This paper contrasts the natural and anthropogenic controls on the conversion of unreactive N2 to more reactive forms of nitrogen (Nr). A variety of data sets are used to construct global N budgets for 1860 and the early 1990s and to make projections for the global N budget in 2050. Regional N budgets for Asia, North America, and other major regions for the early 1990s, as well as the marine N budget, are presented to highlight the dominant fluxes of nitrogen in each region. Important findings are that human activities increasingly dominate the N budget at the global and at most regional scales, the terrestrial and open ocean N budgets are essentially dis- connected, and the fixed forms of N are accumulating in most environmental reservoirs. The largest uncertainties in our understanding of the N budget at most scales are the rates of natural biological nitrogen fixation, the amount of Nr storage in most environmental reservoirs, and the production rates of N2 by denitrification.

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Nitrogen cycles: past, present, and future

Although algal blooms, including those considered toxic or harmful, can be natural phenomena, the nature of the global problem of harmful algal blooms (HABs) has expanded both in extent and its public perception over the last several decades. Of concern, especially for resource managers, is the potential relationship between HABs and the accelerated eutrophication of coastal waters from human activities. We address current insights into the relationships between HABs and eutrophication, focusing on sources of nutrients, known effects of nutrient loading and reduction, new understanding of pathways of nutrient acquisition among HAB species, and relationships between nutrients and toxic algae. Through specific, regional, and global examples of these various relationships, we offer both an assessment of the state of understanding, and the uncertainties that require future research efforts. The sources of nutrients poten- tially stimulating algal blooms include sewage, atmospheric deposition, groundwater flow, as well as agricultural and aquaculture runoff and discharge. On a global basis, strong correlations have been demonstrated between total phos- phorus inputs and phytoplankton production in freshwaters, and between total nitrogen input and phytoplankton pro- duction in estuarine and marine waters. There are also numerous examples in geographic regions ranging from the largest and second largest U.S. mainland estuaries (Chesapeake Bay and the Albemarle-Pamlico Estuarine System), to the Inland Sea of Japan, the Black Sea, and Chinese coastal waters, where increases in nutrient loading have been linked with the development of large biomass blooms, leading to anoxia and even toxic or harmful impacts on fisheries re- sources, ecosystems, and human health or recreation. Many of these regions have witnessed reductions in phytoplankton biomass (as chlorophyll a) or HAB incidence when nutrient controls were put in place. Shifts in species composition have often been attributed to changes in nutrient supply ratios, primarily N:P or N:Si. Recently this concept has been extended to include organic forms of nutrients, and an elevation in the ratio of dissolved organic carbon to dissolved organic nitrogen (DOC:DON) has been observed during several recent blooms. The physiological strategies by which different groups of species acquire their nutrients have become better understood, and alternate modes of nutrition such as heterotrophy and mixotrophy are now recognized as common among HAB species. Despite our increased un- derstanding of the pathways by which nutrients are delivered to ecosystems and the pathways by which they are assimilated differentially by different groups of species, the relationships between nutrient delivery and the development of blooms and their potential toxicity or harmfulness remain poorly understood. Many factors such as algal species presence/ abundance, degree of flushing or water exchange, weather conditions, and presence and abundance of grazers contribute to the success of a given species at a given point in time. Similar nutrient loads do not have the same impact in different environments or in the same environment at different points in time. Eutrophication is one of several mechanisms by which harmful algae appear to be increasing in extent and duration in many locations. Although important, it is not the only explanation for blooms or toxic outbreaks. Nutrient enrichment has been strongly linked to stimulation of some harmful species, but for others it has not been an apparent contributing factor. The overall effect of nutrient over- enrichment on harmful algal species is clearly species specific.

