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.     

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.     

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.     

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.     

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.     

A mathematical model for the peristaltic flow of chyme movement in small intestine

A mathematical model based on viscoelastic fluid (fractional Oldroyd-B model) flow is considered for the peristaltic flow of chyme in small intestine, which is assumed to be in the form of an inclined cylindrical tube. The peristaltic flow of chyme is modeled more realistically by assuming that the peristaltic rush wave is a sinusoidal wave, which propagates along the tube. The governing equations are simplified by making the assumptions of long wavelength and low Reynolds number. Analytical approximate solutions of problem are obtained by using homotopy analysis method and convergence of the obtained series solution is properly checked. For the realistic values of the emerging parameters such as fractional parameters, relaxation time, retardation time, Reynolds number, Froude number and inclination of tube, the numerical results for the pressure difference and the frictional force across one wavelength are computed and discussed the roles played by these parameters during the peristaltic flow. On the basis of this study, it is found that the first fractional parameter, relaxation time and Froude number resist the movement of chyme, while, the second fractional parameter, retardation time, Reynolds number and inclination of tube favour the movement of chyme through the small intestine during pumping. It is further revealed that size of trapped bolus reduces with increasing the amplitude ratio whereas it is unaltered with other parameters.     

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.     

Kinematics of soft-bodied, legged locomotion in Manduca sexta larvae

Caterpillar crawling is distinct from that of worms and molluscs; it consists of a series of steps in different body segments that can be compared to walking and running in animals with stiff skeletons. Using a three-dimensional kinematic analysis of horizontal crawling in Manduca sexta, the tobacco hornworm, we found that the phase of vertical displacement in the posterior segments substantially led changes in horizontal velocity and the segments appeared to pivot around the attached claspers. Both of the motions occur during vertebrate walking. In contrast, vertical displacement and horizontal velocity in the anterior proleg-bearing segments were in phase, as expected for running gaits coupled by elastic storage. We propose that this kinematic similarity to running results from the muscular compression and release of elastic tissues. As evidence in support of this proposal, the compression and extension of each segment were similar to harmonic oscillations in a spring, although changes in velocity were 70 degrees out of phase with displacement, suggesting that the spring was damped. Measurements of segment length within, and across, intersegmental boundaries show that some of these movements were caused by folding of the body wall between segments. These findings demonstrate that caterpillar crawling is not simply the forward progression of a peristaltic wave but has kinetic components that vary between segments. Although these movements can be compared to legged locomotion in animals with stiff skeletons, the underlying mechanisms of caterpillar propulsion, and in particular the contribution of elastic tissues, remain to be discovered.     

Untersuchungen zur Frühontogenie vonNautilus pompilius (Linné)

A series of initial shells ofNautilus pompilius was investigated for morphologic and isotopic evidence of the early ontogenetic development. Shell morphologic features, such as curvature of early shell, cancellate sculpture, suture and grouping of septa, as well as very early shell injuries, color banding and shell/egg dimensions, seem to indicate an interpretation of early ontogeny differing from that which is now generally accepted. The O¹⁸/O¹⁶ and C¹³/C¹² ratios in the early shell and septa of twoNautilus specimens are given. Changes in the carbon isotope content are tentatively correlated with the end of the embryonic period and with environmental changes. Variations in the oxygen isotope content are in part ascribed to migrations from warm to cooler water after a certain stage of development. The size of the body chamber of the young animals can be determined by comparison of the isotope contents in outer shell and septa. Existing ideas concerning early ontogeny ofNautilus are critically discussed.     

