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Biomedical subjects

Svante Winberg

Publications and source records attributed to Svante Winberg.

At least 19 recordsLinked to original sources

Interactions between the neural regulation of stress and aggression.

Socially aggressive interaction is stressful. What is more, social aggression is stressful for both dominant and subordinate animals. Much of the neurocircuitry for stress and aggression overlap. The pattern of neurochemical and hormonal events stimulated by social interaction make it clear that subtle differences in this pattern of response distinguish social rank. The neurotransmitter serotonin (5-HT) responds rapidly to stress, and also appears to play the most important role for inhibitory regulation of aggressive interactions. In addition, the adrenocortical/interrenal steroid hormones corticosterone and cortisol are responsive to stress and influence aggression. However, while 5-HT and glucocorticoids can both be inhibitory to aggression, the relationship between 5-HT and glucocorticoids is not straightforward, and much of the distinctions in function depend upon timing. Neither is inhibitory during the early stressful phase of aggression. This transmitter-hormone combination follows and influences a four-stage functional pattern of effect: (1) predisposed (positively or negatively) toward aggression, (2) motivated toward behavior, (3) responsive to stress (including aggression) and passively allowing aggression, and finally (4) chronically applied 5-HT and glucocorticoids inhibit aggression.

Aggression↗

Does individual variation in stress responses and agonistic behavior reflect divergent stress coping strategies in juvenile rainbow trout?

Individual rainbow trout were transferred to visual isolation in experimental aquaria. As a measure of the speed of acclimation, individual food intake was quantified during the first 6 d following transfer. Following acclimation, aggression was quantified by subjecting the fish to three resident-intruder tests, with 30 d of recovery between the tests. Moreover, between the resident-intruder tests (i.e., two times) the fish were exposed to an unfamiliar environment and their cortisol response was measured. The results of this study show that individuals of juvenile rainbow trout differ distinctly in their response to changes in their environment, and that this diversity in behavior is reflected by consistent behavioral traits displayed by individual fish. These traits have proven to be consistent not only over time but also across situations, revealing two distinct behavioral profiles, in the same manner as shown in studies on proactive and reactive mammals. Our results also show that the reactivity of the hypothalamic-pituitary-interrenal (HPI) axis, when exposed to a stressor, is a consistent physiological trait in juvenile rainbow trout. We found that difference in HPI axis reactivity is linked to the different behavioral profiles. However, HPI axis reactivity could not be linked directly to the singular behavioral traits measured. In other words, we did not find that the consistent behavioral traits shown by the fish were associated with a difference in HPI axis reactivity in the same manner as the reactivity of the hypothalamic-pituitary-adrenocortical axis does in mammals. Taken together, our results show that stress coping strategies akin to what has been described as reactive and proactive stress coping in mammals appear to exist in juvenile rainbow trout.

Adaptation, Psychological↗

Does pulsatile urea excretion serve as a social signal in the gulf toadfish Opsanus beta?

This study evaluated the hypothesis that the pulsatile excretion of urea by toadfish could serve as a social signal. In the first experiment, physiological parameters were measured in pairs of dominant and subordinate toadfish. Subordinate toadfish had elevated concentrations of circulating plasma cortisol, an effect maintained even after cannulation. In the second experiment, one fish of a pair was injected with 14C-urea, and the occurrence of urea pulses during social encounters was documented. Social status did not influence the order of pulsing, that is, whether a dominant or subordinate fish pulsed first during a social encounter. However, in seven out of eight pairs, both toadfish pulsed within 2 h of each other, indicating some form of communication between fish. In the third and final experiment, the response of toadfish to urea (natural or synthetic) was observed. There was a tendency for toadfish to avoid synthetic urea but there was no apparent behavioural response to water containing toadfish urea. Pulsing events do not appear to play an integral role during social encounters as previously hypothesised, but the close timing of pulses in toadfish pairs suggests some transfer of information.

Aggression↗

Avoidance behavior and brain monoamines in fish.

