Mass mortalities due to disease outbreaks have recently affected major taxa in the oceans. For closely monitored groups like corals and marine mammals, reports of the frequency of epidemics and the number of new diseases have increased recently. A dramatic global increase in the severity of coral bleaching in 1997-98 is coincident with high El Niño temperatures. Such climate-mediated, physiological stresses may compromise host resistance and increase frequency of opportunistic diseases. Where documented, new diseases typically have emerged through host or range shifts of known pathogens. Both climate and human activities may have also accelerated global transport of species, bringing together pathogens and previously unexposed host populations. The oceans harbor enormous biodiversity by terrestrial terms (1), much of which is still poorly described taxonomically. Even less well known are the dynamics of intermittent, ephemeral, threshold phenomena such as disease outbreaks. Despite decades of intense study of the biological agents structuring natural communities, the ecological and evolutionary impact of diseases in the ocean remains unknown, even when these diseases affect economically and ecologically important species. The paucity of baseline and epidemiological information on normal disease levels in the ocean challenges our ability to assess the novelty of a recent spate of disease outbreaks and to determine the relative importance of increased pathogen transmission versus decreased host resistance in facilitating the outbreaks. Our objectives here are to review the prevalence of diseases of marine taxa to evaluate whether it can be concluded that there has been a recent increase. We also assess the contributing roles of human activity and global climate, and evaluate the role of the oceans as incubators and conveyors of human disease agents.
Plating methods for estimating survival of indicator organisms, such asEscherichia coli, and water-borne pathogens includingVibrio cholerae, have severe limitations when used to estimate viable populations of these organisms in the aquatic environment. By combining the methods of immunofluorescent microscopy, acridine orange direct counting, and direct viable counting, with culture methods such as indirect enumeration by most probable number (MPN) estimation and direct plating, it was shown that bothE. coli andV. cholerae undergo a "nonrecoverable" stage of existence, but remain viable. Following 2-week incubations in saltwater (5-25%o NaCl) microcosms, total counts, measured by direct microscopic examination of fluorescent antibody and acridine orange stained cells, remained unchanged, whereas MPN estimates and plate counts exhibited rapid decline. Results of direct viable counting, a procedure permitting estimate of substrate-responsive viable cells by microscopic examination, revealed that a significant proportion of the nonculturable cells were, indeed, viable. Thus, survival of pathogens in the aquatic environment must be re-assessed. The "die-off" or "decay" concept may not be completely valid. Furthermore, the usefulness of the coliform and fecal coliform indices for evaluating water quality for public health purposes may be seriously compromised, in the light of the finding reported here.
An environmental isolate (13- 1BB ) of Salmonella enteritidis serogroup C1 was inoculated into sterile Potomac River water microcosms to observe survival and culturability of the organism by employing acridine orange direct count, fluorescent antibody direct count, direct viable count, plate count on veal infusion agar and xylose lysine decarboxylase agar, and indirect enumeration by the most-probable-number method (MPN), using media selective for Salmonella. Loss of culturability on laboratory media was observed within 48 h. However, cultures could be "resuscitated" and cultured on solid media, following addition of nutrients to the microcosms . Cells, resuscitated 4 days after apparent "die-off" (0 colony-forming units (cfu)/mL) using plate count techniques, yielded numbers of cfu in the same order of magnitude as had been observed before the onset of nutrient limitation. Microscopic techniques for direct viable counting indicated that viability is maintained for as long as 60 days after depletion of nutrients, although attempts to culture these cells, by addition of nutrient, after 21 days yielded apparently sterile plates. Thus, longer periods of "dormancy" appear to require conditions other than simple nutrient addition for resumption of cell growth and division.
