Synaptic connections between sensory afferents and the common inhibitory motoneuron in crayfish

The sensory inputs to the common inhibitory motoneuron that innervates every leg muscle of the crayfish Procambarus clarkii (Girard) were analyzed by performing intracellular recordings from its neurite within the neuropil of the 5th thoracic ganglion. Two types of sensory inputs involved in locomotion were studied, those from a movement coding proprioceptor (the coxobasal chordotonal organ) and those from sensory neu rons coding contact forces exerted at the tip of the leg on the substrate (the dactyl sensory afferents). Sinusoidal movements applied to the chordotonal organ strand induced a stable biphasic response in the common inhibitory motoneuron that consisted of bursts of spikes during release and stretch of the strand, corresponding to raising and lowering of the leg, respectively. Using ramp movements imposed on the chordotonal strand, we demonstrated that only movement-coding chordotonal afferents produce excitatory post-synaptic potentials in the common inhibitory motoneuron; these connections are monosynaptic. Mechanical or electrical stimulation of the dactyl sensory afferents resulted in an increase in the tonic discharge of the common inhibitory motoneuron through polysynaptic excitatory pathways. These two types of sensory cues reinforce the central command of the common inhibitory motoneuron and contribute to enhancing its activity during leg movements, and thus facilitate the relaxation of tonic muscle fibres during locomotion.Abbreviations ADR anterior distal root - A Lev anterior levator nerve - CB coxo-basipodite joint - CBCO coxo-basal chordotonal organ - CI common inhibitory motoneuron - Dep depressor nerve - DSA dactyl sensory afferents - EPSP excitatory post-synaptic potential - IN interneuron - MN motoneuron - PDR posterior distal root - P Lev posterior levator nerve - Pro promotor nerve - Rem remotor nerve     

Sensorimotor pathways involved in interjoint reflex action of an insect leg

Coordination of motor output between leg joints is crucial for the generation of posture and active movements in multijointed appendages of legged organisms. We investigated in the stick insect the information flow between the middle leg femoral chordotonal organ (fCO), which measures position and movement in the femur-tibia (FT) joint and the motoneuron pools supplying the next proximal leg joint, the coxa-trochanteral (CT) joint. In the inactive animal, elongation of the fCO (by flexing the FT joint) induced a depolarization in eight of nine levator trochanteris motoneurons, with a suprathreshold activation of one to three motoneurons. Motoneurons of the depressor trochanteris muscle were inhibited by fCO elongation. Relaxation signals, i.e., extension of the FT joint, activated both levator and depressor motoneurons; i.e., both antagonistic muscles were coactivated. Monosynaptic as well as polysynaptic pathways contribute to interjoint reflex actions in the stick insect leg. fCO afferents were found to induce short latency EPSPs in levator motoneurons, providing evidence for direct connections between fCO afferents and levator motoneurons. In addition, neuronal pathways via intercalated interneurons were identified that transmit sensory information from the fCO onto levator and/or depressor motoneurons. Finally, we describe two kinds of alterations in interjoint reflex action: (a) With repetitive sensory stimulation, this interjoint reflex action shows a habituation-like decrease in strength. (b) In the actively moving animal, interjoint reflex action in response to fCO elongation, mimicking joint flexion, qualitatively remained the same sign, but with a marked increase in strength, indicating an increased influence of sensory signals from the FT joint onto the adjacent CT joint in the active animal. © 1997 John Wiley & Sons, Inc. J Neurobiol 33: 891–913, 1997     

A reflex behavior mediated by monosynaptic connections between hair afferents and motoneurons in the larval tobacco hornworm,Manduca sexta

