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Sexual segregation in infant mice: behavioural and neuroendocrine responses to d-amphetamine administration.

Individual differences arise from both genetic and epigenetic factors. The aim of this study was to test whether pups raised in distinct socio-sexual conditions would show different behavioural and neuroendocrine responses to d-amphetamine (AMPH) administration upon placement in a novel environment. This issue was addressed by testing infant CD-1 mouse pups of both sexes at three different developmental ages [3, 8, or 18 postnatal (PND) days]. These pups were raised from birth in all-male, all-female, or mixed-sex litters. AMPH effects were assessed as a function of the hypothalamic-pituitary-adrenal (HPA) axis activational state using litters that were either maternally deprived for 24 h (DEP) or normally kept with the dam (NDEP). A concomitant maternal behaviour score carried out on selected postpartum days showed that mothers taking care of all-male litters were more often involved in Active nursing than those rearing the mixed-sex ones, whereas the latter were found more often Laying still out of the nest. Basal and stress-induced corticosterone (CORT) secretion was increased in unisexually reared pups following maternal deprivation, an effect limited to PND 3. In general, neuroendocrine and behavioural responses to AMPH were found to be dissociated and were affected by sexual segregation only in conjunction with maternal deprivation. On PND 3, AMPH injection (1 or 3 mg/kg, i.p.) decreased CORT secretion in deprived unisexually reared subjects without affecting their behaviour. As a whole, behavioural changes due to unisexual rearing were limited to female subjects. On PND 8, unisexually reared females showed, upon maternal deprivation, a generalized shift to the left in the dose-response curve to AMPH for Crossing behaviour, while on PND 18 AMPH-induced stereotypies were considerably reduced in sexually segregated females, especially following maternal deprivation. Thus, maternal deprivation appeared to "sensitize" the monoaminergic system to an AMPH challenge. The individual behavioural and neuroendocrine profiles shown in response to a stressful challenge suggest that changes in social stimulation early during development might produce subtle shifts in the function of selected central monoaminergic systems.

Aging↗

CCK(B) antagonists protect against anxiety-related behaviour produced by ethanol withdrawal, measured using the elevated plus maze.

The effects of the CCK(B) antagonists, CAM1028 and CI988 and a CCK(A) antagonist, CAM1481, were studied on the anxiety-related behaviour produced by withdrawal from chronic ethanol treatment, using the elevated plus maze. Cessation of chronic ethanol administration produced a profile, in both mice and rats, consistent with increase in anxiety-related behaviour. In mice, SC administration of CAM1028 or CI988 reduced the decrease in the time spent on the open arms, the number of entries into these arms and the increases in the latencies to first open arm entry, after withdrawal from the ethanol treatment. The increases in stretched attend postures and head dips from the closed arms and the central square seen during the withdrawal phase, were also decreased by the CCK(B) antagonists, but the decreases in the number of rears and in general activity were unaffected. The doses of CAM1028 and CI988 tested were 0.1 and 1 mg/kg; for some of the withdrawal-induced changes in behaviour only the 1 mg/kg dose was effective. In contrast, the CCK(A) antagonist, CAM1481, at the same doses, had little effect on the anxiety-related behaviour produced by withdrawal from chronic ethanol treatment, although it did decrease the changes in the number of rears and the head dipping behaviour. In rats, the majority of the changes produced by withdrawal from chronic ethanol treatment were decreased by CAM1028 at 1 mg/kg, although the decreases in open arm entries, rearing behaviour and in overall activity were unaffected. CAM1028, CI988 and CAM1481 had no effects on the behaviour of control mice or rats in the plus-maze. The results show that CCK(B) antagonists were effective in decreasing the majority of the anxiogenic effects of withdrawal from chronic ethanol treatment.

Animals↗

Social behaviour in rats lesioned with ibotenic acid in the hippocampus: quantitative and qualitative analysis.

