Peripheral control of the gain of a central synaptic connection between antagonistic motor neurones in the locust

1996 ◽  
Vol 199 (3) ◽  
pp. 613-625
Author(s):  
T Jellema ◽  
W Heitler
Keyword(s):  
Muscle Contraction ◽  
Sensory Input ◽  
Sense Organ ◽  
Gain Control ◽  
Motor Neurone ◽  
Extensor Muscle ◽  
Chordotonal Organ ◽  
Excitatory Postsynaptic Potential ◽  
Motor Neurones ◽  
Peripheral Sensory

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.

Effects of Afference Sign Reversal on Motor Activity in Walking Stick Insects (Carausius Morosus)

1981 ◽  
Vol 91 (1) ◽  
pp. 179-193
Author(s):  
D. GRAHAM ◽  
U. BÄSSLER
Keyword(s):  
Sensory Input ◽  
Flexor Tendon ◽  
Sense Organ ◽  
Extensor Tendon ◽  
Motor Output ◽  
Chordotonal Organ ◽  
Internal Oscillation ◽  
Stick Insects ◽  
Peripheral Sensory ◽  
Walking Stick

The apodeme of the femoral chordotonal organ of a middle leg can be moved from its normal position close to the extensor tendon and inserted into a clot cut in the flexor tendon. This inverts the output of the sense organ and produces a ‘wrong’ afference. During walking on a pair of light wheels the operated leg either makes walking steps or is raised and extended in a ‘salute’ posture. The coordination is similar to that for an intact animal if the operated leg walks but changes to the middle leg amputee timing when the operated leg ‘salutes’. The transitions between saluting and the normal walking behaviour of the operated leg can be explained if it is assumed that the animal depends heavily upon the C.O. for determining the tibia position during both walking and the saluting behaviour. Motor output to the levators and depressors of the femur and the protractors and retractors of the coxa shows bursting activity during the salute at the frequency of 3–4 Hz. The depressor bursts are also modulated at a frequency of 1 Hz and produce strong regular depressions of the femur which are co-ordinated with the movements of the other legs. The maintenance of regular depressor contractions during the salute shows that an important part of the motor output to the saluting leg (depressor activity) arises from an internal oscillation or rhythmic command which maintains its co-ordinated activity when the normal peripheral sensory input to the leg it is attempting to operate is absent. Retractor activity wanes during the salute suggesting that propulsion is much more dependent upon peripheral input than is the support musculature (depressors), The creation of a ‘wrong’ afference can be used to map the importance of the operated organ in different sub-units of behaviour.

INTEGRATION OF POSITIVE AND NEGATIVE FEEDBACK LOOPS IN A CRAYFISH MUSCLE

1994 ◽  
Vol 187 (1) ◽  
pp. 305-313
Author(s):  
P Skorupski ◽  
P Vescovi ◽  
B Bush
Keyword(s):  
Negative Feedback ◽  
Positive Feedback ◽  
Motor Neurone ◽  
Chordotonal Organ ◽  
Negative Feedback Loops ◽  
Reflex Pathways ◽  
Motor Neurones ◽  
Receptor Organ ◽  
Group 2

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).

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
Keyword(s):  
Nerve Cord ◽  
Body Shape ◽  
Sensory Input ◽  
Body Wall ◽  
Motor Neurone ◽  
The Body ◽  
Burst Duration ◽  
Cycle Period ◽  
Motor Neurones ◽  
Central Oscillator

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.

scholarly journals Reflex Effects of the Femoral Chordotonal Organ Upon Leg Motor Neurones of the Locust

1982 ◽  
Vol 101 (1) ◽  
pp. 265-285 ◽  
Author(s):  
L.H. FIELD ◽  
M. BURROWS
Keyword(s):  
Sense Organ ◽  
Attachment Site ◽  
Joint Effect ◽  
Chordotonal Organ ◽  
Function Generator ◽  
Metathoracic Ganglion ◽  
Motor Neurones ◽  
Direction Of Movement ◽  
The Central Nervous System ◽  
Stimulation Of

The femoral chordotonal organ (FCO) in a hind leg of a locust monitors the position and movement of the tibia about the femur. It consists of a group of sensory neurones embedded in connective tissue attached distally by two structures: the apodeme, which inserts close to the apodeme of the extensor tibiae muscle, and the flexor strand, which inserts at the base of the apodeme of the flexor tibiae muscle. The action of the apodeme and the flexor strand is reciprocal during movements of the tibia; the apodeme is stretched during flexion of the tibia whilst the flexor strand is relaxed. During extension, the apodeme is relaxed and the flexor strand is stretched. To analyse the reflex effects of this sense organ, all other sense organs of a hind leg were denervated. The apodeme of the FCO was then grasped between forceps, severed from its distal attachment site and its movements controlled by a function generator. The flexor strand remained intact and could be stimulated independently by moving the tibia. The different reflex effects mediated by the separate stimulation of the two components of the FCO were revealed by making intracellular recordings from the somata of leg motor neurones in the metathoracic ganglion. A movement stimulus to either component in a way that corresponded to tibial extension, excited flexor tibiae and inhibited extensor tibiae motor neurones. There was also an inter-joint effect whereby extension excited the depressor tarsi and inhibited the levator tarsi motor neurones. A flexion movement had the converse effects on these motor neurones. The effectiveness of the two components was dependent upon the velocity of the stimulus, the set position of the femoro-tibial joint at which the stimulus was applied, the initial direction of movement, and the activity of other neurones in the central nervous system. Slow motor neurones were depolarized more by low velocities of movement, whereas fast ones were depolarized more by high velocities. The two components produced their greatest effects at the set positions where they were most stretched; thus the apodeme was most effective when the joint was flexed, and the flexor strand when it was extended. Elicited movements of the hind legs or apparently spontaneous changes of excitability enhanced or masked the typical response of the motor neurones to stimulation of the FCO, indicating that the effects of this sense organ are not to be viewed as rigid, but as modifiable in the context of the behaviour of the animal. Note:

