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.

Metabolically Healthy Obesity Is a Misnomer: Components of the Metabolic Syndrome Linearly Increase with BMI as a Function of Age and Gender.

OBJECTIVES: We aimed to examine the relationships between body mass index (BMI) and metabolic syndrome (MS) components as a function of age and gender across weight categories. METHODS: This cross-sectional study included 19,328 subjects who participated in a health-screening program. We analyzed 14,093 apparently healthy subjects with a BMI ≥ 18.5 kg/m2 (ranging from 18.5 to 46 kg/m2). RESULTS: At a BMI of 18.5 kg/m2, 16% of subjects had one or more MS components (MS ≥ 1). The number of MS components increased linearly with BMI. The most prevalent components for MS1-4 were hypertension (in men) and increased waist circumference (in women). Among 6391 non-obese subjects with MS = 0, there was a linear increase in blood pressure, glucose, and triglycerides, as well as a decline in high-density lipoprotein cholesterol, as BMI increased. In 2087 subjects with a BMI ≥ 30 kg/m2, a true normometabolic state (MS = 0) was observed in only 7.5%, declining to less than 1% at a BMI ≥ 36 kg/m2 (ATP criteria). Women were metabolically protected relative to men between the ages of 30 and 50 years. CONCLUSIONS: (A) MS components increase linearly with BMI from the lowest normal BMI and continue to increase with age and BMI; (B) metabolically healthy obesity is rare in subjects with a high BMI and declines with age; (C) hypertension is the most common component in men; and (D) in women, MS components are seen at older ages than in men for the same BMI. Metabolic health declines with age and BMI in nearly all subjects with obesity.

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.

Social spatial cognition: social distance dynamics as an identifier of social interactions.

We suggest that socio-spatial behavior, which is an interaction between social and spatial cognition, can be viewed as a set of excursions that originate and end in close proximity to another individual(s). We present an extension of earlier studies that perceived spatial behavior in individual animals as a series of excursions originating from a particular location. We measured here the momentary distance between two individuals (social distance) to differentiate among eight possible types of social excursion originating in a state of proximity between excursion-participants. The defined excursion types are based on whether or not the excursion initiator also concludes the excursion, whether or not the excursion starts and ends at the same location, and the dynamics of the distance between excursion participants. We validated this approach to socio-spatial behavior as a set of excursions using it to analyze the behavior of the two sexes in rodents, of normal vs. stereotyped rats, as well as of different rodent species. Each of these groups displays a prevalent excursion type that reflects a distinct social dynamics. Our approach offers a useful and comprehensive tool for studying socio-spatial cognition, and can also be applied to distinguish among different social situations in rodents and other 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.

Social interaction modulates the intensity of compulsive checking in a rat model of obsessive-compulsive disorder (OCD).

Here we present an empirical study that provides a basis for understanding the impact of the social environment on individuals with mental disorders. Rats treated chronically with the dopamine-agonist quinpirole offer a solid animal model for compulsive behavior that has been comprehensively evaluated and validated in numerous studies. Moreover, the method of behavioral analysis used in the quinpirole rat model has been similarly applied to the analysis of compulsive rituals in OCD patients, revealing similarities to the structure of compulsions in the quinpirole-sensitized rats. Here, we examined how compulsive checking by quinpirole-sensitized rats was modulated by the presence of a partner that was also treated with quinpirole or a partner that was treated with saline, compared to the typical expression of compulsive checking shown by rats tested alone. Our results demonstrate that the presence of a partner does indeed modulate the performance of checking behavior. Specifically, the vigor of compulsive checking was attenuated in the presence of a saline-treated partner, and augmented in the presence of a quinpirole-treated partner. This finding provides compelling evidence that social interactions modulate the expression of compulsive checking in the quinpirole rat model of OCD. This uncovering of the effectiveness of social modulation, indicates the quinpirole preparation as a paradigm for investigating the mechanisms by which the social environment modulates the development and expression of OCD. More generally, it presents a paradigm for the study of the influence of drug effects as a function of social interaction.

The hierarchy of food, sociality, and experience in spatial decision-making by food-deprived rats.

