Ecological characteristics of ninespine stickleback Pungitius pungitius from southern Baffin Island,Canada

Abstract – Stable δ¹³C and δ¹⁵N isotopes, diet and parasites demonstrated that the prey consumed by ninespine stickleback Pungitius pungitius in a small lake on Baffin Island changed during the summer and also revealed intraspecific variation in their ecological niche. In July, there were differences in the diets of male and female ninespine stickleback as indicated by the stable isotopes, differences corroborated by the data on diet composition and the parasite fauna. Differences suggested that the sexes occupied different habitats during spawning. During July, females utilise the shallower littoral areas consuming zooplankton and benthic organisms, while males occupy deeper areas of the littoral zone feeding mainly on pelagic zooplankton. Parasite data support these observations as males had higher infections of copepod‐transmitted parasites than females. There appeared to be no segregation of resources between males and females in late August, although the diet of both male and female ninespine stickleback shifted towards more benthic organisms, compared with July. Differences in δ¹³C isotope, diet composition and infections of co‐occurring parasites demonstrated that sympatric ninespine stickleback and Arctic char Salvelinus alpinus captured in the littoral zone occupied separate niches. Ninespine stickleback preyed mainly on zooplankton and chironomids, while Arctic char consumed a greater variety of prey items, including zooplankton and larger‐sized prey such as insects and ninespine stickleback. The multifaceted approach improved our understanding of the trophic ecology of ninespine stickleback in southern Baffin Island and quantified resource use and dietary overlap with Arctic char.     

Production and predation rates in a cannibalistic Arctic char (Salvelinus alpinus L.) population

Abstract – Production of cannibalistic Arctic char ( Salvelinus alpinus (L.)) and their prey conspecifics were estimated in an allopatric population, situated in an alpine ultraoligotrophic lake in central Norway. Estimates of population abundance were obtained by mark recapture and successive removal experiments. Assuming stable age distribution, we estimated mean yearly biomass (±95% CL) of char in sampled size groups vulnerable to predation (60 to 150 mm) to 7.93 (5.11–14.30) kg · ha⁻¹. Similarly, mean yearly biomass (±95% CL) of cannibalistic char (> 250 mm) was estimated to 0.62 (0.50–1.06) kg · ha⁻¹. Annual production (±95% CL) of char in length groups 60 to 150 mm was estimated to be 4.31 (2.74–8.03) kg · ha⁻¹ · year⁻¹, and production (±95% CL) of cannibalistic char to 0.19 (0.15–0.33) kg · ha⁻¹ · year⁻¹. Depending on the food conversion factor (set to vary from 0.1 to 0.4), the cannibalistic char removed from 10% to 40% of the production of char in sampled size-classes vulnerable to predation yearly. The overall ecological efficiency in energy transfer between the prey and predator population was 4.4%.     

Genetic population structure and variation at phenology-related loci in anadromous Arctic char (Salvelinus alpinus)

The Arctic will be especially affected by climate change, resulting in altered seasonal timing. Anadromous Arctic char (Salvelinus alpinus) is strongly influenced by sea surface temperature (SST) delimiting time periods available for foraging in the sea. Recent studies of salmonid species have shown variation at phenology-related loci associated with timing of migration and spawning. We contrasted genetic population structure at 53 SNPs versus four phenology-related loci among 15 anadromous Arctic char populations from Western Greenland and three outgroup populations. Among anadromous populations, the time period available for foraging at sea (>2°C) ranges from a few weeks to several months, motivating two research questions: (a) Is population structure compatible with possibilities for evolutionary rescue of anadromous populations during climate change? (b) Does selection associated with latitude or SST regimes act on phenology-related loci? In Western Greenland, strong isolation by distance at SNPs was observed and spatial autocorrelation analysis showed genetic patch size up to 450 km, documenting contingency and gene flow among populations. Outlier tests provided no evidence for selection at phenology-related loci. However, in Western Greenland, mean allele length at OtsClock1b was positively associated with the time of year when SST first exceeded 2°C and negatively associated with duration of the period where SST exceeded 2°C. This is consistent with local adaptation for making full use of the time period available for foraging in the sea. Current adaptation may become maladaptive under climate change, but long-distance connectivity of anadromous populations could redistribute adaptive variation across populations and lead to evolutionary rescue.     

