Contributions of structure and innervation pattern of the stick insect extensor tibiae muscle to the filter characteristics of the muscle-joint system

1996 ◽  
Vol 199 (10) ◽  
pp. 2185-2198 ◽  
Author(s):  
U Bässler ◽  
W Stein

It is shown that the low-pass filter characteristics of the muscle­joint system of the femur­tibia joint of the stick insect Cuniculina impigra result from co-contraction of the extensor and flexor tibiae muscles. The most distal region of the extensor muscle, which contains a high percentage of slow muscle fibres, is involved in this co-contraction. This conclusion results from the following evidence. (1) Inertial and friction forces do not affect the characteristics of the low-pass filter of the muscle­joint system. (2) There is some co-contraction of the extensor and flexor muscles during sinusoidal stimulation of the femoral chordotonal organ at high stimulus frequencies. Both muscles generate tonic forces that increase with increasing stimulus frequency and also increase with time from the beginning of stimulation until a plateau is reached. (3) For the extensor muscle, this tonic force is produced by its most distal portion only. (4) Electrical stimulation of the common inhibitory motoneurone (CI1) reduces the tonic force generated in this most distal portion of the extensor muscle. Therefore, CI1 stimulation reduces the amplitude of tibial movement in response to sinusoidal stimulation of the femoral chordotonal organ at stimulus frequencies below 0.5 Hz (over this frequency range, the tibial movement amplitude is a function of the force amplitude produced by the whole extensor muscle and there is no co-contraction), but at chordotonal organ stimulus frequencies of 1 Hz and above, CI1 stimulation increases the tibial movement amplitude (in this case, movement amplitude is limited by the degree of co-contraction of the extensor and flexor muscles). With repeated chordotonal organ stimulation at higher stimulus frequencies, the tibial movement amplitude steadily decreases. This must be a consequence of increasing levels of co-contraction of the extensor and flexor muscles, since at low stimulus frequencies (no co-contraction) there is no reduction in movement amplitude during repeated stimulations. It is concluded that co-contraction of the extensor and flexor tibiae muscles prevents instability in the reflex loop in spite of the high gain necessary for the generation of catalepsy. Therefore, the mechanism described can be considered to be an adaptation to the ecological niche occupied by this animal. The contribution of the distal part of the extensor muscle to this system can be switched off by the CI1 during active movements.

1997 ◽  
Vol 200 (9) ◽  
pp. 1383-1402 ◽  
Author(s):  
R Kittmann

An adaptive gain control system of a proprioceptive feedback system, the femur­tibia control loop, is investigated. It enables the joint control loop to work with a high gain but it prevents instability oscillations. In the inactive stick insect, the realisation of specific changes in gain is described for tibial torque, for extensor tibiae muscle force and for motoneuronal activity. In open-loop experiments, sinusoidal stimuli are applied to the femoral chordotonal organ (fCO). Changes in gain that depend on fCO stimulus parameters (such as amplitude, frequency and repetition rate), are investigated. Furthermore, spontaneous and touch-induced changes in gain that resemble the behavioural state of the animal are described. Changes in gain in motoneurones are always realised as changes in the amplitude of modulation of their discharge frequency. Nevertheless, depending on the stimulus situation, two different mechanisms underlie gain changes in motoneurones. (i) Changes in gain can be based on changes in the strength of the sensorimotor pathways that transmit stimulus-modulated information from the fCO to the motoneurones. (ii) Changes in gain can be based on changes in the mean activity of a motoneurone by means of its spike threshold: when, during the modulation, the discharge of a motoneurone is inhibited for part of the stimulus cycle, then a change in mean activity subsequently causes a change in modulation amplitude and gain. A new neuronal mechanism is described that helps to compensate the low-pass filter characteristics of the muscles by an increased activation, especially by a sharper distribution of spikes in the stimulus cycle at high fCO stimulus frequencies.


