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Dopamine involvement in ACTH-induced grooming behavior.

Low doses of dopamine agonists and antagonists were tested for their effects on the excessive grooming behavior induced by intracerebroventricular (ICV) injections of ACTH1-24. Grooming scores were significantly depressed at doses of haloperidol, metoclopramide, pimozide, and butaclamol that did not decrease locomotor activity. In fact at two doses of haloperidol (0.067 and 0.10 mg/kg), grooming scores were decreased while locomotor activity was increased significantly. Metoclopramide increased grooming scores at a dose reported to block presynaptic dopamine receptors. Apomorphine potentiated the grooming induced by low doses of ACTH. These data support the hypothesis that dopaminergic neurotransmission is necessary for the display of ACTH-induced grooming behavior.

Adrenocorticotropic Hormone↗

Luteinizing hormone releasing hormone (LHRH) in the periaqueductal gray substance increases some subcategories of grooming behavior in male rats.

Locomotion, holeboard exploration and grooming behavior were measured in male rats after injection of LHRH or cerebrospinal fluid (CSF) into the mesencephalic periaqueductal gray substance (PGS). LHRH injection at doses of 75 ng did not alter locomotion and rearing but decreased significantly the scores of head dipping and defecation. Several subcategories of grooming responses were evaluated in a home cage-like environment. Head, body and genital grooming increased significantly after injection of 75 ng LHRH. The frequency of gnawing and head shakes increased as well. Lower doses (50 ng) also raised the scores of head and body grooming and produced a tendency for genital grooming increase. Higher doses (100 ng) did not affect genital grooming responses and produced drowsiness in most of animals. These results demonstrate that some motor activities are selectively modified by the localized administration of LHRH in the PGS.

Animals↗

Regulatory mechanisms underlying novelty-induced grooming in the laboratory rat.

Bouts of pelage cleaning can be readily evoked in laboratory rodents under conditions of exposure to novelty. Such novelty-induced grooming is described as stereotyped and rostro-caudal in its progression. The patterned structure of novelty-induced grooming makes it particularly attractive for research on the organizational and motivational underpinnings of co-ordinated behaviour. Micro-characteristics of stereotyped novelty-induced grooming bouts were studied in 27 female Wistar rats that were exposed to a novel arena with shelters for a period of 15 min. The order of grooming acts within the initial bouts was rostro-caudal, but subsequent bouts became progressively disorganized in their sequencing. The observed pattern of progressive bout disorganization may be attributed to the gradual dearousal from stress. Differences between consecutive bouts in their micro-characteristics suggest that at least some grooming actions emitted within the context of those bouts operate as relatively independent units of behaviour. Those unitary component actions appear to be integrated into protracted pelage cleaning sequences by a separate mechanism. Similar endogenous mechanisms have been proposed for other co-ordinated motor actions, which suggests that the organizational principles identified in the context of novelty-induced grooming may represent general principles that govern co-ordinated behaviour.

Animals↗

Grooming analysis algorithm for neurobehavioural stress research.

Since rodent self-grooming behaviours are elicited by both comfort and stressful conditions, traditional measures such as duration, latency of onset and the number of bouts may be not suitable to dissociate between these opposite conditions. The aim of the current study was to improve and optimize ethological measurement of self-grooming in neurobehavioural stress research enabling differentiation between stress and no-stress situations. This protocol assists in the correct interpretation of animal grooming behaviours and detection of stress by measuring alterations in grooming microstructure in different test situations. While a general pattern of self-grooming uninterrupted cephalocaudal progression is normally observed in no-stress (comfort) conditions in mice and other rodents, the percentage of "incorrect" transitions between different stages and the percentage of interrupted grooming bouts may be used as behavioural marker of stress. The protocol can be a useful tool in neurobehavioural stress research including modelling stress-evoked states, pharmacological screening of potential antistress drugs or behavioural phenotyping of genetically modified animals.

Algorithms↗

Mouse grooming microstructure is a reliable anxiety marker bidirectionally sensitive to GABAergic drugs.

