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Distinct grooming patterns induced by intracerebroventricular injection of CRH, TRH and LHRH in male rats.

This paper reports the effects on grooming, related behaviors and levels of anxiety induced by the hypophysiotropic peptides corticotropin-releasing hormone (CRH, 1 microgram, 0.2 nmol, icv), thyrotropin-releasing hormone (TRH, 100 micrograms, 275 nmol, icv) and luteinizing hormone-releasing hormone (LHRH, 1.5 micrograms, 1.3 nmol, icv) administered into the lateral ventricle of the brain (icv) of adult male rats of a Holtzman-derived colony (N = 15, each group). CRH induced an increase in total grooming scores, whereas LHRH, TRH and vehicle had no effect. CRH strongly increased face and head grooming and induced head shakes. The time spent in rearing and gnawing was significantly decreased. In the plus-maze, CRH reduced the time of exploration in the open arm. TRH increased face grooming and induced body shakes. LHRH had no effect on grooming or rearing behavior. No body or head shakes were observed after LHRH administration. Scoring of individual grooming elements demonstrated differences in action of the three peptides. Although both CRH and TRH increased face grooming, only CRH induced head grooming. Furthermore, CRH induced predominantly head shakes while TRH increased body shake activity. In contrast, CRH was anxiogenic and TRH appeared to induce stereotyped behavior. From the characterization of grooming elements and related responses, we conclude that each hypophysiotropic peptide induces a specific behavioral pattern.

Analysis of Variance↗

Local control of leg movements and motor patterns during grooming in locusts.

This study demonstrates that the thoracic and abdominal nervous system of locusts is sufficient to mediate several site-specific and distinct grooming leg movements. Locusts can use a hindleg or middle leg to groom at least four ipsilateral thoracic and abdominal sites, without input from the brain, subesophageal ganglion, or prothoracic ganglion. The hindleg is used to groom the posterior abdomen, the ventral or posterior hindleg coxa, and the ear; the middle leg is used to groom the anterior hindleg coxa. Grooming movements are often rhythmic and display site-specific intralimb coordination patterns. During grooming of the posterior abdomen or ventral hindleg coxa, for example, hindleg tibial extension occurs nearly simultaneously with femur elevation, in contrast with locust hindleg movements during walking. Electromyographic (EMG) recordings during these movements show that rhythmic bursts of tibial extensor activity occur nearly in-phase with those of trochanteral levators, in contrast to hindleg EMGs during walking. During grooming of the ear, hindleg tibial extension/flexion and tibial extensor/flexor muscle bursts can occur independently of the femur elevation/depression and trochanteral levator/depressor muscle bursts, suggesting that the neural modules controlling tibial and femoral movements can be uncoupled during this behavior. Tibial extension can occur before, or even in the absence of, tibial extensor muscle activity, suggesting that spring-like properties of the leg and energy transfer from femur motion may play important roles in such leg movements. Adjacent legs sometime show coordinated femur movement during grooming with one hindleg, suggesting that grooming may also involve interlimb coordination.

Animals↗

Venom of a parasitoid wasp induces prolonged grooming in the cockroach

The parasitoid wasp Ampulex compressa hunts cockroaches Periplaneta americana, stinging them first in the thorax and then in the head, the sting penetrating towards the subesophageal ganglion. After being stung the cockroach grooms almost continuously for approximately 30 min, performing all the normal components of grooming behavior. This excessive grooming is only seen after the head sting and cannot be attributed to stress, to contamination of the body surface or to systemic or peripheral effects. This suggests that the venom is activating a neural network for grooming. We suggest that the venom induces prolonged grooming by stimulating dopamine receptors in the cockroach, for the following reasons. (1) Reserpine, which causes massive release of monoamines, induces excessive grooming. (2) Dopamine injected into the hemocoel also induces excessive grooming and is significantly more effective than octopamine or serotonin. In addition, the dopamine agonist SKF 82958 induces excessive grooming when injected directly into the subesophageal ganglion. (3) Injection of the dopamine antagonist flupenthixol greatly reduces venom-induced grooming. (4) Dopamine, or a dopamine-like substance, is present in the venom.

