The aetiology of spinal deformity in Atlantic salmon, Salmo salar L.: influence of genetic factors on the frequency and severity in freshwater stages

A radiographic study was carried out on 2-year groups of pre-smolt parr to detect the comparative frequency of vertebral change in a range of defined pedigreed families of high growth rate of Atlantic salmon, Salmo salar L., reared under identical conditions. The observed frequency of deformity was related to the observed deformity level in the seawater parental generation, and to a potentially uncontrolled environmental determinant, i.e. stripping date. High and low parental deformity groups of families were found to produce offspring that had very similar levels of radiographically detectable spinal deformities in the pre-smolt parr, suggesting no genetic link. With regard to stripping date, the overall incidence was nearly half that seen in the preceding year (8.7% cf 16.6%). However, a non-significant trend to fewer radiographic spinal deformities was seen in the later stripping period.     

The aetiology of spinal deformity in Atlantic salmon, Salmo salar L.: influence of different commercial diets on the incidence and severity of the preclinical condition in salmon parr under two contrasting husbandry regimes

A large-scale trial of the effect of different commercial diets on the incidence of preclinical spinal deformation, as assessed by radiography, and the influence of two contrasting rearing systems was carried out. Two sets of three populations of Atlantic salmon, each of 20 000 first feeding fry of identical hatchery origin, created from equal numbers of eggs from 15 different families, were reared under commercial conditions on two different farms. Three commercial (closed formula) extruded fish meal-based diets were used in this study (diets A, B & C). Each diet was fed to one population of 20 000 fish at each site. Fish were fed a percentage of their body weight per day, with feeding rates set at commercial levels, based on water temperature, day length and fish biomass. Additional hand feeding was used to ensure satiation in all tanks. Fish in each tank were bulk-weighed and counted at the beginning and at 2-week intervals throughout the study. The fish were grown for 30 weeks. In addition, phosphorus (P) digestibility was evaluated by in-feed absorption testing in rainbow trout. The morphology of the radiographic lesions conformed to those described previously. Statistical analysis using multivariate regression analysis showed that date of sampling, site and diet were all statistically significant (P < 0.001) on univariable analysis. Farm A had significantly more affected fish than farm B (P < 0.001), which may have been attributable to variation in dissolved oxygen levels. The available dietary P levels were low in each diet. The number of fish affected in the group of fish being fed diet B was significantly lower than in the groups being fed diets A or C (P < 0.001). It appears most likely that the occurrence of preclinical radiographically apparent defects in parr which are believed to lead to the condition known as 'spinal deformity' is predominantly caused by a deficiency of available dietary P in first-feeding fry. The availability of dietary P may also vary considerably between diets formulated using different ingredients. Phytate-P associated with plant ingredients may affect the availability of P as well as other essential dietary nutrients. Additionally, diets for the production of salmonids in fresh water are currently formulated to keep P effluent to a minimum compatible with healthy spinal development. These various factors combine to make it crucial that small Atlantic salmon, especially first-feeding fry, are provided with carefully formulated diets fortified to an adequate level with a high quality source of available P.     

Association of spinal deformity and vaccine-induced abdominal lesions in harvest-sized Atlantic salmon, Salmo salar L

Spinal deformities in Atlantic salmon, Salmo salar L., have been described as a disease of multifactorial origin for which vaccines and time of vaccination have been suggested as risk factors. A vaccine efficacy trial where spinal deformity became evident was continued by the observational study reported here. In the preharvest part of the study 17 months post-sea transfer, there was a prevalence of 11.3% spinal deformity, with deformities present only in one vaccine group indicating a strong vaccine involvement. At slaughter, the prevalence of spinal deformities was 11.7%, and deformed fish had only 62% of normal slaughter weight. Visual analogue scales (VAS) were used for continuous recordings of vaccine-induced abdominal lesions and deformity. A logistic regression model associating presence of spinal deformity with markers of abdominal lesions was developed. The odds ratio for spinal deformity was 5.7 (95% CI: 3.4–9.4) for each unit increase in adhesion score (0–6) and 4.9 (2.9–3.4) for each unit increase in melanin on abdominal organs (0–3). Lesions in the dorsal caudal part of the abdomen gave an odds ratio for spinal deformity of 2.2.     

