Increased turning per unit distance as an area-restricted search mechanism in a pause-travel predator, juvenile plaice, foraging for buried bivalves

During searching, discovery of a prey patch by juvenile plaice Pleuronectes platessa was associated with a change from extensive to intensive search behaviour several moves before an attack on a prey. Intensive search behaviour was characterized by reduced distance of moves, a greater rate of turning per unit distance and shorter pauses between moves. The increase in turn rate was associated with area-restricted seaching, while a decrease in distances moved suggests that plaice search more efficiently for prey when stationary than while moving. The klinokinetic mechanism that appears to regulate search behaviour in juvenile plaice should allow efficient exploitation of a range of prey distribution patterns based on localized cues alone. Such a mechanism is especially useful to a migratory predator, like plaice, whose foraging is subject to time constraints imposed by tidally available feeding areas.     

Saltatory search in a lateral line predator

The dwarf scorpionfish Scorpaena papillosa detected the hydrodynamic signals produced by prey with the mechanosensory lateral line. This species displayed a pause and move search pattern that is consistent with a saltatory search. The pause phase of the search cycle was probably used to detect prey because pauses often ended early in order to initiate an approach at prey and prey were detected throughout the search space. The move phase of the search cycle repositioned the fish so that it moved approximately a third of the reactive distance. Move distance was found to be the most important factor in gaining novel search space. Turning was shown to be relatively unimportant in gaining novel search space with a high frequency of low turn angles made by the fish. The dwarf scorpionfish, however, exhibited a spiralling or looping pattern over a search path exhibiting a turn bias towards either the left or right. The dwarf scorpionfish adopted a search behaviour that is consistent with a saltatory search and efficient for lateral line predation.     

Pausing of flagellar rotation is a component of bacterial motility and chemotaxis.

When bacterial cells are tethered to glass by their flagella, many of them spin. On the basis of experiments with tethered cells it has generally been thought that the motor which drives the flagellum is a two-state device, existing in either a counterclockwise or a clockwise state. Here we show that a third state of the motor is that of pausing, the duration and frequency of which are affected by chemotactic stimuli. We have recorded on video tape the rotation of tethered Escherichia coli and Salmonella typhimurium cells and analyzed the recordings frame by frame and in slow motion. Most wild-type cells paused intermittently. The addition of repellents caused an increase in the frequency and duration of the pauses. The addition of attractants sharply reduced the number of pauses. A chemotaxis mutant which lacks a large part of the chemotaxis machinery owing to a deletion of the genes from cheA to cheZ did not pause at all and did not respond to repellents by pausing. A tumbly mutant of S. typhimurium responded to repellents by smooth swimming and to attractants by tumbling. When tethered, these cells exhibited a normal rotational response but an inverse pausing response to chemotactic stimuli: the frequency of pauses decreased in response to repellents and increased in response to attractants. It is suggested that (i) pausing is an integral part of bacterial motility and chemotaxis, (ii) pausing is independent of the direction of flagellar rotation, and (iii) pausing may be one of the causes of tumbling.     

Long-term effects of suppressing the preratio pause

The preratio pause is a characteristic feature of performances under fixed-ratio schedules of reinforcement, even though the pause is not required by the schedule and it reduces the reinforcement rate. To investigate the reduction of pausing, five rats trained on fixed-ratio schedules were exposed to timeout punishment of pauses that exceeded a specified duration. After a series of 30 punishment sessions, most of the longest pauses were eliminated. For some subjects punishment was withdrawn abruptly, whereas for others a fading procedure was employed. Postpunishment observations then were continued for an additional 60 sessions. The reduced pausing was accompanied by reductions in the positive skew of the baseline distribution of pause durations, and by substantial increases in reinforcement rates. However, the results did not indicate differences as a function of the method of withdrawing the punishment contingency. Although postpunishment performances indicated some degree of recovery in the number of long pauses, performances had stabilized below prepunishment levels when the experiment ended. The results suggest the possibility that reduced pause durations can be self-maintained by the resulting increase in reinforcement rates.     

