PubMed Health⌕ Search

Biomedical subjects

P L Newland

Publications and source records attributed to P L Newland.

At least 19 recordsLinked to original sources

Lateral giant fibre activation of exopodite motor neurones in the crayfish tailfan.

The uropods of decapod crustaceans play a major role in the production of thrust during escape swimming. Here we analyse the output connections of a pair of giant interneurones, that mediate and co-ordinate swimming tail flips, on motor neurones that control the exopodite muscles of the uropods. The lateral giants make short latency output connections with phasic uropod motor neurones, including the productor, the lateral abductor and adductor exopodite motor neurones that we have identified both physiologically and anatomically. On the other hand, tonic motor neurones, including the ventral abductor and reductor exopodite motor neurones, receive no input from the lateral giants. We show that there is no simple reciprocal activation of the phasic opener (lateral abductor) and closer (adductor) motor neurones of the exopodite, but instead both phasic motor neurones are activated in parallel with the productor motor neurone during a tail flip. Our results show that the neuronal pathways activating the tonic and phasic motor neurones of the exopodite are apparently independent, with phasic motor neurones being activated during escape movements and tonic motor neurones being activated during slow postural movements.

Abdomen↗

NADPH-diaphorase histochemistry in the terminal abdominal ganglion of the crayfish.

Nitric oxide (NO) has an important modulatory role on the processing of sensory signals in vertebrates and invertebrates. In this investigation we studied the potential sources of NO in the terminal abdominal ganglion of the crayfish, Pacifastacus leniusculus, using NADPH-diaphorase (NADPHd) histochemistry, with NADPHd acting as a marker for NO synthase (NOS). In the terminal ganglion a mean of 27 strongly labelled NADPHd-positive cell bodies were found, and of these 80% [of stained cell bodies] [corrected] occurred in three regions located in antero-lateral, central and posterior parts of the ganglion. Ventral and antero-ventral commissures as well as specific dorsal and ventral areas of the dendritic neuropil showed positive staining. Intense labelling was seen in the ventro-medial tract, and in the connective between the terminal ganglion and the 5th abdominal ganglion. In addition, some motor neurones and neurones with branches in the sensory commissures were NADPHd positive. Our finding that NADPHd-positive cells occur in consistent patterns in the terminal abdominal ganglion implies that NO may have a role in mechanosensory processing in the crayfish.

Animals↗

Distribution of NADPH-diaphorase-positive ascending interneurones in the crayfish terminal abdominal ganglion.

Previous neuropharmacological studies have described the presence of a nitric oxide-cGMP signalling pathway in the crayfish abdominal nervous system. In this study we have analysed the distribution of putative nitric oxide synthase (NOS)-containing ascending interneurones in the crayfish terminal abdominal ganglion using NADPH-diaphorase (NADPHd) histochemistry. Ascending intersegmental interneurones were stained intracellularly using the fluorescent dye Lucifer yellow and the ganglia containing the stained interneurones subsequently processed for NADPHd activity. Fluorescence persisted throughout histochemical processing. These double-labelling experiments showed that 12 of 18 identified ascending interneurones were NADPHd positive. Thus many ascending interneurones that process mechanosensory signals in the terminal ganglion may contain NOS, and are themselves likely sources of NO which is known to modulate their synaptic inputs. Three clear relationships emerged from our analysis between the effects of NO on the synaptic inputs of interneurones, their output properties and their staining for NADPH-diaphorase. First were class 1 interneurones with no local outputs in the terminal ganglion, the NE type interneurones, which had sensory inputs that were enhanced by NO and were NADPHd positive. Second were class 1 interneurones with local and intersegmental output effects that had sensory inputs that were also enhanced by NO but were NADPHd negative. Third were class 2 interneurones with local and intersegmental outputs that had synaptic inputs that were depressed by the action of NO but were NADPHd positive. These results suggest that NO could selectively enhance specific synaptic connections and sensory processing pathways in local circuits.

Abdomen↗

Opposing actions of nitric oxide on synaptic inputs of identified interneurones in the central nervous system of the crayfish.