Summer habitat use of Arctic char (Salvelinus alpinus) in a small Arctic lake, monitored by acoustic telemetry

Abstract –  Acoustic technologies were applied to describe how landlocked Arctic char from Iqalugaajuruluit Lake, Baffin Island, interact with its lacustrine habitat. Acoustic data from the lake bottom was collected using sonar equipment and substrate types were verified with benthic grabs and mapped in a geographic information system. Arctic char movements during the open water period were recorded from char fitted with acoustic tags. The distribution of the tagged Arctic char in Iqalugaajuruluit Lake was dependent on fish size and related to abiotic factors such as depth, substrate type and depth/temperature, temporally. The volume of water with temperatures below 6 °C during the open water period may be a limiting factor for large char (>400 mm) in small Arctic lakes. The large piscivorous char are found most often in the deepest water over soft substrates and the smaller char which feed on varying proportions of invertebrates and fish were found most often over the more complex substrates such as boulders, pebbles and gravel.     

Effect of androgens on seawater adaptation in Arctic char,Salvelinus alpinus

Sexually immature two-year old Arctic char (Salvelinus alpinus) were implanted with Silastic capsules containing testosterone or 11-ketoandrostenedione in early spring. Seawater adaptability of the hormone-treated and sham-operated fish was tested periodically from May to August using a 48h seawater challenge test with 25‰ seawater. The sham-operated control fish displayed a seasonal pattern in seawater adaptation, showing a good hypoosmoregulatory ability until mid June followed by a marked increase in plasma sodium and magnesium levels in July and August. Gill Na⁺-K⁺-ATPase activity decreased concurrently with the observed decrease in seawater adaptability. Over the same period the androgen-treated fish displayed a similar pattern in seawater adaptability, however, in May and June the plasma sodium levels were significantly higher in both androgen-treated groups compared to the control group. Plasma magnesium regulation was impaired in both androgen-treated groups in August. Gill Na⁺-K⁺-ATPase activity in the testosterone-treated fish was lower in June compared to the control fish, whereas the activity was not affected by 11-ketoandrostenedione treatment. The results show that while androgens impair the hypoosmoregulatory capacity in Arctic char, the seasonal pattern of seawater adaptability is not affected.     

Effect of extender composition and freezing rate on post-thaw motility and fertility of Arctic char, Salvelinus alpinus (L.), spermatozoa

The effects of extender composition and freezing rate on motility and fertility of frozen‐thawed Arctic char, Salvelinus alpinus, spermatozoa were investigated. Three freezing rates, two semen diluents and three cryoprotectants were tested. Semen frozen in 0.3 mol L^?1 glucose diluent with 10% methanol as a cryoprotectant or in a diluent described by Lahnsteiner with 10%N,N‐dimethylacetamide (DMA) resulted in the highest sperm motility. Fertility was the highest for semen frozen in a glucose–methanol extender but was not significantly different than that for semen frozen in Lahnsteiner's diluent with 10% DMA. Dimethyl sulphoxide (DMSO) at 10% was a relatively ineffective cryoprotectant with either semen diluent. Semen frozen at 6 cm above the surface of liquid nitrogen resulted in a higher post‐thaw sperm motility and fertility than semen frozen at 5 cm. The addition of 7% fresh egg yolk to glucose diluent containing methanol or DMSO did not improve the fertility of frozen‐thawed spermatozoa. However, the addition of 7% fresh egg yolk to glucose–DMA extender significantly improved the fertilization percentages of frozen‐thawed spermatozoa. In conclusion, dilution of semen 1:3 in 0.3 mol L^?1 glucose with 10% methanol and freezing 6 cm above the surface of liquid nitrogen (freezing rate of 40±8°C min^?1, mean±SD from ?5 to ?55°C) is a promising protocol for cryopreservation of Arctic char semen.     

