Organic detritus passing from the sea surface through the water column to the sea floor controls nutrient regeneration, fuels benthic life and affects burial of organic carbon in the sediment record1–3. Particle trap systems have enabled the first quantification of this important process. The results suggest that the dominant mechanism of vertical transport is by rapid settling of rare large particles, most likely of faecal pellets or marine snow of the order of >200 μm in diameter, whereas the more frequent small particles have an insignificant role in vertical mass flux4–6. The ultimate source of organic detritus is biological production in surface waters of the oceans. I determine here an empirical relationship that predicts organic carbon flux at any depth in the oceans below the base of the euphotic zone as a function of the mean net primary production rate at the surface and depth-dependent consumption. Such a relationship aids in estimating rates of decay of organic matter in the water column, benthic and water column respiration of oxygen in the deep sea and burial of organic carbon in the sediment record.

Particulate organic carbon flux in the oceans—surface productivity and oxygen utilization

Characteristics, dynamics and significance of marine snow

Heterotrophic bacteria perform two major functions in the transformation of organic matter: They produce new bacterial biomass (bacterial secondary production [BP]), and they respire organic C to inorganic C (bacterial respiration [BR]). For planktonic bacteria, a great deal has been learned about BP and its regulation during the past several decades but far less has been learned about BR. Our lack of knowledge about BR limits our ability to understand the role of bacteria in the carbon cycle of aquatic ecosystems. Bacterial growth efficiency (BGE) is the amount of new bacterial biomass produced per unit of organic C substrate assimilated and is a way to relate BP and BR: BGED (BP)=(BPC BR). Estimates of BGE for natural planktonic bacteria range from <0.05 to as high as 0.6, but little is known about what might regulate this enormous range. In this paper we review the physiological and ecological bases of the regulation of BGE. Further, we assemble the literature of the past 30 years for which both BP and BR were measured in natural planktonic ecosystems and explore the relationship between BGE and BP. Although the relationship is variable, BGE varies systematically with BP and the trophic richness of the ecosystem. In the most dilute, oligotrophic systems, BGE is as low as 0.01; in the most eutrophic systems, it plateaus near 0.5. Planktonic bacteria appear to maximize carbon utilization rather than BGE. A consequence of this strategy is that maintenance energy costs (and therefore maintenance respiration) seems to be highest in oligotrophic systems.

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Bacterial growth efficiency in natural aquatic systems

The redox dye 5-cyano-2,3-ditolyl tetrazolium chloride (CTC) was employed for direct epifluorescent microscopic enumeration of respiring bacteria in environmental samples. Oxidized CTC is nearly colorless and is nonfluorescent; however, the compound is readily reduced via electron transport activity to fluorescent, insoluble CTC-formazan, which accumulates intracellularly. Bacteria containing CTC-formazan were visualized by epifluorescence microscopy in wet-mount preparations, on polycarbonate membrane filter surfaces, or in biofilms associated with optically opaque surfaces. Counterstaining of CTC-treated samples with the DNA-specific fluorochrome 4',6-diamidino-2-phenylindole allowed enumeration of active and total bacterial subpopulations within the same preparation. Municipal wastewater, groundwater, and seawater samples supplied with exogenous nutrients yielded CTC counts that were generally lower than total 4',6-diamidino-2-phenylindole counts but typically equal to or greater than standard heterotrophic (aerobic) plate counts. In unsupplemented water samples, CTC counts were typically lower than those obtained with the heterotrophic plate count method. Reduction of CTC by planktonic or biofilm-associated bacteria was suppressed by formaldehyde, presumably because of inhibition of electron transport activity and other metabolic processes. Because of their bright red fluorescence (emission maximum, 602 nm), actively respiring bacteria were readily distinguishable from abiotic particles and other background substances, which typically fluoresced at shorter wavelengths. The use of CTC greatly facilitated microscopic detection and enumeration of metabolically active (i.e., respiring) bacteria in environmental samples.

Use of a fluorescent redox probe for direct visualization of actively respiring bacteria.

Bacteria and algae are the numerically dominant organisms in the plankton of lakes and the ocean and their metabolism largely controls pelagic energy flow and nutrient cycling. The purpose of this review is to examine the means by which algae and heterotrophic bacteria interact in planktonic systems. In terrestrial systems, bacterial-plant interactions have been well studied in the rhizosphere, the zone in the soil where microorganisms are influenced by plant roots (108). The concept of the "phycosphere" was introduced as a pelagic analogy to the rhizosphere (7). The phycosphere would be the zone surrounding an algal cell within which microorganisms are influenced by algal products. In considering bacterial-algal interactions, then, we should ask ourselves whether a phycosphere exists. I cite specific evidence from the literature for various types of interactions. In doing this I rely heavily on laboratory studies because relatively little field oriented work is available. I include neither the photosynthetic bacteria (e.g. Chlorobiaceae, Rhodospirillaceae) nor the large macroalgae (seaweeds) in this review. The cyanobacteria have both algal and bacterial characteristics, being prokaryotes and capable of oxygenic photosynthesis. As planktonic, aerobic photo-autotrophs the cyanobacteria clearly belong with the phytoplankton in terms of this review.

Interactions between bacteria and algae in aquatic ecosystems

The rates of sedimentation of particulate matter were measured at a station in the Eckernforde Bight, Baltic Sea, using sediment traps. Different parameters (C, N, chlorophyll a, phaeopigments, adenosine triphosphate (ATP), heterotrophic activity (glucose maximum uptake velocity, vmax), biological oxygen demand (BOD), and electron-transport-system (ETS) activity) were measured during a 1-year period to evalnate the composition, vertical distribution, and activity of microorganisms associated with sedimenting particulate matter. Measurements indicated that the organic fractions of particulate sedimenting matter were transformed considerably during sedimentation. The seasonal fluctuations showed higher heterotrophic activity durang summer and lower activity during winter. Laboratory experiments also indicated that an important part of planktonic particulate material was susceptible to mineralization within a short period of time: phytoplankton was mineralized by about 35% and “zooplankton” by about 18% per day, at 20°C; at 5°C mineralization was considerably lower (about 3 and 8%, respectively).

Bacterial activity related to sedimenting particulate matter

The natural heterotrophic utilization of released and particulate photoassimilated compounds by natural phytoplankton populations was followed during the summer of 1975 in the Kiel Bight (Western Baltic Sea). Phytoplankton exudates (2 to 21% of primary production) seem to represent an important substrate for heterotrophic bacteria: uptake rates between 8 and 17,5%/h were observed. Samples incubated with antibiotics indicated that the heterotrophic uptake of phytoplankton exudates during photosynthesis could lead to underestimation of the release rate. Production of 14CO
2was used as an estimate of the mineralization of particulate photoassimilated matter. The mineralization rates measured were in the range of 6 to 20% after 20 h. In order to confirm the quantitative measurements, micro-autoradiographic examinations of the samples were performed.

R. Iturriaga

Papers

Bacterial activity related to sedimenting particulate matter

Observations of heterotrophic activity on photoassimilated organic matter