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Parametric and pharmacological analyses of the enhanced grooming response elicited by the D1 dopamine receptor agonist SKF 38393 in the rat.

The present report investigated several parametric and pharmacological aspects of the enhanced self-grooming behavior of rats following systemic administration of the selective D1 dopamine (DA) receptor agonist SKF 38393. The amount of time that rats spent grooming themselves was measured continuously for 30 min following drug administration to provide a quantitative measure of the drug-induced behavior. SKF 38393 increased the amount of grooming in a dose-dependent manner (0.5-16 mg/kg, SC). The onset of this effect required at least 5 min and it persisted for at least 60 min. The ability of SKF 38393 to enhance grooming was shared by R-SKF 38393, but not S-SKF 38393, consistent with the affinities of these enantiomers for the D1 DA receptor. Unlike SKF 38393, the peripheral D1 agonist fenoldopam (SKF82526) failed to cause an increased grooming response, suggesting a central site of action for elicitation of this behavior. The SKF 38393-induced increase in grooming was competitively antagonized by the D1 selective antagonist SCH 23390 (0.5 mg/kg, SC). Although the D2 DA receptor-selective antagonist eticlopride reduced SKF 38393-elicited grooming, this antagonism appeared to be of a physiological rather than pharmacological nature. When eticlopride was coadministered with the non-selective (mixed) D1/D2 agonist apomorphine, an increase in grooming behavior similar to that produced by SKF 38393 was observed. Inactivation of D1 and D2 DA receptors produced by pretreatment with the irreversible antagonist N-ethoxycarbonyl-2-ethoxy-1,2-dihydroquinoline (EEDQ), at a dose which reduces D1 and D2 receptor density by > or = 50% (8.0 mg/kg, IP), reduced SKF 38393-induced grooming by approximately 50%.(ABSTRACT TRUNCATED AT 250 WORDS)

2,3,4,5-Tetrahydro-7,8-dihydroxy-1-phenyl-1H-3-ben

Grooming behavior in cats with pontile lesions and cats with tectal lesions.

Because an abnormal grooming behavior that is mediated by the superior colliculi is elicited from cats with pontile lesions, an ablation study of these structures was conducted to specify quantitatively the changes in grooming behavior. Cats that underwent the surgical procedure except for the lesion and cats with lesions of the auditory and visual cortices served as control groups. Time-lapse motion pictures of the cats in their home cages were taken, and statistical analyses of the grooming behavior shown on the films indicated that cats with pontile lesions and cats with tectal lesions spent less time grooming, had shorter grooming bouts, and failed to exhibit the normal temporal pattern of grooming behaviors. Other studies revealed that cats with pontile or tectal lesions were deficient in removing tapes stuck on their fur. A sensory-loss hypothesis appeared to account for some of the changes, but a deficit in endogenous control of the grooming behaviors also was indicated. The literature on grooming behavior related to peripheral versus endogenous control was reviewed, and the role of the superior colliculi as a higher order integrative center for complex behaviors is emphasized.

Animals

Biological bases for the integration of appetitive and consummatory grooming behaviors in the cat: a review.

Cats with pontile lesions, frontal neocortical lesions, and thyroidectomized cats display a dissociation of the appetitive and consummatory components of grooming behavior following tactile stimulation of the body surface, an abnormal behavior which waxes and wanes with the seasons of the year. Tryptophan hydroxylase activity and serotonin levels were significantly decreased in the superior colliculi (but not other brain regions) in cats with pontile lesions or frontal neocortical lesions, but not in thyroidectomized cats. Systemic administration of 5-hydroxytryptophan or monoamine oxidase inhibition plus tryptophan administration abolishes the abnormal grooming behavior in each group of cats, and microinjections of 5-hydroxytryptophan or serotonin into the superior colliculi has the same effect, indicating that the change in a serotonergic system is a critical aspect of the abnormal behavior in cats with lesions and that a serotonergic system may also be involved in the genesis of the abnormal grooming behavior in thyroidectomized cats. Functional inactivation of the serotonergic system by p-chlorophenylalanine, LSD, or serotonin receptor blockade does not induce the abnormal grooming behavior in normal cats, indicating that other factors are involved. Cats with lesions and thyroidectomized cats display a rhythmic dysfunction in the excretion of glucocorticoids and glucocorticoid administration abolishes the abnormal grooming behavior, suggesting that glucocorticoids are the other critical factor. Adrenalectomized cats do not display the abnormal grooming behavior, but when adrenalectomized cats are treated with p-chlorophenylalanine, the abnormal behavior appears. Thus, a serotonergic system in the superior colliculi, operating at some level of glucocorticoid function, is involved in the integration of appetitive and consummatory grooming behaviors.