Screamer disease in Atlantic salmon, Salmo salar L., in Chile

Investigation of cranial and spinal deformities in Atlantic salmon smolts from Chile immediately after sea transfer, over two successive crops, demonstrated ankylosis of the mandibular articulation, spinal foreshortening, fracture of vertebrae and rarefaction of osseous and cartilaginous tissues including the operculum. As a result the mouth was permanently agape and opercula could be folded back on themselves. All affected fish had been transferred to sea water at 20 °C plus and fed on particular commercial diets. Fish in cooler areas, or on diets high in vitamin C and phosphorus in the high temperature zone, were clinically normal. The condition has not recurred following dietary adjustment.     

Dietary phosphorus requirement of juvenile Atlantic salmon, Salmo salar L.

The objective of this study was to determine the dietary phosphorus (P) requirement of juvenile Atlantic salmon, Salmon salar L. Triplicate groups of fish (mean initial weight 1.4 g) were fed semipurified, casein-gelatine-based diets containing one of five levels of P (4, 8, 10, 15 and 25 g kg⁻¹) from Ca(H₂PO₄)₂·H₂O, or a commercial feed (17 g kg⁻¹ P) for 9 weeks. Weight gains did not differ significantly among treatment groups fed the experimental diets but were slightly less than gains in fish fed the commercial feed. Feed efficiency (wet weight gain/dry feed consumed) was similar in all groups, averaging 1.45. Availability of dietary P, estimated from apparent retention and apparent digestibility, was 86%. Whole-body P concentrations declined in fish fed diets containing less than 10 g kg⁻¹ P. Fitting a logistic curve to dietary P vs. whole-body P concentrations indicated that a minimum of 11 g kg⁻¹ dietary P (9 g kg⁻¹ digestible P) was required by juvenile Atlantic salmon to maintain whole-body P concentrations at initial levels. Calculation of a dietary requirement using a simple factorial model which incorporated measurements of P availability, feed efficiency and normal whole-body P concentration indicated that the dietary requirement was approximately 10 g kg⁻¹. The dietary requirement established in this study (10–11 g kg⁻¹) is higher than previously reported for Atlantic salmon or other fishes. Possible reasons for the wide range of reported dietary P requirements in fishes are discussed.     

Teratogenicity of elevated egg incubation temperature and egg vitamin A status in Atlantic salmon, Salmo salar L

The present study was undertaken to investigate the possibility that high egg vitamin A (VA) status in combination with elevated egg incubation temperatures may cause deformities in Atlantic salmon, Salmo salar L. Egg batches selected for their total VA concentration were exposed to low (normal, 8 degrees C) or elevated (14 degrees C) egg incubation temperatures. Temperature was the main factor causing bone deformities such as warped gill opercula, fin and jaw deformities, but not for the development of spinal deformities where all groups displayed a 'baseline' occurrence of mild deformity (decreased vertebral size in the cephalic region) and no systematic variation in the occurrence of serious spinal deformities (fused vertebrae). A possible effect of egg incubation temperature fluctuation was found for the groups reared at low temperatures. An indication of a negative effect of elevated egg VA status for the development of organ deformities such as missing septum transversum and situs inversus was found in addition to temperature effects, however, no firm conclusions could be drawn from the present data. The phenotypes for temperature-induced deformities resembled the phenotype of VA-induced deformities, but no clear conclusions on the causality of the deformities found in the present study could be drawn. Egg incubation temperatures, both absolute temperature and temperature variations, should therefore be strictly controlled.     

