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Aquatic macroaggregates (flocs ≥ 0.5 mm) provide an important mechanism for vertical flux of nutrients and organic matter in aquatic ecosystems, yet their role in the transport and fate of zoonotic pathogens is largely unknown. Terrestrial pathogens that enter coastal waters through contaminated freshwater runoff may be especially prone to flocculation due to fluid dynamics and electrochemical changes that occur where fresh and marine waters mix. In this study, laboratory experiments were conducted to evaluate whether zoonotic pathogens (Cryptosporidium, Giardia, Salmonella) and a virus surrogate (PP7) are associated with aquatic macroaggregates and whether pathogen aggregation is enhanced in saline waters. Targeted microorganisms showed increased association with macroaggregates in estuarine and marine waters, as compared with an ultrapure water control and natural freshwater. Enrichment factor estimations demonstrated that pathogens are 2-4 orders of magnitude more concentrated in aggregates than in the estuarine and marine water surrounding the aggregates. Pathogen incorporation into aquatic macroaggregates may influence their transmission to susceptible hosts through settling and subsequent accumulation in zones where aggregation is greatest, as well as via enhanced uptake by invertebrates that serve as prey for marine animals or as seafood for humans.

Harmful algal blooms (HABs) are common in Monterey Bay, CA, and have resulted in repeated closures of shellfish fisheries and the poisoning and death of marine mammals. In the majority of instances, HAB events in this region are first detected by the presence of sick or dying animals. The phrase “cryptic blooms” was adopted to denote the appearance of poisoning at higher trophic levels with no prior evidence of a large phytoplankton bloom. We hypothesize that the onset of many HAB events goes undetected because the bloom is initially concentrated in discrete thin subsurface layers in the water column that are easily missed by conventional sampling and monitoring methods. In this paper, we report on the detection and monitoring of a subsurface layer of phytoplankton in northern Monterey Bay, CA, using a high-resolution, autonomous profiler. This ‘thin layer,’ which measured from 10 cm to 3 m in thickness (85%

Within Monterey Bay, California, USA, the food web transfer of domoic acid (DA), a neurotoxin produced by diatoms of the genus Pseudo-nitzschia, has led to major mortality events of marine mammals and birds. Less visible, and less well known, is whether invertebrates and fish associated with the benthos are also affected by blooms of DA-producing Pseudo-nitzschia spp. This study examines the presence of DA in benthic flatfish offshore of Davenport, California, (37°0′36″N, 122°13′12″W) and within Monterey Bay, California (36°45′0″N, 122°1′48″W), including species that feed primarily in the sediment (benthic-feeding) and species that feed primarily in the water column (benthopelagic-feeding). Flatfish caught between 10 December 2002 and 17 November 2003 at depths of 30–180 m had concentrations of DA in the viscera ranging from 3 to 26 μg DA g−1 of viscera. Although the DA values reported are relatively low, benthic-feeding flatfish were frequently contaminated with DA, especially as compared with the frequency of contamination of flatfish species that feed in the water column. Furthermore, on days in which both benthic-feeding and benthopelagic-feeding flatfish were collected, the former had significantly higher concentrations of DA in the viscera. Curlfin turbot, Pleuronicthys decurrens, the flatfish with both the highest level and frequency of DA contamination, are reported to feed exclusively on polychaetes, suggesting that these invertebrates may be an important vector of the toxin in benthic communities and may pose a risk to other benthic-feeding organisms.

In at least two mass mortality events in Monterey Bay, California, planktivorous fish were implicated as vectors of the neurotoxin domoic acid (DA) from diatoms to sea birds and marine mammals. Whereas the transfer of DA from planktivorous fish to piscivorous predators has been well established, the relationship between toxin levels in plankton-feeding fish and the regional abundance of DA-producing diatoms has not been documented. Here we present data from an extensive field study in which cell densities of toxic Pseudo-nitzschia species and DA levels in anchovies and sardines were measured from samples collected weekly throughout Monterey Bay from 8 October 1999 to 18 October 2000. Four distinct blooms were documented with cell densities ranging from 3.2×103 to 5.0×105 cells l–1. DA was detected in fish viscera samples whenever toxic diatom densities reached ≥103 cells l–1 in surface waters, suggesting that anchovies and sardines regularly consume toxic diatoms when present. Fish contained DA levels above the regulatory limit (20 µg DA g–1 whole fish) only when toxic cell densities exceeded 104 cells l–1. DA was only detected in fish when toxic diatom species were also present in the water, suggesting that the toxin is quickly depurated and that fish are only dangerous vectors during the bloom period. Anchovies appear to be more potent vectors than sardines as they consistently contained more DA than sardines collected simultaneously. Maximum DA levels detected in fish were 1,815 µg DA g–1 in anchovy and 728 µg DA g–1 in sardine viscera samples. In fish with high viscera levels of DA, corresponding body tissues contained 0.2–2.2 µg DA g–1 (0.2±0.1% of the viscera level), suggesting that DA is not accumulated in edible body tissues to levels that threaten human consumers. Results from this study suggest that anchovies may be a valuable indicator species for assessing the risk of DA intoxication to piscivorous sea birds and mammals during toxic Pseudo-nitzschia blooms in Monterey Bay.

