Conditioned discrimination of airborne odorants by garter snakes (Thamnophis radix and T. sirtalis sirtalis)

For snakes, the nasal chemical senses are critical in intraspecific communication and prey recognition. Although it is known that garter snakes can respond differentially to airborne odorants, no previous study has demonstrated that snakes can learn a task with airborne odors as discriminative stimuli. In Experiment 1, 7 plains garter snakes (Thamnophis radix) were trained in a two-choice apparatus to move into a compartment containing lemon-scented chips for a food reward. All 7 snakes improved performance when the first 10 and last 10 trials of the 100 trials of conditioning were compared. Four of the snakes were subsequently trained to move away from the scented compartment into the unscented compartment. The 4 snakes rapidly learned this reversal. In Experiment 2, 7 common garter snakes (T. sirtalis sirtalis) were trained to traverse a two-choice maze with the presence or absence of amyl acetate odor as the conditioned stimulus. The snakes were pretested for odor versus nonodor preference and were trained to go to the initially nonpreferred stimulus. Of the 7 snakes, 5 achieved a predetermined criterion (two training sessions with cumulative correct responding above the .05 confidence level) within 85 trials.     

Neonate plains garter snakes (Thamnophis radix) are attracted to conspecific skin extracts

Several species of snakes aggregate in areas previously marked by conspecifics. However, species differences and sources of such chemical signals have received little attention. Methylene chloride-soluble substances from shed epidermis of adult plains garter snakes attracted neonate conspecifics, whereas similar extracts from a congener and a viperid did not. The functional significance of such conspecific aggregation may relate to location of hibernacula, foraging areas, osmoregulation, or predation avoidance.     

Immobility and supination in garter snakes (Thamnophis elegans) following handling by human predators

A common response of animals to physical restraint is tonic immobility. The authors observed the behavior of garter snakes, Thamnophis elegans, in the field to determine the frequency of immobility as a response to handling. Most snakes fled after release, but the remainder remained immobile, sometimes on their backs (supination), for up to 10 min. Immobility was seen most often in pregnant snakes, which also were more likely than other snakes not to have moved before capture. Failure to move, either before or after capture, might be a consequence of the limited locomotory ability of gravid snakes. However, the lack of observations of interactions between snakes and their natural predators impedes researchers' understanding of the antipredator value, if any, of tonic immobility.     

Laboratory observations of trailing behavior in garter snakes.

Using a Y-maze apparatus, nine garter snakes were trained to follow an earthworm-extract trail. Correct trailing improved significantly as training progressed. All snakes were able to attain a trailing criterion of 8 out of 10 correct trials by the termination of training. For each trial, the number of tongue flicks each snake made was recorded and tongue flick rate calculated. Tongue flick rate was found to be highly characteristic and constant for individual subjects. In addition, running speed and tongue flick rate were found to be significantly correlated.     

Evolutionary persistence of chemically elicited ophiophagous antipredator responses in gartersnakes (Thamnophis sirtalis)

The ability to detect and respond to potential predators is key for the survival of individuals, but this ability is sometimes lost via relaxation of antipredator behavior when prey species are separated from predators. Adult and predator-na?ve neonate gartersnakes (Thamnophis sirtalis) from mainland and insular sites where they do and do not occur with ophiophagous (snake-eating) snakes were tested to determine if responses to such predators have been lost, reduced, or retained, and what might be causing differences in such responses. Our data indicate that, overall, adult snakes from populations syntopic with ophiophagous milksnakes are more responsive to chemical stimuli from milksnakes than adults from areas where they are not syntopic with milksnakes, whereas there were few differences with neonate gartersnakes. Experiments with neonates with or without periodic ophiophagous snake chemical experience over several weeks showed that gartersnakes with such experience became more responsive to ophiophagous cues rather than habituating to them. Such evidence of both genetic and experiential factors underlying the antipredator responses to ophiophagous snakes indicates that antipredator responses have persisted despite separation of predator and prey.     

Snake aggregation pheromones: source and chemosensory mediation in western ribbon snakes (Thamnophis proximus)

Aggregation with conspecifics is a robust phenomenon among snakes that is mediated by chemical cues, although the source of such cues is unknown. Adult western ribbon snakes (Thamnophis proximus) discriminated between skin lipid extracts from conspecifics and those from a sympatric heterospecific snake, the corn snake (Elaphe guttata). Additionally, ribbon snakes were attracted to conspecific skin lipid-marked shelters when paired with control-marked shelters, but no preference was exhibited when shelters marked with E. guttata skin lipids were paired with control-marked shelters. Furthermore, prevention of chemical access to the vomeronasal organs of T. proximus eliminated the ability to respond to conspecific skin lipids. These results indicate that epidermal lipids are the source of chemical cues mediating attraction to shelters marked by conspecifics and that such signals are detected by the vomeronasal system.     

Effect of food competition on aggregation: evidence for social recognition in the plains garter snake (Thamnophis radix).