5-Hydroxytryptophan

Grooming and consort partner selection in a troop of Japanese monkeys (Macaca fuscata).

The Arashiyama West troop of Macaca fuscata consists of 150 monkeys transported from Kyoto, Japan, in February 1972 to their present location near Laredo, Texas. At this site the animals range over 108 acres and during the study period were only minimally provisioned and disturbed. In the report of a 3-year study of consort partner selection it was suggested that yearlong social bonds within the troop appeared to be distinct from consort bonds formed during mating season. Like Sade, we consider grooming to be a good measure of yearround affinitive bonding in macaques. This study compares the pairs formed for consorting with those formed for yearlong grooming activities, by the 94 individuals who formed consort relationships during the mating season of 1973--1974. It was found that yearlong grooming involved pairs of monkeys which were significantly different from those for consorting. Grooming was relatively frequent between related monkeys, while consort partners were seldom related. In addition, yearlong grooming patterns were not disrupted during mating season. These results suggest that consort and grooming activities are behavioral expressions of two important social networks, involving mutually exclusive social bonds.

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

Grooming in the rat as an aftereffect of lateral hypothalamic stimulation.

Grooming occurred as an aftereffect of electrical stimulation at hypothalamic sites that elicited locomotion, drinking, or eating as stimulus-bound behaviors. Removal of food or water, which caused rats to switch from one stimulus-bound behavior to another, produced little or no change in the grooming aftereffect. Stimulation with pulse showed that the absolute refractory period for the neurons responsible for the occurrence of grooming is approximately 1 msec. This value is longer than those reported for drinking, eating, and locomotion. Finally, it was found that with some electrodes, low levels of stimulation elicited grooming directly, during the stimulation; at higher levels grooming occurred only as an aftereffect. It is concluded that grooming is activated through neurons separate from those that produce drinking, eating, or locomotion and that its occurrence as an aftereffect may be due to an interaction between short-lasting inhibitory and longer lasting excitatory effects of the stimulation.

Animals

Intracerebral adrenocorticotropic hormone mediates novelty-induced grooming in the rat.

Intact male rats exhibited more grooming in unfamiliar testing chambers than in their home cages. Hypophysectomized rats showed a much reduced increase in grooming in these testing chambers. Intraventricular injections of antiserum to adrenocorticotropic hormone to intact rats decreased the grooming usually observed in the novel situation, whereas a similar injection of control serum did not produce this effect. Peripheral injections of the antiserum did not affect grooming. Since intraventricularly injected adrenocorticotropic hormone induces excessive grooming, these results suggest that the increased grooming observed in the novel environment may be at least partly due to the release of this hormone directly into the cerebral ventricular system.

Adrenocorticotropic Hormone

Biosocial functions of grooming behavior among the common Indian langur monkey (Presbytis entellus).

An intensive study of the Indian langur monkey (Presbytis entellus) reveals an interplay between biological and social processes. This study which involved captive animals and over 1,500 observation hours suggests that grooming behavior alone cannot be used to measure status differences between the sexes, nor as a means to understand how pathogens carried by ecto-parasites are controlled. It is shown that the type of grooming exhibited in an interactional setting and the region of the body presented by one animal to another for grooming is often determined by the immediate social events and by the nature of the relationship existing between the participants. For these Colobine langur monkeys grooming behavior not only facilitates integration but it also enables an animal to manipulate a tense situation into a peaceful one. Without reference to many of the other patterns of behavior which only relatively, not absolutely differentiate age and sex classes, grooming behavior cannot be entirely understood.

