Rhythmic swimming activity in neurones of the isolated nerve cord of the leech

1976 ◽  
Vol 65 (3) ◽  
pp. 643-668
Author(s):  
W. B. Kristan ◽  
R. L. Calabrese

1. Repeating bursts of motor neurone impulses have been recorded from the nerves of completely isolated nerve cords of the medicinal leech. The salient features of this burst rhythm are similar to those obtained in the semi-intact preparation during swimming. Hence the basic swimming rhythm is generated by a central oscillator. 2. Quantitative comparisons between the impulse patterns obtained from the isolated nerve cord and those obtained from a semi-intact preparation show that the variation in both dorsal to ventral motor neurone phasing and burst duration with swim cycle period differ in these two preparations. 3. The increase of intersegmental delay with period, which is a prominent feature of swimming behaviour of the intact animal, is not seen in either the semi-intact or isolated cord preparations. 4. In the semi-intact preparation, stretching the body wall or depolarizing an inhibitory motor neurone changes the burst duration of excitatory motor neurones in the same segment. In the isolated nerve cord, these manipulations also change the period of the swim cycle in the entire cord. 5. These comparisons suggest that sensory input stabilizes the centrally generated swimming rhythm, determines the phasing of the bursts of impulses from dorsal and ventral motor neurones, and matches the intersegmental delay to the cycle period so as to maintain a constant body shape at all rates of swimming.

1978 ◽  
Vol 75 (1) ◽  
pp. 25-43
Author(s):  
W. O. Friesen ◽  
M. Poon ◽  
G. S. Stent

Four oscillatory interneurones that appear to be the principal components of the central swim oscillator of Hirudo medicinalis have been identified on each side of the segmental ganglia of the ventral nerve cord. During ‘swimming’ episodes of an isolated nerve cord preparation each interneurone undergoes a polarization rhythm that is phase-locked with the impulse burst rhythm of the motor neurones known to drive the swimming movement. Passage of current into any of the interneurones can shift the phase of the swim rhythm. One of the interneurones projects its axon rearward to posterior ganglia and the other three project their axons frontward to anterior ganglia. The oscillatory interneurones are connected both intra- and interganglionically to form a topologically complex intersegmental network of concatenated ring circuits that possess the feature of recurrent cyclic inhibition. Theoretical analysis and electronic analogue models show that the network is inherently oscillatory and can produce both a cycle period and intra- and intersegmental phase relations of its elements that are appropriate for generating the body wave of the swimming movement.


Genetics ◽  
2001 ◽  
Vol 157 (4) ◽  
pp. 1611-1622 ◽  
Author(s):  
Go Shioi ◽  
Michinari Shoji ◽  
Masashi Nakamura ◽  
Takeshi Ishihara ◽  
Isao Katsura ◽  
...  

Abstract Using a pan-neuronal GFP marker, a morphological screen was performed to detect Caenorhabditis elegans larval lethal mutants with severely disorganized major nerve cords. We recovered and characterized 21 mutants that displayed displacement or detachment of the ventral nerve cord from the body wall (Ven: ventral cord abnormal). Six mutations defined three novel genetic loci: ven-1, ven-2, and ven-3. Fifteen mutations proved to be alleles of previously identified muscle attachment/positioning genes, mup-4, mua-1, mua-5, and mua-6. All the mutants also displayed muscle attachment/positioning defects characteristic of mua/mup mutants. The pan-neuronal GFP marker also revealed that mutants of other mua/mup loci, such as mup-1, mup-2, and mua-2, exhibited the Ven defect. The hypodermis, the excretory canal, and the gonad were morphologically abnormal in some of the mutants. The pleiotropic nature of the defects indicates that ven and mua/mup genes are required generally for the maintenance of attachment of tissues to the body wall in C. elegans.