RATIONALE: Neonatal ibotenic acid lesion of the ventral hippocampus was proposed as a relevant animal model of schizophrenia reflecting positive as well as negative symptoms of this disease. Before and after reaching maturity, specific alterations in the animals' social behaviour were found. OBJECTIVE: In this study, social behaviour of ventral hippocampal lesioned rats was analysed. For comparison, rats lesioned either in the ventral hippocampus or the dorsal hippocampus at the age of 8 weeks were tested. METHODS: Rats on day 7 of age were lesioned with ibotenic acid in the ventral hippocampus and social behaviour was tested at the age of 13 weeks. For comparison, adult 8-week-old rats were lesioned either in the ventral or the dorsal hippocampus. Their social behaviour was tested at the age of 18 weeks. RESULTS: It was found that neonatal lesion resulted in significantly decreased time spent in social interaction and an enhanced level of aggressive behaviour. This shift is not due to anxiety because we could not find differences between control rats and lesioned rats in the elevated plus-maze. Lesion in the ventral and dorsal hippocampus, respectively, in 8-week-old rats did not affect social behaviour. CONCLUSIONS: The results of our study indicate that ibotenic acid-induced hippocampal damage per se is not related to the shift in social behaviour. We favour the hypothesis that these changes are due to lesion-induced impairments in neurodevelopmental processes at an early stage of ontogenesis.

Aggression↗

Cognition and behaviour are independent and heterogeneous dimensions in Alzheimer's disease.

Clinical expressions of cognition and behaviour in Alzheimer's disease (AD) patients are heterogeneous. Therefore, assessing the entire range of selective cognitive and behavioural characteristics of dementia in minute detail is extremely important. However, considering that groups of different symptoms may respond to the same pharmacological agent, it is also evident that a correct evaluation of the behaviour requires the grouping of symptoms in fewer syndromes. Thus, the authors have analysed various connections between selective cognitive domains and behavioural symptoms (BPSD) in probable AD outpatients. Two hundred and forty four patients with diagnosis of probable AD, according to DSM-IV and NINCDS-ADRDA criteria were enrolled. The evaluation included the Mini Mental State Examination, the Mental Deterioration Battery, and the Neuropsychiatric Inventory. Treatment with low doses of neuroleptic drugs only was allowed. Principal component analysis condensed the 18 cognitive/behavioural variables in 7 factors namely general-cognitive, constructional abilities, hyperactivity, psychosis, anxiety, mood-excitement and mood-depression/apathy. None of the cognitive domains were included in the behavioural factors and vice-versa. Furthermore, the only BPSD which impaired continuously with progression of disease severity was apathy which was also the most severe symptom. In conclusion, many cognitive and behavioural syndromes exist in patients with AD. However, the results of this study suggest that cognition and behaviour are independent dimensions.

Aged↗

Behavioural strategies used by the hookworms Necator americanus and Ancylostoma duodenale to find, recognize and invade the human host.

The infective third-stage larvae of the hookworms Necator americanus and Ancylostoma duodenale infect their human hosts by active skin invasion, but A. duodenale is in addition capable of oral infection. The behaviour of the larvae when crawling on surfaces has already been described. Here we analyse in various in vitro systems the other behavioural invasion phases: activation, penetration, and orientation within the host. The larvae normally remained in a motionless, energy-saving, resting posture. An activation to sinusoidal locomotion was stimulated in both species by similar cues such as touch, vibration, water currents, heat, light, and chemicals. Human breath in addition stimulated searching and waving ("nictating") behaviour, which facilitates a change-over to the host. Activating cues in air streams were warmth and moisture; CO2 activated only in combination with warmth and/or moisture. Penetration behaviour in both species was stimulated by warmth and skin extracts. The stimulating components of skin extracts were fatty acids, but their stimulating characteristics differed from those inducing schistosome cercarial skin penetration. After penetration into agar substrates, both species showed thermo-orientation, but only A. duodenale followed gradients of serum. The directing serum cues were not amino acids and glucose (the supposed cues for schistosome blood vessel localization), but Ringer's solution attracted the larvae. The host-finding and host-invasion behaviour of both hookworm species is well adapted to the invasion of the human skin, and there seems to be no particular adaptation of A. duodenale behaviour to the oral infection mode. Hookworm host-finding behaviour is not as complex as that of schistosome cercariae but seems well adapted to the ecological conditions in the transmission sites.

Ancylostoma↗

The association of DNA sequence variation at the MAOA genetic locus with quantitative behavioural traits in normal males.

