Rodrigo Almeda

Changes in the C, N, and P cycles by the predicted salps-krill shift in the southern ocean

The metabolic carbon requirements and excretion rates of three major zooplankton groups in the Southern Ocean were studied in February 2009. The research was conducted in the framework of the ATOS research project as part of the Spanish contribution to the International Polar Year. The objective was to ascertain the possible consequences of the predicted zooplankton shift from krill to salps in the Southern Ocean for the cycling of biogenic carbon and the concentration and stoichiometry of dissolved inorganic nutrients. The carbon respiratory demands and NH4-N and PO4-P excretion rates of < 5 mm size copepods, krill and salps were estimated by incubation experiments. The carbon-specific metabolic rates and N:P metabolic quotients of salps were higher than those of krill (furcilia spp. and adults) and copepods, and as expected there was a significant negative relation between average individual zooplankton biomass and their metabolic rates, each metabolic process showing a particular response that lead to different metabolic N:P ratios. The predicted change from krill to salps in the Southern Ocean would encompass not only the substitution of a pivotal group for Antarctic food webs (krill) by one with an indifferent trophic role (salps). In a zooplankton community dominated by salps the respiratory carbon demand by zooplankton will significantly increase, and therefore the proportion of primary production that should be allocated to compensate for the global respiratory C-losses of zooplankton. At the same time, the higher production by salps of larger, faster sinking fecal pellets will increase the sequestration rate of biogenic carbon. Similarly, the higher N and P excretion rates of zooplankton and the changes in the N:P stoichiometry of the metabolic products will modify the concentration and proportion of N and P in the nutrient pool, inducing quantitative and qualitative changes on primary producers that will translate to the whole Southern Ocean ecosystem.

Toxicity of dispersant Corexit 9500A and crude oil to marine microzooplankton

In 2010, nearly 7 million liters of chemical dispersants, mainly Corexit 9500A, were released in the Gulf of Mexico to treat the Deepwater Horizon oil spill. However, little is still known about the effects of Corexit 9500A and dispersed crude oil on microzooplankton despite the important roles of these planktonic organisms in marine ecosystems. We conducted laboratory experiments to determine the acute toxicity of Corexit 9500A, and physically and chemically dispersed Louisiana light sweet crude oil to marine microzooplankton (oligotrich ciliates, tintinnids and heterotrophic dinoflagellates). Our results indicate that Corexit 9500A is highly toxic to microzooplankton, particularly to small ciliates, and that the combination of dispersant with crude oil significantly increases the toxicity of crude oil to microzooplankton. The negative impact of crude oil and dispersant on microzooplankton may disrupt the transfer of energy from lower to higher trophic levels and change the structure and dynamics of marine planktonic communities.

Effects of Crude Oil Exposure on Bioaccumulation of Polycyclic Aromatic Hydrocarbons and Survival of Adult and Larval Stages of Gelatinous Zooplankton

Gelatinous zooplankton play an important role in marine food webs both as major consumers of metazooplankton and as prey of apex predators (e.g., tuna, sunfish, sea turtles). However, little is known about the effects of crude oil spills on these important components of planktonic communities. We determined the effects of Louisiana light sweet crude oil exposure on survival and bioaccumulation of polycyclic aromatic hydrocarbons (PAHs) in adult stages of the scyphozoans Pelagia noctiluca and Aurelia aurita and the ctenophore Mnemiopsis leidyi, and on survival of ephyra larvae of A. aurita and cydippid larvae of M. leidyi, in the laboratory. Adult P. noctiluca showed 100% mortality at oil concentration ≥20 µL L−1 after 16 h. In contrast, low or non-lethal effects were observed on adult stages of A. aurita and M. leidyi exposed at oil concentration ≤25 µL L−1 after 6 days. Survival of ephyra and cydippid larva decreased with increasing crude oil concentration and exposition time. The median lethal concentration (LC50) for ephyra larvae ranged from 14.41 to 0.15 µL L−1 after 1 and 3 days, respectively. LC50 for cydippid larvae ranged from 14.52 to 8.94 µL L−1 after 3 and 6 days, respectively. We observed selective bioaccumulation of chrysene, phenanthrene and pyrene in A. aurita and chrysene, pyrene, benzo[a]pyrene, benzo[b]fluoranthene, benzo[k]fluoranthene, and benzo[a]anthracene in M. leidyi. Overall, our results indicate that (1) A. aurita and M. leidyi adults had a high tolerance to crude oil exposure compared to other zooplankton, whereas P. noctiluca was highly sensitive to crude oil, (2) larval stages of gelatinous zooplankton were more sensitive to crude oil than adult stages, and (3) some of the most toxic PAHs of crude oil can be bioaccumulated in gelatinous zooplankton and potentially be transferred up the food web and contaminate apex predators.