1996 ◽  
Vol 199 (3) ◽  
pp. 613-625
Author(s):  
T Jellema ◽  
W Heitler

The metathoracic fast extensor tibiae (FETi) motor neurone of locusts is unusual amongst insect motor neurones because it makes output connections within the central nervous system as well as in the periphery. It makes excitatory chemical synaptic connections to most if not all of the antagonist flexor tibiae motor neurones. The gain of the FETi-flexor connection is dependent on the peripheral conditions at the time of the FETi spike. This dependency has two aspects. First, sensory input resulting from the extensor muscle contraction can sum with the central excitatory postsynaptic potential (EPSP) to augment its falling phase if the tibia is restrained in the flexed position (initiating a tension-dependent reflex) or is free to extend (initiating a movement-dependent resistance reflex). This effect is thus due to simple postsynaptic summation of the central EPSP with peripheral sensory input. Second, the static tibial position at the time of the FETi spike can change the amplitude of the central EPSP, in the absence of any extensor muscle contraction. The EPSP can be up to 30 % greater in amplitude if FETi spikes with the tibia held flexed rather than extended. The primary sense organ mediating this effect is the femoral chordotonal organ. Evidence is presented suggesting that the mechanism underlying this change in gain may be specifically localised to the FETi-flexor connection, rather than being due to general position-dependent sensory feedback summing with the EPSP. The change in the amplitude of the central EPSP is probably not caused by general postsynaptic summation with tonic sensory input, since a diminution in the amplitude of the central EPSP caused by tibial extension is often accompanied by overall tonic excitation of the flexor motor neurone. Small but significant changes in the peak amplitude of the FETi spike have a positive correlation with changes in the EPSP amplitude, suggesting a likely presynaptic component to the mechanism of gain control. The change in amplitude of the EPSP can alter its effectiveness in producing flexor motor output and, thus, has functional significance. The change serves to augment the effectiveness of the FETi-flexor connection when the tibia is fully flexed, and thus to increase its adaptive advantage during the co-contraction preceding a jump or kick, and to reduce the effectiveness of the connection when the tibia is partially or fully extended, and thus to reduce its potentially maladaptive consequences during voluntary extension movements such as thrusting.


1923 ◽  
Vol 5 (3) ◽  
pp. 327-333 ◽  
Author(s):  
A. R. Moore

1. By the use of preparations of earthworm in which the cutaneous receptors have been anesthetized with a solution of M/8 MgCl2, it is shown that peristalsis can be initiated by tension alone. 2. The receptors of the tension reflex are the intermyal sensory cells of the ventral region of the body wall. 3. It is concluded that Straub obtained the tension reflex because his preparations contained the intermyal receptors; Budington was unable to observe the tension reflex in any preparation from which the intermyal receptors had been removed. 4. Intermyal receptors are the receptors of the following reaction: Passive unilateral tension of the posterior part of an earthworm induces active homolateral tension of the musculature of the anterior segments, and results in the course of progress being brought into line with the enforced orientation of the tail. This reaction is termed the homostrophic reflex. 5. The receptors for the reaction are distributed throughout the entire length of the worm, the effectors are limited to the anterior 15 to 20 segments. The impulse is conducted by the ventral nerve cord. 6. The interaction of the homostrophic reflex and tropisms is considered.


1982 ◽  
Vol 96 (1) ◽  
pp. 161-180 ◽  
Author(s):  
WILLIAM B. KRISTAN

1. Intracellular recordings were made from identified mechanosensory neurones (T and P cells) and longitudinal muscle motor neurones of leeches Hirudo medicinalis and Macrobdella decora while the skin was electrically stimulated to produce local bending responses. 2. The stimulus intensity required to produce local bending was found to activate the mechanosensory neurones at physiological firing rates. For a given stimulation frequency, intracellular activation of the mechanosensory neurones produced the same local bending response as did skin stimulation. Hyperpolarization sufficient to block the propagation of the afferent impulses into the central nervous system eliminated the local bending response to skin stimulation. 3. Stimulating identified longitudinal muscle motor neurones at frequencies observed during the local bending response produced body wall movements similar to those seen in local bending. Hyperpolarization of the motor neurones to block impulse initiation abolished local bending. 4. Mechanosensory neurone to longitudinal muscle motor neurone connexions were demonstrated to be effective and reliable, but polysynaptic for all but the previously documented monosynaptic connexions from mechanosensory neurones onto the L motor neurone (Nicholls & Purves, 1970). 5. It is concluded that the previously identified mechanosensory and motor neurones are exclusively responsible for the local bending response.


1969 ◽  
Vol 50 (1) ◽  
pp. 129-140 ◽  
Author(s):  
M. S. LAVERACK

1. Mechanoreceptors in the body wall of the leech Hirudo are stimulated by deformation of the animal's surface. They respond at all frequencies of stimulation up to about 50-60 Hz. 2. Light flashes, from a microscope lamp or an electronic flash source, are also a potent means of peripheral stimulation. 3. After peripheral stimulation impulses can be recorded in a fast central pathway. This pathway conducts equally well in the posterior to anterior and in the opposite directions. 4. Interference with either the right or left connective linking any two segmental ganglia does not interrupt the rapid conduction of these impulses. 5. Severance of the median connective or Faivre's nerve interrupts conduction. This seems to implicate at least one, and possibly more, of the nerve fibres of this median connective in the rapid transmission of information from the extremities of the body. 6. A slower conducting pathway also exists in the nerve cord.