Monoamine oxidase A (MAOA) catalyses the oxidative deamination of biogenic amines including neurotransmitters, mainly norepinephrine and serotonin in the brain and peripheral tissues. A nonsense mutation in the gene was shown to be involved in a rare X-linked behavioural syndrome, which includes impaired impulse control, aggression and borderline mental retardation (Brunner syndrome). Several recent studies have shown the association of genetic variation of a VNTR in the gene promoter with various pathological behavioural traits. In the present study the association of MAOA genetic variation with a large set of quantitative behavioural traits in normal individuals has been examined. DNA samples from 421 unrelated males were genotyped for 14 SNPs and for the promoter VNTR at the MAOA locus. An additional 16 SNPs were genotyped at apparently neutral loci across the X chromosome to serve as a genomic control for possible false positive associations due to population structure. Behavioural traits were measured using the NEO psychometric questionnaire, which is based on a 5-axis model of personality, and consists of 30 different quantitative traits. There was a robust association of the A2 ("straightforwardness") facet with common allelic variants at the promoter VNTR. Most of the tested traits were not associated with the VNTR despite reasonable power, thus demonstrating that the VNTR influence on quantitative behavioural traits in normal males may be very specific. In contrast, several traits of the C ("conscientiousness") axis were associated with less common SNP-defined haplotypes. Hence, it appears that common genetic variation at the VNTR contributes to the behavioural attribute of "straightforwardness", while rare haplotypes defined by SNPs downstream of the transcription start site may contribute to "conscientiousness". This study is used to address the validation, interpretation and limitation of genetic association studies of quantitative behavioural traits.

Black or African American↗

Effects of hypothalamic lesions upon the sexual and social behaviour of the male common marmoset (Callithrix jacchus).

The sexual and associated behaviour of 10 adult male marmosets was recorded during pair tests with ovariectomized females, before and after bilateral thermal lesions of the hypothalamus. Four sham-lesioned males served as controls. Lesions varied in volume from 1.49 to 3.28 mm3 and extended from the ventromedial hypothalamus to the diagonal band of Broca. Precopulatory behaviours (anticipatory erections, tongue flicking and anogenital investigations of the female) as well as frequencies of mounting, intromission and ejaculation decreased in lesioned males. The greatest suppression of sexual behaviour occurred after lesioning the anterior hypothalamus (AH), beneath the anterior commissure, or at the junction of the AH with the preoptic area (POA). Lesions confined to the POA had less profound behavioural effects. Treating ovariectomized females with estradiol stimulated their proceptivity but had no consistent effects upon the males' behaviour. Lesioned males did not exhibit signs of social withdrawal and frequencies of allogrooming or grooming invitations increased post-operatively. Preliminary studies on intermale aggression indicated that lesions which had the greatest effect upon sexual behaviour also tended to decrease aggressive interactions with other males. Hypothalamic lesions did not affect plasma testosterone levels, except in one male and only one animal showed signs of ill health (weight loss and hypothermia) post-operatively. These results show that damage to the AH or AH-POA junction in male marmosets causes a profound suppression of sexual 'arousal' and copulatory behaviour and that such affects are not due to androgen insufficiency or other, non-specific, side effects of neural damage.

Aggression↗

Spatial memory and stereotypic behaviour of animals in radial arm mazes.

The symmetric radial arm maze, described by Olton in 1976, has developed into an important tool for the study of spatial memory. In a typical test an animal is placed in the centre of the maze, which contains some small piece of food at the end of each arm. The sampling behaviour of the animal is then recorded. In such studies the score (number of choices of arms which still contain food) of the animal is normally compared with the score of an imaginary animal which changes arms entirely at random. In a new method of analysis the non-random score of the animal is split into two parts, one depending on memory and one on stereotypic choice behaviour. Even mild departures from randomness are shown to alter considerably the expected 'random' score in an eight-armed maze. The part of the score claimed to depend on memory was shown to increase when the animals learn to search the maze, the stereotypic part did not. The general effect from stereotypic choice behaviour is shown to result, in most animals, in an increase in the total score. In an eight-armed maze this increase may amount to more than 20% of the total non-random score, even in a well-trained animal. The effect is less pronounced in mazes with 16 arms. It has been proposed that hippocampal lesions produce a stereotypic behaviour. We propose, based on our analysis, that the stereotypic behaviour is not produced but revealed by hippocampal lesions which destroy almost completely the memory-guided behaviour masking the stereotypic behaviour in the intact animal.

Animals↗

Detailed analysis of the effects of apomorphine and d-amphetamine on spontaneous locomotor behaviour of rats as measured in a TV-based, automated open-field system.

