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A 48-h larval development toxicity test using the marine polychaete Galeolaria caespitosa Lamarck (Fam. Serpulidae).

Assessing the risk of effluents and other anthropogenic inputs to the receiving environment is ultimately best done on a site-specific basis, which often requires toxicity tests using organisms relevant to that environment. Additionally, the test species or life stage needs to be available for a reasonable portion of the year to allow temporal fluctuations to be assessed. A 48-h larval development toxicity test using the marine polychaete Galeolaria caespitosa was developed. This test was developed as G. caespitosa releases viable gametes year-round, and the test species is environmentally relevant to the marine system receiving the liquid effluent being evaluated. Toxicity tests were conducted using G. caespitosa from different locations and evaluating the gamete response to copper. All population responses were comparable, with EC50 values ranging from 16 to 40 microg/L copper (as CuCl2, x 2H2O). Toxicity tests were also conducted using G. caespitosa gametes with an effluent produced by a lead smelting operation. The response of the G. caespitosa test with this effluent was compared with three other test methods using two microalgal species, Isochrysis sp. and Nitzschia closterium, and gametes from the mussel Mytilus edulis. The G. caespitosa larval development toxicity test was the most sensitive test to the effluent, with EC50 values ranging from 1-23% effluent, while it ranked second in sensitivity to copper. This test could be applied to other common serpulids worldwide.

Animals↗

Development of marine toxicity data for ordnance compounds.

A toxicity database for ordnance compounds was generated using eight compounds of concern and marine toxicity tests with five species from different phyla. Toxicity tests and endpoints included fertilization success and embryological development with the sea urchin Arbacia punctulata; zoospore germination, germling length, and cell number with the green macroalga Ulva fasciata; survival and reproductive success of the polychaete Dinophilus gyrociliatus; larvae hatching and survival with the redfish Sciaenops ocellatus; and survival of juveniles of the opossum shrimp Americamysis bahia (formerly Mysidopsis bahia). The studied ordnance compounds were 2,4- and 2,6-dinitrotoluene, 2,4,6-trinitrotoluene, 1,3-dinitrobenzene, 1,3,5-trinitrobenzene, 2,4,6-trinitrophenylmethylnitramine (tetryl), 2,4,6-trinitrophenol (picric acid), and hexahydro-1,3,5-trinitro-1,3,5-triazine (RDX). The most sensitive toxicity test endpoints overall were the macroalga zoospore germination and the polychaete reproduction tests. The most toxic ordnance compounds overall were tetryl and 1,3,5-trinitrobenzene. These were also the most degradable compounds, often being reduced to very low or below-detection levels at the end of the test exposure. Among the dinitro- and trinitrotoluenes and benzenes, toxicity tended to increase with the level of nitrogenation. Picric acid and RDX were the least toxic chemicals tested overall.

Animals↗

Effects of organophosphate and carbamate pesticides on acetylcholinesterase and choline acetyltransferase activities of the polychaete Nereis diversicolor.

A toxicity test for organophosphates (OP) and carbamates (C) was improved with the adult ragworm Nereis diversicolor. Animals were maintained in U-shaped glass tubes of 4-mm inner diameter fixed vertically on a plastic plate and placed in glass aquaria. Each tank was covered with glass in order to reduce evaporation and heat dissipation. Temperature varied between 15 and 16 degrees C and salinity was constant (34 per thousand) during the entire length of the experiment. Experiments were performed with a fixed day length of 12 h and seawater was gently aerated. The maintenance system allowed the administration of OP and C compounds via the seawater. An acclimatization period of 48 h was not sufficient to accomodate worms to their artificial burrows; accordingly, we chose to acclimate worms for a week before beginning the exposure. Choline acetyltransferase (ChAT) activity was very low and was not significantly modified by two OP compounds: malathion and parathion-ethyl. ChAT is not a target for these pesticides and should not be used for future studies about OP and C toxicity. On the other hand, inhibitory effects on acetylcholinesterase (AChE) activity were determined at concentrations of 10(-6) M for three OP compounds-malathion, parathion-ethyl, and phosalone-and a carbamate pesticide-carbaryl. We measured only short-term effects and no cumulative effect was determined, the maximum percentage of AChE activity inhibition being between 2 (carbaryl) and 7 (OP compounds) days after exposure and then remaining stable. Mortality occured only after a period of intoxication of 14 days. N diversicolor, which can be easily maintained at the laboratory, seems to be a good candidate for future laboratory studies to test the toxicity of other pollutants.