The planktivorous northern anchovy is a prominent vector of the phycotoxin domoic acid (DA) to organisms at higher trophic levels, including fish-eating seabirds and mammals. Although there are abundant data reporting DA-induced excitotoxic symptoms in higher vertebrates, to date there has been no reported evidence of neurotoxic effects in lower vertebrate vectors such as fish. To explain this apparent lack of toxicity, it has been suggested that DA may not reach the brain in anchovies and/or that fish are not as sensitive neurologically to DA. In the present study, intracoelomic (IC) injection of DA, at doses ranging from 1 to 14 μg DA g−1 total fish weight, resulted in severe neurotoxic symptoms such as spinning, disorientation, inability to school, and mortality, indicating that anchovies are neurologically susceptible and that DA crosses the blood–brain barrier in fish. An ED50 of 3.2 μg DA g−1 total body weight was determined via IC injection of DA in 83 anchovies. Comparable intraperitoneal injection studies with mice, rats, and monkeys report similar DA-induced neurotoxic symptoms at doses near 3.2 μg DA g−1, suggesting a similar neurologic sensitivity and mechanism of toxicity between anchovies and mammals. DA tissue distribution measurements from freshly collected field-exposed anchovies and orally gavaged anchovies indicate that DA uptake from the gastrointestinal tract does occur. Levels as high as 1,175 μg DA g−1 were measured in anchovy viscera, while muscle and brain tissue DA levels were 3 orders of magnitude lower, indicating low but measurable DA uptake. Further evidence is needed to confirm that uptake is sufficient during field events to induce symptoms in anchovies. Our work provides the first reported evidence of neurotoxic symptoms in fish and suggests that anchovies may be affected by DA during toxic diatom blooms. If sufficient uptake occurs, DA-induced neurotoxic symptoms and mortality may make fish easier prey targets, thereby selecting for the highest toxin levels transferred, as well as providing a possible pathway for the transfer of DA to benthic communities.

The planktonic ciliateStrombidium capitatum (Leegaard, 1915) Kahl, 1932 retains functional chloroplasts derived from ingested algal cells. Chloroplast replacement and aging were experimentally investigated in cultured ciliates provided with a cryptophyte (Pyrenomonas salina), a prymnesiophyte (Isochrysis galbana) and a prasinophyte (Pyramimonas sp.) as sources of plastids. All three algae were ingested and chloroplasts from all were retained by the ciliate. Within 15 min of exposure to the cryptophyte, this alga was taken up by the ciliates. Initially, most of the cryptophyte chloroplasts were in intact algal cells in ciliate vacuoles. By 2 h, cryptophyte plastids were commonly found free in the ciliate cytoplasm. WhenS. capitatum was switched from a diet containing cryptophytes to a non-cryptophyte diet, most cryptophyte chloroplasts were diluted out of the ciliates by cell division and/or replaced by non-cryptophyte chloroplasts within 9 h. When the ciliates are not provided with algae, they decrease in size and number. However, the starving ciliate cells contain some chloroplasts for as long as they live (40 h or more). Under these conditions, cryptophyte chloroplasts persist longer than the other chloroplast types. Our observations suggest that chloroplast retention times inS. capitatum depend on the type of chloroplast as well as the availability of phytoplankton containing suitable new chloroplasts, and probably also on the physiological states of the ingested algae and the ciliates. It is interesting that we were not able to grow this ciliate when we provided it only with prey that lacked chloroplasts.

