Laurence P. Madin

Zooplankton vertical migration and the active transport of dissolved organic and inorganic carbon in the Sargasso Sea

The least known component of the “biological pump” is the active transport of carbon and nutrients by diel vertical migration of zooplankton. We measured CO2 respiration and dissolved organic carbon (DOC) excretion by individual species of common vertically migrating zooplankton at the US JGOFS Bermuda Atlantic Time-series Study (BATS) station. The inclusion of DOC excretion in this study builds on published research on active transport by respiration of inorganic carbon and allows a direct assessment of the role of zooplankton in the production of dissolved organic matter used in midwater microbial processes. On average, excretion of DOC makes up 24% (range=5–42%) of the total C metabolized (excreted+respired) and could represent a significant augmentation to the vertical flux that has already been documented for respiratory CO2 flux by migrant zooplankton. Migratory fluxes were compared to other transport processes at BATS. Estimates of combined active transport of CO2 and DOC by migrators at BATS averaged 7.8% and reached 38.6% of mean sinking POC flux at 150 m, and reached 71.4% of mean sinking POC flux at 300 m. DOC export by migrator excretion averaged 1.9% and reached 13.3% of annual DOC export by physical mixing at this site. During most of the year when deep mixing does not occur, diel migration by zooplankton could provide a supply of DOC to the deeper layers that is available for use by the microbial community. A carbon budget comparing migrant zooplankton transport to the balance of fluxes in the 300–600 m depth strata at BATS shows on average that the total migrant flux supplies 37% of the organic carbon remineralized in this layer, and that migrant DOC flux is more than 3 times the DOC flux gradient by diapycnal mixing. New estimates of active transport of both organic and inorganic carbon by migrants may help resolve observed imbalances in the C budget at BATS, but the magnitude is highly dependent on the biomass of the migrating community.

The contribution of microorganisms to particulate carbon and nitrogen in surface waters of the Sargasso Sea near Bermuda

Abstract Seawater samples were collected from the euphotic zone of the Sargasso Sea near Bermuda in August of 1989 and March–April of 1990. Microbial population abundances, chlorophyll concentration, particulate carbon and particulate nitrogen were measured. Calculations were performed to establish the relative and absolute importance of the various microbial assemblages. The choice of conversion factors (g C and N cell −1 , or g C and N μm −3 ) for the microbial populations dramatically affected the estimation of “living” and “detrital” particulate material in the samples, and the relative importance of the various microbial groups. Averaged over all samples on either of the two cruises, microbial biomass constituted a greater proportion of the total particulate carbon and nitrogen during March–April (55% and 63%, respectively), than during August (≈24% and 30%, respectively) using “constrained” conversion factors that were derived. Accordingly, detrital material constituted the bulk of the particulate material during August, but was similar to the amount of microbial biomass during March–April. The bacterial assemblage constituted the largest single pool of microbial carbon (35%) and nitrogen (45%) in the water, and a significant fraction of the total particulate carbon (≈10–20%) and nitrogen (≈15–30%). Phototrophic nanoplankton (microalgae 2–20 μm in size) were second in overall biomass, and often dominated the microbial biomass in the deep chlorophyll maxima that were present during both cruises. The results temper recent assertions concerning the overwhelming importance of bacterial biomass in the oligotrophic Sargasso Sea but still support a major role for these microorganisms in the open ocean as repositories for carbon and nutrients.

Recurrent jellyfish blooms are a consequence of global oscillations

A perceived recent increase in global jellyfish abundance has been portrayed as a symptom of degraded oceans. This perception is based primarily on a few case studies and anecdotal evidence, but a formal analysis of global temporal trends in jellyfish populations has been missing. Here, we analyze all available long-term datasets on changes in jellyfish abundance across multiple coastal stations, using linear and logistic mixed models and effect-size analysis to show that there is no robust evidence for a global increase in jellyfish. Although there has been a small linear increase in jellyfish since the 1970s, this trend was unsubstantiated by effect-size analysis that showed no difference in the proportion of increasing vs. decreasing jellyfish populations over all time periods examined. Rather, the strongest nonrandom trend indicated jellyfish populations undergo larger, worldwide oscillations with an approximate 20-y periodicity, including a rising phase during the 1990s that contributed to the perception of a global increase in jellyfish abundance. Sustained monitoring is required over the next decade to elucidate with statistical confidence whether the weak increasing linear trend in jellyfish after 1970 is an actual shift in the baseline or part of an oscillation. Irrespective of the nature of increase, given the potential damage posed by jellyfish blooms to fisheries, tourism, and other human industries, our findings foretell recurrent phases of rise and fall in jellyfish populations that society should be prepared to face.

Heterotrophic Bacteria and Bacterivorous Protozoa in Oceanic Macroaggregates

Oceanic macroaggregates (marine snow and Rhizosolenia mats) sampled from the Sargasso Sea are associated with bacterial and protozoan populations up to four orders of magnitude greater than those present in samples from the surrounding water. Filamentous, curved, and spiral bacteria constituted a higher proportion of the bacteria associated with the particles than were found among bacteria in the surrounding water. Protozoan populations were dominated numerically by heterotrophic microflagellates, but ciliates and amoebas were also observed. Macroaggregates are highly enriched heterotrophic microenvironments in the oceans and may be significant for the cycling of particulate organic matter in planktonic food chains.

