Sensory representation of a non‐planar joint in the walking legs of the horseshoe crab,Limulus polyphemus

The patello‐tibial joint in the walking leg of the horseshoe crab can move in the flexion‐extension and promotion‐remotion planes with the freedom of movement of a ball‐and‐socket joint. Its mechanical status is represented by at least seven populations of sensory neurons and these include three different types of proprioceptors; articular membrane receptors, muscle length receptors and muscle tension receptors. A population of each of these types of proprioceptor is associated with structures on the anterior and posterior sides of each leg. In this report we describe the representation of joint movement in both planes by two populations of each type of proprioceptor. In general, the articular membrane receptors were most sensitive to joint flexion; receptors on the anterior side of the leg were biased toward promotion, while those on the posterior were biased toward remotion. Muscle length receptor activity was increased by joint extension and was slightly augmented, in posterior and anterior length receptors, by promotion and remotion, respectively. Length receptor tonic activity was a linear function of muscle length for extension but a non‐linear function of muscle flexion. Tension receptor activity was always highly dependent upon active muscular contraction. Tension receptors associated with posterior tibial flexor showed enhanced activity when the joint was promoted; the activity of those associated with the anterior flexor was enhanced by remotion. The relationships between tension receptor activity and muscle length, and between tension receptor activity and joint position, were also explored. While no single receptor population appears capable of representing the status of this joint in both planes, the output of all of the receptor populations studied appears to provide the CNS with unambiguous information about several parameters representing the status of this complex joint.     

Tension receptors on the apodemes of muscles in the walking legs of the crab,Cancer magister †

1. Mechanoreceptors monitoring tension in working muscles are described in the Decapoda Crustacea.

2. The receptors are associated with apodemes of muscles in the walking leg and are well‐developed in the extensor and flexor of the meropodite (Figures 1, 2).

3. The unbranched dendrites of the receptor neurones innervate the tissues surrounding the insertions of the muscle fibres (Figures 3, 4, 5(A)).

4. The receptors show spontaneous activity with the M‐C joint at resting position and this activity increases when the muscle is stretched by holding the joint at a different position (Figure 7).

5. Isometric tension increase in the muscle recruits sensory units (Figures 8, 10(A)) and increases the activity of units firing (Figure 9).

6. Apodeme receptors may be an entirely distinct input channel from chordotonal organs (Figure 10(B,C)). Joint movements produced by a standard muscle stimulus against increasing loads reveal very different responses (Figure 11).

7. Attempts to determine whether chordotonal organs (CP1, Figures 5(B), 6) monitor isometric muscle tension (Figure 12) suggest possible complexities in their dynamic responses.

8. Abbreviations used in this paper are FASN flexor apodeme sensory nerve, EASN extensor apodeme sensory nerve, BASN bender apodeme sensory nerve, and OASN opener apodeme sensory nerve.     

Unusual tension reception in an insect

Muscle tension receptors in animals monitor the tension generated by muscles. This information is important for the initiation and control of movements and for muscle tone in relation to spatial orientation and gravity. Vertebrates have tendon organs located at the musculo-tendinous junction. The number of muscle fibers attached to one receptor is in the range of 3 to 25. In insects by contrast, only a few examples are known where muscle tension is measured by only single receptors embedded in the muscle. All other muscle activity is monitored by a range of other receptors that detect strains on the cuticle or movements of the joints. Here we describe a set of approximately 200 receptor cells located on a single insect muscle. These receptor cells are associated with ovipositor muscle fibers and were preferentially responsive to muscle tension and not muscle length. Although single receptors may respond differently, their summed response to altered muscle tension characterized them as phasic-tonic type receptors. Experimental activation of muscle receptors in animals producing a basic oviposition motor pattern inhibited homonymous muscle activity without resetting the phase of the rhythm. These results suggest a potential role of tension receptors in regulating ovipositor muscle activity and in particular preventing excessive muscle tension during oviposition. The muscle receptors presented here provide the first example of tension measurement in insects by a few hundred receptor cells associated with a single muscle. Their role in motor control and relation to other tension receptors in vertebrates and invertebrates are discussed.     