The fecal pellets of zooplankton are thought to be major carriers of organic matter from surface to deeper waters of the oceans. As the pellets descend, they release soluble components, partially due to breakdown by associated microorganisms. Previous laboratory work of other investigators has suggested that the surface of a fecal pellet rapidly acquires bacteria, which increase in abundance until they and their protozoan consumers disrupt the pellet membrane, spilling contents into the water. In contrast, our field collections of fecal pellets from free-floating particle interceptor traps (from the “Vertex” project off Central California in 1980 and off Mexico in 1981), suggest that microbial decomposition probably is initiated in the sea from inside the fecal pellets. Transmission and scanning electron microscopy indicate that bacterial populations are most abundant in the interior of fecal pellets obtained from the sea, but that the same pellets will acquire the surface bacterial lawn typically observed in laboratory studies if maintained aboard ship. If the fecal pellets are decomposed from the inside, then the principal agents are enteric bacteria or ingested, digestion-resistant bacteria, or both. Such bacteria may differ metabolically from those that colonize the fecal pellet surfaces. Further-more, the abundance of healthy-appearing bacteria inside the pellets suggests that their metabolic activities may produce microhabitats of reduced oxygen tension that could differ considerably from that of the pellet exteriors. Decomposition in these semi-enclosed microenvironments may proceed in a manner not yet predicted by models that attribute decomposition to well-aerated, surface-dwelling bacterial populations on fecal pellets in the sea.

Sinking rates were determined for fecal pellets produced by gelatinous zooplankton (salps, Salpa fusiformis and Pegea socia; pteropods, Corolla spectabilis; and doliolids, Dolioletta gegenbaurii) feeding in surface waters of the California Current. Pellets from the salps and pteropods sank at rates up to 2 700 and 1 800 m d-1, respectively; such speeds exceed any yet recorded for zooplankton fecal pellets. Fecal pellets of salps were rich in organic material, with C:N ratios from 5.4 to 6.2, close to values for living plankton. The relation between volume and sinking rate indicates that salp and pteropod pellets are slightly less dense than those of pelagic Crustacea; moreover, pellet density varied between different collection dates, probably because of differences in composition. In contrast, doliolid pellets sank at rates up to 208 m d-1, a rate much lower than would be expected from pellet size. Thus, density and sinking rates of pellets are much more variable in zooplankton than would be expected from studies of crustaceans alone. Moreover, the extraordinarily high sinking rates of fecal pellets of salps indicates that these tunicates may be disproportionately important in the flux of biogenic materials during periods when they form dense population blooms.

Abundance and production rates were measured on freefloating mats composed of the diatoms Rhizosolenia castracanei and R. imbricata var. shrubsolei in the California Current and boundary waters of the central North Pacific during October, 1980. Mats ranged from 3.0 to 10.6 cm in maximum length and had a mean volume of 6.4 ml. Production rates of the diatom mats averaged 4.0 μg C colony-1 h-1, more than 103 times higher than that of an equal volume of surrounding water. However, because of their low density at these sites, about 1 mat m-3, diatom mats contributed only about 1% of the total primary production. The two large Rhizosolenia species comprised almost all of the phytoplankton biomass within the mats. Rhizosolenia species rarely occurred in seawater between mats, where the phytoplankton community was dominated by a diverse array of nannoplankton. The Rhizosolenia species in the mats appeared to be in healthy condition and contained intracellular bacteria. The very high production rates of the colonies indicate high nutrient demand and, since these particular diatoms lack the symbiotic cyanobacterium Richelia intracellularis, which is implicated in N-fixation in other species of Rhizosolenia, our results indicate that other nutrient sources must be present. We discuss the potential role of the intracellular bacteria in nitrogen fixation.

Salps (mainly Salpa fusiformis and, to a lesser extent, Pegea socia) and a web-building pteropod (Corolla spectabilis) were studied in epipelagic waters of the central California Current. Although both kinds of gelatinous zooplankton trap phytoplankton in a mucus net, a fecal pellet analysis indicated that their diet differs significantly when they feed together, probably because of differences both in the pore sizes of their nets and in their feeding methods. Salps have a finemesh filter, on which they can retain even the smallest phytoplankton; thus, when small coccolithophores are abundant, as they were in our study, salp feces contain such cells and the coccoliths derived from them. In contrast, pteropods feeding in the same area produce fecal pellets consisting chiefly of larger phytoplankton, especially diatoms. Since fecal pellets transport most biogenic material to the deep sea, changes in herbivore species composition at a given geographic location can change the chemistry of materials entering deep water; at our study site, the more salps, the greater the calcite flux, and, the more pteropods, the greater the silica flux. In addition, fecal pellets of both salps and pteropods include partially digested residues of phytoplankton that appear as olive-green spheres, having an ultrastructure identical with that of the socalled olive-green “cells.” Presumably, fecal pellets, after sinking into deep water, ultimately disintegrate. releasing both the viable phytoplankton and the olive-green spheres into aphotic waters. Thus the feces of epipelagic herbivores are likely sources of much of the flora of the deep ocean.