Motor Neurone Coupling in Locust Flight

1968 ◽  
Vol 48 (2) ◽  
pp. 389-404
Author(s):  
JOAN JOHNSTON KENDIG
Keyword(s):  
Sensory Input ◽  
Motor Neurone ◽  
Long Term Effects ◽  
Short Term ◽  
Impulse Frequency ◽  
Locust Flight ◽  
Motor Neurones ◽  
Flight Muscles ◽  
Inhibitory Interactions

1. Techniques are described for recording in locust thoracic ganglia from single units identifiable as the motor neurones of specific flight muscles. 2. There are at least two kinds of excitatory interactions among flight-muscle motor neurones. A spike in one motor neurone may be electrically transmitted to another with little delay but much attenuation. Stimulation of a group of motor neurones produces a second, probably chemically transmitted, potential with a latency of 5-6 msec. 3. No short-term inhibitory interactions between motor neurones were observed. 4. Activity in one motor unit of the flight system has long-term effects on the motor neurones of other units, excitatory in some cases and inhibitory in others. 5. Single impulses in sensory neurones have little effect on motor neurones; sustained sensory input to a motor neurone produces a slow depolarization and increase in impulse frequency. 6. Antidromic impulses in one group of motor neurones can entrain orthodromic impulses in another motor neurone. 7. These data are discussed with reference to the hypothesis that the pattern of locust flight--rhythmic synchronous bursts of synergist activity, strict alternation between antagonists--can be produced by motor neurone interactions alone.

Central connections of sensory neurones from a hair plate proprioceptor in the thoraco-coxal joint of the locust.

1995 ◽  
Vol 198 (7) ◽  
pp. 1589-1601 ◽  
Author(s):  
F Kuenzi ◽  
M Burrows
Keyword(s):  
Stance Phase ◽  
Swing Phase ◽  
Motor Neurone ◽  
Sensory Neurone ◽  
Sensory Neurones ◽  
Selective Stimulation ◽  
Basal Joint ◽  
Motor Neurones ◽  
Individual Motor ◽  
Stimulation Of

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.

The Physiology and Morphology of Median Nerve Motor Neurones in the Thoracic Ganglia of the Locust

1982 ◽  
Vol 96 (1) ◽  
pp. 325-341
Author(s):  
MALCOLM BURROWS
Keyword(s):  
Synaptic Input ◽  
Motor Neurone ◽  
Abdominal Muscles ◽  
Thoracic Ganglia ◽  
Synaptic Potentials ◽  
Metathoracic Ganglion ◽  
Inspiratory Motor ◽  
Motor Neurones ◽  
Chemical Synapses ◽  
The Right

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.

scholarly journals A bipedal mammalian model for spinal cord injury research: The tammar wallaby

F1000Research ◽  
2017 ◽  
Vol 6 ◽  
pp. 921 ◽  
Author(s):  
Norman R. Saunders ◽  
Katarzyna M. Dziegielewska ◽  
Sophie C. Whish ◽  
Lyn A. Hinds ◽  
Benjamin J. Wheaton ◽  
...  
Keyword(s):  
Spinal Cord ◽  
Spinal Cord Injury ◽  
Sensory Input ◽  
Tammar Wallaby ◽  
Muscle Coordination ◽  
Thoracic Spinal Cord ◽  
Hind Limbs ◽  
Cord Injury ◽  
Trunk Posture ◽  
Peripheral Sensory

Background: Most animal studies of spinal cord injury are conducted in quadrupeds, usually rodents. It is unclear to what extent functional results from such studies can be translated to bipedal species such as humans because bipedal and quadrupedal locomotion involve very different patterns of spinal control of muscle coordination. Bipedalism requires upright trunk stability and coordinated postural muscle control; it has been suggested that peripheral sensory input is less important in humans than quadrupeds for recovery of locomotion following spinal injury. Methods: We used an Australian macropod marsupial, the tammar wallaby (Macropus eugenii), because tammars exhibit an upright trunk posture, human-like alternating hindlimb movement when swimming and bipedal over-ground locomotion. Regulation of their muscle movements is more similar to humans than quadrupeds. At different postnatal (P) days (P7–60) tammars received a complete mid-thoracic spinal cord transection. Morphological repair, as well as functional use of hind limbs, was studied up to the time of their pouch exit. Results: Growth of axons across the lesion restored supraspinal innervation in animals injured up to 3 weeks of age but not in animals injured after 6 weeks of age. At initial pouch exit (P180), the young injured at P7-21 were able to hop on their hind limbs similar to age-matched controls and to swim albeit with a different stroke. Those animals injured at P40-45 appeared to be incapable of normal use of hind limbs even while still in the pouch. Conclusions: Data indicate that the characteristic over-ground locomotion of tammars provides a model in which regrowth of supraspinal connections across the site of injury can be studied in a bipedal animal. Forelimb weight-bearing motion and peripheral sensory input appear not to compensate for lack of hindlimb control, as occurs in quadrupeds. Tammars may be a more appropriate model for studies of therapeutic interventions relevant to humans.

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