Auditory acuity in the orientation behaviour of the bushcricket Pachysagella australis walker (Orthoptera,Tettigoniidae, Saginae)

The female tettigoniid Pachysagella australis (Saginae) orients to the call of the conspecific with an angular acuity of ±5°. This acuity is mediated by sound entering to the tympanic receptors through the auditory trachea and the slits exterior to the tympanic membranes. The phonokinetic response of females was filmed in an arena. The slit system was blocked on both anterior and posterior ports with the effect that the female spiralled towards the sound source; blocking the posterior slit alone reduced the auditory acuity as did the partial occlusion of the fore slit, but in both cases the female located the male. Complete blocking of the auditory spiracle caused the insect to spiral towards the unoperated side, whereas reducing the sound input to one side by some 8–12 dB, by plugging the auditory bulla with compacted cotton wool, did not substantially affect the orientation pattern of the insect. Ablation of the tympanic organ on one side caused the female to move in a circular pattern to the unoperated side. A hypothesis is formulated whereby the female, when actively orienting to the calling male, may use the slit port system to gain a high degree of auditory acuity to sound in front of her body axis. When off target she may use the less accurate spiracular input system. This system, with its greater sensitivity to high frequency sounds, may function as an effective anti-predator warning system.     

Sensory interneurons: some observations concerning the physiology and related structural significance of two cells in the crayfish brain

Two large interneurons in the crayfish brain which are sensitive to vibrational stimuli were injected with the fluorescent dye Procion Yellow. The dendritic branching profiles reflect the directional sensitivity of their respective mechanoreceptive fields on the cephalic appendages and integument. One interneuron branches exclusively on the contralateral side of the brain and receives monosynaptic input from the contralateral antenna; the second interneuron branches primarily on the ipsilateral side and is more sensitive to input from ipsilateral receptors although its receptive field is bilateral. The data suggest that these cells are primary and secondary sensory interneurons, respectively.     

Hearing and frequency dependence of auditory interneurons in the parasitoid fly <Emphasis Type="Italic">Homotrixa alleni</Emphasis> (Tachinidae: Ormiini)

The parasitoid tachinid fly Homotrixa alleni detects its hosts by their acoustic signals. The tympanal organ of the fly is located at the prothorax and contains scolopidial sensory units of different size and orientation. The tympanal membrane vibrates in the frequency range of approximately 4–35 kHz, which is also reflected in the hearing threshold measured at the neck connective. The auditory organ is not tuned to the peak frequency (5 kHz) of the main host, the bush cricket Sciarasaga quadrata. Auditory afferents project in the three thoracic neuromeres. Most of the ascending interneurons branch in all thoracic neuromeres and terminate in the deutocerebrum of the brain. The interneurons do not differ considerably in frequency tuning, but in their sensitivity with lowest thresholds around 30 dB SPL. Suprathreshold responses of most neurons depend on frequency and intensity, indicating inhibitory influence at higher intensities. Some neurons respond particularly well at low frequency sounds (around 5 kHz) and high intensities (80–90 dB SPL), and thus may be involved in detection of the primary host, S. quadrata. The auditory system of H. alleni contains auditory interneurons reacting in a wide range of temporal patterns from strictly phasic to tonic and with clear differences in frequency responses.     

Morphological and physiological identification of medulla interneurons in the visual system of the tiger beetle larva

The morphology of visual interneurons in the tiger beetle larva was identified after recording their responses. Stained neurons were designated as either medulla or protocerebral neurons according to the location of their cell bodies. Medulla neurons were further subdivided into three groups. Afferent medulla neurons extended processes distally in the medulla neuropil and a single axon to the brain through the optic nerve. They received their main input from stemmata on the ipsilateral side. Two distance-sensitive neurons, near-by sensitive and far-sensitive neurons, were also identified. Atypical medulla neurons extended their neurites distally in the medulla and proximally to the brain, as afferent medulla neurons, but their input patterns and the shapes of their spikes differed from afferent neurons. Protocerebral neurons sent a single axon to the medulla neuropil. They spread collateral branches in the posterior region of the protocerebrum on its way to the medulla neuropil. They received main input from stemmata on the contralateral side. Medulla intrinsic neurons did not extend an axon to the brain, and received either bilateral or contralateral stemmata input only. The input patterns and discharge patterns of medulla neurons are discussed with reference to their morphology.     

