Association of plasma IgM with body size, histopathologic changes, and plasma chemistries in adult Pacific herring Clupea pallasi

Pacific herring Clupea pallasi immunoglobulin is an IgM-like molecule comprised of heavy and light chains with molecular weights of 79 and 25 to 27 kD, respectively. Purified immunoglobulin was used to generate highly specific polyclonal antibodies for development of a sandwich ELISA. The ELISA was used to quantify total plasma IgM in 602 Pacific herring captured in Prince William Sound and Sitka Sound, Alaska, USA. Plasma IgM concentrations ranged from 0.13 to 5.32 mg ml-1. Using multiple stepwise regression analysis, plasma IgM was highly correlated (p < or = 0.01) with body length, Ichthyophonus hoferi infection, plasma albumin, plasma cholesterol, liver macrophage aggregates, and focal skin reddening. I. hoferi was the only organism significantly associated with plasma IgM. Gender, site, and season (spring vs fall) did not contribute to significant differences in plasma IgM. This study contributes to the understanding of the interaction of body size, plasma chemistries, and pathological changes upon circulating immunoglobulins in fish.     

Growth and emigration of third-stage larvae of Hysterothylacium aduncum (Nematoda: Anisakidae) in larval herring Clupea harengus

The growth and emigration of Hystherothylacium aduncum in laboratory-reared herring larvae Clupea harengus was studied. Experimental infections of 36-day-old herring larvae resulted in 126 hosts infected with 306 H. aduncum larvae. Regression analyses showed a significant worm emigration from the rectum to the head of the fish, accompanied by an increase in worm body length. The emigration was independent of worm intensity, which suggests an ontogenetic process. Some worms departed from this pattern by moving posteriorly or by penetrating into the muscle, and in 5 cases, the larvae were observed to leave living fish. This individual variation has not been observed in previous studies and might be explained by host signals related to condition or development stage. Indirect evidence suggested parasite-induced mortality in the tanks due to the emigrations because only 4 of the 126 infected fish survived 8 days postinfection; the emigration of H. aduncum affected vital organs, such as the heart and brain, and the larvae penetrating or leaving the host's tissues can cause extensive damage to the delicate herring larvae.     

Chronic and persistent viral hemorrhagic septicemia virus infections in Pacific herring

Chronic viral hemorrhagic septicemia virus (VHSV) infections were established in a laboratory stock of Pacific herring Clupea pallasii held in a large-volume tank supplied with pathogen-free seawater at temperatures ranging from 6.8 to 11.6 degrees C. The infections were characterized by viral persistence for extended periods and near-background levels of host mortality. Infectious virus was recovered from mortalities occurring up to 167 d post-exposure and was detected in normal-appearing herring for as long as 224 d following initial challenge. Geometric mean viral titers were generally as high as or higher in brain tissues than in pools of kidney and spleen tissues, with overall prevalence of infection being higher in the brain. Upon re-exposure to VHSV in a standard laboratory challenge, negligible mortality occurred among groups of herring that were either chronically infected or fully recovered, indicating that survival from chronic manifestations conferred protection against future disease. However, some survivors of chronic VHS infections were capable of replicating virus upon re-exposure. Demonstration of a chronic manifestation of VHSV infection among Pacific herring maintained at ambient seawater temperatures provides insights into the mechanisms by which the virus is maintained among populations of endemic hosts.     

Larval and juvenile Pacific herring Clupea pallasii are not susceptible to infectious hematopoietic necrosis under laboratory conditions