The crucian carp performs a typical avoidance behavior when exposed to olfactory cues from injured skin of conspecifics. They swim rapidly to the bottom and hide in available material. This work examines the effects of skin extract exposure and availability of hiding material on this behavior, and concomitant changes in brain monoaminergic activity in crucian carp. Individual fish were exposed to skin extract in aquaria with or without hiding material. Exposure to skin extract resulted in the expected avoidance behavior consisting of rapid movement towards the bottom of the aquarium. This lasted for 1-2 min. Activity then decreased below the level observed before exposure, suggesting a "freezing" type of avoidance behavior. This behavior was independent of availability of hiding material. Brain dopaminergic activity increased in telencephalon and decreased in the brain stem following skin extract exposure, again independent of availability of hiding material. However, fish kept in aquaria without hiding material showed an elevation of serotonergic activity in the brain stem and the optic tectum compared to fish with available hiding material. Absence of hiding material increased serotonergic activity also without exposure to skin extract. In aquaria with hiding material, the fish stirred up a cloud of fine sediments and showed a more pronounced decrease in locomotor activity in agreement with this being a more efficient freezing or immobile avoidance behavior. These results show that basic components of avoidance behavior and related brain changes are present in the fish brain, in accordance with the common phylogenetic roots of avoidance behavior in all vertebrates.

Analysis of Variance↗

Growth hormone-induced stimulation of swimming and feeding behaviour of rainbow trout is abolished by the D1 dopamine antagonist SCH23390.

The effects of GH on various types of behaviour in fish are well documented although the underlying mechanisms are not fully understood. In rainbow trout, an involvement of the brain dopaminergic system in mediating the behavioural effects of GH has been indicated, as GH can alter the brain dopaminergic activity. To further examine the role of the dopaminergic system in the mediation of GH effects on locomotion and foraging, GH- and sham-implanted juvenile rainbow trout were injected with the selective D1 dopamine antagonist SCH23390 or vehicle. Swimming and feeding activity was then studied by direct observation. Brains were thereafter sampled and analysed for the content of serotonin, dopamine and their metabolites in the hypothalamus, optic tectum, cerebellum, telencephalon, and brain stem. GH increased swimming activity as well as feed intake, effects which were abolished by SCH23390. By itself, the antagonist did not affect behaviour, nor did it affect the brain monoamines. In contrast, treatment with GH, with or without SCH23390, decreased the content of the dopamine metabolite homovanillic acid (HVA) in the optic tectum and the cerebellum, as well as the serotonin content (5-HT) in the optic tectum. It is concluded that the D1 dopamine receptor of the dopaminergic system appears to be of importance in the mediation of the effects of GH on behaviour.

Animals↗

Serotonin, but not melatonin, plays a role in shaping dominant-subordinate relationships and aggression in rainbow trout.

The aim of this study was to clarify to what extent the effects of elevated dietary L-tryptophan (Trp) on aggressive behavior and stress responsiveness in rainbow trout are mediated by circulating melatonin and central serotonin (5-HT), respectively. Isolated rainbow trout were paired for 1h a day for 7 days in order to create fish with experience of being dominant and subordinate. Following this week, the fish were tested for aggressive behavior using a resident-intruder test after which they were subjected to one of four treatments: (1) tryptophan, (2) the selective serotonin reuptake inhibitor (SSRI) citalopram, (3) melatonin, and (4) no treatment (controls). After 7 days of treatment, the fish were subjected to a second resident-intruder test. Trp-supplemented feed resulted in a suppression of aggressive behavior in fish with experience of being dominant. Moreover, fish fed Trp-supplemented feed, regardless of social experience, also displayed lower plasma cortisol levels than controls. These effects of elevated dietary Trp were closely mimicked by citalopram treatment, whereas exogenous melatonin had no effect on either aggressive behavior or plasma cortisol. Thus, the effect of elevated dietary Trp on aggressive behavior and stress responses does not appear to be mediated by melatonin even though elevated dietary intake of Trp resulted in an increase in plasma melatonin concentrations.

Aggression↗

Divergence in behavioural responses to stress in two strains of rainbow trout (Oncorhynchus mykiss) with contrasting stress responsiveness.

The aim of this study was to establish whether two lines of rainbow trout divergent for their plasma cortisol response to a standardized stressor would show consistent differences in their behavioural response to a range of challenging situations. Our results show that the high- and low-responding (HR and LR) lines of rainbow trout did not differ in the aggression shown towards an intruder or in their response to the introduction of a novel object to their home environment. However, there was a difference in behaviour between the two selection lines when they were exposed to two unfamiliar environments. These results suggest that the behaviour of the HR and LR fish differs when they are challenged in unfamiliar environments, while their behaviour does not differ when they are challenged in their home environment. These observations are in agreement with studies on mammals that show that individuals with reactive coping styles perform similarly to proactive animals when they are challenged in a familiar environment, while they show different behaviour when they are challenged in unfamiliar environments. Thus, these results provide further evidence that the HR and LR selection lines of rainbow trout exemplify the two different coping styles described in mammals.

Aggression↗

Tryptophan affects both gastrointestinal melatonin production and interrenal activity in stressed and nonstressed rainbow trout.