Grooming is an important part of rodent behavioural repertoire, representing a complex hierarchically ordered cephalo-caudal sequence of patterns sensitive to stress and various drugs. Gamma-aminobutyric acid (GABA) is involved in the regulation of both anxiety and grooming behaviours. This study investigated the predictive validity of grooming behavioural microstructure as a marker of anxiety, by examining the effects of two GABAergic reference compounds, anxiolytic diazepam (0.1 and 0.5 mg/kg i.p.) and anxiogenic pentylenetetrazole (5 and 10 mg/kg i.p.) on mouse grooming. Our data suggest that the percentage of pattern transitions not fitting to the cephalo-caudal progression, and the percentage of interrupted grooming bouts are more reliable behavioural markers of stress bidirectionally sensitive to GABAergic anxiogenic and anxiolytic drugs, compared to the frequency and duration scores. Our study also confirms that detailed ethological analyses of grooming microstructure can be a useful tool in behavioural pharmacology of anxiety.

Animals↗

Increased grooming behavior in mice lacking vitamin D receptors.

Vitamin D is a neuroactive secosteroid with several important functions in the nervous system. Many human and animal findings link alterations in the vitamin D system to various neurological and behavioral disorders. Since grooming is an important element of animal behavior, here we studied whether genetic ablation of vitamin D receptors (VDR) in mice may be associated with altered grooming behaviors. Overall, VDR knockout (VDRko) mice presented longer duration and higher frequency of grooming when tested in the actimeter, open field, elevated plus maze, and horizontal rod tests. Increased grooming did not, however, correlate with unaltered general activity level (actimeter test), anxiety-like behaviors (hole board and elevated plus maze tests), and emotional reactivity index (defecation boli). In general, our results confirm the role of vitamin D and VDR in the regulation of behavior, including grooming, and suggest that increased grooming behavioral phenotype may be associated with genetic ablation of VDR in mutant mice.

Animals↗

Characterization of melanocortin receptor ligands on cloned brain melanocortin receptors and on grooming behavior in the rat.

Since the melanocortin MC3 and melanocortin MC4 receptors are the main melanocortin receptor subtypes expressed in rat brain, we characterized the activity and affinity of nine melanocortin receptor ligands using these receptors in vitro, as well as their activity in a well-defined melanocortin-induced behavior in the rat: grooming behavior. We report here that [D-Tyr4]melanotan-II and RMI-2001 (Ac-cyclo-[Cys4, Gly5, D-Phe7, Cys10]alpha-MSH-NH2) have significantly higher affinity and potency on the rat melanocortin MC4 receptor as compared to the rat melanocortin MC3 receptor. Nle-gamma-MSH (melanocyte-stimulating hormone) was the only ligand with higher affinity and potency on the rat melanocortin MC3 receptor. The potency order of melanocortin MC4 receptor agonists, but not that of melanocortin MC3 receptor agonists, fitted with the potency of these ligands to stimulate grooming behavior, when administered intracerebroventricularly. SHU9119 (Ac-cyclo-[Nle4, Asp5, D-Nal(2)7, Lys10]alpha-MSH-(4-10)-NH2) and RMI-2005 (Ac-cyclo-[Cys4, Gly5, D-Na](2)7, Nal(2)9, Cys10]alpha-MSH-(4-10)-NH2) were able to inhibit alpha-MSH-induced melanocortin receptor activity in vitro, as well as alpha-MSH-induced grooming behavior. Melanotan-II, [Nle4-D-Phe7]alpha-MSH and RMI-2001 were also effective in inducing grooming behavior when administered intravenously. In the absence of purely selective melanocortin MC(3/4) receptor ligands, we demonstrated that careful comparison of ligand potencies in vitro with ligand potencies in vivo, could identify which melanocortin receptor subtype mediated alpha-MSH-induced grooming behavior. Furthermore, blockade of novelty-induced grooming behavior by SHU9119 demonstrated that this physiological stress response is mediated via activation of the melanocortin system.

Amino Acid Sequence↗

Grooming secretions and seasonal adaptations in the blind mole rat (Spalax ehrenbergi).

The postulated thermoregulatory function of grooming in the "blind" mole rat (Spalax ehrenbergi), a solitary, highly aggressive subterranean rodent, was tested by subjecting individuals to extreme hot and cold environments and measuring their grooming frequency and thermoregulatory capacity. It was found that exposure to heat stress during both the winter and the summer failed to elicit grooming in isolated mole rats, even though their body temperature was significantly elevated. Thus, unlike Mongolian gerbils (Meriones unguiculatus), mole rats do not seem to have a mechanism for lowering body temperature by grooming. However, at low temperatures, grooming behavior significantly increased during both seasons, although a decline in body temperature was recorded only during the summer. Because grooming in mole rats expresses Harderian lipids onto the fur, it is suggested that in the winter, under cold stress, this behavior might prevent a drop in body temperature through spreading insulating Harderian lipids.