f Vibrio parahaemolyticus and Vibrio vulnificus, which are native to estuaries globally, are agents of seafood-borne or wound infections, both potentially fatal. Like all vibrios autochthonous to coastal regions, their abundance varies with changes in environmental parameters. Sea surface temperature (SST), sea surface height (SSH), and chlorophyll have been shown to be predictors of zooplankton and thus factors linked to vibrio populations. The contribution of salinity, conductivity, turbidity, and dissolved organic carbon to the incidence and distribution of Vibrio spp. has also been reported. Here, a multicoastal, 21-month study was conducted to determine relationships between environmental parameters and V. parahaemolyticus and V. vulnificus populations in water, oysters, and sediment in three coastal areas of the United States. Because ecologically unique sites were included in the study, it was possible to analyze individual parameters over wide ranges. Molecular methods were used to detect genes for thermolabile hemolysin (tlh), thermostable direct hemolysin (tdh), and tdh-related hemolysin (trh) as indicators of V. parahaemolyticus and the hemolysin gene vvhA for V. vulnificus. SST and suspended particulate matter were found to be strong predictors of total and potentially pathogenic V. parahaemolyticus and V. vulnificus. Other predictors included chlorophyll a, salinity, and dissolved organic carbon. For the ecologically unique sites included in the study, SST was confirmed as an effective predictor of annual variation in vibrio abundance, with other parameters explaining a portion of the variation not attributable to SST.
Although autochthonous vibrio densities are known to be influenced by water temperature and salinity, little is understood about other environmental factors associated with their abundance and distribution. Densities of culturable Vibrio vulnificus containing vvh (V. vulnificus hemolysin gene) and V. parahaemolyticus containing tlh (thermolabile hemolysin gene, ubiquitous in V. parahaemolyticus), tdh (thermostable direct hemolysin gene, V. parahaemolyticus pathogenicity factor), and trh (tdh-related hemolysin gene, V. parahaemolyticus pathogenicity factor) were measured in coastal waters of Mississippi and Alabama. Over a 19-month sampling period, vibrio densities in water, oysters, and sediment varied significantly with sea surface temperature (SST). On average, tdh-to-tlh ratios were significantly higher than trh-to-tlh ratios in water and oysters but not in sediment. Although tlh densities were lower than vvh densities in water and in oysters, the opposite was true in sediment. Regression analysis indicated that SST had a significant association with vvh and tlh densities in water and oysters, while salinity was significantly related to vibrio densities in the water column. Chlorophyll a levels in the water were correlated significantly with vvh in sediment and oysters and with pathogenic V. parahaemolyticus (tdh and trh) in the water column. Furthermore, turbidity was a significant predictor of V. parahaemolyticus density in all sample types (water, oyster, and sediment), and its role in predicting the risk of V. parahaemolyticus illness may be more important than previously realized. This study identified (i) culturable vibrios in winter sediment samples, (ii) niche-based differences in the abundance of vibrios, and (iii) predictive signatures resulting from correlations between environmental parameters and vibrio densities.Vibrio spp. occur naturally in estuarine and marine environments, and two species of this genus, V. vulnificus and V. parahaemolyticus, are responsible for the majority of reported vibrio illnesses in the United States (2). V. vulnificus infections are most commonly associated with the Gulf of Mexico, either via consumption of raw oysters harvested from these waters or wound infections following exposure to seawater. On average, about 50 cases of V. vulnificus septicemia are reported in the United States each year, with a case fatality rate of approximately 50% (31), the highest of any food-borne pathogen. In contrast, V. parahaemolyticus is the most common cause of seafood-associated bacterial gastroenteritis in the United States, with an estimated annual rate of 4,500 cases per year according to the Centers for Disease Control and Prevention. V. parahaemolyticus also causes wound infections, though these are less frequent and less severe compared to those caused by V. vulnificus (5). Primary septicemia can occur following V. parahaemolyticus infection, but it is relatively rare for this pathogen. In the United States, V. parahaemolyticus illness most often results from consumption of r...