Homing experiments with parr, smolt and residents of anadromous Arctic char Salvelinus alpinus and brown trout Salmo trutta: transplantation between neighbouring river systems

Abstract –  Olfaction is decisive for the homeward migration of anadromous salmonids. Two different olfactory hypotheses for explaining how this mechanism works have been proposed (the imprinting and the pheromone hypothesis), and they differ with regard to the origin of the odours that define home. The pheromone hypothesis assumes that fish respond to strain-specific pheromones and that no imprinting is necessary. The imprinting hypothesis assumes that juvenile fish become imprinted to odours of abiotic or biotic origin in natural rivers; the imprinting may be a single event or may occur sequentially during downstream migration as smolts. The two hypotheses were challenged by reciprocal transplantation of parr, residents and smolt of Arctic char ( Salvelinus alpinus ) and brown trout ( Salmo trutta ) between the Salangen and the Løksebotten rivers. Both rivers empty into the Salangen Fjord, Norway, about 10 km apart. Salangen Arctic char released in the Løksebotten River, and Løksebotten brown trout released in the Salangen River (parr, resident and smolt) initiated spontaneous homeward migration without preceding contact with home stream water. This indicates that site imprinting (single or sequential) could not have been involved. In the sea, the released fish apparently integrated into the migratory system of relatives from their river of origin. Recapture rates did not differ for fish allowed to receive single or sequential imprinting.     

Char ecology. Natal homing in sympatric populations of anadromous Arctic char Salvelinus alpinus (L.): roles of pheromone recognition

Abstract –  This study revealed that progeny of sympatric Salangen Arctic char, omitted from any kind of imprinting to home water, returned with high precision to the home river and parental locality upon releases in their native marine environments. Fish did not select river randomly, and the homing precision was the same for progeny whether fertilized in home stream or in foreign water. The progeny studied originate from the lake Øvervatnet and the Salangen River population and were hatched and reared at a hatchery at Voss in southern Norway. Experimental material comprised freshwater resident and smolt released from parr directly. The freshwater resident is partly anadromous exhibiting about 74% smolt when released directly into the sea. Char at all stages (resident, transformed resident, and smolt released from parr directly) returned in significant numbers to the home river system and the site of the specific parental population.     

莱州湾芙蓉岛人工鱼礁区大型底栖动物的营养结构特征

人工鱼礁建设是改善底栖生境、提高底栖动物多样性和资源丰度的重要措施之一,其建设效果与投放时间长短密切相关。为了探究投礁时长对大型底栖动物营养关系的影响,实验应用稳定同位素技术对比分析了莱州湾芙蓉岛海域不同礁龄人工鱼礁区和对照区大型底栖动物及其食源的碳、氮稳定同位素组成,计算了不同区域大型底栖动物的食物基础以及消费者的营养级。结果显示：①长礁龄区、短礁龄区和对照区的大型底栖动物δ¹³C值分别为−25.08‰~−13.34‰、−25.86‰~−17.80‰和−25.39‰~−11.06‰;δ¹⁵N值分别为10.73‰~15.78‰、10.02‰~14.89‰和10.17‰~15.80‰。②相较于短礁龄区和对照区,长礁龄区底栖动物群落营养多样性更高,食物来源多样性水平更高,群落内营养生态位更加多样化,群落结构稳定性较好。③以食性为依据,将大型底栖动物分为浮游生物食者 (Pl)、肉食者 (C)、碎屑食者 (D)和杂食者 (O)等4类摄食功能群,在4类潜在食源 [浮游植物、浮游动物、水体悬浮颗粒有机物 (POM)和沉积物有机物 (SOM)]中,浮游植物碳源对除肉食者外的3类消费者的平均贡献率最高 (31.40%)。④长礁龄区、短礁龄区和对照区的消费者营养级分别为2.00~3.83、2.00~3.49和2.00~3.87;此外在长礁龄区出现了相对多的高营养级捕食者,从而证实长礁龄区的底栖动物群落具有更复杂的营养结构。同时发现,人工鱼礁建设能使大型底栖动物更充分地利用浮游植物碳源,并且能够增加底栖动物群落的营养多样性和丰富度,促进群落中较高营养级动物的增加。本研究结果有助于了解人工鱼礁建设的环境生态效应,为进一步研究底栖生境食物网的物质循环和能量流动提供了基础资料。     

Effect of methanol concentration and thaw rate on the viability and fertility of cryopreserved Arctic char,Salvelinus alpinus (L.), spermatozoa

In two trials, Arctic char (Salvelinus alpinus) semen was frozen in 0.5 mL straws using extenders consisting of 0.3 M glucose and 10%, 12.5% or 15% methanol. Cryopreserved semen was thawed by immersing straws in 25 °C water for 17 s (11.6 °C s^?1) or in 5 °C water for 60 s (3.3 °C s^?1). The viability of the frozen–thawed semen was measured by determining post‐thaw motility and sperm membrane integrity. Two fertility trials were also conducted. There was no effect of trial or thaw rate on post‐thaw sperm viability or fertility. Use of 15% methanol in the extender resulted in the highest overall percentage of sperm motility and fertility. Use of 12.5% methanol as a cryoprotectant resulted in a higher per cent post‐thaw motility and a lower percentage of dead cells than did 10% methanol. Thus, levels of methanol higher than the commonly used 10% are beneficial for cryopreserving Arctic char sperm.     

