Free-operant comparisons of wild and domestic Norway rats.

Wild and domestic rats (Rattus norvegicus) were compared on nondifferential appetitive variable interval responding, discrimination, and discrimination reversal procedures. The effects of strain, sex, deprivation, preexperimental handling, and sessions on response rate were examined. Performances during the 60 days of variable interval training showed prolonged increases over that period for all strain-sex groupings, with domestic subjects responding at higher rates than wild. Males also tended to respond at higher rates than females. During the discrimination procedure wild subjects showed more resistance to extinction than did the domestic subjects, although these differences generally diminished after the 30 days of training. During the discrimination reversal procedure domestic subjects reversed their responding pattern more readily than did the wild subjects, and domestic females reached criterion significantly sooner than domestic males. The effects of deprivation and handling were not significant during the experimental procedures.     

Burrows of wild and albino rats: effects of domestication, outdoor raising, age, experience, and maternal state

Burrows dug by albino rats were compared with those of wild Norway rats in an outdoor pen and in observation chambers in the laboratory. Burrows, in terms of measurements, configurations, or sequential development, were indistinguishable in wild and domestic rats. Burrowing for both wild and domestic rats was unaffected by raising in outdoor burrows, by availability of nesting material, or by pregnancy. Prior experience in burrowing did make it more efficient in a second trial, which suggests that learning may have a limited role in what appears to be a behavior with a strong genetic basis. Feralization of domestic rats in the outdoor pen was especially productive in answering claims of degeneracy in these animals: Albino rats were hardy throughout climatic extremes, they maintained a stable population for two years, they constructed and lived in burrows, and they showed a vaiety of wild-type behaviors.     

Differential effects of environmental enrichment on the open-field behavior of wild and domestic Norway rats.

Early postweaning experience in an enriched environment had a greater influence on the open-field behavior and body weight of wild Norway rats than of their domestic counterparts. Genetic changes accompanying the domestication process may have reduced the relative impact of postweaning experience on the development of the domestic rat's response to changes in its environment.     

Serotonin metabolism and serotonergic receptors in Norway rats selected for low aggressiveness to man

Selection of Norway rats (24–27 generations) for low aggressiveness to man resulting in the loss of aggressive responding to handling markedly influences the brain serotonergic system. In “domesticated” Norway rats levels of serotonin in the midbrain and hypothalamus and 5-hydroxyindole acetic acid in the hypothalamus were higher than in non-selected aggressive rats. The activity of the key enzyme in serotonin biosynthesis, tryptophan hydroxylase, in midbrain of rats with genetically determined lack of aggressiveness to man was higher than in aggressive animals, although there was no difference in tryptophan hydroxylase activity in the hypothalamus. B_max and K_D of [³H]spiperone-specific binding in frontal cortex membranes were increased in tame rats. No significant differences in B_max and K_D were found between “domesticated” and aggressive rats in [³H]serotonin binding in the frontal cortex.     

Selection for reduced aggressiveness towards man and dopaminergic activity in Norway rats

The brain dopaminergic system is involved in the process of long-term selection for reduced aggressive reaction towards man in Norway rats. The dopamine levels in the striatum as well as the nucleus accumbens with the tuberculum olfactorium were significantly lower in domesticated rats than in their wild counterparts. A substantial decrease was found in homovanillic acid level in the n. accumbens and tuberculum olfactorium. Specific binding of [³H]spiperone which labels D-2 dopamine receptors was higher in the mesolimbic structure of tame rats, whereas binding of [³H]SCH 23390 (D-1 receptors) was unchanged in this area. No substantial differences were detected in D-1 and D-2 binding in striatum. Apomorphine (0.3 mg/kg) elicited less locomotion in tame animals, reflecting a decrease of sensitivity of postsynaptic dopamine receptors. Tame rats showed fewer aggressive contacts in a foot-shock test than wild rats and the D-2 receptor antagonist sulpiride (25 mg/kg) significantly decreased the foot-shock aggression only in wild rats. Therefore, domestication, which diminishes defensive behavior and emotional reactivity of animals, is associated with decreases of dopamine level in the striatum, changed metabolism of dopamine in mesolimbic system, and an alteration in density and senstivity of D-2 receptors.     