Vibrio parahaemolyticus is indigenous to coastal environments and a frequent cause of seafood-borne gastroenteritis in the United States, primarily due to raw-oyster consumption. Previous seasonal-cycle studies of V. parahaemolyticus have identified water temperature as the strongest environmental predictor. Salinity has also been identified, although it is evident that its effect on annual variation is not as pronounced. The effects of other environmental factors, both with respect to the seasonal cycle and intraseasonal variation, are uncertain. This study investigated intraseasonal variations of densities of total and pathogenic V. parahaemolyticus organisms in oysters and overlying waters during the summer of 2004 at two sites in the northern Gulf of Mexico. Regression analyses indicated significant associations (P < 0.001) between total V. parahaemolyticus densities and salinity, as well as turbidity in water and in oysters at the Mississippi site but not at the Alabama site. Pathogenic V. parahaemolyticus organisms in Mississippi oyster and water samples were detected in 56% (9 out of 16) and 78% (43 out of 55) of samples, respectively. In contrast, 44% (7 out of 16) of oyster samples and 30% (14 out of 47) of water samples from Alabama were positive. At both sites, there was greater sample-to-sample variability in pathogenic V. parahaemolyticus densities than in total V. parahaemolyticus densities. These data suggest that, although total V. parahaemolyticus densities may be very informative, there is greater uncertainty when total V. parahaemolyticus densities are used to predict the risk of infection by pathogenic V. parahaemolyticus than previously recognized.Vibrio parahaemolyticus is the leading cause of Vibrio-associated gastroenteritis in the United States (15,19,20,32) and has been isolated from oysters, blue crabs, finfish, and planktonic copepods (6,7,18,22). Vibrio infections are most common in individuals living in states bordering the Gulf of Mexico (23) and are usually associated with the consumption of raw shellfish, primarily oysters (9, 20). Recent outbreaks (4,5,10,14) raised the awareness of public health officials concerning shellfish throughout coastal states and prompted the Interstate Shellfish Sanitation Conference to develop an Interim Control Plan for regulating shellfish harvest areas based on V. parahaemolyticus densities in shellfish (31). The Interim Control Plan employs a colony lift technique using DNA probes that target the species-specific thermolabile hemolysin (tlh) gene and the thermostable direct hemolysin (tdh) gene associated with pathogenic V. parahaemolyticus strains (32). A similar DNA probe colony hybridization method has been developed to target the tdh-related hemolysin gene (trh), which is also associated with pathogenic strains of V. parahaemolyticus (27). While these colony lift techniques make it possible to investigate the distribution of pathogenic V. parahaemolyticus strains directly, they have a relatively high limit of detection (LOD). Thus, the colony...
Several Vibrio spp. cause disease in marine fish populations, both wild and cultured. The most common disease, vibriosis, is caused by V. anguillarum. However, increase in the intensity of mariculture, combined with continuing improvements in bacterial systematics, expands the list of Vibrio spp. that cause fish disease. The bacterial pathogens, species of fish affected, virulence mechanisms, and disease treatment and prevention are included as topics of emphasis in this review, * + = 90% of all strains give a positive reaction, (+) = weak and/or delayed positive,-= 90 % of all strains are negative, V = variable reactions, A = acid, AG = acid + gas, R = resistant, S = sensitive; test procedures are given by West & Colwell (1984}. ** arginine dihydrolase, lysine and ornithine decarboxylase The natural habitat of Vibrio spp. is estuarine and marine water and sediment. Vibrios often occur in association with invertebrate and vertebrate animal surfaces and internal organs (Huq et al., 1983; Grimes et al, 1984b}, Stress can compromise the animal host, resulting in a commensal species switching to pathogenicity as the mode of interaction (Grimes et al., 1984c). * Formerly 1/. anguillarum biovar II Rodgers (1981) found V. alginolyticus to be a secondary invader of "red spot", a disease caused by V. "anguillarum. Gauthier & Clement (1979) observed a streptomycin-resistant strain of V. alginolyticus to be present not only in sediment but also water, when the water temperature exceeded 16 °C. However, food chain transmission was not observed when reconstructed marine food chains were studied in the laboratory (Gauthier & Clement, 1979). It is well established that V. alginolyticus can colonize the human gut (Sakazaki et al., 1963). In fact, Hiratsuka et al. (1980) observed the cause of acute enterocolitis in a patient who consumed trout roe contaminated with the same bacteria to be V. alginolyticus. More frequently, V. alginolyticus is associated with skin infections (Blake et al., 1980), such as conjunctivitis and stump ulcer cases described by Schmidt et al. (1979). Even though Blake et al. (1980) reported that little work has been done on virulence mechanisms, it is probable, based on available evidence, that the collagenase and several extracellular proteases produced by V, alginolyticus (Long et al., 1981) participate in the invasion of fish and human skin. Vibrio anguillarum This vibrio, the first recognized to be a fish pathogen, includes strains capable of utilizing arginine and tolerant of 6 % NaC1; some strains tolerate 8 % salt and others are capable of growth in the presence of trace amounts of salt, i.e., they are reported to grow
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