We examined whether a hierarchy existed among attraction to (i) food; (ii) social partner(s); and (iii) familiar feeding location in spatial decision-making in rats. To determine this, lone food-deprived rats were trained to collect baits from 16 salient equispaced objects arranged in a grid layout. Some rats were trained with only the eight objects on the left baited; other rats with only the eight objects on the right baited. After training, dyads of one left-trained and one right-trained rats were tested, with the eight baits now divided into four on the left and four on the right sides of the arena. Rats were free either to go to the familiar objects that had been baited in the lone training trials, regardless of whether these objects were now actually baited; or, alternatively, they could go to baited objects regardless of their side and of the rats' past experience. Rats could also forage individually or together regardless of their past experience or actual bait location. We found that the rats primarily displayed attraction to baits, regardless of their location; followed by a preference to travel together; and then by a significant combined attraction to visit together baited objects (whether familiar or not). Accordingly, attraction to food had the prime influence, sociality the second, and familiarity with the environment the least, in establishing spatial foraging decisions.

The sounds of silence: Barn owl noise in landing and taking off.

A barn owl swooping down generated a quieter, almost silent, noise (acoustic impulses) compared to a louder noise generated by the owl when taking off. These acoustic impulses are at low frequencies which are below the auditory threshold to most rodents. Therefore, rodents are less likely to hear these noises of owl flight. A previous study revealed that rodents exhibit frantic response to an owl taking off (as opposed to their typical freezing response during owl attack). The frantic response could be the result of tactile reception of the air-puffs generated by the owl's wingbeats and may reduce the success in subsequent attacks.

Spatio-temporal organization during group formation in rats.

In the present study, the dynamic process of group formation in eight unfamiliar rats was followed in order to reveal how the group becomes oriented together in time and space, in light of the complexity that accompanies grouping. The focus was on who, where, and when joined together. We found that rats preferred to be in companionship over remaining alone, with all the rats gradually shifting to share the same location as a resting place. Group formation can be viewed as a tri-phasic process, with some rats gradually becoming more social than others, and thus playing a key role in group formation. Starting with seemingly independent traveling, the rats gradually converged to share the same location as a terminal (home base) for roundtrips in the arena. Because such a terminal is considered as the organizer of an individual's spatial behavior, the shared home-base location may be viewed as the organizer of spatial behavior of the entire group. Despite huddling together, the rats continued to travel alone or in duos throughout the 3 h of testing. We suggest that resting together and traveling alone or in duos enabled the maintenance of communal relationship while reducing the complexity involved in traveling in relatively large groups. Taken together, the present results demonstrate the dynamic process during which unfamiliar rats shift from independent to group spatial behavior.

Reproducibility and replicability of rodent phenotyping in preclinical studies.

The scientific community is increasingly concerned with the proportion of published "discoveries" that are not replicated in subsequent studies. The field of rodent behavioral phenotyping was one of the first to raise this concern, and to relate it to other methodological issues: the complex interaction between genotype and environment; the definitions of behavioral constructs; and the use of laboratory mice and rats as model species for investigating human health and disease mechanisms. In January 2015, researchers from various disciplines gathered at Tel Aviv University to discuss these issues. The general consensus was that the issue is prevalent and of concern, and should be addressed at the statistical, methodological and policy levels, but is not so severe as to call into question the validity and the usefulness of model organisms as a whole. Well-organized community efforts, coupled with improved data and metadata sharing, have a key role in identifying specific problems and promoting effective solutions. Replicability is closely related to validity, may affect generalizability and translation of findings, and has important ethical implications.

Consistency of Spatial Representations in Rat Entorhinal Cortex Predicts Performance in a Reorientation Task.

