Water-flow sensitive pedal neurons in Tritonia: role in rheotaxis

Summary We have identified 13 pairs of neurons in the pedal ganglia of the marine nudibranch slug Tritonia diomedea that responded tonically and/or phasically to water-flow directed at the rhinophore sheaths and oral veil tips. Most of the neurons responded equally to inputs from either side of the body, but 6 pairs responded with greater intensity to ipsilateral water-flow stimuli. When stimulated intracellularly in a semi-intact, whole-animal preparation, 4 of these 6 pairs of neurons caused ipsilateral movements that may turn the animal towards that side. These observations suggest a role for these current-sensitive neurons in the previously described orientation to water-currents in Tritonia diomedea.     

Conservation of a Tritonia Pedal peptides network in gastropods

Adults of the nudibranch mollusc Tritonia diomedea crawl using mucociliary locomotion. Crawling is controlled in part by the large Pedal 5 (Pd5) and Pedal 6 (Pd6) neurons that produce Tritonia Pedal peptides (TPeps). TPeps elicit an increase in ciliary beat frequency, thereby increasing crawling speed. In adults of T. diomedea, an extensive network of TPep‐containing neurites adjacent to the basement membrane of the pedal epithelium delivers TPeps to the ciliated cells. In this study, we show that diverse nudibranchs all have a pattern of TPep‐like immunoreactivity similar to that of T. diomedea, with thin tracts of TPep‐like immunoreactive (TPep‐LIR) neurites projecting to the epithelial layer. We also show that members of two non‐nudibranch gastropod species have a pattern of TPep‐innervation similar to that of the nudibranchs. In addition, we characterized two pairs of motor neurons in adults of the nudibranch Armina californica that are possible homologues of the Pd5 and Pd6 cells in T. diomedea. Activity in one of these pairs, the Pedal Peptidergic Dorsal 1 (PPD1) cells, was correlated with mucociliary locomotion. The second pair, the Pedal Peptidergic Ventral 1 cells, shared synchronous synaptic input with the PPD1 cells, a pattern consistent with the shared synaptic input of the T. diomedea Pd5 and Pd6 cells. These findings suggest that the roles of the Pd5 and Pd6 cells as mucociliary motor neurons in nudibranchs are conserved evolutionarily. Additionally, the extensive network of TPep‐LIR neurites seen in the foot of T. diomedea appears likely to be a common feature among gastropods.     

Function of identified nerves in orientation to water flow in Tritonia diomedea

We determined which sensory and motor nerves mediate orientation to flow in the marine slug Tritonia diomedea, and tested the hypothesis that the slug orients to water flow by comparing the intensities of water flow stimulation on each side of its body. Lesion experiments revealed which nerves carried information necessary for flow orientation. The lateral branches of Cerebral Nerve # 2 were the only cerebral nerves necessary for flow orientation. Cutting all cerebral nerves except the lateral branches of Cerebral Nerve # 2 did not eliminate flow orientation. Thus, the lateral branches of Cerebral Nerve # 2 were both necessary and sufficient (among the cerebral nerves) for flow orientation. Denervation of one side of the head by cutting Cerebral Nerves # 1–4 on one side did not eliminate normal flow orientation. We have revised our model of how Tritonia diomedea orients to flow to allow for this unilateral determination of flow direction. Unilaterally cutting Pedal Nerve # 3, which contains many pedal motor axons, reduced turning toward that side, but did not affect final orientation to flow. The ability to detect flow direction was not compro mised by the inability to initially turn towards flow.Abbreviations CeN cerebral nerve - PeN pedal nerve - PlN pleural nerve     

Immunochemical and electrophysiological analyses of magnetically responsive neurons in the mollusc Tritonia diomedea

