Tactile cues compensate for unbalanced vestibular cues during progression on inclined surfaces.

A previous study demonstrated that rodents on an inclined square platform traveled straight vertically or horizontally and avoided diagonal travel. Through behavior they aligned their head with the horizontal plane, acquiring similar bilateral vestibular cues - a basic requirement for spatial orientation and a salient feature of animals in motion. This behavior had previously been shown to be conspicuous in Tristram's jirds. Here, therefore jirds were challenged by testing their travel behavior on a circular arena inclined at 0°-75°. Our hypothesis was that if, as typical to rodents, the jirds would follow the curved arena wall, they would need to display a compensating mechanism to enable traveling in such a path shape, which involves a tilted frontal head axis and unbalanced bilateral vestibular cues. We found that with the increase in inclination, the jirds remained more in the lower section of the arena (geotaxis). When tested on the steep inclinations, however, their travel away from the arena wall was strictly straight up or down, in contrast to the curved paths that followed the circular arena wall. We suggest that traveling along a circular path while maintaining contact with the wall (thigmotaxis), provided tactile information that compensated for the unbalanced bilateral vestibular cues present when traveling along such curved inclined paths. In the latter case, the frontal plane of the head was in a diagonal posture in relation to gravity, a posture that was avoided when traveling away from the wall.

A small step for rats alters spatial behavior: rats on a bi-level arena explore each level separately.

We tested rats on a 'bi-level open-field' whose two halves were separated vertically by an 8-cm step that the rats could easily ascend/descend. We sought to determine what might be the factors that shape traveling in three-dimensional environments; what makes an environment perceived as multileveled; and how are multileveled environments explored compared to two-dimensional environments? We found that rats on the bi-level open-field traveled a greater distance on the lower level compared to the upper one. They also spent a long time at the foot of the step before ascending to the upper level. They established a home-base on one level and a local base on the other one, and explored each level separately. We could not find a particular factor that accounted for the preference for the lower level. We suggest that the momentary egocentric sensation of moving vertically, together with an overall area large enough for exploration, result in perceiving an environment as multilevel. Exploration of such environments is fragmented, and each level is explored relatively independently, as has also been shown in other studies. Regarding the unanswered question of earlier studies concerning what integrates fragmented representations, this is the first study that suggests that in rats, and perhaps also in other rodent species, such integration is achieved by means of home-base behavior, resulting in the establishment of a single comprehensive representation of the multilevel environment.

Can rats and ants exchange information between the horizontal and vertical domains?

Since traveling in nature involves encountering various vertical structures, integration of horizontal and vertical spatial information is required. One form of such integration is to use information acquired in one plane for spatial navigation in another plane. Here we tested whether rats and ants that learned a reward location in a horizontal maze could utilize this information when the maze was rotated to a vertical orientation and vice versa. Rats that were trained in a horizontal Y-maze required more time to reach the reward when the maze was vertically rotated, but they were more accurate in choosing the correct arm. In contrast, rats tested in a horizontal maze after being trained in a vertical maze were less accurate but reached the reward faster. Changes after maze rotation were moderate and non-significant in ants, perhaps since the number of ants arriving at the reward increased over trials, diminishing the effect of maze rotation in ants compared to rats. According to the notion that horizontal spatial information is encoded in more detail than vertical information, the slow performance of rats in the vertical domain could be due to a more physically demanding task whereas their accuracy was due to a preceding detailed horizontal encoding. In contrast, rats in the vertical maze could gather less detailed information and therefore were less accurate in subsequent horizontal trials, where the lower energy cost enabled them to swiftly correct wrong choices. Altogether, the present results provide an indication for transferring spatial information between horizontal and vertical dimensions.

Rodents Prefer Going Downhill All the Way (Gravitaxis) Instead of Taking an Uphill Task.

We directly tested whether, when given the choice to ascend or descend, rodents would favor traveling downwards or upwards. The test incorporated different rodent species that dwell in different habitats and display different life and motor styles. Testing was performed in a three-dimensional Y-maze in which the basis was horizontal and, by rotating it, one arm of the maze could be pointing upwards at a certain angle and the other arm pointed downwards at the same angle. All the tested species displayed a general preference for descent, with rodents from complex habitats being less affected by inclination compared with rodents from flatlands. Unlike laboratory rats, wild species traveled greater distances along the lower compared to the upper maze arm. All the rodents initially tended to travel the entire length of the inclined maze arms, but such complete trips decreased with the increase in inclination. When introduced into the maze from top or bottom, flatland dwellers traveled mainly in the entry arm. Overall, when given the choice to ascend or descend, all the tested species displayed a preference to descend, perhaps as attraction to the ground, where they usually have their burrows.

Keep a level head to know the way ahead: How rodents travel on inclined surfaces?