Competition for food and aggregating behavior were examined in plains garter snakes (Thamnophis radix), in 11 groups of 1 noncompetitor and 2 competitors. Aggregation data were obtained in 48 daily scans. During food competition, success at obtaining food was asymmetrical across pairs. For aggregation, competitors were mostly associated with noncompetitors. After removal of noncompetitors, competitors were more likely to be solitary, but associations between competitors increased. After a 5-month separation, 5 of the original groups were reconstituted; aggregation patterns similar to the 1st experiment were observed. In a 3rd experiment, unfamiliar animals with and without competitive experience did not aggregate preferentially. Garter snakes discriminate on the basis of sex and species; in this study individual relationships (prior competition) affected aggregation, and the data suggest individual recognition both occurs and persists over time.     

Organization of mating behavior in male hamsters (Mesocricetus auratus)

Factor analysis was applied to standard measures of sexual behavior in 73 male hamsters (Mesocricetus auratus) as they interacted with hormone-primed females. The results suggest that 5 factors, or conceptual mechanisms, function in the organization of the behaviors observed in the first 2 copulatory series. Of these, the 3 that relate to the behaviors in the first copulatory series were compared to those emerging from prior analyses of other rodents. These comparisons revealed similarities and differences in factor structure across species. Whereas all of these analyses identify factors related to the initiation and efficiency of copulatory behavior, hamsters seem to differ from other species in the measures that best define these factors. In addition, the copulatory rate factor that has been prominent in previous analyses of rats seems to be absent in hamsters. These results suggest that male sexual behavior in hamsters is organized differently from that in other rodents. In more general terms, they suggest that even species with generally similar copulatory patterns can show significant differences in behavioral organization, in turn suggesting the need for additional factor analytic studies to better establish the extent of these species differences.     

Life-span changes in the copulatory behavior of male rats (Rattus norvegicus)

The copulatory behavior of 5-, 10-, 15-, and 20-month-old sexually naive male rats (Long-Evans) was studied during 2-hr tests with receptive females. There was no apparent change in sexual arousal as measured by latency to initiate copulation across age, with subjects from all groups exhibiting comparable latencies to first mount and first mount with intromission. The numbers of ejaculations achieved were also similar across ages. Significant age differences were found for frequency of mounts, with 20-month-olds having the highest mean frequency. The persistent mounting by older males appeared to account for significant group differences in interintromission interval and ejaculation latency. It is suggested that motor deficits may impair the ability of older males to achieve intromission, increasing the number of mount bouts as well as the number of mounts per bout, thus extending the length of each copulatory series.     

Seasonal changes in open-field behavior in wild male meadow voles (Microtus pennsylvanicus)

Open-field behavior of free-living meadow voles was measured in males held overnight in the laboratory. Movement variables were positively correlated, and had negative associations with grooming and freezing. Parameters including activity, freezing, urinating, and grooming showed annual fluctuations related to the reproductive season. Together with the results of a previous study showing castration of wild voles results in altered open-field behavior, these results emphasize the role of testicular hormones in influencing this behavior. Factor analysis identified an activity component accounting for 39% of the variation, but other parameters were little simplified by this procedure. Most factors cycled annually, and significant variation was found in all eight factors during the 4-year sample. Open-field behavior varied between different reproductive and age segments of the population, and may be related to population processes such as dispersal.     

Sexual Pavlovian conditioned approach behavior in male Japanese quail (Coturnix coturnix japonica)

A localized visual stimulus presented immediately prior to access to a female conspecific stimulated approach behavior in male Japanese quail after several conditioning trials. Development of this conditioned approach behavior was observed with two different types of signal lights, 10-s and 30-s signal durations, large and small experimental chambers, and with male birds housed continually in the conditioning chambers or only placed in the chamber for brief daily sessions. Conditioning also resulted in shorter latencies to initiate copulation in males given access to a female following the signal light, in comparison with males that received access to a female unannounced by the light. Although some aspects of the conditioning experience were retained over a 6-week interval, some loss of the behavior was also observed. The conditioned behavior also decreased with repeated extinction trials, during which the signal light was presented in the absence of access to a female conspecific. The results are discussed in relation to other aspects of the social behavior of Japanese quail and other studies of sexual conditioning.     

Effects of male pre-ejaculatory vocalizations on female receptive behavior in the rat (Rattus norvegicus)

Both male and female rats (Rattus norvegicus) emit ultrasounds during copulation. Bursts of ultrasounds from males that occur as ejaculation approaches are lower in frequency, longer in duration, and louder than those that appear earlier in the ejaculatory series. We attempted to determine if these pre-ejaculatory calls affect female sexual behavior. The behavior of females paired with devocalized males was compared with that of females paired with intact males, and in a second experiment tape-recorded vocalizations were played to females paired with devocalized males. Females were less receptive when paired with devocalized males; they were less likely to remain stationary when males attempted to mount. Playback of both types of calls restored female immobility toward control levels, although pre-ejaculatory calls were more effective than mating ones. Pre-ejaculatory calls restored running and training levels somewhat toward control level but to a lesser extent than female immobility.