Rat 50 kHz Trill Calls Are Tied to the Expectation of Social Interaction.

Rats emit a variety of calls in the 40-80 kHz range (50 kHz calls). While these calls are generally associated with positive affect, it is unclear whether certain calls might be used selectively in certain contexts. To examine this, we looked at ultrasonic calls in 30-40 day old male rats during the expectation of either play or food, both of which are reinforcing. Behavior and vocalizations were recorded while rats were in a test chamber awaiting the arrival of a play partner or food over seven days of testing. Control groups were included for the non-specific effects of food deprivation and social isolation. Play reward led to an increase in 50 kHz vocalizations, generally, with specific increases in trill and "trill with jump" calls not seen in other groups. Expectation of food reward did not lead to a significant increase in vocalizations of any type, perhaps due to the young age of our study group. Further, rats that were food deprived for the food expectation study showed markedly lower calls overall and had a different profile of call types compared to rats that were socially isolated. Taken together, the results suggest that trill-associated calls may be used selectively when rats are socially isolated and/or expecting a social encounter.

Memory for surface objects in an arena by the horse (Equus ferus caballus) under saddle: Evidence for dual process theory of spatial representation.

Place memory, the ability to remember locations, is a feature of many animal species. This episodic-like memory is displayed in the foraging behavior of animals and has been studied in many different kinds of laboratory spatial tasks. A horse stallion, Equus ferus caballus, will create "dung-heaps or stud-piles" by defecation in the same place suggesting that the behavior is central to spatial behavior but to date there has been little investigation of horse olfactory/spatial behavior. The present study describes investigatory behavior of horses for objects on the surface of a riding arena. Horses under saddle approached objects on the arena surface that included small pieces of straw, fur, and paper and larger objects including clumps of debris and were especially interested in dung droppings left by other horses. Once an object was investigated by sniffing, it was usually not approached again during that outing but could be approached anew on the following day. Dung investigatory behavior and place memory were confirmed in a number of structured tests in which test-retest intervals were varied. The results are discussed in relation to the dual process theory that proposes that spatial representations central to adaptive behavior require both allocentric Cartesian spatial information and egocentric episodic-like information.

Does play shape hand use skill in rats?

Hand use is a widespread act in many vertebrate lineages and subserves behaviors including locomotion, predation, feeding, nest construction, and grooming. In order to determine whether hand use is similarly used in social behavior, the present paper describes hand use in the social play of rats. In the course of rough and tumble play sessions, rats are found to make as many as twenty different movements a minute with each hand for the purposes of manipulating a partner into a subordinate position or defending against a partner's attack. The hand movements comprise signaling movements of touching, offensive manipulating of a partner to control a play engagement, and defensive hand movements directed toward blocking, pushing and pulling to parry an attack. For signaling, attack and defense, hand movements have a structure that is similar to the structure of hand movements used for other purposes including eating, but in their contact points on an opponent, they are tailored for partner control. Given the time devoted to play by rats, play likely features the social rat behavior with the most extensive use of hand movements. This extensive use of hand movements for social play is discussed in relation to the ubiquity of hand use in adaptive behavior, the evolution of hand use in the play of mammals, and in relation to extending the multifunctional theory of the purposes of play to include the education of skilled hand movements for various adult functions including as feeding.

A naturalistic method to test depression: Anticipation of play.

The Wistar-Kyoto (WKY) rat was developed as a control for the spontaneous hypertensive rat but has subsequently also been used as a genetic animal model of depression due to its hyper-responsiveness to stress. We used anticipation of social reward (i.e., a play partner) to assess behavioural and vocal differences between the WKY and normal Wistar (WI) rats in the juvenile period. We found marked differences between groups; the WKY rats, were less active, vocalized less, and used significantly fewer types of 50-kHz calls in comparison to their WI counterparts. The animals were re-tested in adulthood and the same differences existed in overall activity, types of vocalizations and the behavioural vocal profiles used by the two groups of animals. These findings provide a robust baseline for an animal model of depression using a social paradigm. This paradigm may be useful to evaluate the efficacy of pharmaceutical interventions as potential treatments of depression in WKY rats.

