Nonspiking Pathways in a Joint-control Loop of the Stick Insect Carausius Morosus

1990 ◽  
Vol 151 (1) ◽  
pp. 133-160 ◽  
Author(s):  
ANSGAR BÜSCHGES
Keyword(s):  
Sensory Information ◽  
Stick Insect ◽  
Chordotonal Organ ◽  
Joint Control ◽  
Control Loop ◽  
Carausius Morosus ◽  
Extensor Motoneurones ◽  
Reflex Activation ◽  
Flexion And Extension ◽  
Stimulation Of

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.

scholarly journals FUNCTIONAL SPECIALIZATION OF THE SCOLOPARIA OF THE FEMORAL CHORDOTONAL ORGAN IN STICK INSECTS

1992 ◽  
Vol 173 (1) ◽  
pp. 91-108 ◽  
Author(s):  
R. Kittmann ◽  
J. Schmitz
Keyword(s):  
Stick Insect ◽  
Sensory Cells ◽  
Chordotonal Organ ◽  
Functional Specialization ◽  
Control Loop ◽  
Sense Organs ◽  
Stick Insects ◽  
Extensor Motoneurones

The femoral chordotonal organ (fCO), one of the largest proprioceptive sense organs in the leg of the stick insect, is important for the control of the femur-tibia joint during standing and walking. It consists of a ventral scoloparium with about 80 sensory cells and a dorsal scoloparium with about 420 sensory cells. The present study examines the function of these scoloparia in the femur-tibia control loop. Both scoloparia were stimulated independently and the responses in the extensor tibiae motoneurones were recorded extra- and intracellularly. The ventral scoloparium, which is the smaller of the two, functions as the transducer of the femur-tibia control loop. Its sensory cells can generate the known resistance reflexes. The dorsal scoloparium serves no function in the femur-tibia control loop and its stimulation elicited no or only minor reactions in the extensor motoneurones. A comparison with other insect leg proprioceptors shows that a morphological subdivision of these organs often indicates a functional specialization.

Processing of Sensory Input from the Femoral Chordotonal Organ by Spiking Interneurones of Stick Insects

1989 ◽  
Vol 144 (1) ◽  
pp. 81-111 ◽  
Author(s):  
ANSGAR BÜSCHGES
Keyword(s):  
Time Course ◽  
Sensory Input ◽  
Stick Insect ◽  
Chordotonal Organ ◽  
Carausius Morosus ◽  
Median Part ◽  
Stick Insects ◽  
Position Sensitivity ◽  
The Right ◽  
Stimulation Of

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.

scholarly journals THE PHYSIOLOGY OF SENSORY CELLS IN THE VENTRAL SCOLOPARIUM OF THE STICK INSECT FEMORAL CHORDOTONAL ORGAN

1994 ◽  
Vol 189 (1) ◽  
pp. 285-292 ◽  
Author(s):  
A Büschges
Keyword(s):  
Sensory Feedback ◽  
Neural Control ◽  
Sensory Information ◽  
Stick Insect ◽  
Sensory Cells ◽  
Chordotonal Organ ◽  
Joint Movement ◽  
Control Loop ◽  
Sensory Neurones ◽  
Control Loops