Journal Article↗

Factors affecting the bioavailability of benzo[a]pyrene from oils in mouse skin: oil viscosity, grooming, activity and its prevention.

In the main study, tritium-labelled benzo[a]pyrene ([3H]BaP) was added to oils of a wide range of viscosity (from 13.5 to ca. 8000 cSt at 40 degrees C) and these were applied once to mouse skin under conditions where grooming was either allowed or prevented. The binding of [3H]BaP to epidermal protein and DNA was assessed. In addition, some studies were conducted to investigate the effect of prior oil exposure on binding levels and to compare binding levels following single and multiple application. It was found in the main study that the binding of [3H]BaP to both DNA and protein was increased as the viscosity of the oil vehicle decreased. Whereas only a twofold difference in DNA binding was found between the lowest and highest viscosity oil vehicle if grooming was allowed, a 14-fold difference was seen if grooming was prevented. This was due to much higher binding levels with low viscosity oils when grooming was prevented. Protein binding showed similar results, although the difference between the grooming and non-grooming situations was slightly less. Whether grooming was prevented or not, both DNA and protein binding of [3H]BaP were found to be inversely proportional to the logarithm of viscosity of the oil vehicle, but when grooming was prevented, the slope for DNA binding was 10 times steeper than when it was permitted. Exposure of the skin to either high or low viscosity oils prior to the application of [3H]BaP in the same oil inhibited the binding to DNA but not to protein.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Molecular transduction mechanisms in ACTH-induced grooming.

Intraventricular administration of ACTH1-24 induces excessive grooming in the rat. Ethogram analysis shows that the peptide does not alter grooming behavior seen in a novel box, but that it prolongs the duration of the grooming bout. Extensive structure-activity studies have been performed which suggest that the active site lies in a region (5-13) of the ACTH molecule. Interestingly, the (1-24) sequence is fully active, whereas (1-10) and (11-24) alone or in combination are inactive, pointing to a specific stereoconformation necessary to induce grooming. However, despite the fact that there are ACTH-and/or alpha-MSH-containing peptidergic neurons, no conclusive evidence is available demonstrating stereospecific, saturable binding sites for these peptides in brain. The analysis of the neural substrate underlying ACTH-induced excessive grooming has been performed by means of electrolytic lesions of specific brain regions and by neuropharmacological manipulations. The data suggest that the periaqueductal gray is the primary target for ACTH and that the activity of neostriatum and accumbens, via a nigro-colliculus-periaqueductal gray pathway, modulates the display of excessive grooming. An important feature of the neural substrate is that it displays single-dose tolerance to the peptide during the first hours after the first peptide injection. It is suggested that the tolerance is a feature of an opioid receptor-containing component of the neural substrate. The molecular mechanism of action of ACTH is complex and may involve different transmembrane signal transduction systems. The peptide decreases the degree of phosphorylation of a neuron-specific, synaptic phosphoprotein B-50 by inhibition of protein kinase C. It is concluded that changes in the degree of phosphorylation of B-50 regulate the activity of the lipid kinase phosphatidylinositol 4-phosphate kinase. Therefore, the B-50 protein seems to be part of a negative feedback loop in the receptor-activated hydrolysis of phosphatidylinositol 4,5-bis-phosphate (PIP2). There is increasing evidence that the molecular mechanism by which ACTH brings about the grooming response involves a change in phosphorylation of B-50. Firstly, the structure-activity relationship of ACTH-induced excessive grooming is nearly identical to that obtained for ACTH-induced inhibition of protein kinase C.(ABSTRACT TRUNCATED AT 400 WORDS)

Adrenocorticotropic Hormone↗

Oxytocin potently enhances novelty-induced grooming behavior in the rat.