Long-term variation in diet of Arctic char, Salvelinus alpinus, and brown trout, Salmo trutta: effects of changes in fish density and food availability

Abstract  The diet of Arctic char, Salvelinus alpinus (L.), and brown trout, Salmo trutta L., was studied in a large Norwegian reservoir over the first four decades (1954–1995) of regulation. Significant changes in diet occurred concurrent with changes in fish density and prey availability. Reduced biomass of fish due to the effect of hydropower development probably led to reduced inter- and intra-specific competition. Consumption of zoobenthos increased for both species during the first decade, fish then became an important food item concurrent with an increase in small-sized Arctic char in the littoral zone. The diet of Arctic char, but not brown trout, then changed radically as the introduced mysid, Mysis relicta Lovén, became an important food item.     

Dietary plasticity of Arctic charr (Salvelinus alpinus) facilitates coexistence with competitively superior European whitefish (Coregonus lavaretus)

Abstract – Habitat use and diet of Arctic charr (Salvelinus alpinus) coexisting with European whitefish (Coregonus lavaretus) and grayling (Thymallus thymallus) were studied in one deep and two relatively shallow subarctic lakes in northern Norway. Stomach content and stable isotope analyses revealed clear and temporally stable resource partitioning between the species in all three lakes. Arctic charr had a wide and flexible trophic niche and was the only piscivorous species. In contrast, whitefish and grayling had remarkably stable planktivorous and benthivorous niches, respectively. In the deepest lake, Arctic charr together with grayling mainly utilised littoral benthos, while piscivory was more prevalent in Arctic charr in the two shallower lakes. In one of the shallow lakes, whitefish was apparently relegated to the inferior profundal niche because of dominance of the littoral by grayling. Our results suggest that Arctic charr may not necessarily need an extensive profundal zone as a refuge, but can coexist with whitefish if a third competing fish species like grayling occurs in the littoral habitat or if profitable small prey fish are available. The study demonstrates that strong dietary plasticity of Arctic charr is instrumental in the observed coexistence with the commonly competitively superior whitefish.     

Dietary plasticity in Arctic charr (Salvelinus alpinus) in response to long‐term environmental change

Corrigan LJ, Winfield IJ, Hoelzel AR, Lucas MC. Dietary plasticity in Arctic charr (Salvelinus alpinus) in response to long‐term environmental change.
Ecology of Freshwater Fish 2011: 20: 5–13. © 2010 John Wiley & Sons A/S Abstract –  In the face of widespread environmental change impacts, there is a need to better understand mechanisms promoting flexibility and resilience of ecosystem components to such change to inform strategies for conservation. Glacial relict species are especially vulnerable to such changes. We investigated the behavioural responses of a native, glacial relict species, Arctic charr (Salvelinus alpinus) to long‐term environmental changes. It was hypothesised that changes in feeding behaviour would occur as a key intermediary to reduction in habitat availability (through climate change and eutrophication) or competitive interactions [with introduced roach (Rutilus rutilus)]. Stomach content analysis was used to assess the diet of 199 charr caught from Windermere, United Kingdom, in the months of March, June, September and November 2003–2007. The results were compared to data from 1940 to 1951 prior to the environmental changes and revealed a marked increase in the contribution of benthic invertebrates in the present diet. Stable isotope analysis confirms the results of the stomach analysis, suggesting that the charr have switched their diet from zooplanktivory towards benthivory. We discuss the possibility that habitat modification and roach population expansion in Windermere have contributed to changes in charr diet. Complementary studies suggest that this diet shift is more likely to be a response to the increasing roach population than to habitat modification; however, further study in Windermere would be required to elucidate the exact mechanism. Long‐term data sets such as these provide information that is useful for determining the directivity of ecological change and the capability of species to respond to change.     