The leg joints of invertebrates are governed by neural control loops that control their position and velocity during movements (for reviews, see Bassler, 1983, 1993). These neural control loops rely on sensory feedback about the position and velocity of the controlled leg joint. In invertebrates, this sensory feedback is provided by external (e.g. hair fields, hair rows) and/or internal sense organs (e.g. chordotonal organs). The femoral chordotonal organ (fCO) serves as the main proprioceptor in the control loop governing the femur-tibia (FT) joint of the insect leg. The fCO measures the position and movement of this joint (e.g. Bassler, 1965, 1993; Burns, 1974; Usherwood et al. 1968; Zill, 1985). Previous investigations have described the physiology of sensory cells within femoral chordotonal organs (e.g. stick insect, Hofmann et al. 1985; Hofmann and Koch, 1985; locust, Matheson, 1990; Matheson and Field, 1990). Numerous investigations have been undertaken into the central processing of sensory information provided by the fCO to gain an insight into the control of FT joint movement during different behavioural tasks, for example during resistance reflexes in the standing animal (locust, Burrows, 1987, 1988; Burrows et al. 1988; stick insect, Bassler, 1988; Buschges, 1989, 1990; Driesang and Buschges, 1993) or during active movements (stick insect, Bassler, 1988; Bassler and Buschges, 1990). Most previous studies have not, however, taken into account the morphological separation of the fCO into two distinct scoloparia in the legs of some species (stick insect, Fuller and Ernst, 1973; Hofmann et al. 1985; Hofmann and Koch, 1985; locust middle leg, Burns, 1974). It has been inferred that the whole fCO supplies position and velocity information about the FT joint. In contrast, recent studies of leg reflexes have shown that only its smaller scoloparium (Fig. 1A), containing approximately one-sixth of the total number of sensory neurones, provides the sensory information that is used by the FT control loop (locust, Field and Pfluger, 1989; stick insect, Kittmann and Schmitz, 1992). These studies did not show what types of sensory neurones are located in the ventral part of the fCO and thus contribute to the FT control loop. We have therefore investigated the physiology of sensory neurones that are located in the ventral scoloparium of the fCO.

A method for selective stimulation of leg chemoreceptors in whole crustaceans

2021 ◽  
Author(s):  
Paolo Solari ◽  
Giorgia Sollai ◽  
Francesco Palmas ◽  
Andrea Sabatini ◽  
Roberto Crnjar
Keyword(s):  
Procambarus Clarkii ◽  
Sensory Information ◽  
Natural Conditions ◽  
Search Behaviour ◽  
Non Invasive ◽  
Animal Survival ◽  
Sensory Motor ◽  
Reflex Activation ◽  
Motor Outputs ◽  
Stimulation Of

The integration of sensory information with adequate motor outputs is critical for animal survival. Here, we present an innovative technique based on a non-invasive closed-circuit device consisting of a perfusion/stimulation chamber chronically applied on a single leg of the crayfish Procambarus clarkii. Using this technique, we focally stimulated the leg inside the chamber and studied the leg-dependent sensory-motor integration involving other sensory appendages, such as antennules and maxillipeds, which remain unstimulated outside the chamber. Results show that the stimulation of a single leg with chemicals, such as disaccharides, is sufficient to trigger a complex search behaviour involving locomotion coupled with the reflex activation of antennules and maxillipeds. This technique can be easily adapted to other decapods and/or other sensory appendages. Thus, it has opened possibilities for studying sensory-motor integration evoked by leg stimulation in whole aquatic animals under natural conditions to supplement, with a direct approach, current ablation/silencing techniques.

Neural mechanisms of adaptive gain control in a joint control loop: muscle force and motoneuronal activity

1997 ◽  
Vol 200 (9) ◽  
pp. 1383-1402 ◽  
Author(s):  
R Kittmann
Keyword(s):  
Muscle Force ◽  
Gain Control ◽  
Open Loop ◽  
Chordotonal Organ ◽  
Joint Control ◽  
Modulation Amplitude ◽  
Control Loop ◽  
Pass Filter ◽  
Low Pass Filter ◽  
Change In Mean

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.

Load Signals Assist the Generation of Movement-Dependent Reflex Reversal in the Femur–Tibia Joint of Stick Insects

2006 ◽  
Vol 96 (6) ◽  
pp. 3532-3537 ◽  
Author(s):  
Turgay Akay ◽  
Ansgar Büschges
Keyword(s):  
Time Course ◽  
Stick Insect ◽  
Motor Output ◽  
Chordotonal Organ ◽  
Frequency Of Occurrence ◽  
Stick Insects ◽  
Extensor Motoneuron ◽  
Motoneuron Activity ◽  
First Time ◽  
Stimulation Of

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.

