An earlier study demonstrated that under-yearling (0+) Atlantic salmon (Salmo salar L.) smolt had a lower vertebral mineral content and mechanical strength and higher prevalence of vertebral deformities than 1+ smolt during the early seawater (SW) phase. The present study aimed to examine if commercial extruded high-energy diets need to be supplemented additional minerals for proper bone mineralization and prevention of bone deformities in fast growing 0 + smolts. We studied vertebral morphology with radiology, and bone mineral content and mechanical strength in 60 g 0+ smolt fed diets with a normal (NM) or elevated (HM) bone mineral (P and Ca) contents from SW transfer (week 0) until 10 times weight increase at week 17. Thereafter, both groups were fed a commercial diet until a mean slaughter weight of 4100 g after 57 week. There were no differences in body weight and length between the dietary groups during the study, while the condition factor differed significantly at the final sampling (NM 1.40; HM 1.29). The most common bone deformity observed was compressions in the tail region of the vertebral column. Lower incidences of vertebral deformities (percent individuals with one or more deformed vertebrae) was observed in the HM group in week 17 (HM 20%; NM 47%) and week 57 (HM 37%; NM 73%), also reflected by higher vertebral length/dorso-ventral diameter ratio in weeks 17 (HM 0.99; MN 0.92) and 57 (HM 0.97; NM 0.88). The HM group had significantly higher vertebral mineral content (HM 550 g kg )1 ; NM 480 g kg )1 ) and mechanical strength (HM 9050 g mm )1 ; NM 4600 g mm )1 ) than the NM group after 8 week feeding. Plasma levels of Ca, P and D-vitamin metabolites recorded in week 8 reflected changes in P homeostasis, but could not explain the preventive effect of the HM diet on development of bone deformities. The results suggest that elevated dietary mineral content during the early SW phase may reduce the prevalence of vertebral deformities in fast growing 0 + salmon smolts. KEY WORDS

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This study investigates the effect of different smolt production strategies on vertebral morphology (radiology), composition (mineral content) and mechanical strength (load-deformation testing) in Atlantic salmon (Salmo salar). Rapid-growing underyearling (0+) smolt were compared with slower-growing yearling (1+) smolt and a reference group of wild smolt (w). The underyearling and yearling smolt were transferred to seawater in October 2002 and May 2003, respectively. The underyearling smolt were reared under continuous light and the yearling smolt under natural light during the first twelve weeks in seawater, at ambient temperatures. Thus, the underyearling smolt hit seawater at 13 °C and were reared at 10-13 °C during the early seawater phase, whereas the yearling smolt hit seawater at 7 °C and were reared at 7-10 °C during the early seawater phase. All groups displayed increased longitudinal growth (up to 9% increase in relative length) of the caudal vertebrae during parr-smolt transformation. However, at transfer to seawater, the underyearling smolt had significantly lower vertebral mineral content (0+ 44%, 1+ 47%, w 50%) and higher incidence of deformed vertebrae (0+ 1.5%, 1+ 0%, w 0%), and at twelve weeks after transfer to seawater significantly lower vertebral mineral content (0+ 36%, 1+41%, w 43%), yield-load (0+6492 g, 1+8797 g, w 9150 g) and stiffness (0+7578 g/mm, 1+ 15,161 g/mm, w 20,523 g/mm), and significantly higher incidence of deformed vertebrae (0+ 2.5%, 1+ 0.3%, w 0%). There was a significant correlation between the mineral content and mechanical properties of the vertebrae. The underyearling smolt had significantly elevated plasma concentrations of total Ca, and P and Ca2+ during the parr-smolt transformation and in the early seawater phase.The results show that underyearling smolt may have an increased risk of developing vertebral deformities. It is possible that this risk can be reduced by postponing the start of the short-day treatment. This will reduce the temperature during smoltification, the temperature and daylength during the early seawater phase, and increase the age at smoltification.

