Stressed mothers - troubled offspring: a study of behavioural maternal effects in farmed Salmo salar.

Mature female Atlantic salmon Salmo salar were given intraperitoneal cortisol implants 1 week prior to stripping to examine the influence of simulated maternal stress on offspring boldness and social dominance. Behavioural tests originally designed to investigate stress responsiveness and coping styles in salmonids (i.e. feeding in isolation, dominance tests and acute confinement) were carried out on the offspring 1·5 years after hatching. In the feeding test, there were no differences between the two treatment groups in total feeding score or number of pellets eaten, but offspring from the cortisol-implanted females made more unsuccessful feeding attempts than offspring from control females. In dominance tests, there was no difference between controls and cortisol-treated fish regarding propensity to become socially dominant. A higher proportion of individuals with bite marks, however, was observed in the cortisol group when compared to controls. Cortisol-treated offspring that gained dominant rank in the dominance tests performed more aggressive acts after stable dominance-subordinate relationships were established compared to control winners. During acute confinement stress, offspring from cortisol-implanted females showed a reduction in the proportion of time they were moving compared to the controls. These results indicate that the maternal endocrine state at spawning affects several aspects of progeny behaviour potentially related to subsequent success and survival in farmed S. salar.

Melanin-based skin spots reflect stress responsiveness in salmonid fish.

Within animal populations, genetic, epigenetic and environmental factors interact to shape individual neuroendocrine and behavioural profiles, conferring variable vulnerability to stress and disease. It remains debated how alternative behavioural syndromes and stress coping styles evolve and are maintained by natural selection. Here we show that individual variation in stress responsiveness is reflected in the visual appearance of two species of teleost fish; rainbow trout (Oncorhynchus mykiss) and Atlantic salmon (Salmo salar). Salmon and trout skin vary from nearly immaculate to densely spotted, with black spots formed by eumelanin-producing chromatophores. In rainbow trout, selection for divergent hypothalamus-pituitary-interrenal responsiveness has led to a change in dermal pigmentation patterns, with low cortisol-responsive fish being consistently more spotted. In an aquaculture population of Atlantic salmon individuals with more spots showed a reduced physiological and behavioural response to stress. Taken together, these data demonstrate a heritable behavioural-physiological and morphological trait correlation that may be specific to alternative coping styles. This observation may illuminate the evolution of contrasting coping styles and behavioural syndromes, as occurrence of phenotypes in different environments and their response to selective pressures can be precisely and easily recorded.

Alterations of the pituitary-gonadal axis in the neonatal blue fox (Alopex lagopus) exposed to prenatal handling stress.

Handling is a stressor for blue foxes. The influence of preterm handling (1 min daily) of vixens was investigated in 68, 10-day-old cubs, 34 male and 34 female. Body, gonadal and pituitary weight, ano-genital distance, gonadal testosterone and oestradiol content and in vitro production, and pituitary LH content were measured in all cubs. The gonads were frozen or incubated in vitro with, or without, added hCG (2.5 IU per sample). The gonadal incubates and homogenates were analysed for testosterone and oestradiol by radioimmunoassay and the pituitary homogenates for LH by immunofluorometric assay. The results indicate that neonatal fox gonads actively produce steroids and that there are significant sex differences in basal steroid production and response of the gonads to hCG, and in pituitary LH content. Maternal stress resulted in a significant reduction of morphometric and hormonal measures of the reproductive system in neonatal blue foxes, with more drastic effects in female cubs. Gonadal weights were lower in cubs of both sexes from stressed vixens (65.7+/-4.3 v 50.6+/-1.8 mg for the ovaries and 23.2+/-1.0 v. 17.7+/-1.0 mg for the testes, control v. stressed animals, P<0.01). The ano-genital distance in female offspring of stressed vixens was reduced (1.1+/-0.04 v. 0.9+/-0.03 cm, P<0.01). Basal ovarian oestradiol and testosterone production were decreased in cubs from prenatally stressed animals in comparison with controls (43.5+/-3.5 v. 32.6+/-3.7 pg ovary(-1) h(-1) and 0.40+/-0.16 v. 0.12+/-0.03 ng ovary(-1) h , P<0.05). Prenatal stress did not affect either pituitary weights or LH content in either sex. There were no significant differences in ano-genital distance, testicular content of testosterone, or in vitro testosterone production between control and treated male cubs. In conclusion, these findings suggest that prenatal handling stress impaired the neonatal reproductive development of the female offspring, but had no marked effects on males. Sex-specific effects of prenatal handling stress on the reproductive development in foxes may be linked with the gender differences in responses of the hypothalamus-pituitary-adrenocortical axis to stress conditions in this species.