Dispersant Corexit 9500A and chemically dispersed crude oil decreases the growth rates of meroplanktonic barnacle nauplii (Amphibalanus improvisus) and tornaria larvae (Schizocardium sp.)

Our knowledge of the lethal and sublethal effects of dispersants and dispersed crude oil on meroplanktonic larvae is limited despite the importance of planktonic larval stages in the life cycle of benthic invertebrates. We determined the effects of Light Louisiana Sweet crude oil, dispersant Corexit 9500A, and dispersant-treated crude oil on the survival and growth rates of nauplii of the barnacle Amphibalanus improvisus and tornaria larvae of the enteropneust Schizocardium sp. Growth rates of barnacle nauplii and tornaria larvae were significantly reduced after exposure to chemically dispersed crude oil and dispersant Corexit 9500A at concentrations commonly found in the water column after dispersant application in crude oil spills. We also found that barnacle nauplii ingested dispersed crude oil, which may have important consequences for the biotransfer of petroleum hydrocarbons through coastal pelagic food webs after a crude oil spill. Therefore, application of chemical dispersants increases the impact of crude oil spills on meroplanktonic larvae, which may affect recruitment and population dynamics of marine benthic invertebrates.

Swimming behavior and prey retention of the polychaete larvae Polydora ciliata (Johnston)

SUMMARY The behavior of the ubiquitous estuarine planktotrophic spionid polychaete larvae Polydora ciliata was studied. We describe ontogenetic changes in morphology, swimming speed and feeding rates and have developed a simple swimming model using low Reynolds number hydrodynamics. In the model we assumed that the ciliary swimming apparatus is primarily composed of the prototroch and secondarily by the telotroch. The model predicted swimming speeds and feeding rates that corresponded well with the measured speeds and rates. Applying empirical data to the model, we were able to explain the profound decrease in specific feeding rates and the observed increase in the difference between upward and downward swimming speeds with larval size. We estimated a critical larval length above which the buoyancy-corrected weight of the larva exceeds the propulsion force generated by the ciliary swimming apparatus and thus forces the larva to the bottom. This modeled critical larval length corresponded to approximately 1 mm, at which, according to the literature, competence for metamorphosis and no more length increase is observed. These findings may have general implications for all planktivorous polychaete larvae that feed without trailing threads. We observed bell shaped particle retention spectra with a minimum prey size of approximately 4 μm equivalent spherical diameter, and we found that an ontogenetic increase in maximum prey size add to a reduction in intra-specific food competition in the various larval stages. In a grazing experiment using natural seawater, ciliates were cleared approximately 50% more efficiently than similar sized dinoflagellates. The prey sizes retainable for P. ciliata larvae covers the microplankton fraction and includes non-motile as well as motile prey items, which is why the larvae are trophically positioned among the copepods and dinoflagellates. Not only do larval morphology and behavior govern larval feeding, prey behavior also influences the feeding efficiency of Polydora ciliata.

Novel insight into the role of heterotrophic dinoflagellates in the fate of crude oil in the sea

Although planktonic protozoans are likely to interact with dispersed crude oil after a spill, protozoan-mediated processes affecting crude oil pollution in the sea are still not well known. Here, we present the first evidence of ingestion and defecation of physically or chemically dispersed crude oil droplets (1–86 μm in diameter) by heterotrophic dinoflagellates, major components of marine planktonic food webs. At a crude oil concentration commonly found after an oil spill (1 μL L−1), the heterotrophic dinoflagellates Noctiluca scintillans and Gyrodinium spirale grew and ingested ~0.37 μg-oil μg-Cdino−1 d−1, which could represent ~17% to 100% of dispersed oil in surface waters when heterotrophic dinoflagellates are abundant or bloom. Egestion of faecal pellets containing crude oil by heterotrophic dinoflagellates could contribute to the sinking and flux of toxic petroleum hydrocarbons in coastal waters. Our study indicates that crude oil ingestion by heterotrophic dinoflagellates is a noteworthy route by which petroleum enters marine food webs and a previously overlooked biological process influencing the fate of crude oil in the sea after spills.

Influence of UVB radiation on the lethal and sublethal toxicity of dispersed crude oil to planktonic copepod nauplii