Intracerebroventricular (i.c.v.) injection of oxytocin was followed by an enhancement of novelty-induced grooming in male and female rats. This effect was dose-dependent, in a dose range of 0.1-10 micrograms. Grooming activity of rats injected i.c.v. with 10 micrograms of oxytocin was 9-fold higher than that of saline-injected controls. The analysis of behavioral element composition revealed an increased occurrence of genital grooming in oxytocin-injected rats. A time-course study revealed a sustained increase in grooming activity of oxytocin-treated rats during 45 min of behavioral testing. Intraperitoneal (i.p.) injection of the dopamine antagonist, haloperidol, totally suppressed oxytocin-enhanced grooming. Furthermore, i.p. injection of the opiate receptor antagonist, naloxone, was followed by an attenuation but not a suppression of grooming enhanced by i.c.v. administration of oxytocin. In addition, a small but significant increase in grooming activity was observed after subcutaneous injection of oxytocin. These results suggest that oxytocin-enhanced grooming behavior involves central mechanisms, e.g. dopamine and opioid transmission in the brain.

Animals↗

ACTH-(1-24) and RX 336-M induce excessive grooming in rats through different mechanisms.

ACTH-(1-24) (0.03-6 micrograms i.c.v.) and RX 336-M (7,8-dihydro-5',6'-dimethylcyclohex-5'-eno-1',2',8',14 codeinone) (1.5-6 mg/kg i.p.) induce dose-related excessive grooming and 'wet-dog' shaking in rats. In the present study, the grooming associated with these compounds was compared and analyzed pharmacologically. Grooming caused by RX 336-M and by ACTH-(1-24) was antagonized when rats were pretreated with comparable doses of morphine (0.5-4 mg/kg s.c.), however, only ACTH-(1-24)-induced grooming was attenuated by naloxone (1 and 10 mg/kg s.c.). ICI 154,129 (N,N-bisallyl-Tyr-Gly-Gly-psi-(CH2S)-Phe-Leu-OH) (30 mg/kg s.c.), a selective delta-opiate receptor antagonist, was ineffective against both ACTH-(1-24) and RX 336-M. Although haloperidol is known to antagonize grooming elicited by ACTH-(1-24) (e.g., Wiegant et al., 1977, European J. Pharmacol. 41, 343), even a high dose of this neuroleptic agent (5 mg/kg s.c.) only partially attenuated grooming caused by RX 336-M. Tolerance developed to the grooming elicited by RX 336-M, and by ACTH-(1-24), but there was no cross-tolerance. Both agents were active in genetically hypotrichotic rats; and, again in such animals, even after numbing the area caudal to the shoulders with lidocaine. Given the divergent results with naloxone, and, possibly, with haloperidol, and the lack of cross-tolerance, we conclude that the excessive grooming induced in rats by ACTH-(1-24) and by RX 336-M is mediated by different mechanisms.

Adrenocorticotropic Hormone↗

Grooming behavior induced by substance P.

The intracerebroventricular (i.c.v.) administration of substance P elicits in rats an excessive grooming that is characterized by body grooming, anogenital grooming and scratching. The total grooming scores displayed by rats treated with substance P hardly exceeded 23% of the theoretical maximal grooming score. Substance P-induced excessive grooming was suppressed by pretreatment with naloxone, haloperidol on neurotensin. It is concluded that substance P induces excessive grooming with a pattern different from that of grooming elicited by other peptides.

Animals↗

Induction of grooming in resting rats by intracerebroventricular oxytocin but not by adrenocorticotropic hormone-(1-24) and alpha-melanocyte-stimulating hormone.