Animals↗

Comparative bioaccumulation of chlorinated hydrocarbons from sediment by two infaunal invertebrates.

Bioaccumulation of chlorinated hydrocarbons (CHs) from field-contaminated sediments by two infaunal invertebrates, Rhepoxynius abronius (a non-deposit feeding amphipod) and Armandia brevis (a nonselective, deposit-feeding polychaete), was examined and species responses were compared. Sediments were selected over a large geographical area of the Hudson-Raritan estuary to assess the potential for bioaccumulation from a typical urban estuary. Unlike polycyclic aromatic hydrocarbons (PAHs) from these sediments, concentrations of CHs in interstitial water (IW) indicated that partition coefficients (Koc) were generally as expected, especially when based on predicted, nonsorbed, interstitial water CH concentrations (IWfree). Correlations between amphipod and polychaete tissue residues revealed that these species were responding similarly to a gradient of CH concentrations in sediment. While tissue residues and BAFloc (lipid/organic carbon normalized bioaccumulation factor) values for the trichlorobiphenyls were similar for both species, accumulation in the polychaete was three to 10 times higher for the more hydrophobic PCBs, which was attributed to differences in the route of exposure. A negative correlation between the bioaccumulation factor (BAF) and total organic carbon (TOC) was found for both species, which was expected according to equilibrium partitioning theory. Because it was assumed that the amphipod was not feeding in these tests and the polychaete was ingesting sediment, comparison of their tissue residues and bioaccumulation factors was useful for highlighting the importance of sediment ingestion, especially for short-term, nonequilibrium exposures. These results may also help elucidate the limitations associated with assessing bioaccumulation and the resultant toxic response in standard 10-day toxicity tests with similar invertebrates.

Animals↗

Polychlorinated biphenyl- and mercury-associated alterations on benthic invertebrate community structure in a contaminated salt marsh in southeast Georgia.

The community structure of a benthic macroinvertebrate assemblage in a contaminated salt marsh was evaluated as part of an ecological characterization of a former chloralkali production facility in Georgia. Sample locations were chosen based on a gradient of the primary contaminants of concern, total mercury and polychlorinated biphenyls (PCBs), primarily Aroclor 1268. Sediment concentrations of Aroclor 1268 ranged from 2.3 to 150 mg/kg dry weight, while mercury concentrations ranged from 15 to 170 mg/kg dry weight in the study area. Mercury and PCBs were determined to be co-located in the sediments. Total organic carbon composition of the sediments was negatively associated with PCB and mercury concentrations. A total of 29 benthic taxa was identified in 49 samples; replicate samples were taken at each of five sampling locations. Mean infaunal density across all sampling locations was estimated at approximately 61,000 to 234,000 organisms m(-2). Overall, polychaetes comprised 57% of the infaunal community with Manayunkia aestuarina as the dominant species. Oligochaetes, nematodes, crustacea, insects, and gastropods comprised 23.0, 18.0, 1.0, 0.7, and 0.2% of the overall benthic community, respectively. Density estimates of individual species between sampling locations showed no consistent patterns in response to pollutants. However, an analysis of higher taxonomic levels revealed some general trends. In uncontaminated areas, the benthic community was dominated by nematodes and oligochaetes, whereas moderate to highly contaminated areas were dominated by polychaetes and a smaller percentage of oligochaetes and nematodes. A trophic analysis of the same data set revealed that the community shifted from an evenly distributed percentage of surface and subsurface feeders in the uncontaminated areas to a community dominated by surface feeders in the more contaminated locations. Carnivores comprised from 0.13 to 0.90% of the trophic structure, with the percentage of carnivores generally decreasing with increasing contamination. Mercury and PCBs were bioaccumulating in representative marsh benthic invertebrates, presenting a potential source of contaminants to marsh consumers. Tissue PCB and tissue mercury concentrations were positively related to sediment PCB and mercury concentrations, respectively. A standard 14-day toxicity test using the amphipod Leptocheirus plumulosus showed no acute toxicity across the sampling locations.

Animals↗

The role of ingestion as a route of contaminant bioaccumulation in a deposit-feeding polychaete.

