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.

Operant conditioning in antlion larvae and its impairment following exposure to elevated temperatures.

Although ambush predators were previously considered limited in their cognitive abilities compared to their widely foraging relatives, there is accumulating evidence it does not hold true. Pit-building antlions are already known to associate vibrations in the sand with the arrival of prey. We used a T-maze and successfully trained antlions to turn right or left against their initial turning bias, leading to a suitable substrate for digging traps. We present here the first evidence for operant conditioning and T-maze solving in antlions. Furthermore, we show that exposure of second instar larvae to an elevated temperature led to impaired retention of what was learned in a T-maze when tested after moulting into the third instar, compared to larvae raised under a more benign temperature. We suggest that climate change, involving an increase in mean temperatures as well as rare events (e.g., heatwaves) might negatively affect the retention of operant conditioning in antlions, alongside known, more frequently studied effects, such as changes in body size and distribution.

Negative effects of fluctuating temperatures around the optimal temperature on reproduction and survival of the red flour beetle.

Whereas the vast majority of animals in nature experience daily or seasonal thermal fluctuations, most laboratory experiments use constant temperatures. We examined the effect of fluctuating temperatures on reproduction and survival under starvation, two important components of fitness. We used the red flour beetle as a model organism, which is a significant pest in grain mills around the world. Fluctuations around the optimal temperature were always negative for the adult survival under starvation. The effect of thermal fluctuations on the number of offspring reaching adulthood was negative as well but increased with the extent of exposure. It was the strongest when the adult parents were kept and the offspring were raised under fluctuating temperatures. However, the later the offspring were exposed to fluctuations during their development, the weaker the effect of fluctuating temperatures was. Moreover, raising the parents under fluctuating temperatures but keeping them after pupation at constant temperatures fully alleviated the negative effects of fluctuations on the offspring. Finally, we demonstrate that keeping the parents a few days under fluctuating temperatures is required to induce negative effects on the number of offspring reaching adulthood. Our study disentangles between the effects of thermal fluctuations experienced during the parental and offspring stage thus contributing to the ongoing research of insects under fluctuating temperatures.

Social isolation causes downregulation of immune and stress response genes and behavioural changes in a social insect.

Humans and other social mammals experience isolation from their group as stressful, triggering behavioural and physiological anomalies that reduce fitness. While social isolation has been intensely studied in social mammals, it is less clear how social insects, which evolved sociality independently, respond to isolation. Here we examined whether the typical mammalian responses to social isolation, e.g., an impaired ability to interact socially and immune suppression are also found in social insects. We studied the consequences of social isolation on behaviour and brain gene expression in the ant Temnothorax nylanderi. Following isolation, workers interacted moderately less with adult nestmates, increased the duration of brood contact, and reduced the time spent self-grooming, an important sanitary behaviour. Our brain transcriptome analysis revealed that only a few behaviour-related genes had altered their expression with isolation time. Rather, many genes linked to immune system functioning and stress response had been downregulated. This probably sensitizes isolated individuals to various stressors, in particular because isolated workers exhibit reduced sanitary behaviour. We provide evidence of the diverse consequences of social isolation in social insects, some of which resemble those found in social mammals, suggesting a general link between social well-being, stress tolerance, and immune competence in social animals.

Factors That Can Affect the Spatial Positioning of Large and Small Individuals in Clusters of Sit-and-Wait Predators.

Shadow competition, the interception of prey by sit-and-wait predators closest to the source of prey arrival, is prevalent in clusters of sit-and-wait predators. Peripheral positions in the cluster receive more prey and should thus be more frequently occupied. Models predicting spatial positioning in groups, however, usually ignore variability among group members. Here, I used a simulation model to determine conditions under which small and large sit-and-wait predators, which differ in their attack range, should differ in their spatial positions in the cluster. Small predators occupied peripheral positions more frequently than large predators at the simulation beginning, while the opposite held true as time advanced. Because of the large and small attack range of large and small predators, respectively, small predators mistakenly relocated away from peripheral positions, while large predators did not relocate fast enough from inferior central positions. Any factor that moderated the frequent relocations of small predators or had the opposite effect on large predators assisted small or large predators, respectively, in reaching the more profitable peripheral positions. Furthermore, any factor elevating shadow competition led to longer occupation of the periphery by large predators. This model may explain why sit-and-wait predators are not homogenously distributed in space according to size.