Toxic effects of petroleum to marine zooplankton have been generally investigated using dissolved petroleum hydrocarbons and in the absence of sunlight. In this study, we determined the influence of natural ultraviolet B (UVB) radiation on the lethal and sublethal toxicity of dispersed crude oil to naupliar stages of the planktonic copepods Acartia tonsa, Temora turbinata and Pseudodiaptomus pelagicus. Low concentrations of dispersed crude oil (1 μL L(-1)) caused a significant reduction in survival, growth and swimming activity of copepod nauplii after 48 h of exposure. UVB radiation increased toxicity of dispersed crude oil by 1.3-3.8 times, depending on the experiment and measured variables. Ingestion of crude oil droplets may increase photoenhanced toxicity of crude oil to copepod nauplii by enhancing photosensitization. Photoenhanced sublethal toxicity was significantly higher when T. turbinata nauplii were exposed to dispersant-treated oil than crude oil alone, suggesting that chemical dispersion of crude oil may promote photoenhanced toxicity to marine zooplankton. Our results demonstrate that acute exposure to concentrations of dispersed crude oil and dispersant (Corexit 9500) commonly found in the sea after oil spills are highly toxic to copepod nauplii and that natural levels of UVB radiation substantially increase the toxicity of crude oil to these planktonic organisms. Overall, this study emphasizes the importance of considering sunlight in petroleum toxicological studies and models to better estimate the impact of crude oil spills on marine zooplankton.

Gender-specific feeding rates in planktonic copepods with different feeding behavior

Planktonic copepods have sexually dimorphic behaviors, which can cause differences in feeding efficiency between genders. Copepod feeding rates have been studied extensively but most studies have focused only on females. In this study, we experimentally quantified feeding rates of males and females in copepods with different feeding behavior: ambush feeding (Oithona nana), feeding-current feeding (Temora longicornis) and cruising feeding (Centropages hamatus). We hypothesize that carbon-specific maximum ingestion rates are similar between genders, but that maximum clearance rates are lower for male copepods, particularly in ambush feeders, where the males must sacrifice feeding for mate searching. We conducted gender-specific functional feeding response experiments using prey of different size and motility. In most cases, gender-specific maximum ingestion and clearance rates were largely explained by the difference in size between sexes, independent of the feeding strategy. However, maximum clearance rates of males were approximately two times higher than for females in the ambush feeding copepod O. nana feeding on an optimal motile prey (Oxyrrhis marina), as hypothesized. We conclude that the conflict between mate searching and feeding can cause significant difference in feeding efficiency between copepod genders in ambush feeders but not in feeding-current and cruising feeders.

How much crude oil can zooplankton ingest? Estimating the quantity of dispersed crude oil defecated by planktonic copepods

We investigated and quantified defecation rates of crude oil by 3 species of marine planktonic copepods (Temora turbinata, Acartia tonsa, and Parvocalanus crassirostris) and a natural copepod assemblage after exposure to mechanically or chemically dispersed crude oil. Between 88 and 100% of the analyzed fecal pellets from three species of copepods and a natural copepod assemblage exposed for 48 h to physically or chemically dispersed light crude oil contained crude oil droplets. Crude oil droplets inside fecal pellets were smaller (median diameter: 2.4-3.5 μm) than droplets in the physically and chemically dispersed oil emulsions (median diameter: 6.6 and 8.0 μm, respectively). This suggests that copepods can reject large crude oil droplets or that crude oil droplets are broken into smaller oil droplets before or during ingestion. Depending on the species and experimental treatments, crude oil defecation rates ranged from 5.3 to 245 ng-oil copepod(-1) d(-1), which represent a mean weight-specific defecation rate of 0.026 μg-oil μg-Ccopepod(1) d(-1). Considering a dispersed crude oil concentration commonly found in the water column after oil spills (1 μl L(-1)) and copepod abundances in high productive coastal areas, copepods may defecate ∼ 1.3-2.6 mg-oil m(-3) d(-1), which would represent ∼ 0.15%-0.30% of the total dispersed oil per day. Our results indicate that ingestion and subsequent defecation of crude oil by planktonic copepods has a small influence on the overall mass of oil spills in the short term, but may be quantitatively important in the flux of oil from surface water to sediments and in the transfer of low-solubility, toxic petroleum hydrocarbons into food webs after crude oil spills in the sea.

Resting eggs in free living marine and estuarine copepods

Marine free living copepods can survive harsh periods and cope with seasonal fluctuations in environmental conditions using resting eggs (embryonic dormancy). Laboratory experiments show that temperature is the common driver for resting egg production. Hence, we hypothesize (i) that seasonal temperature variation, rather than variation in food abundance is the main driver for the occurrence of the resting eggs strategy in marine and estuarine copepod species; and (ii) that the thermal boundaries of the distribution determine where resting eggs are produced and whether they are produced to cope with warm or cold periods. We compile literature information on the occurrence of resting egg production and relate this to spatio-temporal patterns in sea surface temperature and chlorophyll a concentration obtained from satellite observations. We find that the production of resting eggs has been reported for 42 species of marine free living copepods. Resting eggs are reported in areas with high seasonal variation in sea surface temperature (median range 11°C). Temporal variation in chlorophyll a concentrations, however, seems of less importance. Resting eggs are commonly produced to cope with both warm and cold periods and, depending on the species, they are produced at the upper or lower thermal boundaries of a species’distribution.