Sniff, look and loop excursions as the unit of "exploration" in the horse (Equus ferus caballis) when free or under saddle in an equestrian arena.

Spontaneous locomotor behavior in a novel space reveals insights into an animal's world view or Umwelt. For example, in many animal species, spontaneous behavior in a novel environment is parsed into activities at a home base and excursions from the home base. Domestic horses (Equus ferus caballas) are frequently ridden for recreation or in performance events in an equestrian arena but there has been no description of horse behavior in an arena when they are unconstrained and "exploring" or when moving freely under saddle. The present examination of exploration provides insights into horse adaptive behavior more generally as well as insights into horse performance under saddle. Thoroughbred, American Quarter Horse and mixed-breed mares and geldings of various ages, with various degrees of training under saddle, and with varying familiarity with the arena were given 30-min tests in which they were at liberty to explore an equestrian arena. Additional 30-min tests were given in which horses explored the arena containing a tethered partner, or were ridden. Despite breed, sex, age and experience, behavior was similar. A horse spent most of its time near the entrance door of the area where it looked out, paced, and rolled. Periodic excursions formed loops. The outward leg of a loop was slow, often featured sniffing the ground, and ended with a head-raised, ears-forward look toward the far end of the arena. The homeward leg of a loop was made with lowered-head and ears-back and was relatively direct and fast. Successive loops could increase or decrease in size over a test period. If a partner was tethered at the far end of the area, a horse shifted its activity toward the partner. When horses under saddle were asked to make excursions into the arena but otherwise left unconstrained, they made loops, similar to that of freely moving horses. When ridden around the arena they returned more quickly to the near end of the arena than when leaving the near end of the arena. This organized home base/excursion behavior is discussed in relation to horse social structure and to its expression while under saddle.

Avoiding escalation from play to aggression in adult male rats: The role of ultrasonic calls.

Play fighting is most commonly associated with juvenile animals, but in some species, including rats, it can continue into adulthood. Post-pubertal engagement in play fighting is often rougher and has an increased chance of escalation to aggression, making the use of play signals to regulate the encounter more critical. During play, both juvenile and adult rats emit many 50-kHz calls and some of these may function as play facilitating signals. In the present study, unfamiliar adult male rats were introduced in a neutral enclosure and their social interactions were recorded. While all pairs escalated their playful encounters to become rougher, only the pairs in which one member was devocalized escalated to serious biting. A Monte Carlo shuffling technique was used for the analysis of the correlations between the overt playful and aggressive actions performed and the types and frequencies of various 50-kHz calls that were emitted. The analysis revealed that lower frequency (20-30kHz) calls with a flat component maybe particularly critical for de-escalating encounters and so allowing play to continue. Moreover, coordinating calls reciprocally, with either the same call mimicked in close, temporal association or with complementary calls emitted by participants as they engage in complementary actions (e.g., attacking the nape, being attacked on the nape), appeared to be ways with which calls could be potentially used to avoid escalation to aggression and so sustain playful interactions.

Specific 50-kHz vocalizations are tightly linked to particular types of behavior in juvenile rats anticipating play.

Rat ultrasonic vocalizations have been suggested to be either a byproduct of physical movement or, in the case of 50-kHz calls, a means to communicate positive affect. Yet there are up to 14 distinct types of 50-kHz calls, raising issues for both explanations. To discriminate between these theories and address the purpose for the numerous 50-kHz call types, we studied single juvenile rats that were waiting to play with a partner, a situation associated with a high number of 50-kHz calls. We used a Monte-Carlo shuffling procedure to identify vocalization-behavior correlations that were statistically different from chance. We found that certain call types ("split", "composite" and "multi-step") were strongly associated with running and jumping while other call types (those involving "trills") were more common during slower movements. Further, non-locomotor states such as resting and rearing were strongly predictive of a lack of vocalizations. We also found that the various sub-types of USVs can be clustered into 3-4 categories based on similarities in the way they are used. We did not find a one-to-one relationship between any movements and specific vocalizations, casting doubt on the motion byproduct theory. On the other hand, the use of specific calls during specific behaviors is problematic for the affect communication hypothesis. Based on our results, we suggest that ultrasonic calls may serve to coordinate moment-to-moment social interactions.