Infectious hematopoietic necrosis (IHN) leads to periodic epidemics among certain wild and farmed fish species of the Northeast (NE) Pacific. The source of the IHN virus (IHNV) that initiates these outbreaks remains unknown; however, a leading hypothesis involves viral persistence in marine host species such as Pacific herring Clupea pallasii. Under laboratory conditions we exposed specific pathogen-free (SPF) larval and juvenile Pacific herring to 10(3) to 10(4) plaque-forming units (pfu) of IHNV ml(-1) by waterborne immersion. Cumulative mortalities among exposed groups were not significantly different from those of negative control groups. After waterborne exposure, IHNV was transiently recovered from the tissues of larvae but absent in tissues of juveniles. Additionally, no evidence of viral shedding was detected in the tank water containing exposed juveniles. After intraperitoneal (IP) injection of IHNV in juvenile herring with 10(3) pfu, IHNV was recovered from the tissues of sub-sampled individuals for only the first 5 d post-exposure. The lack of susceptibility to overt disease and transient levels of IHNV in the tissues of exposed fish indicate that Pacific herring do not likely serve a major epizootiological role in perpetuation of IHNV among free-ranging sockeye salmon Oncorhynchus nerka and farmed Atlantic salmon Salmo salar in the NE Pacific.     

Discrimination of Baltic herring populations with respect to Anisakis simplex larvae infection

Samples of herring Clupea harengus in the Baltic were collected during the spawning season in western, middle and eastern waters off the Polish coast. Two samples were taken per area, several morphometric characters were measured and some meristic characters (vertebrae and gill rakers) were counted. Infection with Anisakis simplex larvae, total length, age and maturity stage were recorded. Univariate and multivariate statistical tests were used to compare the samples, discriminant analysis was applied to separate herring populations and the fraction of herring populations in each area was estimated. The results obtained indicated that infected fish represented a mixed group of western and central Baltic herring. In the first samples collected from each area, the share of the western population decreased from 40% in the western to 18% in the middle and 13% in the eastern coasts. The share of western herring was higher in the second samples collected in the same area and decreased from 91% in the western coast to 14 and 32% in the middle and eastern ones, respectively.     

Comparison of fatty acid profiles of spawning and non-spawning Pacific herring, Clupea harengus pallasi

Crude lipid and fatty acid composition from liver, intestine, roe, milt and flesh of spawning and non-spawning Pacific herring (Clupea harengus pallasi) were examined to determine the relative effects of spawning on the nutritional value of herring. Depletion of lipid due to spawning condition was significant (P < 0.01) in all organ tissues and flesh of spawning herring. The lipid content ranged from an average of 1.9 to 3.4% (wet weight basis) in different organ tissues of spawning herring, to 10.5 to 16% in non-spawning fish. The fatty acid profile exhibited many differences in the relative distribution of individual fatty acids among organ tissues and between the two fish groups. Oleic acid (C18:1n-9), a major monounsaturated fatty acid (MUFA) found in all tissue lipids, decreased significantly (P < 0.01) in spawning fish. The two monoenes, C20:1n-9 and C22:1n-11, occurred at high concentrations in the flesh but at only minor proportion in the digestive organs and gonads. Spawning herring also had significantly (P < 0.01) higher polyunsaturated fatty acids (PUFA) content in the organ tissues, particularly in the milt and ovary, with docosahexaenoic acid (C22:6n-3, DHA) having the greatest proportion. Among the n-6 fatty acids, only C18:2n-6 and C20:4n-6 occurred at notable amounts and were present in higher proportions in spawning fish. We concluded that although relatively higher n-3 fatty acid content was found in the organ lipids of spawning herring, they are not an energy-dense prey food source due to the fact that both flesh and gonads contain a very low amount of lipid.     

Assessing the accuracy of a polymerase chain reaction test for Ichthyophonus hoferi in Yukon River Chinook salmon Oncorhynchus tshawytscha

