Diseases of American lobsters (Homarus americanus): a review

The American lobster fishery is a significant economic driver in coastal communities of North America. Increasingly, the impacts of infectious disease are recognized as important components and factors in the population ecology and subsequent management of the lobster fishery. Both environmental and anthropogenic factors impact marine diseases. The review herein highlights aspects of several important bacterial, fungal and protistan diseases, including gaffkemia, shell disease, vibriosis, disease caused by species of Lagenidium, Haliphthoros and Fusarium, paramoebiasis and Bumper Car disease. As the global environment continues to change, these diseases could more severely affect both wild caught and impounded lobsters.     

Dominance hierarchies among communally held juvenile lobsters,Homarus americanus

An investigation was conducted to determine the time for establishment of dominance hierarchies, their stability, and their relationship to molting and death of cultured juvenile lobsters, Homarus americanus. Agonistic displays between communally held animals were monitored daily. Interactions between all possible pairs of animals within each group were used to determine formation of a dominance hierarchy. A decrease in the frequency of agonistic interactions was recorded after the first day of the experiment. A stable four‐rank hierarchy was established within 2–3 days. When the dominant animal was molting, the subordinates displayed aggressively toward all members of the group but remained submissive to the dominant. When the dominant animal died, stability was lost and aggressive interactions among subordinates established a new ranking. Molting subordinates were highly susceptible to injury and death. Loss of subordinates did not affect the ranking of individuals.     

Use of injectable potassium chloride for euthanasia of American lobsters (Homarus americanus)

Potassium chloride (KCl: 330 mg/ml) was assessed as an euthanasia agent in American lobsters (Homarus americanus). Two groups of 10 lobsters (408.2 to 849.9 g) were maintained at 11.9 to 12.1 degrees C ('warm') and 1.5 to 2.5 degrees C ('cold') to evaluate the possible effect of ambient temperature on response to KCl. Death was defined as time of cardiac arrest, as viewed and measured by use of ultrasound. The KCl solution was injected (100 mg of KCl/100 g of body weight) at the base of the second walking leg to flood the hemolymph sinus containing the ventral nerve cord with potassium. Disruption of this 'central nervous system' was immediate, followed by cardiac arrest within 60 to 90 seconds. Group median ( +/- SD) baseline heart rate was 42 +/- 14 'warm' and 36 +/- 5 'cold' beats per minute. Time until cardiac arrest ranged from 35 to 90 (57 +/- 18) seconds in the 'warm' group and from 40 to 132 (53 +/- 34) seconds in the 'cold' group. There was no significant difference between group medians for either parameter. Histologic lesions were limited to mild to moderate acute degeneration, characterized by cell swelling, loss of contraction bands, and occasional mild cytoplasmic vacuolation of skeletal muscle at the injection site. Injectable KCl solution was an effective, reliable method for euthanasia of H. americanus.     

Bacteria induced shell disease of lobsters (Homarus americanus)

Chitin degrading species of bacteria in the genera Pseudomonas, Vibrio, and Beneckea were cultured from the lesions of lobsters (Homarus americanus) with a shell disease. A species of bacterium of the genus Vibrio (Beneckea) produced necrosis characteristics of shell disease in experimental lobsters when the integument had been damaged prior to inoculation.     

An unusual heparan sulfate isolated from lobsters (Homarus americanus)

An unusual heparan sulfate was isolated from lobsters (Homarus americanus). The polysaccharide has a composition and properties intermediate to heparin and the more common heparan sulfates. The sulfate and D-glucuronic acid content is high, while anticoagulant activity is low. The major repeating unit appears to consist of D-glucuronic acid and D-glucosamine N-O-disulfate. Some N-acetyl groups are present, the content of L-iduronic acid O-sulfate is low, and monosulfated or nonsulfated disaccharide-repeating units (very common in heparan sulfates) appear to be very rare. The data obtained again emphasize the heterogeneity of heparan sulfates and the need for adequate characterization when dealing with unusual or unexplored sources.     