Age Factors

Effect of 5-HT1C and 5-HT2 receptor stimulation on excessive grooming, penile erection and plasma oxytocin concentrations.

Serotonin (5-HT) receptor agonist-induced excessive grooming, penile erection and oxytocin secretion were studied in chronically cannulated freely moving rats. The 5-HT1C receptor agonist, m-chlorophenylpiperazine (m-CPP), which also binds to other 5-HT receptors, produced dose-dependent excessive grooming, penile erection and increases in circulating oxytocin concentrations. Maximal responses for excessive grooming and penile erection occurred at 0.3-0.6 mg/kg i.v. m-CPP. Higher doses (0.9-2.5 mg/kg i.v.) caused further increases in oxytocin concentrations, but attenuated both behavioral responses. All three responses to m-CPP (0.6 mg/kg) were attenuated by antagonists with high affinity for the 5-HT1C receptor site (mianserin, LY-53857 and metergoline), but not by the 5-HT2 receptor antagonist ketanserin. The 5-HT2/5-HT1C agonist, 1-(2,5-dimethoxy-4-iodophenyl)-2-aminopropane (DOI), increased plasma oxytocin concentrations only. After ketanserin pretreatment, DOI caused penile erection and diminished the oxytocin response. All responses to DOI were blocked completely by pretreatment with LY-53857 plus ketanserin. Excessive grooming and penile erection showed significant bimodal correlations with the oxytocin response. These data suggest that stimulation of 5-HT1C receptors induces excessive grooming, penile erection and increased oxytocin secretion. Stimulation of 5-HT2 receptors causes a further increase in plasma oxytocin concentration, but inhibits both behavioral responses.

Analysis of Variance

Suppressive effect of the dopamine D2 receptor agonist B-HT 920 on rat grooming.

The effect of the D2 agonist B-HT 920 was examined on three behavioural models of induced grooming in the rat. B-HT 920 potently inhibited the grooming elicited by a novel environment, whereas it stimulated the stretching-yawning syndrome. Pretreatment with the selective dopamine D2 receptor antagonist, sulpiride, reversed the phenomenon. When B-HT 920 was administered to rats before water immersion, it similarly antagonized total grooming; wet-dog shakes, detected in these same animals, were potently inhibited. Finally, B-HT 920 displayed inhibitory activity towards adrenocorticotropin hormone-induced excessive grooming. On the basis of these effects, the role of D2 receptor subtypes in the modulation of grooming is discussed.

Analysis of Variance

Effect of the D2-autoreceptor agonist B-HT 958 on both spontaneous and ACTH-induced stretching, yawning and grooming in the rat.

The D2 autoreceptor agonist B-HT 958, intraperitoneally injected into Wistar male rats in a novel environment, significantly increased stretching and yawning (SY) while inhibiting grooming. Pretreatment with the D2 antagonist sulpiride reversed these effects, antagonizing SY and restoring grooming. Similarly, when B-HT 958 was administered to rats in their home cages, it elicited SY and abolished grooming; moreover, when administered before the i.c.v. injection of adrenocorticotropin hormone, dose-dependently enhanced SY and strongly antagonized the typical syndrome of intensified grooming induced by the peptide. The possible relationship between SY and grooming and the involvement of D2 autoreceptors are discussed.

Adrenergic alpha-Agonists

On the ontogenetic and sequential characteristics of bombesin-induced grooming in the infant rat.