Ichthyophonus hoferi Plehn & Mulsow, 1911, is a cosmopolitan, protistan pathogen of marine fishes. It is prevalent in mature returning Chinook salmon Oncorhynchus tshawytscha in the Yukon River watershed, and may be associated with prespawning mortality. We developed and evaluated a polymerase chain reaction (PCR) test for I. hoferi using primers specific to the parasite's small subunit rDNA. The test has a minimum detection limit of approximately 10(-5) parasite spores per reaction and does not cross-react with the closely related salmon parasites Dermocystidium salmonis or Sphaerothecum destruens. Sensitivity and specificity of the PCR test used on somatic muscle and heart tissue for detecting infected fish were determined using 334 Chinook salmon collected from the Yukon River at 2 locations (Tanana and Emmonak) in 2003 and 2004. The true infection status of the fish was determined by testing somatic muscle, heart and kidney tissue using histological evaluation, culture, and PCR. The severity of infection was grouped into 2 categories, light and heavy infection. The probability of detecting a heavily infected fish (sensitivity of the test) was generally much higher than the probability of detecting light infection, suggesting that more than one tissue and/or method should be used to accurately detect light or early infection by I. hoferi. The probability of correctly identifying a negative fish (specificity of the test) was always greater than 94% regardless of the tissue used, infection severity, sampling site or year of collection.     

Feeding, Ichthyophonus sp. infection, distribution and growth history of Norwegian spring-spawning herring in summer

After its collapse in the late 1960s, 1991 was the first year of systematic herring investigations after the Norwegian Sea stock returned to the traditional summer-feeding areas. Of 11 prey groups three dominated the diet; Calanus finmarchicus, Parathemisto spp. and Sebastes spp. There was a high prevalence of Ichthyophonus sp. at stations in the central and western parts of the Norwegian Sea where the cpue was low. This suggests that infected individuals were unable to return from the feeding to the wintering area of the Norwegian fjords. Highest cpue were recorded between 68–73° N and 5–15° E. Age groups 2, 3 and 8 were most frequently caught and age group 8 (the 1983 year class) dominated the samples. Herring occurred in dense concentrations where C. finmarchicus dominated diet. 0-group fish dominated diet at stations along the coast from 66–73° N and out to c . 300 km off the coast. Mature herring overlapped in distribution with 0-group fish and Sebastes spp. contributed largely to herring diet in this area. The first rich year-class after the collapse, the 1983 year class, had a faster growth than rich year classes from past herring periods, but slower than poor ones.     

Dynamics of viral hemorrhagic septicemia, viral erythrocytic necrosis and ichthyophoniasis in confined juvenile Pacific herring Clupea pallasii

Capture of wild, juvenile herring Clupea pallasii from Puget Sound (Washington, USA) and confinement in laboratory tanks resulted in outbreaks of viral hemorrhagic septicemia (VHS), viral erythrocytic necrosis (VEN) and ichthyophoniasis; however, the timing and progression of the 3 diseases differed. The VHS epidemic occurred first, characterized by an initially low infection prevalence that increased quickly with confinement time, peaking at 93 to 98% after confinement for 6 d, then decreasing to negligible levels after 20 d. The VHS outbreak was followed by a VEN epidemic that, within 12 d of confinement, progressed from undetectable levels to 100% infection prevalence with >90% of erythrocytes demonstrating inclusions. The VEN epidemic persisted for 54 d, after which the study was terminated, and was characterized by severe blood dyscrasias including reduction of mean hematocrit from 42 to 6% and replacement of mature erythrocytes with circulating erythroblasts and ghost cells. All fish with ichthyophoniasis at capture died within the first 3 wk of confinement, probably as a result of the multiple stressors associated with capture, transport, confinement, and progression of concomitant viral diseases. The results illustrate the differences in disease ecology and possible synergistic effects of pathogens affecting marine fish and highlight the difficulty in ascribing a single causation to outbreaks of disease among populations of wild fishes.     

Infection of wild fishes by the parasitic copepod Caligus elongatus on the south east coast of Norway