Tailflick escape behavior in larval and juvenile lobsters (Homarus americanus) and crayfish (Cherax destructor)

We examined the escape behavior of larvae and postlarvae of the American lobster (Homarus americanus) and of adult immature (stage ADI) crayfish (Cherax destructor). Responses to standardized water jet stimuli delivered through a pipette were observed and analyzed. Lobster larvae did not respond to stimuli within 60 ms, indicating that they do not have functional giant fibers. The first movement by lobster larvae in response to water jet stimuli was a hyperextension of the abdomen. Larval escape responses also showed very little habituation. Postlarval lobsters and ADI crayfish showed the same range of responses as adult animals. Displacement efficiency of tailflicks exhibited by the different animals and stages was examined and related to the morphology of the animals. A separate behavior from tailflicking by larval lobsters in response to water jet stimuli was also observed. Here, the abdomen was hyperextended and the thoracic appendages were promoted. We termed this behavior a "starburst" response. The features of the tailflicking behavior suggest that it evolved to make the larvae difficult prey to handle for small, slower moving predators, and possibly to allow them to ride the bow waves of faster moving predators.     

Effects of exogenous serotonin on a motor behavior and shelter competition in juvenile lobsters (Homarus americanus)

Three experiments were conducted to determine (1) the pharmacodynamics of 5-hydroxytryptamine in juvenile lobsters; (2) the effects of 5-hydroxytryptamine, using a range of dosages, on a motor behavior used to escape an aversive situation; and (3) the effect of doses that did and did not inhibit this motor behavior on measures of dominance and shelter competition. The fate of 5-hydroxytryptamine in hemolymph over a 60-min post-injection period showed that the concentration fell rapidly to a low plateau that was maintained for at least 1 h. Low doses of 5-hydroxytryptamine did not affect locomotor behavior, but higher doses inhibited it. Dominance and subsequent possession of a shelter were unaffected by a low dose of 5-hydroxytryptamine but a higher dose that inhibited locomotion resulted in lobsters that lost fights and did not secure or retain possession of the shelter. In the context of dominance and shelter competition, we were unable to demonstrate any advantage of the low dose of exogenous 5-hydroxytryptamine and a severe disadvantage with the higher dose. Previous reports of transient increases in aggression in 5-hydroxytryptamine-treated subordinate lobsters did not take into account motor inhibition as a possible critical variable in aggression. Accepted: 14 April 2000     

A preliminary report of vibriosis in cultured American lobsters,Homarus americanus

A vibrio-like bacterium, designated Vibrio sp. (BML 79-078), was isolated from moribund juvenile Amercan lobsters, Homarus americanus. This is the first report of a vibrio-like bacterium being associated with a disease of lobsters. Koch's postulates have been satisfied for this bacterium. In this study, Vibrio sp. (BML 79-078) and Vibrio anguillarum (ATCC 19264) were found to be pathogenic when injected into juvenile lobsters held at 20°C, while Enterobacter aerogenes was not. An endotoxin was found to be associated with the pathogenesis of the disease syndrome caused by Vibrio sp. (BML 79-087).     

20-hydroxyecdysone causes increased aggressiveness in female American lobsters, Homarus americanus

Lobsters become transiently more aggressive before ecdysis. This aggressiveness accompanies an increase in hemolymph titers of 20-hydroxyecdysone (20-HE). Combats between intermolt female lobsters, injected with premolt levels of 20-HE, and larger, saline-injected opponents were videotaped. Aggressive, defensive, and avoidance behaviors were ranked according to aggressiveness in a Rank of Aggression hierarchy, which included opponent-directed and (nonopponent) redirected behaviors. Treated animals performed more and more highly aggressive behaviors than saline-injected controls. Opponents of treated animals performed fewer aggressive behaviors than saline-injected control opponents. Controls performed more defensive behaviors than treated animals, when redirected behaviors were considered. Differences in avoidance behaviors among the four types of combatants were not significant. The total aggressive content was the same in treated and control fights, but the interactions between combatants in the two fights were significantly different. Treated animals were equally as aggressive and defensive as their opponents; controls were relatively less aggressive and more defensive than their opponents. These results correlate with molt-cycle variations in behavior, 20-HE titers, and the effects of 20-HE and molt-differentiated hemolymph on the claw opener muscle. They suggest that 20-HE orchestrates intrinsic, cellular, and nuclear events that produce the molt-cycle transformations in agonistic behavior and aggressive state of lobsters.     