Stress detection at the autotomy plane in the decapod crustacea

Summary The preformed autotomy plane of most reptantian decapods circumnavigates the basi-ischiopodite of the pereiopods. Associated with discrete areas of soft or thin cuticle in the proximity of the breakage plane are two connective chordotonal organs. These organs are sensitive to cuticular stress and are referred to as Cuticular Stress Detectors (CSD).The CSD organs respond to pressure applied to the basi-ischiopodite and upon deformation of the discrete areas of soft cuticle onto which the connective tissue strands of the receptors insert. The CSDs exhibit a wide range of unit activity and both receptors have a similar population of unit types. Some units are active only on application or removal of a force applied to the soft cuticle but a large number of phaso-tonic and tonic units respond to a constant pressure applied to the soft cuticle. The majority of the units respond during application of the stimulus (ON units) but a small proportion of the units increase activity on removal of the stimulus (OFF units).Passively produced tension in the anterior levator (autotomiser) muscle and depressor muscle tendons of the C-B joint is a potent stimulus to both receptors.Both receptors respond to movement of the B-I joint of the Nephropsidean walking leg and to movement of the I-M joint in the pereiopods of other reptantian decapods where the basipodite and ischiopodite have fused. The degree of activity is not directly related to the joint position or direction of movement.During autotomy both receptors respond strongly, particularly CSD 1. CSD 2 also shows increased activity but as the receptor is located distal to the breakage plane the receptor nerve is severed when breakage occurs.After autotomy CSD 1 responds normally to deformation of the soft cuticle but manipulation of the breakage plane membrane or of the regenerating limb bud produces low levels of activity even though the stimulus is gross.The possible functional roles of the receptors are discussed.This work was supported in part by European Science Exchange Programme study visit grants awarded to F. Clarac and W. Wales.     

Interlimb interactions during cyclic in-phase and antiphase movements of arms and legs and their dependence on afferent influences

Coordinated arm and leg movements imply neural interactions between the rhythmic generators of the upper and lower extremities. In ten healthy subjects in the lying position, activity of the muscles of the upper and lower extremities was recorded during separate and joint cyclic movements of the arms and legs with different phase relationships between the movements of the limbs and under various conditions of the motor task. Antiphase active arm movements were characterized by higher muscle activity than during the inphase mode. The muscle activity during passive arm movements imposed by the experimentalist was significantly lower than muscle activity during passive arm movements imposed by the other arm. When loading one arm, the muscle activity in the other, passively moving, arm increased independently from the synergy of arm movements. During a motor task implementing joint antiphase movements of both upper and lower extremities, compared to a motor task implementing their joint in-phase movements, we observed a significant increase in activity in the biceps brahii muscle, the tibialis anterior muscle, and the biceps femoris muscle. Loading of arms in these motor tasks has been accompanied by increased activity in some leg muscles. An increase in the frequency of rhythmic movements resulted in a significant growth of the muscle activity of the arms and legs during their cooperative movements with a greater rate of rise in the flexor muscle activity of the arms and legs during joint antiphase movements. Thus, both the spatial organization of movements and the type of afferent influences are significant factors of interlimb interactions, which, in turn, determine the type of neural interconnections that are involved in movement regulation.     

Sense organs in the femur of the stick insect and their relevance to the control of position of the femur-tibia-joint

Summary The sensory innervation pattern is described for the femur of the middle and the hind legs ofCarausius morosus. — In one of the nerves (F121) extracellular recordings show a unit which mirrors the tension of the flexor tibiae muscle (tension receptor). The tension receptor increases the firing rate of the slow extensor tibiae motoneuron. It measures the tension of one or more muscle fibres of the anterior side near the distal end of the muscle. The anatomical basis of this receptor is uncertain. — Another receptor was found on the ventral side of the distal end of the apodeme of the extensor tibiae muscle (apodeme receptor). Recordings from this receptor could not be obtained inCarausius. But inExtatosoma tiaratum it responded to stretching of the nerve. In the natural position it shows a minimum of excitation in the 90°-position of the femur-tibia-joint and an increase in firing rate for both flexion and extension. — Tactile hairs react phasically and have no special sensitivity for one direction. Two receptors at the dorsal side of the femur-tibia-joint (RDAL and RDPL), which are situated in the same position as inSchistocerca hind legs, react phasically to extension movements and fire tonically in the most extended position of the joint. — The influence of these receptors on the position of the femur-tibia-joint is only weak.Supported by Deutsche Forschungsgemeinschaft     

Different oscillators control the circadian rhythm of eclosion and activity inDrosophila

Summary Anatomical studies on the 6 posterior cirri of the barnacle,Balanus hameri, have revealed the presence of a number of bipolar and multipolar sensory neurons in the coxopodite-basipodite-ramal (CBR) region which appear to function as proprioceptors. The cells are associated with two simple strands which terminate on the surface of the flexor muscles or on the hypodermis.Electrophysiological results suggest that the units show uni-directional responses to movements of the limb segments. No position receptors were identified. The difference in receptor activity to imposed and endogenously generated limb movements suggest that many units normally function as muscle tension receptors.     