A gain-control mechanism for processing of chorus sounds in the afferent auditory pathway of the bushcricket Tettigonia viridissima (Orthoptera; Tettigoniidae)

The representation of alternative conspecific acoustic signals in the responses of a pair of local interneurons of the bushcricket Tettigonia viridissima was studied with variation in intensity and the direction of sound signals. The results suggest that the auditory world of the bushcricket is rather sharply divided into two azimuthal hemispheres, with signals arriving from any direction within one hemisphere being predominantly represented in the discharge of neurons of this side of the auditory pathway. In addition, each pathway also selects for the most intense of several alternative sounds. A low-intensity signal at 45 dB sound pressure level is quite effective when presented alone, but completely suppressed when given simultaneously with another signal at 60 dB sound pressure level. In a series of intracellular experiments the synaptic nature of the intensity-dependent suppression of competitive signals was investigated in a number of interneurons. The underlying synaptic mechanism is based on a membrane hyperpolarisation with a time-constant in the order of 5–10 s. The significance of this mechanism for hearing in choruses, and for the evolution of acoustic signals and signalling behaviour is discussed. Accepted: 20 November 1999     

Central projections of first-order ocellar interneurons in the bug Triatoma infestans (Heteroptera: Reduviidae)

The projections of first-order ocellar interneurons were analyzed in the hematophagous bug Triatoma infestans by cobalt filling. The axons run between the calyces of the mushroom bodies and dorsal of the central body to different regions of the brain and the subesophageal and thoracic ganglia. The interneurons can be grouped into large L cells and small S cells. The L cells have cell bodies ranging from 11.5 to 25 μm and axons ranging from 8 to 25 μm diameter (measured in the ocellar nerve); the S cells have smaller cell bodies of 9 μm or less and axon diameters less than 5 μm. The projections of ten L cells are described in detail; they project to the protocerebral posterior slope (PS), the other ocellus (O), the optic neuropile, and the subesophageal, pro-, meso-, and metathoracic ganglia, either to ipsi- (PS I, II), or contra- (PS IV, V), or bilateral areas. In this case projections occur to the same areas (PSO, PS III) or different areas at each side (PSOE; E = eye). Large-descending (LD) first-order interneurons project to the contralateral posterior slope of the protocerebrum, the deutocerebrum, and subesophageal, pro-, mesa-, and metathoracic areas (LD I-III). Cell bodies are located in the dorsal protocerebral lobes and pars intercerebralis, except the PS II neuron and three LD cells, which are located in the ipsilateral posterior protocerebrum. This is the first report about ocellar pathways in Hemiptera. Their adaptive function is discussed with reference to the bugs' behavior as Chagas disease vectors. © 1996 Wiley-Liss, Inc.     

Etude topographique des interneurones ocellaires et de quelques uns de leurs prolongements chez Periplaneta americana

The topography of the largest ocellar interneurons in the brain of the cockroach Periplaneta americana was shown with cobalt chloride. The ocellar interneurons coloured from one nerve are confined to the ipsilateral side of the pars intercerebralis; their number and their position vary along the ocellar tract. If two ocellar nerves colour from the ocelli, the interneurons show a bilateral symmetry. Only one interneuron runs through the brain between each ocellus and the contralateral connective to the mesothoracic ganglion. When the injection of cobalt chloride is done without any current from the ocellus, the second-order ocellar neurons only are coloured, but when it is done using a current the higher order interneurons are also coloured.Axonal iontophoresis from a cervical connective back into the brain, has revealed that the cellular body of the contralateral higher-order interneuron is situated in the postero-ventral part of the protocerebrum. This pericaryon with a long cellular process is the largest of the ocellar ones (Ø = 50–60 μm). These results are discussed in relation to the ocellar and visual pathways of Schistocerca.     