Fine sand particles enable antlions to build pitfall traps with advanced three-dimensional geometry.

Pit-building antlion larvae are predators that construct pitfall traps in fine sand. We used three-dimensional laser scanning and geometric morphometrics to reveal the shape of antlion pits of two antlion species, analysed the particle size composition of sands from the different natural habitats, and measured the slope angles of the pits of the two species. In most antlions, the pits are structured as a simple inverted cone, as in Myrmeleon hyalinus, studied here. The other antlion studied, Cueta lineosa, constructs a unique pit composed of two inverted truncated cones inserted into one another, which feature substantially steeper walls than the pits of any other antlion studied to date. Pit stability depends on the slope inclination, which oscillates between the maximum angle of stability and the angle of repose. The angles in C. linosa substrates were larger than those in M. hyalinus substrates. One reason for the steeper walls is the greater proportion of fine sand in the natural sand inhabited by C. lineosa However, video-recording revealed that both the natural sand of C. lineosa and the finest sand tested had a higher maximum angle of stability than any of the other substrates studied here. Furthermore, experiments with pits built in different substrates revealed that the shape of the pit is variable and depends on the structure of the sand. Myrmeleonhyalinus displayed a more flexible pit construction behaviour than C. lineosa The present demonstration of such differences in pit characteristics contributes to understanding how these two species co-exist in the same habitat.

Thermal acclimation is not induced by habitat-of-origin, maintenance temperature, or acute exposure to low or high temperatures in a pit-building wormlion (Vermileo sp.).

Wormlions are sit-and-wait insect predators that construct pit-traps to capture arthropod prey. They require loose soil and shelter from direct sun, both common in Mediterranean cities, and explaining their high abundance in urban habitats. We studied different aspects of thermal acclimation in wormlions. We compared chill-coma recovery time (CCRT) and heat-shock recovery time (HSRT) of wormlions from urban, semi-urban and natural habitats, expecting those originating from the urban habitat to be more heat tolerant and less cold tolerant. However, no differences were detected among the three habitats. We then examined whether maintenance temperature affects CCRT and HSRT, and expected beneficial acclimation. However, CCRT was unaffected by maintenance temperature, while temperature affected HSRT in an opposite direction to our prediction: wormlions maintained under the higher temperatures took longer to recover. When testing with two successive thermal shocks, wormlions took longer to recover from both cold and heat shock after applying an initial cold shock. We therefore conclude that cold shock inflicts some damage rather than induces acclimation. Finally, both cold- and heat-shocked wormlions constructed smaller pits than wormlions of a control group. Smaller pits probably translate to a lower likelihood of capturing prey and also limit the size of the prey, indicating a concrete cost of thermal shock. In summary, we found no evidence for thermal acclimation related either to the habitat-of-origin or to maintenance temperatures, but, rather, negative effects of unfavorable temperatures.

The effect of previous experience on trap construction and movement distance in a pit-building predator.

Wormlion larvae are sit-and-wait predators that construct cone-shaped pits in sandy patches to capture prey. Wormlions select microhabitats that feature favorable conditions for pit construction, in a similar way to other trap-building predators, like spiders and antlions. We investigated whether wormlions exhibit an experience-based behavioral plasticity in their pit construction behavior. In a laboratory experiment, pit sizes and relocation distances were compared between larvae that experienced either a period of unfavorable conditions, i.e., surface obstacles, shallow or coarse sand, or a period of favorable conditions, i.e., clear, deep, and fine sand and were able to construct pits undisturbed. We expected that wormlions experiencing improving conditions would build larger pits than those experiencing deteriorating conditions. In addition, we expected that larvae experiencing unfavorable conditions would be less choosy in their new microhabitat and move over shorter distances. We observed a certain effect of recent experience on the trap-building behavior; however, it was not consistent among treatments. Additionally, we detected a correlation between larval body mass, relocation distance, and pit area. These findings might suggest that past experience does not influence wormlion foraging behavior in a simple manner but that different types of experience induce different behavioral responses.

The negative effect of starvation and the positive effect of mild thermal stress on thermal tolerance of the red flour beetle, Tribolium castaneum.