Adrenocorticotropic hormone (ACTH) and alpha-melanocyte-stimulating hormone (alpha-MSH) injected i.c.v. induce so called 'excessive grooming'. Whether these peptides play a role in the initiation of grooming is not clear, since rats will groom even as a consequence of a particular environmental stimulation, such as handling and/or a novel environment. In most studies, therefore, the first 15 min after i.c.v. injection are not examined. Here we report on the effects of slow i.c.v. infusions of ACTH-(1-24), alpha-MSH and oxytocin in resting rats in their home cages. Interestingly, i.c.v. infusions of oxytocin did initiate grooming in a dose-related way. In contrast, i.c.v. infusions of both ACTH-(1-24) and alpha-MSH in resting rats were without effect on grooming. Oxytocin is apparently involved in the initiation of self-grooming in rats, whereas ACTH and alpha-MSH prolonged grooming initiated by other means, e.g. handling procedures and/or a novel environment. We conclude that the effects of alpha-MSH and ACTH on grooming are conditional, depending on the behavioural state (active or resting) of the animal.

Animals↗

Bombesin-induced grooming in the golden hamster.

Lateral cerebroventricular injection of the peptide bombesin (0.01-1.0 micrograms) promptly elicited excessive grooming and scratching behaviors in home-caged male and female golden hamsters. Bombesin-induced grooming persisted throughout a 60-min observation period at doses of 0.1-1.0 micrograms. Grooming with forepaws and mouth was more consistently increased than hindleg scratching behaviors. Dependence of this neuropeptide effect on grooming on muscarinic cholinergic activity was assessed by injecting scopolamine (0.001-1 mg/kg) intraperitoneally 15 min prior to 0.1 microgram bombesin. Excessive grooming induced by centrally administered bombesin was abolished by 0.1 and 1 mg/kg scopolamine, although basal level of grooming was not significantly affected. The findings indicate a cross-species generality of the dependence of bombesin-induced grooming on muscarinic cholinergic activity, and species-specific differences among rodents in the components of excessive grooming elicited by bombesin.

Animals↗

Separation of activation and pattern in grooming development of weaver mice.

The effects of environmental conditions and age on grooming behavior were examined in weaver mutant mice and control littermates. Due to deficits in both the cerebellum and the dopaminergic system, weaver mice provide an opportunity to investigate how both of these systems are involved in grooming. Although homozygous weaver (wv/wv mice display deficiencies in grooming behavior, our results indicate that these effects are both context and age dependent. Overall wv/wv mice spent less time grooming than did controls. However, during the post-swim period wv/wv, after day 13, reached the grooming levels of pre-swim control mice. After day 15 wv/wv mice showed a higher number of post-swim grooming bouts relative to pre-swim, and in fact exceeded the number of bouts performed by controls in either pre- or post-swim conditions. Although controls displayed longer bouts than mutants overall, during the post-swim period wv/wv mice, after day 13, produced bouts as long as the control animals did pre-swim. This could in part reflect activation by previous swimming. Our data indicate these activational effects can be separated from balance or posture problems. From examination of the individual grooming stroke types used by the two groups, it is evident that the strokes used by mutant animals clustered around the early grooming sequence phase. In contrast, some of the later strokes were never used by the wv/wv mice during the entire developmental period studied. Our results emphasize the importance of using multiple measures of an action sequence and testing under different conditions.

Aging↗

The time course of excessive grooming after neuropeptide administration.

In this paper we review the temporal pattern of excessive grooming in the hour or so following the central injection of ACTH1-24 in the rat. Changes in the grooming pattern after specific neuropharmacological manipulations of dopaminergic and opiate-related systems are presented which indicate a differential sensitivity of the grooming responses at different times after injection. The grooming affected by dopaminergic antagonists and opiate agonists and antagonists occurs in the last 30 min of the observation period while that found earlier is unaffected. It is also the grooming in this last 30 min of the observation period term tolerance to central administration of ACTH1-24. In contrast lesions of the central nervous system that affect excessive grooming, i.e., the substantia nigra and the hippocampus, reduce grooming throughout the observation period. The present analysis has provided evidence for dopamine/opiate insensitive and sensitive systems in excessive grooming, and thus temporal aspects are of extreme importance to the understanding of central neuropeptide influences on behavior.