A cautionary note regarding the use of nutritional L-tryptophan to alter aversion-related behaviour in mice.

Nutritional L-tryptophan is reported to have positive effects on the behaviour of several farm animals. The present experiment describes some effects of nutritional L-tryptophan on the aversion-related behaviour of male mice selected for high litter size. The behavioural effects of L-tryptophan were observed in the elevated plus-maze, the light/dark test and a resident-intruder test. The only significant effect was observed in the resident-intruder test, where the total occurrence of exploratory behaviour by the intruder was reduced by L-tryptophan treatment. The present experiment provides a cautionary note advising against supplementation with L-tryptophan to improve animal welfare before the consequences of such treatment are fully understood.

Social facilitation of predatory, sheep-chasing behaviour in Norwegian Elkhounds, grey.

Sheep grazing on unfenced mountain pasture may be attacked by loose dogs, which may chase and kill sheep and separate lambs from their mothers. We have earlier shown that testing dogs individually towards sheep with an electronic dog collar may effectively reduce the chasing propensity 1 year later. The aim of this study was to investigate whether and how a non-chasing and a chasing companion dog influence sheep chasing in test dogs (20 Norwegian Elkhounds, grey), and whether this varies with sex or age. The test dogs' predatory behaviour towards sheep was examined by observation of the dogs in a fenced enclosure with sheep in 5min tests together with, first, a non-chasing (Hamilton stoever, a hound breed) and 2-3 days later a chasing (Border collie) companion dog. Physical contact with sheep was prevented. Initially, the test dogs exhibited a higher chasing motivation towards sheep in tests with a chasing, as compared with a non-chasing, companion (P<0.001). During the entire test, 60% of the dogs attacked sheep when accompanied by the non-chaser. All dogs attacked sheep when the chasing companion was present (P=0.008), although in only 8 of 20 cases the companion dog chased simultaneously with the test dog. In the chasing-companion tests, the attack latency was shorter (P<0.001) than in tests with a non-chasing companion. The attack severity was higher when attacking alone than when attacking together with the chasing companion (P=0.033). In these tests, the higher the attack severity, the shorter was the attack latency (P=0.006). A sequence analysis on chasing-companion tests showed that test dogs generally started with observing or showing interest in sheep, followed by attacks, which increased in severity. No sex differences were observed. In non-chasing companion tests, dogs aged up to 2.5 years exhibited a weaker initial hunting motivation than older dogs (P=0.025), but during the entire test were more inclined to attack (P=0.019). Taken together, our findings indicate that a companion dog showing intentions of predatory behaviour stimulates predatory chase in another dog, while a non-chasing companion has a limited influence on this. In tests certifying dogs for their refrainment of chasing sheep, well trained Border collies approaching sheep on command might be used to reveal the full predatory potential of the dog being tested.

Behavioural differences between three breed groups of hunting dogs confronted with domestic sheep.

When running free in open fields, domestic dogs occasionally display predatory behaviour towards domestic sheep. This has not yet been studied scientifically. The aim of the present study was to investigate the inclination to chase sheep in three breed groups of hunting dogs that are most frequently used in areas with grazing sheep. We studied 41 elkhounds, 29 hare hunting dogs and 68 English setters. Behaviours indicative of motivation for chasing or attacking sheep were examined in three different ways. A path test examined functional traits such as hunting ability, contact willingness, reactivity to sudden noise, and response towards a lone sheep. In a sheep confrontation test, loose-leashed dogs were observed in a fenced enclosure with sheep and given electric shocks through an electronic dog collar if within 1-2m from the sheep. A questionnaire to the dog owners supplied information on their dog's previous experience with sheep and behavioural responses to various types of novel stimuli. No significant sex differences were found. The elkhounds showed the highest interest in a lone sheep in the path test, and displayed the highest initial hunting motivation, the highest percentage of dogs starting a sheep attack, the highest attack severity, and were most frequently given el. shocks. The hare hunting dogs were intermediate, while setters showed the lowest values for these variables. Dogs reported as showing low fearfulness more frequently acted as potential sheep chasers in the tests. Dogs up to 3 years of age showed a more pronounced initial hunting motivation and more frequent attacks than older dogs, although there were no age differences in the number of el. shocks given in the test. The latter may be related to the more frequent abruption of attacks among younger dogs. The main factors predicting a high hunting motivation and attack severity were lack of previous opportunity to chase sheep, low fearfulness towards gunshots and unfamiliar people, and general interest in sheep shown when encountering them. Probability of sheep chasing differed between dog breeds and age groups. Previous experience and certain character traits were indicative of a high predatory motivation towards sheep.