The tarso-pretarsal chordotonal organ as an element in cockroach walking

Many types of sense organs have been demonstrated to show repetitive discharges during walking that could provide informational cues about leg movements and other parameters of locomotion. We have recorded activities of receptors of the distal (tarsal) segments of the cockroach hindleg in restrained and freely moving animals while they were videotaped. These recordings show peaks of activities at the onset and termination of the stance phase. We have morphologically and physiologically identified a joint angle receptor, the tarso-pretarsal chordotonal organ, that contributes to the discharges seen late in stance, prior to the onset of leg flexion in swing. This sense organ encodes the angle and rate of change of the most distal leg joint and specifically discharges when the claws are disengaged from the substrate. Applied displacements of the claws in restrained preparations elicit reflex activation of the tibial flexor muscle and a crossed extensor reflex in the opposite hindleg. These reflexes could function to insure that leg flexion in swing does not occur until the claws are disengaged and to enhance support by the opposite hindleg. Thus, the regular discharges of the chordotonal organ could assure efficient and coordinated muscle contractions and movements during normal, unperturbed walking. Accepted: 2 January 1997     

Motor unit firing patterns of lower leg muscles during isometric plantar flexion with flexed knee joint position

The ankle flexor and extensor muscles are essential for pedal movements associated with car driving. Neuromuscular activation of lower leg muscles is influenced by the posture during a given task, such as the flexed knee joint angle during car driving. This study aimed to investigate the influence of flexion of the knee joint on recruitment threshold-dependent motor unit activity in lower leg muscles during isometric contraction. Twenty healthy participants performed plantar flexor and dorsiflexor isometric ramp contractions at 30 % of the maximal voluntary contraction (MVC) with extended (0°) and flexed (130°) knee joint angles. High-density surface electromyograms were recorded from medial gastrocnemius (MG), soleus (SOL), and tibialis anterior (TA) muscles and decomposed to extract individual motor units. The torque-dependent change (Δpps /Δ%MVC) of the motor unit activity of MG (recruited at 15 %MVC) and SOL (recruited at 5 %MVC) muscles was higher with a flexed compared with an extended knee joint (p < 0.05). The torque-dependent change of TA MU did not different between the knee joint angles. The motor units within certain limited recruitment thresholds recruited to exert plantar flexion torque can be excited to compensate for the loss of MG muscle torque output with a flexed knee joint.     

Somatosensory unit input to the spinal cord during normal walking

Chronic recording techniques in freely walking cats have been used to sample unitary activity from most large myelinated afferent classes. Cutaneous mechanoreceptors are highly sensitive and generate regular activity patterns predictable from their modalities. Knee joint afferents can fire briskly midrange locomotory movements but appear to be influenced by factors other than joint angle. Golgi tendon organs generate activity consistent with sensitivity to active muscle tension. Muscle spindle afferents do not appear to conform to any single functional pattern for all muscles. It is suggested that degree and rate of stretch are sensed by spindles (possibly under dynamic fusimotor bias) in extensor muscles which normally undergo isometric or lengthening contractions whereas rapidly modulated static fusimotor activity is employed to preserve spindle activity during the rapidly shortening contractions of flexor muscles. Both patterns may be represented in different spindles of bifunctional, biarticular muscles such as rectus femoris and sartorius.     

Vibration signals from the FT joint can induce phase transitions in both directions in motoneuron pools of the stick insect walking system