A new look at the comparative physiology of insect and human hearts

Recent electrocardiographic (ECG) studies of insect hearts revealed the presence of human-like, involuntary and purely myogenic hearts. Certain insects, like a small light-weight species of hoverfly (Episyrphus balteatus), have evolved a very efficient cardiac system comprised of a compact heart ventricle and a narrow tube of aorta, which evolved as an adaptation to sustained hovering flights. Application of thermocardiographic and optocardiographic ECG methods revealed that adult flies of this species use the compact muscular heart chamber (heart ventricle) for intensive pumping of insect "blood" (haemolymph) into the head and thorax which is ringed all over with indirect flight musculature. The recordings of these hearts revealed extremely high, record rates of forward-directed, anterograde heartbeat (up to 10Hz), associated with extremely enhanced synchronic (not peristaltic) propagation of systolic myocardial contractions (32.2mm/s at room temperature). The relatively slow, backward-directed or retrograde cardiac contractions occurred only sporadically in the form of individual or twinned pulses replacing occasionally the resting periods. The compact heart ventricle contained bi-directional lateral apertures, whose opening and closure diverted the intracardiac anterograde "blood" streams between the abdominal haemocoelic cavity and the aortan artery, respectively. The visceral organs of this flying machine (crop, midgut) exhibited myogenic, extracardiac peristaltic pulsations similar to heartbeat, including the periodically reversed forward and backward direction of the peristaltic waves. The tubular crop contracted with a periodicity of 1Hz, both forwards and backwards, with propagation of the peristaltic waves at 4.4mm/s. The air-inflated and blindly ended midgut contracted at 0.2Hz, with a 0.9mm/s propagation of the peristaltic contraction waves. The neurogenic system of extracardiac haemocoelic pulsations, widely engaged in the regulation of circulatory and respiratory functions in other insect species, has been replaced here by a more economic, myogenic pulsation of the visceral organs as a light-weight evolutionary adaptation to prolonged hovering flight. Striking structural, functional and even genetic similarities found between the hearts of Episyrphus, Drosophila and human hearts, have been practically utilised for inexpensive testing of new cardioactive or cardioinhibitory drugs on insect heart.     

On the origin of leeches by evolution of development

Leeches are a unique group of annelids arising from an ancestor that would be characterized as a freshwater oligochaete worm. Comparative biology of the oligochaetes and the leeches reveals that body plan changes in the oligochaete-to-leech transition probably occurred by addition or modification of the terminal steps in embryonic development and that they were likely driven by a change in the feeding behavior in the ancestor of leeches. In this review article, developmental changes that are associated with the evolution of several leech-specific traits are discussed. These include (1) the evolution of suckers, (2) the loss of chaetae, (3) the loss of septa, and (4) a fixed number of segments. An altered developmental fate of the teloblast is further proposed to be a key factor contributing to the fixation of the segment number, and the evolutionary change in teloblast development may also account for the loss of the ability to regenerate the lost body segments in the leech.     

Locomotory and other movements of the trunk of Bonellia viridis (Echiura, Bonelliidae)

Trunk irrigatory and locomotory movements in the echiuran Bonellia viridis consist of peristaltic waves of constriction. Irrigation is by antero-posterior waves. Locomotion in the burrow may be produced in three different ways: (a) by antikinetic, (b) by synkinetic, and (c) by antikinetic and synkinetic waves in strict alternation. The animal can turn in the burrow and pass out of narrow holes. The animal can move by peristaltic waves also outside its burrow. The versatility of locomotory modes is related to its inhabiting ready-made burrows.     

Spontaneous motility of the oviduct in the anaesthetized pig

Intraluminal pressure microtransducers were placed at the uterotubal junction, the proximal isthmus, the ampullary-isthmic junction and the mid-ampulla. Spontaneous motility occurred throughout the oestrous cycle in all segments. During oestrus there were regular, high amplitude peristaltic waves in all segments, superimposed on basal activity. On Day 1 of the cycle the pattern was mostly antiperistaltic, presumably related to sperm transport. During the periovulatory period the number of peristaltic and antiperistaltic waves became equal, perhaps in relation to the transport of gametes to the fertilization site. During Day 3 there was no peristaltic activity; the motility patterns of the isthmus and ampullary-isthmic junction were similar (regular phasic contractions of high frequency and amplitude) while the ampullary motility was low. On Day 4, when the eggs enter the uterine lumen, the ampullary-isthmic junction and particularly the isthmus showed strong contraction waves (mostly peristaltic) superimposed on the basal phasic activity. This suggests an active role of the smooth muscle of the lower oviducal segments in ovum descent. During the mid- and late-luteal phases, the isthmus remained motile, with an irregular base line, but lost the pattern of basal contractions that dominated the activity during the first 4 days of the cycle. The ampulla showed low levels of spontaneous motility throughout the rest of the cycle.     