Spatial and spectral dependence of the auditory periphery in the northern leopard frog

We investigated directionalities of eardrum vibration and auditory nerve response in anesthetized northern leopard frogs (Rana pipiens pipiens). Simultaneous measures of eardrum velocities and firing rates from 282 auditory nerve fibers were obtained in response to free-field sounds from eight directions in the horizontal plane. Sound pressure at the external surface of the ipsilateral eardrum was kept constant for each presentation direction (± 0.5 dB). Significant effects of sound direction on eardrum velocity were shown in 90% of the cases. Maximum or minimum eardrum velocity was observed more often when sounds were presented from the lateral and posterior fields, or from the anterior and contralateral fields, respectively. Firing rates of 38% of the fibers were significantly affected by sound direction and maximum or minimum firing rate was observed more frequently when sounds were delivered from the lateral fields, or from the anterior and contralateral fields, respectively. Directionality patterns of eardrum velocity and nerve firing also vary with sound frequency. Statistically significant correlation between eardrum velocity and nerve fiber firing rate was demonstrated in only 45% of the fibers, suggesting that sound transmission to the inner ear through extratympanic pathways plays a non-trivial role in the genesis of directionality of auditory nerve responses.Abbreviations CF characteristic frequency - SVL snout-vent length - TM tympanic membrane     

Cephalic influences on a defensive behaviour in the dogbane tiger moth, Cycnia tenera

ABSTRACT. The dogbane tiger moth ( Cycnia tenera Hübner; Arctiidae) responds to ultrasonic, artificial bat echolocation signals by emitting stereotyped trains of high-frequency, rapidly repeated clicks. By comparing this response in intact and headless moths, the role of protocerebral auditory inter-neurones suggested by other studies was examined. Individual moths were tested intact and decapitated, and their response differences analysed. Response latency and threshold (dB) did not alter with the removal of the head but response duration and responsiveness to stimulus trains were significantly reduced in headless moths. These data are interpreted as suggesting the existence of a reflex arc connecting the moth's tympanic organ (ear) with its sound-producing structure (tymbal), and as providing preliminary evidence that the role of higher-order interneurones is primarily that of response reinforcement.     

Anatomy of the ocellar interneurons of acridid grasshoppers

Summary The anatomy of the small ocellar interneurons in the brain of the acridid grasshopper Schistocerca vaga was revealed by cobalt-filling the three ocellar nerves and subsequent reconstructions from silver-intensified (Timm's method) serial sections.In total, 61 small ocellar interneurons were repeatedly identified with arborizations in many areas of the brain and optic lobe, including in particular the posterior neuropil, ocellar tracts, protocerebral bridge, lobula, ventral bridge and tritocerebral crotch, calyces, and antenno-glomerular tracts.Each ocellar nerve contains the axons of small cells that arborize in the other two ocellar tracts; these tracts are sites of ocellar integration. Direct interactions between the ocelli and compound eyes are suggested by the projections of small ocellar interneurons into the proximal lobula. Small cell arborizations from all three ocelli are distributed across much of the protocerebral bridge, implying a role for the bridge as an ocellar neuropil within the brain.Four of the small interneurons could be seen in whole-mount preparations and are demonstrated to be identical in five species of acridid grasshoppers of two different subfamilies: Schistocerca vaga, S. gregaria, Gastrimargus africanus, Trimerotropis pallidipennis, and Arphia conspersa.     

Responses of the less sensitive acoustic sense cells in the tympanic organs of some noctuid and geometrid moths

The separate impulses contributed by the A1 and A2 acoustic sense cells in the tympanic organs of the noctuids, Autographa pseudogamma and Noctua c.-nigrum, and by the A1, A2, and A3 sense cells in the tympanic organ of the geometrid, Ennomos magnarius, were identified and counted from oscillograms grams made as the moths were exposed to ultrasonic pulses of different intensities. These data were used to construct curves relating the response/intensity characteristics of the less sensitive acoustic sense cells to that of the most sensitive unit, A1. The A2 sense cells of the noctuids were found to be from 20 to 30 dB less sensitive than A1 at sound frequencies to which these ears are most sensitive. In the geometrid it was found that the A2 sense cell was 15 dB less sensitive than A1 and 12 dB more sensitive than A3. Only traces of the response of the fourth geometrid acoustic sense cell (A4) could be identified at high sound intensities. In both noctuids and geometrids the acoustic sensitivity of A2 relative to A1 remained unchanged when tested at selected ultrasonic frequencies between 28 and 50 kHz. This confirms the conclusion that the ears of these moths are incapable of pitch discrimination over this frequency range. Each of the systems had a dynamic range of 40 to 45 dB, that of the geometrid showing greater range overlap of the four A cells and hence greater capacity for sound intensity discrimination.     

