Burrowing,Locomotion and Other Movements of the Echiuran Ochetostoma caudex

Burrowing, iocomotory and other movements of the echiuran Ochetostoma caudex have been examined and discussed. A continuous body cavity enables the worm to undergo peristaltic waves to pump water through the burrow without causing locomotion. The animal is capable of both forward and backward locomotion in its burrow. During forward locomotion, retrograde peristaltic waves are utilized which advance the animal in a step-wise fashion. Pressure changes within the coelom during burrowing, locomotion and during irrigation movements have been measured with the use of electronic recording techniques and the results interpreted in relation to direct visual observation. The structural and functional specializations for burrowing are discussed and compared with the activities of Priapulus caudatus, Sipunculus nudus and Bonellia viridis.     

Peristaltic waves of tubicolous worms and the problem of irrigation in Sabella pavonina

A simple scheme is presented to illustrate four possible kinds of locomotory peristalsis in worm-like animals. The application of this scheme to real animals is discussed. Peristaltic waves may be of constriction or dilatation. A continuous body cavity enables the worm to regulate both speed and direction of travel by controlling the relative tonus of its body wall muscles. Thus peristaltic waves can be used to pump water without causing locomotion.
Sabella irrigates its tube by peristaltic swellings but the coelom and intestine are sub-divided by entire septa. Anatomical and morphological features which allow the shortest, widest segments forming a "piston" to slide down the tube and the narrower elongated segments to grip its walls are considered. In this way the construction of the typical body segment is given a functional explanation.
The functions of septa in annelids are discussed.     

Observations on the locomotion of two British terrestrial planarians (Platyhelminthes, Tricladida)

The locomotion of Microplana terrestris and of M. britannicus is described. Forward locomotion is normally by means of cilia which are confined to the ventral surface of the animals. M. terrestris may however involve stationary peristaltic waves in locomotion. In this case neither muscular nor ciliary forces can alone account for the locomotion and it is necessary that the two mechanisms are combined. Such stationary waves should be distinguished from retrograde and direct locomotory waves. Reversal in both species is by retrograde muscular waves.     

Seismic signal transmission between burrows of the Cape mole-rat,Georychus capensis

Summary Both seismic and auditory signals were tested for their propagation characteristics in a field study of the Cape mole-rat (Georychus capensis), a subterranean rodent in the family Bathyergidae. This solitary animal is entirely fossorial and apparently communicates with its conspecifics by alternately drumming its hind legs on the burrow floor. Signal production in this species is sexually dimorphic, and mate attraction is likely mediated primarily by seismic signalling between individuals in neighboring burrows. Measurements within, and at various distances away from, natural burrows suggest that seismic signals propagate at least an order of magnitude better than auditory signals. Moreover, using a mechanical thumper which could be triggered from a tape recording of the mole-rat's seismic signals, we established that the vertically-polarized surface wave (Rayleigh wave) propagates with less attenuation than either of the two horizontally-polarized waves. Thus, we tentatively hypothesize that Rayleigh waves subserve intraspecific communication in this species.Abbreviations PPM pulses per min - SB simulated burrow - SD standard deviation - SPL sound pressure level     

Machaeridian locomotion

The discovery that machaeridians (class Machaeridia Withers, 1926) are annelids allows their mode of locomotion to be interpreted in the context of the body plan of this phylum. The Plumulitidae were errant epibenthic forms, moving with parapodia. The body of Turrilepadidae and Lepidocoleidae, however, was enclosed largely within the mineralized plates that make up the skeleton. Articulated specimens indicate that these machaeridians were able to burrow like other annelids using peristaltic locomotion. A lepidocoleid specimen indicates that multiple waves of shortened and contracted regions moved over the body. This is in contrast to the mode of locomotion in earthworms and most polychaetes, but similar to peristaltic progression in Polyphysia (Scalibregmidae). Either the rugose sculpture (turrilepadids) and/or the margins of the overlapping shell plates functioned as a burrowing sculpture, allowing forward movement but preventing backwards slipping. A trace from the Devonian Hunsrück Slate associated with a lepidocoleid indicates that considerable flexing of the skeleton was possible, but this is an escape trace and does not represent normal locomotion. Features of the skeleton of machaeridians are convergent on those of molluscs where the shells likewise function in protection and burrowing.     