2006 ◽  
Vol 96 (6) ◽  
pp. 3532-3537 ◽  
Author(s):  
Turgay Akay ◽  
Ansgar Büschges

Reinforcement of movement is an important mechanism by which sensory feedback contributes to motor control for walking. We investigate how sensory signals from movement and load sensors interact in controlling the motor output of the stick insect femur–tibia (FT) joint. In stick insects, flexion signals from the femoral chordotonal organ (fCO) at the FT joint and load signals from the femoral campaniform sensilla (fCS) are known to individually reinforce stance-phase motor output of the FT joint by promoting flexor and inhibiting extensor motoneuron activity. We quantitatively compared the time course of inactivation in extensor tibiae motoneurons in response to selective stimulation of fCS and fCO. Stimulation of either sensor generates extensor activity in a qualitatively similar manner but with a significantly different time course and frequency of occurrence. Inactivation of extensor motoneurons arising from fCS stimulation was more reliable but more than threefold slower compared with the extensor inactivation in response to flexion signals from the fCO. In contrast, simultaneous stimulation of both sense organs produced inactivation in motoneurons with a time course typical for fCO stimulation alone, but with a frequency of occurrence characteristic for fCS stimulation. This increase in probability of occurrence was also accompanied by a delayed reactivation of the extensor motoneurons. Our results indicate for the first time that load signals from the leg affect the processing of movement-related feedback in controlling motor output.


2002 ◽  
Vol 88 (5) ◽  
pp. 2387-2398 ◽  
Author(s):  
Ralph A. DiCaprio ◽  
Harald Wolf ◽  
Ansgar Büschges

Mechanosensory neurons exhibit a wide range of dynamic changes in response, including rapid and slow adaptation. In addition to mechanical factors, electrical processes may also contribute to sensory adaptation. We have investigated adaptation of afferent neurons in the stick insect femoral chordotonal organ (fCO). The fCO contains sensory neurons that respond to position, velocity, and acceleration of the tibia. We describe the influence of random mechanical stimulation of the fCO on the response of fCO afferent neurons. The activity of individual sensory neurons was recorded intracellularly from their axons in the main leg nerve. Most fCO afferents (93%) exhibited a marked decrease in response to trapezoidal stimuli following sustained white noise stimulation (bandwidth = 60 Hz, amplitudes from ±5 to ±30°). Concurrent decreases in the synaptic drive to leg motoneurons and interneurons were also observed. Electrical stimulation of spike activity in individual fCO afferents in the absence of mechanical stimulation also led to a dramatic decrease in response in 15 of 19 afferents tested. This indicated that electrical processes are involved in the regulation of the generator potential or encoding of action potentials and partially responsible for the decreased response of the afferents. Replacing Ca2+ with Ba2+ in the saline surrounding the fCO greatly reduced or blocked the decrease in response elicited by electrically induced activity or mechanical stimulation when compared with control responses. Our results indicate that activity of fCO sensory neurons strongly affects their sensitivity, most likely via Ca2+-dependent processes.


1983 ◽  
Vol 105 (1) ◽  
pp. 127-145 ◽  
Author(s):  
ULRICH BÄSSLER ◽  
U. T. A. WEGNER

The denervated thoracic ventral nerve cord produces a motor output which is similar to that observed in the intact animal during irregular leg movements (seeking movements) or rocking, but not walking. When the nerves to some legs are left intact, and the animal walks on a wheel, the motor output in the protractor and retractor motor neurones of the denervated legs is modulated by the stepping frequency of the walking legs. The modulation is similar to that observed in the motor output to a not actually stepping leg of an intact walking animal. When only the crural nerve of one leg is left intact, stimulation of the trochanteral campaniform sensilli induces protractor and retractor motor output to that leg and the leg behind it. In this case the motor output to the ipsilateral leg is in phase. Stimulation of the femoral chordotonal organ influences activity in motor neurones of the extensor tibiae (FETi and SETi) but not those of the protractor and retractor coxae muscles. In a restrained leg of an intact animal stretching of the femoral chordotonal organ excites FETi and SETi as long as the other legs walk (as in a walking leg) and inhibits FETi and SETi (as in a seeking leg) when the other legs are unable to walk.