Goal-directed behavior can be affected by environmental geometry. A classic example is the rectangular arena reorientation task, where subjects commonly confuse opposite but geometrically identical corners [1]. Until recently, little was known about how environmental geometry shapes spatial representations in a neurobehavioral context [2] (although see [3]). In the present study, we asked: Under what circumstances does the internal cognitive map predict behavior? And when does it fail to do so? To this end, we developed a variant of the classical reorientation task that allows for investigation of temporal dynamics of reorientation. We recorded head-direction (HD) cells and grid cells in the medial entorhinal cortex (MEC) of rats before, during, and after performing the task. MEC cells showed a bimodal response of being either aligned or rotated, relative to the free-foraging open-field sessions. Alignment was remarkably stable between disorientations and indicative of corner choice as a function of current and past alignment of spatial representations. Accordingly, when the cells showed consistent and properly aligned readout across multiple trials, behavioral choices were better predicted by HD and grid cell readout, with a probability of more than 70%. This was not the case when the cells did not show a stable consistent readout. Our findings indicate that entorhinal spatial representations predict corner choice, contingent on the stability and reliability of their readout. This work sets the stage for further studies on the link between the reliability of the neuronal signal and behavior, with implications for many brain systems in many organisms.

From an animal model to human patients: An example of a translational study on obsessive compulsive disorder (OCD).

The application of similar analyses enables a direct projection from translational research in animals to human studies. Following is an example of how the methodology of a specific animal model of obsessive-compulsive disorder (OCD) was applied to study human patients. Specifically, the quinpirole rat model for OCD was based on analyzing the trajectories of travel among different locales, and scoring the set of acts performed at each locale. Applying this analytic approach in human patients unveiled various aspects of OCD, such as the repetition and addition of acts, incompleteness, and the link between behavior and specific locations. It is also illustrated how the same analytical approach could be applicable to studying other mental disorders. Finally, it is suggested that the development of OCD could be explained by the four-phase sequence of Repetition, Addition, Condensation, and Elimination, as outlined in the study of ontogeny and phylogeny and applied to normal development of behavior. In OCD, this sequence is curtailed, resulting in the abundant repetition and addition of acts.

Rats do not eat alone in public: Food-deprived rats socialize rather than competing for baits.

Limited resources result in competition among social animals. Nevertheless, social animals also have innate preferences for cooperative behavior. In the present study, 12 dyads of food-deprived rats were tested in four successive trials, and then re-tested as eight triads of food-deprived rats that were unfamiliar to each other. We found that the food-deprived dyads or triads of rats did not compete for the food available to them at regular spatially-marked locations that they had previously learnt. Rather, these rats traveled together to collect the baits. One rat, or two rats in some triads, lead (ran ahead) to collect most of the baits, but "leaders" differed across trials so that, on average, each rat ultimately collected similar amounts of baits. Regardless of which rat collected the baits, the rats traveled together with no substantial difference among them in terms of their total activity. We suggest that rats, which are a social species that has been found to display reciprocity, have evolved to travel and forage together and to share limited resources. Consequently, they displayed a sort of 'peace economy' that on average resulted in equal access to the baits across trials. For social animals, this type of dynamics is more relaxed, tolerant, and effective in the management of conflicts. Rather than competing for the limited available food, the food-deprived rats socialized and coexisted peacefully.

Curiosity as an approach to ethoexperimental analysis: Behavioral neuroscience as seen by students and colleagues of Bob Blanchard.

This review is a synopsis of an International Behavioral Neuroscience Society (IBNS) symposium which focused on the elements of Behavioral Neuroscience for which Robert J. Blanchard was a Pioneer, Leading Expert, Advocate, Mentor, and Sage. Bob Blanchard's work demonstrably changed our broad understanding of animal behavior, and led the way to experimental design and analysis for studies of animal behavior that helped to clarify the deep complexity and subtleties of behavior. Bob's impact on the field of Behavioral Neuroscience includes the behavior, neurocircuitry, neurochemistry, and pharmacology related to social interactions, aggressive behavior, defensive behaviors, flight, freezing, threat, attack, risk assessment, anxiety disorders, animal models, models of social behavior, and autism. The methods and designs developed by Bob Blanchard over a lifetime have been adopted by scientists around the world, and form a standard of excellence in the field. The article addresses these topics in a way that presents developments in the field, describes the newest research data, and pays tribute to a great scientist and founder of this field of work, Bob Blanchard.