A mass-balance bioaccumulation model was used to examine the bioaccumulation of benzo[a]pyrene (BaP) from ingested sediment by the deposit-feeding polychaete Abarenicola pacifica over a series of experiments employing nine different sediments. Through selective ingestion of fine-grained material, the worm was able to increase the BaP content of ingested sediment by 10-35% above that of the bulk material. During digestion, an average of 5-21% of the ingested BaP was absorbed from gut contents. The relative importance of ingestion as a route of BaP uptake was dependent on the time period of observation. Initial uptake of BaP was postulated to come from absorption of dissolved BaP across the body wall since, after short periods of exposure (<24 h), only 3-38% of observed BaP tissue concentrations could have been derived from ingested material. With time and with increased feeding activity, however, ingested sediment became the major source of BaP to the organism. After 72 h of exposure, 36-119% of the tissue BaP (mean = 77%) was estimated to have been derived from dietary absorption of ingested material, and with continued exposure the dietary route is likely to be as great or greater as the relative significance of the initial uptake from the dissolved phase diminishes. This work and other studies indicate that for many deposit feeders, ingested sediment can be the primary source for the bioaccumulation of hydrophobic toxicants.

Administration, Oral↗

ParaHox gene expression in the polychaete annelid Capitella sp. I.

Hox and ParaHox genes are transcriptional regulators vital for many aspects of embryonic development in bilaterian animals and are considered to have originated from one ancestral proto-Hox/ParaHox cluster. Hox genes are clustered in the genome of both protostomes and deuterostomes, and there is a specific relationship between the position of a gene in the cluster and the position of its expression along the animal body axis (colinearity). It is not clear whether the ParaHox genes Gsx, Xlox, and, Cdx generally exhibit a similar phenomenon since developmental expression for all three ParaHox genes within a single species has not yet been described for any protostome animal. Here we show the spatial and temporal localization for all three ParaHox genes in the polychaete Capitella sp. I, a member of one of the morphologically most diverse and understudied groups within the Metazoa, the Lophotrochozoa. Our data demonstrate that although both CapI-Xlox and CapI-Cdx are regionally expressed in the gut, the three Capitella sp. I ParaHox genes as a group do not perfectly fit predictions of temporal or spatial colinearity. Instead, there is a conservation of expression across species associated with development of particular tissues, and the relative order of initiation of ParaHox gene expression likely reflects the relative order of species-specific tissue development during ontogenesis.

Animals↗

Embryonic expression of HeFoxA1 and HeFoxA2 in an indirectly developing polychaete.

Two forkhead family transcription factors, HeFoxA1 and HeFoxA2, were isolated from the serpulid annelid Hydroides elegans and their transcript distribution were characterized during embryogenesis. HeFoxA1 is first detected in second quartet blastomeres soon after their formation, and later in all vegetal half blastomeres, which comprise ectoderm, endoderm, and mesoderm precursors. HeFoxA1 expression declines first in subtrochal ectoderm and presumptive midgut precursors, as well as apparently in D quadrant blastomeres in advance of any known signaling events. Later, during gastrulation, HeFoxA1 declines in hindgut precursors, and by the end of gastrulation the expression remains active only in foregut precursors. HeFoxA1 is apparently expressed in ectomesoderm cells involved in forming the larva-specific protonephridium (the so-called head kidney). The other ortholog, HeFoxA2, is expressed in a subset of the cells in which HeFoxA1 is expressed during early stages, but later it is largely restricted to the endoderm-ectoderm boundary of the proctodaeum. In addition, HeFoxA2 has a unique expression in two hindgut cells and abutting ectoderm cells located by the imminent anal opening. The combined expression of HeFoxA1 and HeFoxA2 correlates with mesoderm and endoderm expression of their orthologs in other bilaterians.

Amino Acid Sequence↗

Hox gene expression in larval development of the polychaetes Nereis virens and Platynereis dumerilii (Annelida, Lophotrochozoa).