The nervous system of Caenorhabditis elegans is arranged as a series of fibre bundles which run along internal hypodermal ridges. Most of the sensory integration takes place in a ring of nerve fibres which is wrapped round the pharynx in the head. The body muscles in the head are innervated by motor neurones in this nerve ring while those in the lower part of the body are innervated by a set of motor neurones in a longitudinal fibre bundle which joins the nerve ring, the ventral cord. These motor neurones can be put into five classes on the basis of their morphology and synaptic input. At any one point along the cord only one member from each class has neuromuscular junctions. Members of a given class are arranged in a regular linear sequence in the cord and have non-overlapping fields of motor synaptic activity, the transition between fields of adjacent neurones being sharp and well defined. Members of a given class form gap junctions with neighbouring members of the same class but never to motor neurones of another class. Three of the motor neurone classes receive their synaptic input from a set of interneurones coming from the nerve ring. These interneurones can in turn be grouped into four classes and each of the three motor neurone classes receives its synaptic input from a unique combination of interneurone classes. The possible developmental and functional significance of these observations is discussed.


Development ◽  
1999 ◽  
Vol 126 (20) ◽  
pp. 4489-4498 ◽  
Author(s):  
K.M. Knobel ◽  
E.M. Jorgensen ◽  
M.J. Bastiani

During nervous system development, neurons form synaptic contacts with distant target cells. These connections are formed by the extension of axonal processes along predetermined pathways. Axon outgrowth is directed by growth cones located at the tips of these neuronal processes. Although the behavior of growth cones has been well-characterized in vitro, it is difficult to observe growth cones in vivo. We have observed motor neuron growth cones migrating in living Caenorhabditis elegans larvae using time-lapse confocal microscopy. Specifically, we observed the VD motor neurons extend axons from the ventral to dorsal nerve cord during the L2 stage. The growth cones of these neurons are round and migrate rapidly across the epidermis if they are unobstructed. When they contact axons of the lateral nerve fascicles, growth cones stall and spread out along the fascicle to form anvil-shaped structures. After pausing for a few minutes, they extend lamellipodia beyond the fascicle and resume migration toward the dorsal nerve cord. Growth cones stall again when they contact the body wall muscles. These muscles are tightly attached to the epidermis by narrowly spaced circumferential attachment structures. Stalled growth cones extend fingers dorsally between these hypodermal attachment structures. When a single finger has projected through the body wall muscle quadrant, the growth cone located on the ventral side of the muscle collapses and a new growth cone forms at the dorsal tip of the predominating finger. Thus, we observe that complete growth cone collapse occurs in vivo and not just in culture assays. In contrast to studies indicating that collapse occurs upon contact with repulsive substrata, collapse of the VD growth cones may result from an intrinsic signal that serves to maintain growth cone primacy and conserve cellular material.


1996 ◽  
Vol 76 (5) ◽  
pp. 3178-3195 ◽  
Author(s):  
R. M. Johnston ◽  
R. B. Levine

