Athletic anurans: the impact of morphology,ecology and evolution on climbing ability in invasive cane toads

Although primarily terrestrial, cane toads (Rhinella marina) sometimes climb near‐vertical surfaces (tree‐trunks, cliffs, fences) during foraging or dispersal activities. We scored climbing ability (in laboratory trials) of 288 cane toads from four regions in Australia, plus two sites on the island of Hawai'i. We found strong divergence in climbing ability associated not only with sex and relative limb length, but also population of origin. Within each population, longer‐limbed individuals (and hence, males rather than females) were better climbers, although the geographical divergence in climbing ability remained significant even when sex and limb length were included in multivariate regression models. The geographical difference in climbing ability (but not morphology) disappeared when the progeny were raised in captivity under identical conditions, without climbing opportunities. Although influenced by morphology, climbing ability in wild‐caught cane toads appears to be driven primarily by local environmental conditions that facilitate and/or reward arboreal activity.     

When Does Reward Maximization Lead to Matching Law?

What kind of strategies subjects follow in various behavioral circumstances has been a central issue in decision making. In particular, which behavioral strategy, maximizing or matching, is more fundamental to animal''s decision behavior has been a matter of debate. Here, we prove that any algorithm to achieve the stationary condition for maximizing the average reward should lead to matching when it ignores the dependence of the expected outcome on subject''s past choices. We may term this strategy of partial reward maximization “matching strategy”. Then, this strategy is applied to the case where the subject''s decision system updates the information for making a decision. Such information includes subject''s past actions or sensory stimuli, and the internal storage of this information is often called “state variables”. We demonstrate that the matching strategy provides an easy way to maximize reward when combined with the exploration of the state variables that correctly represent the crucial information for reward maximization. Our results reveal for the first time how a strategy to achieve matching behavior is beneficial to reward maximization, achieving a novel insight into the relationship between maximizing and matching.     

Investigating movement in the laboratory: dispersal apparatus designs and the red flour beetle,Tribolium castaneum

The natural dispersal of Tribolium castaneum Herbst (Coleoptera: Tenebrionidae) has been emulated in the laboratory for more than 50 years, using a simple dispersal apparatus. This has typically comprised of a starting container (initial resource or patch) connected by tubing, which contains thread for the animals to climb into a tube and hence to an end container. That is, beetles move to a new viable resource or patch from an inter‐patch zone or non‐viable habitat. We modified this basic apparatus design to test the effect of tubing length and tubing insertion angle on the dispersal rate and proportion of successful dispersers. We expected that the proportion of successful dispersers would be repeatable within each apparatus design, and that increasing tubing length and steepness of the insertion angle would reduce dispersal rate and success across apparatus designs. Dispersal increased linearly through time, similarly so for both males and females. The design with the most vertical tubing insertion angle had a lower proportion of successful dispersers. Tubing length also had a negative relationship with dispersal success (as judged by insects reaching the end container), but a significant reduction in dispersal success was only apparent between the shortest and longest tubing between containers. We suggest that locating and climbing the vertical section of string before they can enter the tubing between containers restricts dispersal and that at higher densities, insects exhibit greater inclination to climb. This type of apparatus has flexible design tolerances and further potential to study the dispersal of other small insect species that primarily use pedestrian locomotion.     

Effects of post-weaning environment on learning in the rat

Rats were reared in three different kinds of post-weaning environment: (1) with litter-mates and a variety of playthings (objects/social), (2) without either litter-mates or playthings (no-objects/isolates), (3) with litter-mates, but without playthings (no-objects/social). The animals were subsequently tested on a variety of learning tasks. In one task, the rats had to remove an obstacle from an alley in order to enter a food compartment; subsequently they were required to remove the obstacle in a different way from the one they had learned. Another task was to open a door leading to a food compartment; when the rats and learned this, the floor of the apparatus was lowered so they had to reach the door by climbing a ladder. Object-deprived/isolates were slower than the objects/social group in the transfer phase of the above tasks, though not in original acquisition. Object-deprived/social animals were not inferior to an objects/social group. Isolates had a higher free feeding weight than social animals, were more active in an open field, and ran faster for food reward when deprived.     