The bilaterian animals are divided into three great branches: the Deuterostomia, Ecdysozoa, and Lophotrochozoa. The evolution of developmental mechanisms is less studied in the Lophotrochozoa than in the other two clades. We have studied the expression of Hox genes during larval development of two lophotrochozoans, the polychaete annelids Nereis virens and Platynereis dumerilii. As reported previously, the Hox cluster of N. virens consists of at least 11 genes (de Rosa R, Grenier JK, Andreeva T, Cook CE, Adoutte A, Akam M, Carroll SB, Balavoine G, Nature, 399:772-776, 1999; Andreeva TF, Cook C, Korchagina NM, Akam M, Dondua AK, Ontogenez 32:225-233, 2001); we have also cloned nine Hox genes of P. dumerilii. Hox genes are mainly expressed in the descendants of the 2d blastomere, which form the integument of segments, ventral neural ganglia, pre-pygidial growth zone, and the pygidial lobe. Patterns of expression are similar for orthologous genes of both nereids. In Nereis, Hox2, and Hox3 are activated before the blastopore closure, while Hox1 and Hox4 are activated just after this. Hox5 and Post2 are first active during the metatrochophore stage, and Hox7, Lox4, and Lox2 at the late nectochaete stage only. During larval stages, Hox genes are expressed in staggered domains in the developing segments and pygidial lobe. The pattern of expression of Hox cluster genes suggests their involvement in the vectorial regionalization of the larval body along the antero-posterior axis. Hox gene expression in nereids conforms to the canonical patterns postulated for the two other evolutionary branches of the Bilateria, the Ecdysozoa and the Deuterostomia, thus supporting the evolutionary conservatism of the function of Hox genes in development.

Animals↗

Molecular phylogenetic evidence of a haplosporidian parasite infecting the polychaete Syllis nipponica (Imajima, 1966).

A haplosporidian parasite was detected in the polychaete Syllis nipponica via the amplification and sequencing of the 18S rDNA locus. This provides additional evidence of haplosporidians in polychaetes, the host group from which the first species of the phylum were described about a century ago. In phylogenetic analyses, the parasite was determined to place among mollusk-specific taxa. As such, separate taxonomic groups for haplosporidians infecting different host phyla should not be anticipated. These findings also underscore the as-yet unknown life cycles of commercially important parasites. Attempts to visualize spores of the parasite directly with scanning electron microscopy did not succeed.

Animals↗

Host-specific performance and host use in the kleptoparasitic marine snail Trichotropis cancellata.

Generalist parasites may disproportionately use certain hosts because of different benefits associated with each host species. I measured the growth rate of the marine snail Trichotropis cancellata, a facultative kleptoparasite that can suspension feed and steal food, on different hosts to determine the relative nutritional benefits of each host. The variation in tentacle (feeding structure) area among the hosts studied had the potential to provide parasitic snails with different amounts of nutrition for growth. In field experiments, suspension-feeding snails isolated from potential hosts grew at a similar rate to snails on brachiopods and significantly more slowly than snails on the following polychaete worms: Serpula columbiana (Serpulidae), Pseudopotamilla ocellata (Sabellidae), Schizobranchia insignis (Sabellidae), and Eudistylia vancouveri (Sabellidae). However, choice among worm hosts affected snail growth rates only in the fall, when phytoplankton levels are low. At this time, snails parasitizing the sabellids Schizobranchia and Eudistylia grew more quickly than snails on Serpula. In the spring and summer, with high levels of phytoplankton, Trichotropis grew at similar rates on all worm species tested. Trichotropis spent approximately the same time stealing food from each worm host species, >50% of the time the worms had their tentacles extended (the difference among hosts was not significant). This finding demonstrates that the similarity of snail growth rates on different worm species is not due to the snails compensating for poor hosts (worms that provide food at a slower rate) by spending more time stealing food. Snails in choice experiments preferred live Serpula to empty Serpula tubes, indicating that at least some of the cue(s) snails use to identify hosts are derived from living host tissues. In choice racks containing live Serpula and live Schizobranchia, snails did not choose one host worm significantly more often than the other. Because Trichotropis grows faster on sabellids than serpulids in the fall, I predicted that snails in nature would infect sabellids more often than other species. However, snails were usually distributed randomly among host species. In the few cases where the snails showed a significant preference among host species, proportionally more snails were found on serpulids than on sabellids or sabellarids. This study is the first to quantify under natural conditions the growth benefits of a kleptoparasite across the range of possible hosts, and implies that factors other than growth rate influence host choice specificity in the marine kleptoparasite T. cancellata.

Animals↗

Movement of carbon among estuarine habitats and its assimilation by invertebrates.