Infectious pancreatic necrosis in Atlantic salmon, Salmo salar L

A clinical and histopathological review was carried out of 21 outbreaks of acute infectious pancreatic necrosis (IPN) in Scottish Atlantic salmon, Salmo salar L., farms (13 marine and eight fresh water) during 1991-2004. A distinctive syndrome was evident in both post-smolts in sea water and fry in fresh water, where liver lesions, which had not previously been associated with IPN, became a consistent finding in addition to the more typical pancreatic and intestinal changes. Initial cases were described in post-smolts in Shetland, but by the end of the period of investigation this type of pathology had extended down the West coast of Scotland and into Ireland. Limited viral strain analysis suggested that similar strains were involved in both fresh water and sea water and that these differed from earlier isolates from rainbow trout, Oncorhynchus mykiss (Walbaum). In fresh water, recovered fish frequently developed a greatly distended intestine associated with accumulation of undigested food. In sea water, after the initial, often significant (50% or more), losses, there were many fish which failed to grow and became chronically emaciated and prone to sea louse infection. Although use of transfer diets containing immune enhancers and the selection of IPN resistant broodstock has reduced losses the disease remains a serious cause of economic loss.     

Plasma protein changes in Atlantic salmon, Salmo salar L., with infectious salmon anaemia

Moderate to severe anaemia and hypoproteinaemia were reported in a Canadian outbreak of 'haemorrhagic kidney syndrome' in Atlantic salmon, later shown to be caused by a variant of infectious salmon anaemia virus (ISAV). The progressive anaemia associated with ISA has been previously reported, but hypoproteinaemia in salmon infected with European isolates of ISA virus has not been well documented. The present study showed a very significant positive correlation between decreasing haematocrit values and total plasma protein concentrations in Atlantic salmon infected with two Canadian and two Norwegian ISA viral isolates. However, variations in the concentration of individual plasma proteins, typical of acute phase responses in higher vertebrates, were not observed.     

The effect of water temperature on vertebral deformities and vaccine-induced abdominal lesions in Atlantic salmon, Salmo salar L

This study investigates the effects of water temperature (T) on vaccine-induced abdominal lesions (i.p. injection with oil-adjuvant vaccine) and vertebral deformities in Atlantic salmon. Quadruple groups of vaccinated (V) or unvaccinated (U) underyearling smolts were reared in tanks under four different temperature regimes for 6 weeks in fresh water (FW) followed by 6 weeks in sea water (SW). The four different T regimes were 10 °C FW-10 °C SW (10-10), 10 °C FW-16 °C SW (10-16), 16 °C FW-10 °C SW (16-10) and 16 °C FW-16 °C SW (16-16). After the temperature regimes were finished, the fish were group-tagged and transferred to a common sea cage for on-growth until harvest size. At termination, weight was significantly affected by both T and V, while lesion score and deformities were affected by T only. The weight difference between the largest and smallest U group was 20.3% (16-10 U: 2.4 kg, 10-16 U: 1.89 kg), while the largest difference between U and V fish within a T regime was 28.7% (16-16 U: 2.1 kg, 16-16 V: 1.5 kg). Fish from the 16-16, 16-10 and 10-16 regimes had a significant higher lesion score than those from the 10-10 regime. Fish from the 10-16 and 16-16 regimes displayed a significantly higher prevalence of vertebral deformities (palpation : 13-27%, radiology: 88-94%) than fish from the 10-10 and 16-10 regimes (palpation: 2-3%, radiology: 27-65%). Vertebra number 26 (located beneath the dorsal fin) was the most frequently affected vertebra in smolts, while vertebra number 43 (located above the anal fin) was most frequently affected in adults.     

Improved recapture rate of vaccinated sea-ranched Atlantic salmon, Salmo salar L.