This paper examined the grooming effects of central injections of the tetradecapeptide bombesin (BN) in the infant rat. Anaesthetized rat pups of 1-20 days of age (n = 7/dose/age) were injected intracerebroventricularly (i.c.v.) with either BN (0.01-1.0 micrograms) or saline (control condition) and placed in heated test cages. Following recovery from the anaesthetic, the subsequent behaviour displayed by the pups was videotaped for the next 60 min. These videotaped responses were then scored for a variety of grooming and other behaviours. Pups of all ages (1-20 days) groomed in response to BN with the 1-day-old pups the least and 20-day-olds the most sensitive to BN. In the 10-day-olds, scratching, in an immature, non-contact form, was elicited by BN. This was replaced by mature, contact type scratching in the sequence of grooming behaviour at 20 days of age. Scratching activities appeared to form a subsystem connected to but nonetheless separate from washing behaviours and this was particularly distinguishable at 20 days of age. Changes in BN-induced grooming appeared to reflect the maturation of the motor capabilities of the developing rat. In conclusion, these results indicate that BN binding sites in the developing rat central nervous system (CNS) are pharmacologically functional from an early stage in ontogeny when there is little or no measurable amount of the peptide itself present in the CNS.

Aging

Stress induced grooming in the rat--an endorphin mediated syndrome.

Exposure of adult male Sprague--Dawley rats to a non-traumatic noise-light stress procedure subsequently increased grooming behavior in a novel environment. The grooming syndrome was marginally facilitated by adrenalectomy and by hypophysectomy. Opiate blockade by naltrexone returned grooming to basal levels. This suggests that stress induced grooming is not dependent upon pituitary-adrenal integrity for its expression, although it may be modulated by the latter. On the other hand this form of grooming may depend upon an endogenous opiate system.

Adrenalectomy

Serotonin, the superior colliculus, and grooming behavior in cats with neocortical lesions.

Previous studies of cats with pontile lesions indicate that a serotonergic deficit exists in the superior colliculi and that this deficit is involved in the genesis of an abnormal grooming behavior. Cats with frontal neocortical lesions exhibit the same serotonergic deficit and the same abnormal grooming behavior. The present study established that the serotonergic deficit is involved in the mediation of the abnormal grooming behavior in cats with frontal neocortical lesions. Microinjections of 5-hydroxytryptophan (5-HTP) and 5-hydroxytryptamine (5-HT) into the superior colliculi abolished or signigicantly reduced the abnormal behavior in cats with frontal neocortical lesions, whereas no effects of 5-HTP were observed after injections into the cerebrospinal fluid above the superior colliculi, into the tegmentum beneath the superior colliculi, or into the medial dorsal nucleus rostral to the superior colliculi. Other substances (tryptophan, noradrenaline, and gamma-amino-butyric acid) had no effect on the abnormal behavior when injected into the superior colliculi. Further evidence implicating a serotonergic deficit in the mediation of the abnormal behavior was obtained by systemic injections: The abnormal behavior was abolished with 5-HTP in cats with frontal neocortical lesions and in adrenalectomized cats that were previously treated with p-chlorophenylalanine.. The present study also demonstrated that the abnormal grooming behavior is induced by frontal neocortical lesions and not by more caudal lesions of the neocortex. The anatomical relations between the frontal neocortex and the superior colliculus and the role of these structures in grooming behavior are discussed.

5-Hydroxytryptophan

Role of glucocorticoids in an abnormal grooming behavior in cats with pontile or frontal neocortical lesions.

Cats with pontile or frontal neocortical lesions display a tactually elicited dissociation of appetitive and consummatory grooming behaviors. Systemic administration of glucocorticoids abolished the abnormal grooming behavior in cats with lesions, even when the stimulatory effect of glucocorticoids on serotonin metabolism was blocked by administration of p-chlorophenylalanine (PCPA). Microinjections of glucocorticoids into the superior colliculi also significantly decreased the abnormal grooming behavior. Adrenalectomized cats did not display the abnormal grooming behavior, but the abnormal behavior did occur in PCPA-treated adrenalectomized cats. Administration of either glucorticoids or 5-hydroxytryptophan abolished the abnormal behavior in PCPA-treated adrenalectomized cats. Thus, it appears that the pontile and frontal neocortical lesions produce deficits in both glucocorticoids and serotonin, and these deficits are necessary and sufficient conditions for inducing the abnormal grooming behavior.