We measured the extent of movement of carbon and its assimilation by invertebrates among estuarine habitats by analysing carbon stable isotopes of invertebrates collected along transects crossing the boundary of two habitats. The habitats were dominated by autotrophs with distinct isotope values: (1) mudflats containing benthic microalgae (mean -22.6, SE 0.6 per thousand) and (2) seagrass and its associated epiphytic algae (similar values, pooled mean -9.8, 0.5 per thousand). Three species of invertebrates were analysed: a palaemonid shrimp, Macrobrachium intermedium, and two polychaete worms, Nephtys australiensis and Australonereis ehlersi. All species had a similar narrow range of isotope values (-9 to -14 per thousand), and showed no statistically significant relationship between position along transect and isotope values. Animals were relying on carbon from seagrass meadows whether they were in seagrass or on mudflats hundreds of metres away. Particulate organic matter collected from superficial sediments along the transects had similar values to animals (mean -11.1, SE 1.3 per thousand) and also showed no significant relationship with position. The isotope values of these relatively immobile invertebrates and the particulate detritus suggest that carbon moves from subtidal seagrass meadows to mudflats as particulate matter and is assimilated by invertebrates. This assimilation might be direct in the case of the detritivorous worm, A. ehlersi, but must be via invertebrate prey in the case of the carnivorous worm, N. australiensis and the scavenging shrimp, M. intermedium. The extent of movement of carbon among habitats, especially towards shallower habitats, is surprising since in theory, carbon is more likely to move offshore in situations such as the current study where habitats are in relatively open, unprotected waters.

Animals↗

To avoid or deter: interactions among defensive and escape strategies in sabellid worms.

Numerous studies demonstrate how sessile marine organisms utilize chemical, structural, and nutritional deterrents to persist in predator-rich environments. Little is known, however, about how mobile, more behaviorally complex species minimize predation by integrating avoidance and deterrence strategies. We investigated this using sabellid polychaete worms from the Caribbean and temperate western Atlantic. Sabellids extend their feather-like radioles beyond their protective tubes for feeding and respiration; the body remains inside the tube and the radioles retract when threatened. We used co-occurring consumers to determine the palatability of radioles and bodies for each of the eight species tested. In addition, we examined chemical or structural traits affecting palatability and evaluated predator escape traits, such as tube strength, speed of radiole retraction, completeness of retraction, and sensitivity to a nearby disturbance. All species had unpalatable radioles that were chemically or structurally defended, but only two species had unpalatable bodies. Thus, most species allocated defenses to tissues that were most exposed to predation. The two species with chemically defended bodies, Bispira brunnea and Bispira variegata, relied less on behavioral escapes than the other species. Their tubes were weak, they did not retract until disturbances were very close, and B. brunnea retracted slowly and incompletely even when touched. Other species generally had stronger tubes and/or retracted when disturbances were farther away. This trade-off of deterrence versus escape even occurred within a single species when populations differed in palatability. Populations of B.variegata from North Carolina and Georgia were chemically deterrent to both temperate and tropical consumers, while populations from Panama and Florida were palatable. The more palatable Panama population retracted in response to distant movement, while the unpalatable North Carolina population did not retract until nearly touched. Thus, most species utilize a combination of predator avoidance and deterrence strategies, but more deterrent populations of species utilized avoidance less.

Adaptation, Biological↗

Complementary DNA cloning and molecular evolution of opine dehydrogenases in some marine invertebrates.

The complete complementary DNA sequences of genes presumably coding for opine dehydrogenases from Arabella iricolor (sandworm), Haliotis discus hannai (abalone), and Patinopecten yessoensis (scallop) were determined, and partial cDNA sequences were derived for Meretrix lusoria (Japanese hard clam) and Spisula sachalinensis (Sakhalin surf clam). The primers ODH-9F and ODH-11R proved useful for amplifying the sequences for opine dehydrogenases from the 4 mollusk species investigated in this study. The sequence of the sandworm was obtained using primers constructed from the amino acid sequence of tauropine dehydrogenase, the main opine dehydrogenase in A. iricolor. The complete cDNA sequence of A. iricolor, H. discus hannai, and P. yessoensis encode 397, 400, and 405 amino acids, respectively. All sequences were aligned and compared with published databank sequences of Loligo opalescens, Loligo vulgaris (squid), Sepia officinalis (cuttlefish), and Pecten maximus (scallop). As expected, a high level of homology was observed for the cDNA from closely related species, such as for cephalopods or scallops, whereas cDNA from the other species showed lower-level homologies. A similar trend was observed when the deduced amino acid sequences were compared. Furthermore, alignment of these sequences revealed some structural motifs that are possibly related to the binding sites of the substrates. The phylogenetic trees derived from the nucleotide and amino acid sequences were consistent with the classification of species resulting from classical taxonomic analyses.