In the tobacco hornworm caterpillar, tactile stimulation of sensory hairs located on the tip of a proleg (the planta) evokes ipsilateral or bilateral retraction of the prolegs in that segment. We have used electrophysiological and anatomical methods to investigate the excitatory neural pathways linking the planta hair afferents and the proleg retractor motoneurons (MNs). An important technical innovation was the development of an isolated proleg and desheathed ganglion preparation that permits rapid and reversible ionic manipulations and drug applications. Action potentials (spikes) in individual planta hair afferents produce time-locked excitatory postsynaptic potentials (EPSPs) in ipsilateral proleg MNs which appear to be chemically-mediated and monosynaptic: the EPSPs have a short and constant latency, they follow afferent spikes without failure, they are reversibly abolished in elevated Mg++ saline, and they persist in saline with elevated Mg++ and Ca++ levels. Planta hair afferents also excite ipsilateral MNs by polysynaptic pathways, and their excitation of contralateral proleg MNs is exclusively polysynaptic. Cobalt-staining of the proleg MNs and planta hair afferents shows that the afferents terminate in ventral neuropil, and the proleg MNs have an unusual ventral projection into this region. The ventral projection is on the ipsilateral side, which is consistent with the electrophysiological finding that time-locked EPSPs are found only from ipsilateral hairs. Two factors that contribute to the strong monosynaptic excitation of proleg MNs by ipsilateral planta hairs are the convergence of many hair afferents onto each MN, and the facilitation shown at each afferent-MN synapse. At least 6 afferents converge on each MN, and at short interspike intervals the afferent-evoked EPSPs are enhanced by as much as 400% by homosynaptic facilitation. The EPSP is abolished reversibly by the cholinergic antagonists curare and atropine, suggesting that the neurotransmitter at the synapse is acetylcholine (ACh). This is of particular interest because the ACh receptors of tobacco-feeding Manduca larvae are reported to be less nicotine-sensitive than those of other insects.     

Parallel processing of mechanosensory inputs from the locust ovipositor by intersegmental and local interneurones

The neural pathways underlying the processing of signals from locust (Schistocerca gregaria) ovipositor hairs by different classes of interneurones are investigated.Spikes in the sensory neurones from these hairs evoke chemically-mediated, unitary EPSPs with a short and constant latency in six identified non-giant projection interneurones with cell bodies in the terminal abdominal ganglion. Five of these interneurones receive direct inputs from the valves ipsilateral to their neuropilar branches, whereas the other receives direct inputs from valves on both sides. The sensory neurone from a single hair makes divergent connections with several interneurones and those from different hairs make convergent connections with a given interneurone. The amplitude of the EPSPs evoked depends on the position of a hair along the proximal-distal axis of the valve, with sensory neurones from more distal hairs generating larger amplitude EPSPs.Deflection of hairs also excites three of the four giant projection interneurones through polysynaptic pathways and some local interneurones in the terminal abdominal ganglion through monosynaptic connections. Branches of non-giant projection interneurones, local interneurones, but not those of the giant interneurones, overlap the axon terminals of the ovipositor hair afferents in the terminal abdominal ganglion.     

The mono- and polysynaptic connections between identified neurons in the system of the passive defensive reflex in the edible snail]

A structure of synaptic connections between the identified sensory and giant command neurons of Helix lucorum was studied. It was found that EPSPs arising in the giant neuron as responses to single action potentials generation in sensory neuron consist of several monosynaptic and several polysynaptic components having different magnitude, latencies, and plasticity. The latencies of monosynaptic components are determined by different presynaptic terminals' lengths.     

Monosynaptic excitation of lateral giant fibres by proprioceptive afferents in the crayfish

Giant interneurones mediate a characteristic `tail flip' escape response of the crayfish, Procambarus clarkii, which move it rapidly away from the source of stimulation. We have analysed the synaptic connections of proprioceptive sensory neurones with one type of giant interneurone, the lateral giant. Spikes in sensory neurones innervating an exopodite-endopodite chordotonal organ in the tailfan, which monitors the position and movements of the exopodite, are followed at a short and constant latency by excitatory postsynaptic potentials in a lateral giant interneurone (LG) recorded in the terminal abdominal ganglion. These potentials are unaffected by manipulation of the membrane potential of LG, by bath application of saline with a low calcium concentration, or by one containing the nicotinic antagonist, curare. The potentials evoked in LG by chordotonal organ stimulation are thus thought to be monosynaptic and electrically mediated. This is the first demonstration that LG receives input from sensory receptors other than exteroceptors in the terminal abdominal ganglion. Accepted: 7 April 1997     