Agonistic behavior between wild and genetically sterile male Norway rats (rattus norvegicus)

Genetically sterile male Norway rats, Rattus norvegicus, were tested in the laboratory to determine both 1) behavioral characteristics, and 2) the ability of sterile males to compete aggressively and sexually with wild Norway rat males. Sterile males were larger in weight, more frequently dominant, won as many fighting encounters, were as aggressive as wild males, and mounted females more frequently. Behavioral activities were similar for both strains when compared under laboratory conditions with no apparent abnormal behavior exhibited by the sterile males. Use of sterile males in biological control programs is discussed.     

How does the ecological foraging behavior of desert kangaroo rats (Dipodomys deserti) relate to their behavior on radial mazes?

Experiment 1 showed that laboratory-reared desert kangaroo rats, like domestic Norway rats, efficiently search for food on a radial arm maze (RAM) by avoiding revisiting arms within a trial. By placing an RAM on the floor so the animals could approach food from any direction, Experiment 2 tested whether efficient search by kangaroo rats was based on tactics of distance minimizing, central-place foraging, trail following, or meandering. In contrast to the dominant trail-following tactic of domestic Norway rats (Hoffman, Timberlake, Leffel, & Gont, 1999), kangaroo rats tended to distance minimize, whether maze arms were present or not. Experiment 3 indicated that kangaroo rats treated a floor configuration of eight food cups as two patches of four, based on beeline travel between patches and meandering within them. We conclude that similar performance in an elevated RAM by different species can be based on different tactics, and we suggest that a laboratory apparatus can be used to cast light on niche-related mechanisms.     

Genotype and environment in the social interactions of wild and domestic “Norway” rats

Wild rats, Rattus norvegicus, (a) trapped as adults or (b) of the second generation in captivity (lab-wild), and domestic rats of two strains, were studied for 28 days in artificial colonies in large cages with attached nest boxes. Controls were kept in mated pairs in small cages. Each colony consisted of six males and six females. The interactions of the males in six colonies of trapped rats were highly “stressful;” 61% died; and most of the survivors lost weight and had greatly enlarged adrenals. In each colony, however, there was a male (an alpha) that gained in weight and spent much time, during the dark hours, in the open on the floor of the cage; and in three colonies there were also other males (“betas”) that gained in weight. The adrenals of alphas and betas weighed about the same as those of the controls. In one of the 12 colonies of domestic rats one male behaved like a wild male; but in the other colonies the males gained in body weight and their adrenal weights resembled those of the controls. In three colonies of lab-wild rats 22% of the males died, but there was no evidence of males of different status. The findings confirm that the “agonistic” behavior of domestic rats is usually much attenuated in comparison with that of the wild type; a number of methodologic implications are discussed.     

Further experiments on the social interactions of domestic “Norway” rats

Domestic rats, Rattus norvegicus, aged nearly one year, were studied in artificial colonies in large cages with attached nest boxes. As in previous experiments on younger domestic rats, but not those on wild rats, the colonies were peaceful. Questions concerning the “aggressive” or “agonistic” behavior of domestic rats are further discussed.     

Perceptual and acoustic evidence for species-level differences in meow vocalizations by domestic cats (Felis catus) and African wild cats (Felis silvestris lybica)

To test for possible anthropogenic selection effects on meows in domestic felids, vocalizations by domestic cats (Felis catus) were compared with cries by their closest wild relative, the African wild cat (Felis silvestris lybica). Comparisons included analysis of acoustic characteristics and perceptual studies with human (Homo sapiens) listeners. The perceptual studies obtained human listener ratings of call pleasantness. Both the acoustic and perceptual comparisons revealed clear species-level differences: The domestic cat meows were significantly shorter in mean duration than the wild cat meows, showed higher mean formant frequencies, and exhibited higher mean fundamental frequencies. Human listeners at all levels of experience and affinity for cats rated domestic cat meows as far more pleasant sounding than wild cat vocalizations. These results are consistent with a model of cat domestication that posits selective pressure on meows based on human perceptual biases.     

Laboratorizing the wild rat (Rattus norvégiens)

Traditional difficulties in procuring wild rats can be overcome by trapping in an unstable environment (e.g., land-fill) where the natives do not fear strange objects such as traps. Techniques for trapping, transferring to, and maintaining in the laboratory are outlined, and apparatus for handling and housing, including the breeding situation, is described. Conclusion: Simple precautions render the wild rat nearly as easy to breed and use in conventional laboratory tasks as domesticated rats.     

