Diel and seasonal variations in abundance, activity, and structure of particle-attached vs free-living bacterial communities were investigated in offshore NW Mediterranean Sea (0-1000 m). Attached bacteria were always less abundant and less diverse but generally more active than free-living bacteria. The most important finding of this study was that the activity of attached bacteria showed pronounced diel variations in the upper mixed water column with higher activities at night. Under mesotrophic conditions, the contribution of attached bacteria to total bacterial activity increased from less than 10% at day time to 83% at night time. At high chlorophyll a concentration, the highest cell-specific activities and contribution to total bacterial activity were due to free-living bacteria at day and to attached bacteria at night. Under summer oligotrophic conditions, free-living bacteria dominated and contributed to the most important part of the bacterial activity at both day and night, whereas attached bacteria were much less abundant but presented the highest cell-specific activities. These diel and seasonal variations in activities were concomitant to changes in bacterial community structure, mainly in the upper layer. The number of attached ribotypes was fairly constant suggesting that particles are colonized by a relatively limited number of ubiquitous ribotypes. Most of these ribotypes were also free-living ribotypes suggesting that attached bacteria probably originate from colonization of newly formed particles by free-living bacteria in the upper layer. These results reinforce the biogeochemical role of attached bacteria in the cycling of particulate organic carbon in the NW Mediterranean Sea and the importance of diel variability in these processes.

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Settling particles were collected from the Ligurian Sea in the northwestern Mediterranean Sea in May 2003 and separated by elutriation into different settling velocity classes (.230, 115-230, 58-115, and ,58 m d 21 ). Particles of the different classes were incubated for 5 d to study their biodegradability. Particulate opal content and organic compound composition (amino acids, pigments, lipids, and carbohydrates) were analyzed initially and at regular time intervals during the incubation period. Most particles (48-67% of total mass) sank at greater than 230 m d 21 and were dominated by large diatom-derived aggregates produced during the spring bloom period. The initial organic composition and the biological lability of these particles varied with settling velocity. The strong phytoplankton signal was visible in all settling velocity classes, while slower settling particles carried with them a greater zooplankton and bacterial signature. As the different class particles decomposed, their compositions changed and became more similar with time, with a dominance of compounds that suggests a more degraded state: the amino acids c-aminobutyric acid and b-alanine, the pigments pyropheophorbide and pheophytin, the deoxysugars fucose and rhamnose, and lipid metabolites (diglycerides and monoglycerides, alcohols, and free fatty acids). Biogenic opal in the particles dissolved faster in more degraded particles than in fresher particles, suggesting that loss of organic matter may expose opal to dissolution. The coupling of settling velocity and decomposition rate measurements shows quantitatively that slower settling particles are quickly degraded and AcknowledgmentsThis research was part of the MedFlux and PECHE (Production and Export of Carbon: Control by Heterotrophs at small temporal scale) programs and was supported by the U.S. National Science Foundation Chemical Oceanography Program (OCE-0136370, OCE-0136318, and OCE-0113687) and the French CNRS (Centre National de la Recherche Scientifique), respectively. Participation of B.M. was funded by ORFOIS (Origin and Fate of Biogetic Particle Fluxes in the ocean) (EVK2-CT2001-00100). We thank Michael Peterson, Lynn Abramson, Jenni Szlosek, Meaghan Askea, and Isabell Putnam for shipboard and laboratory help; David Hirschberg and Michael Peterson for CHN analysis; Claude Mante for help with statistical data treatment; and the captain and crew of the RV Seward Johnson II. We wish to acknowledge the associate editor and two anonymous reviewers for very helpful comments and suggestions on the manuscript. This is MedFlux contribution 7 and MSRC contribution 1318.

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