Décomposition de détritus végétaux dans la zone hyporhéique de cours d'eau forestiers : implication du compartiment fongique

Dans les cours d'eau forestiers, la décomposition des litières végétales constitue un processus écologique fondamental. Ce processus sous l'effet des contraintes abiotiques de la zone hyporhéique (ZH) de ces cours d'eau, ainsi que la quantification de la dynamique des invertébrés et des champignons décomposeurs associés ont été étudiés, le tout inséré dans une vision intégrative à l'échelle de l'écosystème. Les résultats suggèrent de réexaminer le fonctionnement écologique de ces cours d'eau, uniquement basé sur la compréhension des relations trophiques et aux flux de carbone liés à la décomposition des litières situé à la surface du sédiment, en y intégrant la dimension verticale conceptualisée par les "hyporhéologistes". Ces résultats conduisent à repenser la dynamique et le rôle des hyphomycètes aquatiques à une échelle plus large que celle de la colonne d'eau uniquement, et mettent en évidence l'importance de la ZH qui à cet égard restait jusqu'à présent en grande partie négligée.In headwater streams, leaf litter decomposition constitutes a key ecosystem-level process. The objective of this thesis was to characterize leaf decomposition in the hyporheic habitat of streams, quantify the associated dynamics of fungal and invertebrate decomposers together with the effect of abiotic factors, and incorporate this process into a stream ecosystem perspective. Taken as a whole, these findings suggest that the functioning of woodland stream ecosystems, largely based on trophic relationships and carbon dynamics related to leaf decomposition occurring at the sediment surface, should be reconsidered with the incorporation of the vertical dimension conceptualized by "hyporheologists". These results lead to think out the dynamics and role of aquatic hyphomycetes at a broader scale than the stream water column alone, and highlight the crucial function of the hyporheic zone, which up to now remained mostly neglected to this regard

Fungal alteration of the elemental composition of leaf litter affects shredder feeding activity

1. Leaf litter from riparian vegetation provides the main source of matter and energy for food webs of small forest streams. Shredding macroinvertebrates mostly feed on this litter when it has been colonised and conditioned by microorganisms, especially by aquatic hyphomycetes. Since shredders feed selectively, they must make foraging decisions based on the physical and chemical characteristics of the food resource, which can change depending on the identity of fungal species. 2. Here, we addressed the effect of changes in fungal assemblage structure on the elemental composi- tion of oak (Quercus robur) leaf litter and how variation in litter quality affects the feeding of a stream shredder. Leaf discs were incubated in microcosms for 2 weeks, inoculated with various fungal assemblages comprised of three species each, and offered to a shredder (Schizopelex festiva, Trichoptera: Sericostomatidae) as food. 3. This shredder ate more leaves with a high mycelial biomass, which depended on fungal assemblage composition. Leaf litter conditioned by different fungal assemblages resulted in different litter N and P concentrations. Mycelial biomass was positively related to litter P concentration, with the lowest and highest P concentrations differing by 40% at most, but not to litter N concentration, even though the lowest and highest N concentrations differed by as much as 35%. The caddisfly larvae ate more leaves with a low C/P ratio. 4. These findings suggest a key role of litter P concentration in eliciting fungal conditioning effects on shredder-mediated litter decomposition

Effect of acidification on leaf litter decomposition in benthic and hyporheic zones of woodland streams

Anthropogenic acidification has deleterious effects on both structure and functioning of surface water ecosystems. This study examined how it may affect the leaf decomposition rate and the community structure and activity of decomposers in both benthic and hyporheic zones of five headwater streams along an acidification gradient from highly acidic (pH 4.6) to circumneutral (pH 7.4). Overall, responses to acidification in hyporheic zones were less pronounced, but followed the same pattern as in their benthic counterparts. Leaf decomposition was much faster in the circumneutral stream, both in the hyporheic and benthic zones (k = 0.0068 and 0.0534 d−1, respectively), than in the most acidic one (k = 0.0016 and 0.0055 d−1, respectively), and correlated well with the acidic gradient in both compartments. Interestingly, leaf litter decomposition was less affected by acidification in hyporheic compared to benthic compartments, likely due to the relatively low sensitivity of fungi, which were the main decomposers of buried coarse particulate organic matter. These results argue in favour of conserving hyporheic habitats in acidified streams as they can maintain matter and species fluxes that are essential to the ecosystem