The thermal tolerance of a terrestrial insect species can vary as a result of differences in population origin, developmental stage, age, and sex, as well as via phenotypic plasticity induced in response to changes in the abiotic environment. Here, we studied the effects of both starvation and mild cold and heat shocks on the thermal tolerance of the red flour beetle, Tribolium castaneum. Starvation led to impaired cold tolerance, measured as chill coma recovery time, and this effect, which was stronger in males than females, persisted for longer than 2 days but less than 7 days. Heat tolerance, measured as heat knockdown time, was not affected by starvation. Our results highlight the difficulty faced by insects when encountering multiple stressors simultaneously and indicate physiological trade-offs. Both mild cold and heat shocks led to improved heat tolerance in both sexes. It could be that both mild shocks lead to the expression of heat shock proteins, enhancing heat tolerance in the short run. Cold tolerance was not affected by previous mild cold shock, suggesting that such a cold shock, as a single event, causes little stress and hence elicits only weak physiological reaction. However, previous mild heat stress led to improved cold tolerance but only in males. Our results point to both hardening and cross-tolerance between cold and heat shocks.

Foraging syndromes and trait variation in antlions along a climatic gradient.

Behavioral syndromes arise when individual behavior is correlated over time and/or across environmental contexts, often resulting in inter-population behavioral differences. Three main hypotheses have been suggested to explain the evolution of behavioral syndromes. The constraint hypothesis suggests that behaviors originate from a shared mechanism with a strong genetic or physiological basis. In contrast, according to the adaptive hypothesis, behavioral syndromes depend on specific selective pressures in each environment, and thus should evolve when specific behavioral combinations are advantageous. Finally, behavioral syndromes can also arise owing to neutral stochastic processes. We tested here for variation in the foraging syndromes of pit-building antlions originating from different populations along a climatic gradient. Although inter-population variation existed in some traits, foraging syndromes were similar across populations, supporting the constraint hypothesis. These findings suggest that stabilizing selection, acting on the foraging behavior of antlions during their larval phase, outweighs local selection pressures, resulting in "constraint syndromes." We also explored behavioral repeatability of foraging-related traits within and among habitats (natural, novel and disturbed habitats), and detected different levels of repeatability: pit diameter was more repeatable than response time to prey, followed by prey exploitation efficiency. Behavioral repeatability of the same trait differed according to context, suggesting that repeatability is a trait in itself and should not be considered identical even when studying the same behavioral trait.

Search patterns, resource regeneration, and ambush locations impact the competition between active and ambush predators.

Many predators ambush prey rather than pursue them or shift between foraging modes. Active predators typically encounter prey more frequently than ambush predators. I designed a simulation model to examine whether this always holds and how active and ambush predators fare in capturing mobile prey. Prey foraged for clumped resources using area-restricted search, shifting from directional movement before resource encounter to less directional movement afterward. While active predators succeeded more than ambush predators, the advantage of active predators diminished when ambush predators were positioned inside resource patches rather than outside. I investigated the impact of eight treatments and their interactions. For example, regeneration of prey resources increased the difference between ambush predators inside and outside patches, and uncertain prey capture by predators decreased this difference. Several interactions resulted in outcomes different from each factor in isolation. For instance, reducing the directionality level of active predators impacted moderately when applied alone, but when combined with resource regeneration it led to the worst success of active predators against ambush predators inside patches. Ambush predators may not always be inferior to active predators, and one should consider the key traits of the studied system to predict the relative success of these two foraging modes.

Experimental arena settings might lead to misinterpretation of movement properties.

Movement is an important animal behavior contributing to reproduction and survival. Animal movement is often examined in arenas or enclosures under laboratory conditions. We used the red flour beetle (Tribolium castaneum) to examine here the effect of the arena size, shape, number of barriers, access to the arena's center, and illumination on six movement properties. We demonstrate great differences among arenas. For example, the beetles moved over longer distances in clear arenas than in obstructed ones. Movement along the arena's perimeter was greater in smaller arenas than in larger ones. Movement was more directional in round arenas than in rectangular ones. In general, the beetles stopped moving closer to the perimeter and closer to corners (in the square and rectangular arenas) than expected by chance. In some cases, the arena properties interacted with the beetle sex to affect several movement properties. All these suggest that arena properties might also interact with experimental manipulations to affect the outcome of studies and lead to results specific to the arena used. In other words, instead of examining animal movement, we in fact examine the animal interaction with the arena structure. Caution is therefore advised in interpreting the results of studies on movement in arenas under laboratory conditions and we recommend paying attention also to barriers or obstacles in field experiments. For instance, movement along the arena's perimeter is often interpreted as centrophobism or thigmotaxis but the results here show that such movement is arena dependent.