Image analysis of the burrowing mechanisms of Polyphysia crassa (Annelida: Polychaeta) and Priapulus cudatus (Priapulida)

The burrowing mechanisms of two soft-bodied marine invertebrates, Polyphysia crassa (Annelida: Polychaeta) and Priapulus caudatus (Priapulida) were re-examined and extended utilizing computer image analysis. Predetermined points on sequential photographs of animals burrowing in methylcellulose (an artificial medium of high transparency) were digitized and stored as x, y coordinates from which were calculated segment length, width, and volume in addition to producing real and straightened images of the animal on a plotter. Both species are adapted to soft substrata and both utilize a direct peristaltic wave to advance the body into a cavity or loosened area formed by anterior structures. Polyphysia displaces and loosens the substratum with lateral scraping movements of its anterior six segments (the "head region") while Priapulus makes a large cavity anteriorly by the forceful eversion and dilation of the praesoma. Unlike Priapulus, Polyphysia utilizes a different method of locomotion when moving on the mud surface as opposed to burrowing, the former involving two direct peristaltic waves at a time and negligible internal pressures resulting in nearly continuous advance. However when burrowing, Polyphysia , like Priapulus , utilizes a single direct peristaltic wave alternating with phasic pressure pulses which advance the animal in step-wise fashion.     

Fast locomotion in caterpillars

The maximum forward crawling speeds of caterpillars are limited by the hydraulic design of the body and the peristaltic mode of operation of the segmental muscles. High speed locomotory manoeuvers can be achieved by reversing the direction of the normal peristaltic wave (from posterior-anterior to anterior-posterior) although the penalty is a dramatically reduced duty factor of the legs and potential instability. This study describes the suite of reverse gaits available to caterpillars, from reverse walking (the kinematic inverse of normal forward walking), through to reverse galloping (in which all the legs save the claspers are wrenched free of the ground with each step) to recoil-and-roll, a unique form of locomotion in which the insect free-wheels backwards at high-speed. These reverse forms of locomotion are produced primarily in response to threat, involve bilateral activation of the intersegmental muscles and are relatively simple in terms of neural control. The ecological roles of high-speed locomotion are considered in the light of potential predators and the normal habitat and terrain.     

Similarities and differences in the propagation of slow waves and peristaltic waves

The relationship between slow waves and peristaltic reflexes has not been well analyzed. In this study, we have recorded the electrical activity of slow waves together with that generated by spontaneous peristaltic contractions at 240 extracellular sites simultaneously. Recordings were made from five isolated tubular and six sheet segments of feline duodenum superfused in vitro. In all preparations, slow waves propagated as broad wave fronts along the longitudinal axis of the preparation in either the aborad or the orad direction. Electrical potentials recorded during peristalsis (peristaltic waves) also propagated as broad wave fronts in either directions. Peristaltic waves often spontaneously stopped conducting (46%), in contrast to slow waves that never did. Peristaltic waves propagated at a lower velocity than the slow waves (0.98 +/- 0.25 and 1.29 +/- 0.28 cm/s, respectively; P < 0.001; n = 24) and in a direction independent of the preceding slow wave direction (64% in the same direction, 46% in the opposite direction). In conclusion, slow waves and peristaltic waves in the isolated feline duodenum seem to constitute two separate electrical events that may drive two different mechanisms of contraction in the small intestine.     

Auditory evoked blink reflex and posterior auricular muscle response: Observations in patients with HFS and PFS

Our goal was to investigate the characteristics of the auditory brainstem reflexes in patients with hemifacial spasm (HFS) and postparalytic facial syndrome (PFS).The spasm activities and responses by supraorbital and auditory stimuli were recorded from the orbicularis oculi, the posterior auricular and the mentalis muscles in 27 HFS and 21 PFS patients. The results were compared with those of 20 controls.Blink reflex (BR) was obtained by supraorbital stimulation in normal controls and on both sides of HFS and PFS patients whereas sound evoked bilateral auditory blink reflex (ABR) in 96.3%, 90.5% and 100%, respectively. Both BR and ABR showed synkinetic spread on symptomatic sides in all patients. The posterior auricular muscle response (PAMR) was observed bilaterally in 59.3%, 42.9% and 75.0% of groups, respectively. However, there was no synkinetic spread of PAMR.Since PAMR does not show synkinetic spread even in the presence of synkinetic spread of ABR and BR, we may suggest that a distal origin may be responsible of the synkinetic spread, or PAM is probably governed by a smaller nucleus in the brainstem. Thus it may be speculated that its excitability is insufficient to stimulate the ABR nucleus, whereas the reverse process is possible.     