Parallel processing of proprioceptive information in the terminal abdominal ganglion of the crayfish

The processing of proprioceptive information from the exopodite-endopodite chordotonal organ in the tailfan of the crayfish Procambarus clarkii (Girard) is described. The chordotonal organ monitors relative movements of the exopodite about the endopodite. Displacement of the chordotonal strand elicits a burst of sensory spikes in root 3 of the terminal ganglion which are followed at a short and constant latency by excitatory postsynaptic potentials in interneurones. The afferents make excitatory monosynaptic connections with spiking and nonspiking local interneurones and intersegmental interneurones. No direct connections with motor neurones were found.Individual afferents make divergent patterns of connection onto different classes of interneurone. In turn, interneurones receive convergent inputs from some, but not all, chordotonal afferents. Ascending and spiking local interneurones receive inputs from afferents with velocity thresholds from 2–400°/s, while nonspiking interneurones receive inputs only from afferents with high velocity thresholds (200–400°/s).The reflex effects of chordotonal organ stimulation upon a number of uropod motor neurones are weak. Repetitive stimulation of the chordotonal organ at 850°/s produces a small reduction in the firing frequency of the reductor motor neurone. Injecting depolarizing current into ascending or non-spiking local interneurones that receive direct chordotonal input produces a similar inhibition.     

An intracellular study of pudendal afferent inputs onto tail motoneurons in the spinalized cat

Postsynaptic potentials, elicited by stimulation of the sensory pudendal (SPud) and superficial perineal nerves (SPeri) on both sides, were recorded from motoneurons innervating tail muscles in the non-anaesthetized and spinalized cat. The stimulation of SPud and SPeri on both sides predominantly produced excitatory postsynaptic potentials (EPSPs) in all kinds of tail motoneurons (70-95%). The inhibitory postsynaptic potentials (IPSPs) were often observed in motoneurons innervating ventral tail muscles (30-33%). The means of averaged central latencies of EPSPs and IPSPs ranged from 4.3 to 7.3 ms, and from 4.6 to 8.4 ms, respectively. The findings suggests that polysynaptic neuronal pathways from pudendal nerve to tail motoneurons produce tonic activities of all tail muscles to raise the tail in micturation, defecation and sexual movements which are induced by stimulation of pudendal nerves.     

Convergence of forelimb afferent actions on C7-Th1 propriospinal neurones bilaterally projecting to sacral segments of the cat spinal cord

Propriospinal neurones located in the cervical enlargement and projecting bilaterally to sacral segments of the spinal cord were investigated electrophysiologically in eleven deeply anaesthetized cats. Excitatory or inhibitory postsynaptic potentials from forelimb afferents were recorded following stimulation of deep radial (DR), superficial radial (SR), median (Med) and ulnar (Uln) nerves. 26 cells were recorded from C7, 22 from C8 and 3 from Th1 segments. The majority of the cells were located in the Rexed's laminae VIII and the medial part of the lamina VII. In 10 cases no afferent input from the forelimb afferents was found. In the remaining neurones effects were evoked mostly from DR (88%) and Med (63%), less often from SR (46%) and Uln (46%). Inhibitory actions were more frequent than excitatory. The highest number of IPSPs was evoked from high threshold flexor reflex afferents (FRA)--all connections were polysynaptic. However, inhibitory actions were often evoked from group I or II muscle afferents (polysynaptic or disynaptic) and, less frequently, from cutaneous afferents (mostly polysynaptic). Di- or polysynaptic IPSPs often accompanied monosynaptic EPSPs from group I or II muscle afferents. Disynaptic or polysynaptic EPSPs from muscle and cutaneous afferents were also recorded in many neurones, while polysynaptic EPSPs from FRA were observed only exceptionally. Various patterns of convergence in individual neuronal subpopulations indicate that they integrate different types of the afferent input from various muscle and cutaneous receptors of the distal forelimb. They transmit this information to motor centers controlling hind limb muscles, forming a part of the system contributing to the process of coordination of movements of fore--and hind--limbs.     