Pelagic Tunicates: Unique Herbivores in the Marine Plankton

The pelagic tunicates including salps, doliolids, and appendicularians exhibit higher feeding rates, greater retention of bacteria-sized food, and much shorter generation times than other marine planktonic herbivores. Their many unique adaptations enable them to reach high population densities and react with maxiumum flexibility to the unpredictable and patchy environment of the pelagic ecosystem. (Accepted for publication 2 March 1982)

Questioning the Rise of Gelatinous Zooplankton in the World's Oceans

During the past several decades, high numbers of gelatinous Zooplankton species have been reported in many estuarine and coastal ecosystems. Coupled with media-driven public perception, a paradigm has evolved in which the global ocean ecosystems are thought to he heading toward being dominated by “nuisance” jellyfish. We question this current paradigm by presenting a broad overview of gelatinous Zooplankton in a historical context to develop the hypothesis that population changes reflect the human-mediated alteration of global ocean ecosystems. To this end, we synthesize information related to the evolutionary context of contemporary gelatinous Zooplankton blooms, the human frame of reference for changes in gelatinous Zooplankton populations, and whether sufficient data are available to have established the paradigm. We conclude that the current paradigm in which it is believed that there has been a global increase in gelatinous Zooplankton is unsubstantiated, and we develop a strategy for addressing the critical questions about long-term, human-related changes in the sea as they relate to gelatinous Zooplankton blooms.

Diel vertical migration bySalpa aspera and its potential for large-scale particulate organic matter transport to the deep-sea

In mid-summer 1975 throughout the Western Slope Water of the North Atlantic Ocean, massive numbers ofSalpa aspera performed a diel vertical migration of at least 800 m. This resulted in a movement of 85 to 90% of the total zooplankton biomass out of the upper 500 m during the day. Fecal pellet production and losses from this salp population were estimated to contribute approximately 12 mg C m-2 day-1 to the deep planktonic and benthic populations. If all this organic matter reached the deep-sea floor, it would represent over 100% of the daily deep-sea benthic infauna energy requirements.

Is global ocean sprawl a cause of jellyfish blooms

Jellyfish (Cnidaria, Scyphozoa) blooms appear to be increasing in both intensity and frequency in many coastal areas worldwide, due to multiple hypothesized anthropogenic stressors. Here, we propose that the proliferation of artificial structures – associated with (1) the exponential growth in shipping, aquaculture, and other coastal industries, and (2) coastal protection (collectively, “ocean sprawl”) – provides habitat for jellyfish polyps and may be an important driver of the global increase in jellyfish blooms. However, the habitat of the benthic polyps that commonly result in coastal jellyfish blooms has remained elusive, limiting our understanding of the drivers of these blooms. Support for the hypothesized role of ocean sprawl in promoting jellyfish blooms is provided by observations and experimental evidence demonstrating that jellyfish larvae settle in large numbers on artificial structures in coastal waters and develop into dense concentrations of jellyfish-producing polyps.

On the natural history and distribution of oceanic ctenophores

Abstract Oceanic ctenophores show aa great diversity in feeding biology, ranging from Leucothea multicornis , which can eat only small copepods, to species of Ocyropsis , which can capture and ingest small fish and euphausiids. The Cydippida capture prey with tentacles, while most of the Lobata use tentacles and their mucus-covered oral lobes. Leucothea multicornis has elaborate oral lobes, which can function independently of one another; Ocyropsis crystallina feeds by trapping prey in its muscular oral lobes, without use of mucus or tentacles. These two species represent extremes in the feeding biology of the Lobata. The Cestida feed by trapping small prey on tentacles that lie over the sides of the body. Several genera of hyperiid amphipods are closely associated with ctenophores, both as parasites and predators. Medusae and heteropods have also been seen to feed on ctenophores. We have encountered oceanic ctenophores often in abundance on about 75% of the 250 dives we have made in the past three years. Our collections from the North Atlantic and Indian oceans suggest that ctenophores are important predators in the upper waters of the open sea. Because most of the species discussed in this paper are never reported from net plankton collections, they constitute a hitherto ignored component of the open ocean ecosystem.

Production, composition and sedimentation of salp fecal pellets in oceanic waters

Rates of fecal pellet production have been recorded from seven species of oceanic salps feeding on natural diets. Expressed as μg C defecated per mg salp body C per hour, the values range between 3.7 and 27.7. Carbon: nitrogen ratios of the salp fecal pellets average 11.4; the organic matter of the pellets is mainly protein and carbohydrate. Sinking velocities of the pellets are very high, ranging from 320 to 2 238 m d-1 for pellets from three species. However, the pellets sink slower than would be predicted from extrapolation of rates for crustacean pellets, probably due to the shape of the pellets and their density. The high rates of defecation, large size and rapid sedimentation of salp fecal pellets make them likely mechanisms for rapid transport of small particulate matter from surface waters to deep water and the benthos.