Learning from sensory and reward prediction errors during motor adaptation

Voluntary motor commands produce two kinds of consequences. Initially, a sensory consequence is observed in terms of activity in our primary sensory organs (e.g., vision, proprioception). Subsequently, the brain evaluates the sensory feedback and produces a subjective measure of utility or usefulness of the motor commands (e.g., reward). As a result, comparisons between predicted and observed consequences of motor commands produce two forms of prediction error. How do these errors contribute to changes in motor commands? Here, we considered a reach adaptation protocol and found that when high quality sensory feedback was available, adaptation of motor commands was driven almost exclusively by sensory prediction errors. This form of learning had a distinct signature: as motor commands adapted, the subjects altered their predictions regarding sensory consequences of motor commands, and generalized this learning broadly to neighboring motor commands. In contrast, as the quality of the sensory feedback degraded, adaptation of motor commands became more dependent on reward prediction errors. Reward prediction errors produced comparable changes in the motor commands, but produced no change in the predicted sensory consequences of motor commands, and generalized only locally. Because we found that there was a within subject correlation between generalization patterns and sensory remapping, it is plausible that during adaptation an individual''s relative reliance on sensory vs. reward prediction errors could be inferred. We suggest that while motor commands change because of sensory and reward prediction errors, only sensory prediction errors produce a change in the neural system that predicts sensory consequences of motor commands.     

Effect of the post-weaning environment on the climbing behaviour of wild and domestic Norway rats

The climbing behaviour of wild and domestic Norway rats (Rattus norvegicus) was compared after early rearing in three post-weaning environments offering different climbing experiences. Wild rats climbed in the test apparatus even when denied early climbing experience; male domestic rats did not. Early climbing experience increased the climbing scores of both stocks but influenced the climbing proficiency of wild rats only. Treatment differences in climbing behaviour may be related to specific motor experiences gained during development and the effect of early experience on the response to a novel environment (test apparatus). Stock differences in climbing behaviour may reflect a general reduction in motor activity among domestic rats and their reduced sensitivity to stimulus change or novelty.     

Corytibacladius Oliveira,Messias & Santos, 1995, a junior synonym of Limnophyes Eaton, 1875 (Diptera: Chironomidae: Orthocladiinae)

Larvae of the blackfly Simulium noelleri Friederichs aggregate at high populatio n densities on dams and spillways at the outlet of ponds. When displaced into the water column from their point of attachment, larvae can secrete silk threads as "life-lines" which enable them to maintain a link to the substratum an d up which they can climb to regain their original position. Experiments were conducted in a laboratory pond outlet to investigate this use of silk threads, larvae being displaced by means of forceps. It was demonstrated that: (i) the length of thread produced , and the speed of climbing the thread are independent of larval size; (ii) within limits, the speed of climbing was independent of both the length of the thread and the time already spent climbing, and (iii) speed of climb became more rapid as larvae neared the point of attachment. The range of locomotion in blackfly larvae is then discussed.     

Effects of Predation Risk on Vertical Habitat Use and Foraging of Pardosa milvina

Animals face the risk of predation while engaging in regular activities, such as foraging, mate‐seeking, and reproducing. In order to avoid predation, prey can modify behavior to prevent capture. Pardosa milvina may climb in response to chemotactile cues of Hogna helluo, a larger cooccurring wolf spider, to avoid predation. The purpose of this study was to test the effects of the location of predator cues on the climbing response of P. milvina and to test how this antipredator behavior affected foraging success. In experimental arenas, when cues were on the bottom of the containers, P. milvina moved upward, and when cues were on the walls, individuals moved downward. These results suggest that P. milvina respond to H. helluo cues with general avoidance and do not automatically climb in response to the cues. As H. helluo spend most of their time on the ground, P. milvina may avoid predation by spending more time climbing in areas with H. helluo cues. The presence of predator cues significantly decreased foraging by P. milvina. But within the predator cue treatments, climbing ability had no effect on foraging, possibly due to the short height of the feeding arenas. Future studies are needed to determine if climbing by P. milvina in response to cues of H. helluo has direct and indirect negative effects on herbivores in the field.     