Little is known of the action of nitric oxide (NO) at the synaptic level on identified interneurones in local circuits that process mechanosensory signals. Here, we examine the action of NO in the terminal abdominal ganglion of the crayfish Pacifastacus leniusculus, where it has modulatory effects on the synaptic inputs of 17 identified ascending interneurones mediated by electrical stimulation of a sensory nerve. To analyse the role of NO in the processing of sensory signals, we bath-applied the NO donor SNAP, the NO scavenger PTIO, the nitric oxide synthase (NOS) inhibitor l-NAME, the NOS substrate l-arginine, a cyclic GMP (cGMP) analogue, 8-Br-cGMP, and the soluble guanylate cyclase (sGC) inhibitor ODQ. The effects of these chemicals on the synaptic inputs of the interneurones could be divided into two distinct classes. The NO donor SNAP enhanced the inputs to one class of interneurone (class 1) and depressed those to another (class 2). Neither the inactive isomer NAP nor degassed SNAP had any effect on the inputs to these same classes of interneurone. The NO scavenger PTIO caused the opposite effects to those of the NO donor SNAP, indicating that endogenous NO may have an action in local circuits. Preventing the synthesis of NO using l-NAME had the opposite effect to that of SNAP on each response class of interneurone. Increasing the synthesis of endogenous NO by applying l-arginine led to effects on both response classes of interneurone similar to those of SNAP. Taken together, these results suggested that NO was the active component in mediating the changes in amplitude of the excitatory postsynaptic potentials. Finally, the effects of 8-Br-cGMP were similar to those of the NO donor, indicating the possible involvement of a NO-sensitive guanylate cyclase. This was confirmed by preventing the synthesis of cGMP by sGC using ODQ, which caused the opposite effects to those of 8-Br-cGMP on the two response classes of interneurone. The results indicate that a NO--cGMP signal transduction pathway, in which NO regulates transmitter release from mechanosensory afferents onto intersegmental ascending interneurones, is probably present in the local circuits of the crayfish.

Animals↗

Parallel somatotopic maps of gustatory and mechanosensory neurons in the central nervous system of an insect.

Relatively little is still known about the sense of taste, or contact chemoreception, compared with other sensory modalities, despite its importance to many aspects of animal behaviour. The central projections of the sensory neurons from bimodal contact chemoreceptors (basiconic sensilla) were compared with those from mechanosensory tactile hairs located on similar regions of the middle leg of the locust. Basiconic sensilla are multiply innervated, containing one mechanosensory and several chemosensory neurons, whereas tactile hairs are innervated by a single mechanosensory neuron. We show that the sensory neurons from tactile hairs form a complete 3-dimensional somatotopic map in the mesothoracic ganglion. Sensory neurons from hairs located on the coxa projected to a region near the midline of the ganglion with neurons from hairs located on progressively more distal parts of the leg arborizing in successively more lateral regions of neuropil. All the neurons from basiconic sensilla, both mechanosensory and chemosensory, also projected in a similar, strictly somatotopic, manner, and the arbors from these neurons overlapped considerably with those from tactile hairs on equivalent parts of the leg to form a continuous region. Thus, the position of a receptor on the leg is preserved in the central nervous system not only for the mechanosensory neurons from both tactile hairs and basiconic sensilla but also for chemosensory neurons. We could observe no anatomical features or small differences in projection region between sensory neurons from individual basiconic sensilla consistent with differences in modality.

Animals↗

Local movements evoked by chemical stimulation of the hind leg in the locust Schistocerca gregaria.

The behavioural responses of desert locusts, Schistocerca gregaria, to solutions of four behaviourally relevant chemicals (sodium chloride, sucrose, nicotine hydrogen tartrate and lysine glutamate) applied as droplets to the hind tarsus were analysed. All responses following within 1 s of chemical stimulation were local leg avoidance reflexes, and the probability of eliciting such a response increased in a dose-dependent manner with increasing concentration for all the chemicals tested. Chemical identity, however, critically determined the concentration threshold at which the different chemicals became an effective stimulus. For example, a 2.5 mmol l(-)(1) concentration of the secondary plant metabolite nicotine hydrogen tartrate (NHT), a potent feeding deterrent to locusts, was sufficient to evoke avoidance responses in 50 % of cases, whilst for the nutrients, sucrose and lysine glutamate, 250-500 mmol l(-)(1) of the chemical was needed to induce avoidance behaviour in 50 % of the locusts. NaCl was of intermediate effectiveness, with a 50 % response rate occurring at a concentration of approximately 50 mmol l(-)(1). The latency to the start of the response following stimulation was negatively correlated with the concentration of NaCl, but for the other chemicals concentration had no effect on latency. The duration of the avoidance behaviour decreased with increasing concentration for NaCl and more weakly for NHT, but not for the other chemicals. Adding a subthreshold concentration of sucrose to 50 mmol l(-)(1) NaCl decreased the incidence of response compared with 50 mmol l(-)(1) NaCl on its own. Experiments with other mixtures combining NaCl, sucrose and NHT indicate that the frequency and dynamics of the responses to chemical mixtures cannot be simply predicted from the responses to their individual constituents.