Cholinergic control of the walking network in the crayfish Procambarus clarkii

The output of a neuronal network results generally from both the properties of the component neurons and their synaptic relationships. This article aims at synthesizing various results obtained on the neural network generating locomotion in vitro. In the preparation used, consisting of the last three thoracic ganglia (3–5) along with motor nerves from the 5th leg ganglion to the promotor, remotor, levator and depressor muscles, motor nerve recordings generally revealed only tonic activity in several different motoneurons (MNs). However, rhythmic activity can be obtained by the use of cholinergic agents such as the oxotremorine (Oxo) superfused in the bath (5 × 10⁻⁵ M). If Oxo is pressure-ejected locally in the ganglion, it is possible, depending upon the locus where the drug is applied, to elicit a rhythmic activity restricted to a group of antagonistic MNs. To analyze how cholinergic agents are able to induce such rhythmic activity, very small volumes of drug (50–200 pl), were applied close to the recording electrode. Two types of depolarizing response occurred: a fast large amplitude depolarization (5–20 mV) and a long lasting (10 s to several minutes) low amplitude depolarization (1–3 mV). These responses persisted in the presence of TTX and Co²⁺. The transient initial depolarization is a mixed nicotinic and muscarinic voltage-independent response during which the input resistance decreases by 20 to 40%. In contrast, the long lasting component is voltage-dependent, exclusively muscarinic and associated to a 5–10% increase of input resistance due to the closing of a K⁺ conductance that is active at the resting Vm, and totally suppressed at holding potentials below −70 mV. More generally, K⁺ currents activated at resting potential are responsible for membrane potential stability. The injection of TEA, a blocker of the K⁺ currents, through the recording electrode is able to unmask plateaus above a threshold depolarization. These plateaus are TTX-sensitive but persist in the presence of Ca²⁺ channel blockers. Moreover, in 10% of TEA-filled MNs a spontaneous pacemaker activity was revealed. The organization of the locomotor network is also based upon connections between MNs and INs. Within a MN pool, connections are only loosely established, appearing to consist mainly of electrical coupling. Inhibitory synaptic connections between MNs of opposite pools are mediated by chloride channels. However, the neurotransmitter involved could be either GABA or glutamate. Therefore, at the level of a given joint, a basic rhythm occurs due to both motoneuronal membrane properties and motoneuronal connectivity. However, the coordination of all MNs of an entire leg during fictive walking activity requires the involvement of INs. Based upon these data, we propose a two-stage model of the locomotor network organization: a joint motoneuronal level and a whole leg interneuronal level.     

A dye mixture (Neurobiotin and Alexa 488) reveals extensive dye-coupling among neurons in leeches; physiology confirms the connections

Although the neuronal circuits that generate leech movements have been studied for over 30 years, the list of interneurons (INs) in these circuits remains incomplete. Previous studies showed that some motor neurons (MNs) are electrically coupled to swim-related INs, e.g., rectifying junctions connect IN 28 to MN DI-1 (dorsal inhibitor), so we searched for additional neurons in these behavioral circuits by co-injecting Neurobiotin and Alexa Fluor 488 into segmental MNs DI–1, VI–2, DE–3 and VE–4. The high molecular weight Alexa dye is confined to the injected cell, whereas the smaller Neurobiotin molecules diffuse through gap junctions to reveal electrical coupling. We found that MNs were each dye-coupled to approximately 25 neurons, about half of which are likely to be INs. We also found that (1) dye-coupling was reliably correlated with physiologically confirmed electrical connections, (2) dye-coupling is unidirectional between MNs that are linked by rectifying connections, and (3) there are novel electrical connections between excitatory and inhibitory MNs, e.g. between excitatory MN VE-4 and inhibitory MN DI-1. The INs found in this study provide a pool of novel candidate neurons for future studies of behavioral circuits, including those underlying swimming, crawling, shortening, and bending movements.     

