatic hyphomycetes and that fungal decomposers in the hyporheic zone, at least partly, compensate for the role of invertebrate detritivores in the benthic zone. 3. Alder leaves were introduced in a stream either buried in the sediment (hyporheic), buried after 2 weeks of exposure at the sediment surface (benthic-hyporheic), or exposed at the sediment surface for the entire experiment (benthic). Leaf decomposition was markedly faster on the streambed surface than in the two other treatments (2.1-and 2.8-fold faster than in the benthic-hyporheic and hyporheic treatments, respectively). 4. Fungal assemblages were generally less diverse in the hyporheic habitat with a few species tending to be relatively favoured by such conditions. Both fungal biomass and sporulation rates were reduced in the hyporheic treatment, with the leaves subject to the benthic-hyporheic treatment exhibiting an intermediate pattern. The initial 2-week stage in the benthic habitat shaped the fungal assemblages, even for leaves later subjected to the hyporheic conditions. 5. The abundance and biomass of shredders drastically decreased with burial, except for Leuctra spp., which increased and was by far the most common leaf-associated taxon in the hyporheic zone. Leuctra spp. was one of the rare shredder taxa displaying morphological characteristics that increased performance within the limited space of sediment interstices. 6. The carbon budgets indicated that the relative contributions of the two main decomposers, shredders and fungi, varied considerably depending on the location within the streambed. While the shredder biomass represented almost 50% of the initial carbon transformed after 80 days in the benthic treatment, its contribution was <0.3% in the hyporheic one and 2.0% in the combined benthic-hyporheic treatment. In contrast, Correspondence: Julien Cornut, Laboratoire d'écologie fonctionnelle, 2541 mycelial and conidial production in the permanently hyporheic environment accounted for 12% of leaf mass loss, i.e. 2-3 times more than in the two other conditions. These results suggest that the role of fungi is particularly important in the hyporheic zone. 7. Our findings indicate that burial within the substratum reduces the litter breakdown rate by limiting the access of both invertebrate and fungal decomposers to leaves. As a consequence, the hyporheic zone may be an imp...