Feeding kinematics and performance of Hawaiian stream gobies,Awaous guamensis and Lentipes concolor: Linkage of functional morphology and ecology

Distributions of Hawaiian stream fishes are typically interrupted by waterfalls that divide streams into multiple segments. Larvae hatch upstream, are flushed into the ocean, and must climb these waterfalls to reach adult habitats when returning back to freshwater as part of an amphidromous life cycle. Stream surveys and studies of climbing performance show that Lentipes concolor Gill can reach fast‐flowing upper stream segments but that Awaous guamensis Valenciennes reaches only slower, lower stream segments. Gut content analyses for these two species indicate considerable overlap in diet, suggesting that feeding kinematics and performance of these two species might be comparable. Alternatively, feeding kinematics and performance of these species might be expected to differ in relation to the different flow regimes in their habitat (feeding in faster stream currents for L. concolor versus in slower currents for A. guamensis). To test these alternative hypotheses, we compared food capturing kinematics and performance during suction feeding behaviors of A. guamensis and L. concolor using morphological data and high‐speed video. Lentipes concolor showed both a significantly larger gape angle and faster jaw opening than A. guamensis. Geometric models calculated that despite the inverse relationship of gape size and suction pressure generation, the fast jaw motions of L. concolor allow it to achieve higher pressure differentials than A. guamensis. Such elevated suction pressure would enhance the ability of L. concolor to successfully capture food in the fast stream reaches it typically inhabits. Differences in jaw morphology may contribute to these differences in performance, as the lever ratio for jaw opening is about 10% lower in L. concolor compared with A. guamensis, suiting the jaws of L. concolor better for fast opening. J. Morphol. 2009. © 2008 Wiley‐Liss, Inc.     

Interference and Shaping in Sensorimotor Adaptations with Rewards

When a perturbation is applied in a sensorimotor transformation task, subjects can adapt and maintain performance by either relying on sensory feedback, or, in the absence of such feedback, on information provided by rewards. For example, in a classical rotation task where movement endpoints must be rotated to reach a fixed target, human subjects can successfully adapt their reaching movements solely on the basis of binary rewards, although this proves much more difficult than with visual feedback. Here, we investigate such a reward-driven sensorimotor adaptation process in a minimal computational model of the task. The key assumption of the model is that synaptic plasticity is gated by the reward. We study how the learning dynamics depend on the target size, the movement variability, the rotation angle and the number of targets. We show that when the movement is perturbed for multiple targets, the adaptation process for the different targets can interfere destructively or constructively depending on the similarities between the sensory stimuli (the targets) and the overlap in their neuronal representations. Destructive interferences can result in a drastic slowdown of the adaptation. As a result of interference, the time to adapt varies non-linearly with the number of targets. Our analysis shows that these interferences are weaker if the reward varies smoothly with the subject''s performance instead of being binary. We demonstrate how shaping the reward or shaping the task can accelerate the adaptation dramatically by reducing the destructive interferences. We argue that experimentally investigating the dynamics of reward-driven sensorimotor adaptation for more than one sensory stimulus can shed light on the underlying learning rules.     

Distribution and habitat utilization in two atyid shrimps (Crustacea: Decapoda) in rivers of Nuku-Hiva Island (French Polynesia)

The distribution of Atyoida pilipes (Newport, 1847) and Caridina weberi De Man, 1892 was studied at two spatial scales on Nuku-Hiva Island (Marquesas, French Polynesia). At the stream reach scale, the two species were present at 40 and 39 sites, respectively, out of a total of 49 sites sampled. These two taxa represented more than half of the invertebrates collected at 9 sites. There was no difference in distribution between the species at the reach scale. However, clear differences appeared between the two species at the microhabitat scale. A. pilipes was more abundant in lotic habitats, even waterfalls, while C. weberi was associated with accumulations of leaves in lentic habitats. A. pilipes, particularly males, were tolerant to parameters such as water velocity, leaves, and the presence of algae and rocks. C. weberi had a more restricted altitudinal distribution and were scarce upstream from large waterfalls. These differences were linked to differences in climbing ability rather than the effect of chemical parameters, such as oxygen content. Our results suggest that little competition for space or food resources occurs between the two species. Both are feeding on small detritus particles, but through differing mechanisms and in differing habitats.     