11-Hydroxycorticosteroids

Role of thyroid hormones in an abnormal grooming behavior in thyroidectomized cats and cats with pontile lesions.

Thyroidectomized cats and cats with pontile lesions exhibit the same abnormal grooming behavior. Investigations were conducted to determine whether the abnormal grooming behavior in cats with pontile lesions was attributable to a hypothyroid condition. Administration of thyroid hormones abolished the abnormal behavior of thyroidectomized cats but was without effect in cats with pontile lesions. Studies of brain and blood levels of thyroid hormones in cats with pontile lesions failed to detect any differences from normal cats, and normal effects of thyroid hormones on evoked potentials to light flashes and to clicks were obtained in cats with pontile lesions. It was concluded that any thyroid dysfunction that may exist in the cats with pontile lesions was not involved in the genesis of the abnormal grooming behavior. Because the abnormal grooming behavior fluctuates rhythmically during the year, an endocrine hypothesis was entertained as the source of the fluctuations; endocrine concomitants of the abnormal behavior were investigated with longitudinal studies. The urinary excretion of 11-hydroxycorticoids exhibited a significant negative correlation with the abnormal behavior. When compared with that of normal cats, the plasma level of 11-hydroxycorticoids of thyroidectomized cats was significantly decreased. Intramuscular injections of 11-hydroxycorticoids abolished the abnormal behavior in both thyroidectomized cats and cats with pontile lesions. These results considered together with previous findings indicate that thyroidectomy and pontile lesions induce similar changes in 11-hydroxycorticoid and serotonergic functions-changes that are critical features of the physiological bases of the abnormal grooming behavior in both groups.

11-Hydroxycorticosteroids

Cortex, striatum and cerebellum: control of serial order in a grooming sequence.

Rats emit grooming actions in sequences that follow characteristic patterns of serial order. One of these patterns, a syntactic chain, has a particularly stereotyped order that recurs spontaneously during grooming thousands of times more often than could occur by chance. Previous studies have shown that performance of this sequence is impaired by excitotoxin lesions of the corpus striatum. In this study we examined whether the striatum is unique in its importance to this behavioral sequence or whether control of the sequence instead depends equally upon the cortex and cerebellum. In two experiments, a fine-grained behavioral analysis compared the effects of striatal ablation to the effects of motor cortex ablation, ablation of the entire neocortex, or ablation of the cerebellum. Cortical and cerebellar aspiration produced mere temporary deficits in grooming sequences, which appeared to reflect a general factor that was nonsequential in nature. Only striatal damage produced a permanent sequential deficit in the coordination of this syntactic grooming chain. We conclude that the striatum has a unique role in the control of behavioral serial order. This striatal role may be related to a number of sequential disorders observed in human diseases involving the striatum.

Animals

Evidence for a noradrenergic mechanism in the grooming produced by (+)-amphetamine and 4, alpha-dimethyl-m-tyramine (H 77/77) in rats.

Rats were kept on a 12-h light-dark cycle. One hour after the light was switched on, physiological saline, (+)-amphetamine 1 mg/kg, and H 77/77 5 mg/kg were injected s.c.; the number of groomings was counted 1-2 h after the treatments. (+)-Amphetamine and H 77/77 produced increased grooming which was antagonized by the tyrosine hydroxylase inhibitor H 44/68 (250 mg/kg), the dopamine-beta-hydroxylase inhibitor FLA 63 (40), the neuroleptics haloperidol (0.1 and 0.5), and clozapine (1 and 5). The (+)-amphetamine-induced grooming was also antagonized by the NA-receptor blocker aceperone (10) but not by the sedative phenothiazines mepazine (10) and diphenhydramine (20) nor diazepam (1). These results indicate that NA-release is involved in the mediation of (+)-amphetamine- and H 77/77-induced grooming. The inhibition of haloperidol and clozapine is presumably due to NA-receptor blockade.

Amphetamine