Animals↗

Processing of gustatory information by spiking local interneurons in the locust.

Despite the importance of gustation, little is known of the central pathways responsible for the processing and coding of different chemical stimuli. Here I have analyzed the responses of a population of spiking local interneurons, with somata at the ventral midline of the metathoracic ganglion, during stimulation of chemo- and mechanoreceptors on the legs of locusts. Volatile acidic stimuli were used to selectively activate the chemosensory neurons. Different members of the population of local interneurons received depolarizing or hyperpolarizing inputs during chemosensory stimulation. Many of the same interneurons that received chemosensory input also received mechanosensory inputs from tactile hairs on the leg, but others received exclusively mechanosensory inputs. Chemosensory inputs occurred with a short and constant latency, typical of monosynaptic connections. The chemosensory receptive fields of the spiking local interneurons mapped the surface of a hind leg so that spatial information relating to the location of a taste receptor was preserved. The amplitude of potentials in interneurons during chemosensory stimulation varied in a graded manner along the long axis of the leg, thus creating gradients in the chemosensory receptive fields of interneurons. Some interneurons were depolarized to a greater extent by chemical stimuli applied to basiconic sensilla on distal parts of the leg, whereas others were depolarized more by chemical stimulation of more proximal sensilla.

Animals↗

Dynamics of neurons controlling movements of a locust hind leg II. Flexor tibiae motor neurons.

Imposed movements of a proprioceptor that monitors the relative position of the tibia about the femur, the femorotibial chordotonal organ (FeCO), evoke resistance reflexes in the motor neurons that control the movements of the tibia of the locust. The response dynamics of one pool of motor neurons, the flexor tibiae motor neurons, which are located in three groups (anterior, lateral, and posterior), have been analyzed by the Wiener kernel method. First- and second-order kernels that represent the linear and nonlinear responses, respectively, were computed by a cross-correlation between the intracellularly recorded synaptic responses in the motor neurons and the white noise stimulus applied to the FeCO, and were used to define the input-output characteristics of the motor neurons. The posterior fast, intermediate, and slow and the anterior fast and intermediate flexor tibiae motor neurons had biphasic first-order kernels with initial negative phases, indicating that they are velocity sensitive. The falling phases of the kernels had distinct shoulders, indicating that the responses of the motor neurons also had delayed low-pass components, i.e., position sensitivity. The anterior slow flexor motor neuron had a monophasic, low-passed, first-order kernel, indicating that it is position sensitive. The linear component of the motor neuron responses, predicted by convolving the first-order kernels with the stimulus signal, strongly resembled the actual response, whereas the second-order nonlinear component was small, particularly at > 10 Hz. The power spectra of the fast motor neurons showed that they had the highest cutoff frequencies (at > 8 Hz), whereas the slow flexor motor neurons had a gradual roll-off at 1 Hz. The intermediate flexor motor neuron had an intermediate cutoff frequency of approximately 2-3 Hz. The linear responses of the flexor motor neurons could be decomposed into low- and high-frequency components. The high-frequency components (> 10 Hz) were velocity dependent and linear, whereas the low-frequency components (< 10 Hz) were position dependent and nonlinear. The nonlinearity was a signal compression (or half-wave rectification). The results show that although the flexor motor neurons receive many common inputs during FeCO stimulation, each individual has specific dynamic response properties. The responses of the motor neurons are fractionated so that a given individual within the pool will respond best to position, whereas others will respond better to velocity. Likewise, some motor neurons respond best at low frequencies, whereas others respond best at higher frequencies of stimulation.

Animals↗

Convergent chemical and electrical synaptic inputs from proprioceptive afferents onto an identified intersegmental interneuron in the crayfish.