Behavioural changes and aversive conditioning in hunting dogs by the second-year confrontation with domestic sheep.

Domesticated dogs occasionally exhibit predatory behaviour towards domestic sheep when running loose in pasture. Both young and old dogs of either sex may chase sheep. Electronic dog collars applying electric shocks are utilised as one method of training dogs to refrain from attacking sheep. This device is used for a number of other training purposes which have raised concern for the welfare of the dogs being trained. This study aims at testing long-term learning effects of previous sheep tests on sheep chasing in hunting dog breeds (Norwegian elkhounds (grey), English setters, and hare hunting dogs), in particular with use of electronic dog collars, in addition to uncovering potential secondary negative effects on dogs' behaviour and mental stability. The dogs (N=114) were subjected to three tests for two subsequent years, the second year being reported here. Dogs were tested for reactions to different stimuli, including a sheep, in a path test. In a sheep confrontation test, dogs were fenced in with a sheep group and given el. shocks when approaching 1-2m from sheep. A questionnaire to the dog owners reported differences in dogs' behaviour between the years.Dogs showed weaker or delayed behavioural responses in both tests in the second year. No dogs showed interest in or attacked a lone sheep in the path test in the second year, while almost two thirds of them did so the first year. In the sheep confrontation test, the dogs exhibited comparatively hesitant initial hunting motivation the second year, being more evident in dogs which received el. shocks the first year. No dogs chased or attacked sheep as their first response in this test, while half of them did so the first year. The proportion of dogs attacking sheep during the entire test was reduced to almost one fourth. The number of el. shocks administered reduced by the second year, and only one of the dogs that received el. shocks the first year received el. shocks the second year. The owners reported no negative effect on the dogs' behaviour during the year ensuing el. shock treatment. Eighteen of the 24 dogs reported by owners to exhibit behavioural changes lost their previous interest in sheep.The second-year tests indicate that aversive conditioning with the use of electronic dog collar may be an efficient method for reducing the probability of a dog chasing or attacking grazing sheep. No adverse effects were observed with our test procedure.

Variation in behavioural responses of ewes towards predator-related stimuli.

Thirty-two groups of six sheep, classified into three breed categories according to their weight class (L: light, one breed (n=7); M: medium light, two breeds (n=10); H: heavy, three breeds (n=15)) were tested for antipredatory behaviour towards seven stimulus regimes. Tests were performed on 2-5-years-old ewes with lambs inside standardised and fenced home pastures. Stimulus regimes were: man in rain poncho, trolley, ball on trolley (blind stimuli), stuffed wolverine on trolley, stuffed lynx on trolley, stuffed bear on trolley, and man in rain poncho with a dog on leash (carnivore stimuli). The L breed showed the longest recovery time, the longest flight distance and the tightest flocking behaviour. Significant regressions indicate that there were negative linear relationships between sheep weight and recovery time and between sheep weight and flight distance. Carnivore stimuli caused longer recovery times (P<0.001) and longer flight distances (P<0.001) than the blind stimuli. Our results confirm the hypothesis that lighter sheep breeds display stronger antipredatory reactions than heavier breeds.

[Effect of prenatal stress on the pituitary-adrenal axis in blue foxes]. / Vliianie prenatal'nogo stressa na gipofizarno-nadpochechnikovuiu sistemu u pestsov.