Effects of tail loss on the movement patterns of the lizard, Psammodromus algirus

1. Many lizards use caudal autotomy as a defensive strategy. However, subsequent costs related to the alteration of locomotor abilities might decrease the fitness of individuals. In this paper, the movement patterns of spontaneously moving Psammodromus algirus lizards and their escape performance running at high speed were compared before and after tail loss. A control tailed group was also studied to assess the repeatability of locomotor patterns between trials.
2. Tail loss had a significant effect on spontaneous movement patterns. Tailless individuals moved at significantly slower speeds during bursts of locomotion, and distances moved within bursts were significantly reduced. The overall time spent pausing increased, and, as a result, overall speeds decreased to an even greater extent than burst speeds. However, mean durations of individual locomotor bursts and mean pause durations did not change significantly after tail loss.
3. Loss of the tail decreased mean stride length, although the positive relation between stride length and speed was retained.
4. Escape performance was also greatly affected; loss of the tail resulted in substantially reduced attained, maximal and overall escape speeds. These changes resulted in shorter escape distances (the time of the first pause after the initiation of the escape response) because the mean duration of escape responses did not change.
5. The relevance of these alterations for the ecology of this species, and how individuals may compensate for the costs of tail loss, favouring autotomy as an escape strategy, are discussed.     

How to climb a tree: lizards accelerate faster,but pause more,when escaping on vertical surfaces

Many species of lizards effectively traverse both two and three‐dimensional habitats. However, few studies have examined maximum locomotor performance on different inclines. Do maximum acceleration and velocity differ on a level and inclined surface? Do lizards pause more on an inclined surface? To address these questions, Sceloporus woodi lizards (N = 12) were run in the laboratory on a level trackway and a vertical tree trunk. This species is known to frequently utilize both vertical and horizontal aspects of its habitat. Average maximum acceleration on the vertical surface exceeded that on the level surface, although average maximum velocity exhibited the opposite pattern. The average number of pauses during level locomotion was lower compared to vertical locomotion. In addition, the average location of the first pause on the level surface was 0.51 m, which is farther than the average for vertical locomotion where the first pause was at 0.35 m. The combination of performance and pause data suggests that the relative lack of pausing during level locomotion allows individuals to reach higher maximum velocities on level surfaces because they accelerate over greater distances. The increased pausing when moving vertically could be a result of high energetic demands of vertical locomotion, or greater microhabitat complexity as a result of branching and/or refuges. The faster acceleration exhibited during vertical locomotion by S. woodi likely offsets the frequent pauses. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 83–90.     

The movement patterns of lacertid lizards: speed, gait and pauses in Lacerta vivipara

Lacerta vivipara moving across an open space at their normal activity temperature alternate bursts of locomotion with short pauses which tend to occur at the extremes of the limb cycle, i.e. when individual limbs are maximally adducted or retracted and the spinal cord is maximally flexed in the lateral plane. The movement bursts and pauses in adult lizards have mean durations of 0–30 and 012 s, respectively, and within bursts the lizards move at a mean speed of 14–6 cm s^-1. Movement in juvenile lizards is 2–5 times faster (relative to body length) and the pauses are of longer duration (mean = 019 s), giving the locomotion of juveniles a more jerky appearance. Lizards which are chasing crickets increase the speed and the duration of locomo-tory bursts, although the pauses persist. Lizards which are searching for a previously perceived cricket increase pause duration (mean = 0–40 s). Lizards which are fleeing from a sudden disturbance move at almost twice (juveniles) or 3–7 times (adults) the speed of foraging animals: the pauses persist, although at much reduced frequency. Increases in speed result from increases in both stride length (Λ) and stride frequency (n); the ratio Λ/ n appears to remain constant at 006. The significance of these observations is discussed, although the functions of the pauses cannot yet be explained.     

Virtual plankton: A novel approach to the investigation of aquatic predator‐prey interactions

Small‐scale zooplankton swimming behaviors can affect aquatic predator‐prey interactions. Difficulties in controlling prey swimming behavior however, have restricted the ability to test hypotheses relating differences in small‐scale swimming behavior to frequency of predation by fish. We report here a Virtual Plankton (VP) system that circumvents this problem by allowing the observation of fish “preying"on computer‐generated prey images whose size, shape, color and swimming behavior can be precisely controlled. Two experiments were performed in which bluegill sunfish (Lepomis macrochirus) were given a choice of either two VP images, one of which moved twice as fast as the other, or six VP, one of which moved either faster (1.25 x, 1.5 x or 2 x ) or slower (0.5 x) than the other five. Current predator‐prey models based on encounter probabilities and prey visibility predict that moving faster increases predation risk and conversely, moving slower decreases predation risk. In agreement with existing predator‐prey models, in both experiments, fish chose faster moving VP significantly more often than their slower moving neighbors. Contrary to the predictions of existing models, in the second experiment with six VP, the rate at which fish chose a prey image moving half as fast as the five surrounding images did not differ significantly from the rate predicted by chance(l/6). These results suggest that current fish‐zooplankton predation models would benefit by the incorporation of small‐scale swimming behavior and assessments of its influence on overall prey visibility.     