1989 ◽  
Vol 144 (1) ◽  
pp. 81-111 ◽  
Author(s):  
ANSGAR BÜSCHGES

The femoral chordotonal organ (ChO) of the right middle leg of the inactive stick insect Carausius morosus was stimulated by applying movements having a ramp-like time course, while recordings were made from local and interganglionic interneurones in the anterior ventral median part of the ganglion. Position, velocity and acceleration of the movements were varied independently and the interneurones were categorized on the basis of their responses to the changes in these parameters. Position-sensitivity was always accompanied by responses to velocity and/or acceleration. Velocity-sensitive responses were excitatory or inhibitory and were produced by elongation or relaxation, or by both. In some cases, velocity-sensitive neurones were also affected by position and acceleration. Acceleration responses were always excitatory and were often found in neurones that showed no effects of velocity or position. It is inferred that sensory input from different receptors in the ChO is processed by single interneurones. No interneurone in the recording region was found to be directly involved in the resistance reflex of the extensor tibiae motoneurones, elicited by stimulation of the ChO.


1990 ◽  
Vol 151 (1) ◽  
pp. 133-160 ◽  
Author(s):  
ANSGAR BÜSCHGES

In the stick insect Carausius morosus (Phasmida) intracellular recordings were made from local nonspiking interneurones involved in the reflex activation of the extensor motoneurones of the femur-tibia joint during ramp-like stimulation of the transducer of this joint, the femoral chordotonal organ (ChO). The nonspiking interneurones in the femur-tibia control loop were characterized by their inputs from the ChO, their output properties onto the extensor motoneurones and their morphology. Eight different morphological and physiological types of nonspiking interneurones are described that are involved in the femur-tibia control loop. The results show that velocity signals from the ChO are the most important movement parameter processed by the nonspiking interneurones. Altering the membrane potential of these interneurones had marked effects on the reflex activation in the extensor motoneurones as the interneurones were able to increase or decrease the response of the participating motoneurones. The processing of information by the nonspiking pathways showed another remarkable aspect: nonspiking interneurones were found to process sensory information from the ChO onto extensor motoneurones in a way that seems not always to support the generation of the visible resistance reflexes in the extensor tibiae motoneurones in response to imposed flexion and extension movements of the joint. The present investigation demonstrated interneuronal pathways in the joint-control loop that show ‘assisting’ characteristics.


1964 ◽  
Vol 47 (3) ◽  
pp. 487-499 ◽  
Author(s):  
Deforest Mellon ◽  
Donald Kennedy

Intracellular recording techniques were used to study electrical activity in bipolar sensory cells associated with crayfish tactile receptors. Several lines of evidence indicate that spikes evoked by natural stimulation of the receptor originate at a dendritic locus. Although overshooting spikes are recorded in the soma in response to both natural and antidromic stimulation receptor potentials are observed only rarely, and, when present, their amplitude is less than 5 mv. Impulses propagating centrifugally into the soma following antidromic stimulation always exhibit an inflection in the rising phase of the spike; however, orthodromic spikes are usually uninflected. Occasionally, orthodromic responses (in the soma) exhibit rather unusual wave forms. Such spikes evoked by natural stimuli are indistinguishable from those elicited electrically in the dendrite, but they do not resemble antidromic impulses. Because the axonal and dendritic boundaries of the soma have a low safety factor for spike transmission, at high frequencies invasion of the soma by dendritic spikes is impeded and often blocked. The soma region can thus act as a low-pass filter. The significance of this self-limiting mechanism for the behavior of the animal is not known; it is suggested, however, that this impediment is a potentially critical one, and may, in other situations, have encouraged the evolution of alternative arrangements.


1997 ◽  
Vol 77 (6) ◽  
pp. 3297-3310 ◽  
Author(s):  
Philip L. Newland ◽  
Yasuhiro Kondoh