The locomotory activity patterns of a functionally complete colony of Cryptomys hottentotus hottentotus (Rodentia: Bathyergidae)

The locomotory activity patterns of a colony of eight field-captured mole-rats housed in a transparent burrow system were monitored for a total of 240 hours. Movement along the burrow system was recorded for all hours of the 24-hour chronological day for 10 days. Intermittent periods of activity were characteristic for all the mole-rats in the colony. The lack of a distinct sleeping period reflects the asynchrony of each mole-rat's activity cycle with that of other colony members. There are four major peaks of activity during the 24-hour cycle and an absence of a well-defined diurnal or nocturnal phase to the activity cycle.
The nest area is a focal point in the burrow system with individuals spending between 36 and 81% of the day there. There is a positive correlation between the size of a group of sleeping mole-rats in the nest and the frequency of occurrence.     

Locomotory activity of captive Cryptomys hottentotus, (Mammalia: Bathyergidae), a fossorial rodent

Locomotory activity of six mole-rats kept singly and in various sized groups was monitored for a total of 1728 hours by use of photocells, event recorder, and a 21 metre long, transparent burrow system. Movement in tunnelways was recorded for all hours of a 24 hour period. Locomotory activity for individuals and a two member group average14–24% of the total time, while a four member group averaged 57% of the total time. Approximately five rest periods per day averaged 50 minutes duration each. The effects of social isolation, burrow length, diet, and individual variation on locomotory activity of fossorial mammals are discussed.     

Opioids modulate periodicity rather than efficacy of peristaltic waves in the guinea pig ileum in vitro

The influence of opioid receptor blockade by naloxone and opioid receptor activation by opioids on peristalsis was studied in isolated segments of the guinea pig ileum.1. (-)Naloxone, but not (+)naloxone, increased the mean number of peristaltic waves per min within periods of elevated intraluminal pressure. Naloxone tended to modify intermittent peristalsis into ongoing peristalsis, whereas opioids worked in an opposite fashion. 2. Maximum amplitudes of luminal volume displacement during single peristaltic waves were not decreased by opioids. (-)Naloxone, however, applied to non-pretreated segments, decreased transitorily the efficacy of single peristaltic waves to a small, but statistically significant degree 3. Enhancement of peristalsis by naloxone decreased over time, although enough naloxone was present to occupy all opioid receptors. This suggests that opioid receptor blockade induces some compensatory mechanism.     

Courtship herding in the fiddler crab <Emphasis Type="Italic">Uca elegans</Emphasis>

Male and female animals are not always complicit during reproduction, giving rise to coercion. One example of a system that is assumed to involve sexual coercion is the mate herding behaviour of fiddler crabs: males push females towards the home burrow with the goal of forcing copulation at the burrow entrance. We recorded and analysed in detail the courtship behaviour of a North Australian species of fiddler crab Uca elegans. Courtship was composed of four main phases: broadcast waving, outward run, herding and at burrow display. During interactions males produced claw-waving displays which were directed posteriorly towards the female and which varied in timing and structure depending on the courtship phase. We suggest that courtship herding in U. elegans is driven primarily by mate choice for the following reasons, (1) females can evade herding, (2) no other reproductive strategies were observed, (3) males broadcast their presence and accompany courtship with conspicuous claw waves, and (4) the behaviour ends with the female leading the male into the home burrow. As an alternative function for herding in U. elegans we suggest that the behaviour represents a form of courtship guiding, in which males direct complicit females to the correct home burrow.     