Summary The present review sums up and discusses the current literature on occurrence, causation and pathology of vertebral deformities in farmed Atlantic salmon, and also gives a brief introduction into the normal ontogeny and anatomy of the vertebral column of Atlantic salmon. Skeletal development and growth are sensitive processes that can be affected by many factors. Many of these factors can be manipulated under farming conditions, and are thus regarded as risk factors. Several risk factors that relate to environmental conditions and to feed composition have been identified. Elevated temperatures and photoperiod manipulation to speed up growth are likely the most important environmental factors that cause skeletal deformities. Among the nutritional factors, optimal phosphorus nutrition during specific periods, for example after transfer to sea water, appears to be critical for development of deformity at later stages. More research is needed to understand the interdependency of genetics, development, aging, phosphorus nutrition, temperature and photoperiod, in order to establish the best practice procedures for salmon farming that improve fish welfare.

In several terrestrial vertebrates, heat shock (HS) during somitogenesis causes vertebral deformities. To determine if vertebral deformities can occur due to sudden temperature changes during early development in fish, Atlantic salmon embryos were HS treated during somitogenesis. Ten months later these individuals displayed a high prevalence of caudal vertebral column condensations (27-34%). The defects were located caudally of the abdominal cavity, displaying an even distribution in this region independent of time of HS. To determine if HS disturbed vertebral development during somitogenesis, two genes coding for markers of skeletal development were identified, namely, the secreted protein Shh (Sashh) and the transcription factor Twist (Satwist). These proteins are involved in the proliferation and specification of presumptive skeletal cells (sclerotome) in vertebrates. The spatial expression pattern of sashh and satwist in salmon indicated a functional conservation of these proteins. Furthermore, HS embryos displayed expressional disturbance in both sashh and satwist, indicating an effect of HS on sclerotomal cell patterning. However, the HS-protecting ability in embryos seems to be individually regulated because reduction in gene expression was not detected at all stages; in addition, HS did not induce somitic disturbance and vertebral deformity in all embryos.

Suboptimal egg incubation temperature is a risk factor for the development of skeletal deformities in teleosts. Triplicate diploid and triploid Atlantic salmon, Salmo salar L., egg batches were incubated at 6, 8 and 10 °C up until first feeding, whereupon fish were reared on a natural temperature before examination for externally visible skeletal deformities (jaw and spine) and radiographed for vertebral deformities and morphology at the parr stage. Increasing incubation temperatures and triploidy increased the number of fish showing one or more deformed vertebrae. Triploids had significantly higher mean vertebrae cranio-caudal length (L) and dorsal-ventral height (H) ratio at 6 and 10 °C than diploids, but triploidy had no effect on mean vertebrae centra area. Triploids demonstrated an increase in lower jaw deformities with increased incubation temperature, whereas jaw deformities were rare in diploids. Fish incubated at 10 °C had a significantly lower mean vertebral number than fish incubated at 6 °C, and triploids had lower mean vertebral numbers than diploids. Diploid fish with 58 vertebrae had a significantly higher mean vertebral centra area than fish with 59 vertebrae, but vertebral number did not affect the mean vertebral L/H ratio. The results are discussed with respect to the welfare and production of farmed salmonids.

This study describes the long-term effects of surgical ablation of the pineal gland on the spine of 3-yr-old Atlantic salmon (Salmo salar L.) with a mean weight of 3.2 kg. Radiographic examinations showed that 82% of the pinealectomized fish developed marked lateral (scoliosis) and dorso-ventral spinal curvatures. The proportions of the individual vertebral bodies and their mechanical properties were also altered. The stiffness, yield limit and resilience of the vertebral bodies, as measured by compression in the cranio-caudal direction, were significantly lower in the pinealectomized than in the sham-pinealectomized group. Calcium, phosphorous and total mineral content of the vertebral bodies were also significantly lower in the pinealectomized fish, while these parameters were similar in scales in the two groups. Alterations of the spinal curve accompanied by changes in the proportions, mechanical strength and mineral content of the vertebral bodies of the pinealectomized salmon indicate that melatonin has several functions related to vertebral bone growth. As the lesions found in salmon are similar to the spinal malformations observed in avian species and mammals after pinealectomy, this study strengthens the hypothesis of a phylogenetically conserved function of the pineal gland related to skeletal development.