Physiological properties of afferents and synaptic reorganization in the rat cerebellum degranulated by postnatal x-irradiation

Elimination of most granule, basket, and stellate interneurons in the rat cerebellum was achieved by repeated doses of low level x-irradiation applied during the first two weeks of postnatal life. Electrical stimulation of the brain stem and peripheral limbs was employed to investigate the properties of afferent cerebellar pathways and the nature of the reorganized neuronal synaptic circuitry in the degranulated cerebellum of the adult. Direct contacts of mossy fibers on Purkinje cells were indicated by short latency, single spike responses: 1.9 msec from the lateral reticular nucleus of brain stem and 5.4 msec from ipsilateral forlimb. These were shorter than in normal rats by 0.9 and 2.1 msec, respectively. The topography of projections from peripheral stimulation was approximately normal. Mossy fiber responses followed stimulation at up to 20/sec, whereas climbing fiber pathways fatigued at 10/sec. The latency of climbing fiber input to peripheral limb stimulation in x-irradiated cerebellum was 23 ± 8 (SD) msec. In x-irradiated rats, the climbing fiber pathways evoked highly variable extracellular burst responses and intracellular EPSPs of different, discrete sizes. These variable responses suggest that multiple climbing fibers contact single Purkinje cells. We conclude that each type of afferent retains identifying characteristics of transmission. However, rules for synaptic specification appear to break down so that: (1) abnormal classes of neurons develop synaptic connections, i.e., mossy fibers to Purkinje cells; (2) incorrect numbers of neurons share postsynaptic targets, i.e., more than one climbing fiber to a Purkinje cell; and (3) inhibitory synaptic actions may be carried out in the absence of the major inhibitory interneurons, i.e., Purkinje cell collaterals may be effective in lieu of basket and stellate cells.     

Antennal movements induced by odour and central projection of the antennal neurones in the honey-bee

Movements of the antennae induced by odour were investigated. Odour presented to the antenna of one side induced both antennae to move to that side. The EMGs recorded from the flexor muscles of both scapes showed that the latency of the movement of the ipsilateral flagellum when induced by odour was about 71 msec shorter than that of the contralateral flagellum. Recording electrical activities from the antennal nerve showed that there are more than 14 neurones in the antenno-motor externus.The distribution of the antennal nerve in the brain was investigated histologically by the injection of fluorescent dye. Antennal sensory neurones terminated at the glomeruli in the antennal lobe, in the dorsal lobe, in the protocerebrum, and in the commissural part of the suboesophageal ganglion. Injection of the fluorescent dye into the antennal nerve after degeneration of the antennal sensory neurones showed that the antennal motoneurones run in the ventral side of the antennal and dorsal lobes, and terminate in the marginal region of the ipsilateral oesophageal connective.The difference in latency of odour-induced flagellar movements is discussed in relation to the histological results and the unitary responses in the brain reported previously.     

The organization of plurisegmental mechanosensitive interneurons in the central nervous system of the wandering spider Cupiennius salei