The influence of vibratory signals from the femoral chordotonal organ fCO on the activities of muscles and motoneurons in the three main leg joints of the stick insect leg, i.e., the thoraco-coxal (TC) joint, the coxa-trochanteral (CT) joint, and the femur-tibia (FT) joint, was investigated when the animal was in the active behavioral state. Vibration stimuli induced a switch in motor activity (phase transition), for example, in the FT joint motor activity switched from flexor tibiae to extensor tibiae or vice versa. Similarly, fCO vibration induced phase transitions in both directions between the motoneuron pools of the TC joint and the CT joint. There was no correlation between the directions of phase transition in different joints. Vibration stimuli presented during simultaneous fCO elongation terminated the reflex reversal motor pattern in the FT joint prematurely by activating extensor and inactivating flexor tibiae motoneurons. In legs with freely moving tibia, fCO vibration promoted phase transitions in tibial movement. Furthermore, ground vibration promoted stance-swing transitions as long as the leg was not close to its anterior extreme position during stepping. Our results provide evidence that, in the active behavioral state of the stick insect, vibration signals can access the rhythm generating or bistable networks of the three main leg joints and can promote phase transitions in motor activity in both directions. The results substantiate earlier findings on the modular structure of the single-leg walking pattern generator and indicate a new mechanism of how sensory influence can contribute to the synchronization of phase transitions in adjacent leg joints independent of the walking direction.     

Relation between the ankle joint angle and the maximum isometric force of the toe flexor muscles

The purpose of this study was to investigate the relationships between the ankle joint angle and maximum isometric force of the toe flexor muscles. Toe flexor strength and electromyography activity of the foot muscles were measured in 12 healthy men at 6 different ankle joint angles with the knee joint at 90 deg in the sitting position. To measure the maximum isometric force of the toe flexor muscles, subjects exerted maximum force on a toe grip dynamometer while the activity levels of the intrinsic and extrinsic plantar muscles were measured. The relation between ankle joint angle and maximum isometric force of the toe flexor muscles was determined, and the isometric force exhibited a peak when the ankle joint was at 70–90 deg on average. From this optimal neutral position, the isometric force gradually decreased and reached its nadir in the plantar flexion position (i.e., 120 deg). The EMG activity of the abductor hallucis (intrinsic plantar muscle) and peroneus longus (extrinsic plantar muscle) did not differ at any ankle joint angles. The results of this study suggest that the force generation of toe flexor muscles is regulated at the ankle joint and that changes in the length-tension relations of the extrinsic plantar muscle could be a reason for the force-generating capacity at the metatarsophalangeal joint when the ankle joint angle is changed.     

Control of flexor motoneuron activity during single leg walking of the stick insect on an electronically controlled treadwheel

In the present study, motoneurons innervating the flexor tibiae muscle of the stick insect (Cuniculina impigra) middle leg were recorded intracellularly while the single leg performed walking-like movements on a treadwheel. Different levels of belt friction (equivalent to a change in load) were used to study the control of activity of flexor motoneurons. During slow leg movements no fast motoneurons were active, but a recruitment of these neurons could be observed during faster leg movements. The firing rate of slow and fast motoneurons increased with incremented belt friction. Also, the force applied to the treadwheel at different frictional levels was adapted closely to the friction of the treadwheel to be overcome. The motoneurons innervating the flexor tibiae were recruited progressively during the stance phase, with the slow motoneurons being active earlier than the fast (half-maximal spike frequency after 10-15% and 50-60% of the stance phase, respectively). The resting membrane potential was more hyperpolarized in fast motoneurons (64.6 +/- 6.5 mV) than in slow motoneurons (-52.9 +/- 5.4 mV). However, the threshold for the initiation of action potentials was not statistically significantly different in both types of flexor motoneurons. Therefore, action potentials were generated in fast motoneurons after a longer period of depolarization and thus later during the stance phase than in slow motoneurons. We show that motoneurons of the flexor tibiae receive substantial common excitatory inputs during the stance phase and that the difference in resting membrane potential between slow and fast motoneurons is likely to play a crucial role in their consecutive recruitment.     

Functional morphology of accessory circulatory organs in the legs of hemiptera

The circulatory organs in the legs of 32 heteropteran and 2 homopteran species were investigated by means of semithin serial sections. In all species, the leg hemocoel is divided by a thin diaphragm into 2 counter-rotating blood sinuses. This diaphragm twists about an angle of 90° immediately distal to the femoral-tibial joint, thereby forming a kind of chamber that is equipped with a valve flap. Apart from the investigated representatives of the Gerromorpha, a muscle is associated with this chamber. Rhythmic contractions of this muscle compress one sinus, thereby forcing hemolymph from the leg into the thorax. Simultaneously, the other sinus widens and hemolymph is sucked from the thorax into the leg. The “leg heart” muscle generally originates from the anteriodorsal wall of the tibia. Its point of insertion varies between different species. In most of the investigated Hemiptera, this muscle inserts at the tendon of the pretarsal flexor muscle. In others, both attachments are located at the tibial cuticle. This peculiarity and other anatomical facts indicate that in the evolution of these organs in the Hemiptera, one portion of the pretarsal flexor muscle has been recruited for the formation of the leg heart.     

