Sensory and Motor Neurones Responsible for the Local Bending Response in Leeches

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.

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.


1986 ◽  
Vol 126 (1) ◽  
pp. 445-452
Author(s):  
KEITH T. SILLAR ◽  
ROBERT C. ELSON

Intracellular recordings have been made from walking-leg motor neurones of the crayfish, Pacifastacus leniusculus, in isolated preparations of the thoracic ganglia. Some motor neurones display slow depolarizations that can drive bursts of spikes and resemble ‘plateau’ potentials described in other invertebrate and vertebrate neurones. Evidence is presented which suggests that the potentials are regenerative and endogenous to the motor neurones, and are not the result of feedback from a neural network. These potentials can be induced by synaptic inputs from the non-spiking afferent neurones of the thoracic-coxal muscle receptor organ, a basal limb proprioceptor. Reflex input from this receptor is augmented during the active depolarization of the motor neurone. The results are discussed in terms of the control of rhythmic motor output and the central modulation of reflexes in the crayfish's thoracic nervous system.


1995 ◽  
Vol 198 (7) ◽  
pp. 1589-1601 ◽  
Author(s):  
F Kuenzi ◽  
M Burrows

The hair plate proprioceptors at the thoraco-coxal joint of insect limbs provide information about the movements of the most basal joint of the legs. The ventral coxal hair plate of a middle leg consists of group of 10-15 long hairs (70 microns) and 20-30 short hairs (30 microns). The long hairs are deflected by the trochantin as the leg is swung forward during the swing phase of walking, and their sensory neurones respond phasically during an imposed deflection and tonically if the deflection is maintained. Selective stimulation of the long hairs elicits a resistance reflex that rotates the coxa posteriorly and is similar to that occurring at the transition from the swing to the stance phase of walking. The motor neurones innervating the posterior rotator and adductor coxae muscles are excited, and those to the antagonistic anterior rotator muscle are inhibited. By contrast, selective stimulation of the short hairs leads only to a weak inhibition of the anterior rotator. The excitatory effects of the long hairs are mediated, in part, by direct connections between their sensory neurones and particular motor neurones. A spike in a sensory neurone elicits a short-latency depolarising postsynaptic potential (PSP) in posterior rotator and adductor motor neurones whose amplitude is enhanced by hyperpolarising current injected into the motor neurone. When the calcium in the saline is replaced with magnesium, the amplitude of the PSP is reduced gradually, and not abruptly as would be expected if an interneurone were interposed in the pathway. Several sensory neurones from long hairs converge to excite an individual motor neurone, evoking spikes in some motor neurones. The projections of the sensory neurones overlap with some of the branches of the motor neurones in the lateral association centre of the neuropile. It is suggested that these pathways would limit the extent of the swing phase of walking and contribute to the switch to the stance phase in a negative feedback loop that relieves the excitation of the hairs by rotating the coxa backwards.


1982 ◽  
Vol 96 (1) ◽  
pp. 325-341
Author(s):  
MALCOLM BURROWS

Simultaneous intracellular recordings have been made from the two expiratory, and from the two inspiratory motor neurones which have their axons in the unpaired median nerves of the thoracic ganglia. Each motor neurone has an axon that branches to innervate muscles on the left and on the right side of one segment. The expiratory neurones studied were those in the meso- and meta-thoracic ganglia which innervate spiracular closer muscles. The depolarizing synaptic potentials underlying the spikes during expiration are common to the two closer motor neurones in a particular segment. Similarly, during inspiration when there are usually no spikes, the hyperpolarizing, inhibitory potentials are also common to both motor neurones. The synaptic input to the neurones can be derived from four interneurones; two responsible for the depolarizing potentials during expiration and two for the inhibitory potentials during inspiration. The inspiratory neurones studied were those in the abdominal ganglia fused to the metathoracic ganglion which innervate dorso-ventral abdominal muscles. During inspiration the two motor neurones of one segment spike at a similar and steady frequency. The underlying synaptic input to the two is common. During expiration, when there are usually no spikes, the hyperpolarizing synaptic potentials are also common to both neurones. In addition they match exactly the depolarizing potentials occurring at the same time in the closer motor neurones. The same set of interneurones could be responsible. No evidence has been revealed to indicate that the two closer, or the two inspiratory motor neurones of one segment are directly coupled by electrical or chemical synapses. The morphology of both types of motor neurone is distinct from that of other motor neurones in these ganglia. Both types branch extensively in both the left and in the right areas of the neuropile.


