Models of marine molluscan diseases: Trends and challenges.

Disease effects on host population dynamics and the transmission of pathogens between hosts are two important challenges for understanding how epizootics wax and wane and how disease influences host population dynamics. For the management of marine shellfish resources, marine diseases pose additional challenges in early intervention after the appearance of disease, management of the diseased population to limit a decline in host abundance, and application of measures to restrain that decline once it occurs. Mathematical models provide one approach for quantifying these effects and addressing the competing goals of managing the diseased population versus managing the disease. The majority of models for molluscan diseases fall into three categories distinguished by these competing goals. (1) Models that consider disease effects on the host population tend to focus on pathogen proliferation within the host. Many of the well-known molluscan diseases are pandemic, in that they routinely reach high prevalence rapidly over large geographic expanses, are characterized by transmission that does not depend upon a local source, and exert a significant influence on host population dynamics. Models focused on disease proliferation examine the influence of environmental change on host population metrics and provide a basis to better manage diseased stocks. Such models are readily adapted to questions of fishery management and habitat restoration. (2) Transmission models are designed to understand the mechanisms triggering epizootics, identify factors impeding epizootic development, and evaluate controls on the rate of disease spread over the host's range. Transmission models have been used extensively to study terrestrial diseases, yet little attention has been given to their potential for understanding the epidemiology of marine molluscan diseases. For management of diseases of wild stocks, transmission models open up a range of options, including the application of area management, manipulation of host abundance, and use of scavengers and filter feeders to limit the concentration of infective particles. (3) The details of host population processes and pathogen transmission dynamics are blended in models that evaluate the effects of natural selection and/or genetic modification in developing disease resistance in the host population. Application of gene-based models to marine diseases is only now beginning and represents a promising approach that may provide a mechanistic basis for managing marine diseases and their host populations. Overall disease models remain both uncommon and underutilized in addressing the needs for managing molluscan diseases and their host populations.

Infectious diseases affect marine fisheries and aquaculture economics.

Seafood is a growing part of the economy, but its economic value is diminished by marine diseases. Infectious diseases are common in the ocean, and here we tabulate 67 examples that can reduce commercial species' growth and survivorship or decrease seafood quality. These impacts seem most problematic in the stressful and crowded conditions of aquaculture, which increasingly dominates seafood production as wild fishery production plateaus. For instance, marine diseases of farmed oysters, shrimp, abalone, and various fishes, particularly Atlantic salmon, cost billions of dollars each year. In comparison, it is often difficult to accurately estimate disease impacts on wild populations, especially those of pelagic and subtidal species. Farmed species often receive infectious diseases from wild species and can, in turn, export infectious agents to wild species. However, the impact of disease export on wild fisheries is controversial because there are few quantitative data demonstrating that wild species near farms suffer more from infectious diseases than those in other areas. The movement of exotic infectious agents to new areas continues to be the greatest concern.

Widespread survey finds no evidence of Haplosporidium nelsoni (MSX) in Gulf of Mexico oysters.

The advent of molecular detection assays has provided a set of very sensitive tools for the detection of pathogens in marine organisms, but it has also raised problems of how to interpret positive signals that are not accompanied by visual confirmation. PCR-positive results have recently been reported for Haplosporidium nelsoni (MSX), a pathogen of the oyster Crassostrea virginica in 31 of 40 oysters from 6 sites in the Gulf of Mexico and the Caribbean Sea. Histological confirmation of the PCR results was not undertaken, and no haplosporidian has been reported from the numerous histological studies and surveys of oysters in the region. To further investigate the possibility that H. nelsoni is present in this region, we sampled 210 oysters from 40 sites around the Gulf of Mexico and Puerto Rico using PCR and 180 of these using tissue-section histology also. None of the oysters showed evidence of H. nelsoni by PCR or of any haplosporidian by histology. We cannot, therefore, confirm that H. nelsoni is present and widespread in the Gulf of Mexico and the Caribbean Sea. Our results do not prove that H. nelsoni is absent from the region, but taken together with results from previous histological surveys, they suggest that for the purposes of controlling oyster importation, the region should continue to be considered free of the parasite.