Multimodal Convergence of Presynaptic Afferent Inhibition in Insect Proprioceptors

1999 ◽  
Vol 82 (1) ◽  
pp. 512-514 ◽  
Author(s):  
Wolfgang Stein ◽  
Josef Schmitz
Keyword(s):  
Presynaptic Inhibition ◽  
Sensory Information ◽  
Reversal Potential ◽  
Stick Insect ◽  
Chordotonal Organ ◽  
Primary Afferent Depolarization ◽  
Campaniform Sensilla ◽  
Sense Organs ◽  
Position Sensitive ◽  
Conductance Increase

In the leg motor system of insects, several proprioceptive sense organs provide the CNS with information about posture and movement. Within one sensory organ, presynaptic inhibition shapes the inflow of sensory information to the CNS. We show here that also different proprioceptive sense organs can exert a presynaptic inhibition on each other. The afferents of one leg proprioceptor in the stick insect, either the position-sensitive femoral chordotonal organ or the load-sensitive campaniform sensilla, receive a primary afferent depolarization (PAD) from two other leg proprioceptors, the campaniform sensilla and/or the coxal hairplate. The reversal potential of this PAD is about −59 mV, and the PAD is associated with a conductance increase. The properties of this presynaptic input support the hypothesis that this PAD acts as presynaptic inhibition. The PAD reduces the amplitude of afferent action potentials and thus likely also afferent transmitter release and synaptic efficacy. These findings imply that PAD mechanisms of arthropod proprioceptors might be as complex as in vertebrates.

Role of Proprioceptive Signals From an Insect Femur-Tibia Joint in Patterning Motoneuronal Activity of an Adjacent Leg Joint

1999 ◽  
Vol 81 (4) ◽  
pp. 1856-1865 ◽  
Author(s):  
Dietmar Hess ◽  
Ansgar Büschges
Keyword(s):  
Movement Control ◽  
Stick Insect ◽  
Control Mode ◽  
Chordotonal Organ ◽  
Joint Control ◽  
Muscarinic Agonist ◽  
Behavioral State ◽  
Sensory Signals ◽  
Motoneuron Activity

Role of proprioceptive signals from an insect femur-tibia joint in patterning motoneuronal activity of an adjacent leg joint. Interjoint reflex function of the insect leg contributes to postural control at rest or to movement control during locomotor movements. In the stick insect ( Carausius morosus), we investigated the role that sensory signals from the femoral chordotonal organ (fCO), the transducer of the femur-tibia (FT) joint, play in patterning motoneuronal activity in the adjacent coxa-trochanteral (CT) joint when the joint control networks are in the movement control mode of the active behavioral state. In the active behavioral state, sensory signals from the fCO induced transitions of activity between antagonistic motoneuron pools, i.e., the levator trochanteris and the depressor trochanteris motoneurons. As such, elongation of the fCO, signaling flexion of the FT joint, terminated depressor motoneuron activity and initiated activity in levator motoneurons. Relaxation of the fCO, signaling extension of the FT joint, induced the opposite transition by initiating depressor motoneuron activity and terminating levator motoneuron activity. This interjoint influence of sensory signals from the fCO was independent of the generation of the intrajoint reflex reversal in the FT joint, i.e., the “active reaction,” which is released by elongation signals from the fCO. The generation of these transitions in activity of trochanteral motoneurons barely depended on position or velocity signals from the fCO. This contrasts with the situation in the resting behavioral state when interjoint reflex action markedly depends on actual fCO stimulus parameters, i.e., position and velocity signals. In the active behavioral state, movement signals from the fCO obviously trigger or release centrally generated transitions in motoneuron activity, e.g., by affecting central rhythm generating networks driving trochanteral motoneuron pools. This conclusion was tested by stimulating the fCO in “fictive rhythmic” preparations, activated by the muscarinic agonist pilocarpine in the otherwise isolated and deafferented mesothoracic ganglion. In this situation, sensory signals from the fCO did in fact reset and entrain rhythmic activity in trochanteral motoneurons. The results indicate for the first time that when the stick insect locomotor system is active, sensory signals from the proprioceptor of one leg joint, i.e., the fCO, pattern motor activity in an adjacent leg joint, i.e., the CT joint, by affecting the central rhythm generating network driving the motoneurons of the adjacent joint.

