We tested the hypothesis that in the euphotic zone of oligotrophic oceanic waters, the growth-limiting nutrients for bacterioplankton vary over a diel cycle due to competition with phytoplankton, leading to distinct diel patterns in bacterioplankton. During a cruise across the subtropical Atlantic Ocean, a distinct diel periodicity in bacterioplankton was detectable, with highest bacterial abundance in the early morning and declining over the day. A peak in the frequency of dividing cells (FDC, 20 to 25% of the total bacterial community) was recorded at midnight; FDC rates were consistently low during the day and cell volume increased over the day until the early night period. No diel variations in these parameters were detectable in the layers below the deep chlorophyll maximum (DCM). Bio-assay experiments with freshly collected, N+P amended water were started in the early morning. No significant bacterial growth was detectable in the early morning while, in the bio-assays that started at noon, significant bacterial growth was detectable in the N+P treatments. P amendment alone also caused an increase in bacterial abundance although to a lesser extent. Thus, we have evidence that the bacterioplankton in the euphotic layer of the subtropical Atlantic are not limited by the availability of N+P in the morning (most likely limited in C) but at noon when they are competing with phytoplankton for the inorganic N+P sources. Based on the concomitantly measured decline in inorganic N+P in the bio-assays, we estimated the C:N:P ratio of newly produced bacteria. The mean atomic C:N:P ratio for the bacteria in the N+P amended bio-assay experiments started in the early morning was C:N:P = 15:13:1 while the corresponding ratio for bacteria in the bio-assays started at noon was C:N:P = 118:11:1. This diel variation in the C:N:P ratio was caused by the 1 order of magnitude higher bacterial C production at noon as compared to the early morning. Thus, excess uptake of N+P occurs during the morning and luxury C uptake during noon. This plasticity in the C:N:P ratio of bacterioplankton might be a strategy to overcome, at least partly, periods of shortage in a specific nutrient species in the oligotrophic subtropical Atlantic.
MARINE ECOLOGY PROGRESS SERIES
The significance of the tidal zone as nursery area for the brown shrimp Crangon crangon became evident during a study on density and size-distribution of predator populations on a 50 km2 tidal flat in the western Wadden Sea (The Netherlands). A clear picture of the nursery system and its function in the annual cycle of brown shrimp is obtained when nursery population-data are combined with experimental growth rates in juvenile C. crangon and their dependence on temperature and body size. Shrimp brood settles in the area during February-August with main arrival during April, May and June; it leaves the tidal zone, at a body length of ca. 30 mm, during July-September. At the end of the year the majority of the new shrimp generation reaches maturity, after which winter spawning leads to settlement of new brood in spring again. The annual cycle thus obtained fits well in the seasonal pattern of commercial shrimp catches in the area. The predominance of winter spawning and the low effect of summer-egg production is explained by the seasonal temperature cycle and its effect on the development of the juveniles at our latitude, as well as by a specific seasonal pattern of predation pressure on shrimp brood in the tidal zone. Spawning cycles in C. crangon along other parts of the European continental coast are discussed
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