Amino Acid Oxidoreductases↗

Novel antifouling and antimicrobial compound from a marine-derived fungus Ampelomyces sp.

In this study, using a bioassay-guided isolation and purification procedure, we obtained 3-chloro-2,5-dihydroxybenzyl alcohol from a marine-derived Ampelomyces species that effectively inhibited larval settlement of the tubeworm Hydroides elegans and of cyprids of the barnacle Balanus amphitrite. The inhibitive effect on larval settlement was nontoxic and the EC50 of 3-chloro-2,5-dihydroxybenzyl alcohol ranged from 3.19 microg ml-1 to 3.81 microg ml-1 while the LC50 was 266.68 microg ml-1 for B. amphitrite cyprids; EC50 ranged from 0.67 microg ml-1 to 0.78 microg ml-1, and LC50 was 2.64 microg ml-1 for competent larvae of H. elegans, indicating that inhibitive effect of this compound was nontoxic. At a concentration of 50 mug per disc, this compound showed strong inhibitive effects on the growth of 13 out of 15 marine bacterial species tested in disc diffusion bioassay. Overall, the high inhibitory activities against bacteria and larval settlement as well as the non- or low-toxic nature of this compound to the barnacle and polychaete larvae suggest this compound could be a potent antifoulant and/or antibiotic.

Animals↗

Development and Characterization of Microsatellite Markers for the Euryhaline Polychaete Laeonereis acuta (Annelida: Nereididae) in the Southwestern Brazilian Coast.

Laeonereis acuta is a polychaete species typically found at high abundance in estuarine and coastal lagoon environments. Due to its association with polluted habitats, it is commonly used in ecotoxicological studies. Moreover, its occurrence in spatially discontinuous environments with high environmental variability makes it a suitable model for evolutionary studies of local adaptation, genetic landscape, and early stages of speciation. This study aimed to develop primers and characterize microsatellite markers for L. acuta sampled from three coastal lagoons in southwestern Brazil. A total of 10 loci were characterized based on the genotyping of 40 individuals. The number of alleles per locus ranged from 2 to 19. Evidence of null alleles was detected at five loci, although their frequency decreased when coastal lagoons were analyzed separately. When considering all individuals as a single population, five loci showed positive and significant FIS values, and seven loci deviated from Hardy-Weinberg equilibrium. Maric&#xe1; and Guarapina exhibited heterozygote excess at several loci, whereas Jacon&#xe9; showed evidence of population genetic isolation. The 10 microsatellite loci were polymorphic and suitable for population genetic analysis in L. acuta, although these patterns may not necessarily be representative of other geographic regions. These markers may contribute to ecotoxicological studies by clarifying whether physiological responses to pollutants are associated with genetic differentiation among populations. Furthermore, they provide valuable tools for investigating genetic structure and connectivity in discontinuous environments.

Animals↗

Chemical signal mediated premating reproductive isolation in a marine polychaete, Neanthes acuminata (arenaceodentata).

Neanthes acuminata Ehlers (1868) is a monogamous coastal polychaete with male parental care and a high level of sexual selection. We measured the level of prezygotic isolation among allopatric populations of N. acuminata; from the East and West Coast of the USA, a population from Hawaii, and a laboratory culture originating from Los Angeles, CA. All populations were found to preferably mate with members of their own population. Individuals from populations from Atlantic vs. Pacific Ocean failed to pair and to mate, either during the 10 min or 48 hr experiments. Instead, individuals showed high levels of aggressive behavior. Experiments measuring the levels of interpopulation aggression, established that individuals can recognize and discriminate among different populations of N. acuminata on the basis of olfactory cues. Aggressive behavior was induced by exposure of animals to seawater "conditioned" by individuals from the other populations, thus demonstrating the role of olfaction in the detection of "home" populations. The aggressive display was stronger upon exposure to seawater conditioned with "unrelated" populations and especially between Pacific and Atlantic populations.

Aggression↗