Puberty comprises the transition from an immature juvenile to a mature adult state of the reproductive system, i.e. the individual becomes capable of reproducing sexually for the first time, which implies functional competence of the brain-pituitary-gonad (BPG) axis. Early puberty is a major problem in many farmed fish species due to negative effects on growth performance, flesh composition, external appearance, behaviour, health, welfare and survival, as well as possible genetic impact on wild populations. Late puberty can also be a problem for broodstock management in some species, while some species completely fail to enter puberty under farming conditions. Age and size at puberty varies between and within species and strains, and are modulated by genetic and environmental factors. Puberty onset is controlled by activation of the BPG axis, and a range of internal and external factors are hypothesised to stimulate and/or modulate this activation such as growth, adiposity, feed intake, photoperiod, temperature and social factors. For example, there is a positive correlation between rapid growth and early puberty in fish. Age at puberty can be controlled by selective breeding or control of photoperiod, feeding or temperature. Monosex stocks can exploit sex dimorphic growth patterns and sterility can be achieved by triploidisation. However, all these techniques have limitations under commercial farming conditions. Further knowledge is needed on both basic and applied aspects of puberty control to refine existing methods and to develop new methods that are efficient in terms of production and acceptable in terms of fish welfare and sustainability.

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.

In August 1998, 3000 Atlantic salmon Salmo salar L. parr were divided into 7 groups with 2 replicates. Every 6 wk until March of the following year 1 group was vaccinated. One group was held as an unvaccinated control. The fish were transferred to seawater in May 1999, and slaughtered in February 2000. Temperature, fish size and photoperiod at vaccination, and the time between vaccination and sea transfer thus varied among the groups. In all vaccinated groups, growth was reduced for 1 to 2 mo following vaccination. Intra-abdominal lesions developed faster, and stabilised at a higher level in the groups vaccinated early at the highest temperature and the smallest fish size. Growth in seawater was influenced by the time of vaccination. At the end of the experiment, the group vaccinated last (MAR) was the heaviest of the vaccinated groups (4.0 kg), and the group vaccinated first, i.e. in August (AUG) was smallest (3.2 kg). Growth rate in seawater differed only in the summer when specific growth rate was above 1.45 in all groups. There was a correlation between adhesion, condition factor and number of weeks from vaccination to sea transfer. The AUG group had the highest condition factor, with a top level of 1.64 in autumn, and this group also displayed the highest incidence of deformed vertebra. The experiment shows that side effects of vaccination can be significantly reduced when planning the vaccination strategy, by taking environmental factors and fish biology into consideration.

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.