1. Larval crawling is a bilaterally symmetrical behavior that involves an anterior moving wave of motor activity in the body wall muscles in conjunction with sequential movements of the abdominal prolegs and thoracic legs. The purpose of this study was to determine whether the larval CNS by itself and without phasic sensory feedback was capable of producing patterned activity associated with crawling. To establish the extent of similarity between the output of the isolated nerve cord and crawling, the motor activity produced in isolated larval nerve cords was compared with the motor activity from freely crawling larvae. 2. When exposed to the muscarinic receptor agonist pilocarpine (1.0 mM), isolated larval nerve cords produced long-lasting rhythmic activity in the motor neurons that supply the thoracic leg, abdominal body wall, and abdominal proleg muscles. The rhythmic activity evoked by pilocarpine was abolished reversibly and completely by bath application of the muscarinic-receptor antagonist atropine (0.01 mM) in conjunction with pilocarpine (1.0 mM), suggesting that the response was mediated by muscarinic-like acetylcholine receptors. 3. Similar to crawling in intact animals, the evoked activity in isolated nerve cords involved bilaterally symmetrical motor activity that progressed from the most posterior abdominal segment to the most anterior thoracic segment. The rhythmic activity in thoracic leg, abdominal proleg, and abdominal body wall motor neurons showed intrasegmental and intersegmental cycle-to-cycle coupling. The average cycle period for rhythmic activity in the isolated nerve cord was approximately 2.5 times slower than the cycle period for crawling in intact larvae, but not more variable. 4. Like crawling in intact animals, in isolated nerve cords, bursting activity in the dorsal body wall motor neurons occurred before activity in ventral/lateral body wall motor neurons within an abdominal segment. The evoked bursting activity recorded from the proleg nerve was superimposed on a high level of tonic activity. 5. In isolated nerve cords, bursts of activity in the thoracic leg levator/extensor motor neurons alternated with bursts of activity in the depressor/flexor motor neurons. The burst duration of the levator/extensor activity was brief and remained relatively steady as cycle period increased. The burst duration of the depressor/ flexor activity occupied the majority of an average cycle and increased as cycle period increased. The phase of both levator/extensor motor nerve activity and depressor/flexor motor nerve activity remained relatively stable over the entire range of cycle periods. The timing and patterning of thoracic leg motor neuron activity in isolated nerve cords quantitatively resembled thoracic leg motor activity in freely crawling larvae. 6. The rhythmic motor activity generated by an isolated larval nerve cord resembled a slower version of normal crawling in intact larvae. Because of the many similarities between activity induced in the isolated nerve cord and the muscle activity and movements of thoracic and abdominal segments during crawling, we concluded that central mechanisms can establish the timing and patterning of the crawling motor pattern and that crawling may reflect the output of a central pattern generating network.


1995 ◽  
Vol 198 (2) ◽  
pp. 435-456 ◽  
Author(s):  
A Büschges ◽  
J Schmitz ◽  
U Bässler

Bath application of the muscarinic agonist pilocarpine onto the deafferented stick insect thoracic nerve cord induced long-lasting rhythmic activity in leg motoneurones. Rhythmicity was induced at concentrations as low as 1x10(-4) mol l-1 pilocarpine. The most stable rhythms were reliably elicited at concentrations from 2x10(-3) mol l-1 to 5x10(-3) mol l-1. Rhythmicity could be completely abolished by application of atropine. The rhythm in antagonistic motoneurone pools of the three proximal leg joints, the subcoxal, the coxo-trochanteral (CT) and the femoro-tibial (FT), was strictly alternating. In the subcoxal motoneurones, the rhythm was characterised by the retractor burst duration being correlated with cycle period, whereas the protractor burst duration was almost independent of it. The cycle periods of the rhythms in the subcoxal and CT motoneurone pools were in a similar range for a given preparation. In contrast, the rhythm exhibited by motoneurones supplying the FT joint often had about half the duration. The pilocarpine-induced rhythm was generated independently in each hemiganglion. There was no strict intersegmental coupling, although the protractor motoneurone pools of the three thoracic ganglia tended to be active in phase. There was no stereotyped cycle-to-cycle coupling in the activities of the motoneurone pools of the subcoxal joint, the CT joint and the FT joint in an isolated mesothoracic ganglion. However, three distinct 'spontaneous, recurrent patterns' (SRPs) of motoneuronal activity were reliably generated. Within each pattern, there was strong coupling of the activity of the motoneurone pools. The SRPs resembled the motor output during step-phase transitions in walking: for example, the most often generated SRP (SRP1) was exclusively exhibited coincident with a burst of the fast depressor trochanteris motoneurone. During this burst, there was a switch from subcoxal protractor to retractor activity after a constant latency. The activity of the FT joint extensor motoneurones was strongly decreased during SRP1. SRP1 thus qualitatively resembled the motoneuronal activity during the transition from swing to stance of the middle legs in forward walking. Hence, we refer to SRPs as 'fictive step-phase transitions'. In intact, restrained animals, application of pilocarpine also induced alternating activity in antagonistic motoneurone pools supplying the proximal leg joints. However, there were marked differences from the deafferented preparation. For example, SRP1 was not generated in the latter situation. However, if the ipsilateral main leg nerve was cut, SRP1s reliably occurred. Our results on the rhythmicity in leg motoneurone pools of deafferented preparations demonstrate central coupling in the activity of the leg motoneurones that might be incorporated into the generation of locomotion in vivo.


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