Tritonia diomedea uses the Earth’s magnetic field as an orientation cue, but little is known about the neural mechanisms that underlie magnetic orientation behavior in this or other animals. Six large, individually identifiable neurons in the brain of Tritonia (left and right Pd5, Pd6, Pd7) are known to respond with altered electrical activity to changes in earth-strength magnetic fields. In this study we used immunochemical, electrophysiological, and neuroanatomical techniques to investigate the function of the Pd5 neurons, the largest magnetically responsive cells. Immunocytochemical studies localized TPeps, neuropeptides isolated from Pd5, to dense-cored vesicles within the Pd5 somata and within neurites adjacent to ciliated foot epithelial cells. Anatomical analyses revealed that neurites from Pd5 are located within nerves innervating the ipsilateral foot and body wall. These results imply that Pd5 project to the foot and regulate ciliary beating through paracrine release. Electrophysiological recordings indicated that, although both LPd5 and RPd5 responded to the same magnetic stimuli, the pattern of spiking in the two cells differed. Given that TPeps increase ciliary beating and Tritonia locomotes using pedal cilia, our results are consistent with the hypothesis that Pd5 neurons control or modulate the ciliary activity involved in crawling during orientation behavior.     

Properties of central motor neurons exciting locomotory cilia inTritonia diomedea

Summary A pair of large, identifiable neurons (Pd 21), one in each pedal ganglion, can excite previously inactive locomotory cilia on the sole of the foot ofTritonia diomedea (Audesirk, 1978; Fig. 3). These neurons exert their effect via axons which innervate the foot and are probably central motor neurons for pedal cilia. IntactTritonia are stimulated to crawl by the application of 1.5 M NaCl to the tail, and conversely usually stop crawling when the chemosensitive oral veil is touched with food (sea whip,Virgularia sp.). The Pd 21 neurons are excited by 1.5 M NaCl applied externally to the tail, and are inhibited by sea whip touch to the oral veil (Figs. 4 and 5). When aTritonia performs its escape swim, the cilia move strongly, and the Pd 21 neurons fire bursts of spikes in phase with dorsal flexions (Figs. 6 and 7). After a swim, aTritonia rapidly crawls along the substrate; during this time the spiking rate of the Pd 21s is greatly accelerated. Interneurons thought to drive swim bursts produce monosynaptic EPSPs in the Pd 21s (Fig. 8). The Pd 21s are coordinated in their spike activity by synaptic activity which is synchronous in the two neurons regardless of the site of external stimulation, and by electrical coupling between the two cells via axons in a pedal commissure (Figs. 9 and 10). The coupling coefficient for passively conducted potentials is quite high, about 0.15, despite an axon 8 to 12 mm long separating the two cells.Abbreviations BPSP biphasic postsynaptic potential - SW sea water     

Measurment of pedal loading in bicycling: I. Instrumentation

This paper presents a new instrumentation system to precisely measure pedal loads and pedal position. A pedal/dynamometer unit implementing four octagonal strain rings measures all six load components between the foot and pedal. To study the relationship between foot position and loading, the pedal/dynamometer offers three degree-of-freedom adjustability. Pedal position along the pedal arc is precisely described by measuring crank arm angle and relative angle between pedal and crank arm. Linear, continuous rotation potentiometers measure the two angles. Transducer signals are sampled by a digital computer which calculates resultant loads and pedal position as functions of crank arm angle. Transducers are designed to mount on most bicycles without modification. Test subjects ride their own bicycles unconstrained on rollers so that loading data is representative of actual cycling.     

In vivo responses of paired giant mechanoreceptor neurons in Aplysia abdominal ganglion

Two neurons with cell bodies symmetrically located in the abdominal ganglion and giant axons in the left (L1) and right (R1) pleurovisceral connectives of Aplysia californica were examined in vivo and in vitro. Direct stimulation of R1 and L1 in the intact animal does not elicit any observable behavior, suggesting that they are neither motoneurons nor command neurons. These cells respond in vivo to sudden onset mechanical stimulation of widespread regions of the body. R1 and L1 spikes are initiated in at least three different loci: (1) the peripheral axon in the foot, (2) the neuropil of the pleural and/or pedal ganglion, and (3) the neuropil of the abdominal ganglion. Furthermore, R1 and L1 probably have two different mechanisms for spike initiation: (1) sensory (foot), and (2) synaptic (abominal and/or head ganglia). The different loci for spike initiation account for the bidirectional conduction of R1 and L1 spikes. As sensory (mechanoreceptor) neurons, R1 and L1 have peripheral axons in the ipsilateral posterior pedal nerve, show low threshold responses to stimulation of the ipsilateral posterior foot, they are rapidly adapting their responses do not decrease with repetion, and they are not blocked by high Mg⁺⁺/low Ca⁺⁺ solutions. As synaptically-driven neurons, R1 and L1 have widespread bilateral responsiveness, their responses decrease with repetition and their inputs are blocked with high Mg⁺⁺/low Ca⁺⁺ solutions. These neurons integrate sensory and synaptic inputs and conduct bidirectionally, however, their output connections must be specified before their behavioral function can be understood.     