Adrenocorticotropic Hormone↗

Genotype-dependent effect of ACTH1-24 on grooming and yawning in two inbred strains of rats.

It has long been known that the intracerebroventricular administration of ACTH1-24 increases the duration of grooming episodes and the frequency of yawning in rats. The objective of this study was to investigate in what way these episodes are prolonged and whether and to what extent genotype influences such effects. We compared the effect of increasing doses of intracerebroventricular injections of ACTH1-24 on grooming and yawning in males of two inbred strains of Sprague-Dawley rats with distinct yawning frequency, high-yawning (HY) and low-yawning (LY). In LY rats the duration of grooming episodes increased, while in HY rats grooming episodes augmented both in number and duration. In LY rats the duration of grooming components increased likewise, in HY rats however, neither the number nor the duration of the components changed. The grooming rate in both strains of rats was slowed, though more so in LY than in HY rats. Yawning increased in LY rats but not in HY rats. We conclude therefore that ACTH1-24 increases the duration of grooming episodes by slowing the grooming rate according to genotype, and may or may not alter the frequency of yawning.

Adrenocorticotropic Hormone↗

The role of prolactin and testosterone in mediating seasonal differences in the self-grooming behavior of male meadow voles, Microtus pennsylvanicus.

Self-grooming in response to the odors of conspecifics is a form of olfactory communication among meadow voles. The amount of time meadow voles spend self-grooming when they encounter the odors of conspecifics varies seasonally, with males targeting the odors of reproductively active females only during the breeding season. Other odor related behaviors in male voles such as odor preferences for conspecifics and the attractiveness of their odors to conspecifics vary seasonally as well. For male meadow voles, these behaviors are mediated by seasonal variations in testosterone (T) and prolactin (PRL) titers. The objective of this study was to determine whether seasonal differences in the amount of time male meadow voles self-groom in response to odors of conspecifics are mediated by seasonal rhythms in their circulating T and PRL titers. We tested the hypothesis that high titers of both T and PRL are necessary for reproductively active (long-photoperiod; LP) males and sufficient for reproductively quiescent (short-photoperiod; SP) male voles to spend more time self-grooming in response to odors of LP females than to those of other conspecifics. Results of this study demonstrate that high titers of PRL and T are necessary for LP male meadow vole to self-groom more in response to odors of LP females as compared to those of other conspecifics, but were not sufficient to induce SP males to preferentially self-groom to odors of LP females. The endocrine control of self-grooming by LP males appears to depend upon high titers of both PRL and T, which matches the endocrine mediation of other odor related behaviors in male voles. In contrast, the endocrine tissues that underlie self-grooming in SP male meadow voles appear to be refractory to the effects of LP-equivalent titers of PRL and T.

Animals↗

Behavioral effects of anti-muscarinic, anti-serotonergic, and anti-NMDA treatments: hippocampal and neocortical slow wave electrophysiology predict the effects on grooming in the rat.

Previous research has shown that hippocampal and neocortical activation accompanies the postural changes occurring during self-grooming in rats but is absent or reduced during the stereotyped components of grooming, including head-washing and licking or biting. Since electrocortical activation is dependent on ascending cholinergic and serotonergic projections, we hypothesized that central muscarinic and serotonergic blockade would disrupt grooming by degrading cerebral control of changes in posture. Consistent with this, we find that systemic injections of scopolamine: (a) markedly reduce the occurrence of adaptive changes in posture during grooming; (b) reduce the probability of transitions from head-washing to body grooming; (c) reduce both the probability and duration of sequences of body grooming; and (d) do not affect the duration of head-washing or the probability of transitions from washing the snout to washing over the top of the head. Destruction of central serotonergic neurons with intracerebral injections of 5,7-dihydroxytryptamine increases the tendency of scopolamine to shorten the duration and increase the number of separate sequences of grooming. Systemic injections of a NMDA antagonist (CGS 19755) also impair grooming behavior. The data show that blockade of muscarinic and glutamatergic transmission impairs instinctive behavior as well as learned behavior and that the behavioral effects of muscarinic and serotonergic blockade are consistent with data obtained from the study of cortical slow wave electrophysiology.

