Extremely Low Frequency Electromagnetic Fields impair the Cognitive and Motor Abilities of Honey Bees.

Extremely low frequency electromagnetic field (ELF EMF) pollution from overhead powerlines is known to cause biological effects across many phyla, but these effects are poorly understood. Honey bees are important pollinators across the globe and due to their foraging flights are exposed to relatively high levels of ELF EMF in proximity to powerlines. Here we ask how acute exposure to 50 Hz ELF EMFs at levels ranging from 20-100 µT, found at ground level below powerline conductors, to 1000-7000 µT, found within 1 m of the conductors, affects honey bee olfactory learning, flight, foraging activity and feeding. ELF EMF exposure was found to reduce learning, alter flight dynamics, reduce the success of foraging flights towards food sources, and feeding. The results suggest that 50 Hz ELF EMFs emitted from powerlines may represent a prominent environmental stressor for honey bees, with the potential to impact on their cognitive and motor abilities, which could in turn reduce their ability to pollinate crops.

Olfactory selection of Plantago lanceolata by snails declines with seedling age.

BACKGROUND AND AIMS: Despite recent recognition that (1) plant-herbivore interactions during the establishment phase, (2) ontogenetic shifts in resource allocation and (3) herbivore response to plant volatile release are each pivotal to a comprehensive understanding of plant defence, no study has examined how herbivore olfactory response varies during seedling ontogeny. METHODS: Using a Y-tube olfactometer we examined snail (Helix aspersa) olfactory response to pellets derived from macerated Plantago lanceolata plants harvested at 1, 2, 3, 4, 5, 6 and 8 weeks of age to test the hypothesis that olfactory selection of plants by a generalist herbivore varies with plant age. Plant volatiles were collected for 10 min using solid-phase microextraction technique on 1- and 8-week-old P. lanceolata pellets and analysed by gas chromatography coupled with a mass spectrometer. KEY RESULTS: Selection of P. lanceolata was strongly negatively correlated with increasing age; pellets derived from 1-week-old seedlings were three times more likely to be selected as those from 8-week-old plants. Comparison of plant selection experiments with plant volatile profiles from GC/MS suggests that patterns of olfactory selection may be linked to ontogenetic shifts in concentrations of green leaf volatiles and ethanol (and its hydrolysis derivatives). CONCLUSIONS: Although confirmatory of predictions made by contemporary plant defence theory, this is the first study to elucidate a link between seedling age and olfactory selection by herbivores. As a consequence, this study provides a new perspective on the ontogenetic expression of seedling defence, and the role of seedling herbivores, particularly terrestrial molluscs, as selective agents in temperate plant communities.

Coding characteristics of spiking local interneurons during imposed limb movements in the locust.

The performance of adaptive behavior relies on continuous sensory feedback to produce relevant modifications to central motor patterns. The femoral chordotonal organ (FeCO) of the legs of the desert locust monitors the movements of the tibia about the femoro-tibial joint. A ventral midline population of spiking local interneurons in the metathoracic ganglia integrates inputs from the FeCO. We used a Wiener kernel cross-correlation method combined with a Gaussian white noise stimulation of the FeCO to completely characterize and model the output dynamics of the ventral midline population of interneurons. A wide range of responses were observed, and interneurons could be classified into three broad groups that received excitatory and inhibitory or principally inhibitory or excitatory synaptic inputs from the FeCO. Interneurons that received mixed inputs also had the greatest linear responses but primarily responded to extension of the tibia and were mostly sensitive to stimulus velocity. Interneurons that received principally inhibitory inputs were sensitive to extension and to joint position. A small group of interneurons received purely excitatory synaptic inputs and were also sensitive to tibial extension. In addition to capturing the linear and nonlinear dynamics of this population of interneurons, first- and second-order Wiener kernels revealed that the dynamics of the interneurons in the population were graded and formed a spectrum of responses whereby the activity of many cells appeared to be required to adequately describe a particular stimulus characteristic, typical of population coding.

Nitric oxide modulates sodium taste via a cGMP-independent pathway.

Insects, like other animals, require sodium chloride (NaCl) as part of their normal diet and detect it with contact chemoreceptors on the body surface. By adjusting the responsiveness of the chemosensory neurons within these receptors insects can modify the intake of salt and other nutrients, and it has been hypothesized that the responsiveness of chemosensory neurons is regulated by nitric oxide (NO). To identify potential sources of NO in the periphery, the authors applied the NO-sensitive fluorescent probe 4,5-diaminofluorescein and the universal NO synthase antibody, and found that in locusts NO is synthesized within one particular class of cells of the epidermis, the glandular cells, from where it may diffuse to neighboring chemosensory neurons. The effects of NO on chemosensory neurons were investigated by recording from contact chemoreceptors on the leg while perfusing it with drugs that interfere with NO signaling. Results showed that both endogenous and exogenous NO decreased the frequency of action potentials in chemosensory neurons in response to stimulation with NaCl by acting via a cyclic guanosine monophosphate-independent pathway. Variation of the NaCl concentration in the perfusion solution demonstrated that the synthesis of NO in glandular cells depends on the NaCl concentration in the hemolymph. By contrast NO increased the frequency of action potentials in chemosensory neurons in response to sucrose stimulation. The authors suggest that NO released from glandular cells modulates the responsiveness of chemosensory neurons to regulate NaCl intake, and hypothesize that NO may play a key role in the signaling of salt and sugars.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

Processing of tactile information in neuronal networks controlling leg movements of the Locust.

The successful, coordinated, posture and locomotion of any animal requires a precise and continuous adjustment of limb movements by sensory feedback from extero- and proprioceptors associated with the legs. We here review the recent advances in our understanding of how specific local adjustments of the hind legs of the desert locust, Schistocerca gregaria, are made in response to tactile signals from two different classes of exteroceptor on a leg. The aim is to understand particular features of the organization of neuronal networks and how different types of constituent interneurones contribute to the processing of sensory signals. This information can then be used to define the design principles that govern the organization of sensory-motor networks.

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.

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.

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)