New pedal remains of Megaladapis and their functional significance

New remains of Megaladapis from the caves within the Ankarana Range of northern Madagascar and the cave site of Ankilitelo near Toliara in southwestern Madagascar add considerably to the present sample of pedal remains for this genus. Here we describe and analyze the new pedal material and discuss the function of the Megaladapis foot in terms of positional behavior and substrate use. The northern specimens belong to the M. madagascariensis/M. grandidieri group in terms of size and morphology, whereas the new southwestern fossils are assigned to M. madagascariensis. The new specimens demonstrate that the small and intermediate sized M. madagascariensis and M. grandidieri were very similar in anatomy and inferred locomotor function, findings that also support the prior suggestion that they belong to a single widespread subgenus (Megaladapis). The new fossils provide the first examples of many pedal elements and present the first opportunity to analyze the whole pedal complex from associated remains. The foot of Megaladapis is distinctive among primates in numerous features. Intrinsic proportions of the hindlimb indicate that the foot is relatively longer than that of any other primate. The first complete calcanei reveal a large and highly modified hindfoot. The calcaneus is reduced distally, indicating an emphasis on climbing over leaping or quadrupedal walking and running. Proximally, a large, medially directed calcaneal tuberosity suggests both a strong inversion component to plantarflexion and a well-developed abductor mechanism and recalls the calcaneal morphology of the larger lorisines in some respects. Talar shape is consistent with considerable tibial rotation during plantarflexion and dorsiflexion. The subtalar joint is designed to emphasize supination/pronation and medial/lateral rotation over proximodistal translation. The distal tarsals are extremely reduced in length, and they form a high transverse arch and a serial tarsus; this configuration promotes inversion/eversion at the transverse tarsal joint. The phalanges are long and moderately curved, and the hallux is very long, robust, and abducted. Pedal morphology suggests that Megaladapis (subgenus Megaladapis) was well adapted to exploit an arboreal environment. The grasping mechanism of Megaladapis is an extreme modification of the prosimian condition, emphasizing a highly inverted set, mobility in rotation, and a powerful abduction/flexion type grasp using large hallux and the lateral abductor musculature. Such a mechanism insures a secure grasp regardless of the position of the hindlimb or the substrate. These pedal design features contrast with the grasping strategy seen in highly arboreal palaeopropithecids (or “sloth lemurs”), a group that reduces and modifies the hindfoot, culminating in Palaeopropithecus, and emphasizes extrinsic digital flexors in a more hook-like mechanism. Much less is known of M. (Peloriadapis) edwardsi. The larger body size, more gorilla-like talar articular morphology, and anatomy of the proximal fifth metatarsal suggest that this species may have been more terrestrial than the smaller forms, but other aspects of pedal morphology suggest it also exploited arboreal habitats.     

Forewing movements and motor activity during roll manoeuvers in flying desert locusts