Adaptation, Physiological↗

Grooming in Lurcher mutant mice.

Lurcher mutant mice, characterized by degeneration of cerebellar Purkinje cells and granule cells, were compared to normal littermate controls for different facets of grooming and nongrooming behaviors after a brief period of water immersion. By comparison to normal controls, the number and the duration of several grooming components were decreased in Lurcher mutant mice, namely, licking the forelimb, the abdomen, the back, and the hindlimb. By contrast, the number and duration of body-shaking episodes were not reduced. Lurcher mutants had fewer grooming elements for bouts with at least five elements. However, the serial organization of grooming, as determined by the order of appearance of grooming elements, was maintained in Lurcher mutants. These results indicate that the cerebellar cortex is involved in the appearance of various grooming elements but not in the organization of the cephalocaudal sequence.

Analysis of Variance↗

Increased novelty-induced grooming in aged rats: a preliminary observation.

Age-related changes in grooming activity in female Fischer-344 rats were observed in a home cage and in a novel testing chamber for 50 min. Compared to the home cage condition with brief handling treatment, excessive grooming was found in the novel situation. The amount of novelty-induced grooming of aged rats (26-28 months) was about two times that of young animals (6-8 months). This increase in novelty-induced grooming of aged animals was attributable to an increase in the number of grooming bouts, prolongation of each grooming bout, and a slight increase in the duration of licking elements. However, there were no age differences in the percentage of face washing and the duration of face-washing elements. These results are discussed in terms of behavioral aging and age-related changes in peptidergic pathways in the brain.

Aging↗

Ethanol-induced grooming in mice selectively bred for differential sensitivity to ethanol.

In rats, excessive grooming follows the application of several forms of stress. The injection of alcohol may be considered as a stressor, since it increases plasma corticosterone levels. The present study examines the effects of ethanol injections on grooming in two lines of mice selected for differences in their hypnotic response to ethanol, i.e., the long sleep (LS) and short sleep (SS) lines developed at the Institute for Behavior Genetics at the University of Colorado at Boulder. Various subhypnotic doses of ethanol produced excessive grooming in the SS line, but this did not occur in the LS line. The SS mice also displayed more "novelty-induced" grooming compared to the LS mice after repeated exposure to the testing situation. The increase in excessive grooming in both the ethanol-induced and the novelty-induced excessive grooming situations was most apparent in the second half of the observation period.

Animals↗

Neonatal treatment with monosodium glutamate does not alter grooming behavior induced by novelty or adrenocorticotropic hormone.

The increased grooming behavior observed in a novel environment has been attributed to release of peptides derived from proopiomelanocortin (POMC), such as ACTH, alpha-melanocyte-stimulating hormone (alpha-MSH), or beta-endorphin, which themselves can elicit grooming. This is because novelty-induced grooming is attenuated both by hypophysectomy and by antiserum to ACTH injected into the cerebral ventricles. Administration of monosodium glutamate (MSG) to neonatal rats destroys neurons in the arcuate nucleus of the hypothalamus, depleting the brain of POMC peptides, and also hypothalamic dopamine and choline acetyl-transferase activity. Neonatal MSG treatment did not significantly alter the grooming scores of adult rats in either home or novel environments compared to saline-treated animals. There were also no differences between MSG-and saline-treated rats in the grooming scores observed following graded doses of ACTH1-24 (0.2-1.0 micrograms) administered intracerebroventricularly. Thus if increased grooming in the novel environment is due to release into the ventricles of ACTH, alpha-MSH, beta-endorphin, these peptides more likely derive from the pituitary rather than from brain cells, although the failure of the MSG treatment to produce quantitative depletions of cerebral POMC peptides, especially in the brain stem, leaves open the latter possibility.

Adrenocorticotropic Hormone↗

The influence of neurotensin, naloxone, and haloperidol on elements of excessive grooming behavior induced by ACTH.

Naloxone, haloperidol, and neurotensin suppress ACTH-induced grooming. The suppressive effects of naloxone and of haloperidol on ACTH-induced grooming are observed following subcutaneous as well as intracerebroventricular administration. The suppression of ACTH-induced grooming by these drugs is not accompanied by a change in the relative distribution of grooming elements. From previous data and from the results of the present study it is suggested that the underlying substrate involved in ACTH-induced excessive grooming may differ from that of bombesin-induced grooming.

Adrenocorticotropic Hormone↗

Excessive grooming induced by the administration of codeine and morphine.