Litter identity mediates predator impacts on the functioning of an aquatic detritus-based food web

During past decades, several mechanisms such as resource quality and habitat complexity have been proposed to explain variations in the strength of trophic cascades across ecosystems. In detritus-based headwater streams, litter accumulations constitute both a habitat and a resource for detritivorous macroinvertebrates. Because litter edibility (which promotes trophic cascades) is usually inversely correlated with its structural complexity (which weakens trophic cascades), there is a great scope for stronger trophic cascades in litter accumulations that are dominated by easily degradable litter species. However, it remains unclear how mixing contrasting litter species (conferring both habitat complexity and high quality resource) may influence top–down controls on communities and processes. In enclosures exposed in a second-order stream, we manipulated litter species composition by using two contrasting litter (alder and oak), and the presence–absence of a macroinvertebrate predator (Cordulegaster boltonii larvae), enabling it to effectively exert predation pressure, or not, on detritivores (consumptive versus non-consumptive predation effects). Leaf mass loss, detritivore biomass and community structure were mostly controlled independently by litter identity and mixing and by predator consumption. However, the strength of predator control was mediated by litter quality (stronger on alder), and to a lesser extent by litter mixing (weaker on mixed litter). Refractory litter such as oak leaves may contribute to the structural complexity of the habitat for stream macroinvertebrates, allowing the maintenance of detritivore communities even when strong predation pressure occurs. We suggest that considering the interaction between top–down and bottom–up factors is important when investigating their influence on natural communities and ecosystem processes in detritus-based ecosystems

Benthic algae stimulate leaf litter decomposition in detritus-based headwater streams: a case of aquatic priming effect?

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

Allelopathic inhibition of primary producer growth and photosynthesis by aquatic fungi

Autochthonous primary production is generally much reduced in forested headwater streams. Several hypotheses have been proposed for explaining this observation, among them, the low light intensity, or the strong constraints exerted by stream current. Allelopathic inhibition of competitors is a common ecological process in aquatic environments. Aquatic hyphomycetes are known to chemically inhibit bacteria and other fungi (including other aquatic hyphomycetes) but a possible allelopathic effect of aquatic hyphomycetes on primary producers has never been tested. The inhibitory effect of twelve aquatic hyphomycete species was tested on three diatom species. Nine aquatic hyphomycete species exhibited anti-diatom activity. Up to 100% diatom growth inhibition was observed. Our study reveals that such allelopathic interactions might be common in streams and probably involve an array of fungal compounds. We propose that the generally reduced primary production observed in forested headwater streams is, among other factors, due to the inhibition of primary producers by allelopathic compounds released by aquatic hyphomycetes

Rapid characterization of aquatic hyphomycetes by matrix-assisted laser desorption/ionization time-of-flight mass spectrometry

Protein fingerprinting using matrix-assisted laser desorption/ionization time-of-flight mass spec-trometry (MALDI-TOF MS) is a rapid, reliable, and economical method to characterize isolates of terrestrial fungi and other microorganisms. The objective of our study was to evaluate the suitability of MALDI-TOF MS for the identification of aquatic hyphomycetes, a polyphyletic group of fungi that play crucial roles in stream ecosystems. To this end, we used 34 isolates of 21 aquatic hyphomycete species whose identity was confirmed by spore morphology and internal transcribed spacer (ITS1-5.8S-ITS2 = ITS) nuc rDNA sequencing. We tested the efficiency of three protein extraction methods, including chemical and mechanical treatments using 13 different protocols, with the objective of producing high-quality MALDI-TOF mass spectra. In addition to extraction protocols, mycelium age was identified as a key parameter affecting protein extraction efficiency. The dendrogram based on mass-spectrum similarity indicated good and relevant taxonomic discrimination; the tree structure was comparable to that of the phylogram based on ITS sequences. Consequently, MALDI-TOF MS could reliably identify the isolates studied and provided greater taxonomic accuracy than classical morphological methods. MALDI-TOF MS seems suited for rapid characterization and identification of aquatic hyphomycete species