Desert locusts, tethered on a roll torque meter and flying in a wind tunnel are surrounded by an artificial horizon (Fig. 1). Flight motor activity and movement of forewings are monitored continuously. Movements of the artificial horizon elicit roll manoeuvers of the animal with latencies of several seconds; concomitant changes in flight motor pattern and wing movement can be correlated with the animal's roll angle and roll torque. Flight sequences with constant torque and roll angle (steady state) have been analysed with the following results. Wing Kinematics. A phase difference between the movements of the forewings on either side is correlated with roll angle (Fig. 3). Pronation of a forewing is always greater on the side to which the animal rolls, i.e. on the side that produces less lift (Fig. 5). In some experiments the slope of the wing tip path is also decreased (Fig. 5). In both cases, the aerodynamic angle of attack is decreased and the forewing on this side produces less lift. In most experiments, changes in pronation are less pronounced in the contralateral wing (Fig. 11). All factors contribute to a net roll torque that sustains the animal's roll angle. Other kinematic parameters of forewing movement, e.g. wing stroke amplitude, were not found to be correlated with roll angle and torque (Fig. 4). Motor Pattern. Activity of several flight muscles (depressors M97, M98, M99, and M129; elevators M83, M84, and M90) was investigated for changes in burst length and temporal coordination in response to roll stimuli. Most flight muscles fired only once per wing beat cycle in our preparations. Thus, burst length was not found to be correlated with roll angle. Time intervals of firing between all muscle pairs investigated change in correlation with the torque and roll angle (Fig. 9).All mesothoracic muscles are active earlier-relative to the ipsilateral metathoracic subalar muscle M129-during roll to the ipsilateral side than during roll to the contralateral side. Correlations Between Motor and Movement Pattern. The phase of muscle firing within the wing beat cycle varies with roll angle (roll torque). The first basalar M97 and second tergosternal M84 muscles, when referenced e.g. to the upper (M97) or lower (M84) reversal point of the wing tip trajectory, are active earlier on the side the animal turns to (Fig. 10). The onset of the first basalar M97 relative to the beginning of downstroke is correlated with maximum pronation and roll angle (Fig. 11). Mechanisms of Lift Control. Wing pronation, which is very important for lift production is controlled by the central nervous system by altering the phase of muscle activity within the wing beat cycle.     

Characterization of the synaptic actions of an interneuron in the central nervous system of Tritonia

The motor program that drives the swimming behavior of the marine mollusk Tritonia diomedea is generated by three interneuronal populations in the cerebral ganglia. One of these populations, the pair of C2 neurons, is shown to also exert powerful synaptic actions upon most cells in the contralateral pedal ganglion. Intracellular staining with Co²⁺ showed that the C2 neurons projected to the contralateral pedal ganglion as a single unbranched axon, and nearly all contralateral pedal neurons received monosynaptic input from C2. Orthodromic stimulation of most peripheral nerves caused monosynaptic excitation of C2 by afferent sensory cells and, in some cases, monosynaptic inhibition from an unidentified source. C2 neurons produced four types of postsynaptic potential (PSP) on pedal neurons: (1) a fast, Cl^?-mediated inhibition (FIPSP); (2) a fast, Na⁺-mediated excitation (FEPSP); (3) a slow, K⁺-mediated inhibition (SIPSP); and (4) a slow, conductance-decrease excitation (SEPSP). All four could be recorded simultaneously in some pedal neurons. The C2 neurons appear to be high-order, multiaction neurons involved in both the generation of a complex motor program and the coordination of ancillary neuronal activity.     

Involvement of pedal peptide in locomotion in Aplysia: modulation of foot muscle contractions

Pedal peptide (Pep) is a 15-amino-acid neuropeptide that is localized within the Aplysia central nervous system (CNS) predominantly to a broad band of neurons in each pedal ganglion. Pep-neurons were identified by intracellular staining and immunocytology or by radioimmunoassay (RIA) of extracts from identified neurons. RIA reveals that 97% of all Pep-like immunoreactivity (IR-Pep) in pedal nerves is found in the three nerves that innervate the foot. Nearly every Pep-neuron sends an axon out at least one of these three nerves. Application of Pep to foot muscle causes an increase in the amplitude and relaxation rate of contractions driven by nerve stimulation or intracellular stimulation of pedal motor neurons. The increase in relaxation rate was the predominant effect. Intracellular recording in "split-foot" preparations reveals that Pep-neurons increase their overall firing rates and fire in bursts with each step during locomotion. Recovery of IR-Pep from foot perfusate following pedal nerve stimulation increases in a frequency-dependent fashion. Thus it appears that one function of Pep-neurons is to modulate foot muscle contractility during locomotion in Aplysia.     