Rat open field behaviour was measured in a TV-based, automated system. Habituation was evident in saline-treated rats. Most variables measured declined over a 10 min period. Apomorphine affected rat open-field behaviour bimodally, i.e. low doses of apomorphine (0.02-0.08 mg/kg, s.c.) decreased most aspects of rat open-field behaviour, whereas at higher doses (0.2 and 0.5 mg/kg, s.c.) various aspects of open-field behaviour were stimulated. Rearing and average speed, however, were monotonically depressed. Amphetamine (1 and 2 mg/kg) stimulated most aspects of rats open-field behaviour, including rearing. Speed was not affected by amphetamine. Habituation was more pronounced after low doses of apomorphine than after saline treatment and was absent after high doses of apomorphine and after amphetamine. The results represent a detailed analysis of apomorphine and amphetamine effects on rat open-field behaviour and show that the distinct components of this behaviour are affected differentially.

Animals↗

Further analysis of the effects of cholecystokinin octapeptides on avoidance behaviour in rats.

Experiments were performed to examine the acute effects of cholecystokinin octapeptides and fragments on the active and passive avoidance behaviour of rats following peripheral and central administration. Both the sulphated (CCK-8-SE) and non-sulphated cholecystokinin octapeptide (CCK-8-NS) and also the COOH-terminal tetra-, penta-, hexa- and heptapeptides of cholecystokinin octapeptide facilitated the extinction of active avoidance behaviour and retention of passive avoidance behaviour. This latter effect of cholecystokinin octapeptides was reversed by anxiolytic chlordiazepoxide pretreatment, showing that in these test situations cholecystokinin octapeptides are able to modify fear-motivation or arousal of the animals; their effect is at least partly similar to that of the neuroleptic substance haloperidol. Subcutaneous treatment with CCK-8-SE or CCK-8-NS appeared to be 3-10 times more effective than intraperitoneal treatment. Following intracerebroventricular administration, 100-300 times lower doses were needed to cause a behavioural effect similar to that after subcutaneous injection. Microinjection of CCK-8-SE or CCK-8-NS in the fmol dose range into the nucleus accumbens facilitated the extinction of active avoidance behaviour and attenuated the retention of passive avoidance behaviour, while microinjection of these peptides into the central amygdaloid nucleus caused opposite effects on these behavioural tests. It seems that the neuroleptic-like effects of cholecystokinin octapeptides are mediated through the nucleus accumbens, and the opposite action (non neuroleptic-like) through the central amygdaloid nucleus.

Amygdala↗

Hippocampal kindling-induced after-discharge and behavioural depression: immediate and long-term attenuation by opiate antagonists.

Hippocampal kindling results in tonic-clonic convulsions followed by a pronounced period of behavioural depression. The effect of the opiate antagonists, naltrexone and naloxone, on the duration of the after-discharge and behavioural depression was investigated. Naltrexone, injected s.c. 60 min before, significantly reduced the behavioural depression at doses as low as 0.6 mg/kg, but had no effect on the after-discharge even at higher doses. One day later, the behavioural depression was still reduced in some animals and the after-discharge was significantly decreased following the higher doses (0.24-0.48 mg/kg). Naloxone, injected s.c., 10 min before, significantly reduced the behavioural depression at a dose as low as 0.002 mg/kg and also reduced the duration of the after-discharge at some doses. Naloxone did not exert any significant effects 24 h later. Naltrexone, injected i.c. 60 min before kindling in a dose of 1 microgram, significantly attenuated the behavioural depression and had no effect on the after-discharge. The behavioural depression was still attenuated 24 and 48 h later. The involvement of mu-receptor-related endogenous opioid mechanisms in postictal brain processes is suggested. The long-term effects might be related to receptor 'activation' during the immediate period due to an interaction between the antagonist and the kindling experience.

Animals↗

Effect of SCH 23390 and quinpirole on novelty-induced grooming behaviour in spontaneously hypertensive rats and Wistar-Kyoto rats.