Coarse particulate organic matter in the interstitial zone of three French headwater streams

Headwater woodland streams are primarily heterotrophic: they receive substantial inputs of organic matter from the riparian vegetation, while autochthonous primary production is generally low. A substantial part of leaf litter entering running waters may be buried in the streambed because of flooding and sediment movement. Although the general significance of the hyporheic zone for stream metabolism has been reported early, organic matter storage within the sediment of streams has received less attention, with most studies only quantifying accumulations at the streambed surface and ignoring other stream compartments. In the present study, the amounts of three fractions of coarse particulate organic matter (CPOM; > 16, 4–16 and 1–4 mm) were determined in late autumn and early spring in the interstitial and benthic zones of three head- water streams of the Montagne Noire (South-Western France) differing in their substratum grain size. Our findings demonstrated that the total CPOM content in the interstitial zone can be much (up to one order of magnitude) higher than at the sediment surface. The sandy bottomed stream exhibited a higher amount of CPOM (whatever the size fraction) than the two other streams, suggesting that the sediment particle size may be a major determinant of CPOM storage. Given the large amount of organic matter stored in the interstitial zone, this compartment may play an important role for the carbon turnover and associated trophic dynamics in the stream ecosystem

Beyond the water column: aquatic hyphomycetes outside their preferred habitat

Aquatic hyphomycetes have adapted to running waters by their uncommon conidial shape, which facilitates dispersal as well as adherence to plant substrata. However, they have been early and regularly reported to occur in a variety of environments other than their preferred habitat (e.g., in lentic freshwaters, brackish and marine environments, in terrestrial niches such as stream banks, dew, canopy waters and tree holes). In addition, several aquatic hyphomycetes have adapted to a mutualistic lifestyle which may involve plant defence, as endophytes in leaves, gymnosperm needles, orchids and terrestrial roots. There are several lines of evidence suggesting that aquatic hyphomycetes survive under terrestrial conditions due to their sexual states. Although exhibiting higher diversity in pristine streams, aquatic hyphomycetes can survive environmental stress, e.g., pollution or river intermittency. They also inhabit ground and hyporheic waters, where they appear to be subjected to both physical and physiological selection. Appropriate methods including molecular ones should provide a more comprehensive view of the occurrence and ecological roles of aquatic hyphomycetes outside their preferred habitat

The role of organisms in hyporheic processes : gaps in current knowledge, needs for future research and applications

Fifty years after the hyporheic zone was first defined (Orghidan, 1959), there are still gaps in the knowledge regarding the role of biodiversity in hyporheic processes. First, some methodological questions remained unanswered regarding the interactions between biodiversity and physical processes, both for the study of habitat characteristics and interactions at different scales. Furthermore, many questions remain to be addressed to help inform our understanding of invertebrate community dynamics, especially regarding the trophic niches of organisms, the functional groups present within sediment, and their temporal changes. Understanding microbial community dynamics would require investigations about their relationship with the physical characteristics of the sediment, their diversity, their relationship with metabolic pathways, their inter- actions with invertebrates, and their response to environmental stress. Another fundamental research question is that of the importance of the hyporheic zone in the global metabolism of the river, which must be explored in relation to organic matter recycling, the effects of disturbances, and the degradation of contaminants. Finally, the application of this knowledge requires the development of methods for the estimation of hydro- logical exchanges, especially for the management of sediment clogging, the optimization of self-purification, and the integration of climate change in environmental policies. The development of descriptors of hyporheic zone health and of new metrology is also crucial to include specific targets in water policies for the long-term management of the system and a clear evaluation of restoration strategies