Long-term variation in the population structure of Arctic charr, Salvelinus alpinus, and brown trout, Salmo trutta

The effects of induced water level fluctuations and introduction of the mysid Mysis relicta Lovén on population structure of brown trout, Salmo trutta L., and Arctic charr, Salvelinus alpinus (L.), were studied during 1953–1995 in Limingen hydroelectric reservoir, Norway. The main response was a marked reduction in catch‐per‐unit‐effort (CPUE) for trout and charr, probably caused by reduced recruitment following increased variation in water level. For both species, mean length decreased until 1967 and increased thereafter, whereas mean mass‐at‐length increased for the whole period. Both length and mass‐at‐length were negatively correlated with CPUE. The increases in mean length and mass‐at‐length were probably because of reduced competition following the reduced recruitment. Mysis relicta has become an important food item for charr but not for brown trout, but the increases in mean length and mass‐at‐length of charr started prior to the appearance of M. relicta in the charr diet.     

Littoral spawning habitats of three southern Arctic charr (Salvelinus alpinus L.) populations

Abstract Arctic charr populations in southern latitudes are nonmigratory, with all life‐stages limited to freshwater lakes and in‐ or out‐flowing tributaries. Although many of these populations are reported to also spawn in lake littorals, little is known about the physical characteristics of putative spawning grounds. A total of 23 discrete spawning sites within three Irish lakes were located by fyke netting of spawning adults and snorkelling in littoral habitats. Spawning sites were found to be long, narrow strips running parallel to the shore at a maximum depth of 124 cm. Spawning sites were limited to areas of coarse mineral substrate with an adequate (c. 8 cm) depth of clean interstitial spaces. In individual lakes, combined areas of spawning sites made up 0.4–0.7% of available littoral. Egg densities varied considerably between sites (33–900·eggs m^?2) and were significantly correlated with gradient and width of spawning sites. No evidence of redd digging was found. The shallow, localised and restricted nature of spawning grounds makes such populations vulnerable to anthropogenically induced postoviposition changes in surface water level, eutrophication processes such as increased lake sedimentation and elevated nutrient status.     

Trophic polymorphism among Arctic charr Salvelinus alpinus L., from Loch Ericht, Scotland

Abstract— This article examines the morphology and ecology of Arctic charr Salvelinus alpinus in Loch Ericht, Scotland, Gill netting revealed that two forms of charr existed within the loch, a pale form and a colored form. Multivariate analysis of nine size-corrected morphometric measurements of the head region revealed that pale and colored charr differed significantly in the size and shape of the head characteristics and that both groups were easily discriminated on the basis of these characteristics. Stomach contents showed that the colored form fed predominantly on zooplankton and the pale form mainly consumed benthic macro-invertebrates, with larger pale individuals consuming other fish. On the basis of diet and morphology, the terms planktivorous and piscivorous were adopted for the two forms respectively. Length at age also distinguished the two groups, with piscivorous charr being smaller than colored charr up to age 8+ then surpassing the maximum length attained by planktivorous charr at ages of 9+ and above. The results are discussed in relation to other instances of sympatric morphs of Arctic charr.     

Genetic divergence among broodstocks of Arctic charr Salvelinus alpinus in eastern Canada derived from the same founding populations

We examined the amount and distribution of molecular variation at microsatellite loci in 21 broodstocks of Arctic charr Salvelinus alpinus derived from the Fraser River, Labrador and the Nauyuk Lake, Nunavut, Canada. Our goal was to assess the amount of genetic diversity and differentiation as broodstocks are subdivided and propagated in different hatcheries and grow‐out facilities. We observed significant heterogeneity across pairs of ancestral and descendant broodstocks in the mean numbers of alleles at microsatellite loci. We detected a significant decrease in the observed heterozygosity between ancestors and descendants but the amount of decrease did not depend on either the degree of removal from the wild (number of sequential transfers) or the strain (Fraser vs. Nauyuk). Based on allele frequency distributions, there was little genetic evidence of bottlenecks during the creation of subsequent broodstock populations after the initial founding events. All broodstock samples were significantly differentiated from each other but those within the same strain were more similar to each other than to broodstocks from different strains. Broodstocks from the Nauyuk Lake broodstocks showed greater differentiation from each other than did Fraser River broodstocks, which could be attributed to differences in the number of founders.     