Newland, Philip L. and Yasuhiro Kondoh. Dynamics of neurons controlling movements of a locust hind leg. III. Extensor tibiae motor neurons. J. Neurophysiol. 77: 3297–3310, 1997. Imposed movements of the apodeme of the femoral chordotonal organ (FeCO) of the locust hind leg elicit resistance reflexes in extensor and flexor tibiae motor neurons. The synaptic responses of the fast and slow extensor tibiae motor neurons (FETi and SETi, respectively) and the spike responses of SETi were analyzed with the use of the Wiener kernel white noise method to determine their response properties. The first-order Wiener kernels computed from soma recordings were essentially monophasic, or low passed, indicating that the motor neurons were primarily sensitive to the position of the tibia about the femorotibial joint. The responses of both extensor motor neurons had large nonlinear components. The second-order kernels of the synaptic responses of FETi and SETi had large on-diagonal peaks with two small off-diagonal valleys. That of SETi had an additional elongated valley on the diagonal, which was accompanied by two off-diagonal depolarizing peaks at a cutoff frequency of 58 Hz. These second-order components represent a half-wave rectification of the position-sensitive depolarizing response in FETi and SETi, and a delayed inhibitory input to SETi, indicating that both motor neurons were directionally sensitive. Model predictions of the responses of the motor neurons showed that the first-order (linear) characterization poorly predicted the actual responses of FETi and SETi to FeCO stimulation, whereas the addition of the second-order (nonlinear) term markedly improved the performance of the model. Simultaneous recordings from the soma and a neuropilar process of FETi showed that its synaptic responses to FeCO stimulation were phase delayed by about −30° at 20 Hz, and reduced in amplitude by 30–40% when recorded in the soma. Similar configurations of the first and second-order kernels indicated that the primary process of FETi acted as a low-pass filter. Cross-correlation between a white noise stimulus and a unitized spike discharge of SETi again produced well-defined first- and second-order kernels that showed that the SETi spike response was also dependent on positional inputs. An elongated negative valley on the diagonal, characteristic of the second-order kernel of the synaptic response in SETi, was absent in the kernel from the spike component, suggesting that information is lost in the spike production process. The functional significance of these results is discussed in relation to the behavior of the locust.


1980 ◽  
Vol 87 (1) ◽  
pp. 1-22
Author(s):  
JEAN-PIERRE VEDEL

1. Skeletal, muscular and neural organization of the two distal joints (J2 and J3) of the antenna of the rock lobster Palinurus vulgaris has been described. 2. Motor innervation (nine motoneurones) of the two distal joints of the antenna has been determined by anatomical and physiological methods. Extensor and flexor muscles of J2 and J3 are each innervated by one specific excitatory tonic motoneurone. One excitatory phasic motoneurone is common to both the J2 and J3 extensor muscles, another to the J2 and J3 flexor muscles. The J3 extensor muscle also receives a specific phasic motoneurone. An accessory extensor muscle which spans J2 and J3 is innervated by one excitatory motoneurone. A common inhibitory motoneurone innervates the two flexor and the two extensor muscles of J2 and J3. 3. Movements of J2 and J3 are sensed by a proprioceptor (chordotonal organ). Reflex patterns involving this proprioceptor have been extensively studied. Sinusoidal extension-flexion movements imposed on the J3 joint induced intra-segmental reflexes (on the J3 muscle innervations) and inter-segmental reflexes (on the J2 muscle innervations) which exclusively involved the tonic excitatory motoneurones and the common inhibitory motoneurone. 4. Resistance reflexes (activation of the muscle stretched by the imposed movements) occurred whatever the excitability level of the animal and involved both flexor and extensor motoneurones. The motoneurones spiked at a higher frequency when the velocity of the imposed movement was increased. The common inhibitor motoneurone was activated during extension movements. 5. In preparations which became ‘more excitable’, assistance reflexes could be induced by joint stimulations which formerly induced resistance reflexes. Sometimes assistance reflexes could be induced by increasing the velocity of the movements imposed on J3. Assistance reflexes mainly involved extensor motoneurones. 6. The role of the tonic, phasic and inhibitory innervations and the functional significance of resistance and assistance reflexes are discussed in relation to the behavioural role of the rock lobster antenna.


1983 ◽  
Vol 105 (1) ◽  
pp. 297-303 ◽  
Author(s):  
J. C. Montgomery

A method is described of obtaining the relationship between electrical stimulation of the abducens nerve and horizontal eye movement in the dogfish. The stump of the VIth nerve was stimulated intracranially in a fish in which the brain had been removed, but in which the circulation remained intact, and the gills were perfused with sea water. Horizontal rotation of the eye was monitored with an opto-electronic movement detector. Eye rotation was linearly related to stimulus frequency in the 0–20 Hz range, and was maximal at frequencies above 40 Hz. Stimulation of the VIth nerve, with a pulse train whose frequency was modulated sinusoidally between 0 and 20 Hz, produced sinusoidal eye movements. The frequency response of the system approximates a first order low pass filter with a characteristic frequency of 0.23 Hz, and an additional phase lag equivalent to a time delay of approximately 50 ms.


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