/pdf/harmful-algal-blooms-and-eutrophication-nutrient-sources-4ujytujxjq.pdf

Harmful Algal Blooms and Eutrophication: Nutrient Sources, Composition, and Consequences

If stretched end to end, the estimated 1030viruses in the oceans would span farther than the nearest 60 galaxies. This reservoir of genetic and biological diversity continues to yield exciting discoveries and, in this Review, Curtis A. Suttle highlights the areas that are likely to be of greatest interest in the next few years. Viruses are by far the most abundant 'lifeforms' in the oceans and are the reservoir of most of the genetic diversity in the sea. The estimated 1030 viruses in the ocean, if stretched end to end, would span farther than the nearest 60 galaxies. Every second, approximately 1023 viral infections occur in the ocean. These infections are a major source of mortality, and cause disease in a range of organisms, from shrimp to whales. As a result, viruses influence the composition of marine communities and are a major force behind biogeochemical cycles. Each infection has the potential to introduce new genetic information into an organism or progeny virus, thereby driving the evolution of both host and viral assemblages. Probing this vast reservoir of genetic and biological diversity continues to yield exciting discoveries.

Marine viruses — major players in the global ecosystem

Viruses are the most common biological agents in the sea, typically numbering ten billion per litre. They probably infect all organisms, can undergo rapid decay and replenishment, and influence many biogeochemical and ecological processes, including nutrient cycling, system respiration, particle size-distributions and sinking rates, bacterial and algal biodiversity and species distributions, algal bloom control, dimethyl sulphide formation and genetic transfer. Newly developed fluorescence and molecular techniques leave the field poised to make significant advances towards evaluating and quantifying such effects.

/pdf/marine-viruses-and-their-biogeochemical-and-ecological-1lkd7mtgjc.pdf

Marine viruses and their biogeochemical and ecological effects

The discovery that viruses may be the most abundant organisms in natural waters, surpassing the number of bacteria by an order of magnitude, has inspired a resurgence of interest in viruses in the aquatic environment. Surprisingly little was known of the interaction of viruses and their hosts in nature. In the decade since the reports of extraordinarily large virus populations were published, enumeration of viruses in aquatic environments has demonstrated that the virioplankton are dynamic components of the plankton, changing dramatically in number with geographical location and season. The evidence to date suggests that virioplankton communities are composed principally of bacteriophages and, to a lesser extent, eukaryotic algal viruses. The influence of viral infection and lysis on bacterial and phytoplankton host communities was measurable after new methods were developed and prior knowledge of bacteriophage biology was incorporated into concepts of parasite and host community interactions. The new methods have yielded data showing that viral infection can have a significant impact on bacteria and unicellular algae populations and supporting the hypothesis that viruses play a significant role in microbial food webs. Besides predation limiting bacteria and phytoplankton populations, the specific nature of virus-host interaction raises the intriguing possibility that viral infection influences the structure and diversity of aquatic microbial communities. Novel applications of molecular genetic techniques have provided good evidence that viral infection can significantly influence the composition and diversity of aquatic microbial communities.

/pdf/virioplankton-viruses-in-aquatic-ecosystems-30m49tuwrn.pdf

Virioplankton: viruses in aquatic ecosystems

Most of the oceanic reservoir of dissolved organic matter (DOM) is of marine origin and is resistant to microbial oxidation, but little is known about the mechanisms of its formation. In a laboratory study, natural assemblages of marine bacteria rapidly (in <48 hours) utilized labile compounds (glucose, glutamate) and produced refractory DOM that persisted for more than a year. Only 10 to 15% of the bacterially derived DOM was identified as hydrolyzable amino acids and sugars, a feature consistent with marine DOM. These results suggest that microbial processes alter the molecular structure of DOM, making it resistant to further degradation and thereby preserving fixed carbon in the ocean.