Dominance of local sensory signals over inter-segmental effects in a motor system: experiments

Legged locomotion requires that information local to one leg, and inter-segmental signals coming from the other legs are processed appropriately to establish a coordinated walking pattern. However, very little is known about the relative importance of local and inter-segmental signals when they converge upon the central pattern generators (CPGs) of different leg joints. We investigated this question on the CPG of the middle leg coxa?Ctrochanter (CTr)-joint of the stick insect which is responsible for lifting and lowering the leg. We used a semi-intact preparation with an intact front leg stepping on a treadmill, and simultaneously stimulated load sensors of the middle leg. We found that middle leg load signals induce bursts in the middle leg depressor motoneurons (MNs). The same local load signals could also elicit rhythmic activity in the CPG of the middle leg CTr-joint when the stimulation of middle leg load sensors coincided with front leg stepping. However, the influence of front leg stepping was generally weak such that front leg stepping alone was only rarely accompanied by switching between middle leg levator and depressor MN activity. We therefore conclude that the impact of the local sensory signals on the levator?Cdepressor motor system is stronger than the inter-segmental influence through front leg stepping.     

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.     

Central organization of common inhibitory motoneurons in the locust: role of afferent signals from leg mechanoreceptors

Intracellular recordings of mesothoracic common inhibitory neurons (CI₁, CI₂ and CI₃) were made while tactile hairs of the middle legs of locusts (Locusta migratoria) were mechanically stimulated. Generally the three common inhibitory neurons were excited by stimulation of tactile hairs on the ventral and dorsal surface of femur and tibia. The response pattern of all three CI neurons was similar suggesting that they work as a functional unit. Touching hairs on the dorsal surface of tibia and tarsus in some cases led to inhibition of CIs. The connection between sensory cells of tactile hairs and common inhibitory neurons is polysynaptic.To identify interneurons which mediate afferent signals, simultaneous intracellular recordings from CIs and interneurons were made. Different spiking interneurons were identified which made excitatory or inhibitory monosynaptic connections with CIs. Interneurons with inhibitory input to CIs belonged to the ventral midline group of spiking local interneurons. Behavioral and electrophysiological results indicate that reflex movements of the leg are accompanied by activity of CI neurons. Further it appears that CI activity is inhibited when reflex movements of the leg are actively suppressed by the animal.Abbreviations CI common inhibitor - IN interneuron - LY Lucifer Yellow     

Early postnatal maturation in vestibulospinal pathways involved in neck and forelimb motor control

To assess the organization and functional development of vestibulospinal inputs to cervical motoneurons (MNs), we have used electrophysiology (ventral root and electromyographic [EMG] recording), calcium imaging, trans‐synaptic rabies virus (RV) and conventional retrograde tracing and immunohistochemistry in the neonatal mouse. By stimulating the VIIIth nerve electrically while recording synaptically mediated calcium responses in MNs, we characterized the inputs from the three vestibulospinal tracts, the separate ipsilateral and contralateral medial vestibulospinal tracts (iMVST/cMVST) and the lateral vestibulospinal tract (LVST), to MNs in the medial and lateral motor columns (MMC and LMC) of cervical segments. We found that ipsilateral inputs from the iMVST and LVST were differentially distributed to the MMC and LMC in the different segments, and that all contralateral inputs to MMC and LMC MNs in each segment derive from the cMVST. Using trans‐synaptic RV retrograde tracing as well as pharmacological manipulation of VIIIth nerve‐elicited synaptic responses, we found that a substantial proportion of inputs to both neck and forelimb extensor MNs was mediated monosynaptically, but that polysynaptic inputs were also significant. By recording EMG responses evoked by natural stimulation of the vestibular apparatus, we found that vestibular‐mediated motor output to the neck and forelimb musculature became more robust during the first 10 postnatal days, concurrently with a decrease in the latency of MN discharge evoked by VIIIth nerve electrical stimulation. Together, these results provide insight into the complexity of vestibulospinal connectivity in the cervical spinal cord and a cogent demonstration of the functional maturation that vestibulospinal connections undergo postnatally. © 2016 Wiley Periodicals, Inc. Develop Neurobiol 76: 1061–1077, 2016     

