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Preputial gland activity is related to social experience (Hucklebridge, Nowell & Wouters, 1972) and these glands are thought to affect the fighting behaviour of mice through the release of an aggression-promoting pheromone (Mugford & Nowell, 1970). The coagulating glands are thought to be the source of an aggression-inhibiting pheromone (Haug, 1971). The present study provides direct evidence for the function of these glands by assessing the aggression-inducing properties of combinations of the gland secretions with bladder urine or water.
A number of 4-month-old male T.O. albino mice were killed. The preputial and coagulating glands were dissected out and the gland contents were gently squeezed into bladder urine or into the same amount of distilled water. Six test substances were used: (1) bladder urine, (2) bladder urine+preputial gland secretion, (3) bladder urine + coagulating gland secretion, (4) water, (5) water + preputial secretion, and (6) water + coagulating gland secretion.
The 15 mice used
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SUMMARY
The aversive properties of urine of castrated male mice receiving varying doses of testosterone phenylpropionate was assayed using males housed in groups of six as subjects. The aversive efficacy of the urine was found to rise with an increase in androgen levels. There was a delay of 5 days before the exogenous androgen exerted any effect on the aversive properties of the urine. This delay indicated that the aversive factor might be a pheromonal substance released from an androgen-dependent tissue, rather than being an excreted androgen metabolite.
The results are discussed in terms of androgen levels and possible territorial functions.
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Male mouse urine contains an androgen-dependent aversive pheromone which discourages prolonged investigation, by male conspecifics, of an area marked with such urine (Jones & Nowell, 1973a). The source of this pheromone lies in the coagulating glands, whose secretion, when combined with bladder urine, causes avoidance (Jones & Nowell 1973b). The present study describes an attempt to inhibit the release of the aversive pheromone of male mice by treatment with the potent anti-androgen cyproterone acetate. Thus the open-field responses of male mice to the urine of either oil-injected or cyproterone acetate-injected males were compared. Some confirmation of the inhibitory action of cyproterone acetate upon sex accessory organ responses to endogenous androgens was gained by recording the weights of the ventral prostate, preputial and coagulating glands.
Twenty 24-day-old male t.o. albino mice were divided into two groups of ten; the first group received s.c. injections of cyproterone acetate (2 mg/mouse)
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SUMMARY
In an attempt to clarify the role of gonadal function in the release of anti-aggression pheromone into the urine of female mice, two experiments were carried out. Each involved recording the aggressiveness of 'fighter' male mice when presented with castrated male 'opponents' which had been treated either with urine from one of four categories of female donor mice, or with water.
In the first series of experiments it was shown that the urine from oestrous and dioestrous mice contained similar amounts of anti-aggression activity. Both spaying and testosterone propionate (TP) injection of the female urine-donors abolished this activity.
In a second series of experiments urine from spayed donors injected with TP greatly increased the aggression response to a level which exceeded that obtained with the urine of TP-treated intact mice. High doses of oestrogen resulted, in the spayed urine-donors, in an actual increase in aggression response. Both treatments increased clitoral gland weight, and it is concluded that androgen or high levels of oestrogen can stimulate the clitoral gland of the female, a possible consequence being a release of aggression-promoting pheromone.
The aggression-inhibiting pheromone, which depends on the ovary for its release, is probably the product of some other tissue, as yet unidentified.
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The preputial glands are modified sebaceous glands whose size and function are primarily dependent upon the presence of androgens. For this reason they are well developed in the male and vestigial in the female. Their activity is also dependent upon a factor or factors from the pituitary, though the identification of these factors is not yet established (Ebling, Ebling, McCaffery & Skinner, 1971). Experiments by Brain & Nowell (1970) have demonstrated that the isolated mouse, which is more aggressive than the grouped animal, has greater ventral prostate and preputial gland weights, and Mugford & Nowell (1971) have shown that there is a relationship between the presence of the preputial glands and aggressive behaviour, which might be maintained by the release of an aggression-promoting pheromone, in preputial sebum, as a result of hormonal stimulation.
In the course of these latter studies it became obvious that a wide range of preputial activity