Tonkean Macaques Orient Their Food Search From Olfactory Cues Conveyed by Conspecifics

We investigated the ability of monkeys to use referential information about environmental resources from cues passively transmitted by conspecifics. The subjects and their companions were four young males belonging to a group of semifree‐ranging Tonkean macaques (Macaca tonkeana) raised in a 2‐acre wooded park. In a first experiment, a subject had to orient its search strategy from the information obtained from a conspecific. It first had to smell the mouth of a companion having just eaten a food item, and then search for same quality food in the field. The results showed that subjects adjusted their foraging speed according to the value of the reward to be found. They were able to recognize the food resource on the basis of the item previously eaten by their companion. In a second experiment, we asked whether the subjects could anticipate a given food quantity from smelling a companion associated with that given food quantity. When testing whether subjects oriented their search strategy on the basis of companion's identity, they did not appear to use it as a referential cue to food quantity. They nonetheless adapted their foraging speed according to food value. Both experiments indicated that macaques are liable to assess and select food from information extracted from conspecifics, and hence avoid some costs associated with individual food searching.     

The range of passive arm circumduction in primates: Do hominoids really have more mobile shoulders?

Hominoids and lorines are assumed to possess greater shoulder mobility than other primates. This assumption is based on morphological characteristics of the shoulder, rather than on empirical data. However, recent studies have shown that the glenohumeral joint of hominoids is not more mobile than that of other primates (Chan LK. 2007. Glenohumeral mobility in primates. Folia Primatol (Basel) 78(1):1–18), and the thoracic shape of hominoids does not necessarily promote shoulder mobility (Chan LK. 2007. Scapular position in primates. Folia Primatol (Basel) 78(1):19–35). Moreover, lorines differ significantly from hominoids in both these features, thus challenging the assumption that both hominoids and lorines have greater shoulder mobility. The present study aims to test this assumption by collecting empirical data on shoulder mobility in 17 primate species. Passive arm circumduction (a combination of glenohumeral and pectoral girdle movement) was performed on sedated subjects (except humans), and the range measured on the video images of the circumduction. The motion differed among primate species mostly in the craniodorsal directions, the directions most relevant to the animal's ability to brachiate and slow climb. Hylobatids possessed the highest craniodorsal mobility among all primate species studied. However, nonhylobatid hominoids did not have greater craniodorsal mobility than arboreal quadrupedal monkeys, and lorines did not have greater craniodorsal mobility than arboreal quadrupedal prosimians. Nonhylobatid hominoids and lorines had similar craniodorsal mobility, but this was due to a longer clavicle, more dorsal scapula, and lower glenohumeral mobility in the former, and a shorter clavicle, less dorsal scapula, and greater glenohumeral mobility in the latter. This study provides evidence for the reexamination of the brachiation, slow climbing, and vertical climbing hypotheses. Am J Phys Anthropol, 2008. © 2008 Wiley‐Liss, Inc.     

Evolutionary trade‐offs in locomotor capacities in lacertid lizards: are splendid sprinters clumsy climbers?

We tested the hypothesis that an evolutionary trade‐off exists between the capacity to run on level terrain and the ability to climb inclined structures in lacertid lizards. Biomechanical and physiological models of lizard locomotor performance suggest that the morphological design requirements of a ground‐dwelling vs. scansorial life style are difficult to reconcile. This conflict is thought to preclude simultaneous evolution of maximal locomotor performance on level and inclined terrain. This notion has been corroborated by comparative studies on lizard species from other groups (Anolis, Chamaeleo, Sceloporus), but is not supported by our data on 13 species from the family Lacertidae. We found no indication of a negative association between maximal sprint speed of lizards over a level racetrack (indicative of ground‐dwelling locomotor performance), on an inclined stony surface (indicative of climbing performance over rock faces) and inclined mesh surface (indicative of clambering performance among vegetation). Moreover, morphological characteristics associated with fast sprinting capacities (e.g. long hind limbs) apparently enhance, rather than hinder climbing and clambering performance. We conclude that in our sample of lacertid lizards, the evolution of fast sprinting capacity on level terrain has not inflicted major restrictions on climbing and clambering performance.     