Chemosensitivity of crayfish slowly adapting stretch receptors to nicotine

It has recently been demonstrated that slowly adapting stretch receptors (SASRs) in the airways of the dog respond directly to nicotine (Federation Proc. 43: 318, 1984). The purpose of the present experiment was to investigate this chemical effect on an isolated stretch receptor. The crayfish muscle receptor organ was chosen, since crayfish muscle is reported to be insensitive to nicotine or acetylcholine and therefore permits the testing of any direct chemical effect of nicotine on the muscle stretch receptors. The tail was removed and pinned out in a tissue bath, and a stretch receptor organ was surgically isolated. Single-unit SASR extracellular nerve recordings were made while simultaneously measuring tension in the tail. Drugs were prepared in Van Harreveld's solution and administered into the bath kept at 18 degrees C. When resting muscle tension was essentially reduced to zero by cutting both ends of the receptor organ muscle, nicotine (0.07 microM) added to the bath increased receptor activity fourfold. This response was abolished by treatment with hexamethonium (690 microM). In a second group of animals in which the muscle was left intact, nicotine was shown to significantly increase receptor sensitivity to step changes in muscle tension. Once again hexamethonium blocked the response to nicotine. These results demonstrate that the sensitivity of mechanoreceptor can be altered by chemical interaction with nicotinic receptors, which dramatically alter sensory receptor activity.     

Proprioceptor distribution and control of a muscle reflex in the tibia of spider legs

In spiders, retrograde cobalt staining was used to clarify the distribution and detailed innervation of the three types of proprioceptors in the tibio-metatarsal leg joint: internal joint receptors, lyriform slit sense organs, and cuticular spines and hairs. The axons of all these receptors run in just two lateral, ascending nerves, which had previously been associated only with the internal receptors. Each nerve contains several hundred axons ranging in diameter from 0.1 micron to ca. 10 micron. Each slit of the four tibial lyriform organs is innervated by two bipolar sensory neurons. The lateral nerves are entirely sensory and run just beneath the cuticle, a convenient site for electrophysiological recording. We demonstrate simultaneous nerve and muscle recordings from intact spiders; these, in combination with selective sensory ablations, show that a resistance reflex in the flexor metatarsi muscles is elicited by internal joint-receptor units.     

A strand receptor with a central cell body synapses upon spiking local interneurones in the locust

Summary Movements of the femoro-tibial joint of a locust hind leg are monitored by three classes of proprioceptors; a chordotonal organ (Usherwood et al. 1968), multipolar joint receptors (Coillot and Boistel 1968) and a strand receptor innervated by a single afferent with a central cell body (Bräunig 1985). All three classes are excited by imposed or voluntary extension of the tibia. The strand receptor (fe-tiSR) spikes tonically and at a frequency dependent upon the position of the joint whilst the multipolar joint receptors give overlapping information but for a more restricted range. The afferent from the strand receptor makes an excitatory connection with a spiking local interneurone in the midline group of the metathoracic ganglion. The central latency and consistency with which the EPSP follows each sensory spike suggests that the connection is direct. This interneurone also receives convergent inputs from neurones in the chordotonal organ, but not from multipolar joint receptors. Neither the strand receptor nor the multipolar joint receptors apparently synapse upon leg motor neurones that we have tested, in contrast to receptors in the chordotonal organ.     

Effects of movement speed and joint position on knee flexor torque in healthy and post-surgical subjects