https://science.sciencemag.org/content/sci/292/5518/917.full.pdf

Production of Refractory Dissolved Organic Matter by Bacteria

Sedimentary habitats cover most of the ocean bottom and therefore constitute the largest. single ecosystem on earth in spatial coverage, Although only a small fraction of the micro-, meio- and macroscopic benthic organisms that reside in and on sediments have been described and few estimates of total species numbers and biogeographic pattern have been attempted, there is sufficient information on a few species to suggest that sedimentary organisms significantly impact major ecological processes. Benthic organisms contribute to regulation of carbon, nitrogen, and sulfur cycling, water column processes, pollutant distribution and fate, secondary production, and transport, and stability of sediments. Linkages between groups of organisms and the level of functional redundancy is poorly known, however, there is probably substantial redundancy within groups. There is little evidence that biodiversity per se is necessary for benthic systems to contribute to ecosystem services. but because linkages are so poorly known and predictive knowledge confined to a few species, it is not presently possible to predict exactly how species loss will impact these services and ecosystem health. Thus, a precautionary approach of "assume the worst" is advised, and every effort should be made to curtail the species and genetic diversity loss resulting from fishing, pollution, habitat destruction, introduction of non-native (exotic) species, and global warming. Concurrently, scientists must take advantage of exciting, rapidly evolving technology and a rejuvenated interest in biodiversity to provide more concrete and thorough information on benthos and ecosystem processes. [KEYWORDS: Species-diversity; sea; pollution; richness; mesocosm; samples]

The Importance of marine sediment biodiversity in ecosystem processes

Simultaneous determinations of nitrogen gas production, ammonia, and particulate organic nitrogen formation in the coastal sediments of Mangoku-Ura, Simoda Bay, and Tokyo Bay were made by using the 15N-label tracer method. The rate of nitrogen gas production in the sediment surface layer was about 10−2 μg atom of N per g per h, irrespective of the location of the sediments examined. [15N]ammonia and -particulate organic nitrogen accounted for 20 to 70% of the three products, and after several hours of incubation, the major fraction of nondenitrified 15N in Mangoku-Ura and Simoda Bay sediments was recovered as ammonia. In Tokyo Bay sediments, particulate organic nitrogen was produced at a greater rate than was ammonia. The reduction rate data suggest that the pathway of nitrate reduction to ammonia is important in coastal sediments.

Denitrification and ammonia formation in anaerobic coastal sediments.

Abundance of bacteria and tiny DNA-associated particles in the upper layer of Japanese coastal and offshore waters was evaluated by epifluorescence microscopy with 0.015-mum-pore-size Nuclepore filters. The number of tiny DNA-associated particles was compared with the abundance of virus particles estimated by transmission electron microscopy. Although a large variation in virus abundance (1.2 x 10 to 35 x 10 ml) was obtained with the transmission electron microscopy method, the ratio of 4',6-diamidino-2-phenylindole-reactive tiny particles to viruses was in a rather narrow range (1.0 to 1.6), indicating that the majority of the tiny DNA-associated particles identified by epifluorescence microscopy were actually virus particles. This result implies the possibility of using epifluorescence microscopy for the evaluation of virus abundance in marine environments.

Abundance of viruses in marine waters: assessment by epifluorescence and transmission electron microscopy.

Denitrification and nitrification in sediments of Tama Estuary and Odawa Bay, Japan, were investigated by the combined use of a continuous-flow sediment-water system and a 15N tracer technique At Odawa Bay, the nitrification rate was comparable to the nitrate reduction rate, and 70% of the N2 evolved originated from nitrogenous oxides (nitrate and nitrite) which were produced by the action of nitrifying bacteria in the sediments At Tama Estuary, the nitrate reduction rate was 11 to 17 times higher than the nitrification rate, and nitrogenous oxides derived from ammonium accounted for only 6 to 9% of the N2 evolution by denitrification

Isao Koike

Papers

Production of Refractory Dissolved Organic Matter by Bacteria

The Importance of marine sediment biodiversity in ecosystem processes

Denitrification and ammonia formation in anaerobic coastal sediments.

Abundance of viruses in marine waters: assessment by epifluorescence and transmission electron microscopy.

Estimates of Denitrification and Nitrification in Coastal and Estuarine Sediments