Vaccination of sea-ranched Atlantic salmon was conducted in order to investigate if immuno-prophylactic measures could improve their survival. Fish were either vaccinated by bath or injection. A total of 66 000 fish were reared in fresh water at a hatchery on the island of Bornholm and at the presmolt stage were separated in three groups each comprising of 22 000 fish. One group was vaccinated intraperitoneally with a polyvalent vaccine (containing killed Vibrio anguillarum serotype O1 and O2, Yersinia ruckeri and Aeromonas salmonicida ). A second group was bath vaccinated with the corresponding vaccine-components and the third group was used as a non-vaccinated control. One month after vaccination these groups were allocated to three separate net-pens located 500 m from the coastline of the island. After 4 months in the net-pens, 1000 fish from each cage were tagged with Carlin-tags below the dorsal fin. The fish were then released for a migration period in the Baltic Sea. Following a sea period of 40 months (45 months post-vaccination), the recapture rates of the groups were calculated from the returned tags from fishermen. Recapture of the injection vaccinated group was significantly higher (25%) compared with the bath vaccinated fish (14.7%) and the control group (16.8%).     

Water temperature regimes affect cataract development in smolting Atlantic salmon, Salmo salar L.

The effect of fluctuation in water temperature during the freshwater rearing stage on cataract development in Atlantic salmon was investigated. Presmolts were exposed to either constant low or high temperatures, or high temperature with cold-water fluctuations. Temperature fluctuations caused the fastest growth rate and most severe cataracts, whilst low temperature resulted in slow growth and minor cataracts. The fish were then individually marked and maintained together in a seawater net pen for 7 months. Cataract progressed in all groups, fish from the low temperature group showed the fastest subsequent growth, and developed cataracts to the same degree as fish from the other groups. There were no statistically significant differences in selenium-dependent glutathione peroxidase levels in pooled samples of aqueous humour, lens and plasma from fish with or without cataract, except from the last freshwater samples, where cataractous fish showed lower plasma activity. Plasma cortisol and glucose levels did not differ, nor did whole eye riboflavin levels. Higher concentrations of some free amino acids in whole eyes were found in fish with cataract. This study shows that fluctuation in water temperature may cause both increased growth rate and cataract development in Atlantic salmon and that cataract development initiated in the freshwater rearing phase continues after transfers to seawater.     

Chemical pollution as a factor affecting the sea survival of Atlantic salmon, Salmo salar L.

The catch of Atlantic salmon Salmo salar L., from ocean and coastal fisheries since 1960 is described and the decline starting in 1973 noted. This decline continued despite cessation of some fisheries, and evidence indicates a marine locus for the decline. Oceanic pollution has received little attention, and ozone loss has caused an increase in UV radiation reaching the surface of the North Atlantic. Direct effects of this increase include negative impacts on the food chain. Photo-induced toxicity of polycyclic aromatic hydrocarbons (PAH) is a possible effect, but concentrations of these compounds in oceanic waters are low. Chlorinated organic compounds are widely distributed in the North Atlantic, and their persistence, lipophilicity, bioaccumulation potential and toxicity represent a potential hazard to salmon. The suggested causes of decline are likely to be complementary rather than exclusive.     

Heart and skeletal muscle inflammation in Atlantic salmon, Salmo salar L: a new infectious disease

Heart and skeletal muscle inflammation (HSMI) is a disease syndrome of unknown aetiology first observed in farmed Atlantic salmon, Salmo salar, in 1999. In the present study we have demonstrated for the first time that HSMI is an infectious disease. It was induced in Atlantic salmon post-smolts after injection with tissue homogenate from farmed Atlantic salmon previously diagnosed with HSMI. The lesions were also induced in cohabitating salmon given a corresponding injection without tissue homogenate. Six weeks post-challenge the fish that had been injected with tissue homogenate developed a serious epicarditis and myocarditis with mononuclear cell infiltrations in compact and spongy layers of the heart. Similar lesions were found in cohabitants after 10 weeks. The lesions were consistent with samples from field outbreaks of HSMI. No lesions were found in control fish. A viral aetiology is strongly suggested, as no difference in disease induction between an inoculum containing antibiotics and a non-treated inoculum was found. Further investigations are required in order to make conclusions regarding the cause and pathogenesis of HSMI.     

Randomized clinical field trial of a bacterial kidney disease vaccine in Atlantic salmon,Salmo salar L.