Natural Caligus elongatus Nordmann infections of wild coastal fishes on the Norwegian south east coast were monitored at various times of the year from 2002 to 2004. The prevalence for all coastal fish (n = 4427) pooled was 15%, and there were great differences between fish species and seasons. Lumpfish Cyclopterus lumpus L. spawners were the most infected fish, with a prevalence of 61% and a median intensity of 4 lice fish(-1), whereas gadids had a mean prevalence of 19% and a median infection of 1 to 2 lice fish(-1). Sea trout Salmo trutta L. and herring Clupea harengus L. carried C. elongatus at prevalence values of 29 and 21%, respectively. The results were compared with infection data for immature North Sea lumpfish. Lumpfish spawners caught on the coast in March to April had fewer lice than North Sea lumpfish in July. Spawners carried mostly adult lice, as did coastal fish hosts in May to June. The low development rates of lice at low spring temperatures and new genetic data suggest that the May to June adult lice could not have been offspring of the March to April lice, indicating transfer of adult lice to coastal fish. Most coastal fish species appeared to acquire new C. elongatus infections between May to June and September. The relatively high numbers of chalimii on North Sea lumpfish suggest that offshore fish sustain an oceanic population of this louse species.     

Purification and some properties of two creatine kinase isoforms from herring (Clupea harengus) spermatozoa

Creatine kinase (CK, EC 2.7.3.2) isoforms play important role in energy homeostasis and together with easily diffusible compounds like creatine and phosphocreatine maintain a cellular energy buffer and intracellular energy transport system. The CK activity in spermatozoa is the highest from all studied tissues in herring. It was detected that the two CK isoforms, CK1 and CK2, are characteristic only for spermatozoa and are not expressed in other herring tissues. Isolation and purification procedures allowed obtaining purified enzymes with specific activity of the 345 micromol/min/mg for CK1 and 511 micromol/min/mg for CK2. Native Mr's of the CK1 and CK2 determined by gel permeation chromatography were about 330,000 and 90,000, respectively. These results indicate that CK1 form has octameric structure and CK2 is a dimer mostly characteristic for cytosolic CK enzymes. In immunoblotting studies with antisera against CK2, the response was observed for CK2 and there was no response for CK1 and two other isoforms from herring skeletal muscle. These findings make the herring isoforms an interesting model for studies on the fish CK biochemical properties.     

Vulnerability of late larval and early juvenile Atlantic herring, Clupea harengus, to predation by whiting,Merlangius merlangus

Laboratory experiments investigated changes in the vulnerability of herring to predation by whiting during growth from <34 to >80mm in length, Premetamorphic herring (<50mm) failed to react to c . 50% of attacks by the predator, but this was reduced to c . 20% in postmetamorphic fish. Premetamorphic herring failing to react had c. 30% probability of survival due to unforced errors by the predator (unsuccessful attacks that did not elicit a reaction by the prey), but these errors did not occur after herring metamorphosis. The loss of the advantage of lower conspicuousness following metamorphosis was counterbalanced by increased reactivity. Antipredator benefits should increase with schooling behaviour (first evident in experiments at c . 50 mm), so predation mortality should decrease in postmetamorphic herring in the sea. Metamorphosis itself would be associated with high predation risk since conspicuousness is enhanced but reactivity and schooling behaviour are still not fully developed.     

The effect of progressive hypoxia on school structure and dynamics in Atlantic herring Clupea harengus

The effect of progressive hypoxia on the structure and dynamics of herring (Clupea harengus) schools in laboratory conditions was investigated. The length, width and depth of schools of about 20 individuals were measured from video recordings to test the hypothesis that during hypoxia fish schools change their shape and volume. School shape (calculated as the ratios of length/depth, width/depth and length/width) did not change significantly during hypoxia. School length, width, depth, area and volume were all significantly increased at 20% oxygen saturation. Volume, area and width were more sensitive to hypoxia; volume and width were also increased at 25% and area at 30% oxygen saturation. The degree of position changing (shuffling) of individuals within the school was also analysed. Shuffling in normoxia was observed to occur largely through 'O-turn' manoeuvres, a 360( degrees )turn executed laterally to the school that allowed fishes in the front to move to the back. O-turn frequency during normoxia was 0.69 O-turns fish(-1) min(-1) but significantly decreased with hypoxia to 0.37 O-turns fish(-1) min(-1) at 30% oxygen saturation. Shuffling was also investigated by measuring the persistence time of individual herring in leading positions (i.e. the first half of the school). No significant changes occurred during hypoxia, indicating that the decrease in O-turn frequency does not affect shuffling rate during hypoxia, and that position shuffling in hypoxic conditions is mainly due to overtaking or falling back by individual fishes. School integrity and positional dynamics are the outcome of trade-offs among a number of biotic factors, such as food, predator defence, mating behaviour and various physical factors that may impose certain limits. Among these, our results indicate that oxygen level modulates schooling behaviour. Oxygen alters whole-school parameters at oxygen saturation values that can be encountered by herring in the field, indicating that oxygen availability is an important factor in the trade-offs that determine school volume. An increase in school volume in the wild may increase the oxygen available to each individual. However, shuffling rate is not affected by hypoxia, indicating that the internal dynamics of positioning is the result of the balance of other factors, for example related to the nutritional state of each individual fish as suggested by previous studies.     