Generation time and the stability of sex-determining alleles in oyster populations as deduced using a gene-based population dynamics model.

Crassostrea oysters are protandrous hermaphrodites. Sex is thought to be determined by a single gene with a dominant male allele M and a recessive protandrous allele F, such that FF animals are protandrous and MF animals are permanent males. We investigate the possibility that a reduction in generation time, brought about for example by disease, might jeopardize retention of the M allele. Simulations show that MF males have a significantly lessened lifetime fecundity when generation time declines. The allele frequency of the M allele declines and eventually the M allele is lost. The probability of loss is modulated by population abundance. As abundance increases, the probability of M allele loss declines. Simulations suggest that stabilization of the female-to-male ratio when generation time is long is the dominant function of the M allele. As generation time shortens, the raison d'être for the M allele also fades as mortality usurps the stabilizing role. Disease and exploitation have shortened oyster generation time: one consequence may be to jeopardize retention of the M allele. Two alternative genetic bases for protandry also provide stable sex ratios when generation time is long; an F-dominant protandric allele and protandry restricted to the MF heterozygote. In both cases, simulations show that FF individuals become rare in the population at high abundance and/or long generation time. Protandry restricted to the MF heterozygote maintains sex ratio stability over a wider range of generation times and abundances than the alternatives, suggesting that sex determination based on a male-dominant allele (MM/MF) may not be the optimal solution to the genetic basis for protandry in Crassostrea.

Relationship of parasites and pathologies to contaminant body burden in sentinel bivalves: NOAA Status and Trends 'Mussel Watch' Program.

The 1995-1998 database from NOAA's National Status and Trends 'Mussel Watch' Program was used to compare the distributional patterns of parasites and pathologies with contaminant body burdens. Principal components analysis (PCA) resolved five groups of contaminants in both mussels and oysters: one dominated by polycyclic aromatic hydrocarbons (PAHs), one dominated by pesticides, and three dominated by metals. Metals produced a much more complex picture of spatial trends in body burden than did either the pesticides or PAHs. Contrasted to the relative simplicity of the contaminant groupings, PCA exposed a suite of parasite/pathology groups with few similarities between the sentinel bivalve taxa. Thus, the relationship between parasites/pathologies and contaminants differs significantly between taxa despite the similarity in contaminant pattern. Moreover, the combined effects of many contaminants and parasites may be important, leading to complex biological-contaminant interactions with synergies both of biological and chemical origin. Overall, correlations between parasites/pathologies and contaminants were more frequent with metals, frequent with pesticides, and less frequent with PAHs in mussels. In oysters, correlations with pesticides and metals were about equally frequent, but correlations with PAHs were still rare. In mytilids, correlations with metals predominated. Negative and positive correlations with metals occurred with about the same frequency in both taxa. The majority of correlations with pesticides were negative in oysters; not so for mytilids. Of the many significant correlations involving parasites, few involved single-celled eukaryotes or prokaryotes. The vast majority involved multi-cellular eukaryotes and nearly all of them either cestodes, trematode sporocysts, or trematode metacercariae. The few correlations for single-celled parasites all involved proliferating protozoa or protozoa reaching high body burdens through transmission. The tendency for the larger or more numerous parasites to be involved suggests that unequal sequestration of contaminates between host and parasite tissue is a potential mediator. An alternative is that contaminants differentially affect parasites and their hosts by varying host susceptibility or parasite survival.

Assessment of the effectiveness of scup bycatch-reduction regulations in the Loligo squid fishery.