Water and side‐chain embedded π‐turns

Elucidating protein function from its structure is central to the understanding of cellular mechanisms. This involves deciphering the dependence of local structural motifs on sequence. These structural motifs may be stabilized by direct or water‐mediated hydrogen bonding among the constituent residues. π‐Turns, defined by interactions between (i) and (i + 5) positions, are large enough to contain a central space that can embed a water molecule (or a protein moiety) to form a stable structure. This work is an analysis of such embedded π‐turns using a nonredundant dataset of protein structures. A total of 2965 embedded π‐turns have been identified, as also 281 embedded Schellman motif, a type of π‐turn which occurs at the C‐termini of α‐helices. Embedded π‐turns and Schellman motifs have been classified on the basis of the protein atoms of the terminal turn residues that are linked by the embedded moiety, conformation, residue composition, and compared with the turns that have terminal residues connected by direct hydrogen bonds. Geometrically, the turns have been fitted to a circle and the position of the linker relative to its center analyzed. The hydroxyl group of Ser and Thr, located at (i + 3) position, is the most prominent linker for the side‐chain mediated π‐turns. Consideration of residue conservation among homologous sequences indicates the terminal and the linker positions to be the most conserved. The embedded π‐turn as a binding site (for the linker) is discussed in the context of “nest,” a concave depression that is formed in protein structures with adjacent residues having enantiomeric main‐chain conformations. © 2013 Wiley Periodicals, Inc. Biopolymers 101: 441–453, 2014.     

ω‐Turn: A novel β‐turn mimic in globular proteins stabilized by main‐chain to side‐chain CH···O interaction

Mimicry of structural motifs is a common feature in proteins. The 10‐membered hydrogen‐bonded ring involving the main‐chain C?O in a β‐turn can be formed using a side‐chain carbonyl group leading to Asx‐turn. We show that the N? H component of hydrogen bond can be replaced by a C^γ‐H group in the side chain, culminating in a nonconventional C? H···O interaction. Because of its shape this β‐turn mimic is designated as ω‐turn, which is found to occur ～three times per 100 residues. Three residues (i to i + 2) constitute the turn with the C? H···O interaction occurring between the terminal residues, constraining the torsion angles ?_{i + 1}, ψ_{i + 1}, ?_{i + 2} and χ′_{1(i + 2)} (using the interacting C^γ atom). Based on these angles there are two types of ω‐turns, each of which can be further divided into two groups. C^β‐branched side‐chains, and Met and Gln have high propensities to occur at i + 2; for the last two residues the carbonyl oxygen may participate in an additional interaction involving the S and amino group, respectively. With Cys occupying the i + 1 position, such turns are found in the metal‐binding sites. N‐linked glycosylation occurs at the consensus pattern Asn‐Xaa‐Ser/Thr; with Thr at i + 2, the sequence can adopt the secondary structure of a ω‐turn, which may be the recognition site for protein modification. Location between two β‐strands is the most common occurrence in protein tertiary structure, and being generally exposed ω‐turn may constitute the antigenic determinant site. It is a stable scaffold and may be used in protein engineering and peptide design. Proteins 2015; 83:203–214. © 2014 Wiley Periodicals, Inc.     

Redesigning the type II' β‐turn in green fluorescent protein to type I': Implications for folding kinetics and stability

Both Type I' and Type II' β‐turns have the same sense of the β‐turn twist that is compatible with the β‐sheet twist. They occur predominantly in two residue β‐hairpins, but the occurrence of Type I' β‐turns is two times higher than Type II' β‐turns. This suggests that Type I' β‐turns may be more stable than Type II' β‐turns, and Type I' β‐turn sequence and structure can be more favorable for protein folding than Type II' β‐turns. Here, we redesigned the native Type II' β‐turn in GFP to Type I' β‐turn, and investigated its effect on protein folding and stability. The Type I' β‐turns were designed based on the statistical analysis of residues in natural Type I' β‐turns. The substitution of the native “GD” sequence of i+1 and i+2 residues with Type I' preferred “(N/D)G” sequence motif increased the folding rate by 50% and slightly improved the thermodynamic stability. Despite the enhancement of in vitro refolding kinetics and stability of the redesigned mutants, they showed poor soluble expression level compared to wild type. To overcome this problem, i and i + 3 residues of the designed Type I' β‐turn were further engineered. The mutation of Thr to Lys at i + 3 could restore the in vivo soluble expression of the Type I' mutant. This study indicates that Type II' β‐turns in natural β‐hairpins can be further optimized by converting the sequence to Type I'. Proteins 2014; 82:2812–2822. © 2014 Wiley Periodicals, Inc.     