Coactivation of knee flexors during knee extension assists in joint stability by exerting an opposing torque to the anterior tibial displacement induced by the quadriceps. This opposing torque is believed to be generated by eccentric muscle actions that stiffen the knee, thereby attenuating strain to joint ligaments, particularly the anterior cruciate ligament (ACL). However, as the lengths of knee muscles vary with changes in joint position, the magnitude of flexor/extensor muscle force coupling may likewise vary, possibly affecting the capacity for active knee stabilization. The purpose of this study was to assess the effect of changes in movement speed and joint position on eccentric/concentric muscle action relationships in the knees of uninjured (UNI) and post-ACL-surgery (INJ) subjects (n = 14). All subjects were tested for maximum eccentric and concentric torque of the contralateral knee flexors and extensor muscles at four isokinetic speeds (15 degrees-60 degrees x s(-1)) and four joint position intervals (20 degrees-60 degrees of knee flexion). Eccentric flexor torque was normalized to the percentage of concentric flexor torque generated at each joint position interval for each speed tested (flexor E-C ratio). In order to estimate the capacity of the knee flexors to resist active knee extension, the eccentric-flexor/concentric-extensor ratios were also computed for each joint position interval and speed (flexor/extensor E-C ratio). The results revealed that eccentric torque surpassed concentric torque by 3%-144% across movement speeds and joint position intervals. The magnitude of the flexor E-C ratio and flexor/extensor E-C increased significantly with speed in both groups of subjects (P < 0.05) and tended to rise with muscle length as the knee was extended; peak values were generated at the most extended joint position (20 degrees-30 degrees). Although torque development patterns were symmetrical between the contralateral limbs in both groups, between-group comparisons revealed significantly higher flexor/extensor E-C ratios for the INJ group compared to the UNI group (P < 0.05), particularly at the fastest speed tested (60 degrees x s(-1)). The results indicate that joint position and movement speed influence the eccentric/concentric relationships of knee flexors and extensors. The INJ subjects appeared to accommodate to surgery by developing the eccentric function of their ACL and normal knee flexors, particularly at higher speeds and at more extended knee joint positions. This may assist in the dynamic stabilization of the knee at positions where ACL grafts have been reported to be most vulnerable to strain.     

Tumor-promoting phorbol esters inhibit the binding of colony-stimulating factor (CSF-1) to murine peritoneal exudate macrophages

L-cell colony-stimulating factor (CSF-1) is a sialoglycoprotein of molecular weight 70,000 daltons that specifically stimulates macrophage colony formation by single committed cells from normal mouse bone marrow and by various classes of more differentiated tissue-derived mononuclear phagocyte colony-forming cells (Stanley et al., 1978). CSF-1 interacts with target cells by direct and specific binding to membrane receptors (CSF-1 receptors) that are present only on cells of the mononuclear phagocyte series and their precursors. We studied the effect of tumor-promoting phorbol esters on the binding of 125I-labeled CSF-1 (125I-CSF-1) to murine peritoneal exudate macrophages (PEM). Biologically active TPA (12-O-tetradecanoyl phorbol-13-acetate) inhibits the binding of 125I-CSF-1 to its receptor on PEM. This inhibition exhibits temperature, time, and concentration dependence. At 37 degrees C, maximum inhibition occurred at about 10(-7) M; inhibition was 50% at 5 X 10(-9) M. At 0 degrees C, the inhibitory activity of TPA is diminished. The action of TPA on PEM is transient. Treated cells recover their 125I-CSF-1-binding activity whether TPA is later removed or not. The process of recovering CSF-1-binding activity is completely blocked by the addition of cycloheximide. When several phorbol derivatives were tested for their inhibitory activities, only biologically active phorbol esters were found to possess such activities. Furthermore, the inhibitory activities of various phorbol esters are proportional to their tumor-promoting activities. Inhibition appears to be due to a reduction in the total number of available CSF-1 receptors rather than a decrease in receptor affinity.     

Proprioceptive reflex interactions with central motor rhythms in the isolated thoracic ganglion of the shore crab

Summary Experiments were carried out on an isolated central nervous system preparation of the shore crab,Carcinus maenas, comprising the fused thoracic ganglion complex with two proprioceptors of one back leg still attached. These, the thoracic-coxal muscle receptor organ and the coxo-basal chordotonal organ, monitor movement and position of the first and second joints, respectively. Motor activity was recorded extracellularly from the central cut ends of the nerves innervating the promotor and remotor muscles of the thoracic-coxal joint, and the levator and depressor muscles of the coxal-basal joint of the same leg. Simultaneous intracellular recordings were made from central processes of individual motoneurones of each muscle.In the absence of any sensory input, the isolated ganglion exhibited rhythmic bursting in the motor nerve roots, with a slow, usually irregular cycle period of 5–50 s.Both receptor organs had both intra-joint and inter-joint effects on the rhythmically active preparation. In most cases the coxo-basal receptor organ had the greater effect.Resistance reflexes initiated by each of the joint proprioceptors were modulated by the rhythmic activity.It may be concluded that, while the isolated thoracic ganglion of the crab is capable of generating rhythmic motor output, proprioceptive feedback from the two basal joints is important in shaping the motor patterns underlying locomotion. Inappropriate reflexes which would impede active movements about these joints are modulated or reversed so as to permit and even reinforce intended locomotory movements.