Discard reduction is a component of the statutory requirements of the Sustainable Fisheries Act. One species of concern is scup, Stenotomus chrysops, discarded in the Loligo pealei fishery. Initially, regulations were imposed restricting the fishery in time and space to avoid areas and times associated with high scup discarding. Modified gear was required in 2003 on any boat fishing in areas otherwise closed to the fishery to reduce scup discarding. The purpose of this study was to evaluate the success of the 2003 net regulations and the potential influence of time-area closures (GRAs) in achieving a reduction in scup discarding. The regulations are based on three expectations. (1) Reduction in discarding in the Loligo squid fishery will materially reduce total scup discarding. (2) Exclusion of Loligo squid fishing vessels from the GRAs will result in these vessels fishing in areas that inherently produce fewer scup discards without equivalently increasing discarding in other sensitive species. (3) The use of a square-mesh large-mesh section in the extension will reduce scup discarding to the extent that otherwise would be achieved if the boats fished outside the GRAs without the economic cost imposed by redeploying the fleet. Analysis of the NMFS-NEFSC observer database offers no support for the belief that Expectation 1 has been met. Squid catches were too low to sustain a directed fishery in the northern GRA during this study. Thus, had this area been open, limited scup discarding would have occurred. In this study, squid catches averaged 1025 kg tow(-1) in the southern GRA. Thus, had the GRA been open, Loligo fishing would have taken place. Yet, in the 34 tows taken by two vessels, not a single scup was caught. Redeployment of the fleet clearly increased scup discarding in 2003. Thus, Expectation 2 was not met. Field tests demonstrated that the implemented net modification can produce reduced catches of mostly smaller-sized finfish without impairing squid catch, but the data also indicate that this result may not be routinely achieved. Thus, Expectation 3 was not completely met. Implementation of the 2003 net regulation was likely premature, in that the specification was not adequate to guarantee the desired results. The history of the scup discarding issue in the Loligo squid fishery demonstrates that discard reduction cannot be accomplished without adequate prior evaluation of the sources of discards, without the requisite and concomitant experimental evaluation of the results of regulatory reform, and without adequate commercial-scale testing of prospective reforms prior to implementation.

Influence of water allocation and freshwater inflow on oyster production: a hydrodynamic-oyster population model for Galveston Bay, Texas, USA.

A hydrodynamic-oyster population model was developed to assess the effect of changes in freshwater inflow on oyster populations in Galveston Bay, Texas, USA. The population model includes the effects of environmental conditions, predators, and the oyster parasite, Perkinsus marinus, on oyster populations. The hydrodynamic model includes the effects of wind stress, river runoff, tides, and oceanic exchange on the circulation of the bay. Simulations were run for low, mean, and high freshwater inflow conditions under the present (1993) hydrology and predicted hydrologies for 2024 and 2049 that include both changes in total freshwater inflow and diversions of freshwater from one primary drainage basin to another. Freshwater diversion to supply the Houston metropolitan area is predicted to negatively impact oyster production in Galveston Bay. Fecundity and larval survivorship both decline. Mortality from Perkinsus marinus increases, but to a lesser extent. A larger negative impact in 2049 relative to 2024 originates from the larger drop in fecundity under that hydrology. Changes in recruitment and mortality, resulting in lowered oyster abundance, occur because the bay volume available for mixing freshwater input from the San Jacinto and Buffalo Bayou drainage basins that drain metropolitan Houston is small in comparison to the volume of Trinity Bay that presently receives the bulk of the bay's freshwater inflow. A smaller volume for mixing results in salinities that decline more rapidly and to a greater extent under conditions of high freshwater discharge.Thus, the decline in oyster abundance results from a disequilibrium between geography and salinity brought about by freshwater diversion. Although the bay hydrology shifts, available hard substrate does not. The simulations stress the fact that it is not just the well-appreciated reduction in freshwater inflow that can result in decreased oyster production. Changing the location of freshwater inflow can also significantly impact the bay environment, even if the total amount of freshwater inflow does not change.

Are molluscan maximum life spans determined by long-term cycles in benthic communities?

A review of the maximum longevity of bivalves and gastropods indicates that a greater than average number of life spans coincide with the periods of long-term cycles in marine communities. Apparently, long-term cycles exert an important influence on marine communities by affecting the life spans of constituent species. Gastropods and bivalves are affected differently, longevities being determined by some cycles more than others in each group. Overall, molluscan longevities tend to be slightly longer than the corresponding cycle suggesting that there is selection pressure for life spans slightly longer than the cycle controlling recruitment success and generational replacement.