Spatial Working Memory and Reference Memory of Brown Norway and WAG Rats in a Holeboard Discrimination Task

To determine whether working memory (WM) and reference memory (RM) represent different aspects of spatial memory, albino WAG rats and the pigmented Brown Norway rats were tested in the acquisition, retention, and reversal of spatial orientation tasks in the holeboard, which allows the simultaneous assessment of WM and RM. Putative nonmnemonic factors, such as the speed of visiting the holes or the development of a search strategy (preferred sequence of visiting the baited set of holes) were also evaluated because they might influence WM and RM performance. The WM performance of Brown Norway rats was generally worse than that of the WAG rats. The reverse was true for the RM performance. Correlation analysis supported the notion that these two measures are independent. Differences in the speed of visiting the holes and in the development of a preferred sequence of visiting the baited set of holes could not explain the strain differences in WM and RM performance. Because spatial WM and RM appear to be independent measures in the holeboard, this task could be used to investigate whether different neural substrate(s) underlie these two memory components.     

Tyrosine hydroxylase activities in the brains of wild Norway rats and silver foxes selected for reduced aggressiveness towards humans

Tyrosine hydroxylase (TH) activity was measured in brains from Norway rats and silver foxes showing wild type aggressiveness and from their counterparts selected over 20-25 generations for reduced aggressiveness towards humans (tameness). TH activity in the brain stem and cortex was increased in tame animals of both species compared with aggressive counterparts. Selection increased hypothalamic TH activity in foxes, but decreased it in rats. There was no difference in TH activity in corpus striatum between the tame and aggressive animals. Fetal TH activity in the posterior part of the brain was higher in tame than aggressive rats at day 20 of embryogensis. Increased TH activity in the brain stem and cortex of adult aggressive rats was observed after treatment of their mothers with hydrocortisone on the days 16 and 18 of pregnancy. This elevation in TH activity was associated with attenuation of the defense behavior of aggressive rats. The data suggested that alterations in neural TH activity in tame rats and foxes may be part of the neurochemical basis of their behavioral phenotype which is developed by selection. © 1994 Wiley-Liss, Inc.     

Early stimulation and patterned responding in rats

This experiment was designed to determine the extent to which performance differences on learning tasks arising from infantile handling reflect learning or emotionality differences. Handled and nonhandled rats were trained and tested in a controlled operant situation where a response on every other trial yielded reinforcement. A key light was turned on for 10 sec during each trial and was off during the 10-sec intertrial interval. The reinforced (S+) trials alternated in a regular sequence with nonreinforced (S–) trials. The pattern of S+ and S– trials recycled continuously throughout the session of 160 trials. The handled rats emitted more responses during both S+ and S– trials than did the nonhandled rats. However, when they were compared on the percentage of emitted responses that were reinforced, there was no significant differences between the two groups. These results were interpreted to suggest that infantile handling does not directly facilitate associative learning in rats.This research was aided by a grant from the National Research Council of Canada, 67-0247. I thank Linda Easton and Beverly Clark for their help in collecting and analyzing the data.     

The persistence of some effects of handling in infancy on the behaviour of the adult rat

An attempt was made to discover how persistent the effects of handling before weaning are on activity, grooming, emotional elimination and learning in adult rats; whether there is an interaction of the effects of handling and the effects of cage size on the behaviour of rats; and whether the daily removal of the mother during nursing has any effects on the pups.

Since all differences between handled and undisturbed animals found at 50 or 100 days were found at 220 days, it was concluded that the effects of handling in infancy are persistent. The hypothesis that cage size and removal of the mother during nursing significantly affect adult behaviour was not supported by the results.     

Studies of Emotional Reactions VI. Cardiac Responses

Wild-strain male house mice were fostered at birth onto conspecifics, deer mice, or domestic Norway rats. In adulthood, their flesh-eating preferences were tested by allowing them to feed from freshly asphyxiated conspecifics and either deer mice or rats. The mice ate significantly more conspecific flesh than contraspecific flesh, except when the flesh offered was that of the contraspecies upon which the mouse had been fostered at birth. The failure of cross-species-fostered mice to discriminate between their own species and their foster species in flesh-preference tests is attributed to their having learned early in life to respond to the foster species in the same way they normally respond to conspecifics, that is, by approaching them, investigating them, and, when finding them dead, feeding upon them.     