The present experiments were designed to test the hypothesis that elevated dietary levels of l-tryptophan (Trp) result in elevated plasma levels of melatonin and that this increase in plasma melatonin concentration is caused by elevated melatonin production and secretion by the gastro-intestinal-tract (GIT). Feeding juvenile rainbow trout (Oncorhynchus mykiss) Trp-supplemented feed for 7 days resulted in elevated daytime plasma levels of melatonin and reduced poststress plasma cortisol concentrations. Nighttime plasma melatonin concentrations were, however, not affected by elevated dietary Trp. Moreover, stress caused a reduction in daytime plasma levels of melatonin in fish fed Trp-supplemented feed, an effect that was counteracted by treatment with an alpha-receptor antagonist. These results clearly suggest that elevated dietary intake of Trp results in an increase in the GIT production of melatonin in rainbow trout. A suggestion that was further supported by the results from an in vitro experiment demonstrating that addition of Trp to the incubation medium stimulates melatonin production and release by incubated rainbow trout GIT. The results from this study led us to suggest a possible mechanism for melatonin in mediating the effects of elevated dietary Trp on poststress plasma cortisol concentrations and aggressive behavior in rainbow trout.

Animals↗

Divergent stress coping styles in juvenile brown trout (Salmo trutta).

Two distinct stress coping styles, proactive and reactive, have been stated in various animal studies. This divergence in coping has also been indicated in salmonid fish. Here, we test the hypothesis that divergent stress coping styles are identifiable in a sea-ranched brown trout population. To that end, we used a series of tests on individual juvenile brown trout, with each test including a common key aspect of the two different coping styles. Using a clustering method (SAS: PROC FASTCLUS), two groups that clearly differed both in blood chemistry (noradrenalin and adrenalin levels) following confinement and in behavior during hypoxia were identified.

Adaptation, Psychological↗

Socially-mediated differences in brain monoamines in rainbow trout: effects of trace metal contaminants.

Monoaminergic systems play a crucial role in linking behaviour and physiology. Here the physiological and behavioural effects of metal exposure in relation to monoaminergic systems were considered by exposing rainbow trout dyads, demonstrating stable dominance relationships, to cadmium or lead. Fish exposed to 4 microg l(-1) cadmium accumulated more cadmium at the gill than fish held in control water. Fish exposed to 7 microg l(-1) cadmium had higher gill, liver and kidney cadmium concentrations. No significant lead accumulation was seen after exposure to 46 microg l(-1) for 48 h but exposure to 325 microg l(-1) lead caused an increase in gill, liver and kidney lead concentrations. Brain accumulation of both cadmium and lead was only seen after exposure to the highest concentrations. Exposure to 4 or 7 microg l(-1) cadmium, or 46 or 325 microg l(-1) lead for 48 h did not disrupt established dominance hierarchies. As expected with this stable behavioural situation, in control pairs, animals of different social status displayed different physiological profiles. Subordinate fish had higher concentrations of circulating plasma cortisol and telencephalic 5-hydroxyindoleacetic acid/5-hydroxytryptamine (serotonin) (5-HIAA/5-HT) ratios. However, these physiological profiles were affected by metal exposure, with a trend towards higher serotonergic activity in dominant fish. Dominants exposed to 325 microg l(-1) lead had significantly higher hypothalamic 5-HIAA/5-HT ratios when compared with subordinates. The results demonstrate that if stable social hierarchies are established in control water they may not be affected by exposure to cadmium and lead although physiological changes may be evident.

Analysis of Variance↗

Social stress affects circulating melatonin levels in rainbow trout.

In salmonid fishes there are indications that socially subordinate individuals avoid competition with larger, dominant fish by adjusting daily feeding and activity cycles. As in other vertebrates, the pineal organ and its hormone melatonin act as synchronizers of daily rhythms to the external light/dark cycle in salmonids. Social defeat may act as a potent stressor; inducing elevated glucocorticoid secretion and a general behavioral inhibition. Here, we show that social stress also affects circulating melatonin levels in rainbow trout, a species known to display strong dominance hierarchies both in the wild and under captive rearing. Subordinate individuals had significantly higher nighttime melatonin levels than dominant fish or controls. There was no effect of social rank on the much lower melatonin levels observed in animals sampled during the day. Correlations between circulating glucocorticoids and melatonin depended on circadian cycles as well as social context. This study suggests that altered melatonin production contributes to the physiological and behavioral profile of subordinate animals. Social status, and other determinants of the stress level of experimental animals, therefore should be taken into consideration as potential factors influencing the results from in vivo research on this hormone.