Direct excitation of nonspiking local interneurones by exteroceptors underlies tactile reflexes in the locust

Summary Tactile stimulation of a leg of the locustSchistocerca gregaria can lead to specific reflex movements of that leg. At the same time nonspiking interneurones that are presynaptic to the participating motor neurones are excited or inhibited, suggesting that they are directly involved in these reflexes. The afferent pathways mediating these effects have been examined by recording from individual afferents and nonspiking interneurones.Afferent spikes fromtrichoid orcampaniform sensilla on specific regions of a leg evoke chemically-mediated EPSPs with a constant central latency of about 1.5 ms in certain nonspiking interneurones. The branches of an interneurone and the afferents from which it receives inputs overlap in the neuropil of the ganglion.No afferents have been found to evoke IPSPs directly in the nonspiking interneurones. Instead the inhibition is caused by a population of spiking local interneurones that are themselves excited directly by the afferents, and whose spikes evoke IPSPs in certain nonspiking interneurones.The tactile reflexes can involve movements about one or more joints of the leg, and these coordinated responses are explained by the participation of specific nonspiking interneurones that distribute the sensory inputs to the appropriate sets of motor neurones. For example, when hairs on the dorsal surface of a tarsus are touched, the tarsus is levated. This reflex involves nonspiking local interneurones which are excited directly by these hair afferents and which make direct excitatory connections with the single levator tarsi motor neurone.     

Trehalose metabolism in the blue crab Callinectes sapidus: Isolation of multiple structural cDNA isoforms of trehalose-6-phosphate synthase and their expression in muscles

Adult blue crab Callinectes sapidus exhibit behavioral and ecological dimorphisms: females migrating from the low salinity water to the high salinity area vs. males remaining in the same areas. The flesh basal muscle of the swimming paddle shows a dimorphic color pattern in that levator (Lev) and depressor (Dep) of females tend to be much darker than those of males, while both genders have the same light colored remoter (Rem) and promoter (Pro). The full-length cDNA sequence of four structural isoforms of trehalose-6-phosphate synthase (TPS) is isolated from chela muscles of an adult female, C. sapidus. Two isoforms of the C. sapidus TPS encode functional domains of TPS and trehalose-6-phosphorylase (TPP) in tandem as a fused gene product of Escherichia coli Ost A and Ost B. The other two isoforms contain only a single TPS domain. In both males and females, the darker (Lev + Dep) muscles exhibit greater amounts of trehalose, TPS and trehalase activities than the light colored (Rem + Pro). The fact that adult females show higher levels of trehalase activity in the basal muscles and of glucose in Lev + Dep than those of adult males suggests that there may be a metabolic dimorphism. Moreover, the involvement of trehalose in energy metabolism that was examined under the condition of strenuous swimming activity mimicked in adult females demonstrates the intrinsic trehalose metabolism in Lev + Dep, which subsequently results in hemolymphatic hyperglycemia and hyperlactemia. Our data support that trehalose serves as an additional carbohydrate source of hemolymphatic hyperglycemia in this species. Behavioral and ecological dimorphisms of C. sapidus adults may be supported by a functional dimorphism in energy metabolism.     

Activity-dependent induction of facilitation,depression, and post-tetanic potentiation at an insect central synapse