Human somatotropin stimulates the growth of young American lobsters, Homarus americanus (Crustacea, Decapoda)

Larval and postlarval lobsters, injected with human somatotropin (STH, or growth hormone), grew at a more rapid rate than untreated control animals over succeeding molts. Their mean carapace length was significantly longer than that of controls and they were heavier, although the moisture content of treated and control animals was similar. Injected STH increased the growth rate of individual animals by 10 to 20%. This is the first evidence for a growth enhancing effect of human somatotropin on a Crustacean.     

Effect of natural and laboratory diet on O : N ratio in juvenile lobsters (Homarus americanus)

Oxygen consumption and ammonia excretion rates of juvenile lobsters spanning the size range from shelter-restricted to vagile (7-60 mm CL, carapace length) were measured. There are no significant size-dependent break points in the slope of oxygen consumption or nitrogen excretion rates versus weight in freshly caught wild or laboratory fed lobsters. This suggests that there are no inherent changes in energy metabolism in juvenile lobster in the 7-60 mm CL range (stage V to adolescent lobsters), despite major behavioral shifts related to foraging and shelter use. In addition, the O : N ratio of freshly caught wild lobsters (15.7+/-7.4, mean+/-S.D.) is not significantly different from that of lobsters fed brown algae (13.4+/-4.9) or a mixed carbohydrate/protein diet (13.7+/-6.8) in lab. The O : N ratio of wild lobsters is significantly higher than lobsters fed a high protein diet (7.5+/-2.5) in the lab. This suggests that carbohydrates, particularly algal-derived carbohydrates, make up a significant portion of the metabolic fuel of wild juvenile lobsters from settlement through adolescence.     

Excretory calcinosis: a new fatal disease of wild American lobsters Homarus americanus

A significant number of moribund and dead lobsters Homarus americanus were reported to New York state authorities by lobster fishers in Long Island Sound (LIS) during the summer of 2002. Morbid lobsters were characterised by an orange discolouration of the abdomen, lethargy, an excess of epibionts and poor post-capture survival. On necropsy, severe extensive multifocal or diffuse mineralised granulomatous inflammation of the gills and antennal glands was the most striking pathology. In the gills, granulomas often occluded the lumen of filaments, resulting in congestion, ischemia and coagulative necrosis of gill tissues. In the antennal glands, granulomas were concentrated along the border between the coelomosac and labyrinth. No significant pathogens were recovered from diseased individuals. In prechronic individuals, however, it was evident that granulomas were focused around calcium carbonate (aragonite) crystals. This disease may result from anomalously high sea-bottom temperatures in LIS (approximately 23 degrees C) during the summer of 2002 and associated disruptions of the calcium chemistry of lobsters in favour of deposition of minerals in soft tissues. The ultimate cause of death of affected lobsters is probably respiratory failure due to reduced effective surface area of the gills, exacerbated by hypermetabolic temperatures and an abundance of epibionts.     

Circadian rhythms of heart rate in freely moving and restrained American lobsters, Homarus americanus

While circadian rhythms of locomotion have been reported in the American lobster, Homarus americanus, it is unclear whether heart rate is also modulated on a circadian basis. To address this issue, both heart rate and locomotor activity were continuously monitored in light-dark (LD) cycles and constant darkness (DD). Lobsters in running wheels exhibited significant nocturnal increases in locomotor activity and heart rates during LD, and these measures were significantly correlated. In DD, most lobsters exhibited persistent circadian rhythms of both locomotion and heart rate. When heart rate was monitored in restrained lobsters in LD and DD, most animals also demonstrated clear daily and circadian rhythms in heart rate. Overall, this is the first demonstration of circadian rhythms of heart rate in H. americanus, the expression of which does not appear to be dependent on the expression of locomotor activity.     

Circadian rhythms of heart rate in freely moving and restrained American lobsters,Homarus americanus

While circadian rhythms of locomotion have been reported in the American lobster, Homarus americanus, it is unclear whether heart rate is also modulated on a circadian basis. To address this issue, both heart rate and locomotor activity were continuously monitored in light-dark (LD) cycles and constant darkness (DD). Lobsters in running wheels exhibited significant nocturnal increases in locomotor activity and heart rates during LD, and these measures were significantly correlated. In DD, most lobsters exhibited persistent circadian rhythms of both locomotion and heart rate. When heart rate was monitored in restrained lobsters in LD and DD, most animals also demonstrated clear daily and circadian rhythms in heart rate. Overall, this is the first demonstration of circadian rhythms of heart rate in H. americanus, the expression of which does not appear to be dependent on the expression of locomotor activity.     