Flour beetles prefer corners over walls and are slowed down with increasing habitat complexity.

Movement affects all key behaviours in which animals engage, including dispersal and habitat use. The red flour beetle, known as a cosmopolitan pest of stored products, was the subject of our study. We examined whether the beetles preferred corners, walls or open areas, and how turns or obstacles in corridors delayed the beetles' arrival at a target cell. Beetles spent significantly more time in corners than expected by chance, while they spent considerably less time in open areas than expected. However, no significant difference was observed between areas with two or three surrounding walls. This could be attributed to the beetles' stronger attraction to corners than crevices or the insufficient proximity of the third wall to the other two. Movement through the corridor was delayed by turns or obstacles, expressed in arrival probabilities, arrival times, time in the corridor or movement speed. Obstacles on the corridor's perimeter had a stronger effect on the beetle movement than those in the corridor's centre owing to the beetles' tendency to follow walls. The research is important also for applied purposes, such as better understanding beetle movement, how to delay their arrival to new patches, and where to place traps.

Metabolomics analysis of larval secretions reveals a caste-driven nutritional shift in a social wasp colony.

Social wasps exhibit a unique nutritional cycle in which adults feed larvae with prey, and larvae provide adults with larval secretions (LS). LS serves as a vital nutritional source for adults, contributing to the colony's health and reproductive success. The LS nutrient composition has been previously reported in various wasp species, yet these analyses focused solely on worker-destined larvae, overlooking the potential caste designation effects on LS composition. Using metabolomics techniques, we analysed and compared the metabolite and nutrient composition in LS of queen- and worker-destined larvae of the Oriental hornet. We found that queen-destined LS (QLS) contain greater amounts of most metabolites, including amino acids, and smaller amounts of sugars compared to worker-destined LS (WLS). The amino acid-to-sugar ratio in QLS was approximately tenfold higher than in WLS. Thus, as the colony transitions from the production of workers to the production of reproductives, it gradually experiences a nutritional shift that may influence the behaviour and physiology of the adult nest population. This caste-specific metabolite profile and nutrient composition of LS reflect the differences in the diet and physiological requirements of worker- and queen-destined larvae and may play a critical role in caste determination in social wasps.

Behavioral differences between pit-building antlions and wormlions suggest limits to convergent evolution.

Antlions and wormlions are distantly related insect taxa, both digging pits in loose soil and ambushing arthropod prey. Their hunting method, which is rare in the animal kingdom, is a clear example of convergent evolution. There is little research directly comparing the 2 pit-building taxa. Using the same experimental platform to investigate how they respond to biotic and abiotic environmental factors enables an examination of their convergence and its limits. We investigated the response of antlions and wormlions to 3 factors common in their daily life: disturbance to the pits, prey arrival, and conspecific competitors. Although both increased the pit size following disturbance, wormlions increased it faster than antlions. Antlions responded to prey faster than wormlions, but wormlions improved their response time over days. The most diverging response was toward conspecifics. Whereas antlions relocated their pits fast in response to increasing conspecific density, wormlions never relocated. We suggest explanations for the behavioral differences between the taxa. Our results imply that despite the similar hunting method of the 2 taxa they may differ greatly in their behavior, which in turn might have consequences for their habitat use and population dynamics.

Snakes on a slope: strong anti-gravitactic responses and differential habitat use in the Saharan horned viper (Cerastes cerastes) in the Negev desert.

The way species use their habitat dictates their intra- and interspecific interactions. We studied the effects of the microhabitat type and slope on the movement behaviour of the Saharan horned viper (Cerastes cerastes) in its natural habitat. This viper occurs in sand dunes and moves mostly by sidewinding. Additionally, we studied the microhabitat preference of desert rodents-the vipers' main prey. We placed the vipers on different natural dune slopes and recorded their behaviour. We found a strong anti-gravitactic response: vipers moved more frequently towards the top of the dune than in any other direction, despite a decrease in stride length with increasing slope. The foraging-related behaviour of the vipers was concentrated in the dune semi-stable areas rather than its stable or shifting sand areas. We measured rodent activity by placing seed trays in the dune allowing the rodents to collect seeds. Rodent activity was the highest in the shifting sands, closely followed by the semi-stable microhabitat. These results suggest the vipers use the semi-stable microhabitat mainly for foraging and may use the shifting sand areas as commuting routes between such areas. This study may be of use for conservation efforts of psammophilic species in desert dunes.