Agonistic Behavior↗

Stress coping style predicts aggression and social dominance in rainbow trout.

Social stress is frequently used as a model for studying the neuroendocrine mechanisms underlying stress-induced behavioral inhibition, depression, and fear conditioning. It has previously been shown that social subordination may result in increased glucocorticoid release and changes in brain signaling systems. However, it is still an open question which neuroendocrine and behavioral differences are causes, and which are consequences of social status. Using juvenile rainbow trout of similar size and with no apparent differences in social history, we demonstrate that the ability to win fights for social dominance can be predicted from the duration of a behavioral response to stress, in this case appetite inhibition after transfer to a new environment. Moreover, stress responsiveness in terms of confinement-induced changes in plasma cortisol was negatively correlated to aggressive behavior. Fish that exhibited lower cortisol responses to a standardized confinement test were markedly more aggressive when being placed in a dominant social position later in the study. These findings support the view that distinct behavioral-physiological stress coping styles are present in teleost fish, and these coping characteristics influence both social rank and levels of aggression.

Adaptation, Psychological↗

Behavioral and neuroendocrine correlates of displaced aggression in trout.

In humans and other primates, violent actions performed by victims of aggression are often directed toward an individual or object that is not the source of provocation. This psychological phenomenon is often called displaced aggression. We demonstrate that displaced aggression is either rooted in evolutionarily conserved behavioral and neuroendocrine mechanisms, or represent a convergent pattern that has arisen independently in fish and mammals. Rainbow trout that briefly encountered large, aggressive fish reacted with increased aggression toward smaller individuals. There was a strong negative correlation between received aggression and behavioral change: Individuals subjected to intense aggression were subdued, while moderate assaults induced strong agitation. Patterns of forebrain serotonin turnover and plasma cortisol suggest that the presence of socially subordinate fish had an inhibitory effect on neuroendocrine stress responses. Thus, subordinate individuals may serve as stress-reducing means of aggressive outlet, and displaced aggression toward such individuals appears to be a behavioral stress coping strategy in fishes.

Aggression↗

Peripherally administered growth hormone increases brain dopaminergic activity and swimming in rainbow trout.

There is increasing evidence that growth hormone (GH) has important behavioral effects in fish, but the underlying mechanisms are not well understood. To investigate if peripherally administered GH influences the monoaminergic activity of the brain, and how this is correlated to behavior, juvenile rainbow trout were implanted intraperitoneally with ovine GH. Fish were either kept isolated or in groups of five. The physical activity and food intake of the isolated fish were observed after 1 and 7 days, when brains were also sampled. The content of serotonin, dopamine, and noradrenaline and their metabolites in hypothalamus, telencephalon, optic tectum, and brain stem was then analyzed. For fish kept isolated for 7 days following implant, GH increased swimming activity and the levels of the dopamine metabolite 3, 4-hydroxy-phenylacetic acid (DOPAC) were higher in all brain parts examined. In the optic tectum, the levels of the dopamine metabolite homovanillic acid (HVA) were lowered by the GH treatment. One-day GH implant did not affect behavior or monoamine levels of isolated fish. In the fish kept in groups, a 7-day GH implant increased the hypothalamic levels of DOPAC, but not in the other brain parts examined, which may indicate an effect on the brain dopaminergic system from social interactions. It can be concluded that peripherally administered GH may function as a neuromodulator, affecting the dopaminergic activity of the rainbow trout brain, and this is associated with increased swimming activity.

3,4-Dihydroxyphenylacetic Acid↗

Central nervous system actions of growth hormone on brain monoamine levels and behavior of juvenile rainbow trout.

Growth hormone (GH) has been demonstrated to alter the behavior of juvenile salmonids. However, the mechanisms behind this action are not yet understood. In mammals and birds, peripheral GH treatment has been shown to affect monoaminergic activity in the central nervous system, which may be a mechanism whereby GH alters behavior. To investigate if GH may influence behavior directly at the central nervous system, juvenile rainbow trout were injected with GH into the third ventricle of the brain, whereupon physical activity and food intake were observed during 2 h. Thereafter, brains were sampled and the content of serotonin, dopamine, and noradrenaline and their metabolites were measured in hypothalamus, telencephalon, optic tectum, and brainstem. The GH-treated fish increased their swimming activity relative to sham-injected controls, while appetite remained unchanged, compared with sham-injected controls. Analysis of brain content of monoamines revealed that the GH treatment caused a decrease in the dopamine metabolite homovanillic acid in the hypothalamus, indicating a lowered dopaminergic activity. It is concluded that GH may alter behavior by acting directly on the central nervous system in juvenile rainbow trout. Furthermore, GH seems to alter the dopaminergic activity in the hypothalamus. Whether this is a mechanism whereby GH affects swimming activity remains to be clarified.

