Motivational Systems of Agonistic Behavior in Muroid Rodents
Motor Patterns of Submission Page 20



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Many of the motor patterns of submission are the same as those of defense. These include freezing (ie, crouch), flight, defensive upright and sideways and quadrupedal postures, tail-raising, squealing, teeth-chattering, tail-rattling, defense pheromone secretion, and hormone secretion from the adrenal medulla and cortex. Details may be found in the section on defense.

The full submissive posture is characteristic of submission and consists of a variant of freezing in which the animal rolls onto its back with its feet in the air and remains motionless. It is not usually reported as a response to predators, but has been reported as a response to consociate offense in the following species: R norvegicus and Me auratus [Grant and Mackintosh, 1963], M musculus [Scott and Fredericson, 1951], D groenlandicus [ Allin and Banks, 1968], R villosissimus [Begg, 1975], Ps obesus [Daly and Daly, 1975], S fulviventer and hispidus [Peterson and Helland, 1978], P leucopus [Sheppe, 1966], On leucogaster [Clark, 1962], Mi agrestis [Clarke, 1956], O irroratus [Davis, 1972], Cri cricetus [Eibl-Eibesfeldt, 1953a], P maniculatus and californicus [Eisenberg, 1962], Clethrionomys sp var [Johst, 1967], Mi ochrogaster and pennsylvanicus [Krebs, 1970], and Mi pinctorum [Novak and Getz, 1969]. Descriptions of full submissive posture are absent from studies of fighting in species of Lemmus, Meriones, and Apodemus.

In a few muroid rodent species there is a submissive upright posture which includes the immobility of the full submissive posture but the stance of a defensive upright posture, apparently a hybrid form of the two motor patterns. It has been described in M musculus [Grant and Mackintosh, 1963], Mi agrestis [Clarke, 1956], and species of Peromyscus [Eisenberg, 1962] and Apodemus [Montgomery, 1978]. Apparently in the Microtus sp, Apodemus, and M musculus, the submissive upright posture is a substitute for the full submissive posture which is rare by contrast.

Another characteristic motor pattern of submission is a "piping" vocalization, usually ultrasonic in frequency. It consists of a pure, high frequency tone, held constant during long respiratory expirations or perhaps trilled, and repeated on many successive respirations, in almost continuous fashion. It often accompanies a freezing crouch or full submissive posture. In some species such as A flavicollis [Hoffmeyer and Sales, 1977] and R norvegicus [Sales and Pye, 1975], the piping is completely ultrasonic. In R villosissimus, the base frequency is audible at 12 kc (one octave lower than that of R norvegicus), although it is similar in all other respects [Begg, 1975]. Many muroid rodent species produce similar vocalizations in a sexual context, but whether they produce ultrasonic "piping" in submission has not been determined.

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