Diurnal Variations in Cycling Kinematics

Physiological and biomechanical constraints as well as their fluctuations throughout the day must be considered when studying determinant factors in the preferred pedaling rate of elite cyclists. The aim of this study was to monitor the diurnal variation of spontaneous pedaling rate and movement kinematics over the crank cycle. Twelve male competitive cyclists performed a submaximal exercise on a cycle ergometer for 15 min at 50% of their W_max. Two test sessions were performed at 06:00 and 18:00 h on two separate days to assess diurnal variation in the study variables. For each test session, the exercise bout was divided into three equivalent 5‐min periods during which subjects were requested to use different pedal rates (spontaneous cadence, 70 and 90 rev min^?1). Pedal rate and kinematics data (instantaneous pedal velocity and angle of the ankle) were collected. The results show a higher spontaneous pedal rate in the late afternoon than in the early morning (p < 0.001). For a given pedal rate condition, there was a less variation in pedal velocity during a crank cycle in the morning than in the late afternoon. Moreover, diurnal variations were observed in ankle mobility across the crank cycle, the mean plantar flexion observed throughout the crank cycle being greater in the 18:00 h test session (p < 0.001). These results suggest that muscular activation patterns during a cyclical movement could be under the influence of circadian fluctuations.     

Tight turns in stick insects

We investigated insects Carausius morosus walking whilst hanging upside down along a narrow 3 mm horizontal beam. At the end of the beam, the animal takes a 180° turn. This is a difficult situation because substrate area is small and moves relative to the body during the turn. We investigated how leg movements are organised during this turn. A non-contact of either front leg appears to indicate the end of the beam. However, a turn can only begin if the hind legs stand in an appropriate position relative to each other; the outer hind leg must not be placed posterior to the inner hind leg. When starting the turn, both front legs are lifted and usually held in a relatively stable position and then the inner middle leg performs a swing-and-search movement: The leg begins a swing, which is continued by a searching movement to the side and to the rear, and eventually grasps the beam. At the same time the body is turned usually being supported by the outer middle leg and both hind legs. Then front legs followed by the outer middle leg reach the beam. A scheme describing the turns based on a few simple behavioural elements is proposed.     

Increasing protein stability by improving beta‐turns

Our goal was to gain a better understanding of how protein stability can be increased by improving β‐turns. We studied 22 β‐turns in nine proteins with 66–370 residues by replacing other residues with proline and glycine and measuring the stability. These two residues are statistically preferred in some β‐turn positions. We studied: Cold shock protein B (CspB), Histidine‐containing phosphocarrier protein, Ubiquitin, Ribonucleases Sa2, Sa3, T1, and HI, Tryptophan synthetase α‐subunit, and Maltose binding protein. Of the 15 single proline mutations, 11 increased stability (Average = 0.8 ± 0.3; Range = 0.3–1.5 kcal/mol), and the stabilizing effect of double proline mutants was additive. On the basis of this and our previous work, we conclude that proteins can generally be stabilized by replacing nonproline residues with proline residues at the i + 1 position of Type I and II β‐turns and at the i position in Type II β‐turns. Other turn positions can sometimes be used if the φ angle is near ?60° for the residue replaced. It is important that the side chain of the residue replaced is less than 50% buried. Identical substitutions in β‐turns in related proteins give similar results. Proline substitutions increase stability mainly by decreasing the entropy of the denatured state. In contrast, the large, diverse group of proteins considered here had almost no residues in β‐turns that could be replaced by Gly to increase protein stability. Improving β‐turns by substituting Pro residues is a generally useful way of increasing protein stability. Proteins 2009. © 2009 Wiley‐Liss, Inc.     