Handling is a source of stress for farm bred blue foxes. The influence of handling during the late gestation period on the pituitary--adrenal axis was studied in 10-day old male and female blue foxes. Cortisol and progesterone were measured by radioimmunoassay in the plasma, adrenal homogenates, and in vitro incubates from animals of both sexes. Adrenals were incubated in vitro in the absence or presence of ACTH. In addition, the adrenal weight and plasma concentration of ACTH were assessed. In cubs of both sexes, the adrenal weight was decreased after prenatal stress. The plasma concentration of progesterone and the adrenal cortisol in vitro production were elevated in the prenatally stressed female cubs, as compared to the control, along with the adrenal progesterone in vitro production in prenatally stressed male cubs. The adrenal cortisol and progesterone content and plasma ACTH and cortisol concentrations were not affected by prenatal stress. In conclusion, the results of this study suggest that the prenatal stress induced by handling pregnant vixens can affect the pituitary--adrenal axis in neonatal offspring, this effect being more pronounced in female cubs.

[The effect of prenatal stress on steroidogenesis in the gonads of Arctic foxes (Alopex lagopus)]. / Vliianie prenatal'nogo stressa na steroidogenez v gonadakh pestsov.

Handling is a source of stress for farm-bred blue foxes. The influence of handling during the late gestational period was investigated in 10-day old male and female blue foxes. Testosterone and estradiol were measured by RIA in the plasma, gonadal homogenates and in vitro incubates from blue foxes of both sexes. The gonads were incubated in vitro without or with human chorionic gonadotropin. In cubs of both sexes, the gonad weights and ovarian estradiol production were decreased by stress. The testicular testosterone and ovarian estradiol contents were increased in prenatally stressed cubs as compared to the controls. The testicular content and baseline in vitro production of testosterone were not affected by prenatal stress, but the testicular response to human chorionic gonadotropin was higher in the stressed group. This study suggests that prenatal stress induced by handling pregnant vixens may influence gonadal steroidogenesis and this effect was more pronounced in female cubs.

Attitudes to animal-assisted therapy with farm animals among health staff and farmers.

Green care is a concept that involves the use of farm animals, plants, gardens or the landscape in cooperation with health institutions for different target groups of clients. The present study aimed at examining psychiatric therapists' (n = 60) and farmers' (n = 15) knowledge, experience and attitudes to Green care and animal-assisted therapy (AAT) with farm animals for people with psychiatric disorders. Most respondents had some or large knowledge about Green care, but experience with Green care was generally low in both groups. Both farmers and therapists believed that AAT with farm animals could contribute positively to therapy to a large or very large extent, with farmers being significantly more positive. Most of the therapists thought that AAT with farm animals contributes to increased skills in interactions with other humans, with female therapists being more positive than males. Two-thirds of the therapists believed that AAT with farm animals to a large extent could contribute better to mental health than other types of occupational therapy. There were no differences in attitudes to AAT between psychiatrists/psychologists and psychiatric nurses. This study confirms the marked potential of offering AAT services with farm animals for psychiatric patients by documenting positive attitudes to it among psychiatric therapists.

Eye-opening in kittens: effects of light and some biological factors.

Individual variation in the age of eye-opening was studied in 158 kittens. The abiotic and the biotic environments affected eye-opening. Eyes opened earlier in total darkness than in light, and earlier in kittens of young mothers than those of older mothers. Female kittens tended to open their eyes earlier than males, and a paternity effect appeared.

Behavioural differences between laying hen populations selected for high and low efficiency of food utilisation.

1. The effects on behaviour of selecting light-hybrid laying hens for high and low efficiency of food utilisation were investigated. Efficiency of food utilisation was measured as the proportional deviation of observed food consumption from expected food consumption. 2. Videograms of the day-time behaviour of 48 to 53-week-old individually caged hens from the F3 generation were analysed by instantaneous sampling. 3. Low-efficiency hens spent more time food-pecking, walking, pacing, and showing escape and aggressive behaviour than did high-efficiency hens. High-efficiency hens spent more than twice as much time resting and sleeping, and were never observed pacing prior to laying. 4. These differences were larger during the 2 h before laying than at other times of day. 5. Low-efficiency hens had poorer plumage, especially on the neck and the breast, and the poorer the plumage the more agitation they showed. 6. Time spent food-pecking was negatively correlated with laying frequency. 7. Differences in the occurrence of specific behaviour patterns, particularly those symptomatic of pre-laying frustration, partly explained differences in efficiency of food utilisation. Selection for higher efficiency of food utilisation might eliminate hens which are most frustrated prior to laying in cages.