Short-term effects of postnatal manipulation on central beta-adrenoceptor transmission

Neonatal handling is known to induce long-lasting changes in behavioral and neuroendocrine responses to stress. Since the central noradrenergic system participates in the adaptive responses to stressful conditions we have analyzed the effects of postnatal handling on beta-adrenoceptor binding sites and isoprenaline- and forskolin-stimulated cyclic AMP accumulation in cerebral cortex, hippocampus and cerebellum of rats at 1 and 3 months of age. Handled animals showed reduced emotional reactivity and lower ACTH and corticosterone secretion after stress. Binding studies using [(3) H]CGP12-177 revealed increased beta-adrenoceptor binding sites in handled rats in cerebellum and cerebral cortex with no changes in hippocampus, and decreased affinity in all cerebral regions. Handling reduced basal levels of cyclic AMP in hippocampus and cerebellum but not in cerebral cortex. The concentration-response curves of cyclic AMP to isoprenaline were displaced to the right in cerebellum of handled rats without differences in Emax; however, Emax was significantly reduced in cerebral cortex and hippocampus. Direct stimulation of the catalytic subunit of adenylyl cyclase by forskolin reduced the efficiency in hippocampus and cerebellum, but not in cerebral cortex of handled animals. It is concluded that neonatal handling reduces the binding properties of beta-adrenoceptor and its primary biochemical responses in the young rat brain, which may account for the reduced responsiveness to stress attained in the handled rats, and may explain the persistence of the effect. The present study emphasizes the role of the central noradrenergic system in modulating the behavioral and neurendocrine responses to neonatal handling.     

Infantile stimulation and its effects on frustration- and fear-motivated behavior in rats

Infant rats that were either removed from the nest each day (handled) or left undisturbed (nonhandled) were, in adulthood, given 72 food-reinforced runway acquisition trials followed by 24 trials of extinction training with or without shock. Handled and nonhandled control animals were given runway training without food reinforcement. Reinforced rats ran faster than nonreinforced rats, and handled rats ran faster than nonhandled rats during the initial trials of runway acquisition irrespective of the reinforcement condition. Nonhadled rats stopped running sooner than handled rats when shock was introduced in the goalbox, but differences between handled and nonhandled rats given extinction training without shock were small. Results of a second experiment showed no differences between handled and non-handled rats in the magnitude of the depression effect after an incentive shift. It was concluded that infantile handling had little effect on frustration-motivated behavior, but did affect fear-motivated behavior.     

Experiments on neophobia in wild and laboratory rats: a reevaluation.

Two experiments using food-container avoidance as an index of neophobia are reported for two strains of laboratory and one strain of wild rats. In Experiment 1 rats were fed from a single familiar container until their consumption had stabilized. Upon replacing the familiar container with a novel container, the latency of all three strains to begin feeding increased. In Experiment 2 rats were offered a choice between a familiar and a novel container containing identical food. Though there was considerable individual variation among the three strains, the wild strain was more reluctant to eat from the novel container than a hooded laboratory strain, which, in turn, was nore reluctant than an albino laboratory strain. Nonetheless, all three strains showed an initial avoidance of the novel container. It was concluded that both wild and laboratory strains are neophobic and that strain differences are ones of degree, not of kind.     

Early tutoring and adult reproductive behaviour in female domestic canary (Serinus canaria)

We studied the effect of early tutoring on the subsequent sexual preferences and reproductive activity of female domestic canaries (Serinus canaria). Young female canaries were exposed during the first 4 months of life to songs of either domestic or wild canaries. When adult, these females were again exposed to domestic or wild songs. In the first experiment, the sexual responses of the females to unfamiliar domestic and wild songs were quantified with the copulation solicitation display (CSD) assay. In the second experiment, the same females were tested again with modified tutoring songs. In the third experiment, song stimulation of nest-building and egg-laying was studied. Domestic-strain-tutored females gave more CSDs to domestic than to wild songs. In contrast, wild-strain-tutored females showed no sexual preference. We propose that the sexual preference of adult domestic-strain-tutored female canaries for domestic songs is the consequence of learning and categorisation processes. The discrepancy between the results of the domestic-strain-tutored females and those of the wild-strain-tutored females suggests that female canaries have a predisposition to learn songs of their own strain rather than songs of an alien strain. In the third experiment nest-building and egg-laying activities appeared to be unaffected by early tutoring conditions: there was no significant differential effect of the different tutoring and exposure conditions on nest-building and egg-laying scores. Mate attraction and stimulation of females’ reproductive activity appear to be two separate functions of male song, which may have been shaped by different evolutionary constraints. Received: 24 July 1999 / Accepted after revision: 22 February 2000