Summary In spiders the bulk of the central nervous system (CNS) consists of fused segmental ganglia traversed by longitudinal tracts, which have precise relationships with sensory neuropils and which contain the fibers of large plurisegmental interneurons. The responses of these interneurons to various mechanical stimuli were studied electrophysiologically, and their unilateral or bilateral structure was revealed by intracellular staining. Unilateral interneurons visit all the neuromeres on one side of the CNS. They receive mechanosensory input either from a single leg or from all ipsilateral legs via sensory neurons that invade leg neuromeres and project into specific longitudinal tracts. The anatomical organization of unilateral interneurons suggests that their axons impart their information to all ipsilateral leg neuromeres. Bilateral interneurons are of two kinds, symmetric and asymmetric neurons. The latter respond to stimulation of all legs on one side of the body, having their dendrites amongst sensory tracts of the same side of the CNS. Anatomical evidence suggests that their terminals invade all four contralateral leg neuromeres. Bilaterally symmetrical plurisegmental interneurons have dendritic arborizations in both halves of the fused ventral ganglia. They respond to the stimulation of any of the 8 legs. A third class of cells, the ascending neurons have unilateral or bilateral dendritic arborizations in the fused ventral ganglia and show blebbed axons in postero-ventral regions of the brain. Their response characteristics are similar to those of other plurisegmental interneurons. Descending neurons have opposite structural polarity, arising in the brain and terminating in segmental regions of the fused ventral ganglia. Descending neurons show strong responses to visual stimulation. Approximately 50% of all the recorded neurons respond exclusively to stimulation of a single type of mechanoreceptor (either tactile hairs, or trichobothria, or slit sensilla), while the rest respond to stimulation of a variety of sensilla. However, these functional differences are not obviously reflected by the anatomy. The functional significance of plurisegmental interneurons is discussed with respect to sensory convergence and the coordination of motor output to the legs. A comparison between the response properties of certain plurisegmental interneurons and their parent longitudinal tracts suggests that the tracts themselves do not reflect a modality-specific organization.Abbreviations BPI bilateral plurisegmental interneuron - CNS central nervous system - FVG fused ventral ganglia - LT longitudinal tract - PI plurisegmental interneuron - PSTH peristimulus timehistogram - UPI unilateral plurisegmental interneuron     

Synaptic mechanisms of monaural and binaural processing in the locust

(1) Responses of auditory interneurones were recorded intracellularly within the metathoracic ganglion of the locust when stimulating each tympanic membrane with a piezoelectric transducer. Thus, in contrast to conventional sound stimulation, each of the two ears could be activated independently from the other at variable intensities, duration and stimulus onsets. By means of this ‘earphone-like’ stimulation technique the binaural integration properties of auditory interneurons could be analysed. (2) A minority of units (3 out of 43) was affected by input from one side only. Their synaptic input was purely excitatory and the intensity characteristics reflected those of auditory receptor fibres. (3) Most interneurones received input from both ears, each being excitatory or one excitatory or one excitatory and one inhibitory. In some units the unilateral synaptic response already included both an EPSP and an IPSP. As a result of varying temporal interactions between the EPSP and the IPSP within the unilaterally evoked complex response the intensity characteristics differed widely from unit to unit. (4) With binaural simultaneous stimulation the complexity of the postsynaptic responses of most interneurones increased as the synaptic input from both ears coincided at the level of the recorded interneurone. Although both ears were stimulated symmetrically (at the same time and intensity), units were recorded where the latencies of ipsilateral and contralateral synaptic input were different. Contralateral inhibition could either follow or precede ipsilateral excitation and in some cases both EPSP and IPSP had the same latency. On the basis of these findings the binaural synaptic mechanisms of directional coding are discussed and compared with corresponding results under free field stimulus conditions.     

Auditory evoked potential in normal rats and after para-chlorophenylalanine (PCPA) treatment

We have studied the latency behaviour of an early component of the cortical acoustic evoked potentials (EAEP) in albino rats after administration of p-chlorophenylalanine (PCPA), a rather selective tryptophan-hydroxylase inhibitor, at a dose of 100 mg/kg daily for 3 days. The rats were implanted with 3 chronic electrodes: one in the bregma, one in the nasion and 3rd inserted in the periauricular skin. Series of clicks originating from a square pulse of 0.12 msec duration were administrated. Brain responses were amplified by an EEG and averaged by a computer with different post-stimuli analysis times. A first group of 4 rats was tested with clicks of 100 dB (HTL) intensity and brain responses were analysed at 5,10,25,50,100 msec post-stimuli times. Results demonstrate that after PCPA administration there is a latency reduction of EAEP components that have a latency higher than 20 msec. In a second group of 4 rats we have analysed those EAEP components with an intensity of clicks ranging from 60 to 110 dB and results demonstrate that, when PCPA was administered, Latencies of those components were significatively lower than the controls at each stimuli intensity tested. We concluded that 5-HT may influence the acoustic pathways activity and this is according to remarks of other A.A. that found a correlation between acoustic stress and brain 5-HT levels.     