Flow of urine through the ureter: a collapsible, muscular tube undergoing peristalsis

In steady flow through nonuniform collapsible tubes a key concept is the compressive zone, at which flow limitation can occur at both high and low Reynolds numbers. Ureteral peristalsis can be considered as a series of compressive zones, corresponding to waves of active muscular contraction, that move at near-constant speed along the ureter towards the bladder. One-dimensional, lubrication-theory analysis shows that peristalsis can pump urine from kidney into the bladder only at relatively low mean rates of urine flow. Under these circumstances isolated boluses of urine are propelled steadily through the ureter (assumed uniform) by the contraction waves. At higher mean rates of flow the behavior depends on whether the frequency of peristalsis is higher or lower than a critical value. For frequencies above the critical value steady propagation of boluses that are in contact with contraction waves at both ends is possible. As the flow rate rises the urine begins to leak through the contraction waves and steady peristaltic flow breaks down. There is an upper limit to the mean flow rate that can be carried by steady peristalsis, which depends on the mechanical properties of the ureter. At high flow rates the peristaltic contractions do not pump but hinder the flow of urine through the ureter.     

Mutation in<Emphasis Type="Italic"> slowmo</Emphasis> causes defects in<Emphasis Type="Italic"> Drosophila</Emphasis> larval locomotor behaviour

We have identified a mutant slowmotion phenotype in first instar larval peristaltic behaviour of Drosophila. By the end of embryogenesis and during early first instar phases, slowmo mutant animals show a marked decrease in locomotory behaviour, resulting from both a reduction in number and rate of peristaltic contractions. Inhibition of neurotransmitter release, using targeted expression of tetanus toxin light chain (TeTxLC), in the slowmo neurons marked by an enhancer-trap results in a similar phenotype of largely absent or uncoordinated contractions. Cloning of the slowmo gene identifies a product related to a family of proteins of unknown function. We show that Slowmo is associated with mitochondria, indicative of it being a mitochondrial protein, and that during embryogenesis and early larval development is restricted to the nervous system in a subset of cells. The enhancer-trap marks a cellular component of the CNS that is seemingly required to regulate peristaltic movement.     

Evaluation of peristalsis in multiple segments of the guinea-pig isolated small intestine: optimisation of tissue use by refined in vitro methodology

Peristalsis is the aboral movement by which the intestine propels its contents. Since pharmacological research requires an experimental model with which drug-induced modifications of peristalsis can be reliably quantified, we set out to develop and validate an in vitro method for studying peristalsis in multiple gut segments. In our arrangement, up to four 10cm segments isolated from the guinea-pig jejunum and ileum can be set up in parallel and their lumens perfused. Peristalsis was elicited by pressure-evoked wall distension, and the peristalsis-induced changes in the intraluminal pressure were evaluated with software that determined the peristaltic pressure threshold, the frequency, maximal acceleration and amplitude of the peristaltic waves, and the residual baseline pressure. Validation experiments showed that the peristalsis parameters at baseline and after modification by morphine (0.01-10microM) did not differ between segments from the jejunum and ileum, or between segments examined in a consecutive manner. In conclusion, our work succeeded in optimising the use of the guinea-pig jejunum and ileum for multiple recordings of peristalsis in vitro, and in refining the recording and evaluation of peristaltic motility. This system promises to be particularly useful in the pharmacological screening and testing of drugs which modify peristalsis.     

Particle size selection and sediment reworking in a funnel feeder,Leptosynapta tenuis (Holothuroidea,Synaptidae)

The burrowing behavior of the holothurian Leptosynapta tenuis, its lebensspuren, and the types of peristaltic waves used in movement and burrow irrigation are described. Particle size selection is a function of accessability of grains. L. tenuis cannot distinguish between particle sizes but nevertheless ingests small particles in a lesser proportion than that found in the surrounding sediment. Feeding rates are 2900–5500 g · yr⁻¹ · animal⁻¹. These rates are strongly dependent on temperature. Computation of a sediment mixing budget shows that of all the sediment ingested, 51% is ingested from the top 1/2 cm of sediment, 75% from the top 3 cm. The significance of the funnel as a ‘tool’ to exploit the top 1/2 cm of sediment is discussed. L. tenuis differs from other deposit feeders in that (1) it removes small particles from the surface sediment during its reworking activities, (2) it reworks sediment downward as well as upward, (3) its reworking activities tend to increase the stability of the upper 3 cm of the sediment, and (4) because so much sediment is ingested from the top 1/2 cm, it reworks an entire sediment column less efficiently per gram of sediment ingested than many other deposit feeders.     