Synaptic transmission between proprioceptive afferents from a chordotonal organ in the tailfan of the crayfish and an identified ascending interneuron, interneuron A, in the terminal abdominal ganglion was analyzed. Interneuron A is part of a disynaptic pathway from primary afferent neurons to the lateral giant interneuron involved in producing the characteristic ballistic escape behavior of crayfish. Interneuron A received short and long latency excitatory postsynaptic potentials (EPSPs) from chordotonal afferents. Short latency EPSPs occurred with little central synaptic delay, were unchanged by hyperpolarizing current injection of -2 nA, and remained at a constant amplitude when the nervous system was bathed in saline with a low calcium concentration or saline containing the nicotinic antagonist curare. These EPSPs are thus thought to be mediated by electrical transmission. Longer latency potentials were increased in amplitude by hyperpolarizing current injection, reduced in amplitude when the nervous system was bathed in low-calcium saline, and also reduced by bath application of saline containing curare. These potentials are thus thought to be mediated by chemical transmission. The functional significance of the dual modes of transmission at a key synapse in the escape circuitry is discussed.

Afferent Pathways↗

Dynamics of neurons controlling movements of a locust hind leg. III. Extensor tibiae motor neurons.

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 degrees 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.

Animals↗

Cooperative learning in the clinical setting.

The modern clinical practice setting presents nurses with challenges about which they must think critically and develop increasingly autonomous problem-solving approaches. It is essential to provide nursing students with opportunities to practice critical thinking so that they can develop this crucial skill. Cooperative learning strategies are interactive teaching methods that stimulate students to think critically, communicate effectively with peers, and accept responsibility for learning through group process activities. Group care planning is one such cooperative strategy that also promotes a positive attitude about care planning and sharpens time management skills. Cooperative assessment and care planning foster the development of critical thinking and effective problem resolution, preparing students for patient care problems they will likely encounter in future positions.

Education, Nursing, Continuing↗

The central connections and actions during walking of tibial campaniform sensilla in the locust.

Strain acting on the exoskeleton of insects is monitored by campaniform sensilla. On the tibia of a mesothoracic leg of the locust (Schistocerca gregaria) there are three groups of campaniform sensilla on the proximo-dorsal surface. This study analyses the responses of the afferents from one group, their connections with central neurones and their actions during walking. The afferents of the campaniform sensilla make direct excitatory connections with flexor tibiae motor neurones. They also make direct connections with particular spiking local interneurones that make direct inhibitory output connections with the slow extensor tibiae motor neurone. During walking extension movements of the tibiae during stance produce longitudinal tensile forces on the dorsal tibia that peak during mid stance before returning to zero prior to swing. This decline in tension can activate the campaniform sensilla. In turn this would lead to an inhibition of the extensor tibiae motor neurone and an excitation of the flexor tibiae motor neurones. This, therefore, aids the transition from stance to swing. During turning movements, the tibia is flexed and the dorsal surface is put under compression. This can also activate some of campaniform sensilla whose effect on the flexor motor neurones will reinforce the flexion of the tibia.

Animals↗

Presynaptic inhibition of exteroceptive afferents by proprioceptive afferents in the terminal abdominal ganglion of the crayfish.

1. Exteroceptive hairs that are sensitive to water displacement and touch are distributed over the surface of the tailfan of crayfish. We show that the sensory neurons innervating these hairs receive a primary afferent depolarization (PAD) from sensory neurons innervating a proprioceptor that monitors movements of the endopodite and protopodite of the tailfan. This PAD occurs only during high-velocity movements of the exopodite, which are similar to those that occur during swimming. The effects that the proprioceptor mediate are widespread, so that afferents in four sensory nerve roots of the terminal abdominal ganglion, innervating hairs on the protopodite, exopodite, endopodite, and telson, receive a PAD. The PAD is unlikely to be mediated through monosynaptic pathways because there is no anatomic overlap between the central projections of chordotonal afferents and many of the exteroceptive afferents. The depolarization is associated with a conductance increase and can be increased by the injection of hyperpolarizing current or reversed (approximately 10 mV above resting potential) by injection of depolarizing current. The properties of the presynaptic input are, therefore, consistent with being mediated through chemical synapses. This is supported by the observation in the electron microscope that the exteroceptive afferents receive chemical input synapses. The depolarization is mimicked by gamma-aminobutyric acid and reduced by bath application of picrotoxin or bicuculline, suggesting that it is a depolarizing inhibitory postsynaptic potential. The PAD reduces the amplitude of exteroceptive afferent spikes, an action that is thus likely to reduce transmitter release and the efficacy of synaptic transmission.

Action Potentials↗

Cholinergic synaptic transmission between proprioceptive afferents and a hind leg motor neuron in the locust.