5,7-Dihydroxytryptamine↗

Conditioned grooming induced by the dopamine D1-like receptor agonist SKF 38393 in rats.

Stimulation of dopamine D1-like receptors reliably increases grooming in rats and mice. The study examined whether the grooming response elicited by the prototypical D1-like agonist SKF 38393 (8 mg/kg s.c.) could be conditioned to the specific environment in which it occurred. Rats in one group (Paired) received SKF 38393 and rats in another group (Unpaired) received saline in observation boxes outside of their housing room; the rats were then scored for duration and frequency of grooming bouts over 25 min. The ordering of injections was reversed the next day in the rats, housing room. The procedure was repeated twice, with at least one intervening drug-free day, to give three conditioning trials. The D1-like agonist significantly increased grooming on each of the three conditioning trials, without obvious tolerance or sensitization, and the effect tended to persist for the duration of each trial. On the test trial for conditioned grooming, mean grooming duration was significantly greater in the Paired than the Unpaired group, suggesting that SKF 38393-induced grooming had been conditioned to the test environment. This is the first time that drug-elicited grooming has been conditioned to environmental cues.

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

Behavioral assessment of facial pain in rats: face grooming patterns after painful and non-painful sensory disturbances in the territory of the rat's infraorbital nerve.

Noxious stimulation of the rat's face evokes intense face grooming with face wash strokes almost exclusively directed to the stimulated area (e.g. Clavelou et al., Neurosci. Lett., 14 (1989) 3263-3270). Similar asymmetric face grooming behavior has been observed after transection (Berridge and Fentress, J. Neurosci., 6 (1986) 325-330) and chronic constriction of the infraorbital nerve (Vos et al., J. Neurosci., 14 (1994) 2708-2723). In the present study, the relation between unilateral facial pain and asymmetric face grooming was experimentally studied in normal, intact rats: face grooming patterns evoked by non-painful sensory disturbances in the territory of the infraorbital nerve (i.c. unilateral vibrissae clipping, anesthetic infraorbital nerve blockade, application of mineral oil on vibrissae) were compared to those evoked by noxious facial stimulation (s.c. formalin injection in mystacial pad) and those observed in unstimulated control rats, using video-analysis. Only formalin-injected rats displayed significantly more face grooming activity directed to the affected infraorbital nerve territory than unstimulated control rats. Non-painful sensory disturbances (especially mineral oil application) induced an initial bout of directed face grooming; this response was transient and short-lasting. These observations suggest that directed face grooming can be used as a sign of unilateral facial pain in freely moving rodents; unilateral non-painful facial sensory disturbances do not lead to intense and persistent directed face grooming.

Animals↗

Time structure of self-grooming in the rat: self-facilitation and effects of hypothalamic stimulation and neuropeptides.

Specific brain manipulations, such as stimulation of the paraventricular nucleus of the hypothalamus (PVH) or injections of neuropeptides, increase self-grooming in the rat. Such manipulations also affect the different movements that constitute grooming. Using models to assess the time structure of these movements, the authors demonstrate that the rules that control the time structure within grooming are different from the ones that control its initiation. This study also showed that grooming is self-facilitating and that different brain manipulations in the same hypothalamic area induce structurally different kinds of grooming. The authors suggest that this part of the hypothalamus is not only involved in setting priorities to grooming, relative to other behaviors, but is also involved in the timing of different grooming components. These findings suggest that different neural mechanisms may be involved in the initiation and internal time structure of grooming.

Adrenocorticotropic Hormone↗