Seasonal variation in body composition of Arctic char, Salvelinus alpinus, from an ultraoligotrophic alpine lake

Abstract  – Seasonal pattern of energy content was determined in a population of Arctic char, Salvelinus alpinus (L.), living in an ultraoligotrophic alpine lake (altitude 1100 m) located in central Norway (62°5'N). Specific somatic energy content varied between 4.3 and 6.1 kJ · g⁻¹ in spawners and between 4.8 and 6.6 kJ · g⁻¹ in immatures. Corresponding values for specific somatic lipid energy were 0.5–1.6 kJ · g⁻¹ for spawners and 0.7–2.1 kJ · g⁻¹ for immatures. The temporal pattern in storage energy residuals (deviation from mean storage energy, all sampling periods pooled) indicated that total, protein and lipid energy accumulated in the autumn and early winter. This increase was followed by a winter decrease in somatic energy (January to April). A new increase in total somatic energy and somatic lipid energy occurred during late winter (March/April to June), while the lake was still ice covered and water temperatures ranged between 0.5 and 2.5 °C. The observed seasonal pattern of specific energy storage and lipid deposition demonstrated a considerable potential for energy accumulation in Arctic char at low temperatures. It is discussed whether freshwater lakes at high altitudes or latitudes represent a temperature-stabilised and predictable environment in contrast to the variable temperature experienced in the corresponding terrestrial system.     

Temperature–growth patterns of individually tagged anadromous Arctic charr Salvelinus alpinus in Ungava and Labrador,Canada

Individual measurements of annual, or within‐season growth were determined from tag‐recaptured Arctic charr and examined in relation to summer sea surface temperatures and within‐season capture timing in the Ungava and Labrador regions of Eastern Canada. Differences between two years of growth (2010–2011) were significant for Ungava Bay Arctic charr, with growth being higher in the warmer year. Growth of Labrador Arctic charr did not vary significantly among years (1982–1985). Regional comparisons demonstrated that Ungava Arctic charr had significantly higher annual growth rates and experienced warmer temperatures than Labrador Arctic charr. The higher annual growth of Ungava Bay Arctic charr was attributed to the high sea surface temperatures experienced in 2010–2011 and the localised differences in nearshore productivity as compared to Labrador. Within‐season growth rates of Labrador Arctic charr peaked in June, declined towards August and were negatively correlated with the length of time spent at sea and mean experienced sea surface temperatures. A quadratic model relating growth rate to temperature best explained the pattern of within‐season growth. Collectively, results suggest that increases in water temperature may have profound consequences for Arctic charr growth in the Canadian sub‐Arctic, depending on the responses of local marine productivity to those same temperature increases.     

Construction of Aeromonas salmonicida subsp. achromogenes AsaP1‐toxoid strains and study of their ability to induce immunity in Arctic char,Salvelinus alpinus L.

The metalloendopeptidase AsaP1 is one of the major extracellular virulence factors of A. salmonicida subsp. achromogenes, expressed as a 37‐kDa pre‐pro‐peptide and processed to a 19‐kDa active peptide. The aim of this study was to construct mutant strains secreting an AsaP1‐toxoid instead of AsaP1‐wt, to study virulence of these strains and to test the potency of the AsaP1‐toxoid bacterin and the recombinant AsaP1‐toxoids to induce protective immunity in Arctic char. Two A. salmonicida mutants were constructed that secrete either AsaP1_E294A or AsaP1_Y309F. The secreted AsaP1_Y309F‐toxoid had weak caseinolytic activity and was processed to the 19‐kDa peptide, whereas the AsaP1_E294A‐toxoid was found as a 37‐kDa pre‐pro‐peptide suggesting that AsaP1 is auto‐catalytically processed. The LD₅₀ of the AsaP1_Y309F‐toxoid mutant in Arctic char was significantly higher than that of the corresponding wt strain, and LD₅₀ of the AsaP1_E294A‐toxoid mutant was comparable with that of an AsaP1‐deficient strain. Bacterin based on AsaP1_Y309F‐toxoid mutant provided significant protection, comparable with that induced by a commercial polyvalent furunculosis vaccine. Detoxification of AsaP1 is very hard, expensive and time consuming. Therefore, an AsaP1‐toxoid‐secreting mutant is more suitable than the respective wt strain for production of fish bacterins aimed to protect against atypical furunculosis.