Correlation between muscle structure and filter characteristics of the muscle-joint system in three orthopteran insect species

1996 ◽  
Vol 199 (10) ◽  
pp. 2169-2183 ◽  
Author(s):  
D Bässler ◽  
A Büschges ◽  
S Meditz ◽  
U Bässler
Keyword(s):  
Control System ◽  
Locusta Migratoria ◽  
Open Loop ◽  
Corner Frequency ◽  
Joint Control ◽  
Muscle Fibres ◽  
Joint System ◽  
Carausius Morosus ◽  
Muscle Structure ◽  
Stimulation Of

In orthopteran insects, neural networks for joint control exhibit different characteristics due to behavioural specializations. We investigated whether these differences are generated purely by the neuronal networks, or whether characteristics of the muscles or joint architecture (muscle­joint system) are also involved in these behavioural specializations. We compared the properties of the muscle system moving the femur­tibia joint of the middle and hindleg of three species, Carausius morosus, Cuniculina impigra and Locusta migratoria. Four aspects were analysed for the tibial extensor muscle: (i) the frequency-dependence of motoneuronal activity in response to sinusoidal stimulation of the femoral chordotonal organ (fCO), (ii) the muscle structure, (iii) the innervation pattern of the muscle and (iv) the histochemical properties of the muscle fibres. These aspects were compared with the filter characteristics of the open-loop femur­tibia control system and of the muscle­joint system involved. Whereas in both phasmid species (Carausius morosus and Cuniculina impigra) the motoneuronal activity steadily increases with sinusoidal stimulation of the fCO in the frequency range 0.01­5 Hz, in Locusta migratoria there is a decrease in motoneuronal activity between 0.01 and 0.3 Hz. The muscle structure is basically similar in all three species, as the number of singly innervated muscle fibres (supplied by the fast extensor tibiae motor neurone, FETi) decreases from proximal to distal. The number of triply innervated fibres supplied by the FETi, the slow extensor tibiae (SETi) and the common inhibitor 1 (CI1) is maximal in the middle of the muscle, and the number of dually innervated fibres (supplied by SETi, CI1) increases from proximal to distal. Differences between the locust and the two phasmid species exist in the distal portion of the muscle. The phasmid extensor tibiae muscle contains a morphologically distinct bundle of muscle fibres, not present in the locust, which is mostly dually innervated and which is larger in Cuniculina impigra. Similar results were obtained for the histochemical characterisation of the muscle fibres as revealed from their staining for myofibrillar ATPase activity. The number of histochemically identified fast fibres decreased from proximal to distal, while the number of slow fibres increased. In Carausius morosus and Locusta migratoria, the percentage of slow fibres increased by up to 60­70 % at the distal end, while this increase was to almost 100 % in Cuniculina impigra. Apparently, the larger this distal region and the higher the percentage of slow, dually innervated fibres in it, the lower is the upper corner frequency (the stimulus frequency at which the joint control system produces a movement with 70 % of its maximal response amplitude) of the muscle­joint system. In summary, it appears that the upper corner frequency of the open-loop system in Locusta migratoria (<0.05 Hz) results at least in part from properties of the neuronal joint control network, but in Carausius morosus (0.5­1.0 Hz) and Cuniculina impigra (0.1­0.2 Hz) it results from the upper corner frequency of the muscle­joint system.

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
Keyword(s):  
Distal Portion ◽  
Stick Insect ◽  
Extensor Muscle ◽  
Chordotonal Organ ◽  
Movement Amplitude ◽  
Joint System ◽  
Pass Filter ◽  
Low Pass Filter ◽  
Flexor Muscles ◽  
Stimulation Of

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.

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