A randomized, blinded clinical trial was performed to assess the relative effectiveness of five commercial and one experimental vaccine in a population of farmed fish experiencing a bacterial kidney disease (BKD) outbreak that occurred in one study cage that was part of a larger clinical field trial. A total of 6000 uniquely identified Atlantic salmon S1 presmolts were randomly assigned to vaccine groups in the hatchery and transferred to a commercial marine aquaculture site. Repeated sampling events to evaluate growth, inherent physical conditions and health status were carried out over the entire production cycle. During the second summer at sea, the study cage developed an outbreak of BKD that lasted approximately 240 days. The effectiveness of the selected vaccines was evaluated using survival analysis methods. The sole vaccine group offering protection for BKD was found to significantly decrease the hazard of dying (hazard ratio, HR = 0.68, P = 0.018) during the outbreak, compared to the industry standard, vaccine group. Additionally, during the outbreak, fish with a shortened operculum had a significantly decreased hazard (HR = 0.38, P = 0.033) compared to those fish with a normal operculum, while fish with jaw deformities had a significantly increased hazard (HR = 2.55, P = 0.001) compared to fish with normal jaw status.     

Mycobacterium salmoniphilum infection in farmed Atlantic salmon, Salmo salar L

Multiple greyish‐white visceral nodules containing abundant rapidly growing and acid‐fast bacteria, subsequently identified as Mycobacterium salmoniphilum, were detected in moribund and newly dead market‐sized fish during a period of increased mortality in an Atlantic salmon, Salmo salar, farm in western Norway. Isolates cultured from diseased fish were phenotypically consistent with Mycobacterium sp. previously isolated from Atlantic salmon [MT 1890 (= NCIMB13533), MT1892, MT1900 and MT1901] in the Shetland Isles, Scotland. Partial sequences of 16S rDNA, ribosomal RNA internal transcribed spacer (ITS1), 65‐kDa heat‐shock protein (Hsp65) and β subunit of RNA polymerase (rpoB) revealed 97‐99% similarity with M. salmoniphilum type strain ATCC 13758^T. The source of infection was not confirmed. Koch’s postulates were fulfilled following experimental challenge of Atlantic salmon with field isolate NVI6598 ( FJ616988 ). Mortality was recorded in experimentally infected fish; however, the infection remained subclinical in the majority of affected fish over the 131‐day challenge period.     

Histidine nutrition and genotype affect cataract development in Atlantic salmon, Salmo salar L

The aim of this study was to investigate effects of dietary levels of histidine (His) and iron (Fe) on cataract development in two strains of Atlantic salmon monitored through parr-smolt transformation. Three experimental diets were fed: (i) a control diet (CD) with 110 mg kg(-1) Fe and 11.7 g kg(-1) His; (ii) CD supplemented with crystalline His to a level of 18 g kg(-1) (HD); and (iii) HD with added iron up to 220 mg kg(-1) (HID). A cross-over design, with two feeding periods was used. A 6-week freshwater (FW) period was followed by a 20-week period, of which the first three were in FW and the following 17 weeks in sea water (SW). Fish were sampled for weighing, cataract assessment and tissue analysis at five time points. Cataracts developed in all groups in SW, but scores were lower in those fed high His diets (P < 0.05). This effect was most pronounced when HD or HID was given in SW, but was also observed when these diets were given in FW only. Histidine supplementation had a positive effect on growth performance and feed conversion ratio (P < 0.05), whereas this did not occur when iron was added. Groups fed HD or HID had higher lens levels of His and N-acetyl histidine (NAH), the latter showing a marked increase post-smoltification (P < 0.05). The HD or HID groups also showed higher muscle concentrations of the His dipeptide anserine (P < 0.05). There was a strong genetic influence on cataract development in the CD groups (P < 0.001), not associated with tissue levels of His or NAH. The role of His and His-related compounds in cataractogenesis is discussed in relation to tissue buffering, osmoregulation and antioxidation.