Development of spatial receptive-field organization and orientation selectivity in kitten striate cortex.

The functional organization of the receptive field of neurons in striate cortex of kittens from 8 days to 3 mo of age was studied by extracellular recordings. A quantitative dual-stimulus technique was used, which allowed for analysis of both enhancement and suppression zones in the receptive field. Furthermore the development of orientation selectivity was studied quantitatively in the same cells. Already in the youngest kittens the receptive fields were spatially organized like adult fields, with a central zone and adjacent flanks that responded in opposite manner to the light stimulus. The relative suppression in the subzones was as strong as in adult cells. Both simple and complex cells were found from 8 days. The receptive fields were like magnified adult fields. The width of the dominant discharge-field zone and the distance between the positions giving maximum discharge and maximum suppression decreased with age in the same proportions. The decrease could be explained by a corresponding decrease of the receptive-field-center size of retinal ganglion cells. Forty percent of the cells were orientation selective before 2 wk, and the fraction increased to 94% at 4 wk. Cells whose responses could be attenuated to at least half of the maximal response by changes of slit orientation were termed orientation selective. The half-width of the orientation-tuning curves narrowed during the first 5 wk, and this change was most marked in simple cells. The ability of the cells to discriminate between orientations in statistical terms was weak in the youngest kittens due to a large response variability, and showed a more pronounced development than the half-width did. The orientation-tuning curves were fitted by an exponential function, which showed the shape to be adultlike in all age groups. Two kittens were dark reared until recording at 1 mo of age. The spatial receptive-field organization and the orientation selectivity in these kittens were similar to normal-reared kittens at 1 mo. The responsivity of the cells of the dark-reared kittens was lower, and the latency before firing was longer than in the normal-reared kittens of the same age, and these response properties were more similar to those in 1- to 2-wk-old normal kittens. Our results indicate that the spatial organization of the receptive field is innate in most cells and that visual experience is unnecessary for the organization to be maintained and for the receptive-field width to mature during the first month postnatally.(ABSTRACT TRUNCATED AT 400 WORDS)

Handling during pregnancy in the blue fox (Alopex lagopus): the influence on the fetal pituitary-adrenal axis.

Previous studies revealed that handling is a stressor for farmed blue foxes. The present study was designed to examine the effects of a 1-min daily handling stress applied to pregnant blue fox vixens on the function of the fetal pituitary-adrenal system. Plasma concentrations of adrenocorticotropin hormone (ACTH), cortisol, and progesterone, adrenal content of cortisol and progesterone, in vitro adrenal production of these steroids and response to ACTH, and adrenal weights were measured in control (C; n = 73) and stressed (S; n = 58) fetuses. The ACTH levels were lower in stressed fetuses than in the controls (C: males, 128.6 +/- 6.1 pg/ml; females, 165.9 +/- 6.1 pg/ml; S: males, 122.3 +/- 5.4 pg/ml; females, 145.0 +/- 8.1 pg/ml; P < 0.05). In contrast, increased plasma cortisol concentrations in both sexes were demonstrated in stressed compared with control fetuses (C: males, 9.2 +/- 0.4 ng/ml; females, 9.2 +/- 0.4 ng/ml; S: males, 11.8 +/- 0.7 ng/ml; females, 13.2 +/- 0.7 ng/ml; P < 0.00001). The same difference was observed in plasma progesterone concentrations (C: males, 1.54 +/- 0.07 ng/ml; females, 1.49 +/- 0.10 ng/ml; S: males, 1.86 +/- 0.11 ng/ml; females, 1.74 +/- 0.10 ng/ml; P < 0.01). Prenatal stress did not change the baseline adrenal production of cortisol but prevented the cortisol response to ACTH in female fetuses and decreased the progesterone production in both sexes. Additionally, prenatally stressed fetuses of both sexes had significantly lower adrenal weights than controls (C: males, 9.4 +/- 0.3 mg; females, 9.5 +/- 0.4 mg; S: males, 8.1 +/- 0.3 mg; females, 8.2 +/- 0.4 mg; P < 0.001). These results indicate that prenatal handling stress induces a dysregulation of the pituitary-adrenal axis in the fetus and suggest that increased plasma glucocorticoids in the stressed dam can cross the placenta and influence the fetal hypothalamicpituitary-adrenal axis.