Habitat selection by the Spiny-tailed lizard (Uromastyx aegyptia): A view from spatial analysis

Many factors affect the habitat selection for animal species, which in turn may greatly affect their distribution in different ecosystems. Understanding the processes that affect habitat selection is also critical for guiding and managing conservation initiatives. Our study aimed to assess the habitat selection by free-ranging Spiny-tailed lizard (Uromastyx aegyptia) by analyzing a geospatial data connecting its burrow parameters to different habitat characteristics within selected sites in Hail region, Saudi Arabia. We examined evidence and patterns of significant spatial clustering for (366) active burrows by linking their parameters (burrow entrance size, burrow entrance width and burrow entrance height), their reference geographical locations and, two habitat characteristics defined by soil type and vegetation cover. The objective of the analysis was to increase the understanding on the burrows aggregation process in the space and, to describe its possible relation to other spatial habitat configurations. Analysis of distances based on the Nearest Neighbor Index (NNI) and hotspots detection in Nearest neighbor hierarchical clustering (Nnh) suggested twelve (12) spatial clusters located within the study area. In addition, a spatial ordinary least square (OLS) and Poisson regression models revealed significant effects of soil type and vegetation cover on burrow parameters (OLS, p < 0.05; Poisson, p < 0.001), which indicate a strong association between burrows parameters and habitats characteristics. Findings from the study also suggest that other factors such as elevations, highways, and human settlement concentration spots could possibly play a major role in defining burrow spatial aggregation and furthermore have a significant impact on habitat selection.     

Review of diel rhythmic activities in physiological functions of scorpions

In this study I review the subject of biological rhythms in scorpions. This will include only some of the diel rhythmic endogenous physiological functions and not the locomotory rhythmicity, which will be dealt separately. Most studies reported here were conducted on 13 scorpion species that were studied in 39 different studies. Most of these (66.7%) were studies on a single species (Heterometrus fulvipes). Being a scorpionid, it sits and waits near its burrow not being very active, especially the females. The fact that experimenting was carried out irrespective of species diversity, gender, ecological or physiological conditions, and was usually done on animals kept in captivity for some time before the experimenting had started, is a major drawback to this kind of study. Although the main conclusion appears to be that enzymes reached their peak activity at 20.00 h, there are some exceptions showing otherwise that need further study in order to explain them.     

Comparison Between Tidally Driven Groundwater Flow and Flushing of Animal Burrows in Tropical Mangrove Swamps

Tidal groundwater in a mangrove swamp can return to the mangrove creek by one of two mechanisms: (a) it can either flow through the swamp soil due to the water table difference between the creek and the groundwater in the swamp; or (b) it can flow via tidal flushing of animal burrows. This paper compares the magnitude of these two mechanisms for different regions of a mangrove swamp. Direct groundwater flow rates resulting from water stored in the sediment as a consequence of infiltration, especially during and after tidal inundation, were calculated for every square meter in the surface of a mangrove forest from piezometer data. Flow rates of water due to burrow flushing were determined based on published surveys, by estimating the burrow volume and the percentage of the burrow water that is flushed at each tidal inundation. Although direct groundwater flux was found to decrease further away from the creek compared to close to the creek, it was also found to have a similar range as burrow flushing flow. Specifically, direct groundwater flow ranged from 0.004 to 0.04 m³/m²/day, whilst burrow flushing flux ranged from 0.01 to 0.04 m³/m²/day.Considering the errors involved in the experiments and calculations, these ranges can be considered as being the same and neither of the two processes can be considered as negligible compared to the other. As a consequence, surveys of groundwater processes in mangrove areas, and more generally in swamp and tidal areas where animal burrows are present, will need to consider both mechanisms. Investigations of the influence over flushing mechanisms of different residence times of the water in burrows and in the sediment body would also be recommended in order to establish salt and nutrient budget in mangrove swamps.