Development and metamorphosis of the digestive system of larval lobsters,Homarus americanus (Decapoda: Nephropidae)

This study provides a detailed account of the development of the digestive system of larval lobsters (Homarus americanus H. Milne Edwards, 1837) and the morphological changes that occur at metamorphosis. The most dramatic of these changes involves the gastric mill of the cardiac stomach. First-and second-stage lobsters lack the medial and lateral teeth characteristic of the grinding stomach of adult lobsters. Clearly recognizable, heavily cuticularized teeth first appear in the third stage, and accessory lateral teeth do not appear until the fourth stage. In place of the teeth of the gastric mill, first- and second-stage stomachs have a series of pads and ridges which are the apparent rudiments of the teeth. The development of the gastric mill during the larval stages enables lobsters to deal successfully with the more substantial food they encounter in the benthic environment, and corresponds to the drastic change of habitat and diet which occurs at metamorphosis. Confusion about the extent of the midgut and hindgut in larval lobsters has been clarified. The results of this study have shown that the larvae have a long midgut, which lacks a cuticle, and a short hindgut with a cuticular lining, just as in adult lobsters. The junction between midgut and hindgut lies in the sixth abdominal segment in all of the first four stages, as well as in the adult.     

Locomotory activity and exploration behaviour of juvenile European lobsters (Homarus gammarus) in the laboratory

The behaviour of juvenile European lobsters (Homarus gammarus) in the wild is little understood. A laboratory system was designed with a maze-like environment as an alternative to the large technological expenditure of a field study. It provided an apparently endless runway with uniform thigmotactical cues. Juvenile lobsters having a total length of 68 to 115 mm were studied. The lobsters showed an extensive nocturnal locomotory activity. They established home shelters in which they spent the day and covered distances of 1200 to 1600 m during the night. On average, the lobsters performed 136 excursions from their shelters, of which 10% led only to the immediate surroundings of the shelters. Of all the excursions 90% were shorter than 16 m. In some exceptional cases distances of several hundred meters were covered in the maze between shelter visits. Excursions of less than 16 m lasted on average less than 5 min. The frequencies of shelter visits during the dark phase were highest in small lobsters (300 visits) and lowest in larger lobsters (50 visits). The time spent within shelters decreased from 10% to less than 2% with lobster size. A distinct change in behaviour was obvious at a body length of 75 to 80 mm. Smaller lobsters behaved defensively and relied on shelter protection. Larger lobsters were less dependent on shelter protection and thus were able to explore and utilize their environment more intensively.     

Response of American lobsters Homarus americanus to infection with a field isolate of Aerococcus viridans var. homari (Gaffkemia): survival and haematology

American lobsters Homarus americanus were inoculated with a field isolate of the Gram-positive bacterium Aerococcus viridans var. homari, causative agent of gaffkemia, at 1 x 10(6), 1 x 10(4) or 1 x 10(2) colony forming units (CFU) kg(-1) or with sterile 3% NaCl and maintained at 10 or 15 degrees C until they died or were euthanised. Progression of disease in individual animals was monitored daily by total haemocyte count (THC) and haemolymph culture. Post-mortem examinations were performed on all lobsters. Effects of both ambient temperature and infective dose on survival time were observed. Marked bacteraemia occurred in all mortalities. Haemocytopenia (THC < 10 x 10(9) cells l(-1)) preceded death in most, but not all, mortalities.     

Mitotic and meiotic chromosomes of the American lobster Homarus americanus (Nephropidae, Decapoda)

The chromosomes of H. americanus have been characterised by C-banding, fluorochrome banding and restriction endonuclease banding. Thanks to these techniques, it has been possible to identify mitotic and meiotic figures clearly and to study the distribution and structure of heterochromatic regions. Moreover, we have identified small supernumerary chromosomes, variable in number and often asynaptic in first meiotic metaphase.