Movements and burrowing activity in the Antarctic bivalve molluscs<Emphasis Type="Italic"> Laternula elliptica</Emphasis> and<Emphasis Type="Italic"> Yoldia eightsi</Emphasis>

Burrowing was investigated in two Antarctic infaunal bivalve molluscs, Laternula elliptica and Yoldia eightsi, representing amongst the least and most active members of the class Bivalvia in the Southern Ocean. Burrowing rate was expressed via the Burrowing Rate Index (BRI=[³wet weight/time to bury]×10⁴), and produced values of 0.1–10.6 for L. elliptica and 8.8–49.8 for Y. eightsi. These compare with values ranging from 3 to 2,000 for N. American bivalves (mean=222, SE=42.6, n=81), and 200 to 2,200 for Hong Kong bivalves (mean=1,140, SE=346, n=6). Values for the Antarctic species are, therefore, low compared to warmer-water bivalves, and the values below 1 for large L. elliptica are the lowest on record by around ×5. There is no compensation of burrowing activity for low temperature in these species. The relative BRI values for L. elliptica and Y. eightsi reflect the differences in their mode of life, with the former being large, sedentary and suspension-feeding, and the latter being smaller, mobile, ploughing through the sediment and feeding on sediment-surface organic matter. Burrowing in L. elliptica is unexpected, because other members of the Laternulidae do not burrow. This ability is most probably a response to the regular disturbance of sediments in Antarctica by ice, and the strong selective advantage to being able to resume a protected position after disturbance. The burrowing cycle in L. elliptica is composed of three main phases: (1) foot extension and sediment penetration; (2) foot dilation to form an anchor; (3) the drawing down of the shell by contraction of the pedal retractor muscles. Burrowing in Y. eightsi also has three phases: (1) foot extension and penetration of the sediment (digging); (2) rocking movements in the upright position; (3) shell anchorage. In excess of burrowing activity, L. elliptica exhibits a unique suite of movements when exposed at the surface. These comprise levering, where the tips of the siphons are pressed against the sediment to lift the shell from the substratum, looping, where the siphons are extended and rotated and, in the process, translocate the whole animal across the sediment, and jetting, where water is ejected forcibly through the siphons while their tips are directed towards the sediment, lifting part or all of the animal clear of the substratum. In the field, following exhumation by icebergs, these activities serve to place the animal in a favourable position for reburial, which is a clear advantage in disturbed polar environments where predatory nemerteans and asteroids are abundant.     

THE LOCOMOTION OF LIMAX : I. TEMPERATURE COEFFICIENT OF PEDAL ACTIVITY.

Pedal progression of the slug Limax maximus was studied to obtain relations between wave velocity on the sole of the foot, wave frequency, the advance due to a single wave, and the velocity of vertically upward creeping. Each of the first three quantities is directly proportional to the simultaneous velocity of progression. Under comparable conditions, that is when work is done at a constant rate, the frequency of pedal waves is influenced by the temperature according to the equation of Arrhenius, with µ = 10,700 (Q ₁₀ for 11° to 21° = 2.1). The velocity of a single wave must have very nearly the same "temperature characteristic," which is found also in another case of nerve net transmission (in Renilla).     

Morphology of the pedal circulatory system of the marine gastropod Busycon contrarium and its role in locomotion (Gastropoda,Buccinacea)

Summary Dissections, injections, and histological preparations of the foot of Busycon contrarium show that the open circulatory system of the snail foot consists of discrete arteries and veins that anastomose throughout the foot in a pattern similar to the closed circulatory system of annelids and vertebrates. As the major arteries penetrate the foot, their diameter decreases and the thickness of their muscular walls diminishes. Near the ventral surface of the foot, the blood is channeled through the venous sinus, a fine network of discrete spaces delimited by the muscle and connective tissue of the sole. Small nuclei at the edges of some of these vessels indicate that they may have an incomplete endothelial lining. From the venous sinus, blood is channeled into two major veins on either side of the foot that join and return the blood to the kidney.While probably important for expanding the sole region and for maintaining turgor once the foot has expanded, the circulatory system is not isolated from the pedal musculature, but rather forms a continuum with the pedal muscles and the surrounding connective tissue. It is therefore unlikely that the blood functions as the mechanical antagonist to the muscular contractions of locomotion. The results indicate that the role of the pedal circulation in Busycon is intermediate between the fluids in the hydrostatic skeleton of many worm-like animals and the muscular-hydrostat of cephalopod arms and tentacles.