The planktonic ciliateLaboea strobila Lohmann sequesters photosynthetically functional chloroplasts derived from ingested algae. The chloroplasts lie free in the cytoplasm and are most abundant just under the pellicle of the ciliate. The maximum rate of photosynthesis (Pmax) was 925 pg C ciliate-1h-1 (3.7 pg C pg chl.a-1h-1). At saturating irradiance, the amount of carbon fixed h-1 equaled 12.6% of the body carbon of the ciliate. To grow,L. strobila requires both light and algal food. In the absence of food, survival ofL. strobila is significantly longer in the light than in the dark. Based on ingestion rate and photosynthetic rate, we calculate that photosynthesis can make an important contribution to this ciliate's carbon budget even when algal food is plentiful.

Siliceous choanoflagellates and previously undescribed, siliceous cyst-like spheroids (in the size range 2.5 to 15 μm) were found below sea-ice in the Weddell Sea and studied with scanning electron microscopy. Cell counts from water samples obtained over a 10,000 km area indicated that both spheroids and choanoflagellates were abundant in the upper 100 m of the water column, averaging about 105 cells l-1. The large numbers of phagotrophic choanoflagellates suggest that bacterioplankton-picoplankton food chains are important and may indicate the presence of longer, more complex trophic networks than previously considered in Antarctic waters. Circumstantial evidence suggests that the spheroids may be the cysts of choanoflagellates; if so, the production of siliceous cysts by members of this enigmatic taxon may be of considerable interest in discussions of the phylogenetic position of the group in relation to the algae and the metazoans. Similar cysts were found simultaneously in the North Pacific Ocean (see preceding paper: Booth et al. 1980), and recently we have discovered the cysts in the upwelling area of the eastern tropical Pacific Ocean. The occurrence of the cyst in such geographically distant localities suggests that they have a worldwide distribution. Since the minute cysts are easily overlooked on the light microscope, their recognition and further study undoubtedly require the electron microscopic examination of plankton samples.

In the past decade, protozoa have been recognized as ubiquitous residents of the upper waters of the sea, where they have key roles in detrital and producer food webs1–4. In contrast, protozoa are almost unknown from aphotic depths, where food chains presumably are supported by detritus. We show here that ciliate protozoans do accompany sinking detritus to bathypelagic depths. In the productive northeastern Pacific Ocean, the biomass of ciliates on particles settling between the surface and 2,000 m far exceeds the biomass of bacteria, which are the only widely recognized detrital microorganisms in deep pelagic communities. Although absolute numbers of ciliates decline below the euphotic zone, their concentrations (number per g detritus) drop only slightly by 2,000 m. Taxonomic data and patterns of abundance5 indicate that the ciliates are endemics and not remnants of surface colonists, and that the deep sea is a favourable environment for them. Their high concentrations, possibly together with other smaller protozoa, indicate that protists may control bacterial mineralization rates on sinking particles, thereby regulating the vertical distribution of many biologically active elements in the sea.

Diatoms, radiolarians, silicoflagellates, ebridians and archaeomonads are the primary groups of pelagic organisms in siliceous marine deposits, with only the archaemonads regarded as being absent from modern oceans and sediments. They remain a poorly known group of siliceous nanofossils in Cretaceous to Upper Tertiary marine sediments. We present here the first report, from any modern oceanic environment, of pelagic cysts that conform to the descriptions for Archaeomonadaceae. These cysts occurred in seawater, sea ice and marine-surface sediments, demonstrating the group is still extant. Furthermore, their distribution in modern sediment suggests they may be useful guide fossils for sea-ice environments as they occur abundantly only in polar or near-polar water where sea ice is present. Few other microorganisms seem to have such a close and exclusive association with sea ice, thus archaeomonads deserve increased attention as indicators of ice conditions for deep-water marine environments.