An experimental critique of the population carrying capacity concept.

Carrying capacity has multiple definitions, but nowadays, it is mainly referred to as the maximum number of individuals of a particular species sustained by the environment. We examined whether multiple populations of two flour beetle species grown under controlled laboratory conditions reach similar asymptotic population sizes when provided with similar amounts of food resources. We demonstrate that the variation in the asymptotic population sizes was considerably larger than that of the initial food resources and that the latter had no significant effect on the former. Our results experimentally contribute to past literature criticizing the carrying capacity concept, demonstrating that there is no single carrying capacity even under strict laboratory conditions. Therefore, we should not expect to often find "carrying capacities" in nature, where resources fluctuate over time, and interspecific interactions are ubiquitous. We suggest that the classic meaning of carrying capacity should be revisited or saved chiefly for didactic purposes.

The combined role of visual and olfactory cues in foraging by Cataglyphis ants in laboratory mazes.

Foragers use several senses to locate food, and many animals rely on vision and smell. It is beneficial not to rely on a single sense, which might fail under certain conditions. We examined the contribution of vision and smell to foraging and maze exploration under laboratory conditions using Cataglyphis desert ants as a model. Foraging intensity, measured as the number of workers entering the maze and arriving at the target as well as target arrival time, were greater when food, blue light, or both were offered or presented in contrast to a control. Workers trained to forage for a combined food and light cue elevated their foraging intensity with experience. However, foraging intensity was not higher when using both cues simultaneously than in either one of the two alone. Following training, we split between the two cues and moved either the food or the blue light to the opposite maze corner. This manipulation impaired foraging success by either leading to fewer workers arriving at the target cell (when the light stayed and the food was moved) or to more workers arriving at the opposite target cell, empty of food (when the food stayed and the light was moved). This result indicates that ant workers use both senses when foraging for food and readily associate light with food.

The value of spatial experience and group size for ant colonies in direct competition.

Animals often search for food more efficiently with experience. However, the contribution of experience to foraging success under direct competition has rarely been examined. Here we used colonies of an individually foraging desert ant to investigate the value of spatial experience. First, we trained worker groups of equal numbers to solve either a complex or a simple maze. We then tested pairs of both groups against one another in reaching a food reward. This task required solving the same complex maze that one of the groups had been trained in, to determine which group would exploit better the food reward. The worker groups previously trained in the complex mazes reached the food reward faster and more of these workers fed on the food than those trained in simple mazes, but only in the intermediate size group. To determine the relative importance of group size versus spatial experience in exploiting food patches, we then tested smaller trained worker groups against larger untrained ones. The larger groups outcompeted the smaller ones, despite the latter's advantage of spatial experience. The contribution of spatial experience, as found here, appears to be small, and depends on group size: an advantage of a few workers of the untrained group over the trained group negates its benefits.

Ant societies buffer individual-level effects of parasite infections.

Parasites decrease host fitness and can induce changes in host behavior, morphology, and physiology. When parasites exploit social insects, they influence not only infected individuals but also the society as a whole. Workers of the ant Temnothorax nylanderi are an intermediate host for the cestode Anomotaenia brevis. We studied a heavily parasitized population and found that, although parasite infection had strong and diverse consequences for individual workers, colony fitness remained unchanged. At the individual level, we uncovered differences among the three worker types, infected and healthy workers from parasitized colonies and healthy workers from nonparasitized colonies. Infected workers were smaller than healthy ones and had smaller heads as parasite load increased. Behavioral changes extended to all workers from parasitized colonies; such workers were less active than workers in nonparasitized colonies but engaged in more grooming. Healthy workers from parasitized colonies showed behavioral patterns intermediate to those of infected workers and healthy workers from nonparasitized colonies. Despite the lower activity level found in ants from parasitized colonies, an important fitness parameter, per-worker productivity, remained unaltered in parasitized colonies. However, the investment strategies of parasitized colonies changed as their sex ratio became male biased and as male body size increased. In short, ant colonies can buffer the drain of resources by the parasite despite strong effects on individual workers.