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.


1999 ◽  
Vol 202 (16) ◽  
pp. 2151-2159 ◽  
Author(s):  
T. Friedel

Substratum vibrations elicit a fast startle response in unrestrained quiescent desert locusts (Schistocerca gregaria). The response is graded with stimulus intensity and consists of a small, rapid but conspicuous movement of the legs and body, but it does not result in any positional change of the animal. With stimuli just above threshold, it begins with a fast twitch of the hindlegs generated by movements of the coxa-trochanter and femur-tibia joints. With increasing stimulus intensity, a rapid movement of all legs may follow, resulting in an up-down movement of the whole body. The magnitude of both the hindleg movement and electromyographic recordings from hindleg extensor and flexor tibiae muscles increases with stimulus amplitude and reaches a plateau at vibration accelerations above 20 m s(−)(2) (peak-to-peak). Hindleg extensor and flexor tibiae muscles in unrestrained animals are co-activated with a mean latency of 30 ms. Behavioural thresholds are as low as 0. 47 m s(−)(2) (peak-to-peak) at frequencies below 100 Hz but rise steeply above 200 Hz. The response habituates rapidly, and inter-stimulus intervals of 2 min or more are necessary to evoke maximal reactions. Intracellular recordings in fixed (upside-down) locusts also revealed co-activation of both flexor and extensor motor neurones with latencies of approximately 25 ms. This shows that the neuronal network underlying the startle movement is functional in a restrained preparation and can therefore be studied in great detail at the level of identified neurones.


1994 ◽  
Vol 187 (1) ◽  
pp. 305-313
Author(s):  
P Skorupski ◽  
P Vescovi ◽  
B Bush

It is now well established that in arthropods movement-related feedback may produce positive, as well as negative, feedback reflexes (Bassler, 1976; DiCaprio and Clarac, 1981; Skorupski and Sillar, 1986; Skorupski et al. 1992; Vedel, 1980; Zill, 1985). Usually the same motor neurones are involved in both negative feedback (resistance) reflex responses and positive feedback reflexes. Reflex reversal involves a shift in the pattern of central inputs to a motor neurone, for example from excitation to inhibition. In the crayfish, central modulation of reflexes has been described in some detail for two basal limb proprioceptors, the thoracocoxal muscle receptor organ (TCMRO) and the thoracocoxal chordotonal organ (TCCO) (Skorupski et al. 1992; Skorupski and Bush, 1992). Leg promotor motor neurones are excited by stretch of the TCMRO (which, in vivo, occurs on leg remotion) in a negative feedback reflex, but when this reflex reverses they are inhibited by the same stimulus. Release of the TCCO (which corresponds to leg promotion) excites some, but not all, promotor motor neurones in a positive feedback reflex. There are at least two ways in which the reflex control of a muscle may be modulated in this system. Firstly, inputs to motor neurones may be routed via alternative reflex pathways to produce different reflex outputs. Secondly, the pattern of inputs to a motor pool may be inhomogeneous, so that activation of different subgroups of the motor pool causes different outputs. Different crayfish promotor motor neurones are involved in different reflexes. On this basis, the motor neurones may be classified into at least two subgroups: those that are excited by the TCCO in a positive feedback reflex (group 1) and those that are not (group 2). Do these motor neurone subgroups have different effects on the promotor muscle, or is the output of the two promotor subgroups summed at the neuromuscular level? To address this question we recorded from the promotor nerve and muscle in a semi-intact preparation of the crayfish, Pacifastacus leniusculus. Adult male and female crayfish, 8-11 cm rostrum to tail, were decapitated and the tail, carapace and viscera removed. The sternal artery was cannulated and perfused with oxygenated crayfish saline, as described previously (Sillar and Skorupski, 1986).