Animals↗

Time-course of the effect of dietary L-tryptophan on plasma cortisol levels in rainbow trout Oncorhynchus mykiss.

Isolated juvenile rainbow trout were fed a feed supplemented with L-tryptophan (TRP) for 3, 7 or 28 days, after which they were either sampled directly (undisturbed) or subjected to a standardised stressor prior to sampling. Controls (stressed and undisturbed) received the same feed but without any supplementary TRP. Stress resulted in a significant elevation of plasma [cortisol] in fish fed control feed and in fish fed TRP-supplemented feed for 3 and 28 days. However, fish fed TRP-supplemented feed for 7 days did not show any significant elevation of plasma [cortisol] in response to stress. Plasma levels of adrenocorticotropin followed the same general pattern as cortisol. Plasma and brain [TRP] were elevated in fish fed TRP-supplemented feed. The amino acid TRP is the precursor of the monoamine neurotransmitter serotonin (5-hydroxytryptamine, 5-HT) and the brain 5-HT system is known to be involved in the control of the hypothalamic-pituitary-interrenal (HPI) axis. Fish fed TRP-supplemented feed showed elevated levels of 5-hydroxyindoleacetic acid (5-HIAA, a major 5-HT metabolite) in the hypothalamus and optic tectum. However, TRP treatment did not appear to result in any effects on brain dopaminergic activity and the effects on brain norepinephric activity do not support a role of norepinephrine in mediating the effects of TRP on HPI axis reactivity in rainbow trout.

Adrenocorticotropic Hormone↗

Stimulatory and inhibitory effects of 5-HT(1A) receptors on adrenocorticotropic hormone and cortisol secretion in a teleost fish, the Arctic charr (Salvelinus alpinus).

The serotonin(1A) (5-HT(1A)) receptor agonist 8-hydroxy-2-(di-N-propylamino)tetralin (8-OH-DPAT) was administrated to juvenile Arctic charr either by intraperitoneal (i.p.) injections or through a permanent i.p. implanted catheter. 8-OH-DPAT induced an elevation of plasma cortisol in non-stressed fish (receiving the drug through a permanent i.p. catheter), whereas 8-OH-DPAT had the opposite effect when administrated through a standard i.p. injection, dampening the rise in cortisol induced by handling and injection, an effect which was most pronounced at low doses of 8-OH-DPAT. In both cases, plasma levels of adrenocorticotropic hormone followed the same general pattern as plasma cortisol. The results suggests that 5-HT(1A) receptors are involved in the regulation of hypothalamic-pituitary-interrenal axis activity in Arctic charr, but the effect of 5-HT(1A) stimulation seems to vary depending on the state of the fish.

8-Hydroxy-2-(di-n-propylamino)tetralin↗

Effects of cortisol on aggression and locomotor activity in rainbow trout.

Noninvasive administration of cortisol through the diet resulted in relatively rapid (<1.5 h) and highly reproducible increases in plasma cortisol in rainbow trout, comparable to changes seen in fish subjected to substantial stress. Juvenile rainbow trout were reared in isolation for 1 week, before their daily food ration was replaced by a meal of cortisol-treated food corresponding to 6 mg cortisol kg(-1). All fish were observed for 30 min, beginning at 1 or 48 h following the introduction of cortisol-treated food. Additional cortisol (75% of the original dose on Day 2, and 50% on Day 3) was administered to the long-term cortisol-treated group. The resulting blood plasma concentrations of cortisol were similar in short- and long-term treated fish, and corresponded to those previously seen in stressed rainbow trout. Controls were fed similar food without cortisol. Half of the fish from each treatment group (controls and short- and long-term cortisol) were subjected to an intruder test (a smaller conspecific introduced into the aquarium), while half of the fish were observed in isolation. In fish challenged by a conspecific intruder, short-term cortisol treatment stimulated locomotor activity, while long-term treatment inhibited locomotion. Aggressive behavior was also inhibited by long-term cortisol treatment, but not by short-term exposure to cortisol. Cortisol treatment had no effect on locomotor activity in undisturbed fish, indicating that the behavioral effects of cortisol were mediated through interaction with other signal systems activated during the simulated territorial intrusion test. This study demonstrates for the first time that cortisol has time- and context-dependent effects on behavior in teleost fish.

Aggression↗