Grooming behaviour induced by exposure to a novel environment was studied in spontaneously hypertensive rats (SHR) and normotensive Wistar-Kyoto rats (WKY). The dopamine D1 receptor antagonist, SCH 23390, and the dopamine D2 receptor agonist, quinpirole, were used to study brain dopamine systems in these rat strains, via their effects on grooming behaviour. The total grooming behaviour displayed in a 50-min observation period was significantly lower in SHR than in WKY. Except for the paw licking component no differences between the two strains were observed in the separate behavioural elements of grooming behaviour. SCH 23390 and quinpirole were found to suppress novelty-induced grooming behaviour of both strains. In SHR, grooming behaviour was less suppressed by SCH 23390, whereas the suppression by quinpirole was more pronounced than in WKY. These results indicate that there are alterations in central dopamine systems in SHR, probably involving changes both in dopamine D1 and D2 receptor mechanisms in the brain.

Animals↗

"Adaptive" changes in the behaviour of parasitized animals: a critical review.

Changes in host behaviour following infection with parasites are frequently reported in the literature, and are often hypothesized to be adaptive for either host or parasite. However, investigators of such phenomena often use the "adaptation" label for host behavioural changes based on their intuition and not on rigorous criteria. Alterations in host behaviour following infection can only be considered adaptive if they satisfy certain conditions: (1) they must be complex; (2) they must show signs of a purposive design; (3) they are more likely to be adaptations if they have arisen independently in several lineages of hosts or parasites; and (4) they must be shown to increase the fitness of either the host or the parasite. A survey of published examples of host behavioural changes indicates that while some are spectacularly complex and are extremely well-fitted to their presumed function, most are simple increases or decreases in an activity performed prior to infection. There are some suggestions of convergent evolution in behavioural change in distantly related host or parasite groups but more evidence is needed. Finally, most known behavioural changes have not been demonstrated to lead to fitness gains in either hosts or parasites. Few known examples satisfy more than two of the above criteria, and, in general, the adaptive function of changes in host behaviour following infection is in need of more solid proof.

Adaptation, Psychological↗

Evidence that behavioural and electrocortical sleep induced by guanfacine is due to stimulation of alpha 2-adrenoceptors.

The effects of guanfacine and other drugs acting at alpha 1- and alpha 2-adrenoceptors on behaviour, electrocortical activity and ECoG spectrum power were studied in chicks and rats. Guanfacine, given systematically in chicks, produced behavioural and electrocortical slow-wave sleep lasting 100-200 min, depending on the dose; these effects were prevented by yohimbine, a selective antagonist, at alpha 2-adrenoceptors and potentiated by prazosin, a selective antagonist, at alpha 1-adrenoceptors. Similar behavioural and electrocortical effects were obtained after systemic or intraventricular infusion of guanfacine in rats. In addition, a significant increase in total and in lower frequency band (0-4; 4-8 Hz) voltage power was observed. Behavioural and ECoG effects of guanfacine were prevented by phentolamine or yohimbine, whereas prazosin and propranolol were ineffective. Yohimbine itself, given systemically in chicks, produced behavioural stimulation, vocalization, increase in locomotor activity and ECoG desynchronization, with a significant fall in total and 0-3, 3-6, 6-9 and 9-12 Hz voltage power lasting approx. 3 h. Desipramine, an inhibitor of noradrenaline reuptake, produced in chicks behavioural and ECoG arousal, vocalization, pecking, escape responses and aggressive behaviour. In conclusion, the present experiments show that guanfacine sedative effects seem to be mediated predominantly via an activation of presynaptic alpha 2-adrenoceptors and suggest that arousal is due to stimulation of post-synaptic alpha 1-adrenoceptors.

Animals↗

Observation of overt pain behaviour by physicians during routine clinical examination of patients with low back pain.

Observation of overt pain behaviours was carried out by physicians during routine clinical examination of 120 patients with chronic low back pain. Reliable ratings were achieved but only after very careful standardization in an additional 60 pilot patients. Overt pain behaviour was found to be related to other clinical measures of illness behaviour--pain drawing, behavioural symptoms, behavioural signs, use of walking aides and downtime--but did tap an additional dimension. It is concluded that clinical observation of overt pain behaviour can provide useful additional information about illness behaviour in low back pain. Reliable observations can be achieved in a carefully standardized research situation but in routine clinical practice are vulnerable to considerable observer error and bias.

Adult↗

Short-term behavioural effects of neurohypophyseal hormones: pharmacological characteristics.