Previous studies have shown that peripheral injections of the codeinone RX 336-M can induce excessive grooming in rats in an age-dependent fashion. The present experiment demonstrates that codeine, itself, also induces excessive grooming but with apparent equal effectiveness at all ages tested. Because of the structural similarity between codeine and morphine, the effects of intraperitoneally administered morphine were examined as well. Morphine was found to produce a temporal course of excessive grooming quite different from that produced by codeine. Morphine did not affect grooming in the first half (30 min) of the observation period, but accentuated it, briefly, for the next 30-45 min. Intraventricular administration of codeine at doses of 0.3 or 1.0 microgram/3 microliter had no effect on grooming in animals previously shown to demonstrate excessive grooming in response to either dose.

Aging↗

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↗

Body temperature and grooming in the Mongolian gerbil.

The Mongolian gerbil, Meriones unguiculatus, like many other rodents, releases a complex mixture of pigments and lipids from the Harderian gland during a self-groom. The material exits from the external nares of the nose, is mixed with saliva, and spread widely over the pelage. Cold temperatures, especially, are effective in initiating grooming. A self-groom is associated with an increase in body temperature (compensatory thermogenesis in the cold). In addition to acting as a chemosignal, the Harderian material serves two major homeostatic functions: (1) the lipids on the pelage act to insulate the animal against cold and wetness, and (2) the lipids and pigments darken the pelage and increase radiant absorption. Body temperature is thus maintained at a higher level than would otherwise be the case. The amount of Harderian material found on the pelage varies with Harderianectomy, sandbathing, and ambient temperatures. Animals prevented from sandbathing accumulate excess lipids on the pelage, and cold temperatures facilitate the acquisition of lipids on the pelage. Under hot temperatures the grooming of Harderian substances is repressed and the frequency of sandbathing is increased. Thus pelage lipids are reduced in two ways. The amount of Harderian material released during an autogroom is inversely related to the ambient temperature, whereas the amount of saliva used for evaporative cooling is positively related to ambient temperature. The net effect is that pelage lipids are increased and maintained during cold conditions, and are reduced during hot conditions. In hot environments the gerbil switches from the spread of Harderian material for insulation to the spread of saliva for evaporative cooling. The gerbil optimizes its body temperature by varying the frequency of grooming and sandbathing, and by altering the amount of Harderian material and saliva released. Other species living in arid environments may use similar mechanisms to stabilize body temperature. Self-grooming is a critical behavior for meeting thermal needs, and is complexly integrated with related processes.

Adaptation, Physiological↗

Differential thermosensor control of thermoregulatory grooming, locomotion, and relaxed postural extension.

The rat displays three behaviors when surface and/or deep body temperatures rise above their normal levels: grooming, which spreads saliva over the fur to cool by evaporation; locomotion, which results in escape to a cooler environment; and relaxed postural extension, which reduces heat production by muscle activity and increases heat dissipation by increasing body surface area. Each of these behaviors can be elicited in pure or nearly pure form by localized warming of thermosensors in different brain areas. In addition, localized warming of superficial and deep peripheral thermosensors elicits grooming and locomotion, but little or no postural extension. Because of the marked differences in the behaviors induced by localized warming of different sets of thermosensors, it is concluded that sensors are linked to the effector mechanisms for these behaviors by relatively separate and independent pathways rather than a master controller through which sensory signals are integrated to produce a unitary error signal that drives all responses. When more generalized hyperthermia in natural heat stress situations affects mainly superficial thermosensors, as during the initial phase of exposure to ambient heat, grooming and locomotion are predominant and extension is absent or weak. When deep hyperthermia is predominant and superficial hyperthermia is relatively weak, as occurs after escape from ambient heat to thermoneutrality or during and following exercise in a thermally neutral environment, the resulting predominant activation of deep sensors elicits marked postural extension, but little or no grooming or locomotion. When both superficial and deep sensors are hyperthermic, as in the later stages of exposure to ambient heat, all three behaviors are present, although locomotion is somewhat greater and grooming is somewhat less than during predominantly superficial hyperthermia, and extension is somewhat less than during predominantly deep hyperthermia. As a consequence, grooming is maximal during predominantly superficial hyperthermia, locomotion during mixed deep and superficial hyperthermia, and extension during predominantly deep hyperthermia. It is concluded that the relatively fractionated control of the rat's thermoregulatory behaviors by different groups of central and peripheral thermosensors makes possible a differentiation of responses to different distributions of hyperthermia between superficial and deep sensors that optimizes cost-benefit trade-offs in different kinds of heat stress.

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↗