Oral health-related well-being of the long-term hospitalised elderly

Objective: To investigate oral health‐related well‐being of the long‐term hospitalised elderly as reported by their primary nurses in relation to subject's oral health assessed either by primary nurses or by a clinical dental examination. Background data: Little is known about oral health‐related well‐being of the medically compromised, long‐term hospitalised elderly, most of whom are unable to express their feelings and opinions. Materials and methods: A cross‐sectional study using a questionnaire for primary nurses about oral health and oral health‐related well‐being regarding functional, pain/discomfort‐related, and psychosocial limitations of the subjects (n = 255) and assessment of oral health by clinical examination. The total number of limitations and the number of limitations in each category was calculated. Results: Most (77%) of our subjects were unable to eat normal food. Functional limitations dominated followed by psychosocial and pain/discomfort‐related limitations. Overall assessment by each subject's primary nurse ranked oral health of as good for 9% of subjects, as moderate for 44%, and as poor for 47%. Clinical examination‐based assessment ranked oral health as good for 19%, as moderate for 33%, and as poor for 48% of our subjects, with good oral health being ranked as good for more men than women (26% vs. 16%; p = 0.045). Fewer limitations were recorded for those with better oral health assessed both by primary nurse and by clinical examination. Conclusion: More efforts are called for to maintain the oral health of the long‐term hospitalised elderly so as to improve their well‐being.     

Increased effort requirements and risk sensitivity: a comparison of delay and magnitude manipulations

Reward magnitude and delay to reward were independently manipulated in two separate experiments examining risk-sensitive choice in rats. A dual-running wheel apparatus was used and the tangential force resistance required to displace both wheels was low (50g) for half of the subjects, and high (120g) for the remaining subjects. Concurrent FI30-s and FI60-s schedules delivered equivalent amounts of food reward per unit time (i.e. 5 and 10 pellets of food, respectively), and these conditions served as the baseline treatment for all subjects. Variability, either in reward magnitude or delay, was introduced on the long-delay (60s) schedule during the second phase. All subjects were returned to the baseline condition in the third phase, and variability was introduced on the short-delay (30s) interval schedule during phase four. The subjects were again returned to the baseline condition in the fifth and final phase, ultimately yielding a five-phase ABACA design. Original baseline performance was characterized by a slight short-delay interval preference, and this pattern of performance was recovered with each subsequent presentation of the baseline condition. Overall, the data obtained from the reward magnitude and delay-to-reward manipulations were indistinguishable; subjects experiencing low-response effort requirement behaved in a risk-indifferent manner and subjects experiencing high-response effort requirement preferred the variable schedule. Implications for the daily energy budget rule on risk-sensitive foraging are discussed in light of these findings.     

DIFFERENT SPATIAL SCALES OF ADAPTATION IN THE CLIMBING BEHAVIOR OF PEROMYSCUS MANICULATUS: GEOGRAPHIC VARIATION,NATURAL SELECTION,AND GENE FLOW