Summary In Manduca sexta larvae, sensory neurons innervating planta hairs on the tips of the prolegs make monosynaptic excitatory connections with motoneurons innervating proleg retractor muscles. Tactile stimulation of the hairs evokes reflex retraction of the proleg. In this study we examined activity-dependent changes in the amplitude of the excitatory postsynaptic potentials (EPSPs) evoked in a proleg motoneuron by stimulation of individual planta hair sensory neurons. Deflection of a planta hair caused a phasic-tonic response in the sensory neuron, with a mean peak instantaneous firing frequency of >300 Hz, and a tonic firing rate of 10–20 Hz. Direct electrical stimulation was used to activate individual sensory neurons to fire at a range of frequencies including those observed during natural stimulation of the hair. At relatively low firing rates (e.g., 1 Hz), EPSP amplitude was stable indefinitely. At higher instantaneous firing frequencies (>10 Hz), EPSPs were initially facilitated, but continuous stimulation led rapidly to synaptic depression. High-frequency activation of a sensory neuron could also produce post-tetanic potentiation, in which EPSP amplitude remained elevated for several min following a stimulus train. Facilitation, depression, and post-tetanic potentiation all appeared to be presynaptic phenomena. These activity-dependent changes in sensory transmission may contribute to the behavioral plasticity of the proleg withdrawal reflex observed in intact insects.Abbreviations ACh acetylcholine - AChE acetylcholine esterase - CNS central nervous system - EPSP excitatory postsynaptic potential - I _h injected hyperpolarizing current - LTP long-term potentiation - PPR principal planta retractor motoneuron - PTP post-tetanic potentiation - R _in input resistance - V _h hyperpolarized potential - V _m membrane potential - VN ventral nerve - VNA anterior branch of the ventral nerve - V _r resting potential.     

Synaptic potentials of motoneurons of the masseter muscle

Postsynaptic potentials of 93 motoneurons of the masseter muscle evoked by stimulation of different branches of the trigeminal nerve were studied. Stimulation of the most excitable afferent fibers of the motor nerve of the masseter muscle evoked monosynaptic EPSPs with a latent period of 1.2–2.0 msec, changing into action potentials when the strength of stimulation was increased. A further increase in the strength of stimulation produced an antidromic action potential in the motoneurons with a latent period of 0.9 msec. In some motoneurons polysynaptic EPSPs and action potentials developed following stimulation of the motor nerve to the masseter muscle. The ascending phase of synaptic and antidromic action potentials was subdivided into IS and SD components, while the descending phase ended with definite depolarization and hyperpolarization after-potentials. Stimulation of cutaneous branches of the trigeminal nerve, and also of the motor nerve of the antagonist muscle (digastric) evoked IPSPs with a latent period of 2.7–3.5 msec in motoneurons of the masseter muscle. These results indicate the existence of functional connections between motoneurons of the masseter muscle and its proprioceptive afferent fibers, and also with proprioceptive afferent fibers of the antagonist muscle and cutaneous afferent fibers.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 1, No. 3, pp. 262–268, November–December, 1969.     

A sensory map based on velocity threshold of sensory neurones from a chordotonal organ in the tailfan of the crayfish

The central projections of sensory neurones innervating a strand chordotonal organ (CO) in the tailfan of the crayfish, Procambarus clarkii (Girard) have been investigated. The CO monitors movement of the exopodite of the tailfan relative to the endopodite. Intracellular recording and staining were used to characterise the response of the sensory neurones to applied stretches of the chordotonal organ and to reveal their morphology. Two gross morphological types of afferents were found: those that terminated in the terminal (6th) abdominal ganglion on the side ipsilateral to the sensory receptor, and those that had branches in the terminal ganglion and an intersegmental axon that ascended rostrally. Afferents responded to position, velocity and direction of imposed CO displacement. Afferents with particular physiological properties had similar morphologies in different crayfish. Irrespective of their directional responses, afferents had central projection areas dependent upon their velocity thresholds. Many afferents responded only during movement of the CO, and those with the lowest velocity thresholds (2°/s) had branches that projected most anteriorly, while those with progressively higher velocity thresholds (up to 200°/s) projected progressively more posteriorly. Afferents that responded to low velocity ramp movements and spiked tonically projected to more posterior areas of the ganglion than those that responded only to movements.Abbreviations A6SCI sixth abdominal sensory commissure I - CO chordotonal organ - DMT dorsal medial tract - G6 sixth abdominal ganglion - LDT lateral dorsal tract - MDT medial dorsal tract - MVT medial ventral tract - R1–4 nerve roots 1–4 - VLT ventral lateral tract - VMT ventral medial tract