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.     

A mathematical simulation of the ureter: effects of the model parameters on ureteral pressure/flow relations

Ureteral peristaltic mechanism facilitates urine transport from the kidney to the bladder. Numerical analysis of the peristaltic flow in the ureter aims to further our understanding of the reflux phenomenon and other ureteral abnormalities. Fluid-structure interaction (FSI) plays an important role in accuracy of this approach and the arbitrary Lagrangian-Eulerian (ALE) formulation is a strong method to analyze the coupled fluid-structure interaction between the compliant wall and the surrounding fluid. This formulation, however, was not used in previous studies of peristalsis in living organisms. In the present investigation, a numerical simulation is introduced and solved through ALE formulation to perform the ureteral flow and stress analysis. The incompressible Navier-Stokes equations are used as the governing equations for the fluid, and a linear elastic model is utilized for the compliant wall. The wall stimulation is modeled by nonlinear contact analysis using a rigid contact surface since an appropriate model for simulation of ureteral peristalsis needs to contain cell-to-cell wall stimulation. In contrast to previous studies, the wall displacements are not predetermined in the presented model of this finite-length compliant tube, neither the peristalsis needs to be periodic. Moreover, the temporal changes of ureteral wall intraluminal shear stress during peristalsis are included in our study. Iterative computing of two-way coupling is used to solve the governing equations. Two phases of nonperistaltic and peristaltic transport of urine in the ureter are discussed. Results are obtained following an analysis of the effects of the ureteral wall compliance, the pressure difference between the ureteral inlet and outlet, the maximum height of the contraction wave, the contraction wave velocity, and the number of contraction waves on the ureteral outlet flow. The results indicate that the proximal part of the ureter is prone to a higher shear stress during peristalsis compared with its middle and distal parts. It is also shown that the peristalsis is more efficient as the maximum height of the contraction wave increases. Finally, it is concluded that improper function of ureteropelvic junction results in the passage of part of urine back flow even in the case of slow start-up of the peristaltic contraction wave.     

Ultrastructure of the dendritic outer segments of sensory cells in poreless (‘no-pore’) sensilla of insects

Summary Poreless sensilla with inflexible sockets in insects presumably house hygro- and thermoreceptors (type-1, type-2 receptors). The dendritic outer segments of these receptor cells were studied mainly in cryofixed antennae of two species of moth (Antheraea pernyi, A. polyphemus) and one beetle (Aleochara curtula). As a rule two type-1 receptor cells are present. Their dendritic outer segments do not branch. They project into the distal cuticular parts of the sensillum and are in close contact with its four-layered wall. The segments differ in shape and microtubule density. As well, in A. curtula, the microtubules are interconnected by electron-dense material for some distance, thus forming a tubular body-like structure of 1.3 m length. The dendritic outer segment of the single type-2 receptor cell is branched and lamellated. Its lamellae are connected by structures similar to septate junctions, which occupy about 70% of the total surface of the lamellated portion of the dendrite. In tangential sections, the septa appear as parallel strands approximately perpendicular to the long axis of the dendritic segment. The structure of type-1 receptors is discussed with regard to the hypothesis for a mechano-electrical transduction. The possible functions of lamellation and junctional connections in type-2 receptors are discussed.Supported by the Deutsche Forschungsgemeinschaft (SFB 4/G1)     

Spatiotemporal electrical and motility mapping of distension-induced propagating oscillations in the murine small intestine