Patterns of geographic variation in tree-climbing ability of Peromyscus maniculatus were used to examine the influence of spatial variation in natural selection and gene flow on the genetic divergence of climbing behavior among populations. Offspring of adults of two subspecies sampled from 10 localities in montane conifer forest, conifer woodland, and desert scrub/grassland habitats were raised in the laboratory and tested to determine their tree-climbing ability (the maximum diameter artificial rod that a mouse could climb). Comparisons of mean rod-climbing scores revealed that individuals of P. m. rufinus sampled from montane conifer forest and conifer woodland in Arizona were better climbers than P. m. sonoriensis sampled from conifer woodland and desert habitats in Nevada. This result was consistent with the hypothesis that natural selection has produced large-scale adaptation in climbing behavior. However, the climbing ability of P. m. sonoriensis sampled from conifer woodland habitats on isolated mountaintops in Nevada has not evolved in response to natural selection to the degree expected. In addition, populations sampled from desert grassland habitat, adjacent to woodland P. m. rufinus in Arizona, have climbing abilities that are not significantly different from conifer woodland populations. These observations indicate that local adaptation was constrained. An estimate of the heritability of climbing ability (h² = 0.352 ± 0.077) indicates that lack of a response to selection was not due to the absence of additive genetic variation. In addition, regressions of interpopulation differences on the degree of geographic isolation between pairs of populations do not support the hypothesis that gene flow between habitats has constrained evolution. Instead, a combination of historical events and insufficient time to respond to selection appears to have influenced geographic variation and the spatial scale of adaptation in climbing ability.     

How a lack of choice can force ants to climb up waxy plant stems

The aim of this study was to analyse the ability of generalist ants to climb waxy stems and to understand whether the avoidance of such stems is solely based on their reduced ability to adhere to these surfaces. We performed observations of the locomotory behaviour and recorded the traversed distances of Lasius niger ants on flower stems of three plant species covered with epicuticular wax crystals: Anethum graveolens, Dahlia pinnata, and Tagetes patula. Using conventional and cryo-scanning electron microscopy, the surface micromorphology of plant stems and structure of ant attachment organs were examined. The present study demonstrates that the reason ants do not climb wax-covered stems is not because they are incapable of walking on such vertical surfaces, as previously assumed, but rather because they have a choice of other substrates. Presumably, additional locomotory efforts are needed to master climbing on “greasy” stems, and ants need a certain time to adapt to a new quality of the stem surface, and after this period, they can walk on these waxy stems.     

An automated climbing apparatus to measure chemotherapy-induced neurotoxicity in Drosophila melanogaster

We have developed a novel model system in Drosophila melanogaster to study chemotherapy-induced neurotoxicity in adult flies. Neurological deficits were measured using a manual geotactic climbing assay. The manual assay is commonly used; however, it is laborious, time-consuming, subject to human error and limited to observing one sample at a time. We have designed and built a new automated fly-counting apparatus that uses a “video capture-particle counting technology” to automatically measure 10 samples at a time, with 20 flies per sample. Climbing behavior was assessed manually, as in our previous studies, and with the automated apparatus within the same experiment yielding statistically similar results. Both climbing endpoints as well as the climbing rate can be measured in the apparatus, giving the assay more versatility than the manual assay. Automation of our climbing assay reduces variability, increases productivity and enables high throughput drug screens for neurotoxicity.     

Modulation of Motor Area Activity by the Outcome for a Player during Observation of a Baseball Game

Background

Observing competitive games such as sports is a pervasive entertainment among humans. The inclination to watch others play may be based on our social-cognitive ability to understand the internal states of others. The mirror neuron system, which is activated when a subject observes the actions of others, as well as when they perform the same action themselves, seems to play a crucial role in this process. Our previous study showed that activity of the mirror neuron system was modulated by the outcome of the subject''s favored player during observation of a simple competitive game (rock-paper-scissors). However, whether the mirror neuron system responds similarly in a more complex and naturalistic sports game has not yet been fully investigated.

Methodology/Principal Findings

In the present study, we measured the activity of motor areas when the subjects, who were amateur baseball field players (non-pitchers), watched short movie clips of scenes in professional baseball games. The subjects were instructed to support either a batter or a pitcher when observing the movie clip. The results showed that activity in the motor area exhibited a strong interaction between the subject''s supported side (batter or pitcher) and the outcome (a hit or an out). When the subject supported the batter, motor area activity was significantly higher when the batter made an out than when he made a hit. However, such modulation was not apparent when the subject supported the pitcher.

Conclusions/Significance

This result indicates that mirror neuron system activity is modulated by the outcome for a particular player in a competitive game even when observing a complex and naturalistic sports game. We suggest that our inclination to watch competitive games is facilitated by this characteristic of the mirror neuron system.