Ichthyophonus irregularis sp. nov. from the yellowtail flounder Limanda ferruginea from the Nova Scotia shelf

A previously described unusual form of the protistan parasite Ichthyophonus, differing in morphological and developmental features from I. hoferi sensu Plehn & Mulsow, was recovered from yellowtail flounder Limanda ferruginea Storer from the Brown's Bank area of the Nova Scotia shelf. The nuclear gene encoding the rRNA of the small ribosomal subunit was amplified from this unusual form of Ichthyophonus using the polymerase chain reaction, sequenced and aligned with other eukaryote small subunit (ssu)-rDNAs. Inferred phylogenetic trees clearly show that its ssu-rDNA is distinct from those of 2 isolates of I. hoferi sensu Plehn & Mulsow from different hosts and geographical locations (herring in the North Sea, and yellowtail flounder from the Nova Scotia shelf). We consider the unusual form to be a separate species, I. irregularis. The occurrence of a second, distinct type of Ichthyophonus within a single host species raises the possibility that ichthyophoniasis could be produced by different (although related) pathogens, and in some cases, by concurrent infections of the two.     

Differential utilization of energy during wintering and spawning migration in Norwegian spring-spawning herring

The energy utilization in non-feeding Norwegian spring-spawning herring Clupea harengus was examined during periods of wintering in a fjord system in northern Norway and southward spawning migration (800 km distance) to the main spawning area off western Norway. At the onset of wintering the gonadosomatic index (IG) was higher and the amount of mesenteric fat was lower in male than in female herring, whereas the subsequent increase in IG and decrease in mesenteric fat occurred at a higher rate in females than in males. However, no differences were recorded between sexes in fat content (%) of whole fish at any time of the year. The absolute energy loss increased, whereas the relative energy loss decreased, with fish length. Furthermore, the absolute and relative length specific energy losses were two to three and three to four times higher, respectively, during spawning migration than during wintering. The distinct energy loss was attributed mainly to fat metabolism, whereas the storage protein only was transferred from the soma to the gonads and remained relatively constant in whole herring throughout the wintering and spawning migration. © 1999 The Fisheries Society of the British Isles     

Relationship between feeding of blueback herring and the zooplankton community of a Texas reservoir

Following the introduction of blueback herring (Alosa aestivalis), we examined herring food habits and the crustacean zooplankton community in Lake Theo, a 30-hectare reservoir located on a tributary of the Red River in north Texas. Prior to the introduction in spring, 1982, the reservoir contained an established fish community dominated by centrarchids. Blueback herring stomachs and zooplankton were sampled quarterly from summer, 1982 through winter, 1985. Cladocerans accounted for 89.4% of the zooplankton consumed by herring. Median lengths of cladocerans and copepods in the herring diet were significantly greater than those in the reservoir. No appreciable changes in lengths of cladocerans and copepods in the reservoir were observed after blueback herring introduction, but the zooplankton community shifted from cladoceran to copepod domination.     