Neurohypophyseal hormones and related peptides cause behavioural alterations after intracerebroventricular injection in mice. In the present study, these effects, consisting of excessive grooming and scratching, and of escape-directed activity in stressful situations, could easily be distinguished from those of other centrally acting peptides and drugs by means of two different behavioural bioassays. The effects were not antagonized by drugs that block cholinergic or adrenergic receptors, but they were powerfully suppressed by some potent psychotropic agents. Some compounds with strong vasoconstrictor or vasodilatory actions did not mimick or antagonize the behavioural alterations, suggesting that vasoconstriction is not essential for the induction of these effects. A considerable degree of tolerance could be induced and cross-tolerance was observed between different neurohypophyseal hormones. In rats, behavioural alterations caused by oxytocin and vasopressin could be demonstrated as well, but they were by far less pronounced than those observed in mice. For comparison, some data on the behavioural effects of bombesin are included. This peptide caused behavioural alterations similar to those of the neurohypophyseal hormones, but these were apparently mediated by different mechanisms. It is suggested that centrally-released oxytocin and/or vasopressin might be physiologically involved in the regulation of animal behaviour.

Animals↗

The effect of estradiol on the alterations in monoamine-mediated behavioural responses induced by administration of electroconvulsive shocks or imipramine to female rats.

Female rats were treated daily with electroconvulsive shocks (ECS) or imipramine (10 mg/kg) for 10 days. Both types of treatment enhanced behavioural responses mediated by 5-hydroxytryptamine (5-HT) and noradrenaline (NA). A behavioural response mediated by dopamine (DA) was enhanced by electroconvulsive shock-treatment alone. On the other hand, rats treated only once with imipramine exhibited reduced DA-mediated behaviour. Priming the rats with estradiol valerate before starting electroconvulsive shock- or imipramine- treatment did not produce any significant effect on the enhancement in the behavioural response mediated by 5-HT. The enhancement in behaviour mediated by NA caused by electroconvulsive shock was also not altered, but that caused by treatment with imipramine was abolished. Enhancement of behaviour mediated by DA following electroconvulsive shock-treatment was also attenuated, while there was a positive reduction in behaviour mediated by DA in imipramine-treated rats. The two therapeutic approaches to depression, viz., electroconvulsive shock and imipramine, thus produced somewhat different effects on the central functions mediated by monoamines. Furthermore, an estrogen given prior to the treatments differentially altered the influence exerted by electroconvulsive shock and imipramine on monoamine functions in brain. The results may be pertinent to the clinical impression that estrogens produce a partial resistance to the antidepressant efficacy of imipramine-like drugs.

Animals↗

The effects of lesions produced by 5,7-dihydroxytryptamine on 5-hydroxytryptamine-mediated behaviour induced by amphetamine in large doses in the rat.

The role of 5-hydroxytryptamine (5-HT)-containing terminals in the spinal cord and basal ganglia in behavioural responses induced by amphetamine in large doses have been investigated using the neurotoxin for 5-HT, 5,7-dihydroxytryptamine (5,7-DHT). The effects of pretreatment with 5,7-DHT were also examined using the 5-HT agonist, 5-methoxy-N,N-dimethyltryptamine (5-MeODMT). d-Amphetamine (25 mg/kg) induced several classical 5-HT-dependent behavioural responses (head weaving , forepaw treading, hind limb abduction, "wet dog" shakes, Straub tail), together with some classical dopamine (DA)-dependent behaviour and backward locomotion which requires both transmitters. Pretreatment with 5,7-DHT, given into the striatum significantly decreased "wet dog" shakes and virtually abolished backward walking. Pretreatment with 5,7-DHT in the nucleus accumbens or substantia nigra did not significantly alter behaviour. Pretreatment with 5,7-DHT intraspinally did not significantly alter behaviour induced by amphetamine, although a decrease of Straub tail just failed to reach significance (P = 0.056). Similar pretreatment in rats given 5-MeODMT (8 mg/kg) significantly enhanced both Straub tail and tremor but did not alter the other behavioural responses induced by this drug (limb abduction, forepaw treading, head weaving ). The results in general suggest that behavioural responses induced by 5-HT can be classified into 3 groups (a) those requiring striatal 5-HT ("wet dog" shakes and backward locomotion), (b) those requiring spinal 5-HT (Straub tail, tremor) and (c) those requiring neither spinal nor striatal 5-HT (hind limb abduction, head weaving and forepaw treading).

5,7-Dihydroxytryptamine↗