1. The physiology and pharmacology of the connections made by stress-sensitive campaniform sensilla on the proximal dorsal tibia of a locust hind leg with a fast extensor tibiae motor neuron (FETi) have been examined. 2. Mechanical stimulation of a campaniform sensillum on the anterior surface of the tibia elicited bursts of spikes in its afferent and a depolarization in FETi. Each afferent spike was followed at short and constant latency by an excitatory postsynaptic potential (EPSP) in FETi, even at high-frequency stimulation. Electrical stimulation of the extensor tibiae muscle with the tibia fixed elicited an antidromic spike in FETi followed by a compound potential which resulted from the activation of the campaniform sensillum afferent. The connections between the campaniform sensillum and FETi was monosynaptic and chemically mediated. 3. Ionophoresis of ACh into the neuropil depolarized FETi, as did the application of the cholinesterase inhibitor physostigmine. Bath application of the cholinergic agonists, nicotine and muscarine, also depolarized FETi, with nicotine causing a reduction in input resistance, while muscarine caused no detectable change in input resistance. Muscarine and the muscarinic agonist oxotremorine also caused rhythmic depolarizations and bursts of spikes in FETi. These effects were seen in low calcium/high magnesium saline to block synaptic transmission and are therefore due to direct effects on FETi. FETi therefore appears to have both nicotinic and muscarinic receptors. 4. The compound potential in FETi, caused by activation of the campaniform sensillum, was reduced by bath application of nicotinic cholinergic antagonists. However, the muscarinic antagonist scopolamine increased the amplitude of the compound potential.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Dynamics of neurons controlling movements of a locust hind leg: Wiener kernel analysis of the responses of proprioceptive afferents.

1. The response properties of proprioceptive sensory neurons providing input to the local circuits controlling leg movements of the locust have been analysed by the Wiener kernel method. The proprioceptor, the femoral chordotonal organ, encodes the position and movements of the tibia about the femorotibial joint. 2. Intracellular recordings were made from sensory neurons while the apodeme of the organ was moved with a band-limited Gaussian white noise signal with a cutoff frequency of 27, 58, or 117 Hz. To define the input-output characteristics of the neurons, the first- and second-order Wiener kernels were computed by a cross-correlation between the spike response of the afferents and the white noise stimulus. 3. White noise stimulation elicited sustained spiking in 50 out of 54 afferents throughout the 20 s periods of stimulation and recording. The first-order kernels, the linear response properties, of these afferents were of six basic types that were dependent on the cutoff frequency of the white noise stimulus. These included 1) flexion-sensitive afferents that were primarily position sensitive irrespective of stimulus frequency, 2) flexion-sensitive afferents that were position sensitive at low frequencies but also coded velocity at higher frequencies, 3) flexion-sensitive afferents that coded velocity at all stimulus frequencies, 4) flexion-sensitive afferents that coded velocity at low stimulus frequencies but also acceleration at high frequencies, 5) extension-sensitive afferents that coded velocity at all stimulus frequencies, and 6) extension-sensitive afferents that coded velocity at low stimulus frequencies and acceleration at high frequencies. A seventh type contained the four remaining afferents that adapted rapidly to the stimulus within 3-5 s. These were all extension-acceleration sensitive irrespective of stimulus frequency. 4. The gain curves (produced by Fourier transform of the 1st-order kernels) and the power spectra of the linear models (produced by convolving the 1st-order kernels with the white noise) demonstrated that responses in the position-sensitive afferents are representative of a constant gain low-pass filter with a cutoff frequency of approximately 80 Hz, whereas those in the velocity- and acceleration-sensitive afferents are band passed, having peaks at 80 Hz. 5. The main nonlinearity was a signal compression in which the diagonal peak(s) of the second-order nonlinear kernels offset one or more peaks of the first-order kernels and represents a rectification or directional sensitivity of the afferents.(ABSTRACT TRUNCATED AT 400 WORDS)

Afferent Pathways↗

The structure, response properties and development of a hair plate on the mesothoracic leg of the locust.

A hair plate is present on the proximal anterior face of the pro- and mesothoracic tibiae of the legs of the locust Schistocerca gregaria, but not on the metathoracic legs. The hair plate is in a depression of the cuticle and contains about 11 hairs, which are all polarised with their tips pointing towards the dorsal surface of the tibia. The hairs are all of the same trichoid sensilla type and vary in length from 90 to 140 microns. Associated with the hair plate is a pronounced distal extension of the anterior femoral coverplate, the inner face of which is concave, that makes contact with the hairs during flexion and extension movements of the tibia. During postembryonic development, no tibial hair plate hairs are present in the first four larval stages. In fifth-instar larvae just three hairs are present, while the full complement is attained only after the final moult to adulthood. The distal extension of the posterior coverplate is present through all instar stages, becoming more pronounced after each moult. Sensory neurones innervating the hairs of an adult may be divided into two classes on the basis of their responses. The first type responds phasically to imposed deflections and is velocity-sensitive. The second type responds phasotonically and is also sensitive to the velocity of the stimulus but has an additional tonic component sensitive to maintained angular deflections. Both types of afferents are directionally sensitive and respond best to deflections against the natural bend of the hair, equivalent to extension movements of the tibia.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Processing of mechanosensory information from gustatory receptors on a hind leg of the locust.

Gustatory receptors (basiconic sensilla) on the legs of the desert locust, Schistocerca gregaria, are innervated by chemosensory afferents and by a mechanosensory afferent. We show, for the first time, that these mechanosensory afferents form an elaborate detector system with the following properties: 1) they have low threshold displacement angles that decrease with increasing stimulus frequency in the range 0.05-1 Hz, 2) they respond phasically to deflections of the receptor shaft and adapt rapidly to repetitive stimulation, 3) they encode the velocity of the stimulus in their spike frequency and have velocity thresholds lower than 1 degree/s, and 4) they are directionally sensitive, so that stimuli moving proximally towards the coxa elicit the greatest response. The mechanosensory afferents, but not the chemosensory afferents, make apparently monosynaptic connections with spiking local interneurones in a population with somata at the ventral midline of the metathoracic ganglion. They evoke excitatory synaptic potentials that can sum to produce spikes in the spiking local interneurones. Stimulation of the single mechanosensory afferent of a gustatory receptor can also give rise to long lasting depolarizations, or to bursts of excitatory postsynaptic potentials in the interneurones that can persist for several seconds after the afferent spikes. These interneurones are part of the local circuitry involved in the production of local movements of a leg. The mechanosensory afferents from gustatory receptors must, therefore, be considered as part of the complex array of exteroceptors that provide mechanosensory information to these local circuits for use in adjusting, or controlling locomotion.

Adaptation, Physiological↗

Convergence of mechanosensory afferents from different classes of exteroceptors onto spiking local interneurons in the locust.

Tactile stimuli to a leg of a locust are detected by two classes of exteroceptors: trichoid sensilla (tactile hairs), which are purely mechanoreceptors, and basiconic sensilla, which serve a dual role as mechano- and chemoreceptors. The trichoid sensilla have the longer shafts, but because the two types of receptors are intermingled over the surface of a leg, both can be excited when the leg contacts an obstacle. This article analyzes the mechanisms by which the tactile sensory information from these two classes of receptor is collated in the CNS so that spatial information is preserved. The mechanosensory afferents from both types of receptors on a hind leg make convergent connections that are excitatory and apparently direct with the same spiking local interneurons in a midline population of the metathoracic ganglion. The anatomy of the afferent projections suggests that the synapses from the two types of receptor are made onto the same region of branches of an interneuron. The slower conduction velocity of the spikes of the basiconic afferents compared to that of the trichoid afferents means that a mechanical stimulus to the tibia will first produce a depolarization caused by the trichoid afferents, followed up to 20 msec later by a depolarization caused by the basiconic sensilla. Each interneuron is excited by a contiguous and overlapping array of trichoid and basiconic receptors that form its receptive field. Different interneurons have different receptive fields such that the surface of the leg is mapped as a series of overlapping receptive fields. Within a receptive field the contribution of individual receptors can be markedly different: some basiconic receptors evoke large-amplitude EPSPs so that each afferent spike can cause a spike in the interneuron, whereas others generate small EPSPs that must sum with other inputs to evoke a spike. The amplitudes of the EPSPs generated by different receptors within a receptive field form gradients that are distributed according to the axes of the leg. The most effective basiconic and the most effective trichoid sensilla within a particular receptive field can be at the same or at different locations on the leg. Tactile sensory information from different types of receptor on one region of a leg is therefore collated initially by particular spiking local interneurons. This organization ensures summation between the inputs from the different types of receptors and should enhance sensitivity.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