Since the development of knockout animals, the mouse has become an important model to study gastrointestinal motility. However, little information is available on the electrical and contractile activities induced by distension in the murine small intestine. Spatiotemporal electrical mapping and mechanical recordings were made from isolated intestinal segments from different regions of the murine small intestine during distension. The electrical activity was recorded with 16 extracellular electrodes while motility was assessed simultaneously by tracking the border movements with a digital camera. Distension induced propagating oscillatory contractions in isolated intestinal segments. These propagating contractions were dictated by the underlying propagating slow wave with superimposed spikes. The frequencies, velocities, and direction of the propagating oscillations strongly correlated with the frequencies (r = 0.86), velocities (r = 0.84), and direction (r = 1) of the electrical slow waves. N(omega)-nitro-L-arginine methyl ester decreased the maximal diameter of the segment and reduced the peak contraction amplitude of the propagating oscillatory contractions, whereas atropine and verapamil blocked the propagating oscillations. Tetrodotoxin had little effect on the maximal diameter and peak contraction amplitude. In conclusion, distension in the murine small intestine does not initiate peristaltic reflexes but induces a propagating oscillatory motor pattern that is determined by propagating slow waves with superimposed spikes. These spikes are cholinergic and calcium dependent.     

Density gradient system. I. Formation and fractionation of density gradients

The instrumentation of a precise yet flexible system for the formation and fractionation of linear density gradients is described. The use of a special peristaltic pump and gradient-forming chambers permits the rapid formation of both dilute and concentrated solutions into ca. 5 to 300 ml of gradient. This gradient can be delivered to a single centrifuge tube or equally apportioned to as many as 24 tubes.The special peristaltic pump, which is described in the paper that follows, delivers the concentrated solution into the gradient-forming chamber as well as conveying the gradient from the chamber to the special centrifuge tube. A capillary in the wall of the tube conveys the gradient to the bottom, where it is dispersed into 18 equidistant orifices. This dispersion permits the rapid formation and fractionation of the gradient without disruption. A presettable counter coupled to the peristaltic pump permits the gradients to be apportioned into equal volume fractions.Results obtained with both dilute and concentrated gradients are presented.     

A decentralized control scheme for an effective coordination of phasic and tonic control in a snake-like robot

Autonomous decentralized control has attracted considerable attention because it enables us to understand the adaptive and versatile locomotion of animals and facilitates the construction of truly intelligent artificial agents. Thus far, we have developed a snake-like robot (HAUBOT I) that is driven by a decentralized control scheme based on a discrepancy function, which incorporates phasic control. In this paper, we investigate a decentralized control scheme in which phasic and tonic control are well coordinated, as an extension of our previous study. To verify the validity of the proposed control scheme, we apply it to a snake-like robot (HAUBOT II) that can adjust both the phase relationship between its body segments and the stiffness at each joint. The results indicate that the proposed control scheme enables the robot to exhibit remarkable real-time adaptability over various frictional and inclined terrains. These findings can potentially enable us to gain a deeper insight into the autonomous decentralized control mechanism underlying the adaptive and resilient locomotion of animals.     

Neuronal control of heartbeat in the medicinal leech

Summary The leech heartbeat consists of a constriction-dilation rhythm of two lateral heart tubes extending over the length of the body. The beats of the segmental sections of these two tubes are coordinated in such a manner that the heart tube of one body side produces a frontward peristaltic wave while the heart tube on the other body side produces nearly concerted constrictions. This rhythm is metastable, in that left and right heart tubes alternate between peristaltic and concerted constriction modes, with a given mode lasting for tens or hundreds of beat cycles.The constriction-dilation cycles of the segmental heart tube sections are controlled by a set of rhythmically active motor neurons, the heart excitors, or HE cells. A bilateral pair of HE cells is located in all but the two frontmost and the two rearmost segmental ganglia of the ventral nerve cord. Each HE cell innervates via excitatory synapses the circular muscle fibers in the wall of the ipsilateral heart tube section. The activity cycle of the HE cells consists of an active phase, during which they are depolarized and produce a burst of impulses, and an inactive phase during which they are repolarized by a burst of inhibitory synaptic potentials. The intersegmentally coordinated activity cycles of the HE cell set are maintained in an isolated ventral nerve cord. Hence the generation of the heart excitor rhythm does not require sensory feedback.We are indepted to Amy Kelly and King-Wai Yau for advice on the use of the intracellular staining technique and to John Kretz for calling to our attention the existence of an afferent impulse burst rhythm emanating from denervated heart tube preparations. We thank Georgia Harper for excellent technical assistance. This research was supported by Grant GB 31933X from the National Science Foundation and NIH research grants GM17866 and Training Grant GM 01389 from the Institute of General Medical Sciences.