The Behavioral Ecology of Intermittent Locomotion

Most physiological and ecological approaches to animal locomotionare based on steady state assumptions, yet movements of manyanimals are interspersed with pauses lasting from millisecondsto minutes. Thus, pauses, along with changes in the durationand speed of moves, form part of a dynamic system of intermittentlocomotion by which animals adjust their locomotor behaviorto changing circumstances. Intermittent locomotion occurs ina wide array of organisms from protozoans to mammals. It isfound in aerial, aquatic and terrestrial locomotion and in manybehavioral contexts including search and pursuit of prey, matesearch, escape from predators, habitat assessment and generaltravel. In our survey, animals exhibiting intermittent locomotionpaused on average nearly 50% of their locomotion time (range6–94%). Although intermittent locomotion is usually expectedto increase energetic costs as a result of additional expenditurefor acceleration and deceleration, a variety of energetic benefitscan arise when forward movement continues during pauses. Endurancealso can be improved by partial recovery from fatigue duringpauses. Perceptual benefits can arise because pauses increasethe capacity of the sensory systems to detect relevant stimuli.Several processes, including velocity blur, relative motiondetection, foveation, attention and interference between sensorysystems are probably involved. In animals that do not pause,alternative mechanisms for stabilizing the perceptual fieldare often present. Because movement is an important cue forstimulus detection, pauses can also reduce unwanted detectionby an organism's predators or prey. Several models have attemptedto integrate energetic and perceptual processes, but many challengesremain. Future advances will require improved quantificationof the effects of speed on perception.     

On optimal predator search

Search tactics are explored for animals hunting randomly distributed prey in two-dimensional habitats. The predator chooses between two modes of search: continuous travel and alternating pause-travel. A model based on renewal processes is derived for the predator's net rate of energy intake. The model is used to explore the optimal mode of search, search height, pause duration (“giving-up time”) and move length. Energy expenditure for search is assumed to increase from perchtravel over continuous travel to hover-travel. Prey detectability is assumed to be higher for the pausing than for the travelling predator. The following predictions emerge: The relative merits of the different search modes are mainly determined by the relative magnitudes of energy consumption rates and prey detection efficiencies at pausing and travelling. The energetically cheaper among two search modes reaches its highest relative merit at low prey density and detectability. As either increases, a more expensive search tactic may become superior. If the rate of energy expenditure increases considerably at locomotion, pause-travel tactics may be superior to continuous travel. This requires that the search can be performed from sufficient height, because net energy gain decreases rapidly below the optimum search height. It is greater for pause-travel than for continuous travel, and it increases slightly with decreasing prey density, and markedly with increasing prey detectability. With perch-travel or hover-travel tactics, the optimal giving-up time decreases with increasing prey density and detectability. The optimal move length increases with detectability. Empirical evidence coincides qualitatively with several predictions. Possibilities for further tests of the model are discussed, as are observed behavioural and morphological features on which the search model may shed light.     

Acoustic communication in a duetting grasshopper: receiver response variability, male strategies and signal design

In the duetting grasshopper Chorthippus biguttulus, a female's decision to reply to a conspecific male is based on the evaluation of a number of features of the male's song, which consists of uninterrupted syllables separated by pauses. Female responses are tuned to a restricted range of pause durations. However, males produce songs with noisy rather than silent pauses, which should make the measurement of pause durations more difficult for the female. We examined the adaptive value of these noisy pauses by testing female responses to (1) pairs of natural phrases, which differed only with respect to clear or noisy syllable pauses, and (2) synthetic phrases, in which the syllable onset accentuations and noise levels in the pauses were systematically varied. There was considerable variation between females, both in their preference for clear or noisy pauses in natural phrases, and in the optimal combinations of syllable onset accentuations and noise levels in pauses that they preferred in synthetic phrases. The response profiles of individual females were consistent. The experiments with synthetic phrases showed that, on average, females preferred more extreme values of syllable onset accentuations than were present in male songs. Noisy pauses increased the range of syllable pause durations accepted by females. The results suggest that noisy pauses could buffer signallers against the negative consequences of both signal degradation during transmission and extreme receiver choosiness.