Viral hemorrhagic septicemia (VHS) of nonsalmonids

Viral hemorrhagic septicemia (VHS) is a viscerotropic disease of fish that can cause enormous losses in European rainbow trout populations. Although previously thought to be species specific, recent reports have indicated that other nonsalmonid fish species, such as pike, whitefish, grayling, turbot, and herring, can become mortally infected with VHS. This paper reviews several cases of natural VHS-outbreaks in nonsalmonids. Pike, whitefish, grayling, turbot and Pacific herring infected with VHS displayed symptoms typical of hemorrhagic septicemia. The isolation and serological identification of the viruses from all of the infected fish species revealed a close relation to the Egtved-virus strain F1. The virus isolates from diseased pike and rainbow trout were capable of infecting their respective host fish. Experimental infections of pike, whitefish, and grayling fry with Egtved virus strain F1 resulted in high mortalities, with symptoms typical of VHS. Although experiments with older fish suggested an age-dependent decrease in susceptibility, the virus could be isolated from most of the infected fish after several months of experimental infection (carrier status). Histological and electron microscopical findings were comparable to those seen in VHS-infected rainbow trout. The susceptibility of nonsalmonid species to VHS infection is disconcerting with respect to the potential losses of the economically important turbot, Pacific herring, and whitefish, as well as the ecologically valuable grayling which is threatened by extinction. If the VHS susceptible nonsalmonid species can also support the propagation of the Egtved-virus, it could acquire carrier-status and, as a result, represent a “natural” reservoir for the virus.     

Pacific and Atlantic herring produce burst pulse sounds

The commercial importance of Pacific and Atlantic herring (Clupea pallasii and Clupea harengus) has ensured that much of their biology has received attention. However, their sound production remains poorly studied. We describe the sounds made by captive wild-caught herring. Pacific herring produce distinctive bursts of pulses, termed Fast Repetitive Tick (FRT) sounds. These trains of broadband pulses (1.7-22 kHz) lasted between 0.6 s and 7.6 s. Most were produced at night; feeding regime did not affect their frequency, and fish produced FRT sounds without direct access to the air. Digestive gas or gulped air transfer to the swim bladder, therefore, do not appear to be responsible for FRT sound generation. Atlantic herring also produce FRT sounds, and video analysis showed an association with bubble expulsion from the anal duct region (i.e. from the gut or swim bladder). To the best of the authors' knowledge, sound production by such means has not previously been described. The function(s) of these sounds are unknown, but as the per capita rates of sound production by fish at higher densities were greater, social mediation appears likely. These sounds may have consequences for our understanding of herring behaviour and the effects of noise pollution.     

Screening for Viral Hemorrhagic Septicemia Virus in Marine Fish along the Norwegian Coastal Line

Viral hemorrhagic septicemia virus (VHSV) infects a wide range of marine fish species. To study the occurrence of VHSV in wild marine fish populations in Norwegian coastal waters and fjord systems a total of 1927 fish from 39 different species were sampled through 5 research cruises conducted in 2009 to 2011. In total, VHSV was detected by rRT-PCR in twelve samples originating from Atlantic herring (Clupea harengus), haddock (Melanogrammus aeglefinus), whiting (Merlangius merlangus) and silvery pout (Gadiculus argenteus). All fish tested positive in gills while four herring and one silvery pout also tested positive in internal organs. Successful virus isolation in cell culture was only obtained from one pooled Atlantic herring sample which shows that today''s PCR methodology have a much higher sensitivity than cell culture for detection of VHSV. Sequencing revealed that the positive samples belonged to VHSV genotype Ib and phylogenetic analysis shows that the isolate from Atlantic herring and silvery pout are closely related. All positive fish were sampled in the same area in the northern county of Finnmark. This is the first detection of VHSV in Atlantic herring this far north, and to our knowledge the first detection of VHSV in silvery pout. However, low prevalence of VHSV genotype Ib in Atlantic herring and other wild marine fish are well known in other parts of Europe. Earlier there have been a few reports of disease outbreaks in farmed rainbow trout with VHSV of genotype Ib, and our results show that there is a possibility of transfer of VHSV from wild to farmed fish along the Norwegian coast line. The impact of VHSV on wild fish is not well documented.