Animals traveling on a horizontal surface stabilize their head in relation to the substrate in order to gather spatial information and orient. What, however, do they do when traveling on an incline? We examined how three rodent species differing in motor abilities and habitats explore a platform tilted at 0-90°, hypothesizing that they would attempt to maintain bilateral vestibular cues. We found that traveling up or down was mainly straight vertically rather than diagonally, which results in identical bilateral vestibular cues. This was also achieved when traveling horizontally through rotating the head to parallel the horizontal plane. Traveling diagonally up or down was avoided, perhaps due to different bilateral vestibular cues that could hinder orientation. Accordingly, we suggest that maintaining identical bilateral cues is an orientational necessity that overrides differences in motor abilities and habitats, and that this necessity is a general characteristic of animals in motion.

On heights and plains: How rodents from different habitats cope with three-dimensional environments?

Dwelling in a specific habitat requires adaptation to the habitat physical and biological properties in order to maximize fitness. Adaptations that are manifested in the organization of behavior in time and space reflect how the environment is perceived and utilized. Testing species from different habitats in the same laboratory environment can uncover the differences in their behavior and their adaptations to specific habitats. The question posed in this study is that of how two rodent species, one occupying flatlands (Tristram's jird; Meriones tristrami) and the other occupying structured rocky habitats (common spiny mouse; Acomys dimidiatus), differ in the way that they explore the same three-dimensional laboratory environment. Individuals of these two species were introduced into an arena with a five-level ziggurat in the center, and their behavior was followed for 60 min. We found that both species preserved the typical spatiotemporal rodents' behavior of establishing a home-base-a location that is a terminal from which they set out to explore the environment. However, the jirds, which live in flatlands, mainly travelled on the arena floor and the lower levels of the ziggurat; while, in contrast, the spiny mice, which live in rocky habitats and are used to climbing, mostly remained and travelled on the ziggurat, with some of them hardly descending to the arena floor. We suggest that the distinction in spatial behavior between the two species reflects their different motor abilities, different depth perception, and different umvelt (perceived world), in accordance with their different natural habitats.

Social spatial cognition.

Social spatial cognition refers to the interaction between self, place, and partners, with emphasis on the impact of the social environment on spatial behavior and on how individual spatial representations converge to form collective spatial behavior - i.e., common places and routes. Recent studies suggest that in addition to their mental representation (cognitive map) of the physical environment, humans and other animals also have a social cognitive map. We suggest that while social spatial cognition relies on knowledge of both the physical and the social environments, it is the latter hat predominates. This dominance is illustrated here in the modulation of spatial behavior according to dynamic social interactions, ranging from group formation to an attenuation of drug-induced stereotypy through the mere presence of a normal subject. Consequently we suggest that the numerous studies on the biobehavioral controlling mechanisms of spatial behavior (i.e. - the hippocampal formation, animal models for mental disorders) should also consider the social environment rather than solely focusing on the spatial behavior of lone animals.

How do rodents explore a three-dimensional environment? Habitat-dependent and direction-dependent differences.

Many animals are surface-bounded, traveling mostly in two-dimensional (2D) environments. However, those that inhabit structured habitats might also require wayfinding in three-dimensional (3D) environments. Here we forced rodents to ascend or descend when traveling. We tested three species: laboratory rats (a common experimental subject); fat sand rats, which forage while climbing shrubs (representing those used to 3D travel); and Tristram's jirds, which forage in plains (not used to climbing). We examined differences between individuals initially placed on top of the apparatus compared with those placed on its bottom, assuming that this, in addition to the above difference in habitats and motor habits, would influence their spatial behavior. Exploratory activity of top-starting rats and sand rats, but not jirds, differed from bottom starters. Nevertheless, despite the need to continuously ascend or descend, both top- and bottom-starters of the three species displayed the spatio-temporal structure of open-field exploration as previously revealed in a horizontal arena. Specifically, exploration constituted a set of round-trips to a home-base. It is suggested that the preservation of a regular structure of spatial behavior was due to the ability of the tested rodents to mostly maintain a horizontal posture of their head when ascending and descending.

"It's all in their head": hierarchical exploration of a three-dimensional layered pyramid in rats.

Wayfinding in a three-dimensional (3D) environment is intricate, and surface-bounded animals may overcome this complexity by breaking it down into horizontal layers along with the vertical location of each layer. Here, we examined how rats explored a layered pyramid placed in a large open field. We found that exploration presented a hierarchical (or fractal) shape of three types of roundtrips: (1) from the primary home-base to the open-field floor; (2) from the floor up and down the pyramid levels; and (3) from local home-base on each pyramid level. Ascent was slow and interrupted, whereas descent was fast. This difference was a result of level altitude, remaining after data were normalized proportionally to level area. In contrast, the time spent and the distance traveled on each level were dependent on level area, not on level altitude. This structure of spatial behavior accords with multilevel exploration, presenting a relatively independent exploration of each level. The vertical dimension in this experiment thus did not alter the typical spatiotemporal behavior, and the 3D environment was explored by application of the same spatiotemporal approach as that of a horizontal open field. We suggest that this lack of alteration is due to the horizontal posture of the animal's head and trunk during progression on the pyramid. This behavior also seems to fit the bicoding hypothesis, in which the vertical information is virtually contextual (non-metric), and so, when the rat progresses to a new level, it explores it as a newly accessed horizontal floor area.