1971 ◽  
Vol 54 (2) ◽  
pp. 391-404
Author(s):  
JAMES L. LARIMER ◽  
ALAN C. EGGLESTON ◽  
LEONA M. MASUKAWA ◽  
DONALD KENNEDY

1. High-speed cinematography was used to analyse the abdominal movements of crayfish in response to separate stimulation of medial and lateral giant axons. These films showed that the medial giant fibres command complete abdominal flexions with little flaring of the tail appendages. The lateral giants, in contrast, evoked a relatively weak flexion of the middle abdominal segments, accompanied by promotion of the exopodites of the uropods. 2. An examination of the muscles activated by the two types of giant fibres shows that differences in the connexions between the giant fibres and specific motor neurones can account for the behavioural differences observed. 3. The output of the giant fibres was determined in the sixth abdominal ganglion, where their differential effects are most pronounced. The medial giants activate motor neurones whose axons emerge from root 6 of the sixth ganglion. The lateral giants activate motor neurones whose axons emerge via roots 2 and 3, as well as those emerging via root 6. 4. The larger motor neurones associated with the giant axons in the sixth root of the sixth ganglion have been mapped by Procion Yellow injection, and the terminations of the central giant axons in the sixth ganglion have also been determined. The connexions revealed by this technique are consistent with the physiological findings. 5. The evidence suggests that root 6 of the sixth ganglion is homologous with root 3 of the more anterior ganglia. However, the giant motor neurone of the sixth ganglion has not been identified. 6. The medial and lateral giant fibres, and perhaps other specific ‘command’ interneurones, can thus drive specific ensembles of phasic motor neurones to provide a range of stereotyped quick movements. In this respect the organization of the phasic system of interneurones and motor neurones resembles that in the tonic system.


1980 ◽  
Vol 88 (1) ◽  
pp. 249-258
Author(s):  
CHRISTINE E. PHILLIPS

The anatomical and physiological organization of the locust metathoracic flexor tibiae was examined by a combination of intracellular recording and electron microscopy. Nine excitatory motor neurones, three fast, three intermediate and three slow innervate the muscle; each is uniquely identifiable using a combination of physiological response and soma location. A simple spatial distribution of inputs to the muscle from the individual motor neurones was not found. Individual muscle fibres responded to as many as seven of the motor neurones in various combinations. The muscle fibres are heterogeneous, ranging from slow (tonic) to fast (phasic) in a continuum from predominantly phasic proximally to tonicdistally. This is demonstrated by contraction and relaxation rates to directand indirect stimulation, as well as contraction elicited by action potentials in a single flexor motor neurone. The fast and slow contractile properties of the muscle fibres are matched by appropriate ultrastructures. Such a high degree of complexity of neuromuscular innervation as that found in the metathoracic flexor tibiae has not previously been described for an arthropod muscle.


1980 ◽  
Vol 84 (1) ◽  
pp. 103-118
Author(s):  
M. B. Rheuben ◽  
A. E. Kammer

1. Muscles innervated by an identified set of motor neurones were compared between larval and adult stages. 2. The structure of the larval muscle is typically tonic: long sarcomeres, irregular Z-bands, and 10-12 thin filaments around each thick filament. The structure of the adult muscle is phasic: 3-4 micrometers sarcomeres, regular Z-bands, 6-8 thin filaments around each thick filament, and large mitochondrial volume. 3. The tensions produced by these muscles were correspondingly different. The larval twitch was about 7 times slower and the tetanus/twitch ratio 10 times greater than those of the adult. 4. No structural or physiological differences were observed in the neuromuscular junctions of the two stages. 5. The relatively unchanging functional relationship of a single motor neurone with two different muscle fibre types during two developmental stages is compared with the converse situation in which it has been reported that implantation of a different type of motor nerve into a muscle modifies contractile properties.


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