Information Theory Opens New Dimensions in Experimental Studies of Animal Behaviour and Communication.

Over the last 40-50 years, ethology has become increasingly quantitative and computational. However, when analysing animal behavioural sequences, researchers often need help finding an adequate model to assess certain characteristics of these sequences while using a relatively small number of parameters. In this review, I demonstrate that the information theory approaches based on Shannon entropy and Kolmogorov complexity can furnish effective tools to analyse and compare animal natural behaviours. In addition to a comparative analysis of stereotypic behavioural sequences, information theory can provide ideas for particular experiments on sophisticated animal communications. In particular, it has made it possible to discover the existence of a developed symbolic "language" in leader-scouting ant species based on the ability of these ants to transfer abstract information about remote events.

Flexibility and rigidity in hunting behaviour in rodents: is there room for cognition?

Predatory hunting is a complex species-typical behaviour involving different skills, some of which may include learning. This research aims to distinguish between rigid and flexible parts in live-insect hunting behaviour in nine herbivorous and granivorous rodent species, and to find out whether there is room for cognition in this activity. In laboratory experiments, all species studied manifest skilful attacks towards insects in a manner that is typical for specialised predators chasing a fleeing prey. Voles demonstrate a "core" and somewhat primitive scheme of a hunting pattern: approaching a potential victim, biting it, and then seizing and handling. Hamsters display the tendency to start their attacks by actions with paws, but they can achieve success only using teeth as well. Gerbils can successfully use both paws and teeth to start the attack, which brings their hunting behaviour closer to that of specialised rodent predators. We revealed variability in the display of hunting in different species, methods of seizing the prey, and the number of attempts to attack an insect before catching it. We found specific flexible fragments within the "bite-grasp-handle" bouts that can be precursors for adaptive phenotypic variations and include some cognitive attributes. We hypothesise that the divergence and specialisation of predatory behaviour in rodents can be based on the natural fragmentation of the original hunting patterns, that is, on the loss or recombination of particular behavioural elements. We consider a possible link between the fragmentation of hunting behaviour and social learning in different classes of animals and conjecture an intriguing correlation between predatory activity, cognitive skills and personal traits in rodents.

Spatial cognition in the context of foraging styles and information transfer in ants.

Ants are central-place foragers: they always return to the nest, and this requires the ability to remember relationships between features of the environment, or an individual's path through the landscape. The distribution of these cognitive responsibilities within a colony depends on a species' foraging style. Solitary foraging as well as leader-scouting, which is based on information transmission about a distant targets from scouts to foragers, can be considered the most challenging tasks in the context of ants' spatial cognition. Solitary foraging is found in species of almost all subfamilies of ants, whereas leader-scouting has been discovered as yet only in the Formica rufa group of species (red wood ants). Solitary foraging and leader-scouting ant species, although enormously different in their levels of sociality and ecological specificities, have many common traits of individual cognitive navigation, such as the primary use of visual navigation, excellent visual landmark memories, and the subordinate role of odour orientation. In leader-scouting species, spatial cognition and the ability to transfer information about a distant target dramatically differ among scouts and foragers, suggesting individual cognitive specialization. I suggest that the leader-scouting style of recruitment is closely connected with the ecological niche of a defined group of species, in particular, their searching patterns within the tree crown. There is much work to be done to understand what cognitive mechanisms underpin route planning and communication about locations in ants.

Red Wood Ants Display Natural Aversive Learning Differently Depending on Their Task Specialization.

The adaptive benefits of individual specialization and how learning abilities correlate with task performance are still far from being well-understood. Red wood ants are characterized by their huge colonies and deep professional specialization. We hypothesized that red wood ants Formica aquilonia form aversive learning after having negative encounters with hoverfly larvae differently, depending on their task specialization. We tested this hypothesis, first, by examining whether hunters and aphid milkers learn differently to avoid the nuisance of contacts with syrphid larvae, and, second, by analyzing the difference between learning in "field" and laboratory-reared (naïve) foragers. During the first interaction with the syrphid larva in their lives the naïve foragers showed a significantly higher level of aggressiveness than the members of a natural colony. Naïve foragers applied the "mortal grip," "prolonged bites," and "nibbling" toward the enemy with a significantly higher frequency, whereas members of both "field" groups behaved more carefully and tried to avoid encounters with the larva. The aphid milkers, who had a negative experience of interaction with the larva, being "glued" with its viscous secretion, behaved much less aggressively in the follow-up experiments after 10 min and even 3 days, thus exhibiting the shaping of both short- and long-term memories. However, both "field" hunters and naïve foragers demonstrated no signs of aversive learning. These data provide some new insights into the relationship between task specialization and learning performance in ants. Given our previous results, we speculate that scouts and aphid milkers are the most cognitively gifted specialists in red wood ants, whereas hunters and guards are rather brave than smart.

Precise relative-quantity judgement in the striped field mouse Apodemus agrarius Pallas.

Applying the classical experimental scheme of training animals with food rewards to discriminate between quantities of visual stimuli, we demonstrated that not only can striped field mice Apodemus agrarius discriminate between clearly distinctive quantities such as 5 and 10, but some of these mice also exhibit high accuracy in discriminating between quantities that differ only by one. The latter include both small (such as 2 versus 3) and relatively large (such as 5 versus 6, and 8 versus 9) quantities of elements. This is the first evidence of precise relative-quantity judgement in wild rodents. We found striking individual variation in cognitive performance among striped field mice, which possibly reflects individual cognitive variation in natural populations. We speculate that high accuracy in differentiating large quantities is based on the adaptive ability of wild rodents to capture subtle changes in their environment. We suggest that the striped field mouse may be a powerful model species to develop advanced cognitive tests for comparative studies of numerical competence in animals and for understanding evolutionary roots of quantity processing.

Using the Data-Compression Method for Studying Hunting Behavior in Small Mammals.

Using the data-compression method we revealed a similarity between hunting behaviors of the common shrew, which is insectivorous, and several rodent species with different types of diet. Seven rodent species studied displayed succinct, highly predictable hunting stereotypes, in which it was easy for the data compressor to find regularities. The generalist Norway rat, with its changeable manipulation of prey and less predictable transitions between stereotype elements, significantly differs from other species. The levels of complexities of hunting stereotypes in young and adult rats are similar, and both groups had no prior experience with the prey, so one can assume that it is not learning, but rather the specificity of the organization of the stereotype that is responsible for the nature of the hunting behavior in rats. We speculate that rodents possess different types of hunting behaviors, one of which is based on a succinct insectivorous standard, and another type, perhaps characteristic of generalists, which is less ordered and is characterized by poorly predictable transitions between elements. We suggest that the data-compression method may well be more broadly applicable to behavioral analysis.

Myrmica rubra ants are more communicative when young: Do they need experience?

The role of experience in the development of communication in animals is a matter of special interest to many ethologists and psychologists. Ants are known to possess sophisticated and flexible communication systems based mainly on their antennal movements (Reznikova & Ryabko, 2011). However, it is still enigmatic whether young ants need stimulation performances by adults to develop their communication capacities. Experiments with pairwise interactions of Myrmica rubra ants revealed significant differences in individual behavior and the mode of communication in callow (newly emerged) and adult workers. Adult ants are much more mobile than callow ones, and they switch their behavior depending on what partner they interact with, whereas callows behave independently. Adults communicate with callows and queens much longer than with other adults. Both callows and queens seem to be rather attractive to adults, although in different ways. Adults pay close attention to callow ants and initiate prolonged antennal contacts with them, touching their bodies and not leaving them alone. Young (callow) ants appear to be more communicative than adults, and they are equally ready to communicate with each other and with adults. Antennal movements are slow and clumsy in young ants, and they often switch from communication to other activities. It is likely that patterns of antennal movements in callows change gradually. Peculiarities of the mode of communication enable us to speculate that young ants need prolonged contacts with adult nestmates to gain the experience of communication. Some parallels with the development of communication skills in vertebrate species are considered. (PsycINFO Database Record

On the evolutionary origins of differences in sexual preferences.

A novel explanation of the evolutionary process leading to the appearance of differences in sexual preferences is proposed. The explanation is fully general: it is not specific to any particular type of sexual preferences, nor to any species or population. It shows how different sexual preferences can appear in any large group-living population in which sexual selection is sufficiently strong in each sex. The main idea is that the lack of interest toward a member of the opposite sex may be interpreted as a signal of popularity, and thus of reproductive success. It is then boosted by the Fisher runaway process far beyond the point where it becomes costly, resulting in a generalized trait-lack of interest toward the opposite sex. If the interest diverts toward other targets then different sexual preferences emerge. This hypothesis is placed into the context of other works on different sexual preferences in animals; supporting evidence from the literature is reviewed and additional research needed to confirm or refute the hypothesis in any given species is outlined.

Quantity judgments in the context of risk/reward decision making in striped field mice: first "count," then hunt.

We simulated the situation of risky hunting in the striped field mouse Apodemus agrarius in order to examine whether these animals are able to make a choice between small and large quantities of live prey (ants). In the first (preliminary) experiment we investigated to what extent mice were interested in ants as a live prey and how their hunting activity depended on the quantity of these edible but rather aggressive insects. We placed mice one by one into arenas together with ant groups of different quantities, from 10 to 60. Surprisingly, animals, both wild-caught and laboratory-reared, displayed rather skilled predatory attacks: mice killed and ate from 0.37 ± 003 to 4 ± 0.5 ants per minute. However, there was a threshold number of ants in the arenas when rodents expressed signs of discomfort and started to panic, likely because ants bit them. This threshold corresponds to the dynamic density (about 400 individuals per m(2) per min) in the vicinity of anthills and ants' routes in natural environment. In the second experiment mice had to choose between different quantities of ants placed in two transparent tunnels. Ants here served both as food items and as a source of danger. As far as we know, this is the first experimental paradigm based on evaluation of quantity judgments in the context of risk/reward decision making where the animals face a trade-off between the hedonistic value of the prey and the danger it presents. We found that when mice have to choose between 5 vs. 15, 5 vs. 30, and 10 vs. 30 ants, they always tend to prefer the smaller quantity, thus displaying the capacity for distinguishing more from less in order to ensure comfortable hunting. The results of this study are ecologically relevant as they reflect situations and challenges faced by free-living small rodents.

Asymmetry in antennal contacts during trophallaxis in ants.

Behavioural and brain left-right asymmetries are a common feature among the animal kingdom. Lateralization often manifests itself at the population-level with most individuals showing the same direction of lateral bias. Theoretical model based on evolutionary stable strategy predicts that lateralization at the population-level is more likely to characterize social rather than solitary species. Empirical data supporting this hypothesis has been recently obtained in Hymenoptera showing that eusocial honeybees present an asymmetrical use of the antennae: the right antenna is involved in olfactory learning and present more olfactory receptors. However, no evidences about the role of antennal asymmetries in social interactions have been provided so far. Highly social ant species belonging to Formica rufa group are a good model for investigating natural communication because they are able to pass exact information to their nest mates. We applied the "binary tree" experimental paradigm, which allowed us to observe different types of antennal contacts performed by ants out of their nest. To examine possible asymmetrical use of the right and left antenna, we focused on "feeding" (the simplest) contacts where a "donor" ant is exchanging food with a "receiver" ant through trophallaxis. We observed a population-level asymmetry, with the "receiver" ant using the right antenna significantly more often than the left antenna. This study provides the first evidence of lateralization in antennal contacts in ants, and seems to support the hypothesis of mathematical models on the evolution of lateralization suggesting that the alignment of lateralization at the population-level matters in social interactions.