Spectrum of the calling songs, phonotaxis and the auditory system in the cricket Gryllus bimaculatus]

Behavioural experiments with Y-maze showed that phonotaxis in female crickets to male calling songs (CS) depends on the spectrum of the latter. Conservation of the first low-frequency (5 kc. p. s.) component of the spectrum is the necessary and sufficient condition for the development of normal phonotaxis. Signals which in their temporal characteristics are identical to the CS, but their spectrum contains only high-frequency (12.5 kc. p. s.) component, do no evoke positive phonotaxis. High-frequency signals (10-40 kc. p. s.) induce negative phonotaxis of females in the stationary flight. Beginning from the tympanic organ, the auditory system of crickets exhibits distinct differentiation of elements, which provide the analysis of low- and high-frequency signals. Two types of ascending interneurons transmitting information about the sound from the first auditory center to the brain were described in detail. The first type is associated mainly with low-frequency receptors and effectively transmits all that is necessary for the recognition of temporal characteristics of the CS. The second type presumably accounts for the negative phonotaxis. It is associated mainly with high-frequency receptors, exhibits for the negative phonotaxis. It is associated mainly with high-frequency receptors, exhibits significant after-effect, higher sensitivity to sounds of weak intensities, emphasizes the onset of the stimulus effect, and rapidly habituates to repetitive stimulation.     

Cartesian representation of stimulus direction: Parallel processing by two sets of giant interneurons in the cockroach

The cockroachPeriplaneta americana responds to wind puffs by turning away, both on the ground and when flying. While on the ground, the ventral giant interneurons (ventrals) encode the wind direction and specify turn direction, whereas while flying the dorsal giant interneurons (dorsals) appear to do so. We report here on responses of these cells to controlled wind stimuli of different directions. Using improved methods of wind stimulation and of positioning the animal revealed important principles of organization not previously observed.All six cells of largest axonal diameter on each side respond preferentially to ipsilateral winds. One of these cells, previously thought to respond non-directionally (giant interneuron 2), was found to have a restricted directional response (Fig. 3). The organization of directional coding among the ventral giant interneurons is nearly identical to that among the dorsals (Fig. 2). Each group contains, on each side, one cell that responds primarily to wind from the ipsilateral front, another primarily in the ipsilateral rear, and a third responding more broadly to ipsilateral front and rear.These results are discussed in terms of the mechanisms of directional localization by the assembly of giant interneurons.Abbreviations GI giant interneuron - vGI ventral giant interneuron - dGI dorsal giant interneuron - CF 5-carboxyfluorescein - A6 6th abdominal ganglion - TI thoracic interneuron - BED best excitatory direction     

SOUND AND ITS SIGNIFICANCE FOR LABORATORY ANIMALS

1. Several methods of varying accuracy have been used to assess what sounds small laboratory animals such as rodents are capable of hearing. Most rodents can detect sounds from 1000 Hz (the frequency of the Greenwich Time Signal) up to 100000 Hz, depending on the strain, with usually one or more commonly two peaks of sensitivity within this range. Dogs can detect sound most easily from 500 Hz to 55000 Hz, depending on the breed. 2. Rodents also produce sound signals as a behavioural response and for communication in a variety of situations. Ultrasonic calls in the range 22000–70000 Hz are the main communicating pathway during aggressive encounters, mating, and mothering. Similar calls have also been recorded from isolated animals associated with inactivity, rest and possibly even sleep. 3. Very loud sounds cause seizures in rats and mice, or can make them more susceptible to other sounds later in life. This effect is possible even when animals are fully anaesthetized. Sound tends to startle and reduce activity in several species of animal. Even offspring of mice that have been sound-stressed exhibit abnormal behaviour patterns. Sounds also elicit various responses in rats from increasing aggression to making them more tolerant to electric shocks. 4. Levels of sound above 100 dB are teratogenic in several species of animals and several hormonal, haematological and reproductive parameters are disturbed by sounds above 80 dB. When rats are chemically deafened the disturbance to their fertility disappears. Lipid metabolism is disrupted in rats when exposed to over 95 dB of sounds, leading to increases in plasma triglycerides. Atherosclerosis can be produced in rabbits by similar levels of sound. 5. It has also been shown in guinea pigs and cats that hearing damage is governed by the duration as well as the intensity of the sound and is irreversible. Work on chinchillas hs demonstrated that sounds above 